Contents

PREFACE 1
▪ Introduction to Human Biology 1
▪ About OpenStax Resources 1
▪ About Human Biology – An Introduction to the Body’s Organization, Structure, & Function 2
▪ Pedagogical Foundation 2
▪ Subscription site (Biology411.com) with Student Learning and Assessment Resources 2
▪ About the Author 3
▪ Instructor Resources 4

CHAPTER 1
Introduction to Human Biology and the Scientific Method …..6
1.1 Introduction to Human Biology 6
1.2 Structural Organization of the Human Body 7
1.3 Functions of Human Life 12
• Organization 12
• Metabolism 12
• Responsiveness 13
• Movement 14
• Development 14
• Growth 15
• Reproduction 15
• Regulation 15
• Evolution and Adaptations 20
1.4 The Scientific Method 21
• Proposing a Hypothesis 22
• Testing a Hypothesis 23
• Reporting Scientific Work 23
1.5 Career Connection – Technical Writer 25

CHAPTER 2
Chemistry and Life …………………………………………………………………..27
2.1 Introduction 27
2.2 Elements and Atoms: The Building Blocks of Matter 28
• Atoms and Subatomic Particles 29
• Atomic Number and Mass Number 32
• Isotopes 34
• The Behavior of Electrons 35
2.3 Chemical Bonds 37
• Ions and Ionic Bonds 38
• Covalent Bonds 40
▪ Nonpolar Covalent Bonds 41
▪ Polar Covalent Bonds 41
• Water’s Polarity 44
2.4 pH, pH Scale, Acidic, and Basic/Alkaline 45
 2.5 Organic Compounds Essential to Human Functioning 47
• The Chemistry of Carbon 47
• Carbohydrates 49
▪ Monosaccharides 50
▪ Disaccharides 50
▪ Polysaccharides 51
• Lipids 52
▪ Triglycerides 52
▪ Phospholipids 55
▪ Steroids 55
• Proteins 56
▪ Microstructure 57
▪ Shape 58
▪ Enzymes 61
▪ Other Functions 62
• Nucleic Acids 63
▪ Chemical Features 63
▪ Adenosine triphosphate 66
2.6 Career Connection – Pharmaceutical Chemist 67

CHAPTER 3
Cells and Associated Topics …………………………………………………..….69
3.1 Introduction 69
3.2 Microscopy, Cell Theory, and Cell Size 70
3.3 Cell Types 73
• Prokaryotic Cells 74
• Eukaryotic Cells 75
3.4 Components and Structure of the Plasma Membrane 76
3.5 Transport Across the Plasma Membrane 80
• Passive Transport 80
• Active Transport 86
3.6 The Cytoplasm and Cellular Organelles 91
• Cytoplasm 92
• Cytoskeleton 92
• Nucleus 92
• Ribosomes 93
• Mitochondria 94
• Organelles of the Endomembrane System 95
▪ Endoplasmic reticulum 95
▪ Golgi Apparatus 97
3.7 Career Connection – Cytotechnologist 98

CHAPTER 4
DNA, Genes, RNA, and Protein Synthesis …………………………99
4.1 Introduction 99
4.2 Organization of the Nucleus and its DNA 100
4.3 DNA Replication 106
 4.4 From DNA to Protein (Transcription and Translation) 109
• Transcription: DNA to RNA 110
• Translation: mRNA to Protein 112
• Mutations and Their Impact on Proteins 118
4.5 Career Connection – Forensic Scientists and DNA Analysis 119

CHAPTER 5
Digestive System ………………………………………………………………….120
5.1 Introduction 120
5.2 Homeostasis 122
5.3 Tissues 123
5.4 Anatomy of the Digestive System 127
• Oral Cavity and Pharynx 127
• Esophagus 130
• Stomach 131
• Small Intestine 132
• Large Intestine 134
• Accessory Organs 135
5.5 Physical and Chemical Digestion 137
5.6 Physical Digestion of Food 138
5.7 Chemical Digestion of Food 138
5.8 Integrated Functioning Between the Digestive and Endocrine Systems 143
5.9 Absorption 144
5.10 Elimination 145
5.11 Career Connection – Gastroenterologist 146

CHAPTER 6
Energy Considerations .............................................................................................. 147
6.1 Introduction 147
6.2 Energy and the Laws of Thermodynamics 149
6.3 Anabolic and Catabolic Chemical Reactions 151
6.4 Enzymes 153
6.5 ATP and Related Topics 155
6.6 Electrons, Energy, and Electron Carriers 157
6.7 An Overview of Cellular Respiration 159
6.8 Stage 1: Glycolysis 160
6.9 Stage 2: Transition Reaction 161
6.10 Stage 3: Krebs Cycle 163
6.11 Stage 4: The Electron Transport Chain and Oxidative Phosphorylation 164
6.12 Summary ATP Yield from Cellular Respiration 167
6.13 Metabolism without Oxygen: Lactic Acid and Alcohol Fermentation 169
6.14 Summarizing Cellular Respiration and Fermentation 172
6.15 Connections of Carbohydrate, Protein, and Lipid Metabolic Pathways 174
6.16 Calories 175
6.17 Disorders of Metabolism – Phenylketonuria and Prader-Willi Syndrome 178
6.18 Career Connection – Registered Dietician 180
CHAPTER 7
Cardiovascular System I - Blood………………………………………………181
7.1 Introduction 181
7.2 Characteristics of Blood 182
7.3 Functions of Blood 182
• Transportation 182
• Defense 183
• Maintenance of Homeostasis 183
7.4 Composition of Blood 183
7.5 Erythrocytes 185
• Shape and Structure of Erythrocytes 185
• Hemoglobin 186
• Life Cycle of Erythrocytes 189
• Disorders of Erythrocytes 189
7.6 Leukocytes 192
7.7 Platelets (Thrombocytes) and Clotting Factors 194
7.8 Plasma Proteins and Other Solutes 196
7.9 Bone Marrow Sampling and Transplants 196
7.10 The Genetic Basis of the ABO and Rh Blood Types 197
7.11 Blood Type Antibodies, Agglutination, and ABO and Rh Blood Typing 201
7.12 Blood Transfusions and Adverse Reactions 202
7.13 Rh-Incompatibility, Pregnancy, and RhoGAM 205
7.14 Career Connection – Phlebotomy and Medical Lab Technology 206

CHAPTER 8
Cardiovascular System II - Heart…………………………………………...….208
8.1 Introduction 208
8.2 Human Heart and Blood Vessels 209
8.3 Heart Valves 213
8.4 The Cardiac Cycle 217
8.5 Electrocardiogram (ECG or EKG) 222
8.6 Disease of the Heart – Myocardial Infarction 228
8.7 Blood Pressure 229
8.8 Pulse 232
8.9 Career Connection – Cardiologist 233

CHAPTER 9
Cardiovascular System III - Blood Vessels………………………………234
9.1 Introduction 234
9.2 Types of Blood Vessels and Their Relative Structures 234
9.3 Systemic and Pulmonary Circuits 237
9.4 How Blood Flows Through the Body 238
9.5 Exchange of Fluids at the Capillaries and the Lymphatic System 240
9.6 Cardiac Blood Vessels 241
9.7 Strokes 246
 CHAPTER 10
Respiratory System ………………………………………………………….……….248
10.1 Introduction 248
10.2 Organs and Structures of the Respiratory System 249
10.3 The Mechanics of Breathing 254
• Atmospheric and Partial Pressure 254
• Boyle’s Law 256
• Inhalation and Exhalation 257
10.4 Basic Principles of Respiratory Gas Exchange 259
10.5 Oxygen and Carbon Dioxide Transport and Exchange During External and Internal
Respiration 260
• Blood Transport of Oxygen 260
• Blood Transport of Carbon Dioxide 262
• External Respiration of Oxygen and Carbon Dioxide 263
• Internal Respiration of Oxygen and Carbon Dioxide 263
10.6 Medical Issues Associated with the Respiratory System 265
• Asthma 265
• Pneumonia 267
• Carbon Monoxide Poisoning 268
• Sleep Apnea 269
• Decompression Sickness 270
• Acute Mountain Sickness 271
10.7 Career Connection – Respiratory Therapist 273

CHAPTER 11
Endocrine System ……………………………………………………………………..274
11.1 Introduction 274
11.2 Endocrine Signaling 274
11.3 Structures of the Endocrine System 276
• Types of Hormones 277
• How Hormones Work 277
• Regulating Hormone Secretion 280
11.4 The Hypothalamus and Pituitary Gland 281
11.5 The Thyroid Gland, Adrenal Glands, and Gonads 285
• Thyroid Gland 285
• Adrenal Glands 286
▪ Adrenal Cortex 286
▪ Adrenal Medulla 287
• Pancreas 288
• Gonads 289
11.6 Selected Disorders of the Endocrine System 290
• Iodine Deficiency, Hypothyroidism, and Hyperthyroidism 290
• Diabetes Mellitus 291
11.7 Hormones and Athletic Performance 293
11.8 Career Connection – Endocrinologist 293
CHAPTER 12
Urinary System ……………………………………………………………………….....295
12.1 Introduction 295
12.2 Physical Characteristics of Urine 296
12.3 Gross Anatomy of the Urinary System 299
• Kidneys 299
• Ureters 300
• Bladder 301
• Sphincters 302
• Urethra 303
12.4 Internal Gross Anatomy of the Kidney 304
12.5 Microscopic Anatomy of the Kidney 305
12.6 Blood Flow Through the Kidney 308
12.7 Stages of Urine Formation 309
12.8 Hormones that Influence Urine Formation 311
12.9 Career Connections – Dialysis Technician and Nephrologist 312

CHAPTER 13
Cell Division …………………………………………………………………………….…314
13.1 Introduction 314
13.2 Genomic DNA, Chromosomes, and Compaction 315
13.3 The Cell Cycle 317
• Interphase 319
• The Mitotic Phase: Mitosis and Cytokinesis 321
13.4 Cell Cycle Control 323
13.5 Sexual Reproduction and Meiosis Overview 326
• Life Cycle of Sexually Reproducing Organisms 326
• Meiosis Overview 327
13.6 The Process of Meiosis 328
• Meiosis I 328
▪ Prophase I 329
▪ Metaphase I 331
▪ Anaphase I 333
▪ Telophase I and Cytokinesis 333
• Meiosis II 333
▪ Telophase II and Cytokinesis 334
13.7 Comparing Meiosis and Mitosis 336
13.8 Disorders Associated with Cell Division 338
• Cancer 338
▪ Proto-Oncogenes 338
▪ Tumor-Suppressor Genes 339
• Chromosome Number Disorders 343
▪ Aneuploidy 344
▪ Sex Chromosomes Nondisjunction in Humans 345
13.9 Career-Connection – Cytogenetic Technologist 347
CHAPTER 14
Reproductive Systems ………………………………………………………..……..349
14.1 Introduction 349
14.2 Male Reproductive Anatomy 349
14.3 Spermatogenesis 354
14.4 Male Reproductive Hormones 356
14.5 Female Reproductive Anatomy 358
14.6 Oogenesis 362
14.7 The Ovarian Cycle, Menstrual Cycle, and Female Hormonal Regulation 364
14.8 Fertilization and Twins 370
14.9 Development of the Fetal Male and Female Reproductive Systems 370
14.10 Pregnancy, Gestation, and Birth 371
14.11 Contraception and Birth Control 375
14.12 Infertility, IVF, and “Designer Babies” 377
14.13 Career Connection – Reproductive Endocrinologist 386

CHAPTER 15
Skeletal System ...................................................................................... 388
15.1 Introduction 388
15.2 Human Axial Skeleton 389
• The Skull 389
• The Rib Cage 390
• The Vertebral Column 392
• The Auditory Ossicles and the Hyoid Bone 392
15.3 Human Appendicular System 394
• The Pectoral (Upper) Limbs 395
• The Pelvic (Lower) Limbs 396
• The Pectoral Girdle 398
• The Pelvic Girdle 398
15.4 Bone Tissue 399
• Compact Bone Tissue 399
• Spongy Bone Tissue 400
15.5 Bone Classification and Gross Anatomy 401
15.6 Cell Types in Bone 404
15.7 Development and Growth of Bone 405
• Development of Bone 405
• Lengthening of Long Bones 406
• Thickening of Long Bones 406
15.8 Bone Remodeling and Repair 406
15.9 Calcium Homeostasis 409
15.10 Nutrition and Bone Tissue 412
15.11 Diseases and the Skeletal System 414
• Osteoporosis 414
• Osteogenesis Imperfecta 415
15.12 Career Connection – Rheumatologist 416
CHAPTER 16
Muscular System and Movement ..................................................... 418
16.1 Introduction 418
16.2 Skeletal Muscle Structure and Function 419
16.3 Skeletal Muscle Fiber Structure 421
16.4 Skeletal Muscle Contraction and Relaxation 423
16.5 Sources of ATP for Muscle Contraction 435
16.6 Performance-Enhancing Substances 438
16.7 Disorder of the Muscular System – Duchenne Muscular Dystrophy 439
16.8 Career Connection – Physical Therapist 439

CHAPTER 17
Nervous System ..................................................................................... 441
17.1 Introduction 441
17.2 Nervous System Categorization 442
17.3 Nervous Tissue 447
17.4 The General Function of Nervous Tissue 449
17.5 The Action Potential 450
• Electrically Active Cell Membranes 450
• The Membrane Potential 453
• The Action Potential 455
• Propagation of the Action Potential 459
17.6 Communication Between Neurons 462
• Chemical Synapses 462
• Signal Summation 464
17.7 Nervous System Disorders 466
• Neurogenerative Disorders 467
▪ Multiple Sclerosis 467
▪ Alzheimer’s Disease 467
▪ Parkinson’s Disease 469
▪ Amyotrophic Lateral Sclerosis (Lou Gehrig’s Disease) 470
• Neurodevelopmental Disorders 473
▪ Autism 473
▪ Attention Deficit Hyperactivity Disorder (ADHD) 474
• Mental Illnesses 475
▪ Schizophrenia 475
▪ Depression 476
• Other Neurological Disorders 477
▪ Epilepsy 477
▪ Stroke 478
17.8 Career Connection – Neurologist 479

CHAPTER 18
Sensation, Somatosensation, and Special Senses ........................ 480
18.1 Introduction 480
18.2 Sensory Processes 481
• Reception 481
• Transduction 481
• Perception 483
18.3 Somatosensation 484
18.4 Smell and Taste 486
• Odors and Tastes 487
• Odor Reception and Transduction 487
• Taste Reception and Transduction 490
18.5 Hearing 492
• Sound 493
• Sound Reception 494
• Sound Transduction 495
18.6 Vision 497
• Light 497
• Light Reception 499
• Light Transduction 502
18.7 Career Connection – Audiologist, Orthoptist, and Optometrists 503
• Audiologist 503
• Orthoptist and Optometrists 504

CHAPTER 19
Immune and Lymphatic Systems ...................................................... 505
19.1 Introduction 505
19.2 The Innate Immune Response 506
• Barrier Defenses (Physical and Chemical) 507
• Cells of the Innate Immune Response 509
• Inflammatory Response 509
• Soluble Mediators of the Innate Immune Response 513
▪ Cytokines 513
▪ Complement System 514
19.3 The Adaptive Immune Response 515
• Antigen Presenting Cells 515
• T and B Lymphocytes 517
• Immunological Memory 520
• Antibodies 523
• Mechanisms of Acquiring Immunity 524
19.4 The Lymphatic System 527
• Functions of the Lymphatic System 527
• Structure of the Lymphatic System 528
▪ Lymphatic Capillaries 528
▪ Larger Lymphatic Vessels, Trunks, and Ducts 530
19.5 Primary Centers of the Immune System 531
19.6 Disruptions of the Immune System 535
• Immunodeficiency 535
• Hypersensitivities 535
• Allergies 536
• Autoimmunity 537
19.7 Career Connection – Vaccinologist 538
CHAPTER 20
Mendelian Genetics and Associated Topics ................................... 540
20.1 Introduction 540
20.2 Mendelian Genetics 541
• Gregor Mendel 541
• Mendel’s Crosses 542
• A Modern Interpretation of Mendel’s Crosses 546

• Law of Dominance 548

• Using a Punnett Square with Monohybrid Crosses 550
• Law of Segregation 552
• Test Cross 554
• Law of Independent Assortment 555
20.3 Probability Basics 558
• The Product Rule 558
• The Sum Rule 559
20.4 Extensions of Mendel’s Laws 560
• Incomplete Dominance 561
• Codominance 562
• Multiple Alleles 562
• Sex-Linked Traits 565
• Epistasis 568
• Linked Genes Violate the Law of Independent Assortment 571
20.5 Pedigree Analysis 573
• Autosomal Recessive Inheritance 574
• Autosomal Dominant Inheritance 575
• X-Linked Dominant or Recessive Inheritance 576

CHAPTER 21
Evolution and Population Genetics .................................................. 580
21.1 Introduction 580
21.2 A Brief History of the Theory of Evolution by Natural Selection 581
21.3 Understanding Darwin’s Theory of Natural Selection 583
21.4 The Processes of Evolution 586
• Mutation 586
• Meiosis 588
• Genetic Drift 591
• Gene Flow 593
21.5 Evidence of Evolution 593
• Fossils 594
• Anatomy 594
• Embryology 597
• Biogeography 598
• Molecular Biology 599
21.6 Misconception of Evolution 599
• Evolution is Just a Theory 600
• Individuals Evolve 600
• Evolution Explains the Origin of Life 600
• Organisms Evolve on Purpose 601
21.7 Taxonomy, Phylogeny, and Speciation 602
• Taxonomy 602
• Phylogeny and Phylogenetic Trees 603
• Speciation 608
21.8 A Brief Look at Human Evolution 609
• The Evolution of Homo sapiens 609
• Are Humans Still Evolving 610
21.9 Genetic Changes in Populations 611
• Allele Frequencies 612
• Hardy-Weinberg Principle of Equilibrium 612
• Adaptive Evolution 614
• Stabilizing Selection 616
• Directional Selection 616
• Diversifying Selection 616
• No Perfect Organism 617
21.10 Career Connection – Field Biologist 619

CHAPTER 22
Molecular Biology and Biotechnology ............................................. 620
22.1 Introduction 620
22.2 Molecular Biology Techniques 621
• Basic Techniques to Manipulate Genetic Materials (DNA and RNA) 621
• DNA and RNA Extraction 621
• Gel Electrophoresis 621
• Hybridization, Southern Blotting, and Northern Blotting 623
• Cloning 625
• In-vivo (Bacterial) Cloning 625
• In-vitro Cloning: Polymerase Chain Reaction (PCR) 627
• Reproductive Cloning 631
• DNA Sequencing 633
22.3 Biotechnology in Medicine and Agriculture 635
• Genetic Diagnosis and Gene Therapy 636
• Production of Vaccines, Antibiotics, and Hormones 636
• Transgenic Animals 637
• Transgenic Plants 638
22.4 Genomics and Its Applications 640
• Predicting Disease Risk at the Individual Level 640
• Gene Editing 641
• Pharmacogenomics 643

CHAPTER 23
Ecology, Populations, and Biodiversity ........................................... 644
23.1 Introduction 644
23.2 Ecosystems and Energy 645
• Ecosystems and Disturbances 647
• Food Chains and Food Webs 647
• How Organisms Acquire Energy in a Food Web 651
23.3 Biogeochemical Cycles 653
• The Water Cycle 654
• The Carbon Cycle 656
• The Biological Carbon Cycle 657
• The Biogeochemical Carbon Cycle 658
• The Nitrogen Cycle 659
23.4 Population Demographics 661
• Population Size and Density 661
• Estimating Population Size 662
• Species Distribution 663
• Demography 664
23.5 Population Growth and Carrying Capacity 666
• Exponential Growth 667
• Logistic Growth 668
• Carrying Capacity and the Logistic Model 668
• Examples of Logistic Growth 669
23.6 The Human Population 670
• Age Structure, Population Growth, and Economic Development 672
• Long-Term Consequences of Exponential Human Population Growth 674
23.7 What is Biodiversity, and Why is it Important? 675
• Types of Biodiversity 676
• Genetic and Chemical Diversity 676
• Ecosystems Diversity 677
• Current Species Diversity 677
• Importance of Biodiversity 678
• Human Health 679
• Agriculture 680
• Wild Food Sources 682
• Psychological and Moral Value 683
23.8 Threats to Biodiversity 683
• Habitat Loss 683
• Overharvesting 685
• Exotic Species 686
• Climate Change 688
23.9 Preserving Biodiversity 690
• Changing Human Behavior 690
• Conservation in Preserves 691
• Habitat Restoration 694
• The Role of Zoos and Captive Breeding 696
23.10 Career Connections – Biogeographer and Ecologist 697
• Biogeographer 697
• Ecologist 697

Index…………………………………………………………………………… …….…699
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”‡•‘—” ‡•ǣ

x Key Term Activities: ƒ”‹‘—• Šƒ’–‡”Ǧ„‘Ž†‡†–‡”‹‘Ž‘‰›ƒ –‹˜‹–‹‡•ƒ”‡ƒ˜ƒ‹Žƒ„Ž‡

ȋ‡Ǥ‰ǤǡˆŽƒ•Š ƒ”†•ǡ’‹ ǦƒǦŽ‡––‡”ǡˆ‹ŽŽǦ‹Ǧ–Š‡Ǧ„Žƒǡƒ– Š‹‰ǡ ”‘••™‘”†•ǡƒ†‰Ž‘••ƒ”›ȌǤ
–—†‡–• ƒ ‘’›ƒ†’ƒ•–‡–‡”•ƒ††‡ˆ‹‹–‹‘•‹–‘ƒ†‘ —‡––‘ ”‡ƒ–‡ —•–‘
Šƒ’–‡”‰Ž‘••ƒ”‹‡•Ǥ

x Pronunciation of Key Termsǣ–—†‡–•Ž‹‡Ž›™‘̵–”‡‡„‡”–‡”•‹ˆ–Š‡› ƒǯ–

‘””‡ –Ž›’”‘‘— ‡–Š‡ǤŠ‡”‡ˆ‘”‡ǡ˜‹†‡‘•‘ˆ–Š‡ƒ—–Š‘”’”‘‘— ‹‰‡ƒ Š
Šƒ’–‡”ǯ•‡›–‡”•ƒ”‡‹ Ž—†‡†Ǥ

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”‡ˆƒ ‡

x Short, Lecture-Type Videosǣ‘•–ˆ‹‰—”‡•‹‡ƒ Š Šƒ’–‡”ƒ”‡–ƒ‰‰‡†™‹–Šƒ‘–‹ ‡

–Šƒ–•Š‘”–ȋ–›’‹ ƒŽŽ›Ž‡••–Šƒ–‡‹—–‡•Ȍ†‹• —••‹‘•˜‹†‡‘•ƒ”‡ƒ˜ƒ‹Žƒ„Ž‡ǤŠ‡•‡
˜‹†‡‘•ǡ ”‡ƒ–‡†„›–Š‡ƒ—–Š‘”ǡ’”‘˜‹†‡ Ž‡ƒ”ǡ ‘ ‹•‡•—ƒ”‹‡•‘ˆ‹’‘”–ƒ–ƒ†
”‡Ž‡˜ƒ–‹ˆ‘”ƒ–‹‘–‘ˆƒ ‹Ž‹–ƒ–‡•–—†‡–Ž‡ƒ”‹‰Ǥˆ–‡”‡ƒ Š˜‹†‡‘ǡ”‡˜‹‡™
“—‡•–‹‘•ƒ”‡’”‡•‡–‡†ǡƒŽ‘‰™‹–Šƒ†‹• —••‹‘‘ˆ–Š‡ ‘””‡ –ƒ•™‡”•ǤŽŽ˜‹†‡‘•
ƒ”‡ ƒ’–‹‘‡†–‘‡•—”‡ƒ ‡••‹„‹Ž‹–›Ǥ

x Assessment Activities: Š‡•‡ƒ –‹˜‹–‹‡•ƒŽŽ‘™•–—†‡–•–‘ƒ’’Ž›ƒ†‡˜ƒŽ—ƒ–‡–Š‡‹”

Ž‡ƒ”‹‰ƒ†—†‡”•–ƒ†‹‰‘ˆ–Š‡ ‘–‡–ǤŠ”‡‡–›’‡•‘ˆƒ••‡••‡–•ǡ ”‡ƒ–‡†„›
–Š‡ƒ—–Š‘”ǡƒ”‡ƒ˜ƒ‹Žƒ„Ž‡ǣ•‡ŽˆǦƒ••‡••‡–™‘”•Š‡‡–•ȋ †‘ —‡–•Ȍǡ‘Ž‹‡
ƒ –‹˜‹–‹‡•ǡƒ†‘Ž‹‡’”ƒ –‹ ‡“—‹œœ‡•Ǥ—‡•–‹‘–›’‡•‹ Ž—†‡•Š‘”–ƒ•™‡”ǡ
—Ž–‹’Ž‡ Š‘‹ ‡ǡ–”—‡‘”ˆƒŽ•‡ǡ—Ž–‹’Ž‡”‡•’‘•‡•ǡ‡˜‡–•‡“—‡ ‹‰ǡƒ– Š‹‰ǡˆ‹ŽŽǦ
‹Ǧ–Š‡Ǧ„Žƒǡ†”ƒ‰ƒ††”‘’ǡƒ†Š‘–•’‘–ǤŠ‡ƒ –‹˜‹–‹‡•ƒ†’”ƒ –‹ ‡“—‹œœ‡• ƒ„‡
”‡’‡ƒ–‡†•‡˜‡”ƒŽ–‹‡•ˆ‘”ƒ††‹–‹‘ƒŽ’”ƒ –‹ ‡Ǥ

About the Author

‹ŽŽƒ—•Š™ƒǡŽƒ”‘ŽŽ‡‰‡ƒ†ƒ•Š‹‰–‘–ƒ–‡‹˜‡”•‹–›Ȃƒ ‘—˜‡”Ǥ

‹ŽŽ‹ƒȋ‹ŽŽ›Ȍ—•Š™ƒ‹•ƒ‹•–”— –‘”™‹–Š͵ͲΪ›‡ƒ”•‘ˆ• ‹‡ ‡–‡ƒ Š‹‰‡š’‡”‹‡ ‡ƒ–

–Š‡Š‹‰Š• Š‘‘Žǡ ‘—‹–› ‘ŽŽ‡‰‡ǡƒ†—‹˜‡”•‹–›Ž‡˜‡Ž•Ǥ ‡‹•’ƒ••‹‘ƒ–‡ƒ„‘—–• ‹‡ ‡
‡†— ƒ–‹‘ƒ†‡Œ‘›•–Š‡ ŠƒŽŽ‡‰‡•ƒ••‘ ‹ƒ–‡†™‹–Š ”‡ƒ–‹‰ Žƒ••”‡•‘—” ‡•–Šƒ–ƒ”‡™‡ŽŽǦ
†‡•‹‰‡†ǡ‡‰ƒ‰‹‰ǡƒ†‹ˆ‘”ƒ–‹˜‡Ǥ ‡”‡•‹†‡•‹–Š‡„‡ƒ—–‹ˆ—Žƒ ‹ˆ‹ ‘”–Š™‡•–ƒ†
‡Œ‘›•–‹‡™‹–Šˆƒ‹Ž›ǡ‘—–†‘‘”ƒ –‹˜‹–‹‡•ǡƒ†Š‘‡Ǧ”‡Žƒ–‡†’”‘Œ‡ –•Ǥ

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”‡ˆƒ ‡

Instructor Resources

Š‡‹–‡•ƒ”‡ƒ˜ƒ‹Žƒ„Ž‡–‘‹•–”— –‘”•™Š‘ƒ†‘’––Š‹•–‡š–„‘‘ƒ† ‘–ƒ ––Š‡ƒ—–Š‘”

ȋ™ —•Š™ƒ̷„‹‘Ž‘‰›ͶͳͳǤ ‘Ȍ–‘˜‡”‹ˆ›–Š‡‹”‹•–‹–—–‹‘ƒˆˆ‹Ž‹ƒ–‹‘ǣ

Chapter Image PowerPoint Slides: Š‡‘™‡”‘‹–•Ž‹†‡•‹ Ž—†‡ƒŽŽ Šƒ’–‡”ˆ‹‰—”‡•ƒ†

†‡• ”‹’–‹‘•ƒ•ƒ•–ƒ”–‹‰’Žƒ ‡ˆ‘”‹•–”— –‘”•–‘„—‹Ž†–Š‡‹”Ž‡ –—”‡•Ǥ

Reading Study Guides (RSGs):ƒ Š Šƒ’–‡”Šƒ•ƒƒ••‘ ‹ƒ–‡† ǡ™Š‹ ŠŽ‹•–•–Š‡•’‡ ‹ˆ‹ 

–‡”•ƒ†ˆ‹‰—”‡•–Šƒ–•–—†‡–•ƒ”‡”‡•’‘•‹„Ž‡ˆ‘”‘™‹‰ƒ†—†‡”•–ƒ†‹‰ǤŠ‡”‡‹•
ƒŽ•‘ƒŽ‹•–‘ˆ•–—†›“—‡•–‹‘•ǡ™”‹––‡„›–Š‡ƒ—–Š‘”ǡ–Šƒ–‰—‹†‡•–—†‡–•–Š”‘—‰Š–Š‡”‡ƒ†‹‰
ƒ†Š‡Ž’–Š‡’”‘†— –‹˜‡Ž›‡‰ƒ‰‡™‹–Š–Š‡ ‘–‡–Ǥ•™‡”•ƒ”‡ǯ–’”‘˜‹†‡†„‡ ƒ—•‡
†‘‹‰•‘ ƒ†‹‹‹•Š•–—†‡–‡ˆˆ‘”–ƒ†–Š‡Ž‹‡Ž‹Š‘‘†‘ˆ‡‰ƒ‰‹‰™‹–Š‘–Š‡”•–—†‡–•
ƒ†Ȁ‘”–Š‡‹•–”— –‘”–‘†‹• —••–Š‡“—‡•–‹‘•Ǥ

Š‡•‡‰—‹†‡•ƒ†‘–Š‡” ‘—”•‡”‡•‘—” ‡••Š‘—Ž†„‡–Š‡„ƒ•‹•ˆ‘”ƒŽŽƒ••‡••‡–“—‡•–‹‘•ǡ

•‘•–—†‡–•ǯƒš‹‡–›ƒ„‘—–DzŠƒ– ‘–‡–ƒ ”‡•’‘•‹„Ž‡ˆ‘”‘™‹‰ƒ†
—†‡”•–ƒ†‹‰ǫdz‹•”‡†— ‡†Ǥ

ƒ Š ‹•ƒ˜ƒ‹Žƒ„Ž‡ƒ•ƒ‘”††‘ —‡–ǡ•‘‹– ƒ•‡”˜‡ƒ•ƒ•–ƒ”–‹‰’‘‹–ˆ‘”‹•–”— –‘”•

–‘ ”‡ƒ–‡ƒ —•–‘‰—‹†‡–Šƒ–‹•ƒ’’”‘’”‹ƒ–‡ˆ‘”–Š‡‹” Žƒ••ȋ‡•ȌǤ

Lab Simulations: Š‹•–‡š–„‘‘‹• ‘’ƒ–‹„Ž‡™‹–Š™‹†‡˜ƒ”‹‡–›‘ˆ–”ƒ†‹–‹‘ƒŽ —ƒ

‹‘Ž‘‰›Žƒ„ˆ‘”ƒ–•ƒ†ƒ –‹˜‹–‹‡•ǤŠ‡ƒ—–Š‘”Šƒ• ”‡ƒ–‡†ƒ•‡”‹‡•‘ˆŽƒ„•‹—Žƒ–‹‘
Šƒ†‘—–•„ƒ•‡†‘Gizmosǡƒ•‹—Žƒ–‹‘’Žƒ–ˆ‘”‘ˆˆ‡”‡†„›š’Ž‘”‡‡ƒ”‹‰
ȋ‰‹œ‘•Ǥ‡š’Ž‘”‡Ž‡ƒ”‹‰Ǥ ‘ȌǤ”‹ ‹‰‹ˆ‘”ƒ–‹‘‹•ƒ˜ƒ‹Žƒ„Ž‡ƒ–
‰‹œ‘•Ǥ‡š’Ž‘”‡Ž‡ƒ”‹‰Ǥ ‘Ȁƒ„‘—–Ǧ‰‹œ‘•ȀŠ‘™Ǧ–‘Ǧ‰‡–Ǧ‰‹œ‘•Ǥ

ƒ Š•‹—Žƒ–‹‘Šƒ•ƒ‹–”‘†— –‘”›Šƒ†‘—–ǡ™Š‹ Š‰—‹†‡••–—†‡–•–Š”‘—‰Š–Š‡

ˆ‘—†ƒ–‹‘ƒŽ‹ˆ‘”ƒ–‹‘ƒ†–Š‡‡š’‡”‹‡–ȋ•Ȍǡƒ†ƒ”‡’‘”–ˆ‘”–Šƒ–•–—†‡–• ‘’Ž‡–‡
ƒ†•—„‹–ˆ‘”‰”ƒ†‹‰Ǥ‘’‹ •‹ Ž—†‡–Š‡• ‹‡–‹ˆ‹ ‡–Š‘†ǡŠ‘‡‘•–ƒ•‹•ǡ‡œ›‡•ǡ
‡‹‘•‹•ƒ†”‡Žƒ–‡†–‘’‹ •ǡ ‡ŽŽ—Žƒ””‡•’‹”ƒ–‹‘ǡ†‹ˆˆ—•‹‘ǡƒ†‘•‘•‹•Ǥ

‡˜‡”ƒŽ‘ˆ–Š‡•‹—Žƒ–‹‘•ƒ”‡ ƒ•‡•‹™Š‹ Š•–—†‡–•ƒ••—‡–Š‡”‘Ž‡‘ˆƒ

’”‘ˆ‡••‹‘ƒŽƒ†‹’Ž‡‡––Š‡• ‹‡–‹ˆ‹ ‡–Š‘†–‘ƒ•™‡”–‡•–ƒ„Ž‡“—‡•–‹‘•Ǥ

Topic Simulation ActivitiesǣŠ‡ƒ—–Š‘”Šƒ•†‡˜‡Ž‘’‡†Šƒ†‘—–•ˆ‘”™‡„Ǧ„ƒ•‡†ƒ –‹˜‹–‹‡•

‘–Š‡ˆ‘ŽŽ‘™‹‰–‘’‹ •ǣƒ–‘‹ •–”— –—”‡ǡ‹‘•ǡ Š‡‹ ƒŽ„‘†‹‰ǡ’ ǡ‘”‰ƒ‹ ‘Ž‡ —Ž‡•ǡ
„Ž‘‘†–›’‹‰ǡƒ†Š‡ƒ”–Ǧ”‡Žƒ–‡†–‘’‹ •Ǥ

ŽŽ•‹—Žƒ–‹‘†‘ —‡–•ƒ”‡’”‘˜‹†‡†ƒ•‘”†ˆ‹Ž‡•–Šƒ– ƒ„‡”‡˜‹•‡†ƒ•†‡•‹”‡†Ǥ

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”‡ˆƒ ‡

‘—‹–› —„•

’‡–ƒš’ƒ”–‡”•™‹–Š–Š‡ •–‹–—–‡ˆ‘”–Š‡–—†›‘ˆ‘™Ž‡†‰‡ƒƒ‰‡‡–‹
†— ƒ–‹‘ȋ Ȍ–‘‘ˆˆ‡”‘—‹–› —„•‘‘‘•Ȅƒˆ”‡‡’Žƒ–ˆ‘”ˆ‘”
‹•–”— –‘”•–‘•Šƒ”‡ ‘—‹–›Ǧ ”‡ƒ–‡†”‡•‘—” ‡•–Šƒ–•—’’‘”–’‡–ƒš„‘‘•Ǥ
Š”‘—‰Š‘—”‘—‹–› —„•ǡ‹•–”— –‘”• ƒ—’Ž‘ƒ†–Š‡‹”‘™ƒ–‡”‹ƒŽ•‘”
†‘™Ž‘ƒ†”‡•‘—” ‡•–‘—•‡‹–Š‡‹” ‘—”•‡•ǡ‹ Ž—†‹‰ƒ††‹–‹‘ƒŽƒ ‹ŽŽƒ”‹‡•ǡ–‡ƒ Š‹‰
ƒ–‡”‹ƒŽǡ—Ž–‹‡†‹ƒǡƒ†”‡Ž‡˜ƒ– ‘—”•‡ ‘–‡–Ǥ‡‡ ‘—”ƒ‰‡‹•–”— –‘”•–‘Œ‘‹
–Š‡Š—„•ˆ‘”–Š‡•—„Œ‡ –•‘•–”‡Ž‡˜ƒ––‘–Š‡‹”–‡ƒ Š‹‰ƒ†”‡•‡ƒ” Šƒ•ƒ
‘’’‘”–—‹–›–‘‡”‹ Š–Š‡‹” ‘—”•‡•ƒ†‡‰ƒ‰‡™‹–Š‘–Š‡”ˆƒ —Ž–›Ǥ‘”‡ƒ Š–Š‡
‘—‹–› —„•ǡ˜‹•‹–™™™Ǥ‘‡” ‘‘•Ǥ‘”‰ȀŠ—„•Ȁ‘’‡•–ƒšǤ

ͷ
‭ hapter 1: Introduction to Human Biology and the‬
C
‭Scientific Method‬

‭ igure 1.1‬ ‭A basic understanding of medical procedures,‬‭such as measuring blood‬

F
‭pressure, allows you to understand better the information medical professionals collect.‬
‭(credit:‬‭Monthly Checkup‬‭; Bryan Mason; Flickr;‬‭CC‬‭BY 2.0‬‭)‬

‭1.1 Introduction to Human Biology‬

‭ iology‬‭is the science that studies life, so‬‭human‬‭biology‬‭can be defined as the science of‬
B
‭studying humans, which is just one example of a living organism. Although you may be‬
‭taking a course in human biology as a requirement for obtaining your degree, the‬
‭knowledge you gain will serve you well in other aspects of your life. For example, a basic‬
‭understanding of the structure of your body and how it works can benefit your health by‬
‭prompting you to make healthy choices and take appropriate action when signs of illness‬
‭arise. You will also be a more informed consumer of relevant media content and a‬
‭participant in conversations with your medical providers. At some point, everyone will‬
‭have a problem with some aspect of their bodies, and your knowledge can help you to be‬
‭a better person, partner, care provider, or friend.‬
‭ he study of human biology can be subdivided into anatomy and physiology.‬ ‭Anatomy‬‭is‬
T
‭the scientific study of the body’s structures. Some structures are tiny and can only be‬
‭o bserved with a microscope. Other larger ones are easily seen without magnification.‬
‭ uman‬‭physiology‬‭is the scientific study of the chemistry‬‭and physics of the body's‬
H
‭structures and how they work together to support the functions of life, including‬
‭maintaining a sensitive balance of internal conditions called homeostasis.‬

‭6‬
‭Chapter 1: Introduction to Human Biology‬

‭1.2 Structural Organization of the Human Body‬

‭ iving things are highly organized and structured, following a hierarchy that we can‬
L
‭examine on a scale from small to large (‬‭Figure 1.2‬‭).‬
‭ ll matter in the universe is composed of one or more unique pure substances called‬
A
‭elements‬‭; familiar examples include hydrogen, oxygen,‬‭carbon, nitrogen, calcium, and‬
‭iron. The smallest unit of any of these elements is an‬‭atom‬‭, which is composed of‬
‭subatomic particles‬‭such as‬‭protons‬‭,‬‭electrons,‬‭and‬‭neutrons‬‭.‬
‭ toms join to form‬‭molecules‬‭, which are chemical structures‬‭o f at least two atoms held‬
A
‭together by‬‭c hemical bonds‬‭. Names and chemical formulas‬‭o f common molecules include‬
‭water (H‬‭2‬‭O), carbon dioxide (CO‬‭2‬‭), and glucose (C‬‭6‭H ‬ 2‬‭O‭6‬ ‭)‬ .‬
‬ ‭1

‭ any biologically important molecules are‬‭macromolecules‬‭,‬‭large molecules typically‬

M
‭formed by polymerization (a‬‭polymer‬‭is a large molecule‬‭made by combining smaller‬
‭units called‬‭monomers‬‭, which are smaller than macromolecules).‬‭An example of a‬
‭macromolecule is‬‭deoxyribonucleic acid (DNA)‬‭, and‬‭its monomer is a‬‭nucleotide‬
‭(‬‭Figure 1.3‬‭). This macromolecule contains the instructions‬‭for the structure and‬
‭functioning of all living organisms.‬
‭ ‬‭cell‬‭is the smallest independently functioning unit‬‭o f a living organism. Bacteria are‬
A
‭microscopic, independently living organisms. Each bacterium is a single cell. All living‬
‭components of the human anatomy contain cells.‬
‭ human cell typically consists of a flexible membrane enclosing cytoplasm, a water-based‬
A
‭cellular fluid with various small units called‬‭organelles‬‭that have specific functions. For‬
‭example, the‬‭mitochondrion‬‭is an organelle that produces‬‭most of a human cell’s energy.‬
‭Another example of an organelle is the‬‭nucleus‬‭, which‬‭contains most of the DNA in a‬
‭human cell.‬
‭ ‬‭tissue‬‭is a group of many similar cells (though‬‭sometimes composed of a few related‬
A
‭types) that work together to perform a specific function. There are four fundamental‬
‭tissue categories: epithelial, connective, muscle, and neural.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭7‬
‭Chapter 1: Introduction to Human Biology‬

‭ igure 1.2‬ ‭The organization of the body is often‬‭discussed in terms of six or seven‬
F
‭distinct levels of increasing complexity, from the smallest chemical building blocks to a‬
‭unique human organism. The seventh level, macromolecules, isn’t shown in the figure and‬
‭would be included between molecules and organelles. (credit: Levels of Organization in‬
‭Body.jpg‬‭; Wikimedia Commons;‬‭CC BY 4.0‬‭); A link to‬‭a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭8‬
‭Chapter 1: Introduction to Human Biology‬

‭ igure 1.3‬ ‭All molecules, including this DNA molecule,‬‭are composed of various atoms of‬
F
‭different elements joined by chemical bonds. (credit:‬‭ADN animation frame 0001.png‬‭;‬
‭“Brian0918”; Wikimedia Commons;‬‭CC0 1.0‬‭)‬

‭ n‬‭organ‬‭is an anatomically distinct body structure‬‭composed of two or more tissue‬

A
‭types. Each organ performs one or more specific physiological functions. An‬‭organ‬
‭system‬‭is a group of organs that work together to‬‭achieve significant outcomes or meet‬
‭the physiological needs of the organism.‬
‭ his book covers most of the eleven organ systems in the human body (‬‭Figures 1.4a and‬
T
‭1.4b‬‭). Assigning organs to organ systems can be imprecise‬‭since organs that “belong” to‬
‭o ne system can also have functions integral to another system. In fact, most organs‬
‭contribute to more than one system.‬
‭ he organism level is the most complex level of organization. An‬‭organism‬‭is a living being‬
T
‭that can independently perform all physiological functions necessary for life. In‬
‭multicellular organisms, including humans, all cells, tissues, organs, and organ systems‬
‭work together to maintain the organism's life and health.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭9‬
‭Chapter 1: Introduction to Human Biology‬

‭ igure 1.4a‬ ‭Organs that work together are grouped‬‭into organ systems. (credit:‬‭Organ‬
F
‭Systems I.jpg‬‭; Wikimedia Commons;‬‭CC BY 3.0‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭10‬
‭Chapter 1: Introduction to Human Biology‬

‭ igure 1.4b‬ ‭Organs that work together are grouped‬‭into organ systems. (credit:‬‭Organ‬
F
‭Systems II.jpg‬‭; Wikimedia Commons;‬ ‭CC BY 3.0‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭11‬
‭Chapter 1: Introduction to Human Biology‬

‭1.3 Functions of Human Life‬

‭ ach organ system has unique physiological functions to perform. These many functions‬
E
‭are summarized in terms of characteristics that we consider definitive of human life:‬
‭o rganization, metabolism, responsiveness, movement, development, growth, reproduction,‬
‭regulation/homeostasis, and evolution.‬

‭Organization‬
‭ rganisms are highly organized, coordinated structures with one or more cells. Even very‬
O
‭simple, single-celled organisms are remarkably complex: inside each cell, atoms comprise‬
‭molecules. These, in turn, make up cell organelles and other cellular components. In‬
‭multicellular organisms, such as humans, similar cells form tissues. Tissues, in turn,‬
‭collaborate to create organs, which are body structures with distinct functions. Organs‬
‭work together to establish organ systems.‬
‭ he body’s largest organ system is the integumentary system, which includes the skin and‬
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‭its associated structures, such as hair and nails. Skin, the surface tissue, is a barrier that‬
‭protects internal structures and fluids from potentially harmful microorganisms and other‬
‭toxins.‬

‭Metabolism‬
‭ he‬‭first law of thermodynamics‬‭holds that energy‬‭cannot be created nor destroyed but‬
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‭can only change form. Your primary function as an organism is to consume (ingest)‬
‭energy and molecules in the foods you eat, convert some of it into fuel for movement,‬
‭sustain your body functions, and build and maintain your body structures. Two types of‬
‭reactions accomplish this: anabolism and catabolism.‬
‭•‬ ‭ nabolism‬‭is the process of combining smaller, simpler‬‭molecules into larger, more‬
A
‭complex substances. Your body can assemble, by utilizing energy, the complex‬
‭chemicals it needs by combining small molecules derived from the foods you eat.‬

‭•‬ ‭ atabolism‬‭is the process of breaking larger, more‬‭complex substances down into‬
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‭smaller, simpler molecules. Catabolism releases energy. The complex molecules found‬
‭in foods are broken down so the body can use their parts to assemble the structures‬
‭and substances needed for life.‬
‭ hese two processes, taken together, are called metabolism.‬‭Metabolism‬‭is the sum of all‬
T
‭anabolic and catabolic reactions in the body (‬‭Figure‬‭1.5‬‭). Both anabolism and catabolism‬
‭o ccur simultaneously and continuously to keep you alive.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 1: Introduction to Human Biology‬

‭ igure 1.5‬‭Metabolism includes both anabolic and catabolic‬‭reactions.‬‭Anabolic reactions‬

F
‭are building reactions that consume energy. (credit:‬ ‭103 Metabolism.jpg‬‭; Wikimedia‬
‭Commons;‬‭CC BY 4.0‬‭).‬ ‭A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ very cell in your body uses a chemical compound,‬‭adenosine triphosphate‬‭(‬‭ATP‬‭), to‬

E
‭store and release energy. The cell stores energy in the synthesis (anabolism) of ATP and‬
‭then moves the ATP molecules to where energy is needed to fuel cellular activities. The‬
‭ATP is broken down (catabolism), and a controlled amount of energy is released, which‬
‭the cell uses to perform particular functions.‬

‭Responsiveness‬
‭ esponsiveness‬‭is the ability of an organism to adjust‬‭to changes in its internal and‬
R
‭external environments. An example of responsiveness to external stimuli is moving toward‬
‭sources of food and water and away from perceived dangers. Changes in an organism’s‬
‭internal environment, such as increased body temperature, can cause the responses of‬
‭sweating and the dilation of blood vessels in the skin to decrease body temperature, as‬
‭shown by the runners in‬‭Figure 1.6‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 1: Introduction to Human Biology‬

‭ igure 1.6‬ ‭Marathon runners demonstrate two characteristics‬‭o f living‬

F
‭humans—responsiveness and movement. Anatomic structures and physiological‬
‭processes allow runners to coordinate the action of muscle groups and sweat in response‬
‭to rising internal body temperature. (credit: Phil Roeder/Flickr;‬‭Openstax‬‭)‬

‭Movement‬
‭ uman movement includes actions at the joints of the body and the motion of individual‬
H
‭o rgans and even individual cells. As you read these words, red and white blood cells are‬
‭moving throughout your body, muscle cells are contracting and relaxing to maintain your‬
‭posture and to focus your vision, and glands are secreting chemicals to regulate body‬
‭functions. Your body coordinates the action of entire muscle groups to enable you to‬
‭move air into and out of your lungs, push blood throughout your body, and propel the‬
‭food you have eaten through your digestive tract. Consciously, of course, you contract‬
‭your skeletal muscles to move the bones of your skeleton to get from one place to another‬
‭and perform all of your daily activities.‬

‭Development‬
‭ evelopment‬‭is all of the changes the body goes through‬‭in life. Development includes the‬
D
‭process of differentiation, in which unspecialized cells become specialized in structure and‬
‭function to perform specific tasks in the body. Development also consists of growth and‬
‭repair, which involve cell differentiation.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 1: Introduction to Human Biology‬

‭Growth‬
‭ rowth‬‭is the increase in body size. Like all multicellular‬‭o rganisms, humans grow‬
G
‭primarily by increasing the number of existing cells and the amount of non-cellular‬
‭material around cells, such as mineral deposits in bone.‬

‭Reproduction‬
‭ eproduction‬‭is the formation of a new organism from‬‭parent organisms. In humans,‬
R
‭reproduction is accomplished by the male and female reproductive systems. When‬
‭reproduction occurs, DNA-containing genes are passed along to an organism’s offspring.‬
‭These genes ensure that the offspring will belong to the same species and have similar‬
‭characteristics, such as size and shape.‬

‭Regulation/Homeostasis‬
‭ ven the smallest organisms are complex and require multiple regulatory mechanisms to‬
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‭coordinate internal functions, respond to stimuli, and cope with environmental stresses.‬
‭Two examples of internal processes regulated in an organism are nutrient transport and‬
‭blood flow. Organs (groups of tissues working together) perform specific functions, such‬
‭as carrying oxygen throughout the body, removing wastes, delivering nutrients to every‬
‭cell, and cooling the body.‬
‭ ell function requires appropriate conditions, such as proper temperature, pH, and‬
C
‭correct concentrations of various chemicals. These conditions may, however, change from‬
‭o ne moment to the next. Organisms can maintain internal conditions within a narrow‬
‭range almost constantly, despite environmental changes, through‬‭homeostasis‬‭(literally,‬
‭“steady state”). For example, an organism needs to regulate body temperature through‬
‭thermoregulation. Organisms that live in cold climates, such as the polar bear (‬‭Figure 1.7‬‭),‬
‭have body structures that help them withstand low temperatures and conserve body heat.‬
‭Structures that aid this insulation include fur, feathers, blubber, and fat. In hot climates,‬
‭o rganisms have methods (such as perspiration in humans or panting in dogs) that help‬
‭them to shed excess body heat.‬
‭ aintaining homeostasis requires that the body continuously monitor its internal‬
M
‭conditions. From body temperature to blood pressure to levels of certain nutrients, each‬
‭physiological state has a particular‬‭set point‬‭, which‬‭is the physiological value around‬
‭which the normal range fluctuates. A‬‭normal range‬‭is a restricted set of optimally healthy‬
‭and stable values. For example, the set point for average human body temperature is‬
‭approximately 37°C (98.6°F).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 1: Introduction to Human Biology‬

‭ igure 1.7‬ ‭Polar bears and other mammals living in‬‭ice-covered regions maintain their‬
F
‭body temperature by generating heat and reducing heat loss through thick fur and a‬
‭dense layer of fat under their skin. (credit: “longhorn dave”; Flickr;‬‭Openstax‬‭)‬

‭ hysiological parameters, such as body temperature and blood pressure, fluctuate within‬
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‭a normal range slightly above and below those set points. Control centers in the brain and‬
‭o ther parts of the body monitor and react to deviations from homeostasis using‬‭negative‬
‭feedback‬‭, which reverses a change from the set point.‬‭Therefore, negative feedback‬
‭maintains body parameters within their normal range. Maintaining homeostasis by‬
‭negative feedback goes on throughout the body at all times, and understanding negative‬
‭feedback is thus fundamental to understanding human physiology.‬
‭ negative feedback system has three essential components (‬‭Figure 1.8a‬‭). A‬‭sensor‬‭is a‬
A
‭component of a feedback system that monitors a physiological value. This value is‬
‭reported to the‬‭control center‬‭. The control center‬‭is the component in a feedback system‬
‭that compares the value to the normal range. The control center activates an effector if the‬
‭value deviates too much from the set point. An‬‭effector‬‭is the component in a feedback‬
‭system that causes a change to reverse the situation and return the value to the normal‬
‭range.‬
‭ o set the system in motion, a stimulus must drive a physiological parameter beyond its‬
T
‭normal range (beyond homeostasis). This stimulus is “heard” by a specific sensor. For‬
‭example, specific endocrine cells in the pancreas detect excess glucose (the stimulus) in‬
‭the bloodstream to control blood glucose. These pancreatic cells respond to the increased‬
‭blood glucose level by releasing the hormone insulin into the bloodstream. The insulin‬
‭signals skeletal muscle fibers, fat cells, and liver cells to remove excess glucose from the‬
‭bloodstream.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 1: Introduction to Human Biology‬

‭ igure 1.8‬ ‭In a negative feedback loop, a stimulus—a‬‭deviation from a set point—is‬
F
‭resisted through a physiological process that returns the body to homeostasis. (a) A‬
‭negative feedback loop has four essential parts. (b) Negative feedback regulates body‬
‭temperature. (credit:‬‭OpenStax‬‭) A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ s glucose concentration in the bloodstream drops, other pancreatic cells detect the‬
A
‭decrease in concentration—the actual negative feedback—and stop insulin release. These‬
‭actions prevent blood sugar levels from continuing to drop below the normal range.‬
‭ umans have a similar temperature regulation feedback system that promotes heat loss‬
H
‭o r heat gain (‬‭Figure 1.8b‬‭). When the brain’s temperature‬‭regulatory center receives data‬
‭from the sensors indicating that the body’s temperature exceeds its normal range, it‬
‭stimulates a cluster of brain cells called the “heat-loss center.” This stimulation has three‬
‭significant effects:‬
‭•‬ ‭ lood vessels in the skin begin to dilate, allowing more blood from the body core to‬
B
‭flow to the skin's surface, which allows heat to radiate into the environment.‬
‭•‬ ‭As blood flow to the skin increases, sweat glands are activated to increase output. As‬
‭the sweat evaporates from the skin's surface into the surrounding air, it takes heat‬
‭with it.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 1: Introduction to Human Biology‬

‭•‬ ‭ he depth of respiration increases, and a person may breathe through an open‬
T
‭mouth instead of through the nasal passageways, further increasing heat loss from‬
‭the lungs.‬
I‭ n contrast, activation of the brain’s heat-gain center by exposure to cold reduces blood‬
‭flow to the skin, and blood returning from the limbs is diverted into a network of deep‬
‭veins. This arrangement traps heat closer to the body core and restricts heat loss. If heat‬
‭loss is severe, the brain triggers an increase in random signals to skeletal muscles, causing‬
‭them to contract and produce shivering. The muscle contractions of shivering release heat‬
‭while using up ATP. The brain triggers the thyroid gland in the endocrine system to release‬
‭thyroid hormone, which increases metabolic activity and heat production in cells‬
‭throughout the body. The brain also signals the adrenal glands to release epinephrine‬
‭(adrenaline), a hormone that causes the breakdown of glycogen into glucose, which is‬
‭used as an energy source. The breakdown of glycogen into glucose also increases‬
‭metabolism and heat production.‬
‭ lthough negative feedback systems are most common, there is another type called‬
A
‭positive feedback‬‭, which intensifies a change in the‬‭body’s physiological condition rather‬
‭than reversing it. A deviation from the typical range results in more change, and the‬
‭system moves farther from the normal range. Positive feedback in the body is expected‬
‭o nly when there is a definite endpoint. Childbirth and the body’s response to blood loss‬
‭are two examples of positive feedback loops that are normal but are activated only when‬
‭needed.‬
‭ hildbirth at full term is an example of a situation in which the maintenance of the existing‬
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‭body state is not desired. Enormous changes in the mother’s body are required to expel‬
‭the baby at the end of pregnancy. The events of childbirth, once begun, must progress‬
‭rapidly to a conclusion, or the lives of the mother and the baby are at risk.‬
‭The extreme muscular work of labor and delivery results from a positive feedback system‬
‭(‬‭Figure 1.9‬‭).‬
‭ he first labor contractions (the stimulus) push the baby toward the cervix (the lowest‬
T
‭part of the uterus). The cervix contains stretch-sensitive nerve cells that monitor the‬
‭degree of stretching (the sensors). These nerve cells send messages to the brain, which‬
‭causes the pituitary gland at the base of the brain to release the hormone oxytocin into‬
‭the bloodstream. Oxytocin causes stronger contractions of the smooth muscles of the‬
‭uterus (the effectors), pushes the baby further down the birth canal, and causes even‬
‭greater cervix stretching. The cycle of stretching, oxytocin release, and increasingly more‬
‭forceful contractions stops only when the baby is born. At this point, the stretching of the‬
‭cervix halts, ceasing the release of oxytocin.‬

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‭Chapter 1: Introduction to Human Biology‬

‭ igure 1.9‬ ‭A positive feedback loop, such as labor‬‭and delivery, changes the body’s status‬
F
‭rather than causing a return to homeostasis. (credit:‬‭OpenStax‬‭) A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ second example of positive feedback is associated with reversing extreme damage to the‬
A
‭body. Following a penetrating wound, the most immediate threat is excessive blood loss.‬
‭Less blood volume means reduced blood pressure and decreased perfusion (penetration‬
‭o f blood) to the brain and other vital organs. If perfusion is severely reduced, vital organs‬
‭will shut down, and the person will die. The body responds to this potential catastrophe‬
‭by releasing substances in the injured blood vessel wall that begin the process of blood‬
‭clotting. As each step of clotting occurs, it stimulates the release of more clotting‬
‭substances and accelerates the clotting and sealing off the damaged area. Clotting is‬
‭contained in a local site based on the tightly controlled availability of clotting proteins. The‬
‭clotting mechanism is an adaptive, life-saving cascade of events.‬

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‭Chapter 1: Introduction to Human Biology‬

‭Everyday Connection‬

‭Controlled Hypothermia‬

‭ s you have learned, the body continuously engages in coordinated physiological‬

A
‭processes to maintain a stable temperature. However, overriding this system can‬
‭sometimes be helpful or even life-saving.‬

‭ ypothermia is the clinical term for an abnormally low body temperature (hypo- =‬
H
‭“below” or “under”). Controlled hypothermia is clinically induced hypothermia performed‬
‭to reduce the metabolic rate of an organ or a person’s entire body.‬

‭ ontrolled hypothermia is often used, for example, during open-heart surgery because it‬
C
‭decreases the metabolic needs of the brain, heart, and other organs, reducing the risk of‬
‭damage to them. When controlled hypothermia is used clinically, the patient receives‬
‭medication to prevent shivering. The body is cooled to 25–32°C (79–89°F), the heart is‬
‭stopped, and an external heart-lung pump maintains circulation to the patient’s body. The‬
‭heart is cooled further and is maintained at a temperature below 15°C (60°F) for the‬
‭duration of the surgery. This cold temperature helps the heart muscle tolerate its lack of‬
‭blood supply during the surgery.‬

S‭ ome emergency department physicians use controlled hypothermia to reduce damage to‬
‭the heart in patients who have suffered a cardiac arrest. In the emergency department, the‬
‭physician induces coma and lowers the patient’s body temperature to approximately 91°F.‬
‭This condition, maintained for 24 hours, slows the patient’s metabolic rate. The heart's‬
‭workload is reduced because the patient’s organs require less blood.‬

‭Evolution and Adaptations‬

‭ volution can be defined as changes that occur over long periods of time. Organisms‬
E
‭evolve in response to diverse factors such as mutations in the DNA, changes in‬
‭environment, and interactions with each other. One of the best examples of the last factor‬
‭is disease-causing organisms, which must adapt to overcome the defenses of the‬
‭o rganisms they infect. One such organism that has evolved to specialize in infecting‬
‭humans is‬‭Plasmodium‬‭, the organism that causes malaria.‬‭Scientists have suspected for‬
‭quite some time that people with blood type O are less likely to die from severe malaria.‬
‭Recently, the basis for this observation has been determined. Scientists concluded that A‬
‭and B-type blood reacts with a protein excreted by‬‭Plasmodium‬‭and causes severe illness.‬
‭However, type O blood does not react with the protein.‬
‭ ll living organisms exhibit a “fit” to their environment. Biologists refer to this fit as‬
A
‭adaptation‬‭, and it is a consequence of‬‭evolution by‬‭natural selection‬‭, which operates in‬

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‭Chapter 1: Introduction to Human Biology‬

e‭ very lineage of reproducing organisms. Examples of adaptations are diverse and unique,‬
‭from heat-resistant microorganisms that live in boiling-temperature hot springs to the‬
‭tongue length of a nectar-feeding moth that matches the size of the flower from which it‬
‭feeds. All adaptations enhance the reproductive potential of the individuals exhibiting‬
‭them, including their ability to survive and reproduce. Adaptations are not constant. As an‬
‭environment changes, natural selection causes the characteristics of the individuals in a‬
‭population to track those changes.‬
‭ he fact that biology, as a science, has such a broad scope has to do with the tremendous‬
T
‭diversity of life on Earth. Although over one million currently living species of animals‬
‭have been identified, scientists are continually discovering more species as they explore‬
‭ecosystems around the world. The number of currently living species is estimated to be‬
‭between 3 and 30 million.‬
‭ ne source of this diversity is evolution, which is the process of genetic changes in a‬
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‭population or species over time. Evolutionary biologists study the evolution of living‬
‭things in everything from the microscopic world to ecosystems. Using data obtained from‬
‭these studies, it is possible to determine lineages between organisms that had a common‬
‭ancestor at some prior point. In some cases, organisms with physical similarities were‬
‭assumed to be related; however, genetic information has revealed that isn’t necessarily‬
‭true.‬
S‭ cientists face the difficult task of classifying all living things within a unified system called‬
‭taxonomy. They must identify traits common to all organisms and characteristics that help‬
‭distinguish among related groups of animals. The animal classification system‬
‭characterizes animals based on their anatomy, morphology, evolutionary history, features‬
‭o f embryological development, and genetic makeup. This classification scheme is‬
‭constantly developing as new information about species arises. Understanding and‬
‭classifying the great variety of living species helps us better understand how to conserve‬
‭the diversity of life on Earth.‬

‭1.4 The Scientific Method‬

‭ iologists study the living world by posing questions about it and seeking science-based‬
B
‭responses. England’s Sir Francis Bacon is credited with documenting an approach known‬
‭as the‬‭scientific method‬‭to accomplish these tasks.‬ ‭The scientific method is not used only‬
‭by biologists; researchers from almost all fields of study can apply it as a logical, rational‬
‭problem-solving method.‬
‭ he scientific method (‬‭Figure 1.10‬‭) typically starts‬‭with an‬‭observation‬‭, often a problem‬
T
‭to solve, that leads to a question. Let’s think about a simple problem that starts with an‬
‭o bservation and then apply the scientific method to solve the problem. On Monday‬
‭morning, a student arrives at class and quickly discovers that the room is too warm. That‬

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‭Chapter 1: Introduction to Human Biology‬

‭ bservation also describes a problem: the classroom is too warm. The student then asks a‬
o
‭specific‬‭testable question‬‭based on the observation,‬‭such as, "Why is the classroom so‬
‭warm?”‬

‭Proposing a Hypothesis‬
‭ ecall that a‬‭hypothesis‬‭is a suggested answer to‬‭a testable question. To answer a‬
R
‭question, one can propose several hypotheses. For example, one hypothesis might be,‬
‭“The classroom is warm because no one turned on the air conditioning.” A second‬
‭hypothesis might be, “The classroom is warm because there is a power failure, and the air‬
‭conditioning doesn’t work.”‬
‭ nce a hypothesis is selected, then a prediction is stated. A‬‭prediction‬‭is similar to a‬
O
‭hypothesis, but it typically has the format “If . . . then . . . because .” For example, the‬
‭prediction for the first hypothesis might be, “‬‭If‬‭the student turns on the air conditioning,‬
‭then‬‭the classroom will no longer be too warm‬‭because‬‭the cooler air will lower the room‬
‭temperature.”‬

‭ igure 1.10‬ ‭The scientific method consists of a‬‭series of well-defined steps. If the‬
F
‭experimental data do not support a hypothesis, one can propose a new hypothesis to‬
‭evaluate. (credit:‬‭OpenStax‬‭) A link to a video explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 1: Introduction to Human Biology‬

‭Testing a Hypothesis‬
‭ valid hypothesis must be‬‭testable‬‭and‬‭falsifiable‬‭,‬‭meaning that experimental results can‬
A
‭show it is incorrect. Importantly, science does not claim to “prove” anything because‬
‭scientific understandings are always subject to modification with further information. This‬
‭step—openness to disproving ideas—distinguishes sciences from non-sciences. For‬
‭instance, the presence of the supernatural is neither testable nor falsifiable.‬
‭ o test a hypothesis, a researcher will design and conduct one or more‬‭experiments‬‭to‬
T
‭eliminate one or more hypotheses. Each experiment will have one or more variables and‬
‭o ne or more controls. A‬‭variable‬‭is any part of the‬‭experiment that can vary or change‬
‭during the investigation. The‬‭control group‬‭contains‬‭every feature of the experimental‬
‭group, except it is not given the manipulation that the researcher hypothesizes. Therefore,‬
‭if the experimental group's results differ from the control group, the difference must be‬
‭due to the hypothesized manipulation rather than some outside factor.‬
‭ o test the first hypothesis, the student would determine if the air conditioning is on. If the‬
T
‭air conditioning is turned on but does not work, there should be another reason, and the‬
‭student should reject this hypothesis. To test the second hypothesis, the student could‬
‭check if the lights in the classroom are functional. If so, there is no power failure; the‬
‭student should reject this hypothesis. The students should test each hypothesis by‬
‭conducting appropriate experiments.‬
‭ he scientific method may seem too rigid and structured. However, it is understood that,‬
T
‭although scientists often follow this sequence, there is flexibility. Sometimes, an‬
‭experiment leads to conclusions that favor a change in approach. Usually, an experiment‬
‭brings entirely new scientific questions to the puzzle. Many times, science does not‬
‭o perate linearly. Instead, scientists continually draw inferences and make generalizations,‬
‭finding patterns as their research proceeds. Scientific reasoning is more complex than the‬
‭scientific method alone suggests. Notice, too, that the scientific method can be applied to‬
‭solving problems that aren’t necessarily considered “scientific.”‬

‭Reporting Scientific Work‬

‭ hether scientific research is basic or applied, scientists must share their findings for‬
W
‭o ther researchers to expand and build upon their discoveries. Collaboration with other‬
‭scientists—when planning, conducting, and analyzing results—is vital for scientific‬
‭research. For this reason, important aspects of a scientist’s work are communicating with‬
‭peers and disseminating results to peers. Scientists can share results by presenting them‬
‭at a scientific meeting or conference, but this approach can reach only the select few who‬
‭are present. Instead, most scientists present their results in peer-reviewed manuscripts in‬
‭scientific journals.‬‭Peer-reviewed‬‭manuscripts are‬‭scientific papers that a scientist’s‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭23‬
‭Chapter 1: Introduction to Human Biology‬

c‭ olleagues or peers review. These colleagues are qualified individuals, often experts in the‬
‭same research area, who judge whether or not the scientist’s work is suitable for‬
‭publication. The peer review process helps ensure that the research in a scientific paper‬
‭o r grant proposal is original, significant, logical, and thorough. Grant proposals, which are‬
‭requests for research funding, are also subject to peer review. Scientists publish their‬
‭work so other scientists can reproduce their experiments under similar or different‬
‭conditions to expand on the findings. The experimental results must be consistent with the‬
‭findings of other scientists.‬
‭ scientific paper is very different from creative writing. Although creativity is required to‬
A
‭design experiments, there are fixed guidelines for presenting scientific results. First,‬
‭scientific writing must be brief, concise, and accurate. However, it must be detailed enough‬
‭for peers to reproduce the experiments.‬
‭ he scientific paper consists of several specific sections: introduction, materials and‬
T
‭methods, results, and discussion. This structure is sometimes called the “IMRaD” format.‬
‭In the beginning, there is usually an abstract, a concise summary of the entire paper, and‬
‭acknowledgment and reference sections at the end.‬
‭ he‬‭introduction‬‭starts with brief but broad background‬‭information about what is‬
T
‭known in the field. A good introduction also gives the rationale of the work. It justifies the‬
‭work carried out and briefly mentions the end of the paper, where the researcher will‬
‭present the hypothesis or research question driving the research. The introduction refers‬
‭to the published scientific work of others and, therefore, requires citations following the‬
‭journal's style. Using the work or ideas of others without proper citation is plagiarism.‬
‭ he‬‭materials and methods‬‭section includes a complete‬‭and accurate description of the‬
T
‭substances the researchers use and their methods and techniques to gather data. The‬
‭description should be thorough enough to allow another researcher to repeat the‬
‭experiment and obtain similar results, but it isn’t overly detailed. This section will also‬
‭include information on how the researchers made measurements and the types of‬
‭calculations and statistical analyses used to examine raw data. Although the materials and‬
‭methods section accurately describes the experiments, it does not discuss them.‬
S‭ ome journals require a‬‭results‬‭section followed by‬‭a‬‭discussion‬‭section, but it is more‬
‭common to combine both. If the journal does not allow combining both sections, the‬
‭results section simply narrates the findings without any further interpretation. In the‬
‭discussion section, the researchers will interpret the results, describe how variables may‬
‭be related, and attempt to explain the observations. Conducting an extensive literature‬
‭search is indispensable to put the results in the context of previously published scientific‬
‭research. Therefore, researchers include proper citations in this section as well.‬
‭ inally, the‬‭conclusion‬‭section summarizes the importance of the experimental findings.‬
F
‭While the scientific paper almost certainly answers one or more of the researchers' stated‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭24‬
‭Chapter 1: Introduction to Human Biology‬

‭ uestions, any good research should lead to more questions. Therefore, a well-done‬
q
‭scientific paper allows the researchers and others to continue expanding their knowledge‬
‭o f the topic.‬

‭1.5 Career Connection - Technical Writer‬

I‭ f you like science and technical fields and like to write, you may want to explore technical‬
‭writing as a career path. Technical writers usually have a degree in English, journalism, or‬
‭communication. Often, they also have personal knowledge, college coursework, and a‬
‭degree in a specialized field, such as computer science, engineering, medicine, biology,‬
‭agriculture, and other technical fields, such as manufacturing, construction, welding, and‬
‭plumbing; however, companies will usually train technical writers on the subject needed‬
‭and the style in which the writers they employ need to write.‬

‭ lthough the work varies depending on the industry, organization, and specific position,‬
A
‭technical writers typically perform the following tasks:‬

‭●‬ C
‭ reate content: Technical writers create an array of documents, such as product‬
‭information, operating and assembly instructions, “how-to” and “owner’s” manuals,‬
‭technical documentation, business proposals (solicited and unsolicited), lists of‬
‭frequently asked questions (FAQs), grant proposals, and journal articles.‬

‭●‬ R
‭ esearch: Technical writers research to gather the necessary information to write‬
‭accurate, professional, and helpful content.‬

‭●‬ E
‭ dit‬‭:‬ ‭Technical writers edit and standardize content‬‭prepared by other writers in‬
‭their organization.‬

‭●‬ A
‭ dapt content for multiple platforms: Technical writers create paper-based and‬
‭digital content using text, graphics, images, sound, and video to be distributed‬
‭across different platforms, including an organization’s website and social media.‬

‭In addition, technical writers develop and use the following skills:‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭25‬
‭Chapter 1: Introduction to Human Biology‬

‭●‬ W
‭ riting: Technical writers spend extended time writing complicated information in‬
‭clear and concise language.‬

‭●‬ A
‭ udience awareness: Technical writers are highly aware of the audience for their‬
‭writing. They plan, organize, and distribute the content they create with their‬
‭readers, viewers, and users in mind.‬

‭●‬ C
‭ ommunication and collaboration: Technical writers work on teams and‬
‭collaborate with experts, coworkers, and clients.‬

‭●‬ P
‭ roblem-solving: Technical writers often need to figure out how something works‬
‭to write documents their audience can understand.‬

‭●‬ T
‭ ime management: Technical writers often work on multiple projects with tight‬
‭deadlines. Setting priorities to keep projects on track is a crucial skill.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭26‬
‭Chapter 2: Chemistry and Life‬

‭ igure 2.1‬ ‭Different images of sucrose (table sugar):‬‭(a) Magnified sucrose crystals (b) A‬
F
‭“ball and stick” model of sucrose; black balls are carbon atoms, red balls are oxygen‬
‭atoms, and white balls are hydrogen atoms; the lines connecting the balls represent‬
‭covalent bonds (c) a structural model of sucrose showing the element symbols connected‬
‭by lines representing covalent bonds. (credit: a.‬‭Sugar 2xmacro.jpg‬‭; Lauri Andler‬
‭(Phantom);‬‭CC BY-SA 3.0‬ ‭b.‬‭Sucrose molecule 3d model.png‬‭;‬‭Michael Strö ck‬‭;‬‭CC BY-SA 3.0‬
‭c.‬ ‭Sucrose structure formula inkscape.svg‬‭; IMDJack;‬‭CC BY-SA 3.0‬‭)‬

‭2.1 Introduction‬
I‭ n the hierarchy of biological organization, introduced in Chapter 1, the most basic levels‬
‭directly involve chemistry (i.e., subatomic particles, atoms, molecules, and‬
‭macromolecules). Atoms of different elements combine in various combinations to‬
‭comprise all matter, including living things. Some of the most abundant elements in living‬
‭o rganisms include carbon (C), hydrogen (H), nitrogen (N), oxygen (O), sulfur (S), and‬
‭phosphorus (P). Atoms of these elements combine, via chemical bonding, to form the‬
‭biologically essential‬‭macromolecules‬‭, such as nucleic‬‭acids, proteins, carbohydrates, and‬
‭lipids, which are the fundamental components of all living matter. To learn about and‬
‭appreciate the complexity of human biology, one needs to have a basic understanding of‬
‭introductory chemistry that ultimately creates the other levels of biological organization‬
‭(e.g., organelles, cells, tissues, organs, organ systems, and organisms).‬
‭ ll biological processes follow the laws of physics and chemistry. Therefore, to‬
A
‭understand how biological systems work, it is essential to understand the underlying‬
‭physics and chemistry. For example, blood flow within the circulatory system follows the‬
‭laws of physics regulating fluid flow. The breakdown of the large, complex molecules of‬
‭food into smaller molecules—and the conversion of these to release energy in the form of‬
‭adenosine triphosphate (ATP)—is a series of chemical reactions that follow chemical laws.‬

‭27‬
‭Chapter 2: Chemistry‬

‭2.2 Elements and Atoms: The Building Blocks of Matter‬

‭ ll matter in the natural world is composed of one or more of the 92 fundamental‬
A
‭substances called elements. An‬‭element‬‭is a pure substance‬‭(i.e., atoms of only one type)‬
‭distinguished from all other matter because it cannot be created or broken down by‬
‭o rdinary chemical means. While your body can assemble many chemical compounds‬
‭needed for life from their constituent elements, it cannot make elements. They must come‬
‭from the environment. A familiar example of an element that you must take in is calcium‬
‭(Ca). Calcium is essential to the human body; it is absorbed and used for several‬
‭processes, including strengthening bones. When you consume dairy products, your‬
‭digestive system breaks down the food into components small enough to cross into the‬
‭bloodstream. Among these is calcium, which cannot be broken down further because it is‬
‭an element. Therefore, the elemental calcium in cheese is the same as the calcium that‬
‭forms your bones.‬
‭ igure 2.2 shows the elements in the human body and their designated‬‭c hemical‬
F
‭symbols‬‭, beginning with the most abundant: oxygen‬‭(O), carbon (C), hydrogen (H), and‬
‭nitrogen (N). Each element’s name can be replaced by a one—or two-letter chemical‬
‭symbol; you will become familiar with some of these during this course. All the elements‬
‭in your body are derived from the foods you eat and the air you breathe.‬
I‭ n nature, elements rarely occur alone. Instead, they combine to form compounds. A‬
‭compound‬‭is a substance composed of atoms of two or‬‭more elements joined by‬
‭chemical bonds. For example, the compound glucose (C‬‭6‭H ‬ ‬‭12‬‭O‭6
‬ ‬‭) is a vital body fuel that can‬
‭be used to create ATP. It is always composed of the same three elements: carbon,‬
‭hydrogen, and oxygen. Moreover, the elements that make up any given compound always‬
‭o ccur in identical relative amounts. In glucose, there are always six carbon and six oxygen‬
‭units for every twelve hydrogen units. But what, exactly, are these “units” of elements? We‬
‭need to continue our chemistry discussion and learn about atoms to answer this.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭28‬
‭Chapter 2: Chemistry‬

‭ igure 2.2‬ ‭The main elements composing the human‬‭body are shown from most to least‬
F
‭abundant. (credit:‬‭OpenStax‬‭)‬

‭Atoms and Subatomic Particles‬

‭ n‬‭atom‬‭is the smallest quantity of an element that‬‭retains the unique properties of that‬
A
‭element. In other words, an atom of hydrogen is a unit of hydrogen—the smallest amount‬
‭o f hydrogen that can exist. As you might guess, atoms are almost unfathomably tiny. The‬
‭period at the end of this sentence is millions of atoms wide. For additional information‬
‭about atoms, click the link below or scan the QR code to watch the TED-Ed video by Jon‬
‭Bergmann titled “Just How Small is an Atom?”.‬

Just How Small is an Atom?

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭29‬
‭Chapter 2: Chemistry‬

‭ toms are made up of even smaller‬‭subatomic particles‬‭, three types of which are‬
A
‭essential for our discussion:‬‭protons‬‭,‬‭neutrons‬‭, and‬‭electrons‬‭. Protons and neutrons‬
‭have approximately the same mass, about 1.67 × 10‬‭-24‬ ‭grams (i.e., 0.167 followed by 21‬
‭more zeros). Scientists arbitrarily define this amount of mass as one‬‭atomic mass unit‬
‭(amu)‬‭, as‬‭Figure 2.3‬‭shows. Although similar in mass, protons and neutrons differ in‬
‭their electric charge. A proton is positively charged, whereas a neutron is uncharged.‬
‭Therefore, the number of neutrons in an atom contributes significantly to its mass but not‬
‭its charge. For additional information about the discovery of atoms, click the link below or‬
‭scan the QR code to watch the TED-Ed video by Theresa Doud titled “The 2,400-Year‬
‭Search for the Atom”.‬

The 2,400-year search for the atom - Theresa Doud

‭ lectrons are much smaller in mass than protons, weighing only 9.11 × 10‬‭-28‬ ‭grams, or‬
E
‭about 1/1800 of an atomic mass unit. Hence, they do not contribute much to an element’s‬
‭overall atomic mass. Therefore, when considering atomic mass, it is customary to ignore‬
‭the mass of any electrons and calculate the atom’s mass based on the number of protons‬
‭and neutrons alone.‬
‭ lthough not significant contributors to an atom’s mass, electrons contribute substantially‬
A
‭to the atom’s charge, as each electron has a negative charge equal to the proton's positive‬
‭charge. In uncharged, neutral atoms, the number of electrons orbiting (i.e., traveling‬
‭around) the nucleus is the same as the number of protons inside the nucleus. These‬
‭atoms' positive and negative charges cancel each other out, leading to an atom with no net‬
‭charge (i.e., electrically neutral).‬

‭ elative Mass‬
R
‭ harge‬
C ‭(amu)‬ ‭ ocation‬
L
‭Proton‬ ‭+1‬ ‭1‬ ‭nucleus‬
‭Neutron‬ ‭0‬ ‭1‬ ‭nucleus‬
‭Electron‬ ‭–1‬ ‭0‬ ‭o rbitals‬

‭ igure 2.3‬ ‭The relative electrical charge, atomic‬‭mass, and location of subatomic particles‬
F
‭in an atom. Relative mass is measured in atomic mass units (i.e., amu). Orbitals are‬
‭electron pathways that surround an atom’s nucleus.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭30‬
‭Chapter 2: Chemistry‬

‭ igure 2.4‬‭shows two models that can help you visualize the structure of an atom—in this‬
F
‭case, helium (He). In the planetary model, helium’s two electrons are shown circling the‬
‭nucleus in a fixed orbit depicted as a ring. Although this model helps visualize the atomic‬
‭structure, electrons do not travel in fixed orbits. Instead, they whiz around the nucleus‬
‭erratically in a so-called electron cloud.‬
‭ n atom’s protons and electrons carry electrical charges. Protons, with their positive‬
A
‭charge, are designated p‬‭+‭.‬ Electrons, which have a negative charge, are designated e‬‭–‬‭. As‬
‭neutrons have no charge, they are designated as n. Just as a magnet sticks to a steel‬
‭refrigerator because their opposite charges attract, the positively charged protons attract‬
‭the negatively charged electrons. This mutual attraction gives the atom some structural‬
‭stability. The attraction by the positively charged nucleus helps keep electrons from‬
‭straying far. As previously stated, the number of protons and electrons within a neutral‬
‭atom is equal; thus, the atom’s overall charge is balanced.‬

‭ igure 2.4‬‭(a) In the planetary model of atomic structure,‬‭electrons of helium are shown‬
F
‭in fixed orbits, depicted as rings, at a precise distance from the nucleus. (b) In the electron‬
‭cloud model of atomic structure, the electrons of carbon are shown in different locations.‬
‭(credit:‬‭OpenStax‬‭) A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭31‬
‭Chapter 2: Chemistry‬

‭Atomic Number and Mass Number‬

‭ n atom of carbon is unique to carbon, but a proton of carbon is not. One proton is the‬
A
‭same as another, whether found in an atom of carbon, sodium (Na), or iron (Fe). The‬
‭same is true for neutrons and electrons. So, what gives an element its distinctive‬
‭properties—what makes carbon different from sodium or iron? The answer is the unique‬
‭quantity of protons each contains. Carbon, by definition, is an element whose atoms‬
‭contain six protons. No other element has exactly six protons in its atoms. Moreover,‬‭all‬
‭atoms of carbon, whether found in your liver or a lump of coal, contain six protons. Thus,‬
‭the‬‭atomic number‬‭, which is the number of protons in the atom's nucleus, identifies the‬
‭element. Because an atom usually has the same number of electrons as protons, the‬
‭atomic number also determines the usual number of electrons.‬
I‭ n their most common form, many elements contain the same number of neutrons as‬
‭protons. For example, the most common form of carbon has six neutrons and six protons‬
‭for 12 subatomic particles in its nucleus. An element’s‬‭mass number‬‭is the sum of the‬
‭number of protons and neutrons in its nucleus. So, the most common form of carbon’s‬
‭mass number is 12. Remember, electrons have so little mass that they do not appreciably‬
‭contribute to the mass of an atom. Carbon is a relatively light element. Uranium (U), in‬
‭contrast, has a mass number of 238 and is referred to as a heavy metal. Its atomic number‬
‭is 92 (an atom has 92 protons and 92 electrons), but the atom contains 146 neutrons; a‬
‭uranium atom has the most mass of all the naturally occurring elements.‬
‭ he‬‭periodic table‬‭o f the elements, shown in‬‭Figure‬‭2.5‬‭, identifies the 92 elements found‬
T
‭in nature and several larger, unstable elements discovered experimentally. Devised by‬
‭Russian chemist Dmitri Mendeleev (1834–1907) in 1869, the table groups elements that,‬
‭although unique, share certain chemical properties with other elements. For additional‬
‭information, click the link below or scan the QR code to watch the TED-Ed video by Eric‬
‭Rosado titled “Solving the Puzzle of the Periodic Table.”‬

Solving the puzzle of the periodic table - Eric Rosado

‭ he elements are arranged in order of their atomic number, with hydrogen and helium at‬
T
‭the top of the table and the more massive elements below. The periodic table is a helpful‬
‭device because it identifies each element's chemical symbol, atomic number, and mass‬
‭number while organizing elements according to their propensity to react with other‬
‭elements.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭32‬
‭Chapter 2: Chemistry‬

‭ igure 2.5‬ ‭The periodic table shows each element's‬‭atomic mass and atomic number. The‬
F
‭atomic number appears above the symbol for the element, and the approximate atomic‬
‭mass appears below it. (credit:‬‭OpenStax‬‭) A link‬‭to a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭33‬
‭Chapter 2: Chemistry‬

‭ earn more about any element by clicking the link below or scanning the QR code to‬
L
‭watch the TED-Ed resource “Periodic Videos: A Lesson About Every Single Element on the‬
‭Periodic Table.”‬

‭TED-Ed and Periodic Videos‬

‭Isotopes‬
‭ lthough each element has a unique number of protons, it can exist as different isotopes.‬
A
‭An‬‭isotope‬‭is one of the other forms of an element,‬‭distinguished from one another by‬
‭different numbers of neutrons. The standard isotope of carbon is‬‭12‬‭C, commonly called‬
‭carbon twelve.‬‭12‬‭C has six protons and six neutrons‬‭for a mass number of twelve.‬
‭All carbon isotopes have the same number of protons; thus,‬‭13‬‭C has seven neutrons, and‬
‭14‬
‭C has eight neutrons. The different isotopes of‬‭an element can also be indicated with the‬
‭mass number hyphenated (for example, C-12 instead of‬‭12‬‭C). Hydrogen has three common‬
‭isotopes, which are shown in‬‭Figure 2.6‬‭.‬

‭ igure 2.6.‬ ‭This figure shows the isotopes of hydrogen.‬‭Protium, designated‬‭1‬‭H, has one‬
F
‭proton and no neutrons.‬ ‭Deuterium, designated‬‭2‭H ‬
,‬‭has one proton and one neutron.‬
‭Tritium, designated‬‭3‭H
‬
, has one proton and two neutrons.‬ ‭(credit:‬ ‭OpenStax‬‭)‬

‭ n isotope containing more than the usual number of neutrons is called a heavy isotope;‬
A
‭an example is‬‭14‬‭C (carbon-14). Heavy isotopes tend‬‭to be unstable, and unstable isotopes‬
‭are radioactive. A radioactive isotope has a nucleus that readily decays, giving off‬
‭subatomic particles and electromagnetic energy. Different radioactive isotopes (or‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭34‬
‭Chapter 2: Chemistry‬

r‭ adioisotopes) differ in their‬‭half-life‬‭, the time it takes for half of any size sample of an‬
‭isotope to decay. For example, the half-life of tritium—a radioisotope of hydrogen—is‬
‭about 12 years, indicating it takes 12 years for half of the tritium nuclei in a sample to‬
‭decay. Excessive exposure to radioactive isotopes can damage human cells and even cause‬
‭cancer and congenital disabilities, but when exposure is controlled, some radioactive‬
‭isotopes can be helpful in medicine. For additional information, click the link below or‬
‭scan the QR code to watch the TED-Ed video by Pedro Brugarolas titled “Using radioactive‬
‭drugs to see inside your body”:‬

Using radioactive drugs to see inside your body - Pedro Brugar…

‭The Behavior of Electrons‬

I‭ n the human body, atoms do not exist as independent entities. Instead, they constantly‬
‭react with other atoms to form and break down more complex substances. To fully‬
‭understand anatomy and physiology, you must grasp how atoms participate in such‬
‭reactions. The key is understanding the behavior of electrons.‬
‭ lthough electrons do not follow rigid orbits a set distance from the atom’s nucleus, they‬
A
‭stay within defined regions of space called electron shells. An‬‭electron shell‬‭is a layer of‬
‭electrons that encircle the nucleus at a distinct energy level.‬
‭ he atoms of the elements found in the human body have from one to five electron shells,‬
T
‭and all electron shells hold eight electrons except the first shell, which can only contain‬
‭two. This configuration of electron shells is the same for all atoms. The precise number of‬
‭shells depends on the number of electrons in the atom. Hydrogen and helium have just‬
‭o ne and two electrons, respectively. Looking at the periodic table of the elements, you will‬
‭notice that hydrogen and helium are placed alone on either side of the top row; they are‬
‭the only elements with just one electron shell (‬‭Figure‬‭2.7a‬‭). A second shell is necessary to‬
‭hold the electrons in all elements larger than hydrogen and helium.‬
‭ ithium (Li), whose atomic number is 3, has three electrons. Two fill the first electron‬
L
‭shell, and the third spills into a second shell. The second electron shell can accommodate‬
‭as many as eight electrons. With its six electrons, carbon fills its first shell with six‬
‭electrons and half-fills its second. Neon (Ne) fills its two electron shells with ten electrons.‬
‭ gain, looking at the periodic table reveals that all elements in the second row, from‬
A
‭lithium to neon, have just two electron shells. Atoms with more than ten electrons require‬

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‭35‬
‭Chapter 2: Chemistry‬

‭ ore than two shells. These elements occupy the third and subsequent rows of the‬
m
‭periodic table.‬
‭ he factor that most strongly governs the tendency of an atom to participate in chemical‬
T
‭reactions is the number of electrons in its valence shell. A‬‭valence shell‬‭is an atom’s‬
‭o utermost electron shell. If the valence shell is complete, the atom is stable, meaning its‬
‭electrons are unlikely to be pulled away from the nucleus by the electrical charge of other‬
‭atoms. If the valence shell is not full, the atom is reactive, meaning it will tend to react with‬
‭o ther atoms in ways that make the valence shell full. Consider hydrogen, with its one‬
‭electron only half-filling its valence shell. This single electron is likely to be drawn into‬
‭relationships with the atoms of other elements so that hydrogen’s single valence shell is‬
‭stabilized.‬

‭ igure 2.7‬ ‭Electrons orbit the atomic nucleus at‬‭distinct energy levels called electron‬
F
‭shells. (a) With one electron, hydrogen only half-fills its electron shell. Helium also has a‬
‭single shell, but its two electrons fill it. (b) The electrons of carbon fill its first electron‬
‭shell but only half-fill its second. (c) Neon, an element that does not occur in the body, has‬
‭10 electrons, filling both electron shells. (credit:‬ ‭OpenStax‬‭) A link to a video explanation‬
‭o f this figure is available at‬‭Biology411.com‬‭.‬

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‭36‬
‭Chapter 2: Chemistry‬

‭ ll atoms (except hydrogen and helium with their single electron shells) are most stable‬
A
‭when eight electrons are in their valence shell. This principle is called the‬‭octet rule‬‭,‬
‭which states that an atom will give up, gain, or share electrons with another atom to end‬
‭up with eight electrons in its valence shell. For example, oxygen, with six electrons in its‬
‭valence shell, is likely to react with other atoms in a way that adds two electrons to‬
‭oxygen’s valence shell, bringing the number to eight. Covalent bonds are formed when‬
‭two hydrogen atoms share their single electron with oxygen, resulting in a water molecule,‬
‭H‭2‬ ‬‭O (‬‭Figure 2.8‬‭). Incidentally, the name “hydrogen”‬‭reflects its contribution to water‬
‭(hydro- = “water”; -gen = “maker”). Thus, hydrogen is the “water maker.”‬

‭ igure 2.8‬ ‭Two or more atoms may bond to form a‬‭molecule. When two hydrogens and‬
F
‭an oxygen share electrons via covalent bonds, it creates a water molecule. (credit:‬
‭OpenStax‬‭) A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭2.3 Chemical Bonds‬

‭ toms separated by a significant distance cannot link; instead, they must come close‬
A
‭enough for the electrons in their valence shells to interact. But do atoms ever actually‬
‭touch one another? Most physicists would say no because the negatively charged‬
‭electrons in their valence shells repel one another. No force within the human body—or‬
‭anywhere in the natural world—is strong enough to overcome this electrical repulsion. So‬
‭when you read about atoms linking together or colliding, remember that the atoms are‬
‭not merging physically.‬

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‭37‬
‭Chapter 2: Chemistry‬

I‭ nstead, atoms link by forming a‬‭c hemical bond‬‭, a weak or strong electrical attraction‬
‭that holds atoms in the same vicinity. The new grouping is typically more stable—less‬
‭likely to react again—than its component atoms were when they were separate. A more or‬
‭less stable collection of two or more atoms held together by chemical bonds is called a‬
‭molecule‬‭. The bonded atoms may be of the same element,‬‭as in the case of H‬‭2‭,‬ which is‬
‭called molecular hydrogen or hydrogen gas. A molecule comprising two or more atoms of‬
‭different elements is called a chemical compound. Thus, a unit of water, or H‬‭2‬‭O, is a‬
‭compound, as is a single molecule of the gas methane, or CH‬‭4‬‭.‬
‭ or a general introduction to atoms bonding to create molecules, click the link below or‬
F
‭scan the QR code to watch the TED-Ed video by Josh Kurz titled‬
‭“The science of macaroni salad - What’s in a mixture?”:‬

The science of macaroni salad: What's in a mixture? - Josh Kurz

‭ hree types of chemical bonds are important in human physiology because they hold‬
T
‭together substances used by the body for critical aspects of homeostasis, signaling, and‬
‭energy production, to name just a few processes. These are ionic bonds, covalent bonds,‬
‭and hydrogen bonds. For a general introduction to the first two bond types, click the link‬
‭below or scan the QR code to watch the TED-Ed video by George Zaidan and Charles‬
‭Morton titled “How Atoms Bond”:‬

How atoms bond - George Zaidan and Charles Morton

‭Ions and Ionic Bonds‬

‭ ecall that an atom typically has the same number of positively charged protons and‬
R
‭negatively charged electrons. As long as this situation remains, the atom is electrically‬
‭neutral. But when an atom participates in a chemical reaction that results in the donation‬
‭o r acceptance of one or more electrons, the atom will then become positively or negatively‬
‭charged. This situation frequently happens because most atoms “prefer” to have a full‬

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‭38‬
‭Chapter 2: Chemistry‬

‭ alence shell. This outcome can happen either by gaining electrons to fill a‬
v
‭more-than-half-full shell or by giving away electrons to empty a less-than-half-full shell,‬
‭leaving the next smaller electron shell as the new, full valence shell. An atom with an‬
‭electrical charge—positive or negative—is an‬‭ion‬‭.‬
‭ otassium (K), for instance, is an essential element in all body cells. Its atomic number is‬
P
‭19. It has just one electron in its valence shell. This characteristic makes potassium highly‬
‭likely to participate in chemical reactions in which it donates one electron. (It is easier for‬
‭potassium to donate one electron than to gain seven electrons.) The loss will cause the‬
‭positive charge of potassium’s protons to be more influential than the negative charge of‬
‭potassium’s electrons. In other words, the resulting potassium ion will have a positive‬
‭charge. A potassium ion is written K‬‭+‭,‬ indicating‬‭it has lost a single electron. A positively‬
‭charged ion is known as a‬‭c ation‬‭.‬
‭ ow consider fluorine (F), a component of bones and teeth. Its atomic number is nine,‬
N
‭and it has seven electrons in its valence shell. Thus, it is highly likely to bond with other‬
‭atoms so that fluorine accepts one electron (it is easier for fluorine to gain one electron‬
‭than to donate seven electrons). When it does, its electrons will outnumber its protons by‬
‭o ne, and it will have an overall negative charge. The ionized form of fluorine is called‬
‭fluoride and is written as F‬‭–‭.‬ A negatively charged‬‭ion is known as an‬‭anion‬‭.‬
‭ toms with more than one electron to donate or accept will have stronger positive or‬
A
‭negative charges. A cation that has donated two electrons has a net charge of +2. Using‬
‭magnesium (Mg) as an example, this can be written Mg‬‭++‬ ‭o r Mg‬‭2+‬‭. An anion that has‬
‭accepted two electrons has a net charge of –2. For example, the ionic form of selenium‬
‭(Se) is typically written Se‬‭2–‬‭.‬
‭ he opposite charges of cations and anions exert a moderately strong mutual attraction‬
T
‭that keeps the atoms nearby, forming an ionic bond. An‬‭ionic bond‬‭is an ongoing, close‬
‭association between ions of opposite charge. The table salt you sprinkle on your food,‬
‭sodium chloride, owes its existence to ionic bonding.‬‭Figure 2.9‬‭shows sodium commonly‬
‭donates an electron to chlorine, becoming the cation Na‬‭+‭.‬ When chlorine accepts the‬
‭electron, it becomes the chloride anion, Cl‬‭–‭.‬ With‬‭their opposing charges, these two ions‬
‭strongly attract each other.‬
‭ ater is essential to life because it can break the ionic bonds in salts to free the ions. In‬
W
‭fact, in biological fluids, most individual atoms exist as ions. These dissolved ions produce‬
‭electrical charges within the body. The behavior of these ions produces the tracings of‬
‭heart and brain function observed as waves on an electrocardiogram (EKG or ECG) or an‬
‭electroencephalogram (EEG). The electrical activity that derives from the interactions of‬
‭the charged ions is why they are also called electrolytes.‬

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‭39‬
‭Chapter 2: Chemistry‬

‭(d)‬

‭ igure 2.9‬ ‭Ionic bonding: (a) Sodium readily donates‬‭the solitary electron in its valence‬
F
‭shell to chlorine, which needs only one electron to have a full valence shell. (b) The‬
‭o pposite electrical charges of the resulting sodium cation and chloride anion result in the‬
‭formation of a bond of attraction called an ionic bond. (c) The attraction of many sodium‬
‭and chloride ions results in large groupings called crystals, shown in (d). (credit: (a)-(c:)‬
‭OpenStax‬‭; (d)‬‭Michel32nl‬‭;‬‭CC BY-SA 3.0‬‭). A link to‬‭a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭Covalent Bonds‬
‭ nlike ionic bonds formed by the attraction between a cation’s positive charge and an‬
U
‭anion’s negative charge, molecules formed by a‬‭covalent‬‭bond‬‭share electrons in a‬
‭mutually stabilizing relationship. Like next-door neighbors whose kids hang out first at‬

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‭40‬
‭Chapter 2: Chemistry‬

‭ ne home and then at the other, the atoms do not lose or gain electrons permanently.‬
o
‭Instead, the electrons move back and forth between the elements. Because of the close‬
‭sharing of pairs of electrons (one electron from each of two atoms), covalent bonds are‬
‭stronger than ionic bonds. In the following sections, we will distinguish between two‬
‭types of covalent bonds, nonpolar covalent and polar covalent.‬

‭Nonpolar Covalent Bonds‬

‭ onpolar covalent bonds‬‭result from equal sharing‬‭o f electrons, and there are‬
N
‭commonly one or two pairs of shared electrons (‬‭Figure‬‭2.10‬‭). The vital concept to take‬
‭from this is that in covalent bonds, electrons in the outermost valence shell are shared to‬
‭fill the valence shells of both atoms, ultimately stabilizing both of the atoms involved. In a‬
‭single covalent bond‬‭, a pair of electrons is shared;‬‭in a‬‭double covalent bond‬‭, two pairs‬
‭o f electrons are shared.‬
‭ ou can see that the covalent bonds shown in‬‭Figure‬‭2.10‬‭are balanced. The sharing of‬
Y
‭the negative electrons is relatively equal, as is the electrical pull of the positive protons in‬
‭the atoms' nucleus. This is why covalently bonded molecules that are electrically balanced‬
‭in this way are described as‬‭nonpolar‬‭; that is, no‬‭molecule region is either more positive‬
‭o r negative than any other.‬

‭Polar Covalent Bonds‬

‭ roups of legislators with opposite views on a particular issue are often described as‬
G
‭“polarized” by news writers. In chemistry, a‬‭polar‬‭molecule‬‭is a molecule that contains‬
‭regions that have opposite electrical charges. Polar molecules occur when atoms share‬
‭electrons unequally in‬‭polar covalent bonds‬‭.‬
‭ ater is the most familiar example of a polar molecule (‬‭Figure 2.11‬‭). The molecule has‬
W
‭three parts: one atom of oxygen, the nucleus of which contains eight protons, and two‬
‭hydrogen atoms, whose nuclei each contain only one proton. Because every proton exerts‬
‭an identical positive charge, a nucleus with eight protons exerts a charge eight times‬
‭greater than a nucleus with one proton. This means that the negatively charged electrons‬
‭in the water molecule are more strongly attracted to the oxygen nucleus than the‬
‭hydrogen nuclei. Each hydrogen atom’s single negative electron migrates toward the‬
‭oxygen atom, making the oxygen end of their bond slightly more negative than the‬
‭hydrogen end of their bond.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭41‬
‭Chapter 2: Chemistry‬

‭ igure 2.10‬ ‭Examples of nonpolar covalent bonding‬‭(single and double). (credit:‬

F
‭OpenStax‬‭) A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ hat is valid for the bonds is true for the water molecule as a whole; that is, the oxygen‬
W
‭region has a slightly negative charge, and the regions of the hydrogen atoms have a‬
‭slightly positive charge. As shown in‬‭Figure 2.11‬‭,‬‭regions of weak polarity are indicated‬
‭with the Greek letter delta (δ) and a plus (+) or minus (–) sign.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭42‬
‭Chapter 2: Chemistry‬

‭ igure 2.11‬ ‭Polar covalent bonds in a water molecule.‬ ‭(credit:‬‭OpenStax‬‭) A link to a‬

F
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ hese partially charged, or polar, regions are highly likely to interact with charged parts of‬
T
‭o ther polar molecules. For human physiology, the resulting bond is the most important‬
‭formed by water—the‬‭hydrogen bond‬‭(‬‭Figure 2.12‬‭).‬
I‭ ndividual hydrogen bonds are weak and easily broken; however, they occur in vast‬
‭amounts in water and organic polymers, creating a significant force in combination.‬
‭Hydrogen bonds are also responsible for protein folding and zipping the DNA double‬
‭helix together.‬

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‭43‬
‭Chapter 2: Chemistry‬

‭ igure 2.12‬ ‭Notice that the hydrogen bonds occur‬‭between the weakly positive charge‬
F
‭o n the hydrogen atoms and the weakly negative charge on the oxygen atoms. Hydrogen‬
‭bonds are relatively weak, so they are indicated with a dotted (rather than a solid) line.‬
‭(credit:‬‭OpenStax‬‭) A link to a video explanation of‬‭this figure is available at‬
‭Biology411.com‬‭.‬

‭Water’s Polarity‬
‭ ater also attracts or is attracted to other polar molecules and ions. We call a polar‬
W
‭substance that interacts readily with or dissolves in water‬‭hydrophilic‬‭(hydro- = “water”;‬
‭-philic = “loving”). In contrast, nonpolar molecules such as oils and fats do not interact well‬
‭with water, as‬‭Figure 2.13‬‭shows. An excellent example‬‭o f this phenomenon is vinegar and‬
‭o il salad dressing (an acidic water solution). The fat droplets will not stay dissolved, as‬
‭shown below. This result is because the fats are nonpolar. We call such nonpolar‬
‭compounds‬‭hydrophobic‬‭(hydro- = “water”; -phobic =‬‭“fearing”).‬

‭ igure 2.13‬ ‭This macro image of oil and water shows‬‭that oil does not dissolve in water‬
F
‭but forms droplets because it is a nonpolar compound. (credit: Gautam Dogra;‬‭OpenStax‬‭).‬

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‭44‬
‭Chapter 2: Chemistry‬

‭ or additional information, click the link below or scan the QR code to watch the TED-Ed‬
F
‭video by Christina Kleinberg titled “How polarity makes water behave strangely.”‬

How polarity makes water behave strangely - Christina Kleinberg

‭2.4 pH, pH Scale, Acidic, and Basic/Alkaline‬

‭ he‬‭pH‬‭o f a solution indicates its acidity or basicity.‬ ‭You may have used litmus or pH‬
T
‭paper, filter paper treated with a natural water-soluble dye for use as a pH indicator. It‬
‭tests how much acid (acidity) or base (basicity) exists in a solution. You might have even‬
‭used some to test whether the water in a swimming pool is treated correctly. In both cases,‬
‭the pH test measures the concentration of‬‭hydrogen‬‭ions‬‭(i.e., H‬‭+‭)‬ in a given solution.‬
‭ he‬‭pH scale‬‭ranges from 0 to 14 (‬‭Figure 2.14‬‭). A solution with a pH of 7 is considered‬
T
‭neutral‬‭—neither acidic nor basic. Pure water has a‬‭pH of 7. Any pH value below 7.0‬
‭(ranging from 0.0 to 6.9) is classified as‬‭acidic‬‭.‬‭The lower the number below 7, the more‬
‭acidic the solution, or the greater the concentration of H‬‭+‬‭. The concentration of hydrogen‬
‭ions at each pH value is 10 times different than the next pH. For example, a pH 4 solution‬
‭has 10 times more hydrogen ions than a pH 5 solution. A pH 3 solution has 100 times‬
‭more hydrogen ions than a pH 5 solution (i.e., 10 x 10 = 100).‬
‭ ny pH value above 7.0 (7.1 to 14.0) is‬‭basic‬‭o r‬‭alkaline‬‭.‬‭The pH value is increasing‬
A
‭because the concentration of hydrogen ions is decreasing. The pH inside cells (6.8) and‬
‭the pH in the blood (7.4) are both very close to neutral (i.e., pH 7). However, the‬
‭environment in the stomach is very acidic, with a pH of 1 to 2. How do stomach cells‬
‭survive in such an acidic environment? How do they homeostatically maintain the‬
‭near-neutral pH inside them? The answer is that they cannot do it and are constantly‬
‭dying. The stomach continually produces new cells to replace dead ones, which stomach‬
‭acids digest. Scientists estimate that the human body completely replaces the stomach‬
‭lining every seven to ten days.‬

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‭45‬
‭Chapter 2: Chemistry‬

‭ igure 2.14‬ ‭The pH scale measures a solution's hydrogen‬‭ion (H‬‭+‭)‬ concentration. Values‬
F
‭below 7 are classified as acidic, and values above 7 are classified as basic or alkaline. Each‬
‭1 unit change of pH corresponds to a ten-times change in the concentration of H‬‭+‭.‬ For‬
‭example, a pH 4 solution has a ten times greater hydrogen ion concentration than a pH 5‬
‭solution. However, a pH 6 solution has ten times lower ion concentration than the pH 5‬
‭solution and one hundred times lower concentration than the pH 4 solution. (credit:‬
‭OpenStax‬‭) A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

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‭46‬
‭Chapter 2: Chemistry‬

‭2.5 Organic Compounds Essential to Human Functioning‬

‭ rganic compounds‬‭typically consist of groups of carbon‬‭atoms covalently bonded to‬
O
‭hydrogen, usually oxygen, and often other elements. The four types most important to‬
‭human structure and function are carbohydrates, lipids, proteins, and nucleic acids. Before‬
‭exploring these compounds, you need to understand more about the chemistry of carbon.‬

‭The Chemistry of Carbon‬

‭ hat makes organic compounds so common is the chemistry of their carbon core. Recall‬
W
‭that carbon atoms have four electrons in their valence shell and that the octet rule dictates‬
‭that atoms tend to react in such a way as to complete their valence shell with eight‬
‭electrons. Carbon atoms do not complete their valence shells by donating or accepting‬
‭four electrons. Instead, they readily share electrons via covalent bonds.‬
‭ any combinations are possible to fill carbon’s four “vacancies.” Carbon may share‬
M
‭electrons with oxygen, nitrogen, or other atoms in a particular region of an organic‬
‭compound. Moreover, the atoms to which carbon atoms bond may also be part of a‬
‭functional group. A‬‭functional group‬‭is a group of‬‭atoms linked by strong covalent‬
‭bonds that tend to function in chemical reactions as one unit. Four biologically important‬
‭examples are the‬‭hydroxyl‬‭,‬‭c arboxyl‬‭,‬‭amino‬‭, and‬‭phosphate‬‭groups (‬‭Figure 2.15‬‭).‬

‭Functional group‬ ‭Structural formula‬ ‭Importance‬

‭Hydroxyl‬ ‭—O—H‬ ‭ ydroxyl groups are polar. They‬
H
‭are components of all four types‬
‭of organic compounds discussed‬
‭in this chapter. They are involved‬
‭in dehydration synthesis and‬
‭hydrolysis reactions.‬
‭Carboxyl‬ ‭ arboxyl groups are found in fatty‬
C
‭acids, amino acids, and many‬
‭other acids.‬

‭Amino‬ ‭—N—H‬‭2‬ ‭ mino groups are found within‬

A
‭amino acids, the building blocks‬
‭of proteins.‬
‭Phosphate‬ ‭—P—O‬‭4‭2‬ –‬ ‭ hosphate groups are found‬
P
‭within phospholipids and‬
‭nucleotides.‬

‭Figure 2.15‬ ‭Functional groups that are important‬‭in human biology‬

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‭47‬
‭Chapter 2: Chemistry‬

‭ arbon’s affinity for covalent bonding means many distinct and relatively stable organic‬
C
‭molecules readily form larger, more complex molecules. Any large molecule is referred to‬
‭as a‬‭macromolecule‬‭(macro- = “large”), and the four‬‭categories in this section‬
‭(carbohydrates, lipids, proteins, and nucleic acids) all fit this description. However, some‬
‭macromolecules are made up of several “copies” of single units called‬‭monomers‬‭(mono-‬
‭= “one”; -mer = “part”). Like beads in a long necklace, these monomers link by covalent‬
‭bonds to form long‬‭polymers‬‭(poly- = “many”). There‬‭are many examples of monomers‬
‭and polymers among the organic compounds.‬
‭ onomers form polymers by engaging in‬‭dehydration‬‭synthesis‬‭o r‬‭condensation‬
M
‭reactions‬‭(‬‭Figure 2.16‬‭). As noted earlier, this reaction‬‭results in the release of a water‬
‭molecule. Each monomer contributes, with one giving up a hydrogen atom and the other‬
‭giving up a hydroxyl group.‬‭Hydrolysis‬‭divides polymers‬‭into monomers (-lysis =‬
‭“rupture”). The bonds between their monomers are broken via the donation of a water‬
‭molecule, which contributes a hydrogen atom to one monomer and a hydroxyl group to‬
‭the other.‬

‭ igure 2.16‬ ‭An‬‭illustration of dehydration synthesis‬‭and hydrolysis reactions.‬

F
‭Monomers, the basic units for building larger molecules, form polymers (two or more‬
‭chemically bonded monomers). (a) In dehydration synthesis, two monomers are‬
‭covalently bonded in a reaction in which one gives up a hydroxyl group and the other a‬
‭hydrogen atom. A molecule of water is released as a byproduct during dehydration‬
‭reactions. (b) In hydrolysis, the covalent bond between two monomers is split by adding a‬
‭hydrogen atom to one and a hydroxyl group to the other, which requires the contribution‬
‭o f one molecule of water. (credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭48‬
‭Chapter 2: Chemistry‬

‭ or additional information about monomers and polymers discussed in the remainder of‬
F
‭the chapter, click the link below or scan the QR code to watch the TED-Ed video by Jan‬
‭Mattingly titled “From DNA to Silly Putty, the diverse world of polymers”:‬

From DNA to Silly Putty, the diverse world of polymers - Jan Matti…

‭Carbohydrates‬
‭ ‬‭c arbohydrate‬‭is a molecule composed of carbon, hydrogen,‬‭and oxygen; in most‬
A
‭carbohydrates, hydrogen and oxygen are found in the same two-to-one relative‬
‭proportions they have in water. In fact, the chemical formula for a “generic” carbohydrate‬
‭molecule is (CH‬‭2‭O
‬ )‬‭n‬‭. Note this represents a 1:2:1‬‭ratio of carbon, hydrogen, and oxygen.‬

‭ arbohydrates are referred to as saccharides, a word meaning “sugars.” Three forms are‬
C
‭important in the body (‬‭Figure 2.17‬‭).‬‭Monosaccharides‬‭are the monomers of‬
‭carbohydrates.‬‭Disaccharides‬‭(di- = “two”) comprise‬‭two monomers joined via‬
‭dehydration synthesis reactions.‬‭Polysaccharide‬‭s are‬‭polymers and can consist of‬
‭thousands of monomers.‬

‭Figure 2.17‬ ‭Types of Carbohydrates (credit:‬ ‭OpenOregon.pressbooks.pub‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭49‬
‭Chapter 2: Chemistry‬

‭Monosaccharides‬
‭ monosaccharide is a monomer of carbohydrates. Five monosaccharides are important‬
A
‭in the body. Three of these are hexose sugars because they each contain six carbon atoms‬
‭( hex- means six). These are‬‭glucose‬‭,‬‭fructose‬‭, and‬‭galactose‬‭, as shown in‬‭Figure 2.18a‬‭.‬
‭The remaining monosaccharides are the two pentose sugars (i.e.,‬‭ribose‬‭and‬‭deoxyribose;‬
‭Figure 2.18b‬‭), each containing five carbon atoms (pent-‬‭means five).‬

‭Disaccharides‬
‭ disaccharide is a pair of monosaccharides. Disaccharides are formed via dehydration‬
A
‭synthesis, and three disaccharides (‬‭Figure 2.19‬‭) are‬‭important to humans. These are‬
‭sucrose‬‭, commonly referred to as table sugar;‬‭lactose‬‭,‬‭o r milk sugar; and‬‭maltose‬‭, or‬
‭malt sugar. As you can tell from their common names, you consume these in your diet;‬
‭however, your body cannot use them directly. Instead, in the digestive tract, they are split‬
‭into their component monosaccharides via hydrolysis to be absorbed and transported via‬
‭the circulatory system.‬

‭ igure 2.18‬ ‭Five important monosaccharides: (a)‬‭3 Hexoses and (b) 2 Pentoses. (credit:‬
F
‭OpenStax‬‭) A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭50‬
‭Chapter 2: Chemistry‬

‭ igure 2.19‬ ‭Two important disaccharides, sucrose,‬‭and maltose; lactose isn’t shown.‬
F
‭(credit:‬‭OpenStax‬‭)‬

‭Polysaccharides‬
‭ olysaccharides can contain a few to a thousand or more monosaccharides, and three of‬
P
‭them are important to the body:‬
‭•‬ ‭ tarches‬‭are polymers of glucose that plants produce‬‭during photosynthesis, the‬
S
‭process by which oxygen and carbon dioxide are chemically changed to glucose.‬

‭•‬ ‭ lycogen‬‭is also a polymer of glucose, but it is stored‬‭in the tissues of animals,‬
G
‭especially in the muscles and liver. It is not considered a dietary carbohydrate‬
‭because very little glycogen remains in animal tissues after slaughter; however, the‬
‭human body stores excess glucose as glycogen in the muscles and liver.‬

‭•‬ ‭ ellulose‬‭, a polysaccharide that is the primary component‬‭o f the cell wall of green‬
C
‭plants, is the component of plant food referred to as “fiber.” In humans,‬
‭cellulose/fiber is not digestible; however, dietary fiber has many health benefits. It‬
‭helps you feel full, so you eat less; it promotes a healthy digestive tract, and a diet‬
‭high in fiber is thought to reduce the risk of heart disease and possibly some forms of‬
‭cancer.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭51‬
‭Chapter 2: Chemistry‬

‭ ll three of these polysaccharides have the same monomer unit (glucose). What differs‬
A
‭between them is the monomer's arrangement, as shown in‬‭Figure 2.20‬‭.‬

‭ igure 2.20‬‭Three important polysaccharides for humans‬‭are starches, glycogen, and‬

F
‭cellulose/fiber. (credit:‬‭OpenStax‬‭)‬

‭ or additional information, click the link below or scan the QR code to watch the TED-Ed‬
F
‭video by Richard J. Wood titled “How do carbohydrates impact your health”:‬

How do carbohydrates impact your health? - Richard J. Wood

‭Lipids‬
‭ hen discussing human biology, there are three pertinent categories of lipids:‬
W
‭triglycerides (fat), phospholipids, and steroids/sterols.‬

‭Triglycerides‬
‭ ‬‭triglyceride‬‭is one of the most common dietary lipids,‬‭and the type found most‬
A
‭abundantly in body tissues. This compound, commonly called fat, is formed by joining two‬
‭types of molecules (glycerol and fatty acids;‬‭Figure‬‭2.21‬‭):‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭52‬
‭Chapter 2: Chemistry‬

•‭ ‬ ‭ ‬‭glycerol‬‭backbone at the core of triglycerides consists of three carbon atoms.‬

A
‭•‬ ‭Three‬‭fatty acids‬‭, long chains of hydrocarbons with‬‭a carboxyl group at one end,‬
‭extend from each of the carbons of the glycerol.‬

‭ igure 2.21‬ ‭Triglycerides are composed of glycerol‬‭attached to three fatty acids via‬
F
‭dehydration synthesis. Notice that glycerol gives up a hydrogen atom, and the carboxyl‬
‭groups on the fatty acids each give up a hydroxyl group. (credit:‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ riglycerides form via dehydration synthesis. Glycerol gives up hydrogen atoms from its‬
T
‭hydroxyl groups at each bond, and the carboxyl group on each fatty acid chain gives up a‬
‭hydroxyl group. A total of three water molecules are thereby released.‬
‭ atty acid chains with no double carbon bonds anywhere along their length and,‬
F
‭therefore, contain the maximum number of hydrogen atoms are called‬‭saturated fatty‬
‭acids‬‭. These straight, rigid chains pack tightly together‬‭and are solid or semi-solid at room‬
‭temperature (‬‭Figure 2.22a‬‭). Butter and lard are the‬‭fats found in a steak. In contrast, fatty‬
‭acids with one double carbon bond are kinked at that bond (‬‭Figure 2.22b‬‭). These‬
‭monounsaturated fatty acids‬‭cannot pack together tightly‬‭and are liquid at room‬
‭temperature.‬‭Polyunsaturated fatty acids‬‭contain two‬‭o r more double-carbon bonds‬
‭and are liquid at room temperature. Plant oils like olive oil typically contain mono- and‬
‭polyunsaturated fatty acids.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭53‬
‭Chapter 2: Chemistry‬

‭ igure 2.22‬ ‭The level of saturation of a fatty acid‬‭affects its shape. (a) Saturated fatty acid‬
F
‭chains are straight. (b) Unsaturated fatty acid chains are kinked. (credit:‬‭OpenStax‬‭)‬

‭ hereas a diet high in saturated fatty acids increases the risk of heart disease, a diet high‬
W
‭in unsaturated fatty acids is thought to reduce the risk. This observation is especially true‬
‭for the omega-3 unsaturated fatty acids in cold-water fish such as salmon. These fatty‬
‭acids have their first double carbon bond at the third hydrocarbon from the end of the‬
‭fatty acid opposite of the carboxyl group (referred to as the omega end of the molecule).‬
‭ inally, trans fatty acids found in some processed foods, including some stick and tub‬
F
‭kinds of margarine, are thought to be even more harmful to the heart and blood vessels‬
‭than saturated fatty acids.‬‭Trans fats‬‭are created‬‭from unsaturated fatty acids (such as‬
‭corn oil) when chemically treated to produce partially hydrogenated fats.‬
‭ s a group, triglycerides are a significant fuel source for the body. When resting or asleep,‬
A
‭most of the energy used to keep you alive is derived from triglycerides stored in your fat‬
‭(adipose) tissues. Triglycerides also fuel prolonged, slow physical activity such as‬
‭gardening or hiking and contribute a modest percentage of energy for vigorous physical‬
‭exercise. Dietary fat also assists the absorption and transport of the nonpolar fat-soluble‬
‭vitamins A, D, E, and K. Additionally, stored body fat protects and cushions the body’s‬
‭bones and internal organs and acts as insulation to retain body heat.‬
‭ or more information, click the link below or scan the QR code to watch the TED-Ed video‬
F
‭by George Zaidan titled “What is fat?”.‬

What is fat? - George Zaidan

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭54‬
‭Chapter 2: Chemistry‬

‭Phospholipids‬
‭ hospholipids‬‭are similar in structure to triglycerides.‬‭However, instead of having three‬
P
‭fatty acids, a phospholipid contains glycerol, two fatty acid chains, and a phosphate group‬
‭bound to the glycerol, which is attached to the molecule's polar “head” region. (‬‭Figure‬
‭2.23‬‭).‬

‭ igure 2.23‬ ‭An example of a phospholipid (credit:‬‭OpenStax‬‭). A link to a video‬

F
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ ecall that triglycerides are nonpolar and hydrophobic, which is also true for the fatty acid‬
R
‭portion of a phospholipid. However, the head of a phospholipid contains charges on the‬
‭phosphate groups and the nitrogen atom. These charges make the phospholipid head‬
‭hydrophilic. Therefore, phospholipids are said to have hydrophobic tails, containing the‬
‭neutral fatty acids, and hydrophilic heads, containing the charged phosphate groups and‬
‭the nitrogen atom (see the schematic phospholipid to the far right side in‬‭Figure 2.23‬‭)‬

‭Steroids‬
‭ ‬‭steroid‬‭compound (referred to as a sterol) has as‬‭its foundation a set of four‬
A
‭hydrocarbon rings bonded to various other atoms and molecules. Although both plants‬
‭and animals synthesize sterols, the type that makes the most significant contribution to‬
‭human structure and function is‬‭c holesterol‬‭, which‬‭is synthesized by the liver in humans‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭55‬
‭Chapter 2: Chemistry‬

‭and animals and is also present in most animal-based foods (‬‭Figure 2.24‬‭).‬

‭ igure 2.24‬ ‭The structure of cholesterol. (credit:‬‭Cholesterol (chemical structure).svg‬‭;‬

F
‭Wesalius‬‭; Public Domain)‬

‭ holesterol is also a building block of many hormones, such as estrogen, progesterone,‬

C
‭and testosterone. Finally, like phospholipids, cholesterol molecules are found in the cell‬
‭membrane, where their hydrophobic and hydrophilic regions help regulate the flow of‬
‭substances into and out of the cell.‬
‭ or more information, click the link or scan the QR code to watch the TED-Ed video by‬
F
‭Anees Bahji titled “Can steroids save your life?”:‬

Can steroids save your life? - Anees Bahji

‭Proteins‬
‭ ou might associate‬‭proteins‬‭with muscle tissue, but‬‭proteins are critical components of‬
Y
‭all tissues and organs. A protein is an organic molecule composed of‬‭amino acids‬‭linked‬
‭by‬‭peptide bonds‬‭created via dehydration synthesis‬‭between the amino and carboxyl‬
‭functional groups.‬
‭ arbohydrates and lipids are composed of hydrogen, carbon, and oxygen. However, all‬
C
‭proteins also contain nitrogen (N) in addition to these elements.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭56‬
‭Chapter 2: Chemistry‬

‭Microstructure of Proteins‬
‭ n amino acid is a molecule composed of an amino group, a carboxyl group, and a‬
A
‭variable side chain. Just 20 different amino acids contribute to nearly all thousands of‬
‭proteins vital in human structure and function. Body proteins contain a unique‬
‭combination of a few dozen to a few hundred of these 20 amino acid monomers. All 20‬
‭amino acids share a similar structure (‬‭Figure 2.25‬‭).‬‭All consist of a central carbon atom to‬
‭which the following are bonded:‬
•‭ ‬ a‭ hydrogen atom‬
‭•‬ ‭an alkaline (basic) amino functional group NH‬‭2‬ ‭(see‬‭Figure 2.15‬‭)‬
‭•‬ ‭an acidic carboxyl functional group COOH (see‬‭Figure‬‭2.15‬‭)‬
‭•‬ ‭an‬‭R group‬‭called the variable group, of which there‬‭are 20 different types, one for‬
‭each amino acid.‬
‭ otice that all amino acids contain both an acid (the carboxyl group) and a base (the‬
N
‭amino group) (amine = “nitrogen-containing”). For this reason, they make excellent‬
‭buffers, helping the body regulate acid–base balance.‬
‭ hat distinguishes the 20 amino acids from one another is their variable group, which is‬
W
‭referred to as a side chain or an‬‭R group. This group‬‭can vary in size and be polar or‬
‭nonpolar, giving each amino acid its unique characteristics. For example, the side chains of‬
‭two amino acids—cysteine and methionine—contain sulfur. Sulfur does not readily‬
‭participate in hydrogen bonds, whereas all other amino acids do. This variation influences‬
‭the way that proteins containing cysteine and methionine are assembled.‬

‭ igure 2.25‬ ‭The general chemical structure of an‬‭amino acid. (credit:‬‭OpenStax‬‭) A link‬
F
‭to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭57‬
‭Chapter 2: Chemistry‬

‭ s previously stated, amino acids join via dehydration synthesis to form protein polymers‬
A
‭(‬‭Figure 2.26‬‭). The unique bond holding amino acids‬‭together is called a‬‭peptide bond‬‭,‬
‭which is a covalent bond between two amino acids that forms by dehydration synthesis. A‬
‭peptide is a very short chain of amino acids. Strands containing fewer than 100 amino‬
‭acids are generally called polypeptides rather than proteins.‬
‭ he body can synthesize most of the amino acids from components of other molecules;‬
T
‭however, nine cannot be synthesized and must be consumed in the diet. These are known‬
‭as the‬‭essential amino acids‬‭.‬
‭ ree amino acids available for protein construction reside in the amino acid pool within‬
F
‭cells. Structures within cells use these amino acids when assembling proteins. If a‬
‭particular essential amino acid is not available in sufficient quantities in the amino acid‬
‭pool, the synthesis of proteins containing it can slow or even stop.‬

‭ igure 2.26‬ ‭Different amino acids join together‬‭to form peptides, polypeptides, or‬
F
‭proteins via dehydration synthesis. The bonds between the amino acids are peptide‬
‭bonds. (credit:‬‭OpenStax‬‭) A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭The Shape of Proteins‬

J‭ ust as a fork cannot be used to eat soup and a spoon cannot be used to spear meat, a‬
‭protein’s shape is essential to its function. A protein’s shape is determined, most‬
‭fundamentally, by the sequence of amino acids of which it is made (‬‭Figure 2.27a‬‭). This‬
‭sequence is called the‬‭primary structure‬‭o f a protein.‬

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‭58‬
‭Chapter 2: Chemistry‬

‭ lthough some polypeptides exist as linear chains, most are twisted or folded into more‬
A
‭complex‬‭secondary structures‬‭that form when bonding‬‭o ccurs between amino acids with‬
‭different properties at different regions of the polypeptide. The most common secondary‬
‭structure is a spiral called an alpha-helix. If you were to take a length of string and simply‬
‭twist it into a spiral, it would not hold the shape. Similarly, a strand of amino acids could‬
‭not maintain a stable spiral shape without the help of hydrogen bonds, which create‬
‭bridges between different regions of the same strand (‬‭Figure 2.27b‬‭). Less commonly, a‬
‭polypeptide chain can form a beta-pleated sheet, in which hydrogen bonds form bridges‬
‭between different regions of a single polypeptide that has folded back upon itself or‬
‭between two or more adjacent polypeptide chains.‬
‭ he secondary structure of proteins further folds into a compact three-dimensional shape‬
T
‭called the protein’s‬‭tertiary structure‬‭(‬‭Figure 2.27c‬‭).‬‭In this configuration, amino acids‬
‭that are distant in the primary chain can be brought quite close via hydrogen bonds or, in‬
‭proteins containing cysteine, via disulfide bonds. A disulfide bond is a covalent bond‬
‭between sulfur atoms in a polypeptide.‬
‭ wo or more separate polypeptides often bond to form an even larger protein with a‬
T
‭quaternary structure (‬‭Figure 2.27d‬‭). The polypeptide‬‭subunits forming a‬‭quaternary‬
‭structure‬‭can be identical or different. For instance,‬‭hemoglobin, the protein found in red‬
‭blood cells, is composed of four tertiary polypeptides, two of which are called alpha chains‬
‭and two of which are called beta chains.‬
‭ hen proteins are exposed to extreme heat, acids, bases, and certain other substances,‬
W
‭they denature.‬‭Denaturation‬‭is a change in the structure‬‭o f a molecule through physical‬
‭o r chemical means. Denatured proteins lose their functional shape and can no longer‬
‭carry out their jobs. An everyday example of protein denaturation is milk curdling when‬
‭acidic lemon juice is added. Denaturation disrupts weaker chemical bonds, such as‬
‭hydrogen and disulfide bonds. However, peptide bonds are not affected.‬
‭ he contribution of the shape of a protein to its function can hardly be exaggerated. For‬
T
‭example, the long, slender shape of protein strands that make up muscle tissue is essential‬
‭to their ability to contract (shorten) and relax (lengthen). As another example, bones‬
‭contain long threads of a protein called collagen that acts as scaffolding upon which bone‬
‭minerals are deposited.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭59‬
‭Chapter 2: Chemistry‬

‭ igure 2.27‬ ‭The shapes of proteins: (a) The primary‬‭structure is the sequence of amino‬
F
‭acids in the polypeptide chain. (b) The secondary structure, an alpha-helix or a‬
‭beta-pleated sheet, is maintained by hydrogen bonds between amino acids in different‬
‭regions of the polypeptide strand. (c) The tertiary structure occurs due to further folding‬
‭and bonding of the secondary structure. (d) The quaternary structure occurs due to‬
‭interactions between two or more tertiary subunits. The example shown here is‬
‭hemoglobin, a protein in red blood cells (credit:‬‭O‬‭penStax‬‭). A link to a video explanation‬
‭o f this figure is available at‬‭Biology411.com‬‭.‬

‭ or more information, click the link below or scan the QR code to watch the TED video by‬
F
‭David Baker titled “Five challenges we could solve by designing new proteins”:‬

5 challenges we could solve by designing new proteins…

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭60‬
‭Chapter 2: Chemistry‬

‭Proteins Function as Enzymes‬

I‭ f you were trying to type a paper, and every time you hit a key on your laptop there was a‬
‭delay of six or seven minutes before you got a response, you would probably get a new‬
‭laptop. Similarly, without‬‭enzymes‬‭to catalyze (i.e.,‬‭speed up) chemical reactions, the‬
‭human body would be nonfunctional. It functions because enzymes function.‬
‭ nzymatic reactions—chemical reactions catalyzed by enzymes—begin when‬‭substrates‬
E
‭bind to the enzyme. A substrate is a‬‭reactant‬‭in an‬‭enzymatic reaction. This binding‬
‭o ccurs in regions of the enzyme known as‬‭active sites‬‭(‬‭Figure 2.28‬‭). Any given enzyme‬
‭catalyzes just one type of chemical reaction. This characteristic, called specificity, is‬
‭because a substrate with a particular shape and electrical charge can bind only to an‬
‭active site corresponding to that substrate.‬
‭ ue to this jigsaw puzzle-like match between an enzyme and its substrates, enzymes are‬
D
‭known for their specificity. In fact, as an enzyme binds to its substrate(s), the enzyme‬
‭structure changes slightly to find the best fit between the transition state (a structural‬
‭intermediate between the substrate and product) and the active site, just as a rubber glove‬
‭molds to a hand inserted into it. This active-site modification in the presence of substrate,‬
‭along with the simultaneous formation of the transition state, is called‬‭induced fit‬‭.‬
‭ verall, there is a specifically matched enzyme for each substrate and, thus, for each‬
O
‭chemical reaction; however, there is some flexibility. Some enzymes can act on several‬
‭different structurally similar substrates.‬
‭ he binding of a substrate produces an‬‭enzyme–substrate‬‭complex‬‭. Enzymes likely‬
T
‭speed up chemical reactions partly because the enzyme–substrate complex undergoes‬
‭temporary and reversible changes that cause the substrates to orient toward each other‬
‭in an optimal position to facilitate their interaction. These changes promote increased‬
‭reaction speed. The enzyme then releases the product(s) and resumes its original shape.‬
‭The enzyme is then free to engage in the process again and will do so as long as its‬
‭substrate remains.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭61‬
‭Chapter 2: Chemistry‬

‭ igure 2.28‬ ‭According to the induced-fit model,‬‭the enzyme's active site undergoes‬
F
‭conformational changes upon binding with the substrate. (a) Substrates approach active‬
‭sites on enzymes. (b) Substrates bind to active sites, producing an enzyme–substrate‬
‭complex. (c) Changes internal to the enzyme–substrate complex facilitate the interaction of‬
‭the substrates. (d) Products are released, and the enzyme returns to its original form,‬
‭ready to facilitate another enzymatic reaction. (credit:‬‭OpenStax‬‭) A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Other Functions of Proteins‬

‭ dvertisements for protein bars, powders, and shakes say protein is vital in building,‬
A
‭repairing, and maintaining muscle tissue. Still, the truth is that proteins contribute to all‬
‭body tissues, from the skin to the brain cells. Also, some proteins act as hormones,‬
‭chemical messengers that help regulate body functions. For example, growth hormone is‬
‭essential for skeletal growth, among other functions. Proteins in the cell membrane help‬
‭to transport electrolytes in and out of the cell, keeping these ions in a healthy balance.‬
‭ he body can use proteins for energy when carbohydrate and fat intake is inadequate,‬
T
‭and glycogen and adipose tissue stores become depleted. However, since there is no‬
‭storage site for protein except functional tissues, using protein for energy causes tissue‬
‭breakdown and results in body wasting.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭62‬
‭Chapter 2: Chemistry‬

‭Nucleic Acids‬
‭ he fourth organic compound important to human structure and function is the‬‭nucleic‬
T
‭acids‬‭(i.e., DNA and RNA). These polymers are composed‬‭o f‬‭nucleotides‬‭(‬‭Figure 2.29‬‭),‬
‭which are the monomers. A nucleotide is one of a class of organic compounds consisting‬
‭o f three subunits:‬
•‭ ‬ ‭ ne or more phosphate groups‬
o
‭•‬ ‭a pentose sugar (deoxyribose or ribose)‬
‭•‬ ‭a nitrogen-containing base (‬‭adenine‬‭,‬‭c ytosine‬‭,‬‭guanine‬‭,‬‭thymine‬‭, or‬‭uracil‬‭)‬
‭Nucleotides can be assembled into nucleic acids via dehydration synthesis reactions.‬

‭Chemical Features of Nucleic Acids‬

‭ ne way nucleic acids differ in their type of pentose sugar.‬‭Deoxyribonucleic acid‬‭(‬‭DNA)‬
O
‭contains deoxyribose (so-called because it has one less atom of oxygen than ribose), one‬
‭phosphate group, and one nitrogen-containing base. The “choices” of bases for DNA are‬
‭adenine (A), cytosine (C), guanine (G), and thymine (T). The order of these bases in DNA‬
‭specifies the blueprint for building proteins.‬
‭ ibonucleic acid‬‭(‬‭RNA‬‭) is a ribose-containing nucleotide‬‭that helps manifest the genetic‬
R
‭code as protein. RNA contains ribose, one phosphate group, and one nitrogen-containing‬
‭base, but the “choices” of base for RNA are adenine (A), cytosine (C), guanine (G), and‬
‭uracil (U). In other words, if a nucleic acid contains T bases, it is DNA. If it includes U‬
‭bases, then it is RNA.‬
‭ he nitrogen-containing bases adenine and guanine are classified as purines. A‬‭purine‬‭is‬
T
‭a nitrogen-containing base with a double-ring structure, which accommodates several‬
‭nitrogen atoms. The bases cytosine, thymine (found in DNA only), and uracil (found in‬
‭RNA only) are pyrimidines.‬‭Pyrimidines‬‭are nitrogen-containing‬‭bases with a single-ring‬
‭structure.‬
‭ onds formed by dehydration synthesis between the pentose sugar of one nucleic acid‬
B
‭monomer and the phosphate group of another form a “backbone,” from which the‬
‭components’ nitrogen-containing bases protrude. In DNA, two such backbones attach at‬
‭their protruding bases via hydrogen bonds. These twist to form a shape known as a‬
‭double helix‬‭(‬‭Figure 2.30a‬‭). The sequence of nitrogen-containing‬‭bases within a strand‬
‭o f DNA forms the genes that act as a molecular code instructing cells to assemble amino‬
‭acids into proteins. A more detailed diagram of DNA’s structure is shown in‬‭Figure 2.30b‬
‭and includes the‬‭sugar-phosphate backbone‬‭, the complementary‬‭base pairs, and the‬
‭hydrogen bonds between them.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭63‬
‭Chapter 2: Chemistry‬

I‭ n contrast, RNA consists of a single strand of sugar-phosphate backbone studded with‬

‭bases. The types and respective functions of RNA will be discussed in a later chapter.‬

‭ igure 2.29‬ ‭(a) The building blocks of all nucleotides‬‭are one or more phosphate groups,‬
F
‭a pentose sugar, and a nitrogen-containing base. (b) The nitrogen-containing bases of‬
‭nucleotides. (c) The two pentose sugars of DNA and RNA. (credit:‬‭OpenStax‬‭) A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭64‬
‭Chapter 2: Chemistry‬

‭(a)‬ ‭(b)‬

‭ igure 2.30‬ ‭(a)‬‭In the DNA double helix, two strands‬‭attach via hydrogen bonds between‬
F
‭the complementary bases of the component nucleotides. (b) In the structure of the DNA‬
‭double helix, hydrogen bonds connect the complementary bases of the two strands (i.e., A‬
‭and T; G and C). Note that there are two hydrogen bonds between A and T and three‬
‭between G and C. (credit: (a)‬‭OpenStax‬‭; (b)‬‭DNA‬‭molecular structure, showing individual‬
‭nucleotides and bonds.jpg‬‭;‬‭Francescakb‬‭;‬‭CC BY-SA 4.0‬‭).‬ ‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭65‬
‭Chapter 2: Chemistry‬

‭Adenosine Triphosphate‬
‭ he nucleotide‬‭adenosine triphosphate (ATP)‬‭is composed‬‭o f a ribose sugar, an adenine‬
T
‭base, and three phosphate groups‬‭(Figure 2.31‬‭). ATP‬‭is classified as a high-energy‬
‭compound because the two covalent bonds linking its three phosphates store significant‬
‭potential energy. In the body, the energy released from breaking these high-energy bonds‬
‭via hydrolysis helps fuel the body’s activities, from muscle contraction to transporting‬
‭substances in and out of cells to energy-requiring chemical reactions.‬

‭ igure 2.31‬ ‭ATP consists of the nitrogen-containing‬‭base adenine, the sugar ribose, and‬
F
‭three phosphate groups. Adenosine diphosphate (ADP) is missing the gamma phosphate‬
‭group. (credit:‬‭OpenStax‬‭) A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ hen a phosphate group is cleaved from ATP, the products are adenosine diphosphate‬
W
‭(‬‭ADP‬‭) and inorganic phosphate (P‬‭i‭)‬ , as shown in‬‭Figure‬‭2.32‬‭.‬
‭ his chemical reaction is reversible, meaning a phosphate group can be joined with ADP‬
T
‭(via dehydration synthesis) to produce ATP. Energy input is required to‬‭phosphorylate‬
‭(i.e., add a phosphate group) ADP.‬
‭ ells can also transfer a phosphate group from ATP to another organic compound. For‬
C
‭example, when glucose enters a cell, a phosphate group is transferred from ATP, forming‬
‭glucose phosphate (C‬‭6‭H
‬ ‬‭12‬‭O‬‭6‭—
‬ P) and ADP. Once phosphorylated,‬‭glucose can be stored as‬
‭glycogen or metabolized for immediate energy.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭66‬
‭Chapter 2: Chemistry‬

‭ igure 2.32‬ ‭Hydrolysis of ATP to ADP, phosphate,‬‭and energy. (credit:‬‭Hydrolysis of‬

F
‭ATP.png‬‭;‬‭SrKellyOP‬‭;‬‭CC0 1.0‬‭).‬ ‭A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭2.6 Career Connection - Pharmaceutical Chemist‬

‭ harmaceutical chemists are responsible for developing new drugs and trying to‬
P
‭determine the mode of action of both old and new drugs. They are involved in every step‬
‭o f the drug development process. We can find drugs in the natural environment or we can‬
‭synthesize them in the laboratory. In many cases, chemists change potential drugs from‬
‭nature chemically in the laboratory to make them safer and more effective, and sometimes‬
‭synthetic versions of drugs substitute for the version we find in nature.‬

‭ fter a drug's initial discovery or synthesis, the chemist then develops the drug, perhaps‬
A
‭chemically altering it, testing it to see if it is toxic, and then designing methods for efficient‬
‭large-scale production. Then, the process of approving the drug for human use begins.‬
‭The Food and Drug Administration (FDA) handles drug approval in the United States. This‬
‭involves a series of large-scale experiments using human subjects to ensure the drug is‬
‭not harmful and effectively treats its intended condition. This process often takes several‬
‭years and requires the participation of physicians, scientists, and chemists to complete‬
‭testing and gain approval.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭67‬
‭Chapter 2: Chemistry‬

‭ n example of a drug initially discovered in a living organism is Paclitaxel (Taxol), an‬

A
‭anti-cancer drug used to treat breast cancer. This drug was found in the bark of the Pacific‬
‭yew tree. Another example is aspirin, originally isolated from willow tree bark.‬

‭ inding drugs often means testing hundreds of samples of plants, fungi, and other life‬
F
‭forms to see if they contain any biologically active compounds. Sometimes, traditional‬
‭medicine can give modern medicine clues as to where to find an active compound. For‬
‭example, humankind has used willow bark to make medicine for thousands of years,‬
‭dating back to ancient Egypt. However, in the late 1800s, scientists and pharmaceutical‬
‭companies purified and marketed the aspirin molecule, acetylsalicylic acid, for human use.‬

‭ ccasionally, drugs developed for one use have unforeseen effects that allow usage in‬
O
‭o ther, unrelated ways. For example, scientists originally developed the drug minoxidil‬
‭(Rogaine) to treat high blood pressure. When tested on humans, researchers noticed that‬
‭individuals taking the drug would grow new hair. Eventually, the pharmaceutical company‬
‭marketed the drug to men and women with baldness to restore lost hair.‬

‭ pharmaceutical chemist's career may involve detective work, experimentation, and drug‬
A
‭development, all to make human beings healthier.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭68‬
‭Chapter 3: Cells and Associated Topics‬

‭ igure 3.1‬ ‭(a) Nasal sinus cells (viewed with a‬‭light microscope), (b) onion cells (viewed‬
F
‭with a light microscope), and (c)‬‭Vibrio tasmaniensis‬‭bacterial cells (seen through a‬
‭scanning electron microscope) are from very different organisms, yet all share certain‬
‭characteristics of basic cell structure. (credit a: modification of work by Ed Uthman, MD;‬
‭credit b: modification of work by Umberto Salvagnin; credit c: modification of work by‬
‭Anthony D'Onofrio, William H. Fowle, Eric J. Stewart, and Kim Lewis of the Lewis Lab at‬
‭Northeastern University; scale-bar data from Matt Russell;‬‭OpenStax‬‭)‬

‭3.1 Introduction‬
‭ lose your eyes and picture a brick wall. What is the fundamental component used to‬
C
‭build that wall? The answer is an individual brick. Like a brick wall, your body is‬
‭composed of a fundamental building block called “cells.” Think back to Chapter 1 and the‬
‭discussion of the hierarchy of biological organization. You learned that‬‭cells‬‭are the first‬
‭level (beginning with atoms) of organization to demonstrate the characteristics of life. A‬
‭living thing, whether made of one cell (like bacteria) or many cells (like a human), is called‬
‭an organism. Thus, cells are the basic building blocks of all organisms.‬
‭ ou developed from a single fertilized egg cell into a complex organism containing trillions‬
Y
‭o f cells that you see when you look in a mirror. Early, undifferentiated cells differentiated‬
‭and became specialized in their structure and function during this developmental process.‬
‭Some examples include:‬
‭‬
● ‭ pithelial cells protect the body's surface and cover the organs and cavities.‬
E
‭●‬ ‭Bone cells help to support and protect the body.‬
‭●‬ ‭Immune system cells fight invading pathogens.‬
‭●‬ ‭Blood cells carry nutrients and oxygen throughout the body while removing‬
‭carbon dioxide and other wastes.‬

‭69‬
‭Chapter 3: Cells and Associated Topics‬

‭‬
● ‭ uscle cells contract to move bones, digest food, and pump blood.‬
M
‭●‬ ‭Neural cells of the brain and spinal cord receive input, process it, and bring about‬
‭appropriate changes.‬
‭ ifferentiation is typically associated with changes in cells’ size, shape, metabolic activity,‬
D
‭and overall function. Because all cells in the body, beginning with the fertilized egg, contain‬
‭the same DNA, how do the different cell types come to be so different? The answer is‬
‭analogous to a movie script. The various actors in a movie all read from the same script;‬
‭however, they are each only reading their part of the script. Similarly, all cells contain the‬
‭same full complement of DNA, but each cell type only “reads” the DNA portions relevant to‬
‭its function. In biology, this is referred to as unique gene expression and is discussed‬
‭further in Chapter 4.‬
‭ ifferent cell types form specialized‬‭tissues‬‭, the‬‭next level in the hierarchy of biological‬
D
‭o rganization, that work in concert to perform all of the functions necessary for the living‬
‭o rganism. Tissues combine to form‬‭organs‬‭(e.g.,‬ ‭stomach, heart, and brain), and organs‬
‭function in an integrated manner to comprise‬‭organ‬‭systems‬‭(e.g., digestive, circulatory,‬
‭and nervous systems). These systems then work together to form an‬‭organism‬‭(e.g., a‬
‭human being).‬

‭3.2 Microscopy, Cell Theory, and Cell Size‬

‭ ells vary in size. With few exceptions, individual cells cannot be seen with the naked eye,‬
C
‭so scientists study them using microscopes (micro- = “small”; -scope = “to look at”). A‬
‭microscope is an instrument that magnifies an object, and most cell photographs are taken‬
‭with a microscope.‬
‭ he concept of a cell started with microscopic observations of dead cork tissue by scientist‬
T
‭Robert Hooke in 1665. Without realizing their function or importance, Hook coined the‬
‭term “cell” based on the resemblance of the small subdivisions in the cork to monks'‬
‭rooms, called cells (‬‭Figure 3.2a‬‭). About ten years‬‭later, Antonie van Leeuwenhoek‬
‭became the first to observe living and moving cells, bacteria, and protozoa under a‬
‭microscope (‬‭Figure 3.2b‬‭). Later advances in lenses,‬‭microscope construction, and staining‬
‭techniques enabled other scientists to see some components inside cells.‬
‭ y the late 1830s, Matthias Schleiden and Theodor Schwann were studying tissues and‬
B
‭proposed the‬‭unified cell theory‬‭, which states that‬‭all living things are composed of one‬
‭o r more cells, the cell is the basic unit of life, and new cells arise from existing cells.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭70‬
‭Chapter 3: Cells and Associated Topics‬

‭ or additional information on the development of cell theory, click the link or scan the QR‬
F
‭code to watch the TED-Ed video by Lauren Royal-Woods titled “The wacky history of cell‬
‭theory.”‬

The wacky history of cell theory - Lauren Royal-W…

‭(a) (b)‬
‭ igure 3.2‬ ‭a.‬‭Drawings of cells observed by Robert‬‭Hooke. Book in the Golub Collection‬
F
‭o f Antique Microscopes exhibited at the University of California, Berkeley - Berkeley,‬
‭California, USA. b. Illustration from the “Selected Works of Leeuwenhoek…” showing‬
‭animalcules seen in white wine vinegar. (credit:‬ ‭Illustration from "The selected works of‬
‭Leeuwenhoek...";‬‭wellcomeimages.org‬‭;‬‭CC BY 4.0‬‭)‬

‭ s previously stated, many types of cells require magnification to be seen. But how large‬
A
‭is a cell compared to other reference items (e.g., atoms, proteins, human eggs, and chicken‬
‭eggs)?‬ ‭Figure 3.3‬‭provides a visual comparison using‬‭a logarithmic scale.‬
‭ reasonable question is, “Why are individual cells so small?” One answer concerns‬
A
‭surface area, volume, and their ratio (surface area divided by volume). An explanation‬
‭requires some math, so don’t panic! While many cells are somewhat spherical, we will‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭71‬
‭Chapter 3: Cells and Associated Topics‬

‭ isualize them as cubes for this discussion, as the calculations are more straightforward‬
v
‭(‬‭Figure 3.4‬‭). The‬‭surface area‬‭o f an individual side‬‭o f the cube is determined by‬
‭multiplying the length and width (1 mm x 1 mm = 1 mm squared; 1mm‬‭2‬‭). As a cube has six‬
‭sides, the total surface area is calculated by adding the values for each side, which is 6‬
‭mm‬‭2‬‭.‬ ‭Volume‬‭is a cubed calculation determined by‬‭multiplying the object's length, width,‬
‭and height. Therefore, the cube previously discussed has a volume of 1 mm cubed (1 mm‬
‭x 1mm x 1mm = 1mm‬‭3‭)‬ . The‬‭surface area to volume ratio‬‭is shown as the two values,‬
‭separated by a colon (:). In this case, the ratio is 6:1. If the size of each side of the cube is‬
‭doubled (i.e., 2 mm), then the resulting ratio is 24:8, which reduces to 3:1. In other words,‬
‭there is proportionally less surface area as the volume of a cell increases.‬

‭ onsequently, the plasma membrane will not have enough surface area to support the‬
C
‭rate of diffusion required for the increased volume. As a cell grows, it becomes less‬
‭efficient. One way to become more efficient is to divide.‬

‭ igure 3.3‬ ‭The relative sizes of various items on‬‭a logarithmic scale, meaning that each‬
F
‭unit of increase (e.g., 1 to 10 or 10 to 100) represents a 10-fold increase in the measured‬
‭quantity. Remember that pH (discussed in Chapter 2) is also measured using a logarithmic‬
‭scale. Note the approximate size range visible by the naked eye (no additional‬
‭magnification), a light microscope, and an electron microscope. (credit:‬‭OpenStax‬‭). A link‬
‭to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭72‬
‭Chapter 3: Cells and Associated Topics‬

‭ or additional information about the implications of surface area to volume ratio and cell‬
F
‭size, click the link below or scan the QR code‬‭to‬‭watch the TED-Ed video by Murray Gans‬
‭titled “What is the biggest single-cell organism?”.‬

What is the biggest single-celled organism? - Murry Gans

‭ igure 3.4‬ ‭Notice that as a cell increases in size,‬‭its surface area-to-volume ratio‬
F
‭decreases. The cell on the left has a volume of 1 mm‬‭3‭,‬ a surface area of 6 mm‬‭2‬‭, and a‬
‭surface area-to-volume ratio of 6 to 1(i.e., 6:1). In contrast, the cell on the right has a‬
‭volume of 8 mm‬‭3‭,‬ a surface area of 24 mm‬‭2‭,‬ and a surface‬‭area-to-volume ratio of 8:24, or‬
‭3:1. (credit:‬‭OpenStax‬‭) A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭3.3 Cell Types‬

‭ ells fall into two broad categories: prokaryotic and eukaryotic. Bacteria are classified as‬
C
‭prokaryotes‬‭(pro- = “before”; -kary- = “nucleus”),‬‭while cells of animals and plants are‬
‭eukaryotes‬‭(eu- = “true”) and have a nucleus.‬
‭ ells of both categories share four components: 1) a plasma membrane, an outer covering‬
C
‭that separates the cell’s interior from its surrounding environment; 2) cytoplasm,‬
‭consisting of a jelly-like cytosol within the cell in which other cellular components are‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭73‬
‭Chapter 3: Cells and Associated Topics‬

f‭ ound; 3) DNA, the genetic material of the cell; and 4) ribosomes, which synthesize‬
‭proteins. However, prokaryotes differ from eukaryotic cells in several ways, as discussed‬
‭in the next section.‬

‭Prokaryotic Cells‬
‭ prokaryote is a simple, single-celled (unicellular) organism; some examples of these cells‬
A
‭are shown in‬‭Figure 3.5‬‭.‬
‭ prokaryotic cell lacks a nucleus or other membrane-bound organelles, which are‬
A
‭structures in eukaryotic cells with specific functions. Prokaryotic DNA is found in a central‬
‭part of the nucleoid (‬‭Figure 3.6)‬‭, and most DNA is‬‭contained in one circular chromosome.‬
‭Additional genetic information is contained in structures called plasmids, which are tiny,‬
‭circular DNA structures. Plasmids are exchanged between bacterial cells and are‬
‭associated with the phenomenon of antibiotic resistance.‬

‭(a) (b) (c)‬

‭ igure 3.5‬ ‭Three‬‭examples of stained, magnified‬‭bacterial cells: a. Bacillus cereus.‬
F
‭Leifson flagella stain. (credit: (a) CDC/Dr. William A. Clark; Original uploader user‬
‭Marcus007‬‭;‬‭CC0 1.0‬‭(b)‬‭Pseudomonas fluorescens Gram‬‭Stain on Microscope Slide.jpg‬‭;‬‭B.‬
‭Domangue‬‭;‬‭Creative Commons Attribution-Share Alike‬‭4.0‬ ‭(c)‬‭Staphylococcus in‬
‭Magnification of 4000X.jpg‬‭;‬‭Ajay Kumar Chaurasiya‬‭;‬‭Creative Commons Attribution-Share‬
‭Alike 4.0‬‭)‬

‭ rokaryotes typically have a cell wall, which is a rigid structure that contributes to the‬
P
‭shape of the cells and protects them against changes in water pressure in the‬
‭environment.‬‭Antibiotics‬‭such as penicillin interfere‬‭with the synthesis of the cell wall and‬
‭cause bacterial cells to lyse without affecting the human host. Some prokaryotes also have‬

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‭Chapter 3: Cells and Associated Topics‬

s‭ tructures called flagella and pili. Flagella are used for movement, while pili are used to‬
‭exchange genetic material with other bacterial cells.‬

‭ igure 3.6‬ ‭All prokaryotes have chromosomal DNA localized‬‭in a nucleoid, ribosomes, a‬
F
‭cell membrane, and a cell wall. (credit:‬‭OpenStax‬‭)‬

‭Eukaryotic Cells‬
‭ nlike prokaryotic cells, eukaryotic cells have 1) a membrane-bound nucleus, 2)‬
U
‭numerous membrane-bound organelles (e.g., plasma membrane, endoplasmic reticulum,‬
‭Golgi apparatus, and mitochondria), which are responsible for specific functions, and 3)‬
‭several, rod-shaped chromosomes (‬‭Figure 3.7‬‭).‬
‭ or additional information about the possible origins of eukaryotic cells, click the link‬
F
‭below or scan the QR code to watch the TED-Ed video by Adam Jacobson titled “How we‬
‭think complex cells evolved.”‬

How we think complex cells evolved - Adam Jacobson

‭ et's first examine the structure and function of the plasma membrane, the organelle that‬
L
‭separates the inside and outside of a cell.‬

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‭Chapter 3: Cells and Associated Topics‬

‭ igure 3.7‬ ‭This figure shows a generalized eukaryotic‬‭cell's major organelles and other‬
F
‭components. (credit:‬ ‭Figure 04 03 01a.png‬‭; Wikimedia‬‭Commons;‬‭CC BY 4.0‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭3.4 Components and Structure of the Plasma Membrane‬

‭ cell’s‬‭plasma membrane‬‭defines the cell, outlines‬‭its borders, and determines the‬
A
‭nature of its interaction with its environment. Cells exclude some substances, take in‬
‭o thers, and excrete others in controlled quantities. The plasma membrane must be very‬
‭flexible to allow specific cells, such as red and white blood cells, to change shape as they‬
‭pass through narrow capillaries. These are the more obvious plasma membrane functions.‬
‭In addition, the plasma membrane's surface carries markers that allow cells to recognize‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ ne another, which plays a role in the immune response's “self” versus “non-self”‬
o
‭distinction.‬
S‭ cientists identified the plasma membrane in the 1890s and its chemical components in‬
‭1915. The proposed structure of the plasma membrane, referred to as the‬‭fluid mosaic‬
‭model‬‭, was introduced in 1972 by S.J. Singer and Garth‬‭L. Nicolson. This model has‬
‭evolved, but it still best accounts for plasma membrane structure and function as we now‬
‭understand them. The fluid mosaic model describes the plasma membrane structure as a‬
‭mosaic of components—including phospholipids, cholesterol, proteins, and‬
‭carbohydrates—that give the membrane a fluid character (‬‭Figure 3.8‬‭).‬

‭ igure 3.8‬ ‭The fluid mosaic model describes the plasma‬‭membrane as a dynamic‬
F
‭combination of phospholipids, cholesterol, and proteins. Carbohydrates attached to lipids‬
‭(glycolipids) and proteins (glycoproteins) extend from the membrane's outward-facing‬
‭surface. (credit:‬‭OpenStax‬‭). A link to a video explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ he membrane's main fabric comprises‬‭phospholipid‬‭molecules. As discussed in Chapter‬

T
‭2, these molecules consist of a three-carbon glycerol backbone with two fatty acid‬
‭molecules attached to carbons 1 and 2 and a phosphate-containing group attached to the‬
‭third carbon (‬‭Figure 3.9‬‭). This arrangement gives‬‭the overall molecule a head area (the‬

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‭Chapter 3: Cells and Associated Topics‬

‭ hosphate-containing group), which has a polar character or negative charge, and a tail‬
p
‭area (the fatty acids), which has a nonpolar character or no charge.‬

‭ igure 3.9‬ ‭A hydrophilic head and two hydrophobic‬‭tails comprise this phospholipid‬
F
‭molecule. The hydrophilic head group consists of a phosphate-containing group attached‬
‭to a glycerol molecule. The hydrophobic tails, each containing either a saturated or an‬
‭unsaturated fatty acid, are long hydrocarbon chains. (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

I‭ n water, phospholipids arrange themselves so their‬‭hydrophobic tails‬‭face each other‬

‭and their‬‭hydrophilic head‬‭s face out. In this way,‬‭they form a‬‭lipid bilayer‬‭. This‬
‭double-layered phospholipid barrier separates the water and other materials on one side‬
‭from the water and other materials on the other side (‬‭Figure 3.10‬‭).‬
‭ he plasma membranes of cells specializing in absorption are folded into fingerlike‬
T
‭projections called‬‭microvilli‬‭(singular = microvillus);‬‭(‬‭Figure 3.11‬‭). Such cells are typically‬
‭found lining the small intestine, the organ that absorbs nutrients from digested food.‬
‭People with celiac disease have an immune response to gluten, which is a protein found in‬
‭wheat, barley, and rye. The immune response damages microvilli; thus, afflicted individuals‬

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‭Chapter 3: Cells and Associated Topics‬

i‭nefficiently absorb nutrients. This outcome leads to malnutrition, cramping, and diarrhea.‬
‭Patients who have celiac disease must follow a gluten-free diet.‬

‭ igure 3.10‬ ‭The phospholipid bilayer consists of‬‭two adjacent sheets of phospholipids,‬
F
‭arranged tail to tail. The hydrophobic tails associate with one another, forming the interior‬
‭o f the membrane. The polar heads contact the fluid inside and outside of the cell. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ igure 3.11‬ ‭Microvilli, shown here as they appear‬‭o n cells lining the small intestine,‬
F
‭increase the surface area available for absorption. These microvilli are only found on the‬
‭part of the plasma membrane that faces the cavity from which substances will be‬
‭absorbed. (credit "micrograph": modification of work by Louisa Howard;‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 3: Cells and Associated Topics‬

‭ or additional information on cell membranes, click the link below or scan the QR code to‬
F
‭watch the TED-Ed video by Nazzy Pakpour titled “Cell membranes are way more‬
‭complicated than you think.”‬

Cell membranes are way more complicated than you think - Nazz…

‭3.5 Transport Across the Plasma Membrane‬

‭ lasma membranes must allow certain substances to enter and leave a cell while‬
P
‭preventing some harmful materials from entering and some essential materials from‬
‭leaving. In other words, plasma membranes are‬‭selectively‬‭permeable‬‭—they allow some‬
‭substances to pass through but not others. If they were to lose this selectivity, the cell‬
‭would no longer be able to sustain itself and would be destroyed.‬
‭ he membrane’s lipid bilayer structure provides the first level of control. The‬
T
‭phospholipids are tightly packed, and the membrane has a hydrophobic interior. This‬
‭structure causes the membrane to be selectively permeable and allows only substances‬
‭meeting specific criteria to pass through it unaided. In the case of the cell membrane, only‬
‭relatively small, nonpolar materials can move through the lipid bilayer (remember, the‬
‭lipid tails of the membrane are nonpolar). Some examples are other lipids, oxygen and‬
‭carbon dioxide gases, and alcohol. However, water-soluble materials—like glucose, amino‬
‭acids, and electrolytes—need some assistance to cross the membrane because the‬
‭hydrophobic tails of the phospholipid bilayer repel them. All substances that move‬
‭through the membrane do so by one of two general methods, which are categorized‬
‭based on whether or not energy is required.‬‭Passive‬‭transport‬‭is the movement of‬
‭substances across the membrane without cellular energy expenditure. In contrast,‬‭active‬
‭transport‬‭is the movement of substances across the‬‭membrane using energy from‬
‭adenosine triphosphate‬‭(‬‭ATP‬‭).‬

‭Passive Transport‬
I‭ t is necessary to comprehend concentration gradients and diffusion to understand‬‭how‬
‭substances move passively across a cell membrane. A‬‭concentration gradient‬‭is the‬
‭difference in concentration of a substance across a space. Molecules (or ions) will‬
‭spread/diffuse from where they are more concentrated to where they are less‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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c‭ oncentrated until equally distributed in that space (When molecules move this way, they‬
‭are said to move‬‭down‬‭their concentration gradient.).‬‭Diffusion‬‭is the movement of‬
‭particles from an area of higher concentration to an area of lower concentration. A couple‬
‭o f common examples will help to illustrate this concept. Imagine being inside a closed‬
‭bathroom. If a person sprayed a bottle of perfume, the scent molecules would naturally‬
‭diffuse from the spot where they left the bottle to all corners of the bathroom, and this‬
‭diffusion would go on until no more concentration gradient remains.‬
‭ nother example is a spoonful of sugar placed in a cup of tea. Eventually, the sugar will‬
A
‭diffuse throughout the tea until no concentration gradient remains. In both cases, if the‬
‭room is warmer or the tea hotter, diffusion occurs even faster as the molecules bump into‬
‭each other and spread out more quickly than at cooler temperatures. Having an internal‬
‭body temperature of around 98.6‬‭°‬ ‭F thus also aids‬‭in the diffusion of particles within the‬
‭body.‬
‭ henever a substance exists in greater concentration on one side of a semipermeable‬
W
‭membrane, such as the cell membrane, any substance that can move down its‬
‭concentration gradient across the membrane will do so. Consider substances that can‬
‭quickly diffuse through the cell membrane's lipid bilayer, such as the gases oxygen (O‬‭2‭)‬ ‬
‭and CO‬‭2‬‭. O‬‭2‬ ‭generally diffuses into cells because‬‭it is more concentrated outside of them,‬
‭and CO‬‭2‬ ‭typically diffuses out of cells because it‬‭is more concentrated inside. Neither of‬
‭these examples requires any energy on the part of the cell, and therefore they use passive‬
‭transport to move across the membrane. This mechanism of molecules moving across a‬
‭cell membrane from the side where they are more concentrated to the side where they‬
‭are less concentrated requires no energy on the part of the cell. It is a type of passive‬
‭transport called‬‭simple diffusion‬‭(‬‭Figure 3.12‬‭).‬
‭ arge polar or ionic molecules, which are hydrophilic, cannot easily cross the‬
L
‭phospholipid bilayer. Charged atoms or molecules of any size cannot cross the cell‬
‭membrane via simple diffusion as the hydrophobic tails repel the charges in the interior of‬
‭the phospholipid bilayer. Solutes dissolved in water on either side of the cell membrane‬
‭will tend to diffuse down their concentration gradients. Still, because most substances‬
‭cannot pass freely through the cell membrane's lipid bilayer, their movement is restricted‬
‭to‬‭c hannel proteins‬‭and‬‭c arrier proteins‬‭in the membrane.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 3: Cells and Associated Topics‬

‭ igure 3.12‬ ‭The structure of the phospholipid bilayer‬‭allows small, uncharged (i.e.,‬
F
‭nonpolar) substances, such as oxygen and carbon dioxide, and hydrophobic molecules,‬
‭such as lipids, to pass through the cell membrane, down their concentration gradient, by‬
‭simple diffusion. (credit:‬‭OpenStax‬‭). A link to a‬‭video explanation of this figure is available‬
‭at‬‭Biology411.com‬‭.‬

‭ acilitated diffusion‬‭is the process used for those‬‭substances that cannot cross the lipid‬
F
‭bilayer due to their size, charge, or polarity (‬‭Figure‬‭3.13‬‭). A common example of‬
‭facilitated diffusion is the movement of glucose into the cell, which is used to make ATP‬
‭(i.e., cellular metabolism; Chapter 6). Although glucose can be more concentrated outside‬
‭a cell, it cannot cross the lipid bilayer via simple diffusion because it is both large and‬
‭polar. To resolve this, a specialized carrier protein called the glucose transporter will‬
‭transfer glucose molecules into the cell to facilitate its inward diffusion.‬
‭ or example, even though sodium ions (Na‬‭+‭)‬ are highly‬‭concentrated outside of cells,‬
F
‭these electrolytes are charged and cannot pass through the nonpolar lipid bilayer of the‬
‭membrane. Their diffusion is facilitated by membrane proteins that form sodium channels‬
‭(or “pores”) so that Na‬‭+‬ ‭ions can move down their‬‭concentration gradient from outside‬
‭the cells to inside the cells. Many other solutes must undergo facilitated diffusion to move‬
‭into a cell, such as amino acids, or out of a cell, such as wastes. Because facilitated‬
‭diffusion is a passive process, it does not require energy expenditure by the cell.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 3: Cells and Associated Topics‬

‭ igure 3.13‬ ‭(a) Facilitated diffusion of substances‬‭crossing the cell (plasma) membrane‬
F
‭takes place with the help of channel and carrier proteins. Channel proteins are less‬
‭selective than carrier proteins and usually mildly discriminate between their cargo based‬
‭o n size and charge. (b) Carrier proteins are more selective, often only allowing one type of‬
‭molecule to cross. (credit:‬‭OpenStax‬‭). A link to‬‭a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ hannel and carrier proteins transport material at different rates. Channel proteins‬
C
‭facilitate diffusion at a rate of tens of millions of molecules per second. In contrast, carrier‬
‭proteins work at a rate of a thousand to a million molecules per second.‬

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‭83‬
‭Chapter 3: Cells and Associated Topics‬

‭ ater also can move freely across the cell membrane of all cells, either through protein‬
W
‭channels or by slipping between the lipid tails of the membrane itself.‬‭Osmosis‬‭is the‬
‭diffusion of water through a semipermeable membrane (‬‭Figure 3.14‬‭).‬

‭ igure 3.14‬ ‭Osmosis is the diffusion of water through‬‭a semipermeable membrane down‬
F
‭its concentration gradient. If a membrane is permeable to water, though not to a solute,‬
‭water will equalize its concentration by diffusing to the side of lower water concentration‬
‭(and thus the side of higher solute concentration). In the beaker on the left, the solution‬
‭o n the right side of the membrane is hypertonic. (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ he movement of water molecules is not regulated by some cells, so these cells must be‬
T
‭exposed to an environment in which the concentration of‬‭solutes‬‭o utside the cells (in the‬
‭extracellular fluid) is equal to the concentration of solutes inside the cells (in the‬
‭cytoplasm). The term‬‭tonicity‬‭describes the relative‬‭solution concentrations of two‬
‭solutions, for example, inside and outside the cell. Two solutions with the same‬
‭concentration of solutes are said to be‬‭isotonic‬‭.‬‭When cells and their extracellular‬
‭environments are isotonic, the concentration of water molecules is the same outside and‬
‭inside the cells, and the cells maintain their standard shape and function. Water still‬
‭moves in both directions across the membrane, but there is no‬‭net diffusion‬‭(i.e., water‬
‭diffusion into and out of the cell is equal.)‬
‭ smosis occurs when there is an imbalance of‬‭solutes‬‭o utside and inside the cell, and the‬
O
‭solutes‬‭can’t‬‭pass through the cell membrane. A solution‬‭with a higher concentration of‬
‭solutes than another solution is said to be‬‭hypertonic‬‭,‬‭and water molecules tend to‬
‭diffuse‬‭into‬‭a hypertonic solution (‬‭Figure‬‭3.15‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭84‬
‭Chapter 3: Cells and Associated Topics‬

‭(a)‬

‭(b)‬

‭ igure 3.15‬ ‭(a) A hypertonic solution has a solute‬‭concentration higher than another‬
F
‭solution. An isotonic solution has a solute concentration equal to another solution. A‬
‭hypotonic solution has a solute concentration lower than another solution. Net osmosis‬
‭o ccurs towards the hypertonic solution, whether inside or outside the cell. A cell in an‬
‭isotonic solution experiences no net osmosis and retains its shape and function. (Source:‬
‭Cellular Tonicity.png‬‭;‬‭Uploaded by User‬‭Connectivid-D‬‭;‬‭CC BY-SA 4.0‬‭) (b) Illustration of‬
‭tonicity and osmosis using red blood cells placed into hypertonic, isotonic, and hypotonic‬
‭solutions. Red blood cells lack mechanisms commonly found in other cells to prevent them‬
‭from taking on too much water. (credit: Mariana Ruiz Villareal;‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ his observation might seem contradictory, as the hypertonic solution has more solute.‬
T
‭However, osmosis refers to the diffusion of‬‭water‬‭,‬‭not‬‭solute‬‭. While a hypertonic solution‬
‭has a‬‭higher solute concentration‬‭than a second solution,‬‭it has a‬‭lower water concentration‬‭.‬
‭If the membrane is impermeable to the solute but not water, then water will diffuse along‬
‭its concentration gradient (high to low), and thus there will be net osmosis‬‭into‬‭the‬

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‭Chapter 3: Cells and Associated Topics‬

‭ ypertonic cell. However, if too much water diffuses into a cell, the internal pressure can‬
h
‭become so great that the cell will rupture and die. Therefore, maintaining an appropriate‬
‭water balance (i.e., homeostasis) is critical for cells.‬
‭ onversely, if cells have less solute than an outside solution, then the cells are said to be‬
C
‭hypotonic‬‭to the solution. As tonicity is a relative‬‭comparison, this situation can also be‬
‭correctly stated as the outside solution is hypertonic to the cells. Therefore, if cells are‬
‭placed into a hypertonic solution, there will be net osmosis out of the cells, and they will‬
‭shrivel due to water loss.‬
‭ he rate of diffusion, both simple and facilitated, can be influenced by a variety of factors,‬
T
‭including the following:‬
‭•‬ ‭ he extent of the concentration gradient:‬‭The greater‬‭the difference in concentration,‬
T
‭the more rapid the diffusion. The closer the distribution of the material gets to‬
‭equilibrium, the slower the diffusion rate becomes.‬

‭•‬ S‭ olubility:‬‭As discussed earlier, nonpolar or lipid-soluble‬‭materials pass through‬

‭plasma membranes more easily than polar materials, allowing a faster diffusion rate.‬

‭•‬ ‭ istance traveled:‬‭The greater the distance a substance‬‭must travel, the slower the‬
D
‭diffusion rate. This fact places an upper limitation on cell size. A large, spherical cell‬
‭will die because nutrients or waste cannot reach or leave the center of the cell,‬
‭respectively. Therefore, cells must either be small in size, as in the case of many‬
‭prokaryotes, or flattened, as with many single-celled eukaryotes.‬

‭Active Transport‬
I‭ n cases of diffusion (simple or facilitated), the cell expends no energy. Membrane proteins‬
‭that aid in the passive transport of substances do so without using ATP. During active‬
‭transport, ATP is required to move a substance across a membrane, often with the help of‬
‭protein carriers, and usually‬‭against‬‭its concentration‬‭gradient. In other words, energy‬
‭must be expended to overcome diffusion‬‭along‬‭its concentration‬‭gradient.‬
‭ ne of the most common types of active transport involves proteins that serve as pumps.‬
O
‭The word “pump” probably conjures up thoughts of using energy to pump up the tire of a‬
‭bicycle or a basketball. Similarly, energy from ATP is required for these membrane‬
‭proteins to transport substances—molecules or ions—across the membrane, usually‬
‭against their concentration gradients (from an area of low concentration to an area of‬
‭high concentration).‬

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‭86‬
‭Chapter 3: Cells and Associated Topics‬

‭ he‬‭sodium-potassium pump‬‭transports sodium ions (Na‬‭+‬‭) out of a cell while moving‬

T
‭potassium ions (K‬‭+‭)‬ into the cell (‬‭Figure 3.16‬‭). These‬‭pumps are particularly abundant in‬
‭nerve cells, which are constantly pumping out sodium ions and pulling in potassium ions‬
‭to enable electrical “messages” (i.e., nerve impulses) to move along the cells. This process‬
‭accounts for the majority of their ATP usage.‬
‭ wo other forms of active transport, endocytosis and exocytosis, do not involve‬
T
‭membrane carriers.‬‭Endocytosis‬‭(bringing “into the‬‭cell”) is the process of taking in the‬
‭material by enveloping it in a portion of the cell membrane and then pinching off that‬
‭portion of the membrane. Once pinched off, the part of the membrane and its contents‬
‭become an independent, intracellular‬‭vesicle‬‭(i.e.,‬‭a membranous sac).‬

‭ igure 3.16‬ ‭The sodium-potassium pump is found in‬‭many cell (plasma) membranes.‬
F
‭Powered by ATP, the pump moves sodium and potassium ions in opposite directions, each‬
‭against its concentration gradient. In a single pump cycle, three sodium ions are extruded,‬
‭and two potassium ions are imported into the cell. (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ ndocytosis often brings materials into the cell that must be broken down or digested, and‬
E
‭two types are shown in‬‭Figure 3.17‬‭. Phagocytosis (“cell‬‭eating”) is the endocytosis of large‬
‭particles. Many immune cells engage in phagocytosis of invading pathogens, and their job‬
‭is to patrol body tissues for unwanted matter, such as invading bacterial cells, phagocytize‬
‭them, and digest them.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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I‭ n contrast to phagocytosis, pinocytosis (“cell drinking”) brings fluid containing dissolved‬

‭substances into a cell through membrane vesicles.‬
I‭ n contrast with endocytosis,‬‭exocytosis‬‭(taking “out‬‭o f the cell”) is the process of a cell‬
‭exporting material using vesicular transport (‬‭Figure‬‭3.18‬‭). Many cells manufacture‬
‭substances that must be secreted, like a factory manufacturing a product for export. These‬
‭substances are typically packaged into membrane-bound vesicles within the cell. When the‬
‭vesicle membrane fuses with the cell membrane, the vesicle releases its contents into the‬
‭fluid between cells. The vesicle membrane then becomes part of the cell membrane.‬

‭(a) (b)‬

‭ igure 3.17‬ ‭Endocytosis is a form of active transport‬‭in which a cell envelopes‬

F
‭extracellular materials using its cell membrane. (a) In phagocytosis, which is relatively‬
‭nonselective, the cell takes in a large particle. (b) In pinocytosis, the cell takes in small‬
‭particles in a fluid. (credit:‬‭OpenStax‬‭)‬

‭ he stomach and pancreas cells produce and secrete digestive enzymes through‬
T
‭exocytosis (‬‭Figure 3.19‬‭). This process is also used‬‭by endocrine cells to secrete hormones‬
‭that are sent throughout the body and by specific immune cells that secrete large amounts‬
‭o f histamine, a chemical necessary for immune responses.‬

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‭ igure 3.18‬ ‭Exocytosis is much like endocytosis‬‭in reverse. Material destined for export‬
F
‭is packaged into a vesicle inside the cell. The vesicle's membrane fuses with the cell‬
‭membrane and the contents are released into the extracellular space. (credit:‬‭OpenStax‬‭)‬

‭ igure 3.19‬ ‭The pancreatic acinar cells produce‬‭and secrete many digestive enzymes.‬
F
‭The tiny black granules in this electron micrograph are secretory vesicles filled with‬
‭enzymes that will be exported from the cells via exocytosis. LM × 2900. (credit:‬
‭Micrograph provided by the Regents of University of Michigan Medical School © 2012;‬
‭OpenStax‬‭)‬

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‭Concepts Application: Cystic Fibrosis‬

‭ ystic fibrosis‬‭(CF) is a genetic disease affecting‬‭approximately 30,000 people in the‬

C
‭United States, with about 1,000 new cases reported yearly. This disease is most well‬
‭known for its damage to the lungs, causing breathing difficulties and chronic lung‬
‭infections, but it also affects the liver, pancreas, and intestines (‬‭Figure 3.20‬‭). Only about 50‬
‭years ago, the prognosis for children born with CF was bleak—a life expectancy rarely‬
‭over ten years. Today, with advances in medical treatment, many CF patients live into their‬
‭30s.‬

‭ he symptoms of CF result from a malfunctioning membrane ion channel called the cystic‬
T
‭fibrosis transmembrane conductance regulator, or‬‭CFTR‬‭.‬‭In healthy people, the CFTR‬
‭protein is a membrane protein that transports Cl‬‭–‬ ‭ions out of the cell. In a person with CF,‬
‭the CFTR gene is mutated; thus, the cell manufactures a defective channel protein that‬
‭typically is not incorporated into the membrane but is instead degraded by the cell.‬

‭ he CFTR requires ATP to function, making its Cl‬‭–‬ ‭transport a form of active transport.‬
T
‭This characteristic puzzled researchers for a long time because the Cl‬‭–‬ ‭ions flow‬‭down‬
‭their concentration gradient when transported out of cells. Active transport generally‬
‭pumps ions‬‭against‬‭their concentration gradient, but‬‭the CFTR presents an exception to‬
‭this rule.‬

I‭ n normal lung tissue, the movement of Cl‬‭–‬ ‭o ut of‬‭the cell maintains a Cl‬‭–‬‭-rich, negatively‬
‭charged environment immediately outside the cell. This result is significant in the epithelial‬
‭lining of the respiratory system. Respiratory epithelial cells secrete mucus, trapping dust,‬
‭bacteria, and other debris. A cilium (plural = cilia) is one of the hair-like appendages found‬
‭o n certain cells. Cilia on the epithelial cells move the mucus and its trapped particles up‬
‭the airways away from the lungs and toward the outside. To be effectively moved upward,‬
‭the mucus cannot be too viscous; rather, it must have a thin, watery consistency. The‬
‭transport of Cl‬‭–‬ ‭and the maintenance of an electronegative‬‭environment outside the cell‬
‭attract positive ions such as Na‬‭+‬ ‭to the extracellular‬‭space. The accumulation of Cl‬‭–‬ ‭and Na‬‭+‬
‭ions in the extracellular space creates solute-rich mucus with a low concentration of water‬
‭molecules. As a result, through osmosis, water moves from cells and the extracellular‬
‭matrix into the mucus, “thinning” it out. This result is how, in a typical respiratory system,‬
‭the mucus is kept sufficiently watered down to be propelled out of the respiratory system.‬

I‭ f the CFTR channel is absent, Cl‬‭–‬ ‭ions are not transported‬‭o ut of the cell in adequate‬
‭numbers, thus preventing them from drawing positive ions. The absence of ions in the‬
‭secreted mucus results in the lack of a standard water concentration gradient. Therefore,‬
‭there is no osmotic pressure pulling water into the mucus. The resulting mucus is thick‬
‭and sticky, and the ciliated epithelia cannot effectively remove it from the respiratory‬
‭system. Passageways in the lungs become blocked with mucus and the debris it carries, as‬

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s‭ hown in the figure on the next page. Bacterial infections occur more easily because‬
‭bacterial cells are not effectively carried away from the lungs.‬

‭ igure 3.20‬ ‭Cystic fibrosis: Figure A shows the‬‭o rgans that cystic fibrosis can affect.‬
F
‭Figure B shows a cross-section of a normal airway. Figure C shows an airway with cystic‬
‭fibrosis. The widened airway is blocked by thick, sticky mucus that contains blood and‬
‭bacteria. (credit:‬ ‭Cysticfibrosis01.jpg‬‭; National‬‭Heart Lung and Blood Institute (NIH);‬‭CC0‬
‭1.0‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭3.6 The Cytoplasm and Cellular Organelles‬

‭ ow that you have learned that the cell membrane surrounds all cells, you can dive inside‬
N
‭a prototypical human cell to learn about its internal components and functions. All living‬
‭cells in multicellular organisms contain an internal cytoplasmic compartment and a‬
‭nucleus within the cytoplasm.‬‭Cytosol‬‭, the jelly-like‬‭substance within the cell, provides the‬

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f‭ luid medium necessary for biochemical reactions. Eukaryotic cells, including all animal‬
‭cells, also contain various cellular organelles. An‬‭organelle‬‭(“little organ”) is one of several‬
‭different types of membrane-enclosed bodies in the cell, each performing a unique‬
‭function. Just as the various bodily organs work together in harmony to achieve all of a‬
‭human’s functions, the many different cellular organelles work together to keep the cell‬
‭healthy and perform all its essential functions. The organelles and cytosol, taken together,‬
‭compose the cell’s‬‭c ytoplasm‬‭.‬

‭Cytoplasm‬
‭ he cytoplasm is the entire region of a cell between the plasma membrane and the nuclear‬
T
‭envelope (a structure to be discussed shortly). It consists of organelles suspended in the‬
‭gel-like cytosol, the cytoskeleton, and various chemicals. Even though the cytoplasm‬
‭consists of 70 to 80 percent water, it has a semi-solid consistency from the proteins within‬
‭it. However, proteins are not the only organic molecules found in the cytoplasm. Glucose‬
‭and other simple sugars, polysaccharides, amino acids, nucleic acids, fatty acids, and‬
‭glycerol derivatives are also found there. Ions of sodium, potassium, calcium, and many‬
‭o ther elements are also dissolved in the cytoplasm. Many metabolic reactions, including‬
‭protein synthesis, take place in the cytoplasm.‬

‭Cytoskeleton‬
‭ uch like the bony skeleton structurally supports the human body, the cytoskeleton helps‬
M
‭the cells to maintain their structural integrity. The‬‭c ytoskeleton‬‭is a group of fibrous‬
‭proteins located in the cytoplasm that serves a variety of purposes: provides rigidity and‬
‭shape to the cell, facilitates cellular movement, anchors the nucleus and other organelles‬
‭in place, moves vesicles through the cell, pulls replicated chromosomes to the poles of a‬
‭dividing cell, and enable cells within multicellular organisms to move.‬

‭Nucleus‬
‭ he‬‭nucleus‬‭is a cell’s central organelle (review‬‭Figure 3.7‬‭) and typically is the most‬
T
‭prominent organelle (‬‭Figure 3.21‬‭). The nucleus (plural‬‭= nuclei) stores‬‭c hromatin‬‭(a‬
‭combination of DNA and proteins that form‬‭c hromosomes‬‭)‬‭in a gel-like substance called‬
‭the‬‭nucleoplasm‬‭. The‬‭nucleolus‬‭is a condensed region‬‭o f chromatin where ribosome‬
‭subunits are made. The boundary of the nucleus is called the‬‭nuclear envelope‬‭. It‬
‭consists of two phospholipid bilayers (inner and outer) and contains pores that control‬
‭the passage of ions, molecules, and RNA between the nucleoplasm and cytoplasm. The‬
‭nuclear envelope is continuous with the endoplasmic reticulum.‬

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‭ igure 3.21‬ ‭The nucleus and its associated components.‬‭(credit:‬‭OpenStax‬‭). A link to a‬

F
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Ribosomes‬
‭ ibosomes‬‭are the cellular structures responsible‬‭for protein synthesis (‬‭Figure 3.22‬‭).‬
R
‭When viewed through an electron microscope, ribosomes appear as “dots” on the‬
‭endoplasmic reticulum (rough ER) or single, tiny dots that float freely in the cytoplasm.‬
‭Electron microscopy has shown that ribosomes, which are large complexes of protein and‬
‭RNA, consist of two subunits, aptly called large and small. Ribosomes receive their‬
‭“orders” for protein synthesis from the nucleus, where the nitrogenous bases in a strand‬
‭o f DNA (i.e., A, T, G, and C) are used as a template to produce a type of RNA called‬
‭messenger RNA‬‭(‬‭mRNA‬‭). The mRNA travels to the ribosomes,‬‭which use the code‬
‭provided by the sequence of the bases (i.e., A, U, G, and C) in the mRNA to join a series of‬
‭corresponding amino acids to build a protein. Chapter 4 presents a detailed discussion of‬
‭protein synthesis and related topics.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 3.22‬ ‭(a) An illustration of ribosomes attached‬‭to the endoplasmic reticulum. (b)‬
F
‭An electron microscope view similar to the illustration. (credit: (a)‬ ‭Rlawson‬‭at‬‭English‬
‭Wikibooks‬‭;‬‭CC BY-SA 3.0‬ ‭(b)‬‭OpenStax‬‭;‬‭CC BY-SA 3.0‬‭)‬

‭Mitochondria‬
‭ itochondria‬‭(singular = mitochondrion) are often‬‭called the “powerhouses'' or “energy‬
M
‭factories'' of animal cells because they are responsible for making adenosine triphosphate‬
‭(ATP), the cell’s primary energy-carrying molecule.‬‭ATP‬‭represents the short-term stored‬
‭energy of the cell. Cellular respiration is the process of making ATP using the chemical‬
‭energy found in glucose and other nutrients. In mitochondria, this process uses oxygen‬
‭(obtained from the air you inhale) and produces carbon dioxide as a waste product, which‬
‭can be exhaled. Chapter 6 presents a detailed discussion of cellular respiration and‬
‭related topics, while Chapter 10 presents information about the respiratory system.‬
‭ itochondria are oval-shaped, double-membrane organelles (‬‭Figure 3.23‬‭) that have their‬
M
‭own ribosomes and DNA. Each membrane is a phospholipid bilayer embedded with‬
‭proteins. The inner layer has folds called cristae, which increases its surface area. The area‬
‭between the two membranes is called the intermembrane space.‬
‭ ells use ATP constantly, so the mitochondria are always at work. One of the organ‬
C
‭systems in the body that uses enormous amounts of ATP is the muscular system because‬
‭ATP is required to sustain muscle contraction. As a result, muscle cells are packed full of‬
‭mitochondria. Nerve cells, called neurons, also need large quantities of ATP to run their‬
‭sodium-potassium pumps. Therefore, an individual neuron will be loaded with over a‬
‭thousand mitochondria. On the other hand, a bone cell, which is not nearly as‬
‭metabolically active, might only have several hundred mitochondria.‬

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‭ igure 3.23‬ ‭The mitochondria are the cell's energy-conversion‬‭factories. (a) A‬

F
‭mitochondrion is composed of two separate phospholipid bilayer membranes. (b) An‬
‭electron micrograph (EM) of mitochondria. EM × 236,000. (credit: Micrograph provided‬
‭by the Regents of University of Michigan Medical School © 2012;‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Organelles of the Endomembrane System‬

‭ set of three major organelles within the cell forms the‬‭endomembrane system‬‭. These‬
A
‭o rganelles work together to perform various cellular jobs, including producing, packaging,‬
‭and exporting certain cellular products. The organelles of the endomembrane system‬
‭include the endoplasmic reticulum, Golgi apparatus, and vesicles.‬

‭Endoplasmic Reticulum‬
‭ he‬‭endoplasmic reticulum (ER)‬‭is a system of channels‬‭that is continuous with the‬
T
‭nuclear membrane (or “envelope”) covering the nucleus and composed of the same lipid‬
‭bilayer material. The ER provides passages throughout much of the cell that function in‬
‭transporting, synthesizing, and storing materials. The winding structure of the ER results‬
‭in a sizable membranous surface area that supports its many functions (‬‭Figure 3.24‬‭).‬
‭ ndoplasmic reticulum can exist in two forms: rough ER and smooth ER. These two types‬
E
‭o f ER perform some very different functions and can be found in very different amounts‬
‭depending on the type of cell. One of the main functions of the‬‭smooth ER‬‭(SER) is‬
‭synthesizing lipids. The SER synthesizes phospholipids, the main component of biological‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ embranes, and steroid hormones. For this reason, cells that produce large quantities of‬
m
‭hormones, such as those of the female ovaries and male testes, contain large amounts of‬
‭SER. In addition to lipid synthesis, the SER also sequesters (i.e., stores) and regulates the‬
‭concentration of cellular calcium ions (Ca‬‭++‬‭), a function‬‭crucial in nervous system cells‬
‭where Ca‬‭++‬ ‭triggers neurotransmitter release.‬
‭ ough ER‬‭(RER;‬‭Figure 3.24‬‭) is so-called because its‬‭membrane is dotted with ribosomes,‬
R
‭giving the RER a bumpy or studded appearance. The RER functions include synthesizing‬
‭and modifying proteins destined for the cell membrane or exported from the cell.‬
‭Typically, a protein is synthesized within the ribosome and released inside the channel of‬
‭the RER, where sugars can be added to it before it is transported within a vesicle to the‬
‭next stage in the packaging and shipping process: the Golgi apparatus.‬

‭ igure 3.24‬ ‭(a) The ER is a winding network of thin‬‭membranous sacs closely associated‬
F
‭with the cell nucleus. The smooth and rough endoplasmic reticula are very different in‬
‭appearance and function (source: mouse tissue). (b) Rough ER is studded with numerous‬
‭ribosomes (source: mouse tissue). EM × 110,000. (c) Smooth ER (source: mouse tissue).‬
‭EM × 110,510. (Micrographs provided by the Regents of University of Michigan Medical‬
‭School © 2012;‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭The Golgi Apparatus‬

‭ he‬‭Golgi apparatus‬‭is responsible for sorting, modifying,‬‭and shipping off the products‬
T
‭from the rough ER, much like a post office. The Golgi apparatus looks like stacked‬
‭flattened discs, almost like stacks of oddly shaped pancakes. Like the ER, these discs are‬
‭membranous. The Golgi apparatus has two distinct sides, each with a different role. One‬
‭side of the apparatus (i.e.,‬‭cis‬‭face) receives products‬‭in vesicles. These products are‬
‭sorted through the apparatus, chemically modified, and then released from the opposite‬
‭side (i.e.,‬‭trans‬‭face) after being repackaged into‬‭new vesicles. If the product is to be‬
‭exported from the cell, the vesicle migrates to the cell surface, fuses to the cell membrane,‬
‭and the cargo is secreted via exocytosis (‬‭Figure 3.25‬‭).‬ ‭Some of the protein products the‬
‭Golgi packages include digestive enzymes meant to remain inside the cell to break down‬
‭certain materials. The enzyme-containing vesicles released by the Golgi are called‬
‭lysosomes,‬‭and the enzymes function to break down‬‭and digest unneeded cellular‬
‭components, such as a damaged organelle.‬

‭ igure 3.25‬ ‭(a) The Golgi apparatus manipulates‬‭products from the rough ER and also‬
F
‭produces new organelles called lysosomes. Proteins and other ER products are sent to the‬
‭Golgi apparatus, which organizes, modifies, packages, and tags them. Some of these‬
‭products are transported to other cell areas, and some are exported from the cell through‬
‭exocytosis. (b) An electron micrograph of the Golgi apparatus. (credit:‬‭OpenStax‬‭). A link to‬
‭a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭97‬
‭Chapter 3: Cells and Associated Topics‬

‭ ow that you have learned about cell structure click the link below or scan the QR code to‬
N
‭watch a video titled “The Inner Life of the Cell” by Harvard and HHMI. Dr. Chace Tydell, a‬
‭research scientist and instructor, provides a helpful narration. This video presents a‬
‭fascinating and more realistic examination of various aspects of cells.‬

The Inner Life of the Cell by Harvard and HHMI narrated by Tydell

‭3.7 Career Connection - Cytotechnologist‬

‭ ave you ever heard of a medical test called a Pap smear? In this test, a doctor takes a‬
H
‭small sample of cells from a patient's uterine cervix and sends it to a medical lab, where a‬
‭cytotechnologist stains the cells and examines them for any changes that could indicate‬
‭abnormal cell growth or a microbial infection (‬‭Figure‬‭3.26‬‭).‬

‭ ytotechnologists (cyto—= “cell”) are professionals who study cells via microscopic‬
C
‭examinations and other laboratory tests. They are trained to determine which cellular‬
‭changes are within normal limits. Their focus is not limited to cervical cells; they study‬
‭cellular specimens from all organs. When they notice abnormalities, they consult a‬
‭pathologist, a medical doctor who can make a clinical diagnosis.‬

‭ igure 3.26‬ ‭These‬‭uterine cervix cells, viewed through a light microscope, were obtained‬
F
‭from a Pap smear. Normal cells are on the left. The cells on the right are infected with‬
‭human papillomavirus (HPV). Notice that the infected cells are larger; two of these cells‬
‭each have two nuclei instead of one, the usual number. (credit: modification of work by Ed‬
‭Uthman, MD; scale-bar data from Matt Russell;‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭98‬
‭Chapter 4: DNA, Genes, RNA, and Protein Synthesis‬

‭ igure 4.1‬ ‭The central dogma of molecular biology‬‭shows the flow of information from‬
F
‭DNA to RNA (via transcription) and RNA to protein (via translation). Organisms with a‬
‭reverse transcriptase enzyme can use RNA to produce complementary DNA. (credit:‬
‭modified from‬‭Central_dogma_of_molecular_biology.svg‬‭;‬‭Philippe Hupé ;‬‭CC BY-SA 3.0‬‭)‬

‭4.1 Introduction‬
‭ oday, the three letters of the acronym “DNA” have become synonymous with crime‬
T
‭solving, paternity testing, human identification, and genetic testing. These procedures are‬
‭possible because of the mid-twentieth-century discovery that DNA is the genetic material.‬
‭The results of several classic experiments set the stage for an explosion of our knowledge‬
‭about DNA and how it stores and transmits genetic information.‬
‭ NA was first isolated from white blood cells by Friedrich Miescher in the 1860s. Over‬
D
‭fifty years later, Frederick Griffith’s work with strains of the bacterium‬‭Streptococcus‬
‭pneumoniae‬‭provided the first clue that DNA and not‬‭protein (as others argued) is the‬
‭universal molecule of heredity. Griffith’s conclusions were later supported by additional‬
‭bacterial research by Oswald Avery, Colin MacLeod, and Maclyn McCarty and virus‬
‭research by Alfred Hershey and Martha Chase. A short time later, Erwin Chargaff‬
‭determined DNA's adenine, thymine, cytosine, and guanine ratios suggested paired‬
‭relationships (%A = %T and %C = %G). He also found that the percentages of A, T, C, and‬
‭G differ for different species.‬
S‭ ince the rediscovery in 1900 of Gregor Mendel’s work with the inheritance of traits in‬
‭pea plants, the definition of the gene has progressed from an abstract unit of heredity to a‬

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t‭ angible molecular entity capable of replication, expression, and mutation. Genes are‬
‭composed of DNA and are linearly arranged on chromosomes. Genes specify the‬
‭sequences of amino acids, which are the building blocks of proteins. In turn, proteins are‬
‭responsible for orchestrating nearly every cell function. Both genes and the proteins they‬
‭encode are essential to life as we know it.‬

‭4.2 Organization of the Nucleus and its DNA‬

‭ s discussed in Chapter 3, the‬‭nucleus‬‭is the largest‬‭and most prominent‬‭organelle‬‭in‬
A
‭eukaryotic‬‭cells (‬‭Figure 4.2‬‭). The nucleus is generally‬‭considered the cell's control center‬
‭because it stores the genetic instructions (via DNA) for everything a cell will do and all of‬
‭the products it will make. Each cell in your body except for gametes (i.e., sperm and ova)‬
‭contains the complete set of your DNA. Before a cell divides, the DNA must be duplicated‬
‭so that each new cell receives a full complement of DNA.‬
S‭ ome cells in the body, like muscle cells, contain more than one nucleus (‬‭Figure 4.3‬‭),‬
‭which is known as multinucleated. Other cells, like mammalian red blood cells (RBCs), do‬
‭not contain nuclei. RBCs eject their nuclei as they mature, allowing hemoglobin molecules‬
‭to carry oxygen throughout the body (‬‭Figure 4.4‬‭).‬‭Without nuclei, the life span of RBCs is‬
‭short (approximately 115 days), so the body must constantly produce new ones.‬

‭ igure 4.2‬ ‭The nucleus is the control center of‬‭the cell. The nucleus of living cells‬
F
‭contains the genetic material that determines the entire structure and function of that cell.‬
‭(credit:‬‭OpenStax‬‭)‬

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‭ igure 4.3‬ ‭Skeletal muscle cells, unlike cardiac‬‭and smooth muscle cells, contain many‬
F
‭nuclei and are “multinucleated.” These muscle cells are long and fibrous (often called‬
‭muscle fibers). During development, many smaller cells fuse to form a mature muscle‬
‭fiber. The nuclei of the fused cells are conserved in the mature cell, thus imparting a‬
‭multinucleate characteristic to mature muscle cells. LM × 104.3. (credit: Micrograph‬
‭provided by the Regents of University of Michigan Medical School © 2012;‬‭OpenStax‬‭)‬

‭ igure 4.4‬ ‭When erythroblasts transform into mature‬‭red blood cells, their nuclei are‬
F
‭extruded to make room for more hemoglobin. Therefore, mature red blood cells lack‬
‭DNA, so proteins can’t be made to repair themselves or for other functions. The two‬
‭panels here show an erythroblast before and after ejecting its nucleus. (credit:‬
‭modification of micrograph provided by the Regents of University of Michigan Medical‬
‭School © 2012;‬‭OpenStax‬‭)‬

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‭ ike most other cellular organelles, the nucleus is surrounded by a nuclear envelope‬
L
‭membrane. This membranous covering consists of two adjacent lipid bilayers with a thin‬
‭fluid space between them. Spanning these two bilayers are nuclear pores. A‬‭nuclear pore‬
‭is a tiny passageway proteins, RNA, and solutes use to cross between the nucleus and the‬
‭cytoplasm. Proteins called pore complexes line the nuclear pores and regulate the‬
‭transportation of materials into and out of the nucleus (‬‭Figure 4.2‬‭).‬
I‭ nside the nuclear envelope is a gel-like‬‭nucleoplasm‬‭with solutes that include the‬
‭building blocks of nucleic acids (i.e., nucleotides). There may also be a dark-staining mass,‬
‭o ften visible under a simple light microscope, called a nucleolus (plural = nucleoli). The‬
‭nucleolus‬‭is a nucleus region responsible for manufacturing‬‭the RNA necessary to‬
‭construct ribosomes. Once synthesized, newly made ribosomal subunits exit the cell’s‬
‭nucleus through the nuclear pores (‬‭Figure 4.2‬‭).‬
‭ ithin the nucleus are threads of‬‭c hromatin‬‭composed‬‭o f DNA and associated proteins‬
W
‭(‬‭Figure 4.5‬‭). Along the chromatin threads, the DNA‬‭is wrapped around a complex of‬
‭histone proteins‬‭. A‬‭nucleosome‬‭is a single, wrapped‬‭DNA-histone protein complex.‬
‭Multiple nucleosomes along the entire molecule of DNA appear like a beaded necklace, in‬
‭which the string is the DNA and the beads are the associated histones. When a cell divides‬
‭(i.e., mitosis or meiosis), the chromatin condenses into chromosomes to safely transport‬
‭the DNA to the daughter cells.‬
‭ or additional information about DNA and chromosomes, click the link below to watch‬
F
‭the TED-Ed video by‬‭Joe Hanson titled “DNA: The Book‬‭o f You.”‬

DNA: The book of you - Joe Hanson

I‭ n the 1950s, Francis Crick and James Watson worked together at the University of‬
‭Cambridge, England, to determine the structure of DNA (‬‭Figure 4.6‬‭). Maurice Wilkins also‬
‭actively explored this field at King's College at the University of London. In Wilkins’ lab,‬
‭another scientist, Rosalind Franklin, used X-ray diffraction methods to understand DNA‬
‭structure (‬‭Figure 4.7‬‭). Watson and Crick were able‬‭to piece together the puzzle of the‬
‭DNA molecule based on Franklin's data because Crick had also studied this technique.‬
‭Their publication,‬‭A Structure for Deoxyribose Nucleic‬‭Acid‬‭, was published in April 1953.‬

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‭ igure 4.5‬ ‭A DNA double helix comprising two strands‬‭o f complementary bases wraps‬
F
‭around histone protein complexes to form chromatin. Continued bundling and‬
‭condensing of the chromatin produces the fully condensed chromosome in preparation‬
‭for cell division (mitosis or meiosis). (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭ igure 4.6‬ ‭The work of pioneering scientists (a)‬‭James Watson, Francis Crick, and Maclyn‬
F
‭McCarty led to our present-day understanding of DNA. (b) the X-ray diffraction pattern of‬
‭DNA obtained by Rosalind Franklin elucidated its double helix structure. (credit a:‬
‭modification of work by Marjorie McCarty, Public Library of Science;‬‭OpenStax‬‭)‬

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‭(a) (b)‬
‭ igure 4.7‬ ‭(a) Rosalind Elsie Franklin was an English‬‭chemist and X-ray crystallographer‬
F
‭whose work was central to the understanding of the molecular structures of DNA‬
‭(deoxyribonucleic acid), RNA (ribonucleic acid), viruses, coal, and graphite.‬
‭(b) Experimental setup used by Franklin to create Photo 51. The sample consisted of a‬
‭DNA strand stretched across a paperclip and mounted on a piece of cork. X-rays were sent‬
‭through the DNA strand, and the diffracted paths were captured on light-sensitive paper‬
‭to create Photo 51. The "X" in the center of Photo 51 is due to the helical arrangement of‬
‭the DNA molecules in the sample‬‭.‬‭(credit: (a)‬‭Rosalind-franklin-in-paris.jpg‬‭;‬‭CSHL‬‭;‬‭CC‬
‭BY-SA 4.0‬ ‭(‬‭b)‬‭Experimental setup of Photo 51.svg‬‭MagentaGreen‬‭;‬‭CC BY-SA 2.0‬‭). A link to‬
‭a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

I‭ n 1962, James Watson, Francis Crick, and Maurice Wilkins were awarded the Nobel Prize‬
‭in Medicine. Unfortunately, in 1958, Franklin died of ovarian cancer, possibly caused by‬
‭her extensive use of X-rays in her research. Nobel prizes are not awarded posthumously,‬
‭and no more than three recipients can receive a single prize. If Franklin had been alive,‬
‭there is some debate about whether she or Wilkins should have been the third recipient.‬
‭ or additional information about the discovery of DNA’s structure, click the link below to‬
F
‭watch the TED-Ed video by‬‭Clá udio L. Guerra titled‬‭“Rosalind Franklin: DNA’s unsung‬
‭hero.”‬

Rosalind Franklin: DNA's unsung hero - Clá udio L. Guer…

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‭ atson and Crick proposed that DNA consists of two strands that “complement” each‬
W
‭o ther because the molecules that compose the strands fit together and bind to each other,‬
‭creating a double-stranded molecule that looks much like a long, twisted ladder. Each side‬
‭rail of the DNA ladder is composed of alternating sugar and phosphate groups (‬‭Figure‬
‭4.8‬‭). The two sides of the ladder are not identical but are‬‭complementary‬‭. These two‬
‭backbones bond to each other across pairs of protruding bases, with each bonded pair‬
‭forming one “rung,” or cross member of the ladder. The four DNA bases are‬‭adenine (A)‬‭,‬
‭thymine (T)‬‭,‬‭c ytosine (C)‬‭, and‬‭guanine (G)‬‭. Because‬‭o f their shape and charge, the two‬
‭bases that compose a pair always bond together. Adenine always binds with thymine, and‬
‭guanine always binds with cytosine (A to T; G to C).‬

‭(a)‬ ‭(b)‬
‭ igure 4.8‬ ‭Two diagrammatic representations of a‬‭double-stranded DNA molecule with‬
F
‭the sugar-phosphate backbones and hydrogen bonds between the complementary bases‬
‭o f both backbones in the middle. The first figure (a) includes the helical twisting of the‬
‭two strands. The second one identifies a nucleotide, the monomer unit of DNA consisting‬
‭o f deoxyribose (sugar), a phosphate group, and a nitrogen-containing base. (b)‬
‭Nucleotides are joined via dehydration synthesis reactions to create a polynucleotide‬
‭strand with a sugar-phosphate backbone. (credit: (a)‬‭DNA-structure-and-bases.png‬‭;‬
‭Public domain;‬‭CC0 1.0‬ ‭(b)‬‭DNA molecular structure,‬‭showing individual nucleotides and‬
‭bonds.jpg‬‭;‬‭Francescakb‬‭;‬‭CC BY-SA 4.0‬‭). A link to‬‭a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

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‭ round this same time, Austrian biochemist Erwin Chargaff examined the content of DNA‬
A
‭in different species and found that the amounts of adenine, thymine, guanine, and cytosine‬
‭were not found in equal quantities and that they varied from species to species, but not‬
‭between individuals of the same species. He found that the amount of adenine equals the‬
‭amount of thymine, and the amount of cytosine equals the amount of guanine. This‬
‭o bservation is also known as‬‭Chargaff’s rules‬‭. This‬‭finding proved immensely useful‬
‭when Watson and Crick prepared to propose their DNA double helix model.‬

‭4.3 DNA Replication‬

‭ or an organism to grow, develop, and maintain health, cells must reproduce by dividing‬
F
‭to produce two new daughter cells, each with the full complement of DNA as found in the‬
‭o riginal cell. Billions of new cells are made in an adult human every day. A few cell types in‬
‭the body do not divide, including nerve cells, skeletal muscle fibers, and cardiac muscle‬
‭cells. The division time of different cell types varies. Epithelial cells of the skin and‬
‭gastrointestinal lining, for instance, divide very frequently to replace those constantly‬
‭rubbed off the surface by friction.‬
‭ NA replication‬‭is copying DNA that must occur before‬‭cell division. The elucidation of‬
D
‭the structure of the double helix provided a hint as to how DNA makes copies of itself. In‬
‭their 1953 paper, Watson and Crick penned an incredible understatement: "It has not‬
‭escaped our notice that the specific pairing we have postulated immediately suggests a‬
‭possible copying mechanism for the genetic material." With specific base pairs, the‬
‭sequence of one DNA strand can be predicted from its complement. For example, if one‬
‭strand has a region with the sequence AGTGCCT, then the sequence of the complementary‬
‭strand would be TCACGGA.‬
‭ herefore, the double-helix model suggested that the two strands of the double helix‬
T
‭separate during replication, and each strand serves as a template from which the new‬
‭complementary strand is copied. What needed to be clarified was how the replication took‬
‭place. After much debate and experimentation, two scientists in California, Matthew‬
‭Meselson and Franklin Stahl, deduced the general method of DNA replication, called‬
‭semi-conservative replication‬‭, in 1958 (‬‭Figure 4.9‬‭).‬
I‭ t is essential to realize the interrelationship between diagrams of duplicated‬
‭chromosomes (e.g.,‬‭Figure 4.4‬‭) and the duplicated‬‭DNA double helices (‬‭Figure 4.9‬‭).‬
‭Before DNA replication, each chromosome contains one DNA double helix. After DNA‬
‭replication, each chromosome has two identical double helices called‬‭sister chromatids‬‭.‬
‭During cell division (mitosis or meiosis), the sister chromatids will separate and be‬
‭distributed among the two daughter cells so that each one contains a complete set of DNA‬
‭instructions (‬‭Figure 4.10‬‭).‬

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‭(a)‬ ‭(b)‬

‭ igure 4.9‬ ‭Two diagrammatic representations of the‬‭semi-conservative model of DNA‬

F
‭replication. In both cases, the original double helix is opened, and each strand serves as a‬
‭template for producing a complementary strand. The two new daughter DNA molecules‬
‭contain one pre-existing and one newly synthesized strand. Thus, DNA replication is said‬
‭to be “semiconservative.” (credit: (a)‬‭Genomics Education‬‭Programme‬‭;‬‭CC BY 2.0‬ ‭(‬‭b)‬
‭Semiconservative DNA replication.png‬‭;‬‭Eunice Laurent‬‭;‬‭CC BY-SA 4.0‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

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‭ igure 4.10‬ ‭Appearance of a pair of homologous chromosomes‬‭before and after DNA‬

F
‭replication. Before DNA replication, each pair member consists of a single double-helix.‬
‭After replication, each pair member consists of two DNA double helices, each containing‬
‭o ne original strand and one newly synthesized strand, in a manner consistent with‬
‭semi-conservative DNA replication. (credit:‬‭Chromosome‬‭Terminology.png‬‭;‬
‭Christinelmiller‬‭;‬‭CC BY-SA 4.0‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ NA replication is a complex process that involves many components, and it occurs in‬
D
‭three stages:‬‭initiation‬‭,‬‭elongation‬‭, and‬‭termination‬‭:‬
S‭ tage 1: Initiation.‬‭The two complementary strands‬‭are separated, much like unzipping a‬
‭zipper. Special enzymes untwist and separate the two DNA strands by breaking hydrogen‬
‭bonds (i.e., denaturation).‬
S‭ tage 2: Elongation.‬‭Each strand becomes a template‬‭along which a new complementary‬
‭strand is built. The‬‭DNA polymerase‬‭enzyme brings‬‭in the correct bases to complement‬
‭the template strand and synthesize a new strand base by base. This growing strand‬
‭continues to be built until it fully complements the template strand.‬
S‭ tage 3: Termination.‬‭Once the two original strands‬‭are bound to their own finished,‬
‭complementary strands, DNA replication stops, and the two new identical DNA molecules‬
‭are complete.‬
I‭ t is important to note that in eukaryotic cells, DNA replication initiates at many points‬
‭along the DNA of a chromosome and extends bidirectionally. This method allows a‬
‭substantial amount of DNA to replicate relatively quickly so that cell division can proceed.‬

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‭ NA replication must occur precisely so that new cells in the body contain the same‬
D
‭genetic material as their parent cells. Mistakes made during DNA replication, such as‬
‭accidentally adding an inappropriate nucleotide, can render a gene’s product useless.‬
‭Fortunately, most errors are promptly corrected by the proofreading ability of DNA‬
‭polymerase itself or other repair mechanisms. Once the process of DNA replication is‬
‭complete, the cell is ready to divide. You will learn about cell division in Chapter 13.‬
‭ rrors during DNA replication are not the only reason why‬‭mutations‬‭, which are‬
E
‭variations in the nucleotide/base sequence, arise. Mutations also occur because of‬
‭damage to DNA. Such mutations may be of two types: induced or spontaneous.‬‭Induced‬
‭mutations‬‭result from exposure to chemicals, UV rays,‬‭x-rays, or other environmental‬
‭agents.‬‭Spontaneous mutations‬‭o ccur without any exposure‬‭to any environmental agent‬
‭and result from natural reactions within the body.‬
‭ fter a discussion of protein synthesis, the topic of mutations will be revisited so that their‬
A
‭impacts on proteins can be more easily understood.‬

‭4.4 From DNA to Protein (Transcription and Translation)‬

‭ s was stated earlier, DNA provides a “blueprint” for cell structure and physiology. This‬
A
‭term means that DNA contains the information necessary for the cell to build one‬
‭essential molecule type: protein. Most structural components of the cell are made up, at‬
‭least in part, of proteins, and virtually all the functions that a cell carries out are‬
‭completed with the help of proteins. One of the most important classes of proteins is‬
‭enzymes‬‭, which help speed up necessary chemical reactions‬‭inside the cell. These critical‬
‭chemical reactions include building larger molecules from smaller components (such as‬
‭during DNA replication) and breaking down larger molecules into smaller ones (such as‬
‭when harvesting chemical energy from nutrient molecules, such as carbohydrates).‬
‭ he synthesis of proteins consumes more of a cell’s energy than any other metabolic‬
T
‭process. In turn, proteins account for more mass than any other component of living‬
‭o rganisms (except for water).‬
‭ rotein synthesis begins with‬‭genes‬‭, which are functional‬‭segments of DNA that provide‬
P
‭the genetic information necessary to make a product (RNA or protein, depending on the‬
‭gene).‬ ‭Structural genes‬‭provide the codes to construct‬‭proteins via a two-step process‬
‭called‬‭gene expression‬‭, which transforms the information‬‭coded in genes into proteins,‬
‭with an mRNA (i.e., messenger RNA) molecule as the intermediate. The first step of the‬
‭process is transcription, and the second is translation. This flow of information in cells‬
‭from DNA to mRNA (via transcription) and mRNA to protein (via translation) is called the‬
‭central dogma of molecular biology‬‭(‬‭Figure 4.11‬‭).‬ ‭Let’s look at each of these two steps‬
‭in more detail.‬

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‭ igure 4.11‬ ‭This figure illustrates the central dogma‬‭o f molecular genetics. Regions of‬
F
‭DNA, called genes, hold all of the genetic information necessary to build a cell’s proteins.‬
‭The nucleotide sequence (i.e., order of A, T, G, and C bases) of a gene is encoded into‬
‭mRNA (i.e., transcription), which is then decoded (i.e., translation) into an amino acid‬
‭sequence of the gene’s corresponding protein. (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Transcription: DNA to RNA‬

‭ ranscription‬‭involves encoding the information stored‬‭in the sequence of bases on one‬
T
‭o f the two DNA strands (designated as the‬‭template‬‭strand‬‭) into a complementary RNA‬
‭strand. A primary enzyme associated with transcription is‬‭RNA polymerase (Figure‬
‭4.12)‬‭. As previously discussed, DNA polymerase uses‬‭DNA as a template to produce‬
‭complementary DNA, with the rules A to T and G to C. RNA polymerase uses DNA as a‬
‭template to produce complementary RNA. As was stated in Chapter 2, RNA contains the‬
‭base‬‭uracil (U)‬‭instead of T.‬

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‭ reasonable question about this difference is, “Why?” One hypothesis is that because‬
A
‭uracil is less stable than thymine, RNA doesn’t persist as long as DNA in the cell. A‬
‭subsequent question is, “Why is this difference advantageous?”. Remember, RNA is used‬
‭for protein synthesis. Therefore, if a particular protein isn’t needed, having the associated‬
‭RNA persisting wouldn’t be necessary or desirable from a resource conservation‬
‭perspective.‬
I‭ n addition to mRNA, two other types of RNA produced via the transcription of specific‬
‭genes are‬‭ribosomal RNA‬‭(‬‭rRNA‬‭) and‬‭transfer RNA‬‭(‬‭tRNA‬‭;‬‭Figure 4.13‬‭). Ribosomes,‬
‭discussed in Chapter 3, function to produce proteins and contain rRNA as part of their‬
‭structure. tRNAs carry amino acids to the ribosomes for use in protein synthesis.‬
‭Transcription of a gene produces one type of RNA; in other words, a single gene doesn’t‬
‭yield two or three different categories of RNA.‬

‭ igure 4.12‬‭The process of transcription. RNA polymerase‬‭uses the template DNA strand‬
F
‭to produce a complementary, single-stranded RNA. (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

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‭Chapter 4: DNA and Gene Expression‬

‭ igure 4.13‬ ‭A simplified diagram of three types of‬‭RNA (m, t, and r) with the bases (A, C,‬
F
‭G, and U) symbolized as colored bars (credit:‬‭Types‬‭o f RNA‬‭;‬‭Christinelmiller‬‭;‬‭CC BY-SA 4.0‬‭)‬

‭ efore mRNA molecules leave the nucleus and proceed to protein synthesis, they are‬
B
‭modified in several ways. For this reason, it is often called a‬‭pre-mRNA‬‭at this stage. For‬
‭example, your DNA, and thus complementary mRNA, contains long regions called‬
‭non-coding regions‬‭that do not code for amino acids.‬‭Their function is still not completely‬
‭understood, but the process called‬‭splicing‬‭removes‬‭these non-coding regions from the‬
‭pre-mRNA transcript (‬‭Figure 4.14‬‭). A‬‭spliceosome‬‭—a‬‭structure composed of various‬
‭proteins and other molecules—attaches to the mRNA and “splices” or cuts out the‬
‭non-coding regions. The removed segment of the transcript is called an‬‭intron‬‭o r‬
‭intervening sequence. The remaining sequences are called‬‭exons‬‭o r expressed sequences.‬
‭ ther modifications (‬‭5’ cap‬‭and‬‭3’ polyA tail‬‭) are‬‭made to the ends of the mRNA, which‬
O
‭increase stability and facilitate binding with ribosomes. When modifications are complete,‬
‭the remaining molecule is called the mature mRNA. It moves out of the nucleus via the‬
‭nuclear pores and into the cytoplasm for translation.‬

‭Translation: mRNA to Protein‬

‭ ike translating a book from one language into another, the codons on a strand of mRNA‬
L
‭must be interpreted (via‬‭translation‬‭) into the amino‬‭acid alphabet of proteins. The amino‬
‭acids are covalently attached by‬‭peptide bonds‬‭formed‬‭via dehydration synthesis‬
‭reactions, which are catalyzed by ribosomes. The genetic code is a‬‭triplet code‬‭, with each‬
‭RNA‬‭codon‬‭consisting of three consecutive nucleotides‬‭that specify one amino acid or the‬
‭release of the newly formed polypeptide chain. As there are four RNA bases (A, U, G, and‬
‭C) and three bases in a codon, there are 64 possible codons (1/4 x 1/4 x 1/4 = 1/64).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 4.14‬ ‭Three post-transcriptional modifications‬‭to the pre-mRNA‬‭:‬ ‭1) Addition of a‬

F
‭modified guanine nucleotide to the 5’ end, called the 5’ cap; 2) Addition of approximately‬
‭100 to 200 adenine nucleotides to the 3’ end, via an enzyme called poly-A polymerase; 3)‬
‭Remove introns (noncoding regions) and reconnect the exons (coding regions), via the‬
‭spliceosome. These changes produce the mature mRNA that is ready for export from the‬
‭nucleus to the cytoplasm. (credit:‬‭Post-transcriptional‬‭modification of pre-mRNA.png‬‭;‬
‭Kep17‬‭;‬‭CC BY-SA 4.0‬‭).‬ ‭A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

S‭ ixty-one codons specify one of the 20 amino acids, while the remaining three (UAA, UAG,‬
‭and UGA) terminate translation (‬‭Figure 4.15‬‭). While‬‭this figure shows only the three-letter‬
‭abbreviations of amino acids,‬‭Figure 4.16‬‭provides‬‭the complete names.‬

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‭ igure 4.15‬ ‭This figure shows the genetic code for‬‭translating each nucleotide triplet in‬
F
‭mRNA into an amino acid or a termination signal in a protein. (credit: modification of work‬
‭by NIH;‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ igure 4.16‬ ‭Table of abbreviations of amino acids.‬ ‭(credit:‬‭Genomics Education‬

F
‭Programme‬‭;‬‭Flickr;‬‭CC BY 2.0‬‭)‬

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‭ s there are 61 codons that specify amino acids and only 20 amino acids, the genetic code‬
A
‭is said to be‬‭degenerate‬‭; that is, some amino acids‬‭are specified by more than one codon,‬
‭like synonyms you study in your English class (different word, same meaning).‬
‭ or example, CCU, CCC, CCA, and CCG are all codons for proline. Why would degeneracy be‬
F
‭advantageous? If structural genes are mutated, the mRNA would also contain mutations,‬
‭which could specify different amino acids and negatively affect the resulting proteins.‬
‭However, because the code is degenerate, different codons can still result in the same‬
‭amino acid sequence.‬
‭ ranslation also consists of three main stages: initiation, elongation, and termination.‬
T
‭Initiation occurs when a ribosome binds to an mRNA transcript and properly aligns itself‬
‭with the‬‭start codon‬‭(AUG). The elongation stage involves‬‭the recognition of a‬‭tRNA‬
‭anticodon‬‭with the next mRNA codon in the sequence.‬‭Once the anticodon and codon‬
‭sequences are bound (remember, they are complementary base pairs), the tRNA presents‬
‭its amino acid cargo. A peptide bond attaches the growing polypeptide strand to this next‬
‭amino acid. The tRNA molecule then releases the mRNA strand, the mRNA strand shifts‬
‭o ne codon over in the ribosome, and the next appropriate tRNA arrives with its matching‬
‭anticodon. This process continues until a‬‭stop codon‬‭o n the mRNA (UAA, UAG, or UGA) is‬
‭reached, which provides a “stop” message that signals the termination of translation and‬
‭triggers the release of the complete, newly-synthesized protein (‬‭Figure 4.17‬‭).‬
‭ ommonly, several ribosomes will translate an mRNA molecule simultaneously, increasing‬
C
‭the efficiency of protein synthesis. A single ribosome might translate an mRNA molecule in‬
‭approximately one minute, so multiple ribosomes aboard a single transcript could‬
‭produce multiple times the number of the same protein in the same minute. A collection of‬
‭ribosomes translating a single mRNA strand is called a‬‭polysome‬‭o r‬‭polyribosome‬
‭(‬‭Figure 4.18‬‭).‬

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‭ igure 4.17‬ ‭During translation, the mRNA transcript‬‭is “read” by a functional complex‬
F
‭consisting of the ribosome and tRNA molecules. tRNAs bring the appropriate amino acids‬
‭to the growing polypeptide chain by matching their anticodons with codons on the mRNA‬
‭strand. (credit:‬‭OpenStax‬‭). There is an error in‬‭the last part of this figure. When the‬
‭ribosome moves to the next codon (CGA), the previous tRNA doesn’t shift over to the start‬
‭codon. In other words, the tRNA with the AAA anticodon will remain bound to the UUU‬
‭codon, and the tRNA with the GCU anticodon will bind to the CGA codon. A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 4.18‬ ‭This figure is a representation of a‬‭polysome or polyribosome. Numerous‬

F
‭ribosomes (two-part, brown structures) use the same mRNA (blue strand) at the same‬
‭time to produce a protein made of amino acids (colored balls). Note that the length of the‬
‭protein increases when the figure is viewed from left to right, as ribosomes further along‬
‭the mRNA add more amino acids. Also note the sequence of colors, starting with the first‬
‭amino acid (pink colored ball, which represents methionine), is the same. (credit:‬‭Multiple‬
‭Ribosomes Translation Protein Synthesis‬‭; Wikimedia‬‭Commons;‬‭CC BY-SA 4.0‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ he mRNA codons, not the‬‭tRNA anticodons, are used‬‭with the genetic code table to‬
T
‭determine the amino acids inserted (‬‭Figure 4.15‬‭).‬ ‭Also, note the start codon (AUG) is not‬
‭at the very front end of the mRNA; in other words, there is an mRNA sequence in front of‬
‭it that contains other sequence elements that help the small ribosomal subunit to identify‬
‭which AUG functions as the start codon.‬
‭ ow is a good time to click the link below or scan the QR code to watch a Khan Academy‬
N
‭video titled “DNA Replication, RNA Transcription, and Translation.” It summarizes many of‬
‭the concepts discussed in the chapter.‬

DNA replication and RNA transcription and translation | Khan Ac…

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Mutations and Their Impact on Proteins‬

‭ s discussed previously, changes to the base sequence of a structural gene can result in‬
A
‭changes to the amino acid sequence, which would likely impact the protein’s function.‬
‭Changes to a single base pair of DNA are called‬‭point‬‭mutations‬‭, and their impact varies.‬
‭Some point mutations don’t result in amino acid changes due to the degeneracy of the‬
‭genetic code and are, therefore, referred to as‬‭silent‬‭mutations‬‭. These changes usually‬
‭o ccur at the third base of a codon, which often represents the same amino acid as the‬
‭o riginal codon (‬‭Figure 4.15‬‭). Other point mutations,‬‭called‬‭missense mutations‬‭, result in‬
‭replacing one amino acid with another, which may alter the protein's function.‬‭Nonsense‬
‭mutations‬‭are point mutations that produce a stop‬‭codon (UAA, UAG, or UGA),‬
‭prematurely terminate protein synthesis, and likely substantially affect the resulting‬
‭protein’s function. Mutations can also result from adding a base, known as an‬‭insertion‬‭,‬
‭o r removing a base, also known as‬‭deletion‬‭. If an‬‭insertion or deletion alters the‬
‭translational reading frame (a‬‭frameshift mutation‬‭),‬‭the resulting protein is usually‬
‭nonfunctional (‬‭Figure 4.19‬‭).‬

‭ igure 4.19‬ ‭Types of point mutations to structural‬‭genes and their associated impacts on‬
F
‭the resulting proteins. (credit:‬‭OpenStax‬‭). A link‬‭to a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭4.5 Career Connection - Forensic Scientists and DNA Analysis‬

‭ NA evidence was used for the first time to solve an immigration case. The story begins‬
D
‭with a teenage boy returning to London from Ghana to be with his mother. Immigration‬
‭authorities at the airport were suspicious of him, thinking he was traveling on a forged‬
‭passport. After much persuasion, he was allowed to go live with his mother, but the‬
‭immigration authorities did not drop the case against him. Numerous types of evidence,‬
‭including photographs, were provided to the authorities, but deportation proceedings‬
‭were started. Around the same time, Dr. Alec Jeffreys of Leicester University in the United‬
‭Kingdom invented a technique known as DNA fingerprinting. The immigration authorities‬
‭approached Dr. Jeffreys for help. He took DNA samples from the mother and three of her‬
‭children, plus an unrelated mother, and compared the samples with the boy’s DNA.‬
‭Because the biological father was not in the picture, the DNA from the three children was‬
‭compared with the boy’s DNA. He found a match in the boy’s DNA for the mother and his‬
‭three siblings. He concluded that the boy was indeed the mother’s son.‬

‭ orensic scientists analyze many items, including documents, handwriting, firearms, and‬
F
‭biological samples. They analyze the DNA content of hair, semen, saliva, and blood and‬
‭compare it with a database of DNA profiles of known criminals. The analysis includes DNA‬
‭isolation, sequencing, and sequence analysis; most forensic DNA analysis involves‬
‭polymerase chain reaction (PCR) amplification of short tandem repeat (STR) loci and‬
‭electrophoresis to determine the length of the PCR-amplified fragment. Only mitochondrial‬
‭DNA is sequenced for forensics. Forensic scientists are expected to appear at court‬
‭hearings to present their findings. They are usually employed in crime labs of city and‬
‭state government agencies. Geneticists experimenting with DNA techniques work for‬
‭scientific and research organizations, pharmaceutical industries, and college and‬
‭university labs. Students wishing to pursue a career as a forensic scientist should have at‬
‭least a bachelor's degree in chemistry, biology, or physics and preferably some experience‬
‭working in a laboratory.‬

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‭ igure 5.1‬ ‭Eating may be one of the simple pleasures‬‭in life, but digesting even one apple‬
F
‭requires the coordinated work of numerous organs. The digestive processes also‬
‭demonstrate two critical biological principles: 1) integrated functioning, which is‬
‭cooperation between different organ systems, and 2) homeostasis via negative feedback‬
‭control. (credit: “Aimanness Photography”/Flickr;‬‭OpenStax‬‭)‬

‭5.1 Introduction‬
‭ he digestive system is continually at work, yet people seldom appreciate the complex‬
T
‭tasks it performs in a choreographed biological symphony. Consider what happens when‬
‭you eat an apple. Of course, you enjoy the apple’s taste as you chew it, but you overlook‬
‭your digestive system working in the following hours unless something goes amiss and‬
‭you get a stomachache. You may be taking a walk, studying, or sleeping, having forgotten‬
‭about the apple. Still, your stomach and intestines are busy digesting it and absorbing its‬
‭vitamins and other nutrients. Our natural foods consist primarily of proteins, fats, and‬
‭complex carbohydrates. We must convert these macromolecules (see Chapter 2) into‬
‭simple molecules that can be used for required functions, such as energy production and‬
‭storage, growth, and repair. Conversion of the food consumed to the nutrients needed for‬
‭proper functioning is a multi-step process involving two fundamental steps: digestion and‬
‭absorption. During digestion, food particles are broken down, physically and chemically,‬
‭into smaller components; later, they are moved from the gastrointestinal (GI) tract into the‬
‭blood or lymph (via absorption) to be transported throughout the body.‬
‭ s with all body systems, the digestive system does not work in isolation; it functions‬
A
‭cooperatively with the other body systems, referred to as‬‭integrated functioning‬‭.‬
‭Consider, for example, the interrelationship between the digestive and cardiovascular‬
‭systems. Arteries supply the digestive organs with oxygen and processed nutrients, and‬

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‭ eins absorb and transport most of the nutrients absorbed from the digestive tract. These‬
v
‭nutrients are essential to support the heart muscle and vascular tissue functions. The‬
‭interrelationship of the digestive and endocrine systems is also critical. Hormones‬
‭secreted by several endocrine glands and endocrine cells of the pancreas, the stomach,‬
‭and the small intestine contribute to controlling digestion and nutrient metabolism. In‬
‭turn, the digestive system provides the nutrients to fuel endocrine function.‬‭Figure‬‭5.2‬
‭lists examples of other organ systems that contribute to the functioning of the digestive‬
‭system.‬

‭Body system‬ ‭Benefits received by the digestive system‬

‭Cardiovascular‬ ‭ lood supplies digestive organs with oxygen‬

B
‭and processed nutrients.‬
‭Endocrine‬ ‭ ndocrine hormones help regulate secretion‬
E
‭in digestive glands and accessory organs.‬
‭Muscular‬ S‭ keletal muscles support and protect‬
‭abdominal organs; smooth muscle facilitates‬
‭food movement through the digestive tract‬
‭and physical digestion.‬
‭Nervous‬ ‭ eurons help regulate secretions and‬
N
‭muscle contractions in the digestive tract.‬
‭Respiratory‬ ‭ espiratory organs provide oxygen and‬
R
‭remove carbon dioxide.‬
‭Skeletal‬ ‭ ones help protect and support digestive‬
B
‭o rgans.‬

‭Figure 5.2‬ ‭Contributions of other organ systems‬‭to the digestive system‬

I‭ n short, whether you pay attention or not, the organs of the digestive system perform‬
‭their specific functions, allowing you to use the food you eat to keep you going. This‬
‭chapter examines these organs' structure and functions and explores the digestive‬
‭processes' mechanics and chemistry.‬
‭ lick the link below or scan the QR code to watch the TED-Ed video by Emma Bryce‬ ‭titled‬
C
‭“How your digestive system works.” Watch it again after reading the chapter; your‬
‭knowledge and understanding should improve.‬

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How your digestive system works - Emma Bryce

‭ efore we examine the specifics of the digestive system, let’s briefly discuss two other‬
B
‭relevant topics, homeostasis and tissues, which are foundational to our discussions of the‬
‭various organ systems.‬

‭5.2 Homeostasis‬
I‭ n Chapter 1,‬‭homeostasis‬‭is defined as maintaining‬‭internal consistency, one of the‬
‭fundamental characteristics common to living organisms. At this time, we need to learn a‬
‭more technical definition: maintaining‬‭dynamic equilibrium‬‭around a‬‭setpoint‬‭(i.e., a‬
‭defined value) while responding to changing conditions. At first glance, this definition‬
‭might seem somewhat overwhelming, but stop and think about it momentarily. The term‬
‭dynamic means changing; equilibrium refers to a state of balance or consistency, and the‬
‭term set point means a defined value for a particular factor. A specific example will help‬
‭you understand the definition. Your body temperature remains relatively constant,‬
‭around 37C or 98.6F, your set point value. If your temperature climbs above the set point,‬
‭you sweat to cool off; if your temperature drops below the set point, you shiver to warm‬
‭up. Therefore, we physiologically respond to changing conditions to maintain set point‬
‭conditions (i.e., dynamic equilibrium).‬
‭ nother example is that blood glucose levels remain relatively constant because when‬
A
‭they are high, the liver converts some of the excess to glycogen for storage. However,‬
‭when body cells require glucose, glycogen is broken down, and blood glucose levels‬
‭increase (‬‭Figure 5.3‬‭).‬
‭ egative feedback loops are the predominant mechanism for achieving homeostasis in‬
N
‭both body temperature and blood glucose. If the value exceeds the set point, physiological‬
‭action is taken to lower it. These loops result in slight fluctuations above and below the set‬
‭point.‬

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‭ igure 5.3‬ ‭Blood sugar homeostasis is maintained‬‭around a set point by the process of‬
F
‭negative feedback control. If blood sugar levels rise above or drop below the set point‬
‭value, the body responds in such a way as to correct it, and dynamic equilibrium results.‬
‭(credit:‬‭Homeostasis of blood sugar.png‬‭;‬‭Christinelmiller‬‭;‬‭CC0 1.0‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭5.3 Tissues‬
‭ he term‬‭tissue‬‭describes a group of cells found together‬‭in the body that are organized‬
T
‭into layers and function in a coordinated manner. Although there are many types of cells‬
‭in the human body, they are organized into four broad categories of tissues: epithelial,‬
‭connective, muscle, and nervous (‬‭Figure 5.4‬‭). Each‬‭o f these categories is characterized by‬
‭specific functions that contribute to the overall health and maintenance of the body. A‬
‭disruption of the structure is a sign of injury or disease. Such changes can be detected‬
‭through histology, the microscopic study of tissue appearance, organization, and function.‬
‭ pithelial tissue,‬‭also called‬‭epithelium‬‭, describes‬‭the sheets of cells that cover exterior‬
E
‭surfaces of the body (i.e., skin), line internal cavities and passageways, and form certain‬
‭glands. Epithelial tissues provide the body’s first line of protection from physical, chemical,‬
‭and biological wear and tear.‬

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‭ igure 5.4‬ ‭The four types of tissues are exemplified‬‭in nervous tissue, stratified‬
F
‭squamous epithelial tissue, cardiac muscle tissue, and connective tissue. (Micrographs‬
‭provided by the Regents of University of Michigan Medical School © 2012;‬‭OpenStax‬‭)‬

‭ ll substances that enter the body be they from outside the body or from an internal open‬
A
‭surface, such as the interior of the small intestine, must cross the epithelium. Many‬
‭epithelial cells are capable of secretion and release mucous and chemical compounds onto‬
‭their surfaces. For example, cells lining the respiratory tract secrete mucus that traps‬

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i‭ncoming microorganisms and particles; the epithelium of the small intestine releases‬
‭digestive enzymes.‬
‭ onnective tissues‬‭perform essential functions such‬‭as supporting and connecting other‬
C
‭tissues and protecting them. Examples include‬‭bones‬‭that support the body and protect‬
‭o rgans,‬‭tendons‬‭that attach muscles to bones, and‬‭ligaments‬‭that connect bones.‬
‭ lood‬‭, a specialized fluid connective tissue, contains cells that transport oxygen, initiate‬
B
‭clotting, and protect against bacterial and viral infections (‬‭Figure 5.5‬‭). The non-cellular‬
‭portion of blood, called‬‭plasma‬‭, transports nutrients,‬‭respiratory gases, wastes, and‬
‭hormones (e.g., insulin). Another example is‬‭adipose‬‭tissue‬‭, which consists primarily of‬
‭cells called adipocytes that store surplus energy in the form of fat (i.e., triglycerides) and‬
‭contribute to the insulation of the body and temperature homeostasis (‬‭Figure 5.6‬‭).‬

‭ igure 5.5‬ ‭Blood is a fluid connective tissue containing‬‭erythrocytes (red blood cells)‬
F
‭and various leukocytes (white blood cells) circulating in a liquid extracellular matrix called‬
‭plasma. LM × 1600. (Micrograph provided by the Regents of University of Michigan‬
‭Medical School © 2012;‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available‬
‭at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 5: Digestive System‬

‭ igure 5.6‬ ‭Adipose tissue consists of fat cells‬‭with little extracellular matrix. It stores fat‬
F
‭(i.e., triglycerides) for energy and provides insulation. LM × 800. (Micrograph provided by‬
‭the Regents of University of Michigan Medical School © 2012;‬‭OpenStax‬‭)‬

‭ uscle tissue‬‭responds to stimulation by contracting,‬‭which accomplishes different‬

M
‭functions depending on the specific type‬‭. Skeletal‬‭muscle‬‭is attached to bones, and its‬
‭contraction makes possible movement, facial expressions, posture, and other voluntary‬
‭(i.e., under conscious control) movements of the body. Forty percent of your body mass is‬
‭due to skeletal muscle.‬‭Smooth muscle‬‭contraction‬‭is responsible for involuntary‬
‭movements in the internal organs, such as the stomach, intestines, and arteries.‬ ‭Cardiac‬
‭muscle‬‭is found only in the heart, and its contraction‬‭is responsible for moving blood to‬
‭the lungs and the rest of the body (‬‭Figure 5.7‬‭).‬

‭ igure 5.7‬ ‭The three types of muscle tissue function‬‭to contract either under voluntary‬
F
‭(i.e., conscious) or involuntary control. (credit:‬‭Types Of Muscle ku.jpg‬‭;‬
‭scientificanimations.com‬‭;‬‭CC BY-SA 4.0‬‭). A link to‬‭a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ ervous tissue‬‭is composed primarily of‬‭neurons‬‭, which are cells specialized to receive‬
N
‭and transmit electrical impulses from specific areas of the body and send them to‬
‭particular locations to bring about changes, such as muscle contraction (‬‭Figure 5.8‬‭).‬

‭ igure 5.8‬ ‭Nervous tissue obtained from the spinal‬‭cord. The large cell at the center of‬
F
‭the picture is a neuron. The smaller cells, visible as dark dots throughout the tissue, are‬
‭neuroglia that support the neurons. (credit:‬‭Nervous‬‭Tissue: Spinal Cord Motor Neuron‬‭;‬
‭Berkshire Community College Bioscience Image Library‬‭;‬‭CC0 1.0‬‭)‬

‭5.4 Anatomy of the Digestive System‬

‭ he structure of the digestive system can be divided into two parts. The first one is the‬
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‭gastrointestinal (GI) tract‬‭, or alimentary canal,‬ ‭which includes the following regions: the‬
‭o ral cavity, pharynx, esophagus, stomach, small intestine (duodenum, jejunum, and ileum),‬
‭large intestine (cecum, ascending colon, transverse colon, descending colon, and sigmoid‬
‭colon), rectum and anus. The second part is the‬‭accessory organs‬‭, which are not part of‬
‭the GI tract itself but release a variety of secretions into it via ducts and include the‬
‭salivary glands, liver, gallbladder, and pancreas (‬‭Figure 5.9‬‭). We will begin by discussing‬
‭the regions of the GI tract.‬

‭Oral Cavity and Pharynx‬

‭ he‬‭oral cavity,‬‭o r‬‭mouth, is the point of entry of‬‭food into the digestive system via‬
T
‭ingestion‬‭(‬‭Figure 5.10a‬‭). The food consumed is broken‬‭into smaller particles by the‬
‭chewing action of the teeth, with the assistance of the tongue, which primarily contains‬
‭skeletal muscle under voluntary control.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ he chewing and wetting action provided by the teeth and saliva, a water secretion from‬
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‭the‬‭salivary glands‬‭(‬‭Figure 5.10b‬‭), prepare the food‬‭into a mass called the‬‭bolus‬‭for‬
‭swallowing. The tongue helps in swallowing—moving the bolus from the mouth into the‬
‭pharynx. The‬‭pharynx‬‭o pens to two passageways: the‬‭larynx (voice box), which leads to‬
‭the trachea and then the lungs, and the esophagus, which leads to the stomach. The larynx‬
‭has an opening called the‬‭glottis‬‭, which is covered‬‭by a cartilage flap called the‬‭epiglottis‬‭.‬
‭When swallowing, the epiglottis closes the glottis, and food passes into the esophagus, not‬
‭the larynx. If food enters the larynx (i.e., goes “down the wrong tube”), the body responds‬
‭by coughing to clear it. If the obstruction is complete and air can’t pass, a first aid‬
‭procedure called the‬‭Heimlich maneuver‬‭is used to‬‭resolve the blockage.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ ‬‭igure 5.9‬ ‭The organs associated with the digestive‬‭system play integral roles in the‬
F
‭life-sustaining digestion processes. (credit:‬‭OpenStax‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 5.10‬ ‭Food's physical and chemical digestion‬‭begins in the (a) oral cavity. Food is‬
F
‭chewed by teeth and moistened by saliva secreted from the (b) salivary glands. Enzymes in‬
‭the saliva begin to digest starches and fats. With the help of the tongue, the resulting bolus‬
‭moves into the esophagus by swallowing. (credit: modification of work by the National‬
‭Cancer Institute;‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭Esophagus‬
‭ he‬‭esophagus‬‭is a tube approximately ten inches long‬‭that connects the mouth to the‬
T
‭stomach. After being swallowed, the bolus passes through the esophagus. The‬‭smooth,‬
‭involuntary muscles of the esophagus undergo a series of wavelike movements called‬
‭peristalsis‬‭that push the food toward the stomach‬‭(‬‭Figure 5.11‬‭).‬
‭ ings of smooth muscle, called‬‭sphincters‬‭, form valves‬‭in the GI tract to regulate‬
R
‭movement from one region to the next. The gastroesophageal sphincter (a.k.a.‬‭c ardiac‬
‭sphincter‬‭) is located at the juncture of the esophagus‬‭and the stomach. This sphincter‬
‭o pens in response to swallowing and the pressure exerted by the food bolus, and the‬
‭bolus enters the stomach. When there is no swallowing action, this sphincter shuts,‬
‭preventing the stomach contents from traveling up the esophagus.‬‭Acid reflux‬‭o r‬
‭“heartburn” occurs when the acidic (i.e., low pH) digestive juices escape into the‬
‭esophagus and irritate it.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 5.11‬ ‭The esophagus moves food from the pharynx‬‭to the stomach through‬
F
‭involuntary, wavelike contractions of the smooth muscle called peristalsis. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ or additional information about acid reflux, click the link below to watch the TED-Ed‬
F
‭video by‬‭Rusha Modi titled “What causes heartburn?”.‬

What causes heartburn? - Rusha Modi

‭Stomach‬
‭ he‬‭stomach‬‭is a muscular organ about the size of‬‭your fist when empty. However, it can‬
T
‭expand to 20 times its empty size when filled with food. While other regions of the GI tract‬
‭have two layers of smooth muscle (longitudinal and circular), the stomach has a third‬
‭layer (oblique), which allows three-dimensional, powerful contractions to assist with‬
‭physical digestion (‬‭Figure 5.12‬‭). The bolus is transformed‬‭over two to six hours into a‬
‭soupy mixture of partially digested food and acidic secretions called‬‭c hyme‬‭.‬
‭ opular culture tends to refer to the stomach as the location where all digestion takes‬
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‭place. However, this is not true. A vital stomach function is to serve as a temporary holding‬
‭chamber. You can ingest a meal far more quickly than it can be digested and absorbed by‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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t‭ he small intestine. Thus, the stomach holds food and parses only small amounts into the‬
‭small intestine at a time via the‬‭pyloric sphincter‬‭.‬
‭ lthough you might think that the size of a person’s stomach is related to how much food‬
A
‭that individual consumes, body weight does not correlate with stomach size. Instead, when‬
‭you eat more food—such as at holiday dinner—you stretch the stomach more than when‬
‭you eat less.‬

‭ igure 5.12‬ ‭The stomach with various components‬‭labeled. The three layers of smooth‬
F
‭muscle allow the stomach to contact in three dimensions, making it very effective at‬
‭physical/mechanical digestion. The rugae are folds on the interior surface that function to‬
‭increase surface area as well as provide the ability to expand. (credit: modification of‬
‭image‬‭3D Medical Animation Muscular Layers of stomach.jpg‬‭;‬‭scientificanimations.com‬‭;‬‭CC‬
‭BY-SA 4.0‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭Small Intestine‬
‭ he‬‭small intestine‬‭is a tube-like organ, approximately‬‭10 to 15 feet long, with three‬
T
‭regions: duodenum, jejunum, and ileum‬‭(‬‭Figure 5.13‬‭).‬ ‭The‬‭duodenum‬‭receives chyme‬
‭from the stomach through the pyloric sphincter.‬
‭ s discussed later in the chapter, chyme mixes with various secretions that neutralize the‬
A
‭pH and continue chemical digestion along the small intestine. Undigested food is sent to‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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t‭ he colon from the ileum via peristaltic muscle movements. The‬‭ileocecal sphincter‬‭,‬
‭located at the juncture of the ileum and the cecum, regulates the movement of chyme from‬
‭the small intestine into the large intestine.‬
‭ he small intestine has a highly folded surface containing finger-like‬
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‭projections called the‬‭villi‬‭. The surface of each‬‭villus has many‬
‭microscopic projections called‬‭microvilli‬‭(‬‭Figure‬‭5.14‬‭). These‬
‭structures increase the intestine's surface area and increase the‬
‭absorption efficiency of the nutrients from the digested food into‬
‭the blood and lymphatic vessels on the other side.‬

‭ igure 5.13‬ ‭The three regions of the small intestine‬‭are the duodenum, jejunum, and‬
F
‭ileum. (credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ igure 5.14‬ ‭Villi are folds on the small intestine‬‭lining with additional folds at the edge‬
F
‭called microvilli. Both function to increase the surface area to facilitate the absorption of‬
‭nutrients into the blood or lymphatic vessels. (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Large Intestine‬
‭ he human‬‭large intestine‬‭is much smaller in length‬‭compared to the small intestine (5 ft.‬
T
‭vs. 10-15 ft.), so you might wonder why it is called “large.” Its name derives from its‬
‭relatively larger diameter compared to the small intestine (3 in. vs. 1 in.). The large‬
‭intestine has three parts: the‬‭cecum, the colon, and‬‭the rectum‬‭(Figure 5.15),‬‭and the‬
‭primary functions are to extract water and mineral salts from undigested food, synthesize‬
‭specific vitamins, form feces, and store waste material.‬ ‭The‬‭cecum‬‭joins the ileum to the‬
‭colon and is the receiving pouch for the waste matter.‬
‭ ttached to the cecum is the‬‭appendix‬‭, a structure‬‭thought to have been associated with‬
A
‭immune system function at one time and perhaps is no longer necessary. However, at least‬
‭o ne recent report postulates a survival advantage conferred by the appendix: In diarrheal‬
‭illness (e.g., dysentery), the appendix may serve as a bacterial reservoir to repopulate the‬
‭large intestine for those individuals surviving the initial phases of the illness.‬
‭ he‬‭colon‬‭is divided into four regions (ascending,‬‭transverse, descending, and sigmoid)‬
T
‭and is home to many bacteria or “intestinal flora” that aid in the digestive processes. For‬
‭additional information about these bacteria and how our diet impacts them, click the link‬
‭below to watch the TED-Ed video by‬‭Shilpa Ravella‬‭titled “How the food you eat affects‬
‭your gut.”‬

How the food you eat affects your gut - Shilpa Ravella

‭ he‬‭rectum‬‭is the terminal end of the large intestine,‬‭and its primary role is to store feces‬
T
‭until it moves out of the body via the‬‭anus‬‭.‬
‭ he residue of chyme that enters the large intestine contains few nutrients except water,‬
T
‭which is reabsorbed as the residue lingers in the large intestine, typically for 12 to 24‬
‭hours. Thus, it may not surprise you that the large intestine can be removed entirely‬
‭without significantly affecting digestive functioning. For example, in severe cases of‬
‭inflammatory bowel disease, the large intestine can be removed by a procedure known as‬
‭a colectomy. Often, a new fecal pouch can be crafted from the small intestine and sutured‬
‭to the anus; however, if not, an ileostomy can be created by bringing the distal ileum‬
‭through the abdominal wall, allowing collection of the watery chyme in a bag-like adhesive‬
‭appliance.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 5.15‬ ‭The large intestine reabsorbs water‬‭from undigested food and stores waste‬
F
‭material until elimination from the body via the anus (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Accessory Organs‬
‭ he organs discussed above are the organs of the digestive tract through which food‬
T
‭passes. Accessory organs, including the salivary glands (‬‭Figure 5.9b‬‭), liver, gallbladder,‬
‭and pancreas, add secretions to the GI tract via ducts.‬
‭ he‬‭liver‬‭is the largest internal organ in humans,‬‭and it plays a critical role in digesting fats‬
T
‭and detoxifying the blood. The liver produces‬‭bile‬‭,‬‭a digestive juice required to physically‬
‭break down fats into smaller droplets in the duodenum.‬
‭ he‬‭gallbladder‬‭is a small organ that aids the liver‬‭by storing and concentrating bile‬
T
‭(‬‭Figure 5.16‬‭). When chyme-containing fats enter the‬‭duodenum, the bile is secreted from‬
‭the gallbladder into the duodenum via the common bile duct.‬
I‭ f necessary, the gallbladder can be surgically removed. However, the liver will continue to‬
‭produce bile that is secreted into the small intestine via the common bile duct. The bile will‬
‭be less concentrated, so the efficiency of emulsification and subsequent enzyme digestion‬
‭may decrease.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ he‬‭pancreas‬‭produces several enzymes to digest proteins and carbohydrates (‬‭Figure‬

T
‭5.17‬‭). These secretions will travel via ducts that combine with the common bile duct to‬
‭release into the duodenum. Pancreatic secretions also contain‬‭bicarbonate‬‭, a base that‬
‭neutralizes the acidic chyme from the stomach, protects the small intestine from damage,‬
‭and creates an optimal pH environment for the intestinal digestive enzymes.‬

‭ igure 5.16‬ ‭Bile produced by the liver is transported‬‭via the hepatic and cystic ducts to‬
F
‭the gallbladder, where it is concentrated and stored. When needed, the gallbladder‬
‭contracts and forces bile back into the cystic duct, which then travels via the common bile‬
‭duct to be secreted into the duodenum. (credit:‬‭OpenStax‬‭).‬‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 5.17‬ ‭The pancreas has a head, a body, and‬‭a tail. The pancreatic islet cells make‬
F
‭hormones (insulin and glucagon) that regulate blood sugar homeostasis. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭5.5 Physical and Chemical Digestion‬

‭ utrient digestion can be defined as the processes associated with breaking down‬
N
‭macromolecules into smaller units that are absorbed. There are two broad categories of‬
‭digestion: physical and chemical.‬ ‭Physical digestion‬‭,‬‭also referred to as‬‭mechanical‬
‭digestion‬‭, is a process that decreases the size and‬‭increases the surface area of food‬
‭without altering its chemical composition.‬ ‭Chemical‬‭digestion‬‭, also called‬‭enzymatic‬
‭digestion‬‭, involves chemical reactions that change‬‭substances. Enzymes, typically‬
‭proteins, function to speed up (i.e., catalyze) these reactions, which, unlike physical‬
‭digestion, produce new substances with different properties. In other words, tearing a‬
‭piece of paper into small pieces (i.e., a physical change) doesn’t change the chemical‬
‭properties of the paper. However, burning the paper (i.e., a chemical change) does.‬
‭Examples of both types of digestion are discussed later in the chapter.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭5.6 Physical Digestion of Food‬

‭ hysical digestion begins in the mouth with chewing. It continues after swallowing with‬
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‭peristalsis, which consists of sequential, alternating waves of contraction and relaxation of‬
‭smooth muscle. These waves, which occur along the remainder of the GI tract, also play a‬
‭role in mixing food with digestive juices. Peristalsis is so powerful that foods and liquids‬
‭you swallow enter your stomach even if you are standing on your head.‬
‭ nother example of physical digestion involves the action of bile, which is produced by the‬
A
‭liver and stored and concentrated in the gallbladder. Recall that lipids are hydrophobic;‬
‭they do not dissolve in water. Thus, before they are digested in the watery environment of‬
‭the small intestine, large lipid droplets are physically broken down into smaller ones via a‬
‭process called‬‭emulsification‬‭. This change dramatically‬‭increases the surface area‬
‭available for lipid-digesting enzymes, discussed later in the chapter. It is important to note‬
‭that bile isn’t an enzyme; the fat droplets decrease in size but aren’t chemically changed.‬

‭5.7 Chemical Digestion of Food‬

‭ he extensive chemical digestion process begins in the mouth. As food is being chewed,‬
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‭saliva, produced by the salivary glands, mixes with the food. Saliva contains an enzyme‬
‭called‬‭salivary amylase‬‭that catalyzes the conversion‬‭o f dietary starches‬
‭(polysaccharides) into a disaccharide called maltose (Chapter 2). It also includes another‬
‭enzyme called‬‭lingual‬‭lipase‬‭, produced by tongue cells,‬‭that breaks down‬
‭fats/triglycerides. Food does not spend enough time in the mouth to allow all the‬
‭carbohydrates and fats to break down completely, so digestion will continue in the GI tract.‬
‭ nlike ruminants such as sheep and cattle, the human body does not produce enzymes‬
U
‭that can break down most fibrous polysaccharides, such as cellulose. While indigestible‬
‭polysaccharides do not provide nutritional value, they provide dietary fiber, which helps‬
‭propel food through the alimentary canal.‬
‭ o additional enzyme is secreted in the stomach that digests carbohydrates, but some‬
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‭gastric lipase is released. Salivary amylase and lingual lipase will continue catalyzing‬
‭reactions until these proteins are inactivated (i.e., denatured) by the low pH in the‬
‭stomach. Chemical digestion of proteins begins in the stomach via a secreted enzyme‬
‭called‬‭pepsin‬‭, which breaks peptide bonds between‬‭amino acids and cleaves proteins into‬
‭smaller polypeptides. Another cell type secretes hydrogen and chloride ions, which‬
‭combine to form‬‭hydrochloric acid (HCl)‬‭, the stomach‬‭juices' primary acidic component.‬
‭HCl helps to activate the pepsin and creates a highly acidic environment (pH 1.5 to 2.5)‬
‭that also kills many microorganisms in the food.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ hen digesting protein and some fats, pepsin protects the stomach lining from getting‬
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‭digested. Protection is achieved in two ways. First, pepsin is synthesized in an inactive‬
‭form called‬‭pepsinogen‬‭, which does not have the same‬‭enzyme functionality as pepsin.‬
‭Second, the stomach has a thick mucus lining that protects the underlying tissue from the‬
‭action of the digestive juices. Whenever pH levels drop too low, cells in the stomach react‬
‭by suspending HCl secretion and increasing mucous secretions, which is an example of‬
‭homeostasis via a negative feedback loop.‬
‭ s effective as the mucosal barrier is, it is not a “fail-safe” mechanism. Sometimes, gastric‬
A
‭juice eats away at the superficial lining of the stomach mucosa, creating erosions, which‬
‭mostly heal on their own. Deeper and more extensive erosions are called‬‭peptic ulcers‬‭.‬

‭ hy does the mucosal barrier break down? Several factors can interfere with its ability to‬
W
‭protect the stomach lining. Most ulcers are caused by excessive intake of non-steroidal‬
‭anti-inflammatory drugs (NSAIDs), including aspirin, or an infection caused by a type of‬
‭bacteria,‬‭Helicobacter pylori‬‭.‬

‭ ntacids help relieve symptoms of ulcers, such as “burning” pain and indigestion. When‬
A
‭ulcers are caused by NSAID use, switching to other classes of pain relievers allows healing.‬
‭When caused by‬‭H. pylori‬‭infection, antibiotics are‬‭effective. For additional information‬
‭about the discovery that bacteria can cause stomach ulcers, click the link below to watch‬
‭the TED-Ed video by‬‭Rusha Modi titled “The surprising‬‭cause of stomach ulcers.”‬

The surprising cause of stomach ulcers - Rusha Modi

‭ potential complication of ulcers is perforation, in which a hole is created in the stomach‬

A
‭wall, resulting in inflammation of the peritoneum. These ulcers must be surgically‬
‭repaired.‬

I‭ ts numerous digestive functions notwithstanding, only one stomach function is necessary‬

‭to life: the production of a substance called‬‭intrinsic‬‭factor‬‭. The intestinal absorption of‬
‭vitamin B‬‭12‬‭, essential for producing mature red blood‬‭cells and normal neurological‬
‭functioning, cannot occur without intrinsic factor. People who undergo total gastrectomy‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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(‭ stomach removal)—for life-threatening stomach cancer, for example—can survive with‬

‭minimal digestive dysfunction if they receive vitamin B‬‭12‬ ‭injections.‬
I‭ n the stomach, the bolus is transformed (via physical digestion and mixing with‬
‭secretions) into a semi-fluid mixture of partially digested food called‬‭c hyme‬‭. For optimal‬
‭digestion, chyme must be delivered slowly and in small amounts from the stomach. This‬
‭requirement is because chyme from the stomach is typically hypertonic. If large quantities‬
‭were forced all at once into the small intestine, the resulting osmotic water loss (Chapter‬
‭3) from the blood into the intestinal lumen would result in potentially life-threatening low‬
‭blood volume.‬
I‭ n the duodenum, chyme and pancreatic juices mix in an alkaline solution rich in‬
‭bicarbonate, which neutralizes the acidity of chyme and acts as a buffer to stabilize the pH.‬
‭Pancreatic juices also contain a variety of digestive enzymes, including amylase, trypsin,‬
‭nucleases, and lipase.‬‭Pancreatic‬‭amylase‬‭continues‬‭the breakdown of starch and‬
‭glycogen into maltose,‬‭trypsin‬‭continues the digestion‬‭o f polypeptides, and‬‭nucleases‬
‭digest the nucleic acids (DNA and RNA) in the cells we consume. As previously stated, bile‬
‭participates in the physical digestion of fats in a process called‬‭emulsification, where large‬
‭fat droplets (globules) are separated into smaller ones surrounded by bile salts (‬‭Figure‬
‭5.18‬‭). The increased surface area means‬‭pancreatic‬‭lipase‬‭can more efficiently digest‬
‭triglycerides. Even though lipase is released in the saliva and gastric juice, the pancreas is‬
‭the only significant source of this enzyme. Therefore, virtually all lipid digestion occurs in‬
‭the small intestine.‬
‭ he epithelium of the small intestine also produces enzymes called‬‭disaccharidases‬‭(i.e.,‬
T
‭maltase, sucrase, and lactase), which catalyze the hydrolysis of disaccharides (i.e., maltose,‬
‭sucrose, and lactose) into the component monosaccharides (i.e., glucose, fructose, and‬
‭galactose).‬ ‭Maltase‬‭breaks down maltose into glucose,‬‭sucrase‬‭breaks down sucrose (or‬
‭“table sugar”) into glucose and fructose, and‬‭lactase‬‭breaks down lactose (or “milk‬
‭sugar”) into glucose and galactose (‬‭Figure 5.19‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 5.18‬ ‭Emulsification of a fat globule into‬‭smaller fat droplets with more surface‬
F
‭area for efficient digestion by pancreatic lipase. (credit:‬ ‭Emulsification of a fat globule by‬
‭bile salts; Cenveo; pressbooks.ccconline.org;‬‭CC BY‬‭3.0‬‭).‬‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭ igure 5.19‬ ‭Enzymes (amylase, maltase, sucrase,‬‭and lactase) that facilitate carbohydrate‬
F
‭digestion. (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ actose intolerance‬‭is a condition characterized by indigestion caused by dairy products.‬

L
‭It occurs when the absorptive cells of the small intestine do not produce enough lactase,‬
‭the enzyme that digests the milk sugar lactose. In most mammals, lactose intolerance‬
‭increases with age. In contrast, some human populations, most notably Caucasians, can‬
‭maintain the ability to produce lactase as adults.‬

I‭ n people with lactose intolerance, the lactose in chyme is not digested. Bacteria in the‬
‭large intestine ferment the undigested lactose, producing gas. In addition to gas, symptoms‬
‭include abdominal cramps, bloating, and diarrhea. Symptom severity ranges from mild‬
‭discomfort to severe pain; however, symptoms resolve once the lactose is eliminated in‬
‭feces.‬

S‭ ome lactose-free dairy products are available in grocery stores (‬‭Figure 5.20‬‭). In addition,‬
‭dietary supplements (e.g., Lactaid) are available. Taken with food, they provide lactase to‬
‭help digest lactose.‬

‭ igure 5.20‬ ‭Lactaid is a brand of lactose-free milk.‬ ‭Other types of milk (e.g., almond, soy,‬
F
‭and coconut) don’t contain lactose. (credit:‬‭Milk‬‭Aisle‬‭;‬‭AshokaJegroo‬‭;‬‭CC BY-SA 3.0‬‭)‬

‭ lthough the glands of the large intestine secrete mucus, they do not secrete digestive‬
A
‭enzymes. Therefore, chemical digestion in the large intestine occurs exclusively because of‬
‭bacteria in the colon's lumen. Through fermentation, bacteria break down some of the‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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r‭ emaining carbohydrates. This results in the discharge of hydrogen, carbon dioxide, and‬
‭methane gasses that create flatus (gas) in the colon;‬‭flatulence‬‭(i.e., passing gas) is‬
‭excessive flatus. Up to 1500 mL of flatus is produced daily in the colon. More is produced‬
‭when you eat foods such as beans, which are rich in polysaccharides, such as fiber.‬
‭ or additional information about flatulence, click the link below to watch the TED-Ed video‬
F
‭by‬‭Purna Kashyap titled “Why do we pass gas?”.‬

Why do we pass gas? - Purna Kashyap

‭5.8 Integrated Functioning Between the Digestive and Endocrine Systems‬

‭ hat brings about the release of the enzymes previously discussed? In many cases, the‬
W
‭answer is hormones that cause glands to produce and secrete them. The‬‭endocrine‬
‭system‬‭controls the response of the various glands‬‭in the body and the release of‬
‭hormones at the appropriate times. Several important digestive components are under‬
‭hormonal control, which provides an example of integrated functioning. For example, the‬
‭hormone‬‭gastrin,‬‭which is secreted by cells in the‬‭stomach in response to the presence of‬
‭proteins, stimulates the release of hydrochloric acid. The lower pH is necessary to convert‬
‭pepsinogen to pepsin, the active form of the enzyme that hydrolyzes proteins into shorter‬
‭polypeptides. Also, the acidic pH denatures proteins, making them more accessible to‬
‭pepsin.‬
I‭ n the duodenum, a hormone called‬‭secretin‬‭stimulates‬‭the pancreas to produce an‬
‭alkaline bicarbonate solution and deliver it to the duodenum. Secretin acts in tandem with‬
‭another hormone called‬‭c holecystokinin‬‭(CCK). CCK‬‭stimulates the pancreas to produce‬
‭pancreatic juices and the gallbladder to release bile into the duodenum, which is‬
‭responsible for emulsification.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭5.9 Absorption‬
‭ igested food is only useful to the body if it enters the bloodstream and its nutrients are‬
D
‭used. Each day, the GI tract processes up to 10 liters of food, liquids, and GI secretions, yet‬
‭less than one liter enters the large intestine, meaning the majority are reabsorbed. Almost‬
‭all ingested food, 80 percent of electrolytes, and 90 percent of water are absorbed in the‬
‭small intestine (‬‭Figure 5.21‬‭).‬

‭ igure 5.21‬ ‭The‬‭digestive secretions and absorption‬‭o f water. Absorption is a complex‬

F
‭process in which nutrients from digested food are transported from the GI tract into the‬
‭circulatory and lymphatic systems. (credit:‬‭OpenStax‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ ost nutrients are also absorbed from the GI tract's‬‭lumen‬‭(internal opening) and into‬
M
‭the bloodstream through the epithelial cells that comprise the tissue layer adjacent to the‬
‭lumen. Lipids are absorbed into lymphatic vessels and transported to the bloodstream‬
‭just before blood returns to the heart. Although the entire small intestine is involved in‬
‭the absorption of water and lipids, most absorption of carbohydrates and proteins occurs‬
‭in the jejunum. Notably, bile salts and vitamin B‬‭12‬ ‭are absorbed in the ileum. When chyme‬
‭passes from the ileum into the large intestine, it is essentially indigestible food residue‬
‭(mainly plant fibers like cellulose), some water, and millions of bacteria.‬
‭ bsorption can occur by the four transport mechanisms introduced in Chapter 3: (1)‬
A
‭active transport, (2) passive diffusion, (3) facilitated diffusion, and (4) endocytosis. As you‬
‭will recall from Chapter 3, active transport refers to the movement of a substance across a‬
‭cell membrane, going from an area of lower concentration to an area of higher‬
‭concentration (up the concentration gradient). In this type of transport, proteins within‬
‭the cell membrane act as “pumps,” using cellular energy (ATP) to move the substance.‬
‭Passive diffusion refers to the movement of substances from an area of higher‬
‭concentration to an area of lower concentration. In comparison, facilitated diffusion refers‬
‭to the movement of substances from a higher concentration area to an area of lower‬
‭concentration using a carrier protein in the cell membrane. Finally, endocytosis is a‬
‭transportation process in which the cell membrane engulfs material. It requires energy,‬
‭generally in the form of ATP.‬

‭5.10 Elimination‬
‭ he final step in digestion is eliminating undigested food content and waste products. The‬
T
‭semi-solid waste is moved through the colon by peristaltic movements of the muscle and is‬
‭stored in the rectum. As the rectum expands in response to the storage of fecal matter, it‬
‭triggers the neural signals required to stimulate‬‭elimination‬‭. The solid waste is eliminated‬
‭through the anus using peristaltic movements of the rectum.‬
‭ onstipation and diarrhea are two of the most common health concerns that affect‬
C
‭digestion.‬‭Constipation‬‭is a condition where the feces‬‭are hardened because of excess‬
‭water removal in the colon. In contrast, if enough water is not removed from the feces, it‬
‭results in‬‭diarrhea‬‭. Many bacteria, including those‬‭that cause cholera, affect the proteins‬
‭involved in water reabsorption in the colon, resulting in excessive diarrhea.‬
‭ or additional information about constipation, click the link below to watch the TED-Ed‬
F
‭video by‬‭Heba Shaheed titled “What causes constipation?”.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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What causes constipation? - Heba Shaheed

‭5.11 Career Connection - Gastroenterologist‬

‭ gastroenterologist is a medical doctor who specializes in digestive system conditions‬‭.‬
A
‭Gastroenterologists begin as general physicians. They complete three years of medical‬
‭residency after medical school, treating various diseases and conditions. To become‬
‭gastroenterologists, they complete three years of additional study after that. Then, they‬
‭receive special certification. This certification designates them as experts in‬
‭gastrointestinal diseases‬‭and conditions. It also‬‭qualifies them to perform certain exams‬
‭and procedures that general physicians don’t and to interpret the results.‬

‭ pediatric gastroenterologist is a‬‭pediatrician‬‭first,‬‭with extra training in‬

A
‭gastroenterology. Pediatricians spend three years of medical residency practicing general‬
‭pediatric medicine, treating babies, children, and teens for various conditions. Pediatric‬
‭gastroenterologists study for three more years after that to earn their certification. They‬
‭study the gastrointestinal and liver conditions most relevant to growing children, with a‬
‭special emphasis on nutrition. They learn to interpret children’s signs and symptoms and‬
‭perform exams and minor procedures inside their smaller bodies.‬

(‭ credit: Cleveland Clinic Health Library;‬

‭https://my.clevelandclinic.org/health/articles/24198-gastroenterologist‬‭;‬‭Reproduced with‬
‭permission.)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 6: Energy Considerations‬

‭ igure 6.1‬ ‭Metabolism is the sum of all energy-requiring‬‭and energy-consuming‬

F
‭processes of the body. Many factors contribute to overall metabolism, including lean‬
‭muscle mass, the amount and quality of food consumed, and the physical demands placed‬
‭o n the human body. (credit: "tableatny"; Flickr.com;‬‭OpenStax‬‭)‬

‭6.1 Introduction‬
I‭ n childhood, someone may have told you to start the day with a good breakfast to give‬
‭you the energy to get through most of the day. You have heard about a balanced diet with‬
‭fruits and vegetables. What does this mean to your body and physiological processes?‬
‭How do living cells obtain energy?‬
S‭ ome organisms, such as plants, can produce sugar molecules from “scratch” (e.g., water‬
‭and carbon dioxide) through‬‭photosynthesis.‬ ‭In this‬‭process, plants use light (‬‭solar‬
‭energy‬‭) to power the reactions to convert water and‬‭carbon dioxide into glucose‬
‭(‬‭c hemical energy‬‭), which stores this energy in its‬‭chemical bonds. Then, via a series of‬
‭metabolic pathways, collectively called‬‭cellular respiration‬‭,‬‭plants extract the energy‬
‭from glucose and convert it into a form (ATP) that can be used to power its various‬
‭processes (‬‭Figure 6.2‬‭). Oxygen, a by-product of photosynthesis,‬‭is vital for cellular‬
‭respiration.‬
‭ s humans can’t do photosynthesis, we must continually consume nutrients, including‬
A
‭glucose, so that our cells have the building blocks for metabolic processes that release‬
‭energy as ATP. ATP is needed for functions such as manufacturing new proteins, cells, and‬
‭body parts and recycling materials in the cell. All of the chemical reactions inside cells,‬
‭including those that use or release energy, are referred to as the cell’s‬‭metabolism‬‭.‬

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‭Chapter 6: Energy Considerations‬

‭ igure 6.2‬ ‭The summary of the chemical equations‬‭for photosynthesis and cellular‬
F
‭respiration. Note that the products of photosynthesis (glucose and oxygen) are reactants‬
‭for cellular respiration. (credit: Modified from‬‭Cellular Respiration Simple.png‬‭;‬
‭Christinelmiller‬‭;‬‭CC BY-SA 4.0‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ his chapter will introduce you to various aspects of metabolism, such as energy and its‬
T
‭different forms, the physical laws governing energy transfer, anabolic and catabolic‬
‭reactions, and how cells produce energy via aerobic and anaerobic metabolism and‬
‭nutrients. After completing work on this chapter, you will likely think about food and‬
‭eating in a different way!‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 6: Energy Considerations‬

‭6.2 Energy and the Laws of Thermodynamics‬

‭ e define‬‭energy‬‭as the ability to do work. As you‬‭know, energy exists in different forms,‬
W
‭such as electrical, light, and heat. When an object is in motion, energy is associated with it‬
‭because moving objects can enact a change or do work. Think of a wrecking ball. Even a‬
‭slow-moving wrecking ball can do considerable damage to other things. However, a‬
‭wrecking ball that is not in motion cannot perform work. Energy associated with objects in‬
‭motion is called‬‭kinetic energy‬‭.‬
‭ hat if we lift that motionless wrecking ball two stories above a car with a crane? If the‬
W
‭suspended wrecking ball is unmoving, can we associate energy with it? The answer is yes.‬
‭The suspended wrecking ball has associated energy fundamentally different from the‬
‭kinetic energy of objects in motion—this energy form results from the‬‭potential‬‭for the‬
‭wrecking ball to do work. If we release the ball, it will do work. Because this energy type‬
‭refers to the potential to do work, we call it‬‭potential‬‭energy‬‭.‬
‭ hermodynamics studies the relationship between different forms of energy and the‬
T
‭conversion between them. Two laws of thermodynamics govern this study. The first one,‬
‭also called the‬‭law of conservation of energy‬‭, deals‬‭with the total amount of energy in‬
‭the universe. It states that while energy exists in many forms, its total energy is constant.‬
‭In other words, there has always been, and always will be, precisely the same amount of‬
‭energy in the universe. According to the‬‭first law‬‭of thermodynamics‬‭, energy can change‬
‭into different forms but cannot be created or destroyed (‬‭Figure 6.3‬‭).‬
‭ he‬‭second law of thermodynamics‬‭addresses the inefficiency‬‭o f energy conversions‬
T
‭from one form to another. In other words, in every energy transfer, some amount of‬
‭energy is lost in an unusable form (i.e., can’t be captured to do work), and in most cases,‬
‭this form is‬‭heat energy‬‭. For example, when a car’s‬‭engine converts the chemical energy‬
‭in gasoline into mechanical energy, some will be lost as heat that will be detected when‬
‭you put your hand on the car’s hood. Likewise, some energy is lost as heat energy during‬
‭cellular metabolic reactions. This loss is good for warm-blooded creatures (e.g., humans)‬
‭because heat energy helps maintain our body temperature.‬
I‭ n studying the thermodynamics of chemical reactions, the difference between the energy‬
‭content of the reactants and the products is represented by the symbol “∆G” (read “delta‬
‭G”), where G represents “‬‭Gibbs free energy‬‭,” or the‬‭energy available to do work. Delta G‬
‭represents the difference between the products' energy and the reactants' energy (i.e., the‬
‭energy associated with the products minus the energy associated with the reactants).‬
‭Scientists classify reactions with a‬‭negative ∆G value‬‭(∆G < 0), meaning there is a net‬
‭release of energy, as‬‭exergonic reactions‬‭. Think:‬‭ex‬‭ergonic means energy is‬‭ex‬‭iting the‬
‭system. These reactions can be harnessed to perform work inside the cell.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 6: Energy Considerations‬

‭ igure 6.3‬ ‭Shown are two examples of energy being‬‭transferred from one system to‬
F
‭another and transformed from one form to another. Humans can convert the chemical‬
‭energy in food, like this ice cream cone, into kinetic energy (the energy of movement to‬
‭ride a bicycle). Plants can convert light energy from the sun into glucose, a source of‬
‭chemical energy. (credit “ice cream”: modification of work by D. Sharon Pruitt; credit “kids‬
‭o n bikes”: modification of work by Michelle Riggen-Ransom; credit “leaf”: modification of‬
‭work by Cory Zanker;‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ hemical reactions that require energy input rather than releasing energy will have a‬
C
‭positive ∆G value (‬‭∆G > 0) and are classified as‬‭endergonic reactions‬‭. In this case, the‬
‭products have more free energy than the reactants so we can think of them as‬
‭energy-storing molecules.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 6: Energy Considerations‬

‭ igure 6.4‬ ‭Gibbs free energy diagrams for exergonic‬‭and endergonic reactions. In the‬
F
‭first graph, the value of delta G is negative (i.e., less than zero), so the energy associated‬
‭with the products is less than that associated with the reactants. In the second graph, the‬
‭value of delta G is positive (i.e., greater than zero), so the energy associated with the‬
‭products is greater than that associated with the reactants. (credit:‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭6.3 Anabolic and Catabolic Chemical Reactions‬

‭ ne way to classify metabolic chemical reactions is to determine if they break down‬
O
‭molecules (catabolic) or build larger ones (anabolic). The reactions governing the‬
‭breakdown of food to obtain energy are called‬‭c atabolic‬‭reactions‬‭. Conversely,‬‭anabolic‬
‭reactions‬‭use the energy produced by catabolic reactions‬‭to synthesize large molecules‬
‭from smaller ones, such as when the body forms proteins by joining amino acids in a‬
‭specific sequence. Both sets of reactions are critical to maintaining life.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ atabolic reactions break down large organic molecules into smaller molecules, releasing‬
C
‭the energy contained in the chemical bonds (‬‭Figure‬‭6.5‬‭), which means they are exergonic‬
‭(∆G < 0). This energy is harvested so that it can be used to produce ATP.‬
‭ ther energy-storing molecules, such as fats, are also broken down through similar‬
O
‭catabolic reactions to release energy and make ATP.‬
‭ he energy releases (conversions) used to synthesize ATP are not 100 percent efficient‬
T
‭due to the second law of thermodynamics. The energy released is less than the total‬
‭amount contained in the molecule. Approximately 40 percent of energy yielded from‬
‭catabolic reactions is directly transferred to the high-energy molecule ATP, the energy‬
‭currency of the cells, which can be used immediately to power molecular machines that‬
‭support cell, tissue, and organ function. The remaining 60 percent of the energy released‬
‭from catabolic reactions is given off as heat, which tissues and body fluids absorb.‬
I‭ n contrast to catabolic reactions, anabolic reactions involve joining smaller molecules into‬
‭larger ones, which means they are endergonic (∆G > 0). For example, anabolic reactions‬
‭combine monosaccharides to form polysaccharides, fatty acids and glycerol to form‬
‭triglycerides, amino acids to form proteins, and nucleotides to form nucleic acids. These‬
‭processes require energy from ATP molecules generated by catabolic reactions. Anabolic‬
‭reactions create new molecules that form new cells and tissues and revitalize organs.‬

‭ igure 6.5‬ ‭Anabolic pathways require energy to synthesize‬‭large molecules and are‬
F
‭endergonic. Catabolic pathways generate energy by breaking down larger molecules and‬
‭are exergonic. Both types of pathways are required for maintaining the cell’s energy‬
‭balance. (credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭6.4 Enzymes‬
‭ e must consider another critical concept regarding endergonic/anabolic and‬
W
‭exergonic/catabolic reactions. Even exergonic reactions require a small amount of energy‬
‭input before proceeding with their energy-releasing steps. These reactions have a net‬
‭release of energy, meaning they are exergonic but still require some initial energy.‬
‭Activation energy‬‭is the term used to describe this‬‭small amount of energy input‬
‭necessary for all chemical reactions to begin.‬
‭ he activation energy of a particular reaction determines the rate at which it will proceed.‬
T
‭The higher the activation energy, the slower the chemical reaction. Iron rusting, for‬
‭example, illustrates an inherently slow reaction. This reaction occurs slowly over time‬
‭because of its higher activation energy requirement.‬
‭ ike these reactions outside of cells, the activation energy for most cellular reactions is too‬
L
‭high for heat energy to overcome at efficient rates. In other words, their activation‬
‭energies must be lowered for important cellular reactions to occur at appreciable rates‬
‭(number of reactions per unit of time). Scientists refer to this as catalysis. Macromolecules,‬
‭such as proteins, DNA, and RNA, store considerable energy, and their breakdown is‬
‭exergonic. The cell's essential components would disintegrate if cellular temperatures‬
‭provided enough heat energy for these exergonic reactions to overcome their activation‬
‭barriers.‬
‭ substance that helps a chemical reaction to occur is a‬‭c atalyst‬‭, and the particular‬
A
‭molecules that catalyze biochemical reactions are‬‭enzymes‬‭. Almost all enzymes are‬
‭proteins composed of amino acid chains, and they perform the critical task of lowering the‬
‭activation energies of chemical reactions inside the cell (‬‭Figure 6.6‬‭). Enzymes do this by‬
‭their‬‭active sites‬‭binding to the reactant molecules,‬‭called‬‭substrates‬‭, and holding them in‬
‭such a way as to make the chemical bond-breaking and bond-forming processes take‬
‭place more readily (‬‭Figure 6.7‬‭). It is important‬‭to note that enzymes only lower the‬
‭activation energy; the delta G value for the reaction is unchanged.‬

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‭ igure 6.6‬ ‭Enzymes reduce the reaction's activation‬‭energy but do not change the‬
F
‭reaction's free energy. (credit:‬‭OpenStax‬‭). A link‬‭to a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ igure 6.7‬ ‭The interaction between an enzyme and‬‭its substrate. The active site is the‬
F
‭region of an enzyme created by the secondary and tertiary folding of the protein. The site‬
‭is specific for a substrate like a key is specific for a lock. Enzymes are needed for both‬
‭anabolic and catabolic reactions. (credit:‬‭OpenStax‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ or additional information about activation energy and enzymes, click the link below or‬
F
‭scan the QR code to watch the TED-Ed video by Vance Kite‬‭titled “Activation energy:‬
‭Kickstarting chemical reactions.”‬

Activation energy: Kickstarting chemical reactions - Vance Kite

‭6.5 ATP and Related Topics‬

‭ s we learned in previous chapters, living organisms require free energy to power life‬
A
‭processes.‬‭ATP‬‭(adenosine‬‭tri‬‭phosphate) functions‬‭as the primary energy currency for‬
‭cells. Structurally, ATP molecules consist of an adenine base, a ribose sugar, and three‬
‭phosphate groups (‬‭Figure 6.8‬‭).‬

‭ igure 6.8‬ ‭A molecule of ATP contains an adenine‬‭base, a ribose sugar, and three‬
F
‭phosphate groups. If one phosphate group is removed, the resulting molecule is ADP.‬
‭(credit: modified from‬‭230 Structure of Adenosine‬‭Triphosphate (ATP)-01.jpg‬‭; OpenStax‬
‭College;‬‭CC BY 3.0‬‭).‬‭A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

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‭ emember that adenine is a nitrogen-containing base in DNA and RNA, and ribose is a‬
R
‭five-carbon sugar in RNA nucleotides (See Chapters 2 and 4). The chemical bond between‬
‭the second and third phosphate groups, a high-energy bond, represents a cell's most‬
‭significant energy source. It is the first bond that catabolic enzymes break when cells‬
‭require energy to do work. The products of this reaction are a molecule of adenosine‬
‭di‬‭phosphate (ADP) and a lone phosphate group designated‬‭as P‬‭i‬ ‭(inorganic phosphate).‬
‭ATP, ADP, and P‬‭i‬ ‭are constantly cycled through reactions‬‭called the‬‭ATP-ADP cycle‬‭, which‬
‭builds ATP and store energy, and reactions that break down ATP and release energy‬
‭(‬‭Figure 6.9‬‭).‬

‭ igure 6.9‬ ‭The ATP/ADP cycle. (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of this‬
F
‭figure is available at‬‭Biology411.com‬‭.‬

‭ nergy is released when ATP is changed to ADP by removing its terminal phosphate group‬
E
‭via‬‭dephosphorylation‬‭. With its associated energy,‬‭the released phosphate group‬
‭typically binds to another molecule, activates it, and results in cellular work (‬‭Figure 6.10‬‭).‬

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‭ igure 6.10‬ ‭In phosphorylation reactions, the terminal‬‭phosphate of ATP is broken off‬
F
‭and attached to another molecule (e.g., a protein), and ADP is released. Adding a‬
‭phosphate group energizes the molecule and allows work to be accomplished (e.g., active‬
‭transport; Chapter 3). (credit:‬‭OpenStax‬‭)‬

‭ nergy derived from glucose metabolism converts ADP to ATP via adding a phosphate‬
E
‭group (‬‭phosphorylation‬‭). As we explore cellular respiration,‬‭we’ll learn that the two‬
‭ways ATP is regenerated by the cell are substrate-level phosphorylation and oxidative‬
‭phosphorylation.‬
I‭ n‬‭substrate-level phosphorylation‬‭, ATP is regenerated‬‭from ADP as a direct result of‬
‭the chemical reactions in the catabolic pathways. A phosphate group is removed from an‬
‭intermediate reactant in the pathway, and the reaction's free energy is used to add the‬
‭third phosphate to an available ADP molecule, producing ATP.‬
‭ ost ATP is generated in a more complex process called‬‭oxidative phosphorylation‬‭,‬
M
‭which occurs in‬‭mitochondria‬‭(Chapter 3) within a‬‭eukaryotic cell or the plasma‬
‭membrane of a prokaryotic cell. This process generates approximately 90 percent of the‬
‭ATP made during glucose catabolism. It is called oxidative phosphorylation because‬
‭oxygen is involved in the process.‬

‭6.6 Electrons, Energy, and Electron Carriers‬

‭ he transfer of electrons between molecules is necessary because most of the energy‬
T
‭stored in atoms and used to fuel cellular functions is in the form of high-energy electrons.‬
‭The transfer of energy in the form of electrons allows the cell to transfer and use energy‬
‭incrementally—in small packages rather than in a single, destructive burst.‬
‭ he chemical reactions underlying metabolism involve the transfer of electrons from one‬
T
‭compound to another. The electrons in these reactions commonly come from hydrogen‬
‭atoms, which consist of an electron and a proton. A molecule gives up a hydrogen atom in‬
‭the form of a‬‭hydrogen ion (H‬‭+‭)‬ ‬‭and an electron, breaking‬‭the molecule into smaller‬
‭parts. The loss of an electron, called‬‭oxidation‬‭,‬‭releases a small amount of energy; both‬

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t‭ he electron and the energy are then passed to another molecule in the process of‬
‭reduction‬‭o r gaining an electron. These two reactions‬‭always happen together (i.e., an‬
‭oxidation-reduction reaction).‬
I‭ n living systems, a small class of compounds, called‬‭coenzymes‬‭, functions as electron‬
‭shuttles: They bind and carry high-energy electrons between compounds in pathways. The‬
‭principal electron carriers we will consider are derived from the B vitamin group and are‬
‭derivatives of nucleotides. These compounds can be easily reduced (that is, they accept‬
‭electrons) or oxidized (they lose electrons). Nicotinamide adenine dinucleotide (NAD) is‬
‭derived from vitamin B3, niacin.‬‭NAD‬‭+‬ ‭is the oxidized‬‭form of the molecule;‬‭NADH‬‭is the‬
‭reduced form of the molecule after it has accepted two electrons and a proton, which are‬
‭the equivalent of a hydrogen atom with an extra electron (‬‭Figure 6.11‬‭).‬

‭ igure 6.11‬ ‭The oxidized form of the electron carrier‬‭(NAD‬‭+‭)‬ is on the left, and the‬
F
‭reduced form (NADH) is on the right. The nitrogenous base in NADH has one more‬
‭hydrogen ion and two more electrons than in NAD‬‭+‬‭(see‬‭the red boxes at the top of the‬
‭figure). (credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

S‭ imilarly, flavin adenine dinucleotide (‬‭FAD‬‭) is derived‬‭from vitamin B2, also called‬
‭riboflavin. Its reduced form is‬‭FADH‬‭2‭.‬ NAD‬‭+‬ ‭and FAD‬‭are extensively used in energy‬
‭extraction from sugars, as we will observe in the remainder of the chapter.‬

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‭6.7 An Overview of Cellular Respiration‬

‭ s discussed previously, cellular respiration is the phrase used to describe a series of‬
A
‭metabolic reactions that break down glucose in the presence of oxygen to produce carbon‬
‭dioxide, water, and energy as heat and ATP:‬
‭C‬‭6‭H
‬ ‬‭12‬‭O‬‭6‬ ‭+ 6 O‬‭2‬ ‭-------------> 6 CO‬‭2‬ ‭+ 6 H‬‭2‬‭0 + Energy‬‭(ATP and Heat)‬

‭ he numbers in front of oxygen, carbon dioxide, and water are called‬‭coefficients‬‭, which‬
T
‭indicate the numbers of each of these molecules involved in the reaction, so it follows the‬
‭law of conservation of matter. In other words, the number of atoms of each element on‬
‭the reactant side of the equation must equal the number on the product side. Subscripts‬
‭in a chemical formula cannot be changed because compounds contain specific ratios of‬
‭elements; however, coefficients can be changed. For example, based on the chemical‬
‭equation above, there are 12 hydrogen atoms on the reactant side of the equation, all‬
‭from the one molecule of glucose (If a number isn’t present in front of a molecular‬
‭formula, it is understood to be one.). On the product side, there are 6 molecules of water,‬
‭each with two hydrogen atoms, for a total of 12 hydrogen atoms. The same balance is‬
‭true for the numbers of carbon and oxygen atoms, as shown in the table below:‬

‭Element‬ ‭Number of Atoms‬ ‭Number of Atoms‬

‭(Reactants)‬ ‭(Products)‬

‭Carbon‬ ‭6‬ ‭6‬

‭Hydrogen‬ ‭12‬ ‭12‬

‭Oxygen‬ ‭18‬ ‭18‬

‭ ased on this summary chemical equation, one glucose molecule and six oxygen‬
B
‭molecules will chemically react to produce six molecules of carbon dioxide and six water‬
‭molecules. Energy is released as ATP and heat, indicating an exergonic reaction.‬
‭ ellular respiration occurs in four stages: glycolysis, transition reaction, Krebs cycle, and‬
C
‭the electron transport chain. To bring structure to the discussion, we can answer the‬
‭following seven questions for each of the first three stages of cellular respiration:‬
‭ .‬
1 ‭ here in the cell is it occurring?‬
W
‭2.‬ ‭What carbon-containing molecule enters?‬
‭3.‬ ‭What carbon-containing molecule leaves?‬
‭4.‬ ‭Is carbon dioxide produced?‬

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‭Chapter 6: Energy Considerations‬

‭ .‬ I‭ s ATP used?‬
5
‭6.‬ ‭Is ATP produced?‬
‭7.‬ ‭Are reduced coenzymes (NADH and FADH‬‭2‬‭) produced?‬

‭6.8 Stage 1: Glycolysis‬

‭ lycolysis‬‭is the first stage in the pathway to break‬‭down glucose to extract free energy. It‬
G
‭is important to note that glycolysis occurs in the cytoplasm of eukaryotic cells. Like all‬
‭metabolic pathways, glycolysis occurs in steps, each catalyzed by a different enzyme. In‬
‭this stage, the six-carbon glucose molecule is split into two, three-carbon‬‭pyruvate‬
‭molecules by investing two ATP molecules to energize the separation. (Don’t worry; the‬
‭cell will get the investment of ATP back. It’s like the stock market: You have to invest money‬
‭to, hopefully, make money!) As glucose is metabolized further, bonds are rearranged‬
‭through a series of steps, and free energy is released to form ATP via substrate-level‬
‭phosphorylation of ADP with available phosphate (P‬‭i‬‭)‬‭molecules.‬
‭ igh-energy electrons and hydrogen atoms pass to NAD‬‭+‬‭,‬‭reducing it to NADH. Although‬
H
‭two molecules of ATP were invested to destabilize glucose at the beginning of the process,‬
‭four molecules of ATP are formed by substrate-level phosphorylation, resulting in a net‬
‭gain of two ATP and two NADH molecules for the cell (‬‭Figure 6.12‬‭).‬
I‭ n summary, glycolysis occurs in the cytoplasm. One glucose molecule (with six carbon‬
‭atoms) enters, two molecules of pyruvate (each with three carbon atoms) leave, no carbon‬
‭dioxide is produced, two net ATP are produced, and two NADH are produced.‬
I‭ f oxygen is available, then the two pyruvate molecules will continue to the next stage of‬
‭cellular respiration, which is the transition reaction.‬

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‭Chapter 6: Energy Considerations‬

‭ igure 6.12‬ ‭In glycolysis, a six-carbon glucose‬‭molecule is converted into two pyruvate‬
F
‭molecules, each with three carbon atoms. Two net ATP (four produced minus two‬
‭invested) and two NADH are produced (credit:‬‭OpenStax‬‭).‬‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭6.9 Stage 2: Transition Reaction‬

I‭ n the next stage of cellular respiration, called the‬‭transition reaction‬‭, each of the two‬
‭molecules of pyruvate produced from glycolysis undergoes three changes (‬‭Figure 6.13‬‭):‬

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‭Chapter 6: Energy Considerations‬

‭ igure 6.13‬ ‭During the transition reaction, three‬‭steps occur to convert each pyruvate‬
F
‭into acetyl CoA. In the process, carbon dioxide is released, and one molecule of NADH is‬
‭formed. (credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ hey are transformed into an‬‭acetyl group‬‭containing‬‭two carbon atoms attached to a‬

T
‭coenzyme A‬‭(CoA) carrier molecule, creating‬‭acetyl‬‭CoA‬‭. During this conversion, a‬
‭molecule of CO‬‭2‬ ‭and two high-energy electrons are‬‭removed. NAD+ picks up the electrons‬
‭and a hydrogen ion, and the resulting NADH carries the electrons to the next stage for ATP‬
‭production (‬‭Figure 6.14‬‭).‬

‭ igure 6.14‬ ‭In the transition reaction, each pyruvate‬‭molecule (pyruvic acid) produces‬
F
‭o ne carbon dioxide molecule, one NADH molecule, and one molecule of acetyl CoA (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at biology411.com.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 6: Energy Considerations‬

I‭ n summary, the transition reaction occurs when pyruvate, produced by glycolysis, is‬
‭transported from the cytoplasm into the mitochondria. Two pyruvate molecules (each‬
‭with three carbon atoms) enter, two acetyl CoA (each with two carbon atoms) leave, two‬
‭carbon dioxide are produced, no ATP is produced, and two NADH are produced.‬

‭6.10 Stage 3: Krebs Cycle‬

‭ he third stage of cellular respiration goes by several names, including the‬‭c itric acid‬
T
‭c ycle‬‭and the‬‭Krebs cycle‬‭, after Hans Krebs, who first‬‭identified the steps in the pathway‬
‭in the 1930s in pigeon flight muscles. These reactions occur in the mitochondria, and the‬
‭term cycle is used because the eight steps of the cycle are a series of chemical reactions‬
‭that regenerate the first molecule (called oxaloacetate) that combines with the acetyl‬
‭group. This pathway will harvest the remainder of the extractable energy from what‬
‭began as a glucose molecule.‬
‭ rocessing of the two acetyl CoA molecules in the Krebs cycle will result in the production‬
P
‭o f a total of six NADH, two FADH‬‭2‬‭, and two ATP (‬‭Figure‬‭6.15‬‭). Also note that six carbon‬
‭dioxide molecules have been released at this point, accounting for the six carbons in the‬
‭starting glucose molecule. The high-energy NADH and FADH‬‭2‬ ‭produced in this stage will‬
‭be used with the other NADH (produced via glycolysis and the transition reaction) in the‬
‭last stage of cellular respiration (electron transport chain) to make the majority of ATP‬
‭molecules via oxidative phosphorylation.‬

‭ igure 6.15‬ ‭In the Krebs cycle, each molecule of‬‭acetyl-CoA (with two carbon atoms) is‬
F
‭broken down to yield 2 CO‬‭2‭.‬ Additionally, 3 NADH,‬‭1 FADH‬‭2‭,‬ and 1 ATP are produced. As‬
‭the transition reaction produces two molecules of acetyl CoA, the Krebs cycle yields 4 CO‬‭2‭,‬ ‬
‭6 NADH, 2 FADH‬‭2‬‭, and 2 ATP. (credit:‬‭OpenStax‬‭). A‬‭link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 6: Energy Considerations‬

‭6.11 Stage 4: The Electron Transport Chain and Oxidative Phosphorylation‬

‭ ou have just read about three stages in glucose catabolism: glycolysis, the transition‬
Y
‭reaction, and the Krebs cycle. To this point, a net yield of only four ATP has been produced‬
‭via substrate-level phosphorylation, about 10% of the total observed ATP yield of one‬
‭glucose molecule via cellular respiration. Most of the ATP generated during cellular‬
‭respiration is not produced by substrate-level phosphorylation but is instead derived from‬
‭a process called oxidative phosphorylation, which is part of the fourth stage, called the‬
‭electron transport chain‬‭. This stage is the only part‬‭o f glucose metabolism that directly‬
‭uses the oxygen taken in via the respiratory system and transported to cells via the‬
‭circulatory system.‬
‭ nderstanding the mitochondria’s basic structure is essential before discussing the events‬
U
‭o f the last stage. Mitochondria have two phospholipid bilayers, an inner membrane and‬
‭an outer membrane, which create inner and outer mitochondrial compartments (‬‭Figure‬
‭6.16‬‭).‬

‭ igure 6.16‬ ‭The basic structure of mitochondria is‬‭shown. The two phospholipid‬
F
‭membranes form an inner and outer compartment within the organelle. (credit:‬
‭modification of‬‭Mitochondrie.svg‬‭;‬‭Tatoute‬‭;‬‭CC BY-SA‬‭3.0‬‭). A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭ he electron transport chain begins by moving electrons obtained from the reduced‬
T
‭coenzymes (NADH and FADH‬‭2‬‭) through a series of electron‬‭carriers embedded in the‬
‭inner mitochondrial membrane. There are multiple copies of this series of electron‬
‭carriers in each mitochondrion, and the energy from the electrons being transferred‬
‭through the chain components is used to pump hydrogen ions via active transport from‬
‭the inner to the outer compartment. This action causes hydrogen ions to accumulate‬
‭within the outer compartment and decrease the pH of this region. Oxygen, the‬‭terminal‬

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‭Chapter 6: Energy Considerations‬

‭electron acceptor‬‭, will join with hydrogen ions to produce metabolic water (‬‭Figure 6.17‬‭).‬

‭ igure 6.17‬ ‭The electron transport chain is a series‬‭o f electron transporters embedded‬
F
‭in the inner mitochondrial membrane that shuttles electrons from NADH and FADH‬‭2‬ ‭to‬
‭molecular oxygen, the terminal electron acceptor. In the process, protons (H‬‭+‭)‬ are pumped‬
‭from the inner to the outer compartment, and oxygen is reduced to form water. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ herefore, a concentration gradient forms in which hydrogen ions diffuse from the outer‬
T
‭mitochondrial compartment into the inner one by passing through a channel protein‬
‭associated with an enzyme called‬‭ATP synthase‬‭. This‬‭complex protein acts as a tiny‬
‭generator, turned by the force of the hydrogen ions diffusing through it from higher to‬
‭lower concentration. Turning parts of this molecular machine facilitate phosphorylating‬
‭ADP to produce ATP (‬‭Figure 6.18‬‭).‬
‭ he overall result of this stage of cellular respiration is the production of ATP from the‬
T
‭energy of the electrons removed from hydrogen atoms, which were initially part of a‬
‭glucose molecule. At the end of the pathway, the electrons join an oxygen molecule to form‬
‭oxygen ions. The extra electrons on the oxygen attract hydrogen ions (protons) from the‬
‭surrounding medium, and water is formed. Thus, oxygen is the final electron acceptor in‬
‭the electron transport chain. A summary of the electron transport chain and the coupled‬
‭action of ATP synthase in oxidative phosphorylation is shown in‬‭Figure 6.19‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 6.18‬ ‭ATP synthase is a complex molecular‬‭machine that uses a proton (H‬‭+‭)‬ ‬
F
‭gradient to form ATP from ADP and inorganic phosphate (Pi). (credit: modification of‬
‭work by Klaus Hoffmeier;‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ igure 6.19‬ ‭Oxidative phosphorylation (credit:‬‭Modified from‬‭OpenStax‬‭). A link to a‬

F
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭6.12 Summary ATP Yield from Cellular Respiration‬

I‭ n the electron transport chain, the energy potential associated with the electrons from‬
‭NADH is greater than that of FADH‬‭2‬ ‭because of the‬‭point where donated electrons enter‬
‭the transport chain. This difference means that the ATP yield of NADH is greater than‬
‭FADH2 (3 versus 2 ATP per molecule, respectively) in oxidative phosphorylation. This‬
‭difference is important when calculating one glucose molecule's ATP yield from cellular‬
‭respiration. The total ATP produced via substrate phosphorylation and oxidative‬
‭phosphorylation of one glucose molecule is shown in‬‭Figure 6.20‬‭.‬

‭No. of ATP Produced by the‬

‭ o. of ATP‬
N ‭No. of‬ ‭Processing of Reduced‬
‭Produced by‬ ‭ educed‬
R ‭Coenzymes During Oxidative‬ ‭Total Number‬
‭Substrate-Level‬ ‭Coenzymes‬ ‭Phosphorylation (3 ATP per‬ ‭of ATP‬
‭Phosphorylation‬ ‭Produced‬ ‭NADH; 2 ATP per FADH‬‭2‭)‬ ‬ ‭Produced‬
‭Stage‬

‭Glycolysi‬‭s‬ ‭2 ATP‬ ‭2 NADH‬ ‭6 ATP‬ ‭8 ATP‬

‭ ransition‬
T ‭0 ATP‬ ‭2 NADH‬ ‭6 ATP‬ ‭6 ATP‬
‭Reaction‬

‭Krebs Cycle‬ ‭2 ATP‬ ‭ NADH‬

6 ‭ 8 ATP‬
1 ‭24 ATP‬
‭2 FADH‬‭2‬ ‭4 ATP‬

‭38 ATP‬

‭ igure 6.20‬ ‭Determining the net ATP yield from cellular‬‭respiration of one glucose‬
F
‭molecule. A link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

I‭ n some cases, depending on the tissue type, the transport of NADH produced via‬
‭glycolysis into the mitochondria requires the input of energy in the form of 2 ATP. In that‬
‭case, the total ATP yield is reduced from 38 to 36 per molecule of metabolized glucose.‬
‭This reduction is shown in Figure 6.21, along with additional details about the overall‬
‭reduced coenzyme (NADH and FADH‬‭2‭)‬ and ATP yields.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 6: Energy Considerations‬

‭ igure 6.21‬ ‭Summary of cellular respiration of a‬‭glucose molecule. (credit:‬‭OpenStax‬‭).‬ ‭A‬

F
‭link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 6: Energy Considerations‬

‭6.13 Metabolism without Oxygen: Lactic Acid and Alcohol Fermentation‬

I‭ n cellular respiration, ATP is produced using the energy of high-energy electrons carried‬
‭by NADH or FADH‬‭2‬ ‭to the electron transport chain.‬‭If oxygen isn’t present, then cellular‬
‭respiration doesn’t occur, as the terminal electron acceptor isn’t available. However,‬
‭glycolysis isn’t affected and will continue if NAD+ is available and a net yield of 2 ATP per‬
‭glucose is produced. Therefore, the 2 NADH made during glycolysis must be reoxidized to‬
‭NAD‬‭+‭,‬ as the electron transport chain isn’t operating.‬ ‭How is this done? Processes that‬
‭use an organic molecule, not oxygen, to regenerate NAD‬‭+‬ ‭from NADH are collectively‬
‭called‬‭fermentation‬‭.‬
‭ he fermentation method used by animals and certain bacteria, such as those in yogurt, is‬
T
‭lactic acid fermentation and is represented by the following simplified equation:‬
‭Pyruvate + NADH —-----> Lactic acid (lactate) + NAD‬‭+‬
‭ his type of fermentation is also routinely used in mammalian red blood cells, which do‬
T
‭not have mitochondria, and in skeletal muscle with an insufficient oxygen supply to allow‬
‭cellular respiration to continue (that is, in muscles used to the point of fatigue).‬
I‭ n muscles,‬‭lactic acid‬‭accumulation must be removed‬‭by blood circulation, which is‬
‭accomplished via the liver. Here, the lactic acid can be used to create glucose and then‬
‭stored as glycogen. The association of glycolysis and lactic acid fermentation is shown in‬
‭Figure 6.22,‬‭and the chemical change of pyruvate to‬‭lactate is shown in‬‭Figure 6.23‬‭.‬
S‭ uch lactic acid accumulation was once believed to cause muscle stiffness, fatigue, and‬
‭soreness, although more recent research disputes this hypothesis. Once the lactic acid has‬
‭been removed from the muscle and circulated to the liver, it can be reconverted into‬
‭pyruvic acid and further catabolized for energy.‬

‭ igure 6.22‬ ‭In lactic acid (lactate) fermentation,‬‭the end‬

F
‭product of glycolysis (pyruvate) is reduced to lactate to‬
‭regenerate NAD‬‭+‭.‬ (credit:‬‭OpenStax‬‭). A link to a‬‭video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 6.23‬ ‭The reduction of pyruvate to lactate‬‭accompanied by the oxidation of NADH‬

F
‭to NAD‬‭+‭,‬ which can sustain glycolysis and generate‬‭two net ATP from each glucose. credit:‬
‭modification of‬‭D-LDH reaction.svg‬‭;‬‭Miguelferig‬‭; Wikimedia‬‭Commons;‬‭CC0 1.0‬‭). A link to‬
‭a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ nother fermentation process yeast cells use is‬‭alcohol‬‭fermentation‬‭(‬‭Figure 6.24‬‭). In‬

A
‭the first reaction, pyruvate loses a carbon atom as carbon dioxide, producing a‬
‭two-carbon molecule called acetaldehyde. In the second reaction, NADH gives up two‬
‭electrons and a hydrogen ion to NAD‬‭+,‬ ‭and acetaldehyde‬‭is converted to ethanol.‬

‭ igure 6.24‬ ‭Ethanol fermentation reactions (credit:‬‭modified from‬‭Pyruvate decarb‬

F
‭1.svg‬‭;‬‭Cwernert‬‭; vectorized by‬‭GKFX‬‭;‬‭CC BY-SA 3.0‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ thanol‬‭is a component of alcoholic beverages, such‬‭as wine and beer (‬‭Figure 6.25‬‭). The‬
E
‭ethanol tolerance of yeast is variable, ranging from about 5 percent to 21 percent,‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ epending on the yeast strain and environmental conditions. In other words, when the‬
d
‭alcohol content of a fermenting solution reaches a specific concentration, the yeast cells‬
‭can’t survive, and fermentation ceases.‬

‭ igure 6.25‬ ‭Fermentation of grape juice into wine‬‭produces CO‬‭2‬ ‭as a byproduct, which is‬
F
‭why these tanks have valves to release pressure inside the tanks. (credit:‬‭OpenStax‬‭)‬

‭ hile students are familiar with alcohol and alcoholic beverages, far fewer students are‬
W
‭aware of how alcohol impacts the body. For additional information about this topic, click‬
‭the link or scan the QR code to watch the TED-Ed video by Judy Grisel‬‭titled “How does‬
‭alcohol cause hangovers?”.‬

How does alcohol cause hangovers? - Judy Grisel

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 6: Energy Considerations‬

‭6.14 Summarizing Cellular Respiration and Fermentation‬

‭ he previous sections provide substantial detail about these two forms of metabolism,‬
T
‭which can be overwhelming! To facilitate understanding, it is helpful to summarize the‬
‭content in different ways. The first method is via a comparison table (‬‭Figure 6.26‬‭), and‬
‭the second method is via a flow chart (‬‭Figure 6.27‬‭).‬

‭Process‬ ‭→‬ ‭Cellular Respiration‬ ‭Fermentation‬

‭Item‬ ‭↓‬

‭1. Process location(s)‬ ‭Cytoplasm and mitochondria‬ ‭Cytoplasm only‬

‭2. Stages Involved‬ ‭ lycolysis, transition‬

G ‭Glycolysis only‬
‭reaction, Krebs cycle, and‬
‭electron transport chain‬

‭3. Final electron acceptor‬ ‭Oxygen (forms water)‬ ‭Lactic acid or ethanol‬‭*‬

‭4. Net ATP yield per glucose‬ ‭38‬‭**‬ ‭2‬

‭ . The final‬
5 ‭Carbon dioxide‬ ‭Lactic acid or ethanol‬‭*‬
‭carbon-containing compound‬
‭produced‬

*‭ ‬‭Product depends on the organism considered.‬

‭**‬‭The yield can be 36 or 38 ATP, depending on how‬‭the NADH produced in glycolysis‬
‭transfers electrons into the mitochondria.‬

‭Figure 6.26‬ ‭A comparison table of cellular respiration‬‭and fermentation.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 6.27‬ ‭A flow-chart comparison of cellular‬‭respiration and fermentation. (credit:‬

F
‭OpenStax‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 6: Energy Considerations‬

‭6.15 Connections of Carbohydrate, Protein, and Lipid Metabolic Pathways‬

‭ he most common nutrient metabolized for ATP production is glucose. Complex‬
T
‭carbohydrates (e.g., starch and glycogen) can be hydrolyzed to yield glucose monomers.‬
‭However, what about triglycerides and proteins? Fortunately, especially for students,‬
‭there isn’t a completely different process for using these nutrients. Instead, they undergo‬
‭chemical processes to change them into intermediates that are present in glucose‬
‭metabolism (‬‭Figure 6.28‬‭). For example, the carbon‬‭chains of fatty acids can be broken‬
‭into two-carbon fragments and converted into acetyl CoA. In the case of proteins, the‬
‭amino group (-NH‬‭2‭)‬ can be removed from amino acids,‬‭and the resulting molecule is‬
‭converted into pyruvate, acetyl CoA, or intermediates in the Krebs cycle, depending on the‬
‭particular amino acid. The removed‬‭amino group‬‭is‬‭first converted into‬‭ammonia‬‭(NH‬‭3‭)‬ ‬
‭molecules. Then, the liver combines them with carbon dioxide to produce‬‭urea‬‭, a waste‬
‭product secreted via the urine.‬
I‭ t is important to remember that excess carbohydrates and fats not immediately needed‬
‭by the body can be converted into long-term storage forms. For example, excess‬
‭carbohydrates can be stored as glycogen in the liver and muscles, and excess triglycerides‬
‭can be stored as fat in adipose tissue. If amino acids are in excess, they aren’t stored as‬
‭proteins but undergo chemical processes to convert them to carbohydrates or fats.‬

‭ igure 6.28‬ ‭Three principal nutrient sources used‬‭to produce ATP (complex‬
F
‭carbohydrates, fats, and proteins). (credit:‬‭OpenStax‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭6.16 Calories‬
I‭ n the preceding discussion, relative energy potential is described by ATP yield. However,‬
‭food labels use the term “Calories” instead of ATP to describe the energy potential in a‬
‭product serving (‬‭Figure 6.29‬‭). How is the number‬‭o f Calories determined? The first‬
‭method, called proximate analysis, uses different chemical methods to determine the‬
‭number of grams of carbohydrates, fat, and protein in a food serving. Then, the total‬
‭Calories are calculated using the conversion of one gram of carbohydrates or protein to‬
‭four Calories, while one gram of fat contains nine Calories. The second method, called‬
‭bomb calorimetry, measures the heat released when a dried food sample is incinerated in‬
‭a device submerged in water (‬‭Figure 6.30‬‭). The amount‬‭o f heat released is estimated by‬
‭the change in water temperature after incineration. A‬‭Calorie‬‭is formally defined as the‬
‭energy required to change the temperature of 1 kg of water by 1 °C.‬

‭ igure 6.29‬ ‭An example of a nutrition facts label.‬‭The number of Calories per serving is‬
F
‭approximately determined by multiplying the grams of carbohydrates, fats, and proteins‬
‭by their respective per-gram Caloric content (4 Calories/gram of carbohydrate and‬
‭proteins; 9 Calories/gram of fat). (credit:‬ ‭BruceBlaus‬‭;‬‭CC BY-SA 4.0‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 6: Energy Considerations‬

‭ igure 6.30‬ ‭Diagram of a bomb calorimeter apparatus‬‭used to determine the Caloric‬

F
‭content of a dried food sample. (credit:‬‭Bomb Calorimeter‬‭Diagram.png‬‭;‬‭Lisdavid89‬‭;‬‭CC‬
‭BY-SA 3.0‬‭).‬‭A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ or additional information about bomb calorimetry, click the link below to watch the video‬
F
‭titled “The science of a calorimeter.”‬

The science of a calorimeter

‭ n average, a person needs 1500 to 2000 Calories daily to sustain daily activities. The total‬
O
‭number of Calories one needs depends on body mass, age, height, gender, activity level,‬
‭and daily exercise. If exercise is a regular part of one’s day, more Calories are required. As‬
‭a rule, people underestimate the number of Calories ingested and overestimate the‬
‭amount they burn through exercise. This error can lead to the ingestion of too many‬
‭Calories per day. The accumulation of an extra 3500 Calories adds one pound of weight. If‬
‭an excess of 200 Calories per day is ingested, one extra pound of body weight will be‬
‭gained every 18 days. An extra 20 pounds can be gained over a year at that rate. Of‬
‭course, this increase in Calories could be offset by increased exercise.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭176‬
‭Chapter 6: Energy Considerations‬

‭ or additional information about Calories, click the link below or scan the QR code to‬
F
‭watch the TED-Ed video by‬‭Emma Bryce titled “What‬‭is a Calorie?”.‬

What is a calorie? - Emma Bryce

‭ or additional information about exercise and metabolism, click the link below or scan the‬
F
‭QR code to watch the TED-Ed video by Dr. Jen Gunter‬‭titled “Can exercise actually boost‬
‭your metabolism?”.‬

Can Exercise Actually "Boost" Your Metabolism? | Body Stuff with…

‭Everyday Connection‬

‭ etabolism and Obesity‬

M
‭Obesity in the United States is epidemic. The rate of obesity has been steadily rising since‬
‭the 1980s. In the 1990s, most states reported that less than 10 percent of their‬
‭populations were obese, and the state with the highest rate reported that only 15 percent‬
‭o f their population was considered obese. By 2010, the U.S. Centers for Disease Control‬
‭and Prevention reported that nearly 36 percent of adults over 20 years old were obese,‬
‭and an additional 33 percent were overweight, leaving only about 30 percent of the‬
‭population at a healthy weight. These studies find the highest levels of obesity are‬
‭concentrated in the southern states. They also find the level of childhood obesity is rising.‬

‭ besity is defined by the body mass index (BMI), which measures a person's weight in‬
O
‭kilograms divided by height in meters. The normal, or healthy, BMI range is between 18‬
‭and 24.9 kg/m‬‭2‬‭. Overweight is defined as a BMI of‬‭25 to 29.9 kg/m‬‭2‬‭, and obesity is‬
‭considered to be a BMI greater than 30 kg/m‬‭2‬‭. Obesity‬‭can arise from several factors,‬
‭including overeating, poor diet, a sedentary lifestyle, limited sleep, genetic factors, and‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 6: Energy Considerations‬

e‭ ven diseases or drugs. Severe obesity (morbid obesity) or long-term obesity can result in‬
‭serious medical conditions, including coronary heart disease; type 2 diabetes;‬
‭endometrial, breast, or colon cancer; hypertension (high blood pressure); dyslipidemia‬
‭(high cholesterol or elevated triglycerides); stroke; liver disease; gallbladder disease; sleep‬
‭apnea or respiratory diseases; osteoarthritis; and infertility. Research has shown that‬
‭losing weight can help reduce or reverse the complications associated with these‬
‭conditions.‬

‭6.17 Disorders of Metabolism - Phenylketonuria and Prader-Willi Syndrome‬

‭ henylketonuria‬‭(‬‭PKU‬‭) is a genetic disease that affects‬‭about 1 in every 15,000 births in‬

P
‭the United States. People afflicted with PKU lack sufficient functional phenylalanine‬
‭hydroxylase, an enzyme that converts‬‭the amino acid‬‭phenylalanine‬‭into tyrosine,‬
‭another amino acid. Because of this enzyme issue, phenylalanine levels rise to toxic levels‬
‭in the body, damaging the central nervous system and brain. Symptoms include delayed‬
‭neurological development, hyperactivity, mental retardation, seizures, skin rash, tremors,‬
‭and uncontrolled movements of the arms and legs. Pregnant women with PKU are at high‬
‭risk for exposing the fetus to too much phenylalanine, which can cross the placenta and‬
‭affect fetal development. Babies exposed to excess phenylalanine in utero may present‬
‭with heart defects, physical and mental retardation, and microcephaly. Every infant in the‬
‭United States and Canada is tested at birth to determine whether PKU is present. The‬
‭earlier a modified diet is begun, the less severe the symptoms will be. To avoid symptoms‬
‭and damage, the person must follow a strict diet low in phenylalanine. Phenylalanine is‬
‭found in high concentrations in artificial sweeteners, including aspartame. Therefore,‬
‭these sweeteners must be avoided. Some animal products and certain starches are also‬
‭high in phenylalanine, and the intake of these foods should be carefully monitored.‬

‭ rader-Willi Syndrome (PWS)‬‭is a genetic disorder‬‭that results in persistent feelings of‬

P
‭intense hunger and reduced rates of metabolism. Typically, affected children have to be‬
‭supervised around the clock to ensure they do not excessively eat. Currently, PWS is the‬
‭leading genetic cause of morbid obesity in children, and it is associated with a number of‬
‭cognitive deficits and emotional problems (‬‭Figure‬‭6.31‬‭).‬

‭ hile genetic testing can be used to diagnose, several behavioral diagnostic criteria are‬
W
‭associated with PWS. From birth to 2 years of age, lack of muscle tone and poor sucking‬
‭behavior may serve as early signs of PWS. Developmental delays are seen between the‬
‭ages of 6 and 12, and excessive eating and cognitive deficits associated with PWS usually‬
‭o nset a little later.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 6: Energy Considerations‬

‭ igure 6.31‬ ‭Eugenia Martínez Vallejo, depicted in‬‭this 1680 painting, may have had‬
F
‭Prader-Willi syndrome. At just eight years old, she weighed approximately 120 pounds‬
‭and was nicknamed “La Monstrua” (the monster). (credit:‬‭OpenStax‬‭)‬

‭ hile the exact mechanisms of PWS are not fully understood, there is evidence that‬
W
‭affected individuals have hypothalamic abnormalities. This observation is not surprising,‬
‭given the hypothalamus’s role in regulating hunger and eating. However, as you will learn‬
‭in the next section of this chapter, the hypothalamus is also involved in regulating sexual‬
‭behavior. Consequently, many individuals suffering from PWS fail to reach sexual maturity‬
‭during adolescence.‬

‭ here is no current treatment or cure for PWS. However, if weight can be controlled in‬
T
‭these individuals, then their life expectancies are significantly increased (historically,‬
‭sufferers of PWS often died in adolescence or early adulthood). Advances in the use of‬
‭various psychoactive medications and growth hormones continue to enhance the quality‬
‭o f life for individuals with PWS.‬

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‭6.18 Career Connection - Registered Dietitian‬

‭ besity is a worldwide health concern, and many diseases, such as diabetes and heart‬
O
‭disease, are becoming more prevalent because of obesity. This fact is one of the reasons‬
‭why people increasingly seek out registered dietitians for advice. Registered dietitians help‬
‭plan nutrition programs for individuals in various settings. They often work with patients‬
‭in healthcare facilities, designing nutrition plans to treat and prevent diseases. For‬
‭example, dietitians may teach a patient with diabetes how to manage blood sugar levels by‬
‭eating the correct types and amounts of carbohydrates. Dietitians may also work in‬
‭nursing homes, schools, and private practices.‬

‭ o become a registered dietitian, one must earn at least a bachelor’s degree in dietetics,‬
T
‭nutrition, food technology, or a related field. In addition, registered dietitians must‬
‭complete a supervised internship program and pass a national exam. Those who pursue‬
‭careers in dietetics take courses in nutrition, chemistry, biochemistry, biology,‬
‭microbiology, and human physiology. Dietitians must become experts in the chemistry and‬
‭physiology (biological functions) of food (proteins, carbohydrates, and fats).‬

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‭Chapter 7: Cardiovascular System I - Blood‬

‭ igure 7.1‬ ‭A single drop of blood contains millions‬‭o f cells. The scanning electron‬
F
‭micrograph shows (from left to right) a red blood cell, platelet, and white blood cell.‬
‭(credit:‬‭OpenStax‬‭)‬

‭7.1 Introduction‬
‭ ur large, complex bodies need blood to deliver nutrients and remove wastes from our‬
O
‭trillions of cells. These tasks are accomplished via the circulatory system, which includes‬
‭three fundamental components: blood, heart, and blood vessels. The heart pumps blood‬
‭throughout the body using a network of blood vessels. This chapter focuses on blood and‬
‭related topics, while the following two focus on the heart and blood.‬
‭ lood, a type of connective tissue, is composed of a‬‭cellular portion‬‭and a‬‭noncellular‬
B
‭portion‬‭called the‬‭matrix‬‭. The components of the‬‭cellular portion, called‬‭formed‬
‭elements‬‭, include‬‭red‬‭blood cells (RBCs)‬‭o r‬‭erythrocyte‬‭s,‬‭white blood cells (WBCs)‬‭o r‬
‭leukocytes‬‭, and cell fragments called‬‭platelets‬‭o r‬‭thrombocytes‬‭. The blood matrix, called‬
‭plasma‬‭, consists mainly of water but also contains‬‭proteins, salts, lipids, glucose, and‬
‭dissolved respiratory gasses (oxygen and carbon dioxide). Plasma suspends the blood‬
‭cells and platelets, which enables them to circulate throughout the body within the vessels.‬
‭ he lifespan of the formed elements is typically relatively brief. Although one type of‬
T
‭leukocyte called memory cells can survive for years, most erythrocytes, leukocytes, and‬
‭platelets usually live only a few hours to a few months. Thus, the body must form new‬

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‭ lood cells and platelets quickly and continuously. When you donate a unit of blood‬
b
‭during a blood drive (approximately 475 mL, or about one pint), your body typically‬
‭replaces the donated plasma within 24 hours, but it takes about four to six weeks to‬
‭replace the blood cells. This situation restricts the possible donation frequency.‬

‭7.2 Characteristics of Blood‬

‭ hen you think about blood, its color is likely the first characteristic that comes to mind.‬
W
‭Blood that has just taken up oxygen in the lungs is bright red, and blood that has released‬
‭oxygen in the tissues is a dusky red with a bluish tint. As discussed later, this is because‬
‭hemoglobin‬‭is a pigment that changes color depending‬‭o n how much oxygen it carries.‬
‭ he average blood temperature is slightly higher than the normal body‬
T
‭temperature—about 38 °C (or 100.4 °F), compared to 37 °C (or 98.6 °F) for an internal‬
‭body temperature reading. However, minimal daily variations are normal. Although the‬
‭surface of blood vessels is relatively smooth, as blood flows through them, it experiences‬
‭some friction and resistance, especially as vessels age and lose their elasticity, which‬
‭produces heat. This phenomenon is the reason for its slightly higher temperature.‬
‭ he pH of blood averages about 7.4; however, it can range from 7.35 to 7.45 in a healthy‬
T
‭person. Blood is more basic (alkaline) on a chemical scale than pure water, with a pH of‬
‭7.0. Blood contains several buffers that help to maintain pH homeostasis.‬
‭ lood constitutes approximately eight percent of adult body weight. Adult males typically‬
B
‭have approximately five to six liters of blood, while females have about four to five liters.‬

‭7.3 Functions of Blood‬

‭ he primary function of blood is to transport oxygen and nutrients to body cells and‬
T
‭remove wastes from them. However, blood’s other functions include defense and‬
‭maintenance of homeostasis.‬

‭Transportation‬
‭ utrients from your food are absorbed in the digestive tract (Chapter 5). Most of these‬
N
‭travel via the bloodstream directly to the liver, where they are processed and released‬
‭back into the bloodstream for delivery to body cells. Oxygen from the air you breathe‬
‭diffuses into the blood at interfaces between blood vessels and lung structures. The‬
‭oxygenated blood returns to the heart and is pumped out to the rest of the body.‬
‭Moreover, endocrine glands scattered throughout the body release their products, called‬

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‭ ormones, into the bloodstream, which carries them to distant target cells. Blood also‬
h
‭picks up cellular wastes and byproducts and transports them to various organs for‬
‭removal. For example, blood moves carbon dioxide to the lungs for exhalation from the‬
‭body, and waste products are transported to the kidneys and liver for excretion from the‬
‭body in the form of urine or bile derivatives.‬

‭Defense‬
‭ umerous types of WBCs protect the body from external threats, such as disease-causing‬
N
‭bacteria that have entered the bloodstream in a wound. Other WBCs seek out and destroy‬
‭internal threats, such as cells with mutated DNA that could multiply to become cancerous‬
‭o r body cells infected with viruses.‬
‭ hen vessel damage results in bleeding, blood platelets and specific proteins dissolved in‬
W
‭the plasma interact to block the ruptured areas of the blood vessels involved. This action‬
‭protects the body from further blood loss.‬

‭Maintenance of Homeostasis‬
‭ ecall that body temperature is regulated via a classic‬‭negative feedback loop‬‭. If you‬
R
‭exercised on a warm day, your rising core body temperature would trigger several‬
‭homeostatic mechanisms, including increased blood transport from your core to your‬
‭body periphery, which is typically cooler. As blood passes through the skin's vessels, heat‬
‭would dissipate to the environment, and the blood returning to your body's core would be‬
‭cooler. In contrast, blood is diverted from the skin on a cold day to maintain a warmer‬
‭body core. In extreme cases, this may result in frostbite.‬
‭ lood also helps to maintain the body's chemical balance. Proteins and other compounds‬
B
‭in the blood act as buffers, regulating the pH of body tissues and the water content of‬
‭body cells.‬

‭7.4 Composition of Blood‬

‭ ou have probably had blood drawn from a vein in your arm, which was then sent to a lab‬
Y
‭for analysis. Some of the most common blood tests—for instance, those measuring lipid or‬
‭glucose levels in plasma—determine which substances are present within blood and in‬
‭what quantities. Other blood tests check for the composition of the blood itself, including‬
‭the amounts and types of cells.‬

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‭ ne such test, called‬‭hematocrit‬‭, measures the percentage of erythrocytes in a blood‬

O
‭sample. It is performed by spinning the blood sample in a specialized‬‭centrifuge‬‭, which‬
‭causes the heavier elements suspended within the blood sample to separate from the‬
‭lightweight, liquid plasma (‬‭Figure 7.2‬‭). Because the‬‭densest formed elements in blood are‬
‭the erythrocytes, these settle at the very bottom of the centrifuged tube. A pale, thin layer‬
‭composed of the remaining formed elements of blood is located above the erythrocytes.‬
‭These are the leukocytes and the platelets. This layer is called the‬‭buffy coat‬‭because of its‬
‭color; it typically constitutes less than 1 percent of a blood sample. Above the buffy coat is‬
‭the blood plasma, usually a pale, straw-colored fluid comprising the remainder of the‬
‭sample.‬

‭ igure 7.2‬ ‭The cellular elements of blood include‬‭a vast number of erythrocytes (RBC)‬
F
‭and comparatively fewer leukocytes (WBC) and thrombocytes (platelets). Plasma is the‬
‭fluid in which the cells and platelets are suspended. A blood sample spun in a centrifuge‬
‭reveals that plasma is the lightest component. It floats at the top of the tube, separated‬
‭from the heaviest elements, the erythrocytes, by a buffy coat of leukocytes and platelets.‬
‭Hematocrit is the percentage of the total sample that is composed of erythrocytes.‬
‭Depressed and elevated hematocrit levels are shown for comparison. (credit:‬‭OpenStax‬‭).‬
‭A link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

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‭7.5 Erythrocytes‬
‭ rythrocytes are the most common formed element: A single drop of blood contains‬
E
‭millions of erythrocytes and just thousands of leukocytes. Specifically, males have about‬
‭5.4 million erythrocytes per microliter (‬‭µ‬‭L) of blood,‬‭and females have approximately 4.8‬
‭million per‬‭µ‬‭L. Erythrocytes are estimated to make‬‭up about 25 percent of the total cells in‬
‭the body. As you can imagine, they are microscopic cells with a mean diameter of only‬
‭about 7–8 micrometers (‬‭µ‭m ‬ ) (Figure 7.3). The primary‬‭functions of erythrocytes are to‬
‭pick up inhaled oxygen from the lungs, transport it to the body’s tissues, pick up some‬
‭(about 24 percent) carbon dioxide waste in the tissues, and transport it to the lungs for‬
‭exhalation. Erythrocytes usually remain within the blood vessels. However, leukocytes‬
‭typically leave the blood vessels to perform their defensive functions.‬

‭Shape and Structure of Erythrocytes‬

‭ s an erythrocyte matures in the red bone marrow, it loses its nucleus and most of its‬
A
‭o ther organelles. As these cells lack mitochondria, they must rely on anaerobic‬
‭metabolism/fermentation for ATP (Chapter 6). This means mature erythrocytes do not‬
‭utilize any of the oxygen they are transporting, so they can deliver it all to the tissues.‬
‭ rythrocytes are‬‭biconcave disks‬‭, plump at their periphery and very thin in the center‬
E
‭(‬‭Figure 7.3‬‭). Since they lack most organelles, there is more interior space for the presence‬
‭o f the hemoglobin molecules that transport gasses, as you will see shortly. The biconcave‬
‭shape also provides a greater surface area where gas exchange can occur relative to its‬
‭volume.‬
‭ apillaries‬‭are incredibly narrow, slowing the passage‬‭o f the erythrocytes and providing‬
C
‭an extended opportunity for gas exchange. However, the space within capillaries can be so‬
‭small that, despite their small size, erythrocytes may have to fold in on themselves to make‬
‭their way through. Fortunately, their structural proteins are flexible, allowing them to‬
‭bend over themselves to a surprising degree and then spring back again when they enter‬
‭a wider vessel.‬

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‭ igure 7.3‬ ‭Erythrocytes, called red blood cells,‬ ‭are biconcave discs with very shallow‬
F
‭centers. This shape optimizes the surface area-to-volume ratio, facilitating gas exchange. It‬
‭also enables them to fold up as they move through narrow blood vessels. (Credit:‬
‭OpenStax‬‭)‬

‭Hemoglobin‬
‭ emoglobin is a large molecule consisting of proteins and iron (an element). It consists of‬
H
‭four folded chains of a protein called‬‭globin‬‭, with‬‭two of the chains designated alpha (α1‬
‭and α2) and the other two as beta (β1 and β2;‬‭Figure‬‭7.4a‬‭). Each of these globin‬
‭molecules is bound to a red pigment molecule called‬‭heme‬‭, which contains an ion of iron‬
‭(Fe‬‭+2‬‭;‬‭Figure 7.4a‬‭and‬‭b‬‭).‬
‭ ach iron ion in the heme can bind to one oxygen molecule (i.e., O‬‭2‭)‬ ; therefore, each‬
E
‭hemoglobin molecule can transport four oxygen molecules. An individual erythrocyte may‬
‭contain about 300 million hemoglobin molecules and, thus, can bind to and transport up‬
‭to 1.2 billion oxygen molecules.‬
I‭ n the capillaries associated with the lungs, hemoglobin in the RBCs picks up oxygen,‬
‭which binds to the iron ions, forming‬‭oxyhemoglobin‬‭.‬‭The bright red, oxygenated‬
‭hemoglobin travels to the body tissues, releasing some oxygen molecules, becoming‬
‭darker red‬‭deoxyhemoglobin‬‭. Oxygen release depends‬‭o n the need for oxygen in the‬
‭surrounding tissues, so hemoglobin rarely, if ever, leaves all of its oxygen behind.‬

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‭ igure 7.4‬ ‭(a) A hemoglobin molecule contains four‬‭globin proteins, each bound to one‬
F
‭molecule of the iron-containing pigment heme. (b) A single erythrocyte can hold 300‬
‭million hemoglobin molecules and, thus, more than 1 billion oxygen molecules. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

I‭ n the capillaries, carbon dioxide enters the bloodstream. About 75% dissolves in the‬
‭plasma, forming‬‭bicarbonate ions‬‭. The remaining 25%‬‭binds to the amino acids in‬
‭hemoglobin, forming a molecule known as‬‭c arbaminohemoglobin‬‭.‬‭The hemoglobin‬
‭carries carbon dioxide from the capillaries back to the lungs, releasing it to pick up‬
‭oxygen.‬
I‭ n patients with insufficient hemoglobin, the tissues may not receive sufficient oxygen,‬
‭resulting in anemia, a condition discussed later in the chapter. In determining the‬
‭oxygenation of tissues, the value of the most significant interest in healthcare is the‬
‭percent saturation‬‭; that is, the percentage of hemoglobin‬‭sites occupied by oxygen in a‬
‭patient’s blood. Clinically, this value is commonly referred to simply as “percent sat.”‬
‭ ercent saturation is typically monitored using a device known as a pulse oximeter, which‬
P
‭is applied to a thin part of the body, usually the tip of the patient’s finger. The device sends‬
‭two different wavelengths of light (one red, the other infrared) through the finger and‬
‭measures the light with a photodetector as it exits. Hemoglobin absorbs light differentially‬
‭depending upon its saturation with oxygen. The machine calibrates the amount of light the‬
‭photodetector receives against the amount absorbed by the partially oxygenated‬

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‭ emoglobin and presents the data as percent saturation. Standard pulse oximeter‬
h
‭readings range from 95–100 percent. Lower percentages reflect‬‭hypoxemia‬‭o r low blood‬
‭oxygen. The term hypoxia is more generic and simply refers to low oxygen levels.‬
I‭ n response to hypoxemia, less oxygen will exit the vessels supplying the kidney, resulting‬
‭in hypoxia in the tissue fluid of the kidney, where oxygen concentration is monitored. Cells‬
‭within the kidney secrete a hormone called‬‭erythropoietin‬‭(‬‭EPO‬‭), which increases‬
‭erythrocyte production and restores oxygen levels.‬

I‭ n a classic negative feedback loop, as oxygen saturation rises, EPO secretion falls, and vice‬
‭versa, thereby maintaining homeostasis. Populations dwelling at high elevations, with‬
‭inherently lower oxygen levels in the atmosphere, naturally maintain a hematocrit higher‬
‭than people living at sea level. Consequently, people traveling to high elevations may‬
‭experience symptoms of hypoxemia, such as fatigue, headache, and shortness of breath,‬
‭for a few days after their arrival. In response to hypoxemia, the kidneys secrete EPO to‬
‭step up the production of erythrocytes until homeostasis is achieved once again. To avoid‬
‭the symptoms of hypoxemia, also known as altitude sickness, mountain climbers typically‬
‭rest for several days to a week or more at a series of camps situated at increasing‬
‭elevations to allow EPO levels and erythrocyte counts to rise.‬
‭ PO is a banned substance in most organized sports. Still, it is also used medically to treat‬
E
‭certain anemia, specifically those triggered by certain types of cancer and other disorders‬
‭in which increased erythrocyte counts and oxygen levels are desirable.‬

‭Everyday Connection‬

‭ lood Doping‬
B
‭In its original intent, the term blood doping was used to describe the practice of injecting‬
‭supplemental RBCs by transfusion into an individual, typically to enhance performance in‬
‭a sport. Additional RBCs would deliver more oxygen to the tissues, providing extra aerobic‬
‭capacity, clinically called VO‬‭2‬ ‭max. The source of‬‭the cells was either from the recipient‬
‭(autologous) or from a donor with compatible blood (homologous).‬

‭ his practice was aided by the well-developed harvesting, concentrating, and freezing‬
T
‭techniques of the RBCs that could be later thawed and injected yet still retain their‬
‭functionality. These practices are considered illegal in virtually all sports and run the risk‬
‭o f infection, significantly increasing the viscosity of the blood and the potential for‬
‭transmission of blood-borne pathogens if the blood is collected from another individual.‬

‭ ith the development of synthetic EPO in the 1980s, it became possible to provide‬
W
‭additional RBCs by artificially stimulating RBC production in the bone marrow. Originally‬

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‭ eveloped to treat patients suffering from anemia, renal failure, or cancer treatment,‬
d
‭recombinant DNA technology can generate large quantities of EPO. Synthetic EPO is‬
‭injected under the skin and can increase hematocrit for many weeks. It may also induce‬
‭polycythemia and raise hematocrit to 70 or greater. This increased viscosity raises the‬
‭resistance of the blood and forces the heart to pump more powerfully; in extreme cases, it‬
‭has resulted in death. Other drugs, such as cobalt II chloride, have been shown to increase‬
‭natural EPO gene expression. Blood doping has become problematic in many sports,‬
‭especially cycling. Lance Armstrong, the winner of seven Tour de France and many other‬
‭cycling titles, was stripped of his victories and admitted to blood doping in 2013.‬

‭Life Cycle of Erythrocytes‬

‭ rythrocyte production in the marrow occurs at a staggering rate of more than two‬
E
‭million cells per second. For this production to occur, several raw materials must be‬
‭present in adequate amounts. These include the same nutrients essential to the‬
‭production and maintenance of any cell, such as glucose, lipids, and amino acids. However,‬
‭erythrocyte production also requires several trace elements, such as iron.‬
‭ rythrocytes live up to 120 days in circulation, after which the worn-out cells are removed‬
E
‭by a type of cell called a‬‭macrophage‬‭, located primarily‬‭within the bone marrow, liver, and‬
‭spleen. The components of the degraded erythrocytes’ hemoglobin are further processed‬
‭as follows:‬
‭•‬ ‭ lobin, the protein portion of hemoglobin, is broken down into amino acids, which‬
G
‭can be sent back to the bone marrow to produce new erythrocytes.‬

‭•‬ ‭ he iron contained in the heme portion of hemoglobin may be stored in the liver or‬
T
‭spleen or carried through the bloodstream to the red bone marrow for recycling into‬
‭new erythrocytes.‬

‭•‬ ‭ he non-iron portion of heme is degraded into waste products that are either moved‬
T
‭to the liver and used as a component of bile (Chapter 5) or eliminated via feces or‬
‭urine.‬

‭Disorders of Erythrocytes‬
‭ he size, shape, and number of erythrocytes and the number of hemoglobin molecules‬
T
‭can majorly impact a person’s health.‬‭Anemia‬‭is the‬‭general condition when the number‬
‭o f RBCs or hemoglobin is deficient. More than 400 types of anemia exist, and more than‬

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‭3.5 million Americans suffer from this condition.‬

‭ nemias are divided into three major groups: those caused by blood loss, those caused by‬
A
‭faulty or decreased RBC production, and those caused by excessive destruction of RBCs.‬
‭The effects of the various anemias are widespread because reduced numbers of RBCs or‬
‭hemoglobin will result in lower levels of oxygen delivered to body tissues. Since oxygen is‬
‭required for cellular respiration and ATP production, anemia produces fatigue, lethargy,‬
‭and an increased risk of infection. An oxygen deficit in the brain impairs the ability to‬
‭think clearly and may prompt headaches and irritability.‬
‭ lood loss anemias are relatively straightforward. In addition to bleeding from wounds or‬
B
‭o ther lesions, these forms of anemia may be due to ulcers, hemorrhoids, inflammation of‬
‭the stomach (gastritis), and some gastrointestinal tract cancers. The excessive use of‬
‭aspirin or other nonsteroidal anti-inflammatory drugs, such as ibuprofen, can trigger‬
‭ulceration and gastritis. Excessive menstruation and loss of blood during childbirth are‬
‭also potential causes.‬
‭ nemias caused by faulty or decreased RBC production include sickle cell anemia, iron‬
A
‭deficiency anemia, vitamin deficiency anemia, and bone marrow and stem cell diseases.‬
‭•‬ ‭ characteristic change in the shape of erythrocytes is seen in‬‭sickle cell anemia‬‭.‬
A
‭This genetic disorder is caused by the production of an abnormal type of hemoglobin,‬
‭called hemoglobin S, which delivers less oxygen to tissues and causes erythrocytes to‬
‭assume a sickle (or crescent) shape, especially at low oxygen concentrations (‬‭Figure‬
‭7.5‬‭). These abnormally shaped cells can then become‬‭lodged in narrow capillaries‬
‭because they cannot fold in to squeeze through, blocking blood flow to tissues and‬
‭causing various serious problems, from painful joints to delayed growth and even‬
‭blindness and strokes. Sickle cell anemia is a genetic condition mainly observed in‬
‭individuals of African descent.‬

‭•‬ I‭ ron deficiency anemia‬‭is the most common type and‬‭results when the amount of‬
‭available iron is insufficient to allow the production of sufficient heme, a component‬
‭o f hemoglobin. This condition can occur in individuals with a diet deficient in iron and‬
‭is especially common in teens and children, as well as in vegans and vegetarians. Iron‬
‭deficiency anemia may be caused by either an inability to absorb and transport iron‬
‭o r slow, chronic bleeding.‬

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‭ igure 7.5‬ ‭Sickle cell anemia is caused by a mutation‬‭in one of the hemoglobin genes.‬
F
‭Erythrocytes produce an abnormal type of hemoglobin, which causes the cell to take on a‬
‭sickle or crescent shape. (credit: Janice Haney Carr;‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ or additional information about sickle cell anemia, click the link below or scan the QR‬
F
‭code to watch the TED-Ed video by Amber M. Yates‬‭titled‬‭“How this disease changes the‬
‭shape of your cells.”‬

How this disease changes the shape of your cells - Amber M. Yates

‭•‬ ‭ itamin deficiency anemias‬‭generally involve insufficient‬‭vitamin B12 and folate.‬

V
‭Pernicious anemia is caused by poor vitamin B12 absorption and is often seen in‬
‭patients with Crohn’s disease (a severe intestinal disorder often treated by surgery),‬
‭surgical removal of the intestines or stomach (common in some weight loss‬
‭surgeries), intestinal parasites, and AIDS.‬

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‭•‬ ‭ arious disease processes can also interfere with the production and formation of‬
V
‭RBCs and hemoglobin:‬

‭–‬ ‭ halassemia‬‭is an inherited condition typically found‬‭in individuals from the‬

T
‭Middle East, the Mediterranean, Africa, and Southeast Asia. In this condition,‬
‭the maturation of the RBCs does not proceed normally.‬

‭–‬ ‭ plastic anemia‬‭is a condition that results from insufficient‬‭numbers of RBC‬

A
‭stem cells. Aplastic anemia is often inherited or triggered by radiation,‬
‭medication, chemotherapy, or infection.‬

‭–‬ ‭ ead exposure from industrial sources or dust from paint chips of‬
L
‭iron-containing paints or pottery that has not been correctly glazed may also‬
‭destroy the red marrow.‬
I‭ n contrast to anemia, an elevated erythrocyte count is called‬‭polycythemia‬‭and is‬
‭detected in a patient’s elevated hematocrit. It can occur temporarily in a dehydrated‬
‭person; the plasma volume falls when inadequate water intake or excessive water losses.‬
‭As a result, the hematocrit rises. For reasons mentioned earlier, a mild form of‬
‭polycythemia is chronic but expected in people living at high altitudes. Some elite athletes‬
‭train at high elevations specifically to induce this phenomenon. Finally, a type of bone‬
‭marrow disease called‬‭polycythemia vera‬‭(from the‬‭Greek vera = “true”) causes excessive‬
‭production of immature erythrocytes. Polycythemia vera can dangerously elevate blood‬
‭viscosity, raising blood pressure and making it more difficult for the heart to pump blood‬
‭throughout the body. It is a relatively rare disease that occurs more often in men than‬
‭women and is more likely to be present in elderly patients over 60 years of age.‬

‭7.6 Leukocytes‬
‭ eukocytes (white blood cells; WBC) make up approximately 1% of the volume of the cells‬
L
‭in the blood. The primary role of leukocytes is very different from that of red blood cells;‬
‭they are primarily involved in the immune response to identify and target pathogens, such‬
‭as invading bacteria, viruses, and other foreign organisms. White blood cells are formed‬
‭continually; some only live for hours or days, but some types, called memory cells, live for‬
‭years.‬
‭ hite blood cells differ significantly from red blood cells in appearance. They have nuclei‬
W
‭and organelles but don’t contain hemoglobin. The different types of white blood cells are‬
‭identified by their appearance after staining, which involves adding special dyes to the‬
‭cells and examining them with a microscope (‬‭Figure‬‭7.6‬‭).‬

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‭ ne of the most distinctive characteristics of leukocytes is their movement. Whereas‬

O
‭erythrocytes spend their days circulating within the blood vessels, leukocytes routinely‬
‭leave the bloodstream to perform defensive functions in the body’s tissues. For‬
‭leukocytes, the vascular network is simply a highway they travel and soon exit to reach‬
‭their final destination.‬
‭ nce they have exited the capillaries, some leukocytes will take up fixed positions in bone‬
O
‭marrow, the spleen, the thymus, or other organs. Others will move about through the‬
‭tissue spaces very much like amoebas (small, single-celled organisms), continuously‬
‭extending their plasma membranes, sometimes wandering freely, and sometimes moving‬
‭toward the direction in which chemical signals draw them. The attracting of leukocytes is‬
‭caused by injured or infected cells and nearby leukocytes releasing the equivalent of a‬
‭chemical “911” call, attracting more leukocytes to the site.‬

‭,‬
‭ igure 7.6‬‭(a) A lobe-shaped nucleus and the presence‬‭o f stained granules in the‬
F
‭cytoplasm characterize granulocytes. (b) Agranulocytes lack stained granules in their‬
‭cytoplasm and include lymphocytes and monocytes. (credit:‬‭OpenStax‬‭)‬

‭ s was the case with RBC, excessive or diminished numbers of WBC are associated with‬
A
‭specific diseases. Elevated WBC numbers are associated with diseases such as‬‭lymphoma‬
‭(in the lymphatic system) and‬‭leukemia‬‭(in the bone‬‭marrow). Some forms of lymphoma‬
‭and leukemia progress slowly and respond well to treatment. Others tend to progress‬
‭quickly and require aggressive treatment, without which they are rapidly fatal. For‬
‭additional information about leukemia, click the link or scan the QR code to watch the‬
‭TED-Ed video by Danilo Allegra and Dania Puggioni‬‭titled “What is leukemia?”.‬

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What is leukemia? - Danilo Allegra and Dania Puggio…

‭ eukopenia describes conditions that cause a decrease in the number of WBCs due to‬
L
‭illness, immunosuppression, or other factors. If too few leukocytes exist in an individual,‬
‭the person is more susceptible to various infections.‬

‭7.7 Platelets (Thrombocytes) and Clotting Factors‬

‭ latelets are not cells but membrane-surrounded fragments (2-4 μm in diameter) of the‬
P
‭cytoplasm of a megakaryocyte cell (‬‭Figure 7.7‬‭). These‬‭fragments contain small vesicles‬
‭but don’t have nuclei or other organelles. Each megakaryocyte releases 2000–3000‬
‭platelets during its lifespan. Following platelet release, megakaryocyte remnants, which‬
‭are little more than a cell nucleus, are consumed by macrophages.‬

‭ igure 7.7‬ ‭(a) Platelets are small cell fragments‬‭created from a megakaryocyte. (b)‬
F
‭Platelets and fibrin form a plug, or clot, to help temporarily close a hole in a blood vessel‬
‭until it is fully repaired. (credit:‬‭OpenStax‬‭)‬

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‭ lood must‬‭c lot‬‭to heal wounds and prevent excess‬‭blood loss. Platelets are attracted to‬
B
‭the wound site, where they adhere by extending many projections and releasing their‬
‭contents. These contents activate other platelets and interact with other factors, which‬
‭convert‬‭fibrinogen‬‭, a water-soluble protein in blood‬‭serum, into‬‭fibrin‬‭(a non-water‬
‭soluble protein), causing the blood to clot. Many clotting factors require‬‭vitamin K‬‭to‬
‭work, and vitamin K deficiency can lead to problems with blood clotting. For additional‬
‭information about blood clotting and wound healing, click the link below or scan the QR‬
‭code to watch the TED-Ed video by Sarthak Sinah‬‭titled‬‭“How a wound heals itself.”‬

How a wound heals itself - Sarthak Sinha

‭ hrombocytosis is a condition caused by too many platelets, which may trigger the‬
T
‭formation of unwanted blood clots (‬‭thrombosis‬‭), a‬‭potentially fatal disorder. If there are‬
‭insufficient platelets, called thrombocytopenia, blood may not clot properly, and excessive‬
‭bleeding may result from even minor wounds.‬
‭ dditional substances, collectively called‬‭c lotting‬‭factors‬‭, must be present for clots to‬
A
‭form. Another reason for the failure of blood to clot is the inadequate production of‬
‭functional amounts of one or more of these factors.‬‭Hemophilia‬‭is a genetic disease, and‬
‭the most common form of it is hemophilia A, which accounts for approximately 80% of‬
‭cases. Patients with hemophilia bleed from even minor internal and external wounds and‬
‭leak blood into joint spaces after exercise and into urine and stool. Regular infusions of‬
‭clotting factors isolated from healthy donors can help prevent bleeding in hemophilia‬
‭patients.‬
‭ mong aspirin's many known biochemical activities is its role as an anticoagulant. Aspirin‬
A
‭is very effective at inhibiting the aggregation of platelets. It is routinely administered‬
‭during a heart attack or stroke to reduce the adverse effects. Physicians sometimes‬
‭recommend that patients at risk for cardiovascular disease take a low dose of aspirin daily‬
‭as a preventive measure. However, aspirin can also lead to serious side effects, including‬
‭increasing the risk of ulcers. A patient must consult a physician before beginning any‬
‭aspirin regimen.‬

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‭7.8 Plasma Proteins and Other Solutes‬

‭ bout 7% of the plasma volume—nearly all that is not water—consists of proteins. The‬
A
‭three primary categories of plasma proteins (proteins unique to the plasma) are as‬
‭follows:‬
‭•‬ ‭ lbumin‬‭is the most abundant of the plasma proteins.‬‭The liver manufactures‬
A
‭albumin molecules, which function as binding proteins and transport vehicles for‬
‭fatty acids and steroid hormones. Recall that lipids are hydrophobic; however, their‬
‭binding to albumin enables their transport in the watery plasma. Albumin is also the‬
‭most significant contributor to the osmotic pressure of blood; that is, its presence‬
‭holds water inside the blood vessels and draws water from the tissues, across blood‬
‭vessel walls, and into the bloodstream. This, in turn, helps to maintain both blood‬
‭volume and blood pressure.‬

‭•‬ ‭ lobulin‬‭is the second most common plasma protein.‬‭One type, called gamma‬
G
‭globulins, is associated with the immune system and is better known as antibodies or‬
‭immunoglobulins.‬

‭•‬ ‭ ibrinogen is the least abundant plasma protein, and it is associated with blood‬
F
‭clotting, as previously discussed.‬

I‭ n addition to proteins, plasma contains a wide variety of other substances, including‬

‭electrolytes (e.g., sodium, potassium, and calcium ions), dissolved gasses (e.g., oxygen and‬
‭carbon dioxide), various nutrients (e.g., vitamins, lipids, glucose, and amino acids), and‬
‭metabolic wastes (e.g., ammonia and urea). These nonprotein solutes combined contribute‬
‭approximately one percent to the total plasma volume.‬

‭7.9 Bone Marrow Sampling and Transplants‬

S‭ ometimes, a healthcare provider will order a bone marrow biopsy, a diagnostic test of a‬
‭sample of‬‭red bone marrow‬‭, or a bone marrow transplant,‬‭a treatment in which a‬
‭donor’s healthy bone marrow—and its stem cells—replaces the faulty bone marrow of a‬
‭patient. These tests and procedures are often used to assist in the diagnosis and treatment‬
‭o f various severe forms of anemia, such as thalassemia major and sickle cell anemia, as‬
‭well as some types of cancer, specifically leukemia.‬
I‭ n the past, when a bone marrow sample or transplant was necessary, the procedure‬
‭would have required inserting a large-bore needle into the pelvic bones (‬‭Figure 7.8‬‭). This‬
‭location was preferred because it is close to the body surface, making it more accessible,‬

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a‭ nd it is relatively isolated from most vital organs. Unfortunately, the procedure is quite‬
‭painful.‬
‭ ow, direct sampling of bone marrow can often be avoided. In many cases, stem cells are‬
N
‭isolated in just a few hours from a sample of a patient’s blood.‬

‭ igure 7.8‬ ‭Bone marrow being removed via aspiration‬‭from the pelvic bones. (credit:‬
F
‭Bone marrow aspiration.jpg‬‭;‬‭Andrew Ratto‬‭; WikiMedia‬‭Commons;‬‭CC BY 2.0‬‭).‬

‭ or an individual requiring a‬‭bone marrow transplant‬‭,‬‭a matching donor is essential to‬

F
‭prevent the immune system from destroying the donor cells—a phenomenon known as‬
‭tissue rejection. To treat patients with these transplants, it is first necessary to eliminate‬
‭the patient’s diseased marrow through radiation or chemotherapy. Stem cells in the donor‬
‭bone marrow are then intravenously infused into the recipient, where they will travel via‬
‭the bloodstream and establish themselves in the bone marrow.‬

‭7.10 The Genetic Basis of the ABO and Rh Blood Types‬

‭ hen some people hear the phrase “blood type,” four terms come to mind: Types A, B, AB,‬
W
‭and O. But what do these letters represent? To adequately answer this, we need to review‬
‭some basic genetic concepts.‬

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I‭ n Chapter 4, you learned that chromosomes are structures made of DNA wrapped‬
‭around histone proteins (i.e., nucleosomes). A human somatic cell (i.e., nonreproductive‬
‭cell) has 23 pairs of chromosomes, with one chromosome of each pair from each parent.‬
‭Genes‬‭are segments of DNA that contain information‬‭to make something, such as a‬
‭protein, via a two-step process (transcription and translation).‬
‭ umans have approximately 18,500 genes scattered over the 22 homologous pairs of‬
H
‭autosomal chromosomes and the X and Y sex chromosomes. You have two copies for‬
‭most genes, one from each of your parents. Suppose a particular gene is located on‬
‭chromosome 15. In that case, the first copy will be present on chromosome 15,‬
‭contributed by the mother, and the second copy will be on chromosome 15, contributed‬
‭by the father. Fertilization resulted in both copies being present in the same cell.‬
‭ gene that influences a particular trait, such as blood type, can have more than one form;‬
A
‭the different forms are called‬‭alleles‬‭. Alleles of‬‭a gene differ in some way concerning the‬
‭nucleotide base sequence (i.e., the order of A, T, G, and C bases) on a strand. Therefore,‬
‭transcription and translation will result in proteins with different amino acid sequences‬
‭and possibly different expressions. In the case of blood type, there are three alleles‬
‭designated‬‭A‭,‬‬‭B‭,‬ and‬‭o‬‭. The reason the‬‭o‬‭allele is‬‭lowercase will be explained shortly.‬
‭ he combination of two alleles for a given gene in a diploid organism is called its‬
T
‭genotype‬‭. The observable traits that result from‬‭the expression of these alleles are called‬
‭phenotypes‬‭. For example, if an individual has two‬‭alleles associated with the disease‬
‭sickle cell anemia (the genotype), then their erythrocytes will be sickle-shaped (the‬
‭phenotype).‬
I‭ n the case of blood type, one gene, located on chromosome 9, is responsible for‬
‭determining the phenotype. You possess two alleles for this gene, one obtained from your‬
‭mother via her chromosome and the other from your father via his chromosome.‬
‭ s there are three alleles for this gene and each person has two alleles, there are six‬
A
‭possible genotypes:‬‭AA‬‭,‬‭AB‬‭,‬‭Ao‬‭,‬‭BB‬‭,‬‭Bo‬‭, and‬‭oo‬‭. If‬‭both alleles are the same, the genotype is‬
‭said to be‬‭homozygous‬‭(i.e.,‬‭AA‬‭,‬‭BB‬‭,‬‭oo‬‭); if the two‬‭alleles are different, then it is classified‬
‭as‬‭heterozygous‬‭(i.e.,‬‭AB‬‭,‬‭Ao‬‭,‬‭Bo‬‭).‬
I‭ n the case of blood type, the phenotype is determined only by the genotype of this gene.‬
‭However, multiple genes and non-genetic factors (e.g., diet, health, etc.) are involved in‬
‭determining the phenotype for other traits, such as height and weight.‬
‭ lleles of a gene interact to determine the phenotype. Two possible allelic interactions are‬
A
‭dominant and‬‭recessive. If an allele is‬‭dominant‬‭,‬‭its expression will mask/hide the‬
‭expression of another allele, which is said to be‬‭recessive‬‭. Dominant alleles are frequently‬
‭represented by upper case letters (e.g.,‬‭A‬‭and‬‭B‭)‬ ,‬‭and recessive alleles by lower case‬

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l‭etters (e.g.,‬‭o‭)‬ . Therefore, if the genotype contains an‬‭A‬‭o r‬‭B‬‭allele with an‬‭o‬‭allele, then‬
‭the‬‭A‬‭o r‬‭B‬‭allele will determine the phenotype (e.g.,‬‭type A or B). The O phenotype is only‬
‭o bserved when an individual has the homozygous genotype for the‬‭o‬‭allele (i.e.,‬‭oo‬‭).‬
‭Please note that alleles are italicized, but the phenotype isn’t.‬
‭ he interaction between the‬‭A‬‭and‬‭B‬‭alleles is considered‬‭codominant‬‭, meaning neither‬
T
‭o f these alleles dominates the other one, and both are expressed in the phenotype.‬
‭Therefore, if the blood type genotype is‬‭AB‬‭, the phenotype‬‭is AB.‬
‭ hat is the function of the proteins produced by the expression of the‬‭A‬‭and‬‭B‬‭alleles of‬
W
‭the blood type gene? The answer is they function as enzymes, called glycosyltransferases,‬
‭to add sugars to proteins embedded in the RBC membrane. The transferase produced by‬
‭the expression of the‬‭A‬‭allele differs from that produced‬‭by the‬‭B‬‭allele by a small number‬
‭o f amino acids. These differences result in the two transferases adding different sugars to‬
‭the membrane proteins. The transferase produced by the expression of the‬‭o‬‭allele is not‬
‭functional, and no additional sugar is added.‬
‭ hese carbohydrate-protein complexes function as “identifying flags” of sorts, called‬
T
‭antigens‬‭, for the immune system and help distinguish‬‭between self and non-self or‬
‭foreign. In other words, antigens provide a way for the immune system to identify and‬
‭destroy non-native cells, such as bacteria or viruses in the body, that aren’t supposed to‬
‭be present because of their non-self or foreign antigens. If a person’s blood type‬
‭genotype has an‬‭A‬‭allele, then the RBC will have the‬‭A antigen; if the‬‭B‬‭allele is present, the‬
‭B antigen will be present. If both the‬‭A‬‭and‬‭B‬‭alleles‬‭are present, then both A and B‬
‭antigens will be present. If the genotype is homozygous for the o allele (i.e.,‬‭oo‬‭), neither A‬
‭nor B antigens will be present (‬‭Figure 7.9‬‭). In other‬‭words, type O blood is characterized‬
‭by the absence of A and B antigens, not the presence of an O antigen.‬
‭ second group of blood antigens is the‬‭Rh group‬‭,‬‭first discovered in a type of primate‬
A
‭called a rhesus macaque. Although dozens of Rh antigens have been identified, only one,‬
‭designated D, is clinically significant. This antigen is a protein, not a carbohydrate, as in‬
‭the ABO system. Individuals that have at least one copy of the functional‬‭Rh D‬‭allele (i.e.,‬
‭Rh D‬‭+‬‭/‬‭Rh D‬‭+‬‭o r‬‭Rh D‬‭+‬‭/‬‭Rh D‬‭-‬‭) have the Rh D protein/antigen‬‭in the RBC membrane and are‬
‭phenotypically classified as Rh+; Individuals without a functional allele don’t have the‬
‭protein and are phenotypically classified as Rh-. In other words, the‬‭Rh D‬‭+‬ ‭allele‬
‭dominates the‬‭Rh D‬‭-‬ ‭allele.‬

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‭ igure 7.9‬ ‭A summary of the blood type groups (A,‬‭B, AB, and O) and their associated‬
F
‭antigens. (credit:‬‭ABO blood type.svg‬‭; InvictaHOG;‬‭WikiMedia Commons;‬‭CC0 1.0‬‭).‬ ‭A link‬
‭to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ ote that the Rh group is distinct from the ABO group, so that any individual may have or‬
N
‭lack this Rh antigen, no matter their ABO blood type. When identifying a patient’s ABO and‬
‭Rh blood type, the Rh group is designated by adding the word positive or negative (i.e.,‬‭Rh‬
‭positive‬‭and‬‭Rh negative‬‭) to the ABO type. For example,‬‭A positive (A‬‭+‭)‬ means ABO group‬
‭A blood with the Rh antigen present, and AB negative (AB‬‭−‭)‬ means ABO group AB blood‬
‭without the Rh antigen.‬

‭ BO and Rh phenotype classifications for four ethnic groups in the United States are‬
A
‭presented in‬‭Figure 7.10‬‭. Note the variation in the‬‭frequencies of the blood types among‬
‭these groups. For example, 25% of Asian Americans have blood type B+, while the‬
‭percentage among African, Caucasian, and Latino/Latina Americans is 18%, 9%, and 9%,‬
‭respectively.‬

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‭ lood‬
B ‭Caucasian-‬ ‭ atino/Latina-‬
L
‭Type‬ ‭African-Americans‬ ‭Asian-Americans‬ A‭ mericans‬ ‭Americans‬

‭A‭+‬ ‬ ‭24‬ ‭27‬ ‭33‬ ‭29‬

‭A‭−‬ ‬ ‭2‬ ‭0.5‬ ‭7‬ ‭2‬
‭+‬
‭B‬ ‭18‬ ‭25‬ ‭9‬ ‭9‬
‭B‭−‬ ‬ ‭1‬ ‭0.4‬ ‭2‬ ‭1‬
‭+‬
‭AB‬ ‭4‬ ‭7‬ ‭3‬ ‭2‬
‭−‬
‭AB‬ ‭0.3‬ ‭0.1‬ ‭1‬ ‭0.2‬
‭O‭+‬ ‬ ‭47‬ ‭39‬ ‭37‬ ‭53‬
‭−‬
‭O‬ ‭4‬ ‭1‬ ‭8‬ ‭4‬

‭ igure 7.10‬ ‭Distribution of ABO and Rh blood types‬‭within four ethnic groups in the‬
F
‭United States.‬

‭ .11 Blood Type Antibodies, Agglutination, and Antibody-Based ABO and Rh‬
7
‭Blood Typing‬
I‭ f cells or viruses with foreign antigens are present in the blood, the immune system will‬
‭respond to destroy them, thus protecting you. One type of response is the production and‬
‭secretion of antibodies (also called immunoglobulins) by a type of leukocytes.‬ ‭Antibodies‬
‭are proteins that recognize specific antigens, bind to them, and facilitate the destruction of‬
‭the associated foreign cells or viruses.‬
‭ sually, the body must be exposed to a foreign antigen before an antibody is produced,‬
U
‭which is valid for the Rh blood group. However, it is not the case for the ABO blood group.‬
‭Individuals with type A blood without prior exposure to incompatible blood have‬
‭preformed antibodies to the B antigen circulating in their blood plasma. These antibodies‬
‭called anti-B antibodies, will cause clumping of the foreign blood cells, called‬
‭agglutination,‬‭and subsequent destruction, called‬‭hemolysis,‬‭if they ever encounter‬
‭erythrocytes with B antigens. Similarly, an individual with type B blood has performed‬
‭anti-A antibodies. Individuals with type AB blood, which has both antigens, do not have‬
‭preformed antibodies to either. People with type O blood lack antigens A and B on their‬
‭erythrocytes, but anti-A and anti-B antibodies circulate in their blood plasma.‬

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‭ commonly used method to determine a person’s ABO and Rh blood types is to add‬
A
‭commercially-prepared antibodies to drops of blood and check if agglutination occurs.‬
‭For example, if purified anti-A antibodies are added to a blood drop, and agglutination‬
‭o ccurs, then one can conclude that A antigens must be present on the erythrocytes. The‬
‭same general process is used to test for the presence of B and Rh antigens, using anti-B‬
‭and anti-D antibodies (‬‭Figure 7.11‬‭)‬

‭ igure 7.11‬ ‭The blood typing card contains three‬‭reaction sites or wells. One is coated‬
F
‭with an anti-A antibody, one with an anti-B antibody, and one with an anti-D antibody‬
‭(tests for the presence of Rh factor D). Mixing a drop of blood and saline into each well‬
‭enables the blood to interact with the type-specific antibodies. Agglutination of RBCs in a‬
‭given site indicates a positive identification of the blood antigens, such as A and Rh‬
‭antigens for blood type A‬‭+‭.‬ (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of this figure‬
‭is available at‬‭Biology411.com‬‭.‬

‭7.12 Blood Transfusions and Adverse Reactions‬

‭ ‬‭blood transfusion‬‭is a medical procedure in which‬‭blood from one individual,‬
A
‭designated the donor, is transferred to an individual, designated the recipient. Common‬
‭reasons for blood transfusions include treating anemia and blood loss due to accidents,‬
‭surgery, or childbirth.‬
‭ lood transfusions in humans were risky until Karl Landsteiner, an Austrian biologist and‬
B
‭physician, discovered the major human blood groups in 1900. Until then, physicians did‬
‭not understand that death sometimes followed blood transfusions, when the type of‬
‭donor blood infused into the patient was incompatible with the patient’s blood. With the‬
‭discovery of blood groups, it became possible for the first time to match patient-donor‬
‭blood types and prevent transfusion reactions and deaths.‬

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‭ ore than 50 antigens have been identified on erythrocyte membranes, but the most‬
M
‭significant in terms of their potential harm to patients during blood transfusions are the‬
‭ABO and Rh blood groups.‬
‭ ransfusion reactions are avoided by transfusing only matching blood types; that is, a type‬
T
‭B‭+‬ ‬ ‭recipient should ideally receive blood only from‬‭a type B‬‭+‬ ‭donor, and so on. When‬
‭acute bleeding threatens the patient’s life, there may not be time to identify blood type. In‬
‭these cases, blood from a‬‭universal donor‬‭—an individual‬‭with type O‬‭−‬ ‭blood—may be‬
‭transfused. Recall that type O erythrocytes do not display A or B antigens. Thus, anti-A or‬
‭anti-B antibodies circulating in the patient’s blood plasma will not encounter any‬
‭erythrocyte surface antigens on the donated blood and, therefore, will not be provoked‬
‭into a response. One problem with this designation of the universal donor is if the O‬‭−‬
‭individual had prior exposure to Rh antigen, Rh antibodies might be present in the‬
‭donated blood. Also, introducing type O blood into an individual with type A, B, or AB‬
‭blood will introduce antibodies against both A and B antigens, as these continuously‬
‭circulate in the type O blood plasma. This may cause problems for the recipient, but‬
‭because the volume of blood transfused is much lower than that of the patient’s blood, the‬
‭adverse effects of the relatively few infused plasma antibodies are typically limited. Rh‬
‭factor also plays a role in transfusion reactions. If Rh‬‭−‬ ‭individuals receiving blood have‬
‭had prior exposure to Rh antigen, antibodies for this antigen may be present in the blood‬
‭and trigger agglutination to some degree. Although it is always preferable to type a‬
‭patient’s blood before transfusing, this is not always possible in an actual life-threatening‬
‭emergency, and these procedures may be implemented.‬
‭ patient with blood type AB‬‭+‬ ‭is known as the‬‭universal‬‭recipient‬‭. This patient can‬
A
‭theoretically receive any type of blood because the patient’s blood—having both A and B‬
‭antigens on the erythrocyte surface—does not produce anti-A or anti-B antibodies. In‬
‭addition, an Rh‬‭+‬ ‭patient can receive both Rh‬‭+‬ ‭and‬‭Rh‬‭−‬ ‭blood. However, remember that the‬
‭donor’s blood will contain circulating antibodies, again with possible negative implications.‬
‭Figure 7.12‬‭summarizes the blood types and compatibilities‬‭for transfusions.‬

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‭ igure 7.12‬ ‭A summary of the characteristics of‬‭the blood types in the ABO blood group‬
F
‭and compatible types for transfusions. See the text for more details about the universal‬
‭donor and recipient. (credit:‬‭OpenStax‬‭). A link‬‭to a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ or additional information about blood transfusions, click the link below or scan the QR‬
F
‭code to watch the TED-Ed video by Bill Schutt‬‭titled‬‭“How do blood‬
‭transfusions work?”.‬

‭How do blood transfusions work? - Bill Schutt‬

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‭7.13 Rh-Incompatibility, Pregnancy, and RhoGam‬

I‭ n contrast to the ABO group antibodies, which are pre-formed, antibodies to the Rh‬
‭antigen are produced only in Rh‬‭−‬ ‭individuals after‬‭exposure to the antigen. This process,‬
‭called sensitization, occurs following a transfusion with Rh-incompatible blood or, more‬
‭commonly, with the birth of an Rh‬‭+‬ ‭baby to an Rh‬‭−‬ ‭mother. Problems are rare in a first‬
‭pregnancy since the baby’s Rh‬‭+‬ ‭cells rarely cross‬‭the placenta (the organ of gas and‬
‭nutrient exchange between the baby and the mother). However, during or immediately‬
‭after birth, the Rh‬‭−‬ ‭mother can be exposed to the‬‭baby’s Rh‬‭+‬ ‭cells (‬‭Figure 7.13‬‭). Research‬
‭has shown this occurs in 13−14% of such pregnancies. After exposure, the mother’s‬
‭immune system begins to generate anti-Rh antibodies. If the mother should conceive‬
‭another Rh‬‭+‬ ‭baby, the Rh antibodies she has produced‬‭can cross the placenta into the fetal‬
‭bloodstream and destroy the fetal RBCs. This condition, known as‬‭hemolytic disease of‬
‭the newborn (HDN)‬‭o r erythroblastosis fetalis, may‬‭cause anemia in mild cases. Still,‬
‭agglutination and hemolysis can be so severe that the fetus may die in the womb or‬
‭shortly after birth without treatment.‬
‭ drug known as‬‭RhoGAM‬‭, short for Rh immune globulin,‬‭can temporarily prevent the‬
A
‭development of Rh antibodies in the Rh‬‭−‬ ‭mother, thereby‬‭averting this potentially serious‬
‭disease for the fetus. RhoGAM antibodies destroy any fetal Rh‬‭+‬ ‭erythrocytes that may‬
‭cross the placental barrier. RhoGAM is typically administered to Rh‬‭−‬ ‭mothers during‬
‭weeks 26−28 of pregnancy and within 72 hours following birth. It has proven remarkably‬
‭effective in decreasing the incidence of HDN. Earlier, we noted that the incidence of HDN in‬
‭an Rh‬‭+‬ ‭subsequent pregnancy to an Rh‬‭−‬ ‭mother is about‬‭13–14% without preventive‬
‭treatment. Since the introduction of RhoGAM in 1968, the incidence has dropped to about‬
‭0.1% in the United States.‬
‭ or an additional review of several topics in this chapter, click the link below or scan the‬
F
‭QR code to watch the TED-Ed video by Natalie S. Hodge‬‭titled “Why do blood types‬
‭matter?”.‬

Why do blood types matter? - Natalie S. Hodge

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‭ igure 7.13‬ ‭(a) The first exposure of an Rh‬‭-‬ ‭mother‬‭to Rh‬‭+‬ ‭erythrocytes during‬
F
‭pregnancy induces sensitization. Anti-Rh antibodies begin to circulate in the mother’s‬
‭bloodstream. A second exposure occurs with a subsequent pregnancy with an Rh‬‭+‬ ‭fetus in‬
‭the uterus. Maternal anti-Rh antibodies may cross the placenta and enter the fetal‬
‭bloodstream, causing agglutination and hemolysis of fetal erythrocytes. (b) If the Rh‬‭-‬
‭mother is injected with RhoGam (anti-Rh antibodies) during pregnancy or soon after the‬
‭birth of a child, the antibodies will bind to the fetal Rh‬‭+‬ ‭cells and prevent a maternal‬
‭immune response that could affect a subsequent fetus with Rh+ cells. (credit:‬‭OpenStax‬‭).‬
‭A link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭7.14 Career Connection - Phlebotomy and Medical Lab Technology‬

‭ hlebotomists are professionals trained to draw blood (phleb- = “a blood vessel”; -tomy =‬
P
‭“to cut”). When more than a few drops of blood are required, phlebotomists perform a‬
‭venipuncture, typically of a surface vein in the arm. They perform a capillary stick on a‬
‭finger, an earlobe, or an infant's heel when only a tiny quantity of blood is required.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ n arterial stick is collected from an artery to analyze blood gases. After collection, the‬
A
‭blood may be analyzed by medical laboratories or used for transfusions, donations, or‬
‭research. While many allied health professionals practice phlebotomy, the American‬
‭Society of Phlebotomy Technicians issues certificates to individuals passing a national‬
‭examination, and some large labs and hospitals hire individuals expressly for their skill in‬
‭phlebotomy.‬

‭Medical or clinical laboratories employ various individuals in technical positions:‬

‭•‬ ‭ edical technologists (MT), also known as clinical laboratory technologists (CLT),‬
M
‭typically hold a bachelor’s degree and certification from an accredited training‬
‭program. They perform various tests on various body fluids, including blood. The‬
‭information they provide is essential to primary care providers in determining a‬
‭diagnosis, monitoring the course of a disease, and responding to treatment.‬

‭•‬ ‭ edical laboratory technicians (MLT) typically have an associate’s degree but may‬
M
‭perform duties similar to those of an MT.‬

‭•‬ ‭ edical laboratory assistants (MLA) spend most of their time processing samples and‬
M
‭carrying out routine assignments within the lab. Clinical training is required, but a‬
‭degree may be optional to obtain a position.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 8: Cardiovascular System II - Heart‬

‭ igure 8.1‬ ‭The human heart is a muscular pump that‬‭keeps the body continually‬
F
‭supplied with blood. (credit: Patrick J. Lynch;‬‭OpenStax‬‭)‬

‭8.1 Introduction‬
‭ ost animals are complex multicellular organisms that require a mechanism for‬
M
‭transporting nutrients throughout their bodies and removing waste products. The‬
‭circulatory system supplies the cells, tissues, and organs with oxygen and nutrients. It also‬
‭removes carbon dioxide and wastes, which are byproducts of cellular respiration (Chapter‬
‭6).‬
‭ t the core of the human circulatory system is the heart. Although the term “heart” is an‬
A
‭English word, cardiac (heart-related) terminology can be traced back to the Latin term‬
‭“kardia.” Cardiology is the study of the heart, and cardiologists are the physicians who deal‬
‭primarily with the heart.‬
‭ he heart is the first functional organ to develop in human embryos. It begins beating and‬
T
‭pumping blood around day 21 or 22, a mere three weeks after fertilization. In adults, the‬
‭heart is approximately the size of a clenched fist and protected beneath the rib cage. The‬
‭heart, made of specialized and unique cardiac muscle, pumps blood throughout the body.‬
‭Heart contractions are driven by self-generated electrical impulses that the brain and‬
‭hormones help to regulate.‬
‭ nderstanding the heart’s basic anatomy and function is essential to understanding the‬
U
‭body’s circulatory and respiratory systems. In this chapter, you will explore the‬
‭remarkable pump that propels the blood into the vessels. There is no better word to‬
‭describe the function of the heart other than “pump” since its contraction develops the‬

‭208‬
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‭ ressure that ejects blood into the pulmonary vessels, which take blood to the lungs and‬
p
‭back to the heart and the aorta, which sends blood out the body. Although the term‬
‭“pump” suggests a mechanical device made of steel and plastic, the anatomical structure is‬
‭essentially a living, sophisticated muscle. As you read this chapter, try to keep these twin‬
‭concepts in mind: pump and muscle.‬

‭8.2 Human Heart and Blood Vessels‬

‭ he human heart is a perfect example of the relationship between structure and function.‬
T
‭The four-chambered heart, with its unique cardiac muscle, one-way valves, and associated‬
‭blood vessels, is designed to transport vital oxygen to the body cells for cellular‬
‭respiration and remove carbon dioxide from tissues, a waste product of this process.‬
‭ he heart pumps blood through the three divisions of the circulatory system: the‬
T
‭coronary circuit‬‭(vessels that serve the heart),‬‭pulmonary‬‭c ircuit‬‭(heart and lungs), and‬
‭systemic circuit‬‭(systems of the body), as shown in‬‭Figure 8.2‬‭. Coronary circulation takes‬
‭blood directly from the main artery (aorta) coming from the heart. For pulmonary and‬
‭systemic circulation, the heart must pump blood to the lungs or the rest of the body.‬
I‭ n humans, the heart is divided into four chambers: two atria and two ventricles. There is‬
‭o ne atrium and one ventricle on the right side and one atrium and one ventricle on the‬
‭left side. Blood in the two sides is prevented from mixing by the septum, which separates‬
‭the sides. The‬‭atria‬‭are the chambers that receive‬‭blood from blood vessels, and the‬
‭ventricles‬‭are the chambers that pump blood out of‬‭the heart to either the pulmonary‬
‭circuit (right side) or the systemic circuit (the left side).‬
‭ lood vessels, the third component of the circulatory system, will be discussed in detail in‬
B
‭Chapter 9. However, let’s briefly mention three types of blood vessels: arteries, veins, and‬
‭capillaries.‬ ‭Arteries‬‭take blood away from the heart,‬‭and‬‭veins‬‭bring blood toward the‬
‭heart. In almost all cases, arteries carry blood with a high oxygen content (called “red‬
‭blood”), while veins carry blood with a low oxygen content (called “blue blood”). The‬
‭exceptions are the‬‭pulmonary arteries‬‭, which carry‬‭blue blood from the right ventricle to‬
‭the lungs, and the‬‭pulmonary veins‬‭, which carry red‬‭blood from the lungs to the left‬
‭atrium.‬‭Capillaries‬‭are where the exchange of substances‬‭between the blood and the cells‬
‭o f tissues occurs.‬
‭ wo of the three remaining blood vessels important to our discussion of the heart are the‬
T
‭superior and inferior vena cava. Both are veins and return blue blood from the body to‬
‭the heart. The‬‭superior vena cava‬‭transports blood‬‭from above the heart (i.e., superior‬
‭to the heart). In contrast, the‬‭inferior vena cava‬‭transports blood from below the heart‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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(‭ i.e., inferior to the heart). The third one is the‬‭aorta‬‭, which carries the red blood from‬
‭the heart to the rest of the body.‬

‭ igure 8.2‬ ‭The human circulatory system is divided‬‭into systemic, pulmonary, and‬
F
‭coronary circuits. Blood is pumped from veins of the systemic circuit via the superior and‬
‭inferior vena cava into the heart's right atrium and then into the right ventricle. Blood‬
‭then enters the pulmonary circuit and is oxygenated by the lungs. From the pulmonary‬
‭circuit, blood re-enters the heart through the left atrium. Blood re-enters the systemic‬
‭circuit through the aorta from the left ventricle and is distributed to the rest of the body.‬
‭The coronary circuit, which provides blood to the heart, is not shown. (credit:‬‭OpenStax‬‭).‬
‭A link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ ecause the right side of the heart sends blood to the pulmonary circuit (i.e., the lungs),‬
B
‭there is a shorter distance to pump, which means that the muscle wall on the right side of‬
‭the heart is not as thick as the left side, which must have enough pressure to pump blood‬
‭throughout the entire body.‬
‭ he‬‭right atrium‬‭receives deoxygenated (i.e., blue)‬‭blood from the superior vena cava,‬
T
‭which drains blood from the jugular vein that comes from the brain and from the veins‬
‭that come from the arms, as well as from the inferior vena cava, which drains blood from‬
‭the veins that come from the lower organs and the legs (‬‭Figure 8.3a‬‭). This blue blood‬
‭then passes to the‬‭right ventricle‬‭through the‬‭atrioventricular‬‭valve‬‭o r the‬‭tricuspid‬

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P
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‭ alve‬‭, a flap of connective tissue that opens in only one direction to prevent the backflow‬
v
‭o f blood. After it is full, the right ventricle pumps the blood past the‬‭pulmonary‬
‭semilunar valve‬‭to the‬‭pulmonary arteries‬‭and then‬‭to the lungs for re-oxygenation. The‬
‭left atrium‬‭then receives the oxygen-rich, red blood‬‭from the lungs via the‬‭pulmonary‬
‭veins‬‭. This blood passes through the‬‭bicuspid valve‬‭o r‬‭mitral valve‬‭(the atrioventricular‬
‭valve on the left side of the heart) to the‬‭left ventricle,‬‭where the blood is pumped past‬
‭the‬‭aortic semilunar valve‬‭and into the aorta, the‬‭major artery of the body, taking‬
‭oxygenated blood to the organs and muscles of the body. This pumping pattern is called‬
‭double circulation‬‭and is found in all mammals.‬
‭ he heart comprises three layers: the epicardium, the myocardium, and the endocardium‬
T
‭(‬‭Figure 8.3‬‭). The‬‭myocardium‬‭consists of the heart‬‭muscle cells that make up the middle‬
‭layer and the bulk of the heart wall. The contraction of the cells in this layer generates the‬
‭force to move blood to the lungs (the pulmonary circuit) or to the rest of the body (the‬
‭systemic circuit).‬
‭ he heart has blood vessels that supply the heart muscle with blood (‬‭Figure 8.3b‬‭). The‬
T
‭coronary arteries‬‭branch from the aorta and surround‬‭the heart's outer surface like a‬
‭crown. They diverge into capillaries, where the heart muscle is supplied with oxygen,‬
‭before converging into the‬‭coronary veins‬‭to take‬‭the deoxygenated blood back to the‬
‭right atrium, where it will be re-oxygenated through the pulmonary circuit.‬

‭ he heart muscle will die without a steady supply of oxygen the blood provides.‬
T
‭Atherosclerosis‬‭is the blockage of an artery by the‬‭buildup of fatty plaques. Because of‬
‭the narrow diameter of the coronary arteries and their function in serving the heart itself,‬
‭atherosclerosis can be deadly in these arteries. The slowdown of blood flow and‬
‭subsequent oxygen deprivation that results from atherosclerosis causes severe pain,‬
‭known as‬‭angina‬‭, and complete blockage of the arteries‬‭will cause a‬‭myocardial‬
‭infarction (MI)‬‭and‬‭the death of cardiac muscle tissue.‬‭An MI is commonly known as a‬
‭heart attack‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 8.3‬ ‭(a) The heart is primarily made of a thick‬‭muscle layer called the myocardium,‬
F
‭surrounded by membranes. One-way valves separate the four chambers. (b) Blood vessels‬
‭o f the coronary system, including the coronary arteries and veins, keep the heart muscle‬
‭oxygenated. (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭8.3 Heart Valves‬

‭ s previously discussed, the heart has two types of valves: atrioventricular (AV) and‬
A
‭semilunar. Both types function to prevent blood from flowing in the opposite direction. In‬
‭Figure 8.4‬‭, the heart has been sectioned in the horizontal plane (i.e., side to side), which is‬
‭different from the vertical plane sectioning (i.e., top to bottom) seen in‬‭Figure 8.3‬‭. Let's‬
‭more closely examine the structure of these valves and their mechanisms of action.‬

‭ igure 8.4‬ ‭With the atria and major vessels removed‬‭by horizontal sectioning, all four‬
F
‭valves are visible: the two AV valves (tricuspid and bicuspid) and the two semilunar valves‬
‭(pulmonary and aortic). The tricuspid, pulmonary, and aortic valves have three flaps (or‬
‭structural components), while the bicuspid valve has two. All four valves are shown in the‬
‭closed position, preventing blood backflow. In the case of the AV valves, backflow is‬
‭prevented from the ventricles into the atria. In the case of the semilunar valves, backflow‬
‭is prevented from the pulmonary trunk into the right ventricle (on the right side) and‬
‭from the aorta into the left ventricle (on the left side). (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

I‭ n‬‭Figure 8.5a‬‭, the two atrioventricular valves are‬‭o pen, and the two semilunar valves are‬
‭closed. This occurs when both atria and ventricles are relaxed and when the atria contract‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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t‭ o pump blood into the ventricles.‬‭Figure 8.5b‬‭shows a frontal view. Although only the left‬
‭side of the heart is illustrated, the right side is virtually identical, with the AV valve opening‬
‭and the semilunar valve closing.‬

‭ igure 8.5‬ ‭(a) A horizontal section through the heart‬‭illustrates the four heart valves. The‬
F
‭two atrioventricular valves are open; the two semilunar valves are closed. The atria and‬
‭vessels have been removed. (b) A vertical section through the heart illustrates blood flow‬
‭through the mitral valve. When the mitral valve is open, it allows blood to move from the‬
‭left atrium to the left ventricle. The aortic semilunar valve is closed to prevent the‬
‭backflow of blood from the aorta to the left ventricle. While the right side isn’t shown, the‬
‭right AV (bicuspid) valve would also be open, and the pulmonary semilunar valve would‬
‭also be closed. (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 8.6a‬‭shows the atrioventricular valves closed while the two semilunar valves are‬
F
‭o pen. This occurs when the ventricles contract to eject blood into the pulmonary arteries‬
‭and the aorta. Closure of the two AV valves prevents blood from being forced back into‬
‭the atria. This stage is presented from a frontal view in‬‭Figure 8.6b‬‭. Again, although only‬
‭the left side of the heart is shown, the situation on the right side is similar.‬

‭ igure 8.6‬ ‭(a) A horizontal section through the‬‭heart illustrates the four heart valves‬
F
‭during ventricular contraction. The two AV valves are closed, but the two semilunar valves‬
‭are open. The atria and vessels have been removed. (b) A vertical section view shows the‬
‭closed mitral (bicuspid) valve that prevents the backflow of blood into the left atrium. The‬
‭aortic semilunar valve is open to eject blood into the aorta. While the right side isn’t‬
‭shown, the right AV (bicuspid) valve would also be closed, and the pulmonary semilunar‬
‭valve would be open. (credit:‬‭OpenStax‬‭). A link to‬‭a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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I‭ n‬‭Figures 8.5‬‭and‬‭8.6‬‭note the presence‬‭of chordae tendineae,‬‭strands of fibrous tissue‬

‭that connect the leaflets of the AV valves (tricuspid and bicuspid) to papillary muscle‬
‭tissue in the ventricles (‬‭Figure 8.7‬‭). These strands‬‭prevent the valve leaflets from opening‬
‭into the atria during ventricular contractions and blood from leaking back into the atria.‬

‭ igure 8.7‬ ‭The chordae tendineae, which extend from‬‭leaflets of AV valves to the‬
F
‭papillary muscle in the ventricles. These fibrous strands anchor the valve leaflets and‬
‭prevent them from prolapsing into the atria when the ventricles contract, thus maintaining‬
‭correct blood flow through the heart. (credit:‬ ‭Muscolopapillare+cordetendinee.jpg‬‭;‬
‭E.Faccio P.Saccheri; Wikimedia Commons;‬‭CC BY-SA 3.0‬‭).‬ ‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

I‭ f valves don’t close properly, then blood can leak and decrease the efficiency of correct‬
‭flow. For example, if the left AV (mitral valve) doesn’t close properly during ventricular‬
‭contraction, then instead of blood flowing only from the left ventricle through the left‬
‭semilunar (aortic) valve, some blood will leak back into the left atrium. This condition is‬
‭referred to as mitral valve prolapse (‬‭Figure 8.8‬‭).‬‭If there is minimal leakage, then no‬
‭intervention may be necessary. However, surgery may be needed to correct it in more‬
‭severe cases.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 8.8‬ ‭If the left AV valve, also known as the‬‭mitral valve, doesn’t correctly close‬
F
‭during ventricular contraction, then instead of blood only moving through the left‬
‭semilunar (aortic) valve into the aorta, some will leak back into the left atria. (credit:‬
‭modification of‬‭Mitral Valve Prolapse.png‬‭;‬‭BruceBlaus‬‭;‬‭Wikimedia Commons;‬‭CC BY-SA 4.0‬‭)‬

‭8.4 The Cardiac Cycle‬

‭ he heart's primary purpose is to pump blood through the body; it does so in a repeating‬
T
‭sequence of events called the cardiac cycle. The‬‭c ardiac‬‭c ycle‬‭is the coordination of the‬
‭filling and emptying of the heart of blood by electrical signals that cause the heart muscles‬
‭to contract and relax. The human heart beats over 100,000 times per day! In each cardiac‬
‭cycle, the heart contracts (‬‭systole‬‭), pushing out‬‭the blood and pumping it to the lungs or‬
‭through the body; this is followed by a relaxation phase (‬‭diastole‬‭), where the heart fills‬
‭with blood, as illustrated in‬‭Figure 8.9‬‭. The atria‬‭contract simultaneously, forcing blood‬
‭through the atrioventricular valves into the ventricles. When the ventricles contract, the‬
‭atrioventricular valves close, which produces a monosyllabic “‬‭lup‬‭” sound. Following a‬
‭brief delay, the ventricles contract simultaneously, forcing blood through the semilunar‬
‭valves into the aorta and the pulmonary arteries for systemic and pulmonary circulation,‬
‭respectively. When the ventricles relax, the semilunar valves close, which produces a‬
‭monosyllabic “‬‭dup‬‭” sound.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 8.9‬ ‭During (a) cardiac diastole, the heart‬‭muscle is relaxed, and blood flows into‬
F
‭the heart. During (b) atrial systole, the atria contract, pushing blood into the ventricles.‬
‭During (c) atrial diastole, the ventricles contract, forcing blood out of the heart. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ or additional information about the heart and its function, click the link below or scan‬
F
‭the QR code to watch the TED-Ed video by‬‭Edmond Hui‬‭titled “How the heart actually‬
‭pumps blood.”‬

How the heart actually pumps blood - Edmond Hui

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭218‬
‭Chapter 8: Heart‬

‭ ne of the simplest yet effective diagnostic techniques applied to assess the state of a‬
O
‭patient’s heart is‬‭auscultation‬‭, or listening to the‬‭internal sounds, using a stethoscope.‬
‭Figure 8.10‬‭shows the parts of a‬‭stethoscope‬‭, and‬‭Figure 8.11‬‭shows the proper‬
‭placement of the stethoscope bell during the procedure. It is common practice for the‬
‭clinician to ask the patient to breathe deeply. This procedure allows for listening to airflow‬
‭and may also amplify heart murmurs, a term used to describe an unusual sound coming‬
‭from the heart caused by the turbulent flow of blood. Inhaling increases blood flow into‬
‭the right side of the heart and may increase the sounds of right-sided heart murmurs.‬
‭Partially exhaling restricts blood flow into the left side of the heart and may increase the‬
‭sounds of left-sided heart murmurs.‬

‭ igure 8.10‬ ‭The parts of a stethoscope. The clinician‬‭inserts the earplugs and the bell at‬
F
‭various locations on the patient’s chest during auscultation. (credit: Freeimages.com;‬‭CC0‬
‭1.0‬‭)‬

‭ or additional information about the development of the stethoscope, click the link below‬
F
‭o r scan the QR code to watch the TED-Ed video by Jessica Oreck titled “How the‬
‭stethoscope was invented.”‬

How the stethoscope was invented | Moments of Vision 7 - Jessica…

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 8: Heart‬

‭ igure 8.11‬ ‭Proper placement of the stethoscope‬‭bell facilitates auscultation or listening‬

F
‭to the sounds of the heart during the cardiac cycle. A different valve can be heard at each‬
‭o f the four locations on the chest. (credit:‬‭OpenStax‬‭)‬

‭ he heart pumping is a function of the‬‭c ardiac muscle‬‭cells that comprise the‬

T
‭myocardium. These distinctive muscle cells are striated like skeletal muscle but pump‬
‭rhythmically and involuntarily like smooth muscle (‬‭Figure 8.12‬‭). The cardiac muscle cells‬
‭can initiate an electrical impulse at a fixed rate that spreads rapidly from cell to cell to‬
‭trigger the contractions associated with the beating heart. This unique property is known‬
‭as‬‭autorhythmicity‬‭; neither smooth nor skeletal muscle‬‭can do this. Even though cardiac‬
‭muscle has autorhythmicity, heart rate is regulated by the endocrine and nervous systems.‬
‭ he heart's internal pacemaker regulates the autonomous contractions of cardiac muscle‬
T
‭cells using electrical signals. The internal pacemaker starts at the‬‭sinoatrial (SA) node‬
‭near the right atrium's wall (‬‭Figure 8.13‬‭). Electrical‬‭charges spontaneously pulse from‬
‭the SA node, causing the two atria to contract in unison. The pulse reaches a second node,‬
‭called the‬‭atrioventricular (AV) node‬‭, between the‬‭right atrium and right ventricle,‬
‭where it pauses for approximately 0.1 seconds before spreading to the walls of the‬
‭ventricles. From the AV node, the electrical impulse enters the bundle of His, then to the‬
‭left and right bundle branches extending through the septum between the ventricles.‬
‭Finally, the Purkinje fibers conduct the impulse from the bottom of the heart up the walls‬
‭o f the ventricles, which causes them to contract. This pause allows the atria to empty into‬
‭the ventricles before the ventricles pump out the blood.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 8: Heart‬

‭ igure 8.12‬ ‭Cardiac muscle is composed of striated‬‭muscle cells found in cardiac tissue.‬
F
‭(credit: modification of work by Dr. S. Girod, Anton Becker; scale-bar data from Matt‬
‭Russell;‬‭OpenStax‬‭)‬

‭ igure 8.13‬ ‭Specialized conducting components of‬‭the heart include the sinoatrial (SA)‬
F
‭node, the atrioventricular (AV) node, the bundle of His, the right and left bundle branches,‬
‭and the Purkinje fibers. (credit:‬‭OpenStax‬‭). A link‬‭to a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 8: Heart‬

‭8.5 Electrocardiogram (ECG or EKG)‬

‭ he electrical impulses in the heart produce electrical currents that flow through the body.‬
T
‭Carefully placing electrodes on the skin at specific locations allows recording of the heart's‬
‭complex electrical signal (‬‭Figure 8.11‬‭) as an‬‭electrocardiogram‬‭(ECG‬‭o r‬‭EKG; Figure‬
‭8.14).‬‭The K comes from kardiology, which is the German‬‭term for cardiology. It is‬
‭essential to understand how this figure is associated with the events of the cardiac cycle,‬
‭which include atrial systole, atrial diastole, ventricular systole, ventricular diastole, and‬
‭complete diastole (when both atria and ventricles are relaxed).‬ ‭Figure 8.15‬‭explains this‬
‭association.‬

‭ igure 8.14‬ ‭An ECG that shows the repetitive pattern‬‭o f electrical changes associated‬
F
‭with regular heart contractions and relaxations. (credit:‬ ‭Normal ECG 2.svg‬‭;‬ ‭Madhero88‬‭;‬
‭CC BY-SA 3.0‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ n ECG has several prominent points: The P wave, the QRS complex, and the T wave‬
A
‭(Figure 8.16)‬‭. The small‬‭P wave‬‭represents the initiation‬‭o f atrial systole or contraction of‬
‭the atria. The large‬‭QRS complex‬‭includes the relaxation‬‭o f the atria (atrial diastole) and‬
‭contraction of the ventricles (ventricular systole). Compared to the P wave, a more‬
‭significant electrical signal is necessary because of the increased thickness of the‬
‭ventricular myocardial wall. Lastly, the‬‭T wave‬‭represents‬‭the relaxation of the ventricles‬
‭(ventricular diastole). The interval between the T wave and the next P wave represents‬
‭complete, or common, diastole when the atria and ventricles are relaxed.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 8: Heart‬

‭ igure 8.15‬ ‭The heart's beating is regulated by‬‭an electrical impulse that causes the‬
F
‭characteristic reading of an ECG. The signal is initiated at the SA node. The signal then (a)‬
‭spreads to the atria, causing them to contract (i.e., atrial systole). The signal is (b) delayed‬
‭at the AV node before passing it on to the (c) heart apex. The delay allows the atria to‬
‭relax (atrial diastole) before the (d) ventricles contract (ventricular systole). The final part‬
‭o f the ECG cycle prepares the heart for the next beat (ventricular diastole and complete‬
‭diastole). (credit:‬‭Opentextbc.ca‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 8: Heart‬

‭ igure 8.16‬ ‭The relationship between the cardiac‬‭cycle and ECG. Initially, both the atria‬
F
‭and ventricles are relaxed (complete diastole). The P wave represents the initiation of the‬
‭events that will result in atrial contraction (systole). Atrial systole extends until the QRS‬
‭complex, at which point the atria relax. The QRS complex represents the initiation of the‬
‭events that will result in ventricular contraction. The T wave represents the beginning of‬
‭ventricular relaxation. The period after the T wave until the initiation of the next P wave is‬
‭complete diastole, when both atria and ventricles are relaxed. (credit:‬‭OpenStax‬‭). A link to‬
‭a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Everyday Connection‬

E‭ xternal Automated Defibrillators‬

‭If the heart’s electrical activity is severely disrupted, cessation of electrical activity or‬
‭fibrillation may occur. In fibrillation, the heart beats in a wild, uncontrolled manner, which‬
‭prevents it from being able to pump effectively. Atrial fibrillation is a serious condition, but‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 8: Heart‬

a‭ s long as the ventricles continue to pump blood, the patient’s life may not be in‬
‭immediate danger. Ventricular fibrillation is a medical emergency requiring life support‬
‭because the ventricles do not pump blood effectively. In a hospital setting, it is often‬
‭described as “code blue.” If untreated for as little as a few minutes, ventricular fibrillation‬
‭may lead to brain death. The most common treatment is defibrillation, which uses special‬
‭paddles to apply a charge to the heart from an external electrical source to establish a‬
‭normal rhythm. A defibrillator (‬‭Figure 8.17‬‭) effectively‬‭stops the heart so that the SA‬
‭node can trigger a normal conduction cycle. Because of their effectiveness in‬
‭re-establishing a regular rhythm, external automated defibrillators (EADs) are being‬
‭placed in areas frequented by large numbers of people, such as schools, restaurants, and‬
‭airports. These devices contain simple and direct verbal instructions that can be followed‬
‭by nonmedical personnel in an attempt to save a life.‬

‭ igure 8.17‬ ‭Two types of defibrillators. (a) An‬‭external automatic defibrillator can‬
F
‭reestablish a normal sinus rhythm in a person with fibrillation. (b) Defibrillator paddles‬
‭are more commonly used in hospital settings. (credit:‬‭OpenStax‬‭)‬

‭Everyday Connection‬

‭ PR‬
C
‭The position of the heart in the torso between the vertebrae and sternum allows‬
‭individuals to apply an emergency technique known as‬‭c ardiopulmonary resuscitation‬
‭(‬‭CPR‬‭) if a patient’s heart stops beating. By applying‬‭pressure with the flat portion of one‬
‭hand on the sternum in the area between the line at T4 and T9 (‬‭Figure 8.18‬‭), it is possible‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 8: Heart‬

t‭ o manually compress the blood within the heart enough to push some of the blood within‬
‭it into the pulmonary and systemic circuits. This is particularly critical for the brain, as‬
‭irreversible damage and neurons die within minutes of blood flow loss. Current standards‬
‭call for chest compression at least 5 cm deep and at a rate of 100 compressions per‬
‭minute. At this stage, the emphasis is on performing high-quality chest compressions‬
‭rather than providing artificial respiration. CPR is generally performed until the patient‬
‭regains spontaneous contraction or is declared dead by an experienced healthcare‬
‭professional.‬

‭ hen performed by untrained or overzealous individuals, CPR can result in broken ribs‬
W
‭o r sternum, inflicting additional severe damage on the patient. If the hands are placed too‬
‭low on the sternum, it is also possible to manually drive the xiphoid process into the liver,‬
‭which may prove fatal for the patient. Proper training is essential. This proven‬
‭life-sustaining technique is so valuable that virtually all medical personnel and concerned‬
‭members of the public should be certified and routinely recertified in its application.‬
‭Various colleges, hospitals, the American Red Cross, and some commercial companies‬
‭o ffer CPR courses. They usually include the practice of the compression technique on a‬
‭mannequin.‬

‭ igure 8.18‬ ‭If the heart should stop, CPR can maintain‬‭blood flow until the heart‬
F
‭resumes beating. By applying pressure to the sternum, the blood within the heart will be‬
‭squeezed out and into circulation. Proper positioning of the hands on the sternum to‬
‭perform CPR would be between the lines at T4 and T9, which indicate specific thoracic‬
‭vertebrae. (credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭226‬
‭Chapter 8: Heart‬

I‭ t is essential to understand how the events of an ECG correlate with not only heart‬
‭contractions and relaxations but also blood flow through it.‬ ‭Figure 8.19‬‭shows these‬
‭relationships.‬

‭ igure 8.19‬ ‭An illustration of the interrelationship‬‭between the cardiac cycle, blood flow‬
F
‭through the heart, and an ECG. Begin at the area labeled “Start” and move clockwise.‬
‭(credit: Modified from‬‭Phases of the Cardiac Cycle.jpg‬‭;‬‭OpenStax College;‬‭CC BY 3.0‬‭). A‬
‭link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭227‬
‭Chapter 8: Heart‬

‭8.6 Disease of the Heart - Myocardial Infarction‬

‭ yocardial infarction (MI) is the formal term for what is commonly referred to as a‬‭heart‬
M
‭attack‬‭. It usually results from a lack of blood flow‬‭and oxygen to a heart region, resulting‬
‭in the death of the cardiac muscle cells. An MI often occurs when a coronary artery is‬
‭blocked by the buildup of atherosclerotic plaque consisting of lipids, cholesterol and fatty‬
‭acids, and white blood cells, primarily macrophages. It can also occur when a portion of an‬
‭unstable atherosclerotic plaque travels through the coronary arterial system and lodges in‬
‭o ne of the smaller vessels. The resulting blockage restricts blood and oxygen flow to the‬
‭myocardium and causes tissue death.‬

I‭ n the case of acute MI, there is often sudden pain beneath the sternum called‬‭angina‬
‭pectoris‬‭, usually radiating down the left arm in males‬‭but not in female patients. Until this‬
‭anomaly between the sexes was discovered, many female patients suffering MIs were‬
‭misdiagnosed and sent home. In addition, patients typically present with difficulty‬
‭breathing and shortness of breath, irregular heartbeat, nausea and vomiting, sweating,‬
‭anxiety, and fainting, although not all of these symptoms may be present. Many of the‬
‭symptoms are shared with other medical conditions, including anxiety attacks and simple‬
‭indigestion, so differential diagnosis is critical. It is estimated that between 22 and 64‬
‭percent of MIs present without symptoms.‬

‭ n MI can be confirmed by examining the patient’s ECG. In addition, echocardiography or‬

A
‭cardiac magnetic resonance imaging may be employed. Standard blood tests indicating an‬
‭MI include elevated levels of creatine kinase and cardiac troponin, both of which are‬
‭released by damaged cardiac muscle cells.‬

I‭ mmediate treatments for MI are essential and include administering supplemental‬

‭oxygen, aspirin that helps to break up clots, and nitroglycerine administered under the‬
‭tongue to facilitate its absorption. Despite its unquestioned success in treatments and use‬
‭since the 1880s, the mechanism of nitroglycerine still needs to be understood.‬
‭Longer-term treatments include injections of agents that dissolve the clot, the‬
‭anticoagulant heparin, balloon angioplasty, and stents to open blocked vessels and bypass‬
‭surgery to allow blood to pass around the site of a blockage. If the damage is extensive,‬
‭coronary replacement with a donor heart or coronary assist device, a sophisticated‬
‭mechanical device that supplements the heart's pumping activity, may be employed.‬
‭Despite the attention, the development of artificial hearts to augment the severely limited‬
‭supply of heart donors has proven less than satisfactory but will likely improve in the‬
‭future.‬

S‭ ignificant risk factors for MI include cardiovascular disease, age, smoking, high blood‬
‭levels of low-density lipoprotein (LDL, often referred to as “bad” cholesterol), low levels of‬
‭high-density lipoprotein (HDL, or “good” cholesterol), hypertension, diabetes mellitus,‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ besity, lack of physical exercise, chronic kidney disease, excessive alcohol consumption,‬
o
‭and use of illegal drugs.‬

‭ or additional information about myocardial infarctions, click the link below or scan the‬
F
‭QR code to watch the TED-Ed video by Krishna Sudhir titled “What happens during a‬
‭heart attack?”.‬

What happens during a heart attack? - Krishna Sudhir

‭8.7 Blood Pressure‬

‭ lood pressure is one of the critical parameters measured on virtually every patient in‬
B
‭every healthcare setting.‬ ‭Blood pressure (BP)‬‭is‬‭the pressure exerted by blood on the‬
‭walls of a blood vessel that helps to push blood through it.‬‭Systolic blood pressure‬
‭measures blood pressure on vessels while the heart is beating.‬‭Diastolic blood pressure‬
‭measures the pressure in the vessels between heartbeats.‬
‭ he technique used today was developed more than 100 years ago by a pioneering‬
T
‭Russian physician, Dr. Nikolai Korotkoff. Turbulent blood flow through the vessels can be‬
‭heard as a soft ticking while measuring blood pressure, known as‬‭Korotkoff sounds‬‭. The‬
‭technique of measuring blood pressure requires using a‬‭sphygmomanometer‬‭(i.e., blood‬
‭pressure cuff) and a stethoscope (‬‭Figure 8.20‬‭). The‬‭method is as follows:‬
‭•‬ ‭ he clinician wraps an inflatable cuff tightly around the patient’s arm at about the‬
T
‭heart level.‬

‭•‬ ‭ he clinician squeezes a rubber pump to inject air into the cuff, raising pressure‬
T
‭around the artery and temporarily stopping blood flow into the patient’s arm.‬

‭•‬ ‭ he clinician places the stethoscope in the inner bend of the patient’s elbow and,‬
T
‭while gradually allowing air within the cuff to escape, listens for the Korotkoff sounds.‬
‭ lthough there are five recognized Korotkoff sounds, only two are typically recorded.‬
A
‭Initially, no sounds are heard since there is no blood flow through the vessels, but as air‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ ressure drops, the cuff relaxes, and blood flow returns to the arm.‬
p
‭As shown in‬‭Figure 8.21‬‭, the first Korotkoff sound‬‭indicates systolic pressure. As more air‬
‭is released from the cuff, blood can flow freely through the brachial artery, and all sounds‬
‭disappear. The point at which the last sound is heard is recorded as the patient’s diastolic‬
‭pressure. The optimal systolic blood pressure is 120 mmHg, and the optimal diastolic‬
‭blood pressure is 80 mmHg. Most hospitals and clinics now have automated equipment‬
‭for measuring blood pressure that uses the same fundamental principles.‬

‭ igure 8.20‬ ‭Using a sphygmomanometer (blood pressure‬‭cuff) and stethoscope to‬

F
‭determine a patient’s blood pressure. The silver valve in the clinician’s right hand is closed‬
‭when the cuff pressure is increased by pumping the black bulb in the same hand. When‬
‭the pressure is at approximately 180 millimeters of mercury (mm Hg), the valve is slowly‬
‭o pened to release pressure on a blood vessel in the arm called the brachial artery. The‬
‭stethoscope allows the clinician to listen for Korotkoff sounds, associated with sounds of‬
‭blood first moving through the blood vessel (systolic pressure) and when the blood is‬
‭freely flowing through the blood vessel (diastolic pressure). The blood pressure value is‬
‭reported as a ratio (e.g., 120/80), which is the systolic divided by diastolic pressures (e.g.,‬
‭noted as “120 over 80”). (credit:‬‭agilemktg1‬‭; Flickr)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭agilemktg1‬

‭ igure 8.21‬ ‭Determining systolic and diastolic blood‬‭pressure using a‬

F
‭sphygmomanometer and a stethoscope. At the pressure associated with point A (e.g., 120‬
‭mm Hg), the first Korotkoff sound is heard, and the systolic pressure is recorded. The cuff‬
‭pressure is decreased slowly by continuing to release air until the last sound is heard and‬
‭the diastolic pressure is recorded (e.g., 80 mm Hg).‬ ‭(credit:‬‭Korotkow English.jpg‬‭;‬
‭PhilippN‬‭; Wikimedia;‬‭CC BY 3.0‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ any factors (e.g., hormones, stress, exercise, eating, sitting, and standing) can affect‬
M
‭blood pressure.‬ ‭Hypertension‬‭, or consistently high‬‭blood pressure (e.g., systolic pressure‬
‭o f 130 mm Hg or more; diastolic pressure of 90 mm Hg or more), is a disorder that affects‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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a‭ pproximately 30% or more of the US adult population. However, unless blood pressure‬
‭is checked regularly, the condition may not be promptly detected, and long-term physical‬
‭consequences that aren’t associated with immediately visible symptoms can result. This is‬
‭why hypertension is referred to as “the silent killer.”‬

‭ or additional information about blood pressure and related topics, click the link below or‬
F
‭scan the QR code to watch the TED-Ed video by‬‭Wilfred‬‭Manzano titled “How blood‬
‭pressure works.”‬

How blood pressure works - Wilfred Manzano

‭8.8 Pulse‬
‭ fter blood is ejected from the heart, elastic fibers in the arteries help maintain high‬
A
‭pressure as they expand to accommodate the blood and then recoil (i.e., snap back). This‬
‭expansion and recoiling effect, known as the‬‭pulse‬‭,‬‭can be palpated (felt) manually or‬
‭measured electronically.‬
‭ ecause pulse indicates heart rate, which is recorded as beats per minute (bpm), it is‬
B
‭measured clinically to provide clues to a patient’s state of health. Both the rate and the‬
‭strength of the pulse are significant clinically. A high or irregular pulse rate can be caused‬
‭by physical activity or other temporary factors, but it may also indicate a heart condition.‬
‭The pulse strength suggests the strength of ventricular contraction and the amount of‬
‭blood being ejected from the left ventricle. If the pulse is strong, then systolic pressure is‬
‭high. If it is weak, systolic pressure has fallen, and medical intervention may be warranted.‬
‭ ulse can be palpated manually by placing the tips of the fingers across an artery that‬
P
‭runs close to the body surface and pressing lightly. This procedure is typically performed‬
‭using the radial artery in the wrist or the common carotid artery in the neck. However,‬
‭various commercial electronic devices are also available to measure pulse.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭8.9 Career Connection - Cardiologist‬

‭ ardiologists are medical doctors who specialize in the diagnosis and treatment of‬
C
‭diseases of the heart. After completing four years of medical school, cardiologists‬
‭complete a three-year residency in internal medicine, followed by an additional three or‬
‭more years in cardiology. Following these ten years of medical training and clinical‬
‭experience, they qualify for a rigorous two-day examination administered by the Board of‬
‭Internal Medicine that tests their academic training and clinical abilities, including‬
‭diagnostics and treatment. After successful completion of this examination, a physician‬
‭becomes a board-certified cardiologist. Some board-certified cardiologists may be invited‬
‭to become a Fellow of the American College of Cardiology (FACC). This professional‬
‭recognition is awarded to outstanding physicians based on merit, including exceptional‬
‭credentials, achievements, and community contributions to cardiovascular medicine.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 9: Cardiovascular System III -Blood Vessels‬

‭ igure 9.1‬ ‭While most blood vessels are located‬‭deep from the surface and are not‬
F
‭visible, the upper limb's superficial veins indicate the extent, prominence, and importance‬
‭o f these structures to the body. (credit:‬‭Arm veins‬‭- 20090522.jpg‬‭;‬‭Colin Davis‬‭; Wikimedia‬
‭Commons;‬‭CC BY 2.0‬‭)‬

‭9.1 Introduction‬
I‭ n this chapter, you will learn about the vascular part of the cardiovascular system, that is,‬
‭the vessels that transport blood throughout the body and provide the physical site where‬
‭gases, nutrients, and other substances are exchanged with body cells.‬

‭9.2 Types of Blood Vessels and Their Relative Structures‬

‭ here are five categories of blood vessels: artery, arteriole, capillary, venule, and vein‬
T
‭(‬‭Figure 9.2‬‭). An‬‭artery‬‭is a blood vessel that carries‬‭blood away from the heart, where it‬
‭branches into ever-smaller vessels. Eventually, the smallest arteries, vessels called‬
‭arterioles‬‭, further branch into tiny‬‭c apillaries‬‭,‬‭where nutrients and wastes are‬
‭exchanged, and then combine with other vessels that exit capillaries to form‬‭venules‬‭,‬
‭small blood vessels that carry blood to a‬‭vein‬‭, a‬‭larger blood vessel that returns blood to‬
‭the heart’s right atrium.‬

‭ he different types of blood vessels vary slightly in their structures, but they share the‬
T
‭same general features. Arteries and arterioles have thicker walls than veins and venules‬
‭because they are closer to the heart and receive blood surging at a far greater pressure‬

‭234‬
‭Chapter 9: Blood Vessels‬

(‭ ‬‭Figure 9.3‬‭). Each type of vessel has a‬‭lumen‬‭—a hollow passageway through which blood‬
‭flows. Arteries have smaller lumens than veins, a characteristic that helps maintain blood‬
‭pressure moving through the system. Together, their thicker walls and smaller diameters‬
‭give arterial lumens a more rounded appearance in cross-section than the lumens of‬
‭veins.‬

‭ igure 9.2‬ ‭The five types of blood vessels: arteries,‬‭arterioles, capillaries, venules, and‬
F
‭veins. (credit:‬‭Circulation diagram labeling the different‬‭types of blood vessels.png‬‭;‬‭David‬
‭Nascari and Alan Sved‬‭;‬‭CC BY-SA 4.0‬‭). A link to a‬‭video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ apillary beds‬‭contain a large number of capillaries.‬‭Capillaries are narrow-diameter‬

C
‭tubes that can fit red blood cells through in a single file. They are the sites for exchanging‬
‭nutrients, waste, and respiratory gases (e.g., oxygen and carbon dioxide) with tissues at‬
‭the cellular level. Fluid crosses from the capillaries into the spaces between the cells of the‬
‭tissues, called the interstitial space (‬‭Figure 9.4‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 9.3‬ ‭Arteries and veins share the same general‬‭features, but the walls of arteries‬
F
‭are much thicker to deal with the higher pressure of the blood that flows through them‬
‭and the need to maintain blood pressure. The micrograph shows the relative differences‬
‭in thickness. (Micrograph provided by the Regents of the University of Michigan Medical‬
‭School © 2012;‬‭OpenStax‬‭)‬

‭ igure 9.4.‬ ‭The general structure of a capillary.‬ ‭Blood flows through it from the arterial‬
F
‭end (i.e., from an arteriole) and exits at the venous end (i.e., into a venule). Substances‬
‭(e.g., nutrients, respiratory gases, and waste products) can cross the capillary membrane‬
‭and either leave or enter it. (credit:‬ ‭Compartments‬‭o f Capillary Fluid Movement‬‭;‬
‭pressbooks.ccconline.org/). A link to a video explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ asoconstriction, the narrowing of the blood vessels, and‬‭vasodilation‬‭, the widening of‬
V
‭the blood vessels, regulate the movement of materials at the capillaries; this is also‬
‭important in overall blood pressure regulation.‬
‭ hen blood passes through capillaries and enters the venules, the pressure initially‬
W
‭exerted upon it by heart contractions has diminished. In other words, compared to‬
‭arteries, venules and veins withstand a much lower pressure from the blood that flows‬
‭through them. Their walls are considerably thinner, and their lumens are larger in‬
‭diameter, allowing more blood to flow with less resistance.‬

‭9.3 Systemic and Pulmonary Circuits‬

‭ rteries and veins transport blood in two distinct circuits: the‬‭systemic circuit‬‭and the‬
A
‭pulmonary circuit‬‭(‬‭Figure 9.5‬‭).‬

‭ igure 9.5‬ ‭The pulmonary circuit moves blood from‬‭the right side of the heart to the‬
F
‭lungs and back to the heart. The systemic circuit moves blood from the left side of the‬
‭heart to the head and body and returns it to the right side of the heart to repeat the cycle.‬
‭The arrows indicate the direction of blood flow, and the colors show the relative oxygen‬
‭concentration levels. Notice the pulmonary arteries carry blue blood, and the pulmonary‬
‭veins carry red blood, which is the opposite of all other arteries and veins. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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S‭ ystemic arteries provide oxygen-rich blood to the body’s tissues. The blood returned to‬
‭the heart through systemic veins has less oxygen since much of the oxygen carried by the‬
‭arteries has been delivered to the cells to use in cellular respiration (Chapter 6). In‬
‭contrast, in the pulmonary circuit, arteries carry blood low in oxygen exclusively to the‬
‭lungs for gas exchange. Pulmonary veins then return freshly oxygenated blood from the‬
‭lungs to the heart’s left atrium to be pumped back into the systemic circulation.‬

‭9.4 How Blood Flows Through the Body‬

‭ lood is pushed through the body by the action of the pumping heart. With each rhythmic‬
B
‭pump, blood is forced under high pressure and velocity away from the heart, initially‬
‭along the main artery, the aorta. In the aorta, the blood travels at approximately 12 in/sec.‬
‭As blood moves into the arteries, arterioles, and ultimately to the capillary beds, the rate‬
‭o f movement slows dramatically to about 0.01 in/sec, one thousand times slower than the‬
‭rate of movement in the aorta.‬
‭ he slow rate of travel through the capillary beds, which reach almost every cell in the‬
T
‭body, assists with gas, nutrient, and waste exchange and promotes fluid diffusion into the‬
‭interstitial space (i.e., interstitial fluid). Blood flow through the capillary beds is regulated‬
‭depending on the body’s needs and is directed by nerve and hormone signals. For‬
‭example, after a large meal, most of the blood is diverted to the stomach by vasodilation of‬
‭vessels of the digestive system and vasoconstriction of other vessels. During exercise,‬
‭blood is diverted to the skeletal muscles through vasodilation, while blood to the digestive‬
‭system is diminished through vasoconstriction.‬
‭ fter the blood has passed through the capillary beds to the venules, veins, and finally to‬
A
‭the superior or inferior venae cavae, the rate of flow increases again but is still much‬
‭slower than the initial rate in the aorta. Blood primarily moves in the veins by the‬
‭rhythmic movement of smooth muscle in the vessel wall and by the action of the skeletal‬
‭muscle as the body moves. Because most veins must move blood against the pull of‬
‭gravity, blood is prevented from flowing backward in the veins by‬‭one-way valves‬‭.‬
‭Because skeletal muscle contraction aids venous blood flow (‬‭Figure 9.6‬‭), getting up and‬
‭moving frequently after long periods of sitting is essential so that blood will not pool in‬
‭the extremities.‬
S‭ imultaneously, valves inferior to the contracting muscles close; thus, blood should not‬
‭seep back down toward the feet. Military recruits are trained to flex their legs slightly‬
‭while standing at attention for prolonged periods. Failure to do so may allow blood to‬
‭pool in the lower limbs rather than return to the heart. Consequently, the brain will not‬
‭receive enough oxygenated blood, and the individual may lose consciousness. ‬

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‭ igure 9.6‬ ‭The contraction of skeletal muscles surrounding‬‭a vein compresses the blood‬
F
‭and increases the pressure in that area. This action forces blood closer to the heart, where‬
‭venous pressure is lower. Note that one-way valves ensure that blood flows only in the‬
‭proper direction. (credit:‬‭OpenStax‬‭)‬

‭ aricose veins‬‭are a condition in which veins become‬‭enlarged because the valves no‬
V
‭longer close properly, allowing blood to flow backward. Varicose veins are often most‬
‭prominent on the legs (‬‭Figure 9.7‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 9: Blood Vessels‬

‭ igure 9.7‬ ‭Varicose veins are a condition caused‬‭by the failure of one-way valves in veins‬
F
‭that permit blood to pool in them. This condition is most common in the lower extremities,‬
‭such as the legs. (credit: Thomas Kriese;‬‭OpenStax‬‭)‬

‭9.5 Exchange of Fluids at the Capillaries and the Lymphatic System‬

‭ roteins and other large solutes cannot leave the capillaries. The loss of the watery‬
P
‭plasma creates a hypertonic solution within the capillaries, especially near the venules.‬
‭Remember, from Chapter 3, there is net osmosis towards the hypertonic solution in these‬
‭cases. A pressure gradient will also exist in this region, which favors the movement of‬
‭fluids into the lower-pressure venule end of the capillaries. These factors cause about‬
‭85% of the plasma that leaves the capillaries to eventually move back into the capillaries‬
‭near the venules. The remaining 15% of blood plasma drains out from the interstitial fluid‬
‭into nearby‬‭lymphatic vessels‬‭(‬‭Figure 9.8‬‭) and is‬‭now called‬‭lymph fluid‬‭. It passes‬
‭through lymph nodes before it returns to the heart via the vena cava.‬‭Lymph nodes‬‭are‬
‭specialized organs that filter the lymph by percolation through a maze of connective tissue‬
‭filled with white blood cells. The white blood cells remove infectious agents, such as‬
‭bacteria and viruses, to clean the lymph before it returns to the bloodstream. If your‬
‭body is dealing with an infection, one response is swelling of lymph nodes. This is why‬
‭your clinician may check your lymph nodes to see if they are enlarged, which indicates an‬
‭infection.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ he lymph fluid ultimately returns to the heart and rejoins the returning blood near the‬
T
‭venae cavae junction, entering the heart's right atrium.‬

‭ igure 9.8‬ ‭Fluid from the capillaries moves into‬‭the interstitial space and lymph‬
F
‭capillaries by diffusion down a pressure gradient and osmosis. Out of 7,200 liters‬
‭(approximately 1900 gallons) of fluid pumped by the average heart daily, over 1,500 liters‬
‭(400 gallons) are filtered via the lymphatic system. (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭9.6 Cardiac Blood Vessels‬

‭ he heart has blood vessels that supply the heart muscle with blood (‬‭Figure 9.9‬‭). In‬
T
‭o ther words, blood moving through the heart chambers doesn’t meet its circulatory‬
‭needs. The‬‭coronary arteries‬‭branch from the aorta‬‭and surround the heart's outer‬
‭surface like a crown. They diverge into capillaries, where the heart muscle is supplied with‬
‭oxygen and nutrients before converging into the‬‭c ardiac‬‭veins‬‭to take the deoxygenated‬
‭blood back to the right atrium. Here, blood will be re-oxygenated via the pulmonary‬
‭circuit.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 9: Blood Vessels‬

‭ igure 9.9‬ ‭Figure showing the arteries and veins‬‭associated with heart circulation. The‬
F
‭heart tissue has been removed to facilitate this view. (credit: Illustration by J. M. Bourgery‬
‭from Atlas of Human Anatomy and Surgery / Atlas d'antomie Humaine et de Chirurgie by‬
‭Jean Marc Bourgery (1797-1849) Los Angeles: Taschen, 2005. Atlas Case QM 25 .B67‬
‭2005; Crossett Library, Flickr)‬

‭ ike other tissues, the heart muscle will die without a steady blood supply to its arteries.‬
L
‭Atherosclerosis‬‭is the blockage of an artery by the‬‭buildup of fatty plaques and is the‬
‭result of coronary artery disease (‬‭Figures 9.10 and‬‭9.11‬‭). Because of the narrow size of‬
‭the coronary arteries and their function in serving the heart itself, atherosclerosis can be‬
‭deadly in these arteries. The slowdown of blood flow and subsequent oxygen deprivation‬
‭resulting from atherosclerosis causes severe pain, known as‬‭angina pectoris‬‭, and‬
‭complete blockage of the arteries will cause‬‭myocardial‬‭infarction (MI;‬‭Chapter 8),‬
‭commonly known as a heart attack.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 9: Blood Vessels‬

‭ igure 9.10‬ ‭(a) Illustration of a plaque's effect‬‭in narrowing an artery's diameter and‬
F
‭diminishing blood flow. (b) A microscopic view of an artery with a plaque in it. (credit:‬
‭2113ab Atherosclerosis.jpg‬‭; OpenStax College;‬‭CC BY‬‭3.0‬‭). A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭ wo options to treat atherosclerosis of the coronary arteries are bypass surgery and‬
T
‭balloon angioplasty. In the case of‬‭bypass surgery‬‭,‬‭referred to as coronary artery bypass‬
‭graft (CABG), the blockage is “bypassed” using a grafted artery from another location in‬
‭the body (e.g., leg, chest, or arm). The number of bypasses necessary is the basis of‬
‭adjectives you may have heard to describe the CABG surgery: single, double, triple, and‬
‭quadruple (‬‭Figure 9.12‬‭). While it would seem that‬‭removing a small artery from these‬
‭locations might be problematic, other arteries in the areas can compensate, and there isn’t‬
‭an issue.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 9: Blood Vessels‬

‭ igure 9.11‬ ‭In this coronary angiogram (X-ray),‬‭the dye makes two blockages of the‬
F
‭coronary arteries visible. Such blockages can lead to decreased blood flow and insufficient‬
‭oxygen delivered to the cardiac tissues. Uncorrected can lead to cardiac muscle death via a‬
‭myocardial infarction (i.e., heart attack). (credit:‬‭OpenStax‬‭)‬

‭ igure 9.12‬ ‭Four types of coronary artery bypass‬‭surgery (single, double, triple, and‬
F
‭quadruple) are characterized by the number of points of bypass needed to restore‬
‭adequate circulation to the heart. The artery used in the procedure is frequently taken‬
‭from another location in the body. (credit:‬ ‭Blausen.com‬‭staff (2014). "‬‭Medical Gallery of‬
‭Blausen Medical 2014‬‭".‬‭WikiJournal of Medicine‬‭1‬‭(2).‬‭DOI‬‭:‬‭10.15347/wjm/2014.010‬‭.‬‭ISSN‬
‭2002-4436‬‭.‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 9: Blood Vessels‬

I‭ n the case of‬‭balloon angioplasty‬‭, a catheter is inserted into the coronary artery and‬
‭directed to the point of blockage. The balloon is inflated, temporarily blocking blood flow‬
‭through the artery, mimicking symptoms of an MI. The expanded balloon pushes the‬
‭blockage to the vessel's side, restoring most circulation through the artery (‬‭Figure 9.13‬‭).‬
‭Sometimes, a stent (mesh tube made of metal or other materials) may be inserted into the‬
‭artery to provide structural support and prevent it from narrowing again (‬‭Figure 9.14‬‭).‬

‭ igure 9.13‬ ‭Demonstration of the balloon angioplasty‬‭procedure to restore more‬

F
‭normal circulation in a partially blocked coronary artery. (credit:‬‭PTCA NIH.gif‬‭;‬‭National‬
‭Institutes of Health;‬‭CC0 1.0‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 9: Blood Vessels‬

‭ igure 9.14‬ ‭Using a stent to provide structural‬‭support to a coronary artery, preventing‬

F
‭it from narrowing. (credit: Blausen.com staff (2014). "‬‭Medical Gallery of Blausen Medical‬
‭2014‬‭". WikiJournal of Medicine 1 (2).‬‭DOI‬‭:‬‭10.15347/wjm/2014.010‬‭)‬

‭ or more information about both treatments, click the link below or scan the QR code to‬
F
‭watch the TED-Ed video by Krishna Sudhir titled “What happens during a heart attack?”.‬

What happens during a heart attack? - Krishna Sudhir

‭9.7 Strokes‬
‭ heart attack (MI) occurs when the heart muscle is deprived of oxygen, and the tissue‬
A
‭dies. A‬‭stroke‬‭is another medical condition that‬‭is associated with oxygen deprivation.‬
‭However, a stroke impacts blood flow to the brain, not the heart. There are two primary‬
‭stroke classifications: hemorrhagic and ischemic. A‬‭hemorrhagic stroke‬‭o ccurs when a‬
‭blood vessel supplying the brain with freshly oxygenated blood leaks or ruptures, and‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 9: Blood Vessels‬

t‭ here is bleeding into the brain. An‬‭ischemic stroke‬‭, the more common type, occurs when‬
‭brain cells are deprived of oxygen due to a brain artery blockage and is frequently‬
‭associated with blood clots.‬
‭ or more information about strokes, click the link below or scan the QR code to watch the‬
F
‭TED-Ed video by‬‭Vaibhav Goswami titled “What happens‬‭during a stroke?”‬

What happens during a stroke? - Vaibhav Goswami

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 10.1‬ ‭The “thin” air at high elevations, meaning‬‭it contains less oxygen, can strain‬
F
‭the human respiratory system. This is the reason why climbers at high altitudes typically‬
‭use bottled oxygen. (credit: “bortescristian”; Flickr;‬‭OpenStax‬‭)‬

‭10.1 Introduction‬
‭ old your breath. Really! See how long you can hold your breath as you continue reading.‬
H
‭How long did you do it? Chances are it will take less than a minute before you begin to feel‬
‭very uncomfortable. A typical human cannot survive without breathing for more than‬
‭three minutes, and even if you wanted to hold your breath longer, your autonomic‬
‭nervous system would take control, and you would breathe. This outcome is because‬
‭every cell in the body needs oxygen for cellular respiration, specifically oxidative‬
‭phosphorylation (Chapter 6), to produce adenosine triphosphate (ATP). In the chemical‬
‭equation for cellular respiration, oxygen is used as a reactant, and carbon dioxide is‬
‭released as a waste product. You may be surprised to learn that although oxygen is a‬
‭critical need for cells, the accumulation of carbon dioxide primarily drives your need to‬
‭breathe!‬
‭ xygen is inhaled (inhalation), and carbon dioxide is exhaled (exhalation) via the‬
O
‭respiratory system, which includes muscles to move air into and out of the lungs,‬
‭passageways through which air moves, and microscopic gas exchange surfaces covered by‬
‭capillaries. The circulatory system transports gases from the lungs to tissues throughout‬
‭the body and vice versa.‬
‭ arious diseases such as asthma, emphysema, chronic obstructive pulmonary disorder‬
V
‭(COPD), and lung cancer can affect the respiratory system. These conditions affect the gas‬
‭exchange process, resulting in difficulty breathing and other health challenges.‬

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‭10.2 Organs and Structures of the Respiratory System‬

‭ uring‬‭inhalation‬‭, air enters the body through the‬‭nostrils‬‭and moves into the‬‭nasal‬
D
‭c avity‬‭(‬‭Figure 10.2‬‭). As air passes through the nasal‬‭cavity, the air is warmed to body‬
‭temperature and humidified. The respiratory tract is coated with mucus to seal the tissues‬
‭from direct contact with air. The air crossing these surfaces picks up water, as mucus‬
‭contains substantial water. This helps ensure the air is similar to the body's conditions,‬
‭reducing any damage that cold, dry air can cause. Particulate matter floating in the air is‬
‭removed in the nasal passages via mucus and cilia, finger-like extensions of the epithelial‬
‭cells of the surface (Chapter 4). Warming, humidifying, and removing particles are critical‬
‭protective mechanisms that prevent damage to the trachea and lungs. Thus, inhalation‬
‭serves several purposes besides bringing oxygen into the respiratory system.‬
‭ rom the nasal cavity, air passes through the‬‭pharynx‬‭(throat) and the‬‭larynx‬‭(voice box)‬
F
‭as it makes its way to the‬‭trachea‬‭(‬‭Figures 10.2 and‬‭10.3‬‭). The primary function of the‬
‭trachea is to funnel the inhaled air to the lungs and the exhaled air back out of the body.‬
‭The human trachea is a cylinder, about 4 to 5 inches long and 1 inch in diameter. It sits in‬
‭front of the esophagus and extends from the larynx into the chest cavity, dividing into the‬
‭two‬‭primary bronchi‬‭in the middle of the chest. It‬‭is made of incomplete rings of cartilage‬
‭and smooth muscle. The trachea is lined with mucus-producing cells and ciliated epithelia.‬
‭The cilia propel foreign particles trapped in the mucus toward the pharynx. The cartilage‬
‭strengthens and supports the trachea to keep the passage open. The smooth muscle can‬
‭contract, decreasing the trachea’s diameter, which causes expired air to rush upwards‬
‭from the lungs at a great force. The forced exhalation helps expel mucus when we cough.‬
‭Smooth muscle can contract or relax depending on stimuli from the external environment‬
‭o r the body’s nervous system.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 10.2‬ ‭Air enters the respiratory system through‬‭the nasal cavity, and the pharynx‬
F
‭then passes through the trachea and into the bronchi, bringing air into the lungs. (credit:‬
‭2301 Major Respiratory Organs.jpg‬‭; OpenStax College;‬‭CC BY 3.0‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 10.3‬ ‭The trachea and bronchi are made of‬‭incomplete rings of cartilage (white)‬
F
‭and smooth muscle (red). (credit: modification of work by Gray's Anatomy;‬‭OpenStax‬‭). A‬
‭link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ he primary bronchi lead to the right and left lungs, which are not identical. The right lung‬
T
‭is larger and contains three lobes, whereas the smaller one has two (‬‭Figure 10.4‬‭). The‬
‭muscular‬‭diaphragm‬‭, which facilitates breathing, marks‬‭the end of the thoracic cavity and‬
‭separates it from the abdominal cavity below. Many students first learn about the‬
‭diaphragm when experiencing hiccups. For additional information, click the link on the‬
‭next page or scan the QR code to watch the TED-Ed video by‬‭John Cameron titled “Why do‬
‭we hiccup?”.‬

Why do we hiccup? - John Cameron

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 10: Respiratory System‬

‭ he esophagus, discussed in the digestive system (Chapter 5), passes through the‬
T
‭diaphragm to connect with the stomach in the abdominal cavity. The aorta and inferior‬
‭vena cava also pass through the diaphragm, as discussed in Chapters 8 and 9.‬
‭ ach primary bronchus divides into secondary bronchi, then tertiary bronchi, and so on,‬
E
‭creating smaller and smaller diameter‬‭bronchioles‬‭as they split and spread through the‬
‭lung (‬‭Figure 10.3‬‭). Like the trachea, the bronchi‬‭are made of cartilage and smooth muscle.‬
‭At the bronchioles, the cartilage is replaced with elastic fibers. In humans, bronchioles‬
‭with a diameter smaller than 0.5 mm are called‬‭respiratory‬‭bronchioles‬‭. They lack‬
‭cartilage and, therefore, rely on inhaled air to support their shape. As the passageways‬
‭decrease in diameter, the relative amount of smooth muscle increases.‬

‭ igure 10.4‬ ‭The trachea splits into the right and‬‭left primary bronchi, which lead to the‬
F
‭lungs. The right lung is made of three lobes and is larger. To accommodate the heart, the‬
‭left lung is smaller and has only two lobes. Each lung lobe is connected to the primary‬
‭bronchus on that side via a secondary bronchus. (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ he respiratory bronchioles subdivide into several alveolar ducts, surrounded by‬

T
‭numerous alveoli and alveolar sacs. The‬‭alveolar sacs‬‭resemble bunches of grapes‬
‭attached to the end of the bronchioles (‬‭Figure 10.5‬‭).‬‭Gas exchange occurs only in alveoli,‬
‭which consist of a thickness of a single cell. This thin surface is optimal for diffusion‬
‭(Chapter 3). Alveoli are in direct contact with the circulatory system's capillaries (also‬
‭o ne-cell thick). Such intimate contact ensures that oxygen will diffuse from alveoli into the‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ lood and be distributed to the body's cells. In addition, the carbon dioxide produced by‬
b
‭cells as a waste product will diffuse from the blood into the alveoli to be exhaled.‬
‭ he lungs have a sponge-like consistency because there are so many alveoli (~300 million‬
T
‭per lung) within each alveolar sac and so many sacs at the end of each alveolar duct. This‬
‭o rganization produces a vast surface area that is available for gas exchange. This large‬
‭surface area, combined with the thin-walled nature of the alveolar and capillary cells,‬
‭allows gases to diffuse across them quickly.‬

‭ igure 10.5‬ ‭Terminal bronchioles are connected by‬‭respiratory bronchioles to alveolar‬

F
‭ducts and alveolar sacs. Each alveolar sac contains 20 to 30 spherical alveoli and appears‬
‭like a bunch of grapes. Air flows into the alveolar sacs and then circulates into the alveoli,‬
‭where gas exchange occurs with the pulmonary capillaries. Remember from Chapter 9‬
‭that the blood in the pulmonary arteries is low oxygen and high carbon dioxide (blue‬
‭blood), and blood in the pulmonary veins is high oxygen and low carbon dioxide (red‬
‭blood). (credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭10.3 The Mechanics of Breathing‬

‭ uring inhalation, the lungs expand with air (higher oxygen content and lower carbon‬
D
‭dioxide content), and oxygen diffuses across the lung’s surface and enters the‬
‭bloodstream in the pulmonary capillaries. In contrast, carbon dioxide diffuses from the‬
‭bloodstream into the lungs. During exhalation, the lungs expel air (lower oxygen and‬
‭higher carbon dioxide content) and decrease lung volume. The process of human‬
‭breathing will be explained in the following sections.‬

‭Atmospheric and Partial Pressure‬

‭ as movement into or out of the lungs depends on the gas's pressure, a force exerted by a‬
G
‭substance in contact with another.‬‭Atmospheric pressure‬‭is the force exerted by gases in‬
‭the air surrounding any given surface, such as the body.‬ ‭Partial pressure‬‭is the portion‬
‭o f atmospheric pressure contributed by each component in a mixture of gases. For‬
‭example, our atmosphere consists primarily of nitrogen (N), oxygen (O), water (H‬‭2‭O ‬ ), and‬
‭carbon dioxide (CO‬‭2‭)‬ . The total pressure exerted by‬‭the mixture is the sum of the partial‬
‭pressures of the components (P‬‭Atmosphere‬ ‭= P‬‭N‬ ‭+ P‬‭O‬ ‭+ P‬‭H2O‬ ‭+ P‬‭CO2‬‭).‬
‭ tmospheric and partial pressures can be expressed using the unit atmosphere,‬
A
‭abbreviated atm, or in‬‭millimeters of mercury (mm‬‭Hg‬‭). One atm equals 760 mmHg, the‬
‭atmospheric pressure at sea level. The most common source of familiarity with the units‬
‭o f mm Hg is measuring blood pressure (Chapter 8) or listening to a weather report‬
‭describing barometric pressure in units of inches of mercury. For example, you may hear”‬
‭the barometer is rising” or “the barometer is falling.” A‬‭barometer‬‭is a device that‬
‭measures pressure based on the height of a column of mercury that it supports (‬‭Figure‬
‭10.6‬‭). The higher the atmospheric pressure, the greater‬‭the height of the mercury column‬
‭supported. The partial pressures of atmospheric gases are presented in‬‭Figure 10.7‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 10.6‬ ‭A simple barometer used to measure atmospheric pressure. The greater the‬
F
‭pressure, the higher the column of mercury supported. (credit: a modified form of‬
‭MercuryBarometer.svg‬‭;‬‭Danomagnum‬‭;‬‭CC0 1.0‬‭). A link‬‭to a video explanation of this figure‬
‭is available at‬‭Biology411.com‬‭.‬

‭ ercent of the total‬

P ‭ artial pressure‬
P
‭ as‬
G ‭composition‬ ‭(mm Hg)‬
‭Nitrogen (N‬‭2‭)‬ ‬ ‭78.6‬ ‭597.4‬
‭Oxygen (O‬‭2‬‭)‬ ‭20.9‬ ‭158.8‬
‭Water (H‬‭2‭O
‬ )‬ ‭0.4‬ ‭3.0‬
‭Carbon dioxide (CO‬‭2‭)‬ ‬ ‭0.04‬ ‭0.3‬
‭Others‬ ‭0.06‬ ‭0.5‬
‭ otal‬
T ‭100%‬ ‭760.0‬
‭composition/total‬
‭atmospheric pressure‬

‭ igure 10.7‬ ‭The partial pressures of the component‬‭atmospheric gases and‬

F
‭determination of the total atmospheric pressure. As oxygen is 20.9% of the total air‬
‭composition, its partial pressure is obtained by multiplying the total partial pressure (760‬
‭mmHg) by the decimal form of the percentage (0.209). The obtained value is 158.8‬
‭mmHg.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 10: Respiratory System‬

‭ or additional information about barometers and related topics, click the link below or‬
F
‭scan the QR code to watch the TED-Ed video by Asaf Bar-Yosef titled “The history of the‬
‭barometer and how it works.”‬

The history of the barometer (and how it works) - Asaf Bar-Yos…

‭Boyle’s Law‬
‭ o understand gas exchange in the lungs, it is necessary to understand the underlying‬
T
‭principles of gases and their behavior. One of these principles was developed by Robert‬
‭Boyle, a research scientist, and is called‬‭Boyle’s‬‭Law‬‭. This law states that gas pressure (P)‬
‭and volume (V) are inversely proportional in a closed space and constant temperature.‬
I‭ n other words, as gas volume decreases, gas pressure increases proportionately and vice‬
‭versa (‬‭Figure 10.8‬‭). Boyle’s Law is represented by‬‭the mathematical equation‬ ‭P‬‭1‭V
‬ ‬‭1‬ ‭=‬
‭P‬‭2‭V ‬
‬ ‭2‬,
‭ where 1 indicates initial conditions, and 2‬‭
i ndicates final conditions.‬

‭ igure 10.8‬ ‭Data from Robert Boyle’s original 1662‬‭experiment shows that pressure and‬
F
‭volume are inversely related. No units are given, as Boyle used arbitrary units in his‬
‭experiments. (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭256‬
‭Chapter 10: Respiratory System‬

‭ et’s consider a specific example of Boyle’s Law. Assume a gas container's initial pressure‬
L
‭( P‬‭1‬‭) and volume (V‬‭1‬‭) are 760 mm Hg and 2 gallons,‬‭respectively. The container's final‬
‭volume (V‬‭2‬ ‭) is reduced to 1 gallon (i.e., reduced‬‭by a factor of ½). Therefore, P‬‭2‬ ‭would‬
‭equal P‬‭1‬‭*V‬‭1‬‭/V‬‭2‭,‬ which is (760 mm Hg) * (2 gallons)/1‬‭gallon, which is 1,520 mm Hg.‬
‭Therefore, the pressure in the final conditions (P‬‭2‭)‬ ‬‭is twice as much as in the initial‬
‭conditions (P‬‭1‭)‬ . Note that the inverse of 1/2 is‬‭2/1. In other words, when the gas volume‬
‭is decreased by half, the pressure increases by the inverse proportion; in this example, it‬
‭doubles.‬

‭Inhalation and Exhalation‬

‭ he relationship between gas pressure and volume, defined by Boyle’s Law, helps explain‬
T
‭breathing mechanics associated with inhalation and exhalation. During inhalation, the‬
‭volume of the thoracic cavity increases as a result of contraction of the diaphragm‬
‭(flattens) and the intercostal muscles, which are connected to the rib cage (expands the‬
‭rib cage) (‬‭Figure 10.10a‬‭). Because the pressure in‬‭the thoracic cavity decreases, the‬
‭environment's air pressure is now greater than in the thoracic cavity, so air moves along‬
‭its pressure gradient (higher to lower) and rushes into the respiratory passages and the‬
‭lungs. Upon exhalation, the intercostal muscles relax, returning the chest wall to its‬
‭o riginal position (‬‭Figure 10.10b‬‭). The diaphragm also‬‭relaxes and moves higher into the‬
‭thoracic cavity. This action increases the pressure within the thoracic cavity relative to the‬
‭environment, and air rushes out of the respiratory passages and lungs. The air movement‬
‭o ut of the lungs is passive; no muscles contract to expel the air.‬
‭ or additional information about breathing mechanics, click the link below to watch the‬
F
‭TED-Ed video by‬‭John Cameron titled “How breathing works.”‬

How breathing works - Nirvair Kaur

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭257‬
‭Chapter 10: Respiratory System‬

‭ igure 10.9‬ ‭An example of Boyle's Law, which states‬‭that gas pressure (P) and volume‬
F
‭(V) are inversely proportional at a constant temperature. The initial conditions were‬
‭changed by reducing the container volume by half (2 gallons to 1 gallon), which resulted‬
‭in the pressure doubling (760 mm Hg to 1520 mm Hg). (credit:‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

I‭ f any condition diminishes a person’s ability to breathe (e.g., surgery, infections, lung‬
‭diseases, heart failure, or pulmonary edema), the person will experience some degree of‬
‭respiratory distress. In such situations, a device called a‬‭ventilator‬‭(breathing machine)‬
‭may be used to assist the person.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 10.10‬ ‭The lungs, chest wall, and diaphragm‬‭are all involved in breathing; (a)‬
F
‭inhalation and (b) exhalation. (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ or additional information about ventilators, click the link below or scan the QR code to‬
F
‭watch the TED-Ed video by‬‭Alex Gendler titled “How‬‭do ventilators work?.”‬

How do ventilators work? - Alex Gendler

‭10.4 Basic Principles of Respiratory Gas Exchange‬

‭ he other primary function of the lungs is oxygen and carbon dioxide exchange. As‬
T
‭discussed previously, cells need oxygen during cellular respiration and produce carbon‬
‭dioxide, which is a waste product.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ as exchange during respiration occurs primarily through‬‭simple diffusion‬‭(Chapter 3).‬

G
‭Diffusion is a process in which transport is driven by a concentration (or partial pressure)‬
‭gradient. Gas molecules move from a region of high concentration (high partial pressure)‬
‭to a low concentration (low partial pressure). Blood that is low in oxygen concentration‬
‭and high in carbon dioxide concentration undergoes gas exchange with air in the lungs.‬
‭The air in the lungs has a higher concentration of oxygen than that of oxygen-depleted‬
‭blood and a lower concentration of carbon dioxide. These concentration (or partial‬
‭pressure) gradients allow oxygen to diffuse from the alveolar sacs into the pulmonary‬
‭capillaries and carbon dioxide from the pulmonary capillaries into the alveolar sacs.‬
I‭ n our discussion of diffusion (Chapter 3), you learned that molecules move from an area‬
‭o f higher concentration to a lower concentration. In the case of gases, they move from‬
‭higher partial pressure (measured in mmHg) to lower partial pressure. In other words,‬
‭oxygen and carbon dioxide move with their concentration gradients, which are also partial‬
‭pressure gradients. As with concentration gradients, the more significant the partial‬
‭pressure difference between the two areas, the more rapid the gas movement. Because‬
‭both oxygen and carbon dioxide are small, nonpolar molecules (Chapter 2), they freely‬
‭diffuse directly across the phospholipid bilayer of the cells’ plasma membranes.‬
‭ as exchange occurs at two sites in the body: in the lungs, where oxygen is picked up and‬
G
‭carbon dioxide is released, and in the tissues, where oxygen is released, and carbon‬
‭dioxide is picked up.‬‭External respiration‬‭is the‬‭exchange of gases with the external‬
‭environment and occurs between the lungs' alveoli and the pulmonary capillaries.‬
‭Internal respiration‬‭is the exchange of gases with‬‭the internal environment and occurs in‬
‭the tissues.‬

‭ 0.5 Oxygen and Carbon Dioxide Transport in the Blood and Exchange During‬
1
‭External and Internal Respiration‬

‭Blood Transport of Oxygen‬

‭ ost oxygen molecules are carried from the lungs to the body’s tissues by a specialized‬
M
‭transport system, which relies on erythrocytes—the red blood cells. As discussed in‬
‭Chapter 7, erythrocytes contain‬‭hemoglobin‬‭, which‬‭binds oxygen molecules (‬‭Figure‬
‭10.11‬‭).‬‭Heme‬‭is the‬‭iron‬‭-containing portion that chemically‬‭binds oxygen. Each‬
‭hemoglobin molecule contains four iron-containing heme molecules, and because of this,‬
‭o ne hemoglobin molecule can carry up to four molecules of oxygen.‬
‭ s oxygen diffuses across the respiratory membrane from the alveoli (review‬‭Figure‬
A
‭10.5‬‭) to the capillary, it also diffuses into the‬‭red blood cell and is bound by hemoglobin.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 10.11‬ ‭Each erythrocyte contains approximately‬‭250 - 300 million molecules of‬
F
‭hemoglobin. The figure shows one molecule composed of four protein subunits, each‬
‭containing one iron ion as part of the heme group (represented by the green disk).‬
‭(credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ he following reversible chemical reaction describes the production of the final product,‬
T
‭oxyhemoglobin‬‭(HbO‬‭2‭)‬ , which is formed when oxygen‬‭binds to hemoglobin (Hb)‬
‭𝐻𝑏‬‭‬ + ‭‬‭𝑂‬‭2‬ ↔ ‭𝐻𝑏‬‭𝑂‬‭2‬

‭ xyhemoglobin is a bright red-colored molecule that contributes to the bright red color of‬
O
‭oxygenated blood. Venous blood, associated with‬‭deoxyhemoglobin‬‭(Hb), is darker red‬
‭(‬‭Figure 10.12‬‭). In other words, oxyhemoglobin becomes‬‭deoxyhemoglobin at the‬
‭systemic capillaries when it releases oxygen.‬
‭ hen considering the blood as a whole, the percent of the available heme units bound to‬
W
‭oxygen at a given time is called‬‭hemoglobin saturation‬‭.‬‭A hemoglobin saturation of 100‬
‭percent means that every heme unit in the body's erythrocytes is bound to oxygen. In a‬
‭healthy individual with normal hemoglobin levels, hemoglobin saturation generally ranges‬
‭from 95 percent to 99 percent.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 10.12‬ ‭Comparison of an arterial (oxyhemoglobin)‬‭blood sample and venous‬

F
‭(deoxyhemoglobin) blood sample. Notice that the oxyhemoglobin (‬‭𝐻𝑏‬‭𝑂‬‭2‭)‬ is a bright red‬
c‭ olor compared to the darker red color of deoxyhemoglobin (‬‭𝐻𝑏‬‭). (credit:‬‭Arterial and‬
‭Venous Blood Model‬‭; Yui.kubo;‬‭CC BY-NC-SA 4.0‬‭)‬

‭Blood Transport of Carbon Dioxide‬

‭ arbon dioxide molecules are transported in the blood from body tissues to the lungs by‬
C
‭two primary methods. First, carbon dioxide can enter red blood cells and reversibly bind‬
‭to hemoglobin to form a‬‭c arbaminohemoglobin‬‭molecule.‬‭Therefore, when it reaches the‬
‭lungs, the carbon dioxide can freely dissociate from the hemoglobin, diffuse into the‬
‭alveoli, and be expelled from the body via exhalation. This form transports a relatively‬
‭small amount of carbon dioxide.‬
‭ ost carbon dioxide molecules are transported by a second method called the‬
M
‭bicarbonate buffer system‬‭. In this system, carbon‬‭dioxide diffuses into the red blood‬
‭cells.‬‭Carbonic anhydrase (CA)‬‭, an enzyme within the‬‭red blood cells, quickly catalyzes‬
‭the reaction that converts carbon dioxide into carbonic acid (H‬‭2‭C
‬ O‬‭3‭)
‬ . Carbonic acid is an‬
−
‭unstable intermediate molecule that immediately splits into‬‭bicarbonate‬(‭𝐻𝐶𝑂‬‭3‬) ‭and‬
‭ ydrogen (H‬‭+‭)‬ ions. Since carbon dioxide is quickly‬‭converted into bicarbonate ions, this‬
h
‭reaction allows for the continued uptake of carbon dioxide into the blood down its‬
‭concentration gradient. It also results in the production of H‬‭+‬ ‭ions, which could alter the‬
‭pH of the blood, making it more acidic (Chapter 2). However, hemoglobin binds to the‬
‭free H‬‭+‬ ‭ions and thus limits pH changes.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ he newly synthesized bicarbonate ion is transported from the red blood cell into the‬
T
‭plasma. When the blood reaches the lungs, the bicarbonate ion is transported back into‬
‭the red blood cell, the H‬‭+‬ ‭ion dissociates from the‬‭hemoglobin, and the two bind together‬
‭to recreate the carbonic acid intermediate, which is converted back into carbon dioxide‬
‭through the enzymatic action of CA. The carbon dioxide produced is expelled through the‬
‭lungs during exhalation:‬

‭CA‬

‭CO‬‭2‬ ‭+ H‬‭2‭O
‬ ↔ H‬‭2‭C
‬ O‬‭3‬ ‭↔‬ ‭HCO‬‭3‬‭-‬ ‭+ H‬‭+‬
‭(carbonic acid ) (bicarbonate)‬

‭External Respiration of Oxygen and Carbon Dioxide‬

I‭ n‬‭external respiration‬‭, the pulmonary arteries carry‬‭deoxygenated blood into the lungs‬
‭from the heart, where it branches and eventually becomes the capillary network‬
‭composed of pulmonary capillaries. These pulmonary capillaries create the respiratory‬
‭membrane with the alveoli (‬‭Figure 10.13‬‭). As the blood‬‭is pumped through this capillary‬
‭network, gas exchange occurs due to partial pressure differences in carbon dioxide and‬
‭oxygen between the alveoli and the blood in the pulmonary capillaries. Carbon dioxide‬
‭diffuses from the pulmonary capillaries into the alveolar sacs, where it will be exhaled‬
‭from the body. Oxygen diffuses from the alveolar sacs into the pulmonary capillaries,‬
‭where the majority binds with deoxyhemoglobin to create oxyhemoglobin. Oxygenated‬
‭hemoglobin returns to the heart's left atrium via the pulmonary veins.‬

‭Internal Respiration of Oxygen and Carbon Dioxide‬

I‭ nternal respiration is the gas exchange that occurs at the level of body tissues (‬‭Figure‬
‭10.14‬‭). Like external respiration, internal respiration‬‭o ccurs as simple diffusion due to‬
‭partial pressure gradients. However, the gradients are opposite of those present in the‬
‭respiratory membranes in the lungs. Oxygen dissociates from hemoglobin, diffuses out of‬
‭the blood, crosses the interstitial space, and enters the cells of the tissues, where it is used‬
‭for cellular respiration and ATP production. Carbon dioxide, a waste product of this‬
‭process, will diffuse from the tissues’ cells into the interstitial fluid and enter the blood‬
‭(Chapter 6).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 10.13‬ ‭In external respiration, oxygen diffuses‬‭across the respiratory membrane‬
F
‭from the alveolus to the capillary, whereas carbon dioxide diffuses from the capillary into‬
‭the alveolus. Correction: (credit:‬‭Respiratory Cardiovascular‬‭Junction‬‭;‬‭Cenveo;‬‭CC BY 3.0‬‭).‬
‭A link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 10: Respiratory System‬

‭ igure 10.14‬ ‭During internal respiration, oxygen‬‭diffuses out of the capillary and into‬
F
‭tissue cells, whereas carbon dioxide diffuses out of cells and into the capillary. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ he blood moves back to the heart and then to the lungs for gas exchange during external‬
T
‭respiration. This cycle of external respiration, followed by internal respiration,‬
‭continuously repeats.‬

‭10.6 Medical Issues Associated with the Respiratory System‬

‭Asthma‬

‭ sthma‬‭is an example of an obstructive condition affecting‬‭lung air passageways in adults‬

A
‭and children. Approximately 8.2 percent of adults (18.7 million) and 9.4 percent of‬
‭children (7 million) in the United States have asthma. In addition, asthma is the most‬
‭frequent cause of hospitalization in children.‬

‭ sthma is a chronic disease characterized by inflammation and bronchospasms‬

A
‭(constriction of the smooth muscle associated with the bronchioles), which can inhibit air‬
‭from entering the lungs. In addition, excessive mucus secretion can occur, further‬
‭contributing to the narrowing of the airways. Bronchospasms occur periodically and lead‬
‭to an “asthma attack.” An attack may be triggered by environmental factors such as dust,‬
‭pollen, pet hair, or dander, changes in the weather, mold, tobacco smoke, and respiratory‬
‭infections, or by exercise and stress.‬

S‭ ymptoms of an asthma attack involve coughing, shortness of breath, wheezing, and chest‬
‭tightness. Symptoms of a severe asthma attack, which require immediate medical attention,‬
‭include difficulty breathing, resulting in blue (cyanotic) lips or face, confusion, drowsiness,‬
‭rapid pulse, sweating, and severe anxiety. The severity of the condition, frequency of‬
‭attacks, and identified triggers influence the type of medication an individual may require.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 10: Respiratory System‬

‭ onger-term treatments are used for those with more severe asthma. Short-term,‬
L
‭fast-acting drugs that treat an asthma attack are typically administered via an inhaler.‬
‭Asthma medications can be administered via a nebulizer for young children or individuals‬
‭with difficulty using an inhaler.‬

‭ or additional information about asthma, click the link below or scan the QR code to watch‬
F
‭the TED-Ed video by‬‭Christopher E. Gaw titled “How‬‭does asthma work?.”‬

How does asthma work? - Christopher E. Gaw

‭ igure 10.15‬ ‭A visual comparison of the bronchioles‬‭(normal versus asthmatic). Note‬

F
‭the thickened airway wall and mucus in the affected passageway. (credit:‬‭pnash‬‭; Flickr;‬‭CC‬
‭BY-NC-ND 2.0‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Pneumonia‬

‭ neumonia‬‭is a general term for infections of the‬‭lungs that lead to inflammation and‬
P
‭accumulation of fluids and white blood cells in the alveoli (‬‭Figure 10.16‬‭). Pneumonia can‬
‭be caused by bacteria, viruses, fungi, and other organisms, although most pneumonias are‬
‭bacterial in origin. As the alveoli fill with fluids and white blood cells, air exchange‬
‭becomes impaired, and patients experience respiratory distress. In addition, pneumonia‬
‭can lead to pleurisy, an infection of the pleural membrane surrounding the lungs, making‬
‭breathing very painful.‬ ‭Antibiotics remain the mainstay‬‭treatment for bacterial‬
‭pneumonia, although antibiotic-resistant bacterial strains are becoming more prevalent‬
‭and problematic.‬

‭ igure 10.16‬ ‭Pneumonia caused by bacteria (e.g.,‬‭Streptococcus pneumoniae‬‭). (a).‬

F
‭pneumonia affecting part of the left lung. (b). healthy alveoli (air sacs). (c). alveoli filled‬
‭with mucus. (credit:‬‭Lobar pneumonia illustrated.jpg‬‭;‬‭nhlbi.nih.gov/health/pneumonia;‬
‭CC0 1.0‬‭).‬ ‭A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭For additional information a‬‭bout pneumonia, click the link below or scan the QR code to‬
‭ atch the TED-Ed video by‬‭Eve Gaus and Vanessa Ruiz‬‭titled “Why is pneumonia so‬
w
‭dangerous?”.‬

Why is pneumonia so dangerous? - Eve Gaus and Vanessa Ruiz

‭Carbon Monoxide Poisoning‬

‭ hile carbon dioxide can readily associate and dissociate from hemoglobin, other‬
W
‭molecules, such as carbon monoxide (CO), cannot. Carbon monoxide has a greater affinity‬
‭for hemoglobin than oxygen. Therefore, when carbon monoxide is present, it binds to‬
‭hemoglobin preferentially over oxygen. As a result, oxygen cannot bind to hemoglobin,‬
‭very little oxygen is transported through the body, and‬‭c arbon monoxide poisoning‬
‭results (‬‭Figure 10.17‬‭). Carbon monoxide is a colorless,‬‭o dorless gas, making it difficult to‬
‭detect. It is essential to have a carbon monoxide detector installed in your home if you‬
‭have a natural gas, oil, or propane heating source or appliances. Short-term exposure to‬
‭carbon monoxide can cause headaches, confusion, and nausea; long-term exposure can‬
‭cause brain damage or death. Administering 100 percent (pure) oxygen is the usual‬
‭treatment for carbon monoxide poisoning because it speeds up carbon monoxide‬
‭separation from hemoglobin. A blood transfusion is also a possible remedy.‬

‭ igure 10.17‬ ‭As the percent of carbon monoxide (CO)‬‭increases in the blood (X-axis), the‬
F
‭oxygen saturation of hemoglobin decreases (Y-axis). (credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Sleep Apnea‬

‭ leep apnea‬‭is a chronic disorder that can occur in‬‭children or adults and is characterized‬
S
‭by the cessation of breathing during sleep. These episodes may last for several seconds or‬
‭several minutes and may differ in the frequency with which they are experienced. Sleep‬
‭apnea leads to poor sleep, which is reflected in the symptoms of fatigue, evening napping,‬
‭irritability, memory problems, and morning headaches. In addition, many individuals with‬
‭sleep apnea experience a dry throat in the morning after waking from sleep, possibly due‬
‭to excessive snoring.‬

‭ bstructive sleep apnea is the most common type of sleep apnea. It is caused by an airway‬
O
‭o bstruction during sleep, which can occur at different points in the airway, depending on‬
‭the underlying cause. For example, the tongue and throat muscles of some individuals‬
‭with obstructive sleep apnea may relax excessively, causing the muscles to push into the‬
‭airway. Another example is obesity, a known risk factor for sleep apnea, as excess adipose‬
‭tissue in the neck region can push the soft tissues toward the lumen of the airway, causing‬
‭the trachea to narrow.‬

‭ diagnosis of obstructive sleep apnea is usually made during a sleep study, where the‬
A
‭patient is monitored in a sleep laboratory for several nights or via a test at home. The‬
‭patient’s blood oxygen levels, heart rate, respiratory rate, and blood pressure are‬
‭monitored, as are brain activity and air volume inhaled and exhaled. Sleep apnea‬
‭treatment commonly includes using a‬‭continuous positive‬‭airway pressure (CPAP)‬
‭machine‬‭during sleep. The CPAP machine has a mask‬‭that covers the nose, or the nose‬
‭and mouth, and forces air into the airway at regular intervals. This pressurized air can‬
‭help gently force the airway to remain open, allowing more normal ventilation. Other‬
‭treatments include lifestyle changes to decrease weight, eliminate alcohol and other sleep‬
‭apnea–promoting drugs, and changes in sleep position.‬

‭ or additional information about snoring and sleep apnea, click the link below or scan the‬
F
‭QR code to watch the TED-Ed video by‬‭Alayna Vaughan‬‭titled “The sleep disorder you‬
‭might not know you have.”‬

The sleep disorder you might not know you have - Alayna Vaughan

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Decompression Sickness‬
‭ ecompression sickness‬‭(DCS) occurs when gases dissolved‬‭in the blood or other body‬
D
‭tissues are no longer dissolved following a reduction in pressure on the body. This‬
‭condition affects underwater divers who surface from a deep dive too quickly, and it can‬
‭affect pilots flying at high altitudes in planes with unpressurized cabins. Divers often call‬
‭this condition “the bends,” a reference to joint pain that is a symptom of DCS.‬

I‭ n all cases, DCS is brought about by a reduction in barometric pressure. At high altitudes,‬
‭barometric pressure is much less than on Earth’s surface because pressure is produced‬
‭by the weight of the column of air above the body pressing down on the body. The‬
‭substantial pressures on divers in deep water are likewise from the weight of a column of‬
‭water pressing down on the body. For divers, DCS occurs at standard barometric pressure‬
‭(at sea level). Still, it is brought on by the relatively rapid decrease of pressure as divers‬
‭rise from the high-pressure conditions of deep water to the now low, by comparison,‬
‭pressure at sea level.‬

I‭ n DCS, gases dissolved in the blood (primarily nitrogen) come rapidly out of solution,‬
‭forming bubbles in the blood and other body tissues. This occurs because when the‬
‭pressure of a gas over a liquid is decreased, the amount of gas that can remain dissolved‬
‭in the liquid also decreases. Air pressure keeps your normal blood gases dissolved in the‬
‭blood. When pressure is reduced, less gas remains dissolved. You can see this effect when‬
‭you open a carbonated drink. Removing the bottle's seal reduces the gas pressure over‬
‭the liquid. This, in turn, causes bubbles as dissolved gases (in this case, carbon dioxide)‬
‭come out of solution in the liquid.‬

‭ he most common symptoms of DCS (e.g., joint pain, headaches, and vision disturbances)‬
T
‭o ccur in 10 to 15 percent of cases. Left untreated, very severe DCS can result in death.‬
‭Immediate treatment is with pure oxygen. The affected person is then moved into a‬
‭hyperbaric chamber (‬‭Figure 10.18‬‭), a reinforced, closed‬‭chamber pressurized to greater‬
‭than atmospheric pressure. It treats DCS by repressurizing the body to remove pressure‬
‭gradually. Because the‬‭hyperbaric chamber‬‭introduces‬‭oxygen to the body at high‬
‭pressure, it increases the oxygen concentration in the blood. This replaces some of the‬
‭nitrogen in the blood with oxygen, which is easier to tolerate out of solution.‬

‭ nother example of using a hyperbaric chamber is the treatment of anaerobic bacterial‬

A
‭infections created by bacteria that cannot or prefer not to live in the presence of oxygen.‬
‭An increase in blood and tissue levels of oxygen helps kill the anaerobic bacteria‬
‭responsible for the infection, as oxygen is toxic to anaerobic bacteria. The chamber‬
‭stimulates the healing process for wounds and grafts by increasing the energy production‬
‭needed for repair. Increasing oxygen transport allows cells to ramp up cellular respiration‬
‭and thus ATP production, the energy required to build new structures.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 10.18‬ ‭An example of a hyperbaric chamber,‬‭which is used to treat DCS (credit:‬
F
‭“komunews”; Flickr;‬‭OpenStax‬‭)‬

‭Acute Mountain Sickness‬

‭ cute mountain sickness‬‭(AMS), or altitude sickness,‬‭is a condition that results from‬

A
‭acute exposure to high altitudes due to the low partial pressure of oxygen at high‬
‭altitudes. AMS typically can occur at 2400 meters (8000 feet) above sea level due to low‬
‭blood oxygen levels, as the body has difficulty adjusting to the low partial pressure of‬
‭oxygen (‬‭Figure 10.19‬‭). Hemoglobin saturation is lower‬‭at high altitudes than at sea level.‬
‭For example, hemoglobin saturation is about 67 percent at 19,000 feet above sea level,‬
‭whereas it reaches approximately 98 percent at sea level.‬

I‭ n severe cases, AMS can cause pulmonary or cerebral edema. Symptoms of AMS include‬
‭nausea, vomiting, fatigue, lightheadedness, drowsiness, feeling disoriented, increased‬
‭pulse, and nosebleeds. The only treatment for AMS is descending to a lower altitude;‬
‭however, pharmacologic treatments and supplemental oxygen can improve symptoms.‬

‭ cclimatization can prevent AMS, which is the adjustment the respiratory system makes‬
A
‭due to chronic exposure to a high altitude. Over time, the body adjusts to accommodate‬
‭the lower partial pressure of oxygen. The low partial pressure of oxygen at high altitudes‬
‭results in a lower oxygen saturation level of hemoglobin in the blood. In turn, the tissue‬
‭levels of oxygen are also lower. As a result, the kidneys are stimulated to produce the‬
‭hormone‬‭erythropoietin‬‭(‬‭EPO‬‭), which stimulates the‬‭production of erythrocytes,‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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r‭ esulting in a greater number of circulating erythrocytes in an individual at a high altitude‬

‭over a long period. With more red blood cells, there is more hemoglobin to help transport‬
‭the available oxygen. Even though there is low saturation of each hemoglobin molecule,‬
‭more hemoglobin will be present, and therefore more oxygen in the blood. Over time, this‬
‭allows the person to partake in physical exertion without developing AMS.‬

‭Altitude‬ ‭Atmospheric‬ ‭Partial pressure of‬

‭Example location‬ ‭(feet above sea level)‬ ‭pressure‬‭(mm Hg)‬ ‭oxygen‬‭(mm Hg)‬

‭ ew York City, NY‬

N ‭‬
0 ‭ 60‬
7 ‭ 59‬
1
‭Boulder, Colorado‬ ‭5000‬ ‭632‬ ‭133‬
‭Aspen, Colorado‬ ‭8000‬ ‭565‬ ‭118‬
‭Pikes Peak,‬ ‭14,000‬ ‭447‬ ‭94‬
‭Colorado‬
‭Denali (Mt.‬ ‭20,000‬ ‭350‬ ‭73‬
‭McKinley), Alaska‬
‭Mt. Everest, Tibet‬ ‭29,000‬ ‭260‬ ‭54‬

‭ igure 10.19‬ ‭Partial pressure values of oxygen at‬‭different altitudes. Note that as the‬
F
‭altitude increases, the partial pressure of oxygen decreases.‬

‭ or additional information about AMS, click the link or scan the QR code to watch the‬
F
‭TED-Ed video by‬‭Andrew Lovering titled “What happens to your body at the top of Mount‬
‭Everest?.”‬

What happens to your body at the top of Mount Everest - …

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭10.7 Career Connection - Respiratory Therapist‬

‭ espiratory therapists or respiratory practitioners evaluate and treat patients with lung‬
R
‭and cardiovascular diseases. They work as part of a medical team to develop patient‬
‭treatment plans. Respiratory therapists may treat premature babies with underdeveloped‬
‭lungs, patients with chronic conditions such as asthma, or older patients suffering from‬
‭lung diseases such as emphysema and chronic obstructive pulmonary disease (COPD).‬
‭They may operate advanced equipment such as compressed gas delivery systems,‬
‭ventilators, blood gas analyzers, and resuscitators. Specialized programs to become a‬
‭respiratory therapist generally lead to a bachelor’s degree with a respiratory therapist‬
‭specialty. Because of the growing aging population, career opportunities for respiratory‬
‭therapists are expected to remain strong.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 11.1‬ ‭While glands in our bodies naturally‬‭produce hormones, some can also be‬
F
‭made using recombinant DNA (rDNA) technology. For example, insulin is produced by‬
‭specialized pancreas cells via transcription and translation of the gene. However, when‬
‭the insulin gene was isolated and inserted into the bacterial genome, these cells produced‬
‭the hormone, which is isolated and purified. Humulin is a commercially available insulin‬
‭product used by people with diabetes to regulate blood sugar levels, and an insulin pen‬
‭for injecting a recombinant insulin, Humulin”, is shown above. (credit:‬‭Human insulin 100‬
‭IU-1ml pen yellow background (02).jpg‬‭;‬‭Wesalius‬‭;‬‭CC‬‭BY 4.0‬‭)‬

‭11.1 Introduction‬
‭ ommunication is when a sender transmits signals to one or more receivers to control‬
C
‭and coordinate actions. In the human body, two major organ systems participate in‬
‭relatively “long-distance” communication: the nervous system (via‬‭neural signaling‬‭) and‬
‭the endocrine system (via‬‭endocrine signaling‬‭). Together,‬‭these two systems are‬
‭primarily responsible for maintaining homeostasis in the body.‬

‭11.2 Endocrine Signaling‬

‭ he‬‭endocrine system‬‭uses chemical signaling to bring‬‭about changes. These signals,‬
T
‭called‬‭hormones‬‭, are produced by cells, tissues, and‬‭o rgans and secreted into the‬
‭extracellular fluid. The hormones are then transported throughout the body, primarily via‬
‭the bloodstream, where they bind to‬‭receptors‬‭o n target‬‭cells. A hormone’s‬‭target cells‬
‭have appropriate receptors to attach to the hormone, like a key that fits a lock. Once a‬
‭hormone binds to a receptor, a chain of events initiates the target cell’s response (‬‭Figure‬
‭11.2‬‭).‬

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‭ igure 11.2‬‭The interaction between chemical messengers‬‭(e.g., hormones; 1) and cell‬

F
‭membrane-bound receptors (2). If successful, then a detailed response is initiated (3)‬
‭(Credit:‬‭Receptor (Biochemistry).svg‬‭);‬‭Wyatt Pyzynsk‬‭i‬‭;‬‭CC BY-SA 4.0‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ ndocrine signaling requires more time than neural signaling to prompt a response in‬
E
‭target cells, though the precise amount of time varies, depending on the hormone. For‬
‭example, the hormones released when confronted with a dangerous or frightening‬
‭situation, the‬‭fight-or-flight response‬‭, occur by‬‭releasing the adrenal‬
‭hormones—‬‭epinephrine‬‭and‬‭norepinephrine‬‭—within seconds.‬‭The two hormones‬
‭dilate blood vessels, increase the heart and respiratory rate, and suppress the digestive‬
‭and immune systems. These responses boost the body’s oxygen transport to the brain‬
‭and muscles, thereby improving the body’s ability to fight or flee (flight). In contrast, it‬
‭may take up to 48 hours for target cells to respond to certain reproductive hormones.‬
I‭ n addition, endocrine signaling is typically less specific than neural signaling. Depending‬
‭o n the target cells involved, the same hormone may play a role in various physiological‬
‭processes. For example, the hormone‬‭oxytocin (OT)‬‭promotes uterine contractions in‬
‭women in labor. It is also important in breastfeeding and may be involved in the sexual‬
‭response and in feelings of emotional attachment in both males and females.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭11.3 Structures of the Endocrine System‬

‭ ndocrine glands‬‭are the major players in this system.‬‭The primary function of these‬
E
‭ductless glands is to secrete their hormones directly into the surrounding interstitial fluid‬
‭and then into the capillaries. Once in the blood, the circulatory system transports‬
‭hormones throughout the body. The endocrine system includes the hypothalamus,‬
‭pituitary, thymus, thyroid, adrenal, and pineal glands (‬‭Figure 11.3‬‭). Some of these glands‬
‭have both endocrine and nonendocrine functions. For example, the pancreas contains‬
‭cells that function in digestion and cells that secrete the hormones insulin and glucagon,‬
‭which regulate blood glucose levels.‬
‭ ther organs that contain cells that secrete hormones are the heart, kidneys, stomach,‬
O
‭small intestine, liver, skin, female ovaries, and male testes. Moreover, some tissues (e.g.,‬
‭adipose and bone) also have endocrine functions.‬

‭ igure 11.3‬ ‭Endocrine glands and cells are located‬‭throughout the body and play an‬
F
‭essential role in homeostasis. (credit:‬‭OpenStax‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ he ductless endocrine glands should not be confused with the body’s‬‭exocrine system‬‭,‬
T
‭whose glands release their secretions through ducts. Examples of exocrine glands include‬
‭the sebaceous and sweat glands of the skin and the salivary glands. As previously noted,‬
‭the pancreas also has an exocrine function: most of its cells secrete pancreatic juice‬
‭through the pancreatic and accessory ducts to the lumen of the small intestine.‬

‭Types of Hormones‬
‭ here are four basic classes of hormones: amino acid-derived, peptide-derived,‬
T
‭protein-derived, and lipid-derived (‬‭Figure 11.4‬‭).‬‭Amino acid-derived (Amine)‬
‭hormones‬‭are relatively small molecules that consist of a single, modified amino acid (e.g.,‬
‭tryptophan or tyrosine) and include the adrenal gland hormones epinephrine and‬
‭norepinephrine.‬ ‭Peptide-derived hormones‬‭consist‬‭o f relatively short polypeptide‬
‭chains, including pituitary hormones, antidiuretic hormone, and oxytocin.‬‭Protein-derived‬
‭hormones‬‭consist of relatively longer polypeptide‬‭chains and include growth and‬
‭follicle-stimulating hormones.‬ ‭Lipid-derived hormones‬‭are structurally similar to‬
‭cholesterol and include steroid hormones such as estrogen, progesterone, testosterone,‬
‭and cortisol.‬
‭ ll amine, peptide, and protein hormones are water-soluble (‬‭hydrophilic‬‭), whereas‬
A
‭steroid hormones are‬‭hydrophobic‬‭. Because blood is‬‭water-based, steroid hormones‬
‭must travel to their target cell bound to transport proteins. This more complex structure‬
‭extends the half-life of steroid hormones much longer than those derived from amino‬
‭acids (i.e., the time required for half the hormone concentration to be degraded.). For‬
‭example, the lipid-derived hormone cortisol has a half-life of approximately 60 to 90‬
‭minutes. In contrast, the amino acid–derived hormone epinephrine has a half-life of about‬
‭o ne minute.‬

‭How Hormones Work‬

‭ ven though hormones released by endocrine glands can travel long distances through‬
E
‭the blood and come into contact with many different cell types, they only affect target cells‬
‭with the necessary receptors. These receptors can be located in the cytoplasm‬
‭(‬‭c ytoplasmic receptors‬‭) or the plasma membrane (‬‭membrane-bound‬‭receptors‬‭).‬
‭Lipid-derived steroid hormones can enter the cell by diffusing across the plasma‬
‭membrane and binding to receptors in the cytoplasm or nucleoplasm, which interact with‬
‭the nuclear DNA to regulate gene transcription (‬‭Figure‬‭11.5‬‭).‬
‭ ipid-insoluble hormones (amine, peptide, and protein) bind to receptors on the plasma‬
L
‭membrane surface of target cells and trigger a signaling pathway to change the cell’s‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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a‭ ctivities. The hormone is called the‬‭first messenger,‬‭and the cellular component‬

‭associated with the signaling pathway is called the‬‭second messenger‬‭. The second‬
‭messenger directly triggers the intracellular activity and carries out the specific effects‬
‭associated with the hormone, such as the breakdown of glycogen into glucose monomers.‬

‭ igure 11.4‬ ‭The four classes of hormone types.‬ ‭(credit:‬‭OpenStax‬‭). A link to a video‬
F
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭:‬

‭ igure 11.5‬ ‭A steroid hormone directly initiates‬‭the production of proteins within a‬

F
‭target cell. The hormone binds to its receptor in the cytosol, forming a receptor-hormone‬
‭complex. The receptor-hormone complex attaches to the target gene on DNA. Transcribing‬
‭the gene creates a messenger RNA that is translated into the desired protein within the‬
‭cytoplasm. (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ he second messenger most hormones use is‬‭c yclic adenosine‬‭monophosphate‬

T
‭(cAMP)‬‭. In the cAMP second messenger system, a water-soluble‬‭hormone binds to its‬
‭receptor in the cell membrane (Step 1 in‬‭Figure 11.6‬‭).‬‭This receptor is associated with an‬
‭intracellular component called a G protein, and binding of the hormone activates the‬
‭G-protein component (Step 2). The activated G protein, in turn, activates an enzyme called‬
‭adenylyl cyclase (Step 3), which converts adenosine triphosphate (ATP) to cyclic AMP or‬
‭cAMP (Step 4). As the second messenger, cAMP activates a type of enzyme called a protein‬
‭kinase in the cytoplasm (Step 5). Activated protein kinases initiate a reaction in which a‬
‭phosphate group is added to numerous cellular proteins, including other enzymes (Step‬
‭6).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ xamples of hormones that use cAMP as a second messenger include luteinizing hormone,‬
E
‭glucagon, which plays a role in regulating blood glucose levels, and thyroid-stimulating‬
‭hormone, which causes the release of T‬‭3‬ ‭and T‬‭4‬ ‭from‬‭the thyroid gland.‬

‭ igure 11.6‬ ‭Water-soluble hormones cannot diffuse‬‭through the cell membrane. These‬
F
‭hormones must bind to a surface cell-membrane receptor. The receptor then initiates a‬
‭cell-signaling pathway within the cell involving G proteins, adenylyl cyclase, the secondary‬
‭messenger cyclic AMP (cAMP), and protein kinases. In the final step, these protein kinases‬
‭phosphorylate proteins in the cytoplasm, adding a phosphate group to them. This action‬
‭activates proteins in the cell that carry out the changes specified by the hormone. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭Regulating Hormone Secretion‬

‭ ormone levels must be tightly controlled to maintain homeostasis. The body maintains‬
H
‭this control by balancing hormone production and degradation.‬‭Negative‬‭feedback loops‬
‭regulate the synthesis and secretion of most hormones in response to various stimuli.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ his type of feedback loop is characterized by inhibiting the further secretion of a‬

T
‭hormone in response to adequate levels of that hormone. This allows blood hormone‬
‭levels to be regulated within a narrow range. An example of a negative feedback loop is‬
‭the release of the thyroid hormones T‬‭3‬ ‭and T‬‭4‬‭, which‬‭are directed by the hypothalamus‬
‭and pituitary gland (‬‭Figure 11.7‬‭). As the concentrations‬‭o f the thyroid hormones rise,‬
‭they signal the hypothalamus and anterior pituitary gland to reduce the secretion of TRH‬
‭and TSH, respectively, which prevents additional T‬‭3‬ ‭and T‬‭4‬ ‭secretion.‬

‭ igure 11.7‬ ‭An example of a negative‬

F
‭feedback loop using the thyroid system.‬
‭The hypothalamus secretes TRH, which‬
‭causes the anterior pituitary gland to‬
‭secrete TSH, which causes the thyroid‬
‭gland to secrete T‬‭3‬ ‭and T‬‭4‬‭. These two‬
‭hormones “feedback” to the anterior‬
‭pituitary gland and hypothalamus to‬
‭diminish the secretion of their respective‬
‭hormones (TSH and TRH). (credit:‬
‭OpenStax‬‭). A link to a video explanation‬
‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭11.4 The Hypothalamus and Pituitary Gland‬

‭ he hypothalamus–pituitary complex (‬‭Figure 11.8‬‭) is‬‭the “command center” of the‬
T
‭endocrine system. The‬‭pituitary gland‬‭is located at‬‭the base of the brain and is attached‬
‭to another region called the‬‭hypothalamus‬‭via a structure‬‭called the infundibulum. The‬
‭anterior pituitary gland‬‭receives products called‬‭releasing hormones‬‭from the‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ ypothalamus. These releasing hormones then stimulate the secretion of six hormones,‬
h
‭which are discussed later in this chapter.‬
‭ he‬‭posterior pituitary gland‬‭is an extension of the‬‭brain that releases two hormones:‬
T
‭antidiuretic hormone (ADH)‬‭and‬‭oxytocin‬‭(‬‭OT‬‭), which‬‭are produced by the‬
‭hypothalamus. Again, the posterior pituitary does not produce hormones but instead‬
‭stores hormones produced by the hypothalamus (ADH and OT). From this location, these‬
‭hormones are released into the bloodstream.‬

‭ igure 11.8‬ ‭The hypothalamus region lies below‬‭the thalamus and connects to the‬
F
‭pituitary gland by the stalk-like region called the infundibulum. The pituitary gland‬
‭consists of an anterior and posterior lobe, with each lobe secreting different hormones in‬
‭response to signals from the hypothalamus (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ he anterior pituitary produces a variety of hormones, including the following: growth‬

T
‭hormone (GH), thyroid-stimulating hormone (TSH), adrenocorticotropic hormone (ACTH),‬
‭prolactin (PRL), follicle-stimulating hormone (FSH), and luteinizing hormone (LH).‬
‭ rowth hormone (GH)‬‭controls muscle and bone growth‬‭rates by influencing protein‬
G
‭synthesis (Chapter 4). It also influences cellular metabolic functions. Athletes or other‬
‭individuals can use human GH to increase muscle mass, which will be discussed later as‬
‭part of a discussion of performance-enhancing drugs (PEDs).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ hyroid-stimulating hormone‬‭(‬‭TSH‬‭) controls the release‬‭o f two hormones from the‬

T
‭thyroid gland (triiodothyronine (‬‭T‬‭3‭)‬ and thyroxine‬‭(‬‭T‬‭4‬‭)), which control the body’s‬
‭metabolic rate required to perform essential functions.‬‭Iodine‬‭is needed to produce both‬
‭hormones; lacking this element in the diet can lead to a condition called goiter. This‬
‭condition and the use of iodized salt to meet this dietary need are discussed later in this‬
‭chapter.‬
‭ drenocorticotropic hormone (ACTH)‬‭causes the release‬‭o f several types of substances,‬
A
‭including a‬‭mineralocorticoid‬‭hormone called‬‭aldosterone‬‭and the three main‬
‭glucocorticoid hormones‬‭: cortisol, corticosterone,‬‭and cortisone. Aldosterone promotes‬
‭the reabsorption of sodium ions (Na‬‭+1‬‭) and excretion‬‭o f potassium ions (K‬‭+1‬‭) to maintain‬
‭homeostatic concentrations. The three glucocorticoids maintain glucose levels within a‬
‭normal range between meals.‬
‭ rolactin (PRL)‬‭is produced by both sexes but is known‬‭primarily for its role in females,‬
P
‭as it stimulates the development of mammary glands in breasts and milk production after‬
‭childbirth. There is uncertainty about the specific functions of PRL in males. With respect‬
‭to the reproductive system, it may function to optimize testosterone production and,‬
‭therefore, sperm production.‬
‭ ollicle Stimulating Hormone (FSH)‬‭and‬‭Luteinizing‬‭Hormone (LH)‬‭are secreted in‬
F
‭response to a particular releasing hormone from the hypothalamus, called‬
‭gonadotropin-releasing hormone (GnRH‬‭), that is secreted‬‭at the onset of puberty. FSH‬
‭and LH both target the gonads, the ovaries in females, and the testes in males. While both‬
‭sexes secrete FSH and LH, their actions differ depending on sex. FSH stimulates the‬
‭production and maturation of‬‭gametes‬‭, or sex cells,‬‭which are‬‭ova‬‭(eggs) in females and‬
‭sperm‬‭in males. In females, FSH also promotes the‬‭growth and development of‬‭follicles‬‭,‬
‭which are structures in the ovaries containing ova. Follicles release‬‭estrogen‬‭and‬
‭progesterone‬‭both before and after the release of‬‭ova. In females, LH triggers ovulation,‬
‭which is when a follicle ruptures and releases the ovum, as well as the production of‬
‭estrogen and progesterone. In males, LH stimulates the production of‬‭testosterone‬‭by the‬
‭testes.‬
‭ s stated previously, ADH and OT are secreted from the posterior pituitary gland. ADH‬
A
‭functions to control water levels in the body by triggering the kidneys' reabsorption of‬
‭water. In females, OT stimulates uterine contraction during childbirth and milk let-down‬
‭during nursing. In males, it stimulates vas deferens and prostate gland contractions during‬
‭ejaculation.‬
‭ summary of information about hormones released by the posterior and anterior‬
A
‭pituitary glands is presented in‬‭Figure 11.9‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 11.9‬ ‭Summary of posterior and anterior pituitary‬‭gland hormone information‬

F
‭(credit:‬‭Major Pituitary Hormones.jpg‬‭; OpenStax College;‬‭CC BY 3.0‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭11.5 The Thyroid Gland, Adrenal Glands, Pancreas, and Gonads‬

‭Thyroid Gland‬
‭ he‬‭thyroid gland‬‭is located in the neck, just below‬‭the larynx, and in front of the trachea,‬
T
‭as shown in‬‭Figure 11.10‬‭. It is a butterfly-shaped‬‭gland with two lobes. It has a dark red‬
‭color due to its extensive vascular system. When the thyroid swells due to dysfunction, it‬
‭can be felt under the skin of the neck.‬

‭Figure 11.10‬ ‭This illustration shows the location‬‭o f the thyroid gland. (credit:‬‭OpenStax‬‭)‬

‭ s stated earlier, thyroid gland cells synthesize the hormones‬‭T‬‭3‬ ‭and‬‭T‬‭4‬ ‭(the number‬
A
‭indicates the atoms of iodine present), which increase the rates of mitochondrial ATP‬
‭production via increased metabolic rate and cellular respiration.‬
‭ or additional information about thyroid hormone’s role in regulating metabolism, click‬
F
‭the link below to watch the TED-Ed video by Emma Bryce titled “How does the thyroid‬
‭manage your metabolism?”.‬

How does the thyroid manage your metabolism? - Emma Bryce

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Adrenal Glands‬
‭ he‬‭adrenal glands‬‭are associated with the kidneys, and one gland is on top of each‬
T
‭kidney (‬‭Figure 11.11)‬‭. The adrenal glands consist‬‭o f an outer adrenal cortex and an inner‬
‭adrenal medulla, which secrete different hormones.‬

‭ igure 11.11‬ ‭The location of the adrenal glands on‬‭top of the kidneys is shown. (credit:‬
F
‭modification of work by NCI;‬‭OpenStax‬‭)‬

‭Adrenal Cortex‬
‭ he‬‭adrenal cortex‬‭produces mineralocorticoids, glucocorticoids,‬‭and small amounts of‬
T
‭sex hormones (androgens;‬‭Figure 11.8‬‭). The principal‬‭mineralocorticoid is‬‭aldosterone,‬
‭which regulates the concentration of Na‬‭+‬ ‭and K‬‭+‬ ‭ions‬‭in urine, sweat, pancreas, and saliva.‬
‭Decreased sodium ion concentrations, blood volume, or blood pressure stimulate‬
‭aldosterone release.‬
‭ he three main glucocorticoids are cortisol, corticosterone, and cortisone. The‬
T
‭glucocorticoids stimulate the synthesis of glucose and convert non-carbohydrates, such as‬
‭fats, to glucose by liver cells. They also promote the release of fatty acids from adipose‬
‭tissue. These hormones increase blood glucose levels to maintain them within a normal‬
‭range between meals. They are secreted in response to ACTH, and negative feedback‬
‭loops regulate their levels.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ ndrogens are sex hormones produced in small amounts by the adrenal cortex in both‬
A
‭males and females. They do not affect sexual characteristics and may supplement sex‬
‭hormones released from the gonads (testes and ovaries).‬

‭ igure 11.12‬ ‭Shows the adrenal cortex and medulla‬‭regions diagrammatically and‬
F
‭histologically. Note that the adrenal cortex is subdivided into three areas based on‬
‭different tissue types, but it isn’t necessary to learn the associated names. ANS is an‬
‭abbreviation for the autonomic nervous system. (credit:‬‭The Adrenal Glands.jpg‬‭;‬
‭OpenStax College;‬‭CC BY 3.0‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭Adrenal Medulla‬
‭ he‬‭adrenal medulla‬‭contains two types of secretory‬‭cells (‬‭Figure 11.8‬‭): one that‬
T
‭produces epinephrine (adrenaline) and another that makes norepinephrine‬
‭(noradrenaline). Epinephrine is the primary adrenal medulla hormone, accounting for 75‬
‭to 80 percent of its secretions. Epinephrine and norepinephrine are associated with the‬
‭“flight or fight” response and increase heart rate, breathing rate, cardiac muscle‬
‭contractions, blood pressure, and blood glucose levels. They also accelerate the‬
‭breakdown of glucose in skeletal muscles and stored fats in adipose tissue in anticipation‬
‭o f energy demands on the body.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Pancreas‬
‭ he‬‭pancreas‬‭, illustrated in‬‭Figure 11.13‬‭, is an elongated‬‭o rgan between the stomach and‬
T
‭the first part of the small intestine (duodenum). It contains both exocrine cells that release‬
‭digestive enzymes (Chapter 5) and endocrine cells that release hormones.‬

‭ igure 11.13‬ ‭The pancreas is adjacent to the duodenum‬‭and under the stomach. Some of‬
F
‭its cells (acinar cells) produce digestive enzymes, while others (islet cells) produce insulin‬
‭and glucagon. (credit:‬‭Exocrine and Endocrine Pancreas.jpg‬‭;‬‭OpenStax College;‬‭CC BY‬
‭3.0‬‭).‬‭A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ he endocrine cells of the pancreas form clusters called‬‭pancreatic islets‬‭, which contain‬
T
‭cells that produce the two hormones‬‭insulin‬‭and‬‭glucagon‬‭,‬‭which regulate blood glucose‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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l‭evels to maintain homeostasis (‬‭Figure 11.14‬‭). As blood glucose levels decline, glucagon is‬
‭released to raise the blood glucose levels by increasing rates of glycogen breakdown and‬
‭glucose release by the liver. When blood glucose levels rise, such as after a meal, insulin is‬
‭released to lower blood glucose levels by increasing the rate of glucose uptake in most‬
‭body cells and by increasing glycogen synthesis in skeletal muscles and the liver.‬

‭ igure 11.14‬ ‭Insulin and glucagon regulate blood‬‭glucose levels. (credit:‬‭OpenStax‬‭). A‬

F
‭link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Gonads‬
‭ he‬‭gonads‬‭—the male testes and female ovaries—produce‬‭steroid hormones. The‬‭testes‬
T
‭produce androgens, testosterone being the most prominent, which allow for the‬
‭development of secondary sex characteristics and the production of sperm cells. The‬
‭ovaries‬‭produce estrogen and progesterone, which cause‬‭secondary sex characteristics‬
‭and prepare the body for childbirth. These hormones are discussed in the context of their‬
‭reproductive system functions in Chapter 14.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭11.6 Selected Disorders of the Endocrine System‬

‭Iodine Deficiency, Hypothyroidism, and Hyperthyroidism‬

‭ s discussed previously, dietary iodine is required to synthesize T‬‭3‬ ‭and T‬‭4‭.‬ However, for‬
A
‭much of the world’s population, foods do not provide adequate levels of this mineral‬
‭because the amount varies according to the level in the soil in which the food was grown,‬
‭as well as the irrigation and fertilizers used. Marine fish and shrimp tend to have high‬
‭levels because they concentrate iodine from seawater, but many people in landlocked‬
‭regions lack access to seafood. Thus, iodized salt is the primary source of dietary iodine in‬
‭many countries. Fortification of salt with iodine began in the United States in 1924, and‬
‭international efforts to iodize salt in the world’s poorest nations continue today.‬

‭ ietary iodine deficiency can impair the ability to synthesize T‬‭3‬ ‭and T‬‭4‭,‬ leading to various‬
D
‭severe disorders, generally classified as‬‭hypothyroidism‬‭.‬‭TSH is secreted increasingly‬
‭when T‬‭3‬ ‭and T‬‭4‬ ‭cannot be produced. As a result of‬‭this hyperstimulation, the overall size of‬
‭the thyroid gland increases, and a condition called‬‭goiter‬‭results (‬‭Figure 11.15‬‭). A goiter‬
‭is only a visible indication of the deficiency. Other iodine deficiency disorders include‬
‭impaired growth and development, decreased fertility, and prenatal and infant death.‬
‭Moreover, iodine deficiency is the primary cause of preventable mental retardation‬
‭worldwide. Neonatal hypothyroidism (cretinism) is characterized by cognitive deficits,‬
‭short stature, and sometimes deafness and muteness in children and adults born to‬
‭iodine-deficient mothers during pregnancy.‬

‭Figure 11.15‬ ‭Goiter (credit: “Almazi”; Wikimedia‬‭Commons;‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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I‭ n areas of the world with access to iodized salt, dietary deficiency is rare. Instead, thyroid‬
‭gland inflammation is the more common cause of low blood levels of thyroid hormones.‬

I‭ n contrast,‬‭hyperthyroidism‬‭—an abnormally elevated‬‭blood level of thyroid‬

‭hormones—is often caused by a pituitary or thyroid tumor. In‬‭Graves’ disease‬‭, the‬
‭hyperthyroid state results from an autoimmune reaction in which antibodies‬
‭overstimulate the follicle cells of the thyroid gland. Hyperthyroidism can lead to an‬
‭increased metabolic rate, excessive body heat and sweating, diarrhea, weight loss, tremors,‬
‭and increased heart rate. The person’s eyes may bulge (called exophthalmos) as‬
‭antibodies produce inflammation in the soft tissues of the orbits. The person may also‬
‭develop a goiter.‬

‭Diabetes Mellitus‬

‭ ysfunction of insulin production and secretion, as well as the target cells’ responsiveness‬
D
‭to insulin, can lead to a condition called‬‭diabetes‬‭mellitus‬‭. An increasingly common‬
‭disease, diabetes mellitus has been diagnosed in more than 18 million adults in the United‬
‭States and more than 200,000 children. It is estimated that up to 7 million more adults‬
‭have the condition but have not been diagnosed. In addition, approximately 79 million‬
‭people in the US are estimated to have pre-diabetes, a condition in which blood glucose‬
‭levels are abnormally high but not yet high enough to be classified as diabetes.‬

‭ here are two primary forms of diabetes mellitus.‬‭Type 1 (juvenile) diabetes‬‭is an‬
T
‭autoimmune disease affecting the beta cells of the pancreas. Specific genes are recognized‬
‭to increase susceptibility. The beta cells of people with type 1 diabetes do not produce‬
‭insulin; thus, synthetic insulin must be administered by injection or infusion. This form of‬
‭diabetes accounts for less than five percent of all diabetes cases.‬

‭ ype 2 (adult onset) diabetes‬‭accounts for approximately‬‭95 percent of all cases. It is‬
T
‭acquired, and lifestyle factors such as poor diet, inactivity, and pre-diabetes significantly‬
‭increase a person’s risk. About 80 to 90 percent of people with type 2 diabetes are‬
‭overweight or obese. In type 2 diabetes, cells become resistant to the effects of insulin. In‬
‭response, the pancreas increases insulin secretion, but the beta cells become exhausted‬
‭over time. In many cases, type 2 diabetes can be reversed by moderate weight loss,‬
‭regular physical activity, and consumption of a healthy diet; however, if blood glucose‬
‭levels cannot be controlled, affected individuals will eventually require insulin.‬

‭ wo of the early manifestations of diabetes are excessive urination and excessive thirst.‬
T
‭The kidneys are responsible for filtering glucose from the blood. Excessive blood glucose‬
‭draws water into the urine, and as a result, the person eliminates an abnormally large‬
‭quantity of urine with substantial glucose. Using body water to dilute the urine leaves the‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ ody dehydrated, so the person is unusually and continually thirsty. The person may also‬
b
‭experience persistent hunger because the body cells cannot access the glucose in the‬
‭bloodstream.‬

‭ ver time, persistently high glucose levels in the blood injure tissues throughout the body,‬
O
‭especially those of the blood vessels and nerves. Inflammation and injury of the lining of‬
‭arteries lead to atherosclerosis and an increased risk of heart attack and stroke. Damage‬
‭to the microscopic blood vessels of the kidney impairs kidney function and can lead to‬
‭kidney failure. Damage to blood vessels that serve the eyes can lead to blindness. Blood‬
‭vessel damage also reduces circulation to the limbs, whereas nerve damage leads to a loss‬
‭o f sensation, called neuropathy, particularly in the hands and feet. Together, these changes‬
‭increase the risk of injury, infection, and tissue death (necrosis), contributing to a high rate‬
‭o f toe, foot, and lower leg amputations in people with diabetes.‬

‭ ncontrolled diabetes can also lead to a dangerous form of metabolic acidosis called‬
U
‭ketoacidosis. If cells are deprived of glucose, they increasingly rely on fat stores for fuel.‬
‭However, in a glucose-deficient state, the liver is forced to use an alternative lipid‬
‭metabolism pathway that increases the production of ketone bodies (or ketones), which‬
‭are acidic. The build-up of ketones in the blood causes ketoacidosis, which—if left‬
‭untreated—may lead to a life-threatening “diabetic coma.” Together, these complications‬
‭make diabetes the seventh leading cause of death in the United States.‬

‭ iabetes is diagnosed when lab tests reveal that blood glucose levels are higher than‬
D
‭normal, a condition called hyperglycemia. Diabetes treatment depends on the type, the‬
‭severity of the condition, and the ability of the patient to make lifestyle changes. As noted‬
‭earlier, moderate weight loss, regular physical activity, and a healthy diet can reduce blood‬
‭glucose levels. Some patients with type 2 diabetes may be unable to control their disease‬
‭with these lifestyle changes and will require medication. Historically, the first-line‬
‭treatment of type 2 diabetes was insulin. Research advances have resulted in alternative‬
‭o ptions, including medicines that enhance pancreatic function.‬

‭ or additional information about diabetes and related topics, click the link below to watch‬
F
‭the TED-Ed video by Duncan C. Ferguson titled “What did dogs teach humans about‬
‭diabetes?”.‬

What did dogs teach humans about diabetes? - Duncan C. Ferguson

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭11.7 Hormones and Athletic Performance‬

‭ he endocrine system can be exploited for illegal or unethical purposes. A prominent‬

T
‭example of this is the use of drugs by professional athletes.‬

S‭ ome athletes attempt to boost their performance by using artificial hormones that‬
‭enhance muscle performance.‬‭Anabolic steroids‬‭, a form‬‭o f the male sex hormone‬
‭testosterone, are one of the most widely known performance-enhancing drugs (PEDs).‬
‭Steroids help build muscle mass. Other hormones used to enhance athletic performance‬
‭include erythropoietin, which triggers the production of red blood cells, and human‬
‭growth hormone, which can help build muscle mass.‬

‭ he use of performance-enhancing drugs is banned by all major collegiate and‬

T
‭professional sports organizations in the United States because they impart an unfair‬
‭advantage to athletes who take them. In addition, the drugs can cause significant and‬
‭dangerous side effects. For example, anabolic steroid use can increase cholesterol levels,‬
‭raise blood pressure, and damage the liver. Altered testosterone levels (both too low or‬
‭too high) have been implicated in causing structural damage to the heart and increasing‬
‭the risk for cardiac arrhythmias, heart attacks, congestive heart failure, and sudden death.‬
‭Paradoxically, steroids can have a feminizing effect in males, including shriveled testicles‬
‭and enlarged breast tissue. In females, their use can cause masculinizing effects such as an‬
‭enlarged clitoris and facial hair growth. Both sexes' use can promote increased aggression‬
‭(commonly known as “roid-rage”), depression, sleep disturbances, severe acne, and‬
‭infertility.‬

‭11.8 Career Connection - Endocrinologist‬

‭ ndocrinology is a specialty in the field of medicine that focuses on the treatment of‬
E
‭endocrine system disorders. Endocrinologists—medical doctors specializing in this‬
‭field—are experts in treating diseases associated with hormonal systems, ranging from‬
‭thyroid disease to diabetes mellitus. Endocrine surgeons treat endocrine disease through‬
‭the removal, or resection, of the affected endocrine gland.‬

‭ atients referred to endocrinologists may have signs and symptoms or blood test results‬
P
‭that suggest excessive or impaired functioning of an endocrine gland or endocrine cells.‬
‭The endocrinologist may order additional blood tests to determine whether the patient’s‬
‭hormonal levels are abnormal or if they may stimulate or suppress the function of the‬
‭suspect endocrine gland. Then, the patient may have blood taken for analysis. Treatment‬
‭varies according to the diagnosis. Some endocrine disorders, such as type 2 diabetes, may‬
‭respond to lifestyle changes such as modest weight loss, adoption of a healthy diet, and‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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r‭ egular physical activity. Other disorders may require medication, such as hormone‬
‭replacement, and routine monitoring by the endocrinologist. These include disorders of‬
‭the pituitary gland that can affect growth and disorders of the thyroid gland that can‬
‭result in various metabolic problems.‬

S‭ ome patients experience health problems due to the expected hormone decline that can‬
‭accompany aging. These patients can consult with an endocrinologist to weigh the risks‬
‭and benefits of hormone replacement therapy intended to boost their natural levels of‬
‭reproductive hormones.‬

I‭ n addition to treating patients, endocrinologists may be involved in research to improve‬

‭the understanding of endocrine system disorders and develop new treatments for these‬
‭diseases.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 12: Urinary System‬

‭ igure 12.1‬‭A patient is undergoing dialysis due to‬‭impaired kidney function. Blood travels‬
F
‭through a tube immersed in a solution, with the tube walls being semipermeable‬
‭membranes. The solution is made to remove urea from the blood through diffusion. The‬
‭semi-permeable tube wall allows urea through but keeps the larger blood components,‬
‭such as proteins and blood cells, within the tube. The “cleaned” blood is eventually‬
‭returned to the body. (credit:‬‭Patient receiving‬‭dialysis 03.jpg‬‭;‬‭Anna Frodesiak‬‭;‬‭CC0 1.0‬‭)‬

‭12.1 Introduction‬
‭ he urinary system has roles you are likely aware of, such as cleansing the blood and‬
T
‭ridding the body of wastes. However, this system also has other equally important‬
‭functions, such as pH homeostasis, blood pressure regulation, and endocrine functions.‬
‭The kidneys also perform the final synthesis step of vitamin D production, which helps to‬
‭absorb and maintain calcium and phosphorus.‬
I‭ f the kidneys fail, these functions are compromised or lost altogether, and homeostasis is‬
‭devastatingly affected. The impacted individual might experience weakness, lethargy,‬
‭shortness of breath, anemia, widespread edema (swelling), heart arrhythmias, and more.‬
‭Each of these functions is vital to your well-being and survival. The urinary system,‬
‭controlled by the nervous system, also stores urine until a convenient time for disposal‬
‭and then provides the anatomical structures to transport this waste liquid to the outside‬

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‭ f the body. Failure of nervous control or the anatomical structures leading to a loss of‬
o
‭control of urination results in a condition called incontinence.‬
‭ his chapter will help you to understand the anatomy of the urinary system and how it‬
T
‭enables the physiologic functions critical to homeostasis. It is best to think of the kidney as‬
‭a regulator of plasma makeup rather than simply a urine producer. As you read each‬
‭section, ask yourself: “What happens if this does not work?” This question will help you to‬
‭understand how the urinary system maintains homeostasis and affects all the body's‬
‭o ther systems and the quality of one’s life.‬
‭ efore reading the chapter, it will be helpful to gain a general overview of the urinary‬
B
‭system. Click the link or scan the QR code to watch the TED-Ed video by Emma Bryce‬
‭titled “How do your kidneys work?”‬

How do your kidneys work? - Emma Bryce

‭12.2 Physical Characteristics of Urine‬

‭ he urinary system’s ability to filter the blood resides in the approximately 2 to 3 million‬
T
‭tufts of specialized capillaries, called‬‭glomeruli‬‭,‬‭distributed more or less equally between‬
‭the two kidneys. Because the glomeruli filter the blood, based chiefly on particle size,‬
‭larger elements (e.g., blood cells, platelets, antibodies, and albumen) can’t pass through‬
‭these capillaries into structures called‬‭nephrons‬‭,‬‭which are the functional unit of the‬
‭kidney (i.e., they produce urine). All other solutes, such as ions, amino acids, vitamins, and‬
‭wastes, are filtered to create a fluid, called filtrated, that has a composition similar to‬
‭plasma. The glomeruli produce about 200 liters (approximately 52 gallons) of this filtrate‬
‭daily, yet you excrete less than two liters, or half a gallon, of waste, called‬‭urine‬‭. This‬
‭means that the body reabsorbs most filtrate.‬
‭ rine composition is dynamic or changing, depending on water intake, exercise,‬
U
‭environmental temperature, nutrient intake, and other factors (‬‭Figure 12.2‬‭). Some‬
‭characteristics, such as color and odor, are rough descriptors of your state of hydration.‬
‭For example, if you exercise or work outside and sweat a lot, your urine will turn darker‬
‭and produce a slight odor, even if you drink plenty of water. Athletes are often advised to‬
‭consume water until their urine is clear, which is good advice. However, it takes time for‬
‭the kidneys to process body fluids and store urine in the bladder. Another way of looking‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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a‭ t this is that the quality of the urine produced is the average over the time it takes to‬
‭make that urine. Producing clear urine may take only a few minutes if you drink a lot of‬
‭water or several hours if you are working outside and not drinking much.‬
‭ rinalysis‬‭(urine analysis) often provides clues to renal disease. Typically, only traces of‬
U
‭protein are found in urine; when higher amounts are found, damage to the glomeruli is‬
‭the likely cause.‬
‭ he color of urine is determined primarily by the breakdown products of red blood cell‬
T
‭destruction. The liver converts the “heme” of hemoglobin into water-soluble forms that‬
‭can be excreted into the bile and indirectly into the urine. Certain foods like beets, berries,‬
‭and fava beans may also affect urine color. A kidney stone or cancer of the urinary system‬
‭may produce sufficient bleeding to manifest as pink or even bright red urine.‬

‭Characteristic‬ ‭ ormal values‬

N
‭Color‬ ‭ ale yellow to deep amber‬
P
‭Odor‬ ‭Odorless‬
‭Volume‬ ‭750–2000 mL/24 hour‬
‭pH‬ ‭4.5–8.0‬
‭Protein‬ ‭None or trace‬
‭Ketones‬ ‭None‬
‭Blood‬ ‭None‬
‭Glucose‬ ‭None‬

‭Figure 12.2‬ ‭Selected urine characteristics and the‬‭associated normal range of values.‬

‭ iseases of the liver or obstructions of bile drainage from the liver impart a dark “tea” or‬
D
‭“cola” hue to the urine. Dehydration produces darker, concentrated urine with a slight‬
‭ammonia odor. Most of the‬‭ammonia‬‭produced from protein‬‭breakdown is converted‬
‭into‬‭urea‬‭by the liver, so ammonia is rarely detected‬‭in fresh urine. The strong ammonia‬
‭o dor you may notice in bathrooms or alleys is due to the breakdown of urea into‬
‭ammonia by bacteria in the environment. About one in five people detect a distinctive‬
‭o dor in their urine after consuming asparagus; other foods, such as onions, garlic, and‬
‭fish, can impart their harmless aromas.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ rine volume varies considerably, but the normal range is one to two quarts per day. The‬
U
‭kidneys must produce a minimum urine volume of about 0.5 quarts per day to rid the‬
‭body of wastes. The output below this level may result from severe dehydration or renal‬
‭disease. Excessive urine production may be due to diabetes mellitus or diabetes insipidus.‬
‭In‬‭diabetes mellitus‬‭, blood glucose levels exceed‬‭the number of available sodium-glucose‬
‭transporters in the kidney, and glucose appears in the urine. The osmotic nature of‬
‭glucose attracts water, leading to its loss in the urine. In the case of‬‭diabetes insipidus‬‭,‬
‭insufficient antidiuretic hormone (ADH) release or inadequate numbers of ADH receptors,‬
‭meaning that too few water channels are inserted into the cell membranes that line tubes‬
‭in the kidney that transport urine from the nephrons ultimately to the ureters. Insufficient‬
‭numbers of water channels reduce water reabsorption, which results in high volumes of‬
‭very dilute urine.‬

‭Figure 12.3‬ ‭Urine color and its association with‬‭hydration levels. (credit:‬‭OpenStax‬‭)‬

‭ he urine's pH (hydrogen ion concentration) can vary more than 1000-fold, from a low of‬
T
‭4.5 to a maximum of 8.0. Diet can influence pH; meats lower the pH, whereas citrus fruits,‬
‭vegetables, and dairy products raise the pH. Chronically high or low pH can lead to‬
‭disorders, such as the development of kidney stones. For more information about this‬
‭topic, click the link or scan the QR code to watch the TED-Ed video by Arash Shadman‬
‭titled “What causes kidney stones?”.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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What causes kidney stones? - Arash Shadman

‭ etones‬‭are byproducts of fat metabolism. Finding‬‭ketones in the urine suggests that the‬
K
‭body uses fat as an energy source instead of glucose. In diabetes mellitus, when there is‬
‭not enough insulin (type I diabetes mellitus) or because of insulin resistance (type II‬
‭diabetes mellitus), there is plenty of glucose. Still, without the action of insulin, the cells‬
‭cannot take it up, so it remains in the bloodstream. This outcome forces cells to use fat as‬
‭their energy source, which produces excess ketones as byproducts. These excess ketones‬
‭will appear in the urine. Ketones may also be present if the diet has a severe deficiency of‬
‭proteins or carbohydrates.‬
‭ o blood should be present in the urine. Although blood may sometimes appear in urine‬
N
‭samples due to menstrual contamination, this is abnormal. Now that you understand the‬
‭typical characteristics of urine, the next section will introduce you to the various organs of‬
‭the urinary system and their respective functions.‬

‭12.3 Gross Anatomy of the Urinary System‬

‭ he urinary system organs include the kidneys, ureters, bladder, sphincters, and urethra‬
T
‭(‬‭Figure 12.4‬‭). Let’s look at each of these organs‬‭in more detail.‬

‭Kidneys‬
‭ he‬‭kidneys‬‭lie on either side of the spine and are‬‭well protected by muscle, fat, and ribs.‬
T
‭They are roughly the size of your fist, and the male kidney is slightly larger than the female‬
‭kidney. The kidneys are well vascularized, receiving about 25 percent of the oxygenated‬
‭blood pumped by the heart to the systemic circulation. The kidneys receive freshly‬
‭oxygenated (red) blood from the renal arteries, filter it to produce urine, and return the‬
‭blue blood to the inferior vena cava via the renal veins.‬
‭ n top of each kidney is an‬‭adrenal gland‬‭, and its‬‭cortex layer produces and secretes‬
O
‭aldosterone. This hormone, as discussed in Chapter 11, plays essential roles in sodium‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭reabsorption, water reabsorption, and urine concentration.‬

‭Ureters‬
‭ he‬‭ureters‬‭are approximately 11-inch long tubes that‬‭exit the kidney and empty urine‬
T
‭into the‬‭urinary bladder. As urine passes through‬‭the ureter, it does not passively drain‬
‭into the bladder but is propelled by waves of smooth muscle contractions called‬
‭peristalsis‬‭(‬‭Figure 12.5‬‭). The attachment of the ureters‬‭to the bladder creates a one-way‬
‭valve that allows urine into the bladder but prevents its movement back into the ureter.‬

‭ igure 12.4‬ ‭The gross anatomy of the urinary system.‬‭The kidneys filter blood received‬
F
‭from the renal arteries to produce urine that is transported to the bladder via the ureters,‬
‭stored, and then eliminated through the urethra. The renal veins return the filtered blood‬
‭to general circulation. (credit:‬‭Urinary System Organs‬‭;‬‭Cenveo;‬‭CC BY 3.0‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 12.5‬ ‭A magnified view of the histology of‬‭the ureter. The lumen of the ureter is‬
F
‭the internal opening through which urine will move from the kidneys to the bladder.‬
‭Smooth muscle contractions help transport urine through the lumen. LM × 128. (credit:‬
‭Micrograph provided by the Regents of the University of Michigan Medical School © 2012;‬
‭OpenStax‬‭)‬

‭Bladder‬
‭ he‬‭bladder‬‭is located in the pelvic region and collects‬‭urine from both ureters (‬‭Figure‬
T
‭12.6‬‭). Due to its cellular composition, this organ‬‭can stretch to accommodate up to 500 -‬
‭600 mL of urine in adults. The structure of the bladder is similar between males and‬
‭females. However, there are some differences. Males have a prostate gland just below the‬
‭bladder, which surrounds the‬‭urethra‬‭. This gland produces‬‭secretions that make up the‬
‭non-cellular fluid of semen. In females, the bladder is in front of the uterus, which is the‬
‭o rgan where fetal development and pregnancy occur. During late pregnancy, the‬
‭bladder’s urine capacity is reduced due to compression by the enlarging uterus, resulting‬
‭in an increased frequency of urination. Chapter 14 (Reproductive Systems) discusses the‬
‭prostate gland and the uterus in more detail.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 12.6‬ ‭Comparison of the bladder in males and‬‭females. Note the presence of the‬
F
‭prostate gland in males. (‬‭NIH Image Gallery;‬‭Flickr).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭Sphincters‬
‭ rination occurs when the‬‭bladder empties (voids)‬‭and is controlled by smooth muscles‬
U
‭associated with the internal and external urethral‬‭sphincters‬‭. These are located at the‬
‭juncture of the base of the bladder and the urethra (‬‭Figure 12.7‬‭). This process involves‬
‭an interplay of involuntary and voluntary actions by the sphincters. When bladder volume‬
‭reaches about 150 mL, an urge to urinate is sensed but is easily overridden. Voluntary‬
‭control of urination relies on consciously preventing the relaxation of the external‬
‭urethral sphincter to maintain urinary continence. As the bladder fills, subsequent urges‬
‭become more challenging to ignore. Ultimately, the voluntary constraint will fail with‬
‭resulting incontinence (i.e., involuntary urination), which will occur as bladder volume‬
‭approaches 300 to 400 mL.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 12.7‬ ‭Location of the urinary sphincter muscles‬‭between the bladder and the‬
F
‭urethra. These muscles, under both voluntary and involuntary control, regulate the‬
‭movement of urine from the bladder into the urethra. (credit:‬ ‭Urinary Sphincter.png‬‭;‬
‭BruceBlaus‬‭;‬‭CC BY-SA 4.0‬‭)‬

‭ or additional information about control of urination, click the link below or scan the QR‬
F
‭code to watch the TED-Ed video by Heba Shaheed titled “Is it bad to hold your pee?”.‬

Is it bad to hold your pee? - Heba Shaheed

‭ rethra‬
U
‭As shown in‬‭Figure 12.7‬‭, the urethra is the structure‬‭that transports urine from the‬
‭bladder to the outside of the body. It is the only organ of the urinary system that shows‬
‭any significant difference between males and females. In males, the urethra is‬

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a‭ pproximately seven inches long and extends from the base of the bladder to the end of‬
‭the penis. A second key difference is that the male urethra functions not only in the‬
‭urinary system but also in the reproductive system, as it transports semen. In females, the‬
‭urethra is much shorter (approximately 1.5 inches compared to 7 to 8 inches in males)‬
‭and only transports urine. The shorter length in females makes it less of a barrier to fecal‬
‭bacteria than the longer male urethra. It is the most likely explanation for women's‬
‭greater incidence of urinary tract infections (UTIs).‬

‭12.4 Internal Gross Anatomy of the Kidney‬

‭ sectioned kidney reveals the‬‭c apsule‬‭, which is the‬‭o utermost layer and contains‬
A
‭connective tissue that provides protection and support for the organ (‬‭Figure 12.8‬‭).‬
‭Internally, the kidney has three regions—an outer‬‭cortex‬‭, a‬‭medulla‬‭in the middle, and‬
‭the‬‭renal pelvis‬‭at the innermost part of the kidney,‬‭from which urine drains into the‬
‭ureters. The renal cortex is granular due to the presence of nephrons—the functional‬
‭unit of the kidney that filters blood and produces urine. The medulla consists of multiple‬
‭pyramidal tissue masses called‬‭renal pyramids,‬‭and‬‭there are, on average, approximately‬
‭eight of them in each kidney. In between the pyramids are spaces called renal columns‬
‭through which the blood vessels pass. Urine drains through structures called‬‭collecting‬
‭ducts‬‭into the minor calyces, the major calyces, and‬‭the renal pelvis region. The renal‬
‭pelvis leads to the ureter on the outside of the kidney.‬

‭ igure 12.8‬ ‭The internal structure of the kidney‬‭is shown. (credit: modification of work‬
F
‭by NCI;‬‭OpenStax‬‭). A link to a video explanation of‬‭this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭12.5 Microscopic Anatomy of the Kidney‬

‭ he naked eye cannot see the renal structures that conduct the kidney's essential work, so‬
T
‭they are only with a microscope's aid. As mentioned previously, the functional unit of the‬
‭kidney is the nephron, illustrated in‬‭Figure 12.9‬‭.‬‭Each kidney contains over one million‬
‭nephrons that dot the renal cortex, giving it a granular appearance when sectioned‬
‭lengthwise. A nephron comprises a‬‭renal corpuscle‬‭,‬‭a‬‭renal tubule‬‭, and the associated‬
‭capillary network.‬

‭ igure 12.9‬ ‭The nephron is the functional unit of‬‭the kidney. The glomerulus and‬
F
‭convoluted tubules are located in the kidney cortex, and collecting ducts are found in the‬
‭pyramids of the medulla. Note the close association between the nephron and elements of‬
‭the circulatory system. (credit: modification of work by NIDDK;‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ he renal corpuscle in the renal cortex consists of a network of capillaries known as the‬
T
‭glomerulus‬‭and the capsule, a cup-shaped chamber that‬‭surrounds it, called‬‭Bowman's‬
‭c apsule‬‭(‬‭Figure 12.10‬‭). When blood passes through‬‭the glomerulus, 10 to 20 percent of‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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t‭ he plasma filters through the capillaries to be captured by Bowman’s capsule. The filtrate‬
‭then moves through the rest of the nephron. As mentioned, these glomerular capillaries‬
‭filter blood based primarily on particle size.‬
‭ he renal tubule is a long and convoluted (twisted) structure that extends from the‬
T
‭glomerulus and is divided into three parts based on function. The first part is called the‬
‭proximal convoluted tubule (PCT)‬‭because it is near‬‭the glomerulus; it stays in the renal‬
‭cortex. This tubule portion adjacent to the lumen is lined with epithelial cells with‬
‭finger-like projections (microvilli). These projections create a large surface area to‬
‭maximize the absorption and secretion of solutes (Na‬‭+‭,‬ ‬‭Cl‬‭–‬‭, glucose, etc.). As these‬
‭epithelial cells actively transport ions across their membranes, they possess a high‬
‭concentration of mitochondria to produce sufficient ATP.‬

‭ igure 12.10‬ ‭A diagram of an isolated nephron and‬

F
‭collecting duct without the blood vessels included in the‬
‭previous figure. (credit:‬‭OpenStax‬‭). A link to a‬‭video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ he second part is called the‬‭loop of Henle‬‭because‬‭it forms a loop (with descending and‬
T
‭ascending limbs) that goes through the renal medulla. The descending and ascending‬
‭portions of the loop of Henle are just continuations of the same tubule. They run adjacent‬
‭and parallel to each other after making a hairpin turn (loop of Henle) at the deepest point‬
‭o f their descent. Cellular differences in structure along this region result in differential‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ ermeability to solutes and water, which are essential to forming urine and maintaining‬
p
‭homeostasis.‬
‭ he third part of the renal tubule is called the‬‭distal‬‭convoluted tubule (DCT),‬‭also‬
T
‭located in the renal cortex. The DCT, the last part of the nephron, connects and empties its‬
‭contents into collecting ducts that line the medullary pyramids. The cells of the DCT also‬
‭pump ions against their concentration gradient, so they contain relatively large numbers‬
‭o f mitochondria, although fewer than in the PCT.‬
‭ he collecting ducts join with the nephrons but are not technically part of them. Each duct‬
T
‭collects filtrate from several nephrons for final modification. They are lined epithelial cells‬
‭that contain receptors for‬‭ADH‬‭. When stimulated by‬‭ADH, these cells insert‬‭aquaporin‬
‭c hannel proteins‬‭into their membranes, which, as their‬‭name suggests, allow water to‬
‭pass from the duct lumen through the cells and into the interstitial spaces to be recovered‬
‭by the circulatory system (‬‭Figure 12.11‬‭). This process‬‭allows for the recovery of large‬
‭amounts of water from the filtrate back into the blood, which causes the urine to become‬
‭more concentrated. In the absence of ADH, these channels are not inserted, resulting in‬
‭water excretion in the form of more dilute urine.‬

‭ igure 12.11‬ ‭An aquaporin channel inserted into‬‭the cell membrane (phospholipid‬
F
‭bilayer; Chapter 3). Positively charged amino acids inside the channel prevent the leakage‬
‭o f electrolytes (ions) across the cell membrane while allowing water to move due to‬
‭o smosis (credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭12.6 Blood Flow Through the Kidney‬

‭ ecause the kidney filters blood, its network of blood vessels is an essential component of‬
B
‭its structure and function. Renal blood supply starts with the‬‭renal arteries‬‭branching‬
‭from the aorta. It ends with the exiting of the‬‭renal‬‭veins‬‭to join the‬‭inferior vena cava‬
‭(‬‭Figure 12.12‬‭). The branch of the circulatory system‬‭that enters the glomerulus is called‬
‭the‬‭afferent arteriole,‬‭and the branch that exits‬‭the glomerulus is called the‬‭efferent‬
‭arteriole‬‭.‬
‭ he network of capillaries within the glomerulus is called the glomerular capillary bed.‬
T
‭Once the efferent arteriole exits the glomerulus, it forms the‬‭peritubular capillary‬
‭network‬‭, which surrounds and interacts with parts‬‭o f the renal tubule. As the glomerular‬
‭filtrate progresses through the nephron, these capillary networks recover most of the‬
‭solutes and water and return them to circulation.‬
‭ he blood flow through the kidney must be appropriate to allow for filtration. The kidneys‬
T
‭are very effective at regulating the blood flow rate over a wide range of blood pressures,‬
‭which is essential to consistent filtration. Your blood pressure will decrease when you‬
‭relax or sleep and increase when exercising. However, despite these changes, the filtration‬
‭rate through the kidney will change very little.‬

‭ igure 12.12‬ ‭The two capillary beds are‬

F
‭shown in this figure. The afferent arteriole‬
‭leads to the glomerulus (first bed) inside of‬
‭Bowman’s capsule (glomerular capsule); the‬
‭efferent arteriole transports blood from the‬
‭first bed to the peritubular capillaries (second‬
‭bed). (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭12.7 Stages of Urine Formation‬

‭ aving reviewed the anatomy and microanatomy of the urinary system, you can now‬
H
‭focus on the physiology of urine formation. Nephrons take a simple blood filtrate and‬
‭modify it into urine. Many changes occur in the nephron's different parts before the urine‬
‭is created for disposal. The principle task of the nephron population is to balance the‬
‭plasma to homeostatic set points and excrete potential toxins in the urine. They do this by‬
‭accomplishing three principal functions—filtration, reabsorption, and secretion. First, the‬
‭nephrons filter blood that runs through the capillary network in the glomerulus. All‬
‭solutes, except large proteins, are filtered out into the glomerulus by‬‭glomerular‬
‭filtration‬‭. Second, the filtrate is collected in the‬‭renal tubules. Most of the solutes get‬
‭reabsorbed in the PCT by‬‭tubular reabsorption‬‭. In‬‭the loop of Henle, the filtrate‬
‭continues to exchange solutes and water with the renal medulla and the peritubular‬
‭capillary network. Water is also reabsorbed during this step. Then, additional solutes,‬
‭wastes, and certain drugs are secreted into the kidney tubules during‬‭tubular secretion‬‭,‬
‭which is essentially the opposite process of tubular reabsorption. The collecting ducts‬
‭collect filtrate from the nephrons, which is ultimately moved into the renal pelvis, which‬
‭connects to the ureter. This entire process is illustrated in‬‭Figure 12.13‬‭.‬
‭ echanisms by which substances move across membranes for reabsorption or secretion‬
M
‭include active transport, simple diffusion, facilitated diffusion, and osmosis (Chapter 3).‬
‭Active transport‬‭utilizes energy, usually ATP, to‬‭move a specific substance across a‬
‭membrane from a low to a high concentration. An example is the active transport of‬
‭sodium ions (Na‬‭+‭)‬ out of a cell and potassium ions‬‭(K‬‭+‬‭) into a cell by the‬
‭membrane-bound, sodium-potassium ion pump, which is a protein. Both ions are moved‬
‭in opposite directions from a lower to a higher concentration.‬ ‭Simple diffusion‬‭moves‬
‭small, nonpolar substances directly across the cell membrane from a higher to a lower‬
‭concentration. No ATP is required for simple diffusion, as the concentration differences‬
‭provide energy.‬ ‭Facilitated diffusion‬‭is similar‬‭to simple diffusion in that a substance‬
‭moves down its concentration gradient. The difference is that it requires specific‬
‭membrane receptors or channel proteins for movement. The movement of glucose and, in‬
‭certain situations, Na‬‭+‬ ‭ions are examples of facilitated‬‭diffusion.‬‭Osmosis‬‭is the diffusion of‬
‭water through channel proteins called aquaporins.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 12.13‬ ‭Each part of the nephron performs a‬‭different function in filtering waste‬
F
‭and maintaining homeostatic balance. (1) The glomerulus forces small solutes out of the‬
‭blood by pressure. (2) The proximal convoluted tubule reabsorbs ions, water, and‬
‭nutrients from the filtrate into the interstitial fluid and actively transports toxins and drugs‬
‭from the interstitial fluid into the filtrate. (3) The descending loop of Henle is lined with‬
‭cells containing aquaporins that allow water to pass from the filtrate into the interstitial‬
‭fluid. (4) In the thin part of the ascending loop of Henle, Na‬‭+‬ ‭and Cl‬‭–‬ ‭ions diffuse into the‬
‭interstitial fluid. These ions are transported into the interstitial fluid in the thick part.‬
‭Because salt but not water is lost, the filtrate becomes more dilute as it travels up the limb.‬
‭(5) In the distal convoluted tubule, K‬‭+‬ ‭and H‬‭+‬ ‭ions‬‭are selectively secreted into the filtrate,‬
‭while Na‬‭+‬‭, Cl‬‭–‬‭, and HCO‬‭3‭–‬ ‬ ‭ions are reabsorbed to maintain‬‭pH and electrolyte balance in the‬
‭blood. (6) The collecting duct reabsorbs solutes and water from the filtrate, forming dilute‬
‭urine. (credit: modification of work by NIDDK;‬‭OpenStax‬‭).‬‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭12.8 Hormones That Influence Urine Formation‬

‭ s discussed, two hormones influencing total body water are aldosterone and ADH.‬
A
‭Aldosterone‬‭, often called the “salt-retaining hormone,”‬‭is synthesized and released from‬
‭the adrenal cortex. It promotes Na‬‭+‬ ‭reabsorption by‬‭the nephron, promoting the retention‬
‭o f water. Almost all of the sodium in the blood is reclaimed by the renal tubules under the‬
‭influence of aldosterone. Because sodium is constantly reabsorbed by active transport‬
‭and water follows sodium to maintain osmotic balance, aldosterone manages sodium and‬
‭water levels in body fluids. Patients who have‬‭Addison's‬‭disease‬‭have a failing adrenal‬
‭cortex and cannot produce aldosterone. They lose sodium in their urine constantly, which‬
‭can be fatal if the supply is not replenished.‬
‭ rogesterone is a steroid structurally similar to aldosterone. It binds to the aldosterone‬
P
‭receptor, weakly stimulates Na‬‭+‬ ‭reabsorption, and‬‭increases water recovery. This process‬
‭is unimportant in men due to low levels of circulating progesterone. However, when‬
‭progesterone levels increase, it may cause increased water retention during women’s‬
‭menstrual cycles.‬
‭ iuretics‬‭are drugs that can increase water loss by‬‭interfering with the recapture of‬
D
‭solutes and water from the forming urine. They are often prescribed to lower blood‬
‭pressure. Coffee, tea, and alcoholic beverages are familiar diuretics. ADH, released by the‬
‭posterior pituitary, works to do the exact opposite. It promotes water recovery, decreases‬
‭urine volume, and maintains plasma solute concentrations and blood pressure. It does so‬
‭by stimulating the insertion of aquaporin proteins into the cell membrane of specific‬
‭collecting duct cells to form water channels, allowing the movement of water from the‬
‭lumen of the collecting duct into the interstitial space in the medulla of the kidney by‬
‭o smosis. From there, it enters the capillaries to return to the circulation. ADH also acts as‬
‭a vasoconstrictor to narrow blood vessels and increase blood pressure during bleeding.‬
‭ iuretics may be prescribed for hypertension to reduce blood volume and blood‬
D
‭pressure. The most frequently prescribed diuretic for this purpose is hydrochlorothiazide.‬
‭It inhibits the Na‬‭+‭/‬ Cl‬‭-‬ ‭transporter in the DCT and‬‭collecting duct. The result is a loss of Na‬‭+‬
‭with water following passively via osmosis.‬
‭ nother example of a diuretic is the indigestible sugar mannitol, which is most often‬
A
‭administered to reduce brain swelling after head injury. However, it is not the only sugar‬
‭that can produce a diuretic effect. In cases of poorly controlled diabetes mellitus, glucose‬
‭levels exceed the capacity of the tubular glucose transporters, resulting in glucose in the‬
‭urine. The unrecovered glucose becomes a potent diuretic.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭12.9 Career Connections - Dialysis Technician and Nephrologist‬

‭Dialysis Technician‬

‭ ialysis‬‭is a medical process of removing wastes and‬‭excess water from the blood by‬
D
‭diffusion and ultrafiltration (‬‭Figure 12.14‬‭). When‬‭kidney function fails, dialysis is done to‬
‭rid the body of wastes artificially. This is a vital process to keep patients alive. Sometimes,‬
‭the patients undergo artificial dialysis until they are eligible for a kidney transplant.‬
‭Dialysis is a life-long necessity in others who are not candidates for kidney transplants.‬

‭ ialysis technicians typically work in hospitals and clinics. While some roles in this field‬
D
‭include equipment development and maintenance, most dialysis technicians work in direct‬
‭patient care. Their on-the-job duties, which typically occur under the direct supervision of‬
‭a registered nurse, focus on providing dialysis treatments. This can include reviewing‬
‭patient history and current condition, assessing and responding to patient needs before‬
‭and during treatment, and monitoring the dialysis process. Treatment may include taking‬
‭and reporting a patient’s vital signs and preparing solutions and equipment to ensure‬
‭accurate and sterile procedures.‬

‭ igure 12.14‬ ‭An overview of the process of dialysis.‬

F
‭(credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Nephrologist‬

‭ ephrology is a subspecialty of internal medicine. To become a nephrologist, students‬

N
‭complete medical school and additional training to become certified in internal medicine.‬
‭Two or more additional years are spent studying kidney disorders and their‬
‭accompanying effects on the body.‬

‭ nephrologist studies and deals with diseases of the kidneys—both those that cause‬
A
‭kidney failure (such as diabetes) and the conditions produced by kidney disease (such as‬
‭hypertension). Blood pressure, blood volume, and changes in electrolyte balance come‬
‭under the purview of a nephrologist.‬

‭ ephrologists usually work with other physicians who refer patients to them or consult‬
N
‭with them about specific diagnoses and treatment plans. Patients are typically referred to‬
‭a nephrologist for symptoms such as blood or protein in the urine, very high blood‬
‭pressure, kidney stones, or renal failure.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 13: Cell Cycle and Cell Division‬

‭ igure 13.1‬ ‭A‬‭Bright field microscopy image of root‬‭meristem of onion showing cells in‬
F
‭interphase or one of the stages of mitosis: prophase, metaphase, anaphase, or telophase‬
‭(Magnification: 1600x)‬‭. (credit:‬ ‭root meristem of‬‭o nion (cells in prophase, metaphase,‬
‭anaphase, telophase).jpg‬‭;‬‭Dr. Josef Reischig, CSc.‬‭;‬‭CC BY-SA 3.0‬‭)‬

‭13.1 Introduction‬
‭ he ability to reproduce is a fundamental characteristic of all organisms. Although many‬
T
‭unicellular organisms (e.g., bacteria) produce genetically identical clones of themselves‬
‭through‬‭asexual reproduction‬‭, most multicellular organisms‬‭reproduce regularly using‬
‭another method—‬‭sexual reproduction.‬‭This highly evolved‬‭method involves the‬
‭production by parents of two‬‭haploid‬‭cells (‬‭gametes‬‭;‬‭sperm and ovum) and the fusion of‬
‭the two haploid cells via‬‭fertilization‬‭to form a‬‭single,‬‭diploid‬‭cell called a‬‭zygote‬‭(‬‭Figure‬
‭13.1‬‭)—a genetically unique organism.‬
I‭ n our species, billions of cell divisions are required to produce a complex, multicellular‬
‭human comprising trillions of cells. Thus, the original zygote is the ancestor of all cells in‬
‭the body. However, cell reproduction is still necessary once a human is fully grown. All‬
‭multicellular organisms use cell division to grow, maintain, and repair cells and tissues.‬
‭Cell division is closely regulated, and the occasional failure of this regulation can have‬
‭life-threatening consequences.‬

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‭13.2 Genomic DNA, Chromosomes, and Compaction‬

‭ efore discussing the steps a cell must undertake to replicate, a deeper understanding of‬
B
‭the structure and function of a cell’s genetic information is necessary. A cell’s DNA,‬
‭packaged as a double-stranded DNA molecule(s), is called its‬‭genome‬‭. In eukaryotes, the‬
‭genome consists of double-stranded linear DNA molecules packaged into structures called‬
‭c hromosomes‬‭(‬‭Figure 13.2‬‭). Each eukaryotic species‬‭has a characteristic number of‬
‭chromosomes in the nuclei of its cells. A human gamete has 23 chromosomes, while each‬
‭human body cell, called a‬‭somatic cell‬‭, has 46 chromosomes.‬
‭ he chromosome content of a gamete, called the‬‭haploid‬‭number‬‭, is designated by the‬
T
‭letter‬‭n‬‭and represents one set of chromosomes. For‬‭example, in humans, n=23. A‬
‭somatic cell contains two sets of chromosomes, one from the sperm and one from the egg,‬
‭and is designated as‬‭2n‬‭(‬‭diploid number‬‭). Therefore,‬‭in humans, 2n=46.‬

‭ igure 13.2‬ ‭There are 46 chromosomes (2 sets of 23‬‭each) in each human somatic cell.‬
F
‭The condensed chromosomes are viewed within the nucleus (top), removed from a cell in‬
‭mitosis, spread out on a slide (right), and artificially arranged according to length (left);‬
‭an arrangement like this is called a‬‭karyotype‬‭. In‬‭this image, the chromosomes were‬
‭exposed to fluorescent stains to differentiate the different chromosomes. A staining‬
‭method, "chromosome painting,” employs fluorescent dyes that highlight chromosomes in‬
‭various colors. As there are two X chromosomes in the karyotype, we can conclude the‬
‭chromosomes are from a female somatic cell. (credit: National Human Genome‬
‭Project/NIH;‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ atched pairs of chromosomes in a diploid organism are called‬‭homologous‬

M
‭c hromosomes‬‭(‬‭Figure 13.3‬‭). Homologous chromosomes‬‭are the same length and have‬
‭specific nucleotide segments called‬‭genes‬‭in the same‬‭chromosomal location or‬‭loci‬
‭(singular: locus). Genes, the functional units of chromosomes, determine specific‬
‭characteristics (‬‭phenotypes‬‭) by coding for specific‬‭proteins (Chapter 4; transcription and‬
‭translation). Traits are the variations of those characteristics. For example, hair color is a‬
‭characteristic with different traits (e.g., blonde, brown, and black).‬

‭ igure 13.3‬ ‭An example of haploid (n) versus diploid‬‭(2n) chromosome numbers. The‬
F
‭haploid cell (n=3) contains one member of each homologous pair present in the diploid‬
‭cell (2n=6). (credit:‬ ‭Haploid vs. diploid.svg‬‭;‬‭Ehamberg‬‭;‬‭CC BY-SA 3.0‬‭).‬ ‭A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ ach member of a homologous pair of chromosomes originates from a different parent;‬

E
‭therefore, the genes' sequences are not necessarily identical. The phenotypic variation of‬
‭individuals within a species is due, at least in part, to the specific combination of the genes‬
‭inherited from both parents. Chapter 7 includes a discussion of the ABO and Rh blood‬
‭typing systems. Refer back to that content to refresh your knowledge of the following‬
‭terms: genotype, alleles, phenotype, homozygous, heterozygous, dominant, recessive, and‬
‭codominant.‬

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‭Chapter 13: Cell Cycle and Cell Division‬

‭ inor variations of traits, such as blood type, eye color, and handedness, contribute to the‬
M
‭natural variation found within a species. However, the difference is less than one percent‬
‭o f the entire DNA sequence from any pair of human homologous chromosomes. The‬‭sex‬
‭c hromosomes‬‭X and Y are the single exception to the‬‭rule of homologous chromosome‬
‭uniformity: Other than a small amount of homology necessary to accurately produce‬
‭gametes, the genes found on the X and Y chromosomes are different.‬
I‭ f the DNA from all 46 chromosomes in a human cell nucleus were laid out end to end, it‬
‭would measure approximately 6.5 feet; however, its diameter would be only about‬
‭0.00000008 inches! Considering that the size of a typical human cell is about 0.0004‬
‭inches, DNA must be tightly packaged to fit in the cell’s nucleus, which is smaller than the‬
‭size of the cell. At the same time, it must also be readily accessible for the genes to be‬
‭expressed (i.e., transcription and translation). How are chromosomes compacted to‬
‭achieve this outcome?‬
I‭ n the first level of‬‭c hromosome compaction‬‭, short‬‭stretches of the DNA double helix‬
‭wrap around a core of eight histone proteins at regular intervals along the entire length of‬
‭the chromosome (‬‭Figure 13.4‬‭). The DNA-histone complex‬‭is part of the‬‭c hromatin‬‭. Each‬
‭bead-like histone-DNA complex is called a‬‭nucleosome‬‭.‬‭The second level of compaction‬
‭o ccurs as the nucleosomes and the DNA between them are coiled into a thicker chromatin‬
‭fiber. This coiling further shortens the chromosome, about 50 times shorter than the‬
‭extended form. In the third level of packing, a variety of fibrous proteins is used to pack‬
‭the chromatin. These fibrous proteins also ensure that each chromosome in a‬
‭non-dividing cell occupies a particular area of the nucleus that does not overlap with any‬
‭o ther chromosome.‬

‭13.3 The Cell Cycle‬

‭ ou have read numerous times in this chapter about the importance and prevalence of cell‬
Y
‭division. While a few cells in the body do not undergo cell division (such as gametes, red‬
‭blood cells, most neurons, and some muscle cells), most somatic cells divide regularly. As‬
‭mentioned, a somatic cell is a general term for a body cell; all human cells, except gametes,‬
‭are somatic cells.‬
‭ ells in the body replace themselves over the lifetime of a person. For example, the cells‬
C
‭lining the gastrointestinal tract must be frequently replaced when constantly “worn off” by‬
‭food movement through the gut. But what triggers a cell to divide, and how does it‬
‭prepare for and complete cell division? The‬‭cell cycle‬‭is an ordered series of events‬
‭involving cell growth and division that produces two new daughter cells. Cells on the path‬
‭to cell division proceed through a series of precisely timed and carefully regulated stages‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 13: Cell Cycle and Cell Division‬

‭ f growth, DNA replication, and nuclear and cytoplasmic division that ultimately produces‬
o
‭two genetically identical (clone) cells.‬

‭ igure 13.4‬ ‭Double-stranded DNA wraps around histone‬‭proteins to form nucleosomes‬

F
‭that appear as “beads on a string.” The nucleosomes are coiled into a thicker chromatin‬
‭fiber. When a cell undergoes mitosis, the chromosomes condense/compact even further.‬
‭Notice that the fully condensed chromosome is duplicated because such compaction only‬
‭o ccurs before cell division (mitosis or meiosis) when the duplicated chromosome’s sister‬
‭chromatids will be separated. (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ ach “turn” of the cell cycle can be divided into‬‭the‬‭interphase‬‭and the‬‭mitotic phases‬‭.‬
E
‭Interphase is the period of the cell cycle during which the cell is not dividing but is‬
‭preparing for this event. The majority of cells are in interphase most of the time. The‬
‭mitotic phase includes mitosis and cytokinesis.‬‭Mitosis‬‭is the division of nuclear genetic‬
‭material (chromosomes), during which the cell nucleus breaks down, and two new, fully‬
‭functional nuclei are formed.‬‭Cytokinesis‬‭divides‬‭the cytoplasm into two distinctive cells.‬

‭Interphase‬
I‭ nterphase is subdivided into G‬‭1‭,‬ S, and G‬‭2‬‭subphases. During‬‭G‬‭1‬ ‭(gap 1 phase), the cell‬
‭accumulates the building blocks of chromosomal DNA and the associated proteins and‬
‭accumulates sufficient energy reserves to replicate each chromosome in the nucleus‬
‭(‬‭Figure 13.5‬‭). For cells that will divide again,‬‭G‭1‬ ‬ ‭is followed by DNA replication during the‬
‭S‬‭(synthesis)‬‭phase‬‭. In other words, each chromosome‬‭replicates to produce two sister‬
‭chromatids joined at the centromere (‬‭Figure 13.6‬‭).‬

‭ igure 13.5‬ ‭The two major cell cycle phases include‬‭the mitotic phase and interphase.‬
F
‭The first phase includes mitosis (designated M), which is nuclear division, and cytokinesis‬
‭(designated C) when the cytoplasm divides and two daughter cells are produced. During‬
‭interphase, the cell grows and performs all of its normal functions. Interphase is further‬
‭subdivided into G‬‭1‬‭, S, and G‬‭2‬ ‭phases. The DNA is‬‭replicated during the S phase of‬
‭interphase (credit:‬ ‭The Cell Cycle.svg‬‭;‬‭George Welle‬‭r‬‭;‬‭CC BY-SA 3.0‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 13: Cell Cycle and Cell Division‬

‭ fter synthesis, the cell proceeds through the‬‭G‬‭2‬ ‭(gap 2) phase, replenishing its energy‬
A
‭stores and synthesizing proteins necessary for chromosome manipulation and movement.‬
‭Some cell organelles are duplicated, and the cell's cytoskeleton, which provides structural‬
‭support, is dismantled to provide resources for the mitotic phase.‬

‭ igure 13.6‬ ‭A pair of duplicated homologous chromosomes‬‭with their attached sister‬

F
‭chromatids produced by DNA replication during the S phase of interphase. The red and‬
‭blue colors distinguish the parental origin of each homolog in the pair. Make sure to‬
‭differentiate between the concepts of sister chromatids (one chromosome and its exact‬
‭copy attached during mitosis) and homologous chromosomes (two paired chromosomes‬
‭inherited separately, one from each parent). (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ ells' durations of the G‬‭1‬‭, S, and G‬‭2‬ ‭phases vary‬‭the most. A cell might spend several hours‬
C
‭o r many days in this phase. The S phase typically lasts 8-10 hours, and the‬‭G‬‭2‬ ‭phase‬
‭approximately five hours.‬
‭ ot all cells that begin the G‬‭1‬ ‭phase of interphase‬‭will continue through the cell cycle.‬
N
‭Instead, they will enter a phase designated‬‭G‬‭0‭,‬ a‬‭“resting” cell cycle phase. Cells that have‬
‭temporarily stopped dividing and are resting (a common condition) and cells that have‬
‭permanently ceased dividing (such as nerve cells) are said to be in G‬‭0‭.‬ In some cases, cells‬
‭transitioning from G‬‭1‬ ‭to G‬‭0‬ ‭can return to G‬‭1‬ ‭and continue‬‭through the subsequent cell‬
‭cycle events.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭The Mitotic Phase: Mitosis and Cytokinesis‬

‭ he mitotic phase of the cell typically takes one to two hours to complete. During this time,‬
T
‭a cell undergoes two major processes. First, it completes mitosis, during which the‬
‭contents of the nucleus are equitably pulled apart and distributed between its two halves.‬
‭Second, cytokinesis divides the cytoplasm into two new cells.‬
‭ fter interphase, mitosis is subdivided into five major stages: prophase, prometaphase,‬
A
‭metaphase, anaphase, and telophase (‬‭Figure 13.7‬‭).‬
‭ rophase‬‭is the first phase of mitosis, during which the loosely packed chromatin coils‬
P
‭and condenses into visible chromosomes. Each chromosome becomes visible during‬
‭prophase with its identical sister chromatid attached, forming the familiar X-shape. The‬
‭mitotic spindle, consisting of the centrioles and the microtubules, begins to organize, and‬
‭the nuclear envelope also disintegrates in preparation for separating the chromatids.‬
‭ rometaphase‬‭is the second phase of mitosis and occurs‬‭near the end of prophase when‬
P
‭microtubules invade the nuclear area from the mitotic spindle. The nuclear membrane has‬
‭disintegrated, and the microtubules attach themselves to the centromeres that adjoin pairs‬
‭o f sister chromatids (‬‭Figure 13.8‬‭). The‬‭kinetochore‬‭is a protein structure on the‬
‭centromere, the attachment point between the mitotic spindle and the sister chromatids.‬
‭This stage is called late prophase or “prometaphase” to indicate the transition between‬
‭prophase and metaphase.‬
‭ etaphase‬‭is the third stage of mitosis. During this‬‭stage, the sister chromatids, with their‬
M
‭attached microtubules, line up along a linear plane in the middle of the cell. A metaphase‬
‭plate forms between the centrosomes at either end of the cell. The metaphase plate is the‬
‭plane that goes through the center of the spindle on which the sister chromatids are‬
‭positioned. The microtubules are now poised to pull apart the sister chromatids and bring‬
‭o ne from each pair to each side of the cell.‬
‭ naphase‬‭is the fourth stage of mitosis. Anaphase‬‭o ccurs over a few minutes when the‬
A
‭pairs of sister chromatids are separated, forming individual chromosomes again. These‬
‭chromosomes are pulled to opposite ends of the cell by their kinetochores as the‬
‭microtubules shorten. Each end of the cell receives one partner from each pair of sister‬
‭chromatids, ensuring that the two new daughter cells will contain identical genetic‬
‭material.‬
‭ elophase‬‭is the final stage of mitosis. Telophase‬‭is characterized by forming two new‬
T
‭daughter nuclei at either end of the dividing cell. These newly formed nuclei surround the‬
‭genetic material, which uncoils such that the chromosomes return to loosely packed‬
‭chromatin. The mitotic spindle breaks apart, and each new cell receives its complement of‬
‭DNA, organelles, membranes, and centrioles. At this point, the cell begins to split in half as‬
‭cytokinesis begins.‬

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‭ igure 13.7‬ ‭Mitosis is divided into five stages—prophase,‬‭prometaphase, metaphase,‬

F
‭anaphase, and telophase. The pictures at the bottom were taken by fluorescence‬
‭microscopy (hence, the black background) of cells artificially stained by fluorescent dyes:‬
‭blue fluorescence indicates DNA (chromosomes), and green fluorescence indicates‬
‭microtubules (spindle apparatus). (credit “mitosis drawings”: modification of work by‬
‭Mariana Ruiz Villareal; credit “micrographs”: modification of work by Roy van Heesbeen;‬
‭“cytokinesis micrograph”; Wadsworth Center/New York State Department of Health;‬
‭scale-bar data from Matt Russell). A link to a video explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 13.8‬ ‭During prometaphase, mitotic spindle‬‭microtubules from opposite poles‬

F
‭attach to each sister chromatid at the kinetochore. In anaphase, the connection between‬
‭the sister chromatids breaks down, and the microtubules pull the chromosomes toward‬
‭o pposite poles. (credit:‬‭OpenStax‬‭)‬

‭ ytokinesis occurs due to the action of a contractile band made up of microfilaments that‬
C
‭form around the cell's midline. As the band shortens, a “fissure” called the‬‭c leavage‬
‭furrow‬‭is created. The band continues to shorten,‬‭deepening the furrow until the two‬
‭parts finally separate, producing genetically identical daughter cells.‬
I‭ t can be understandably challenging for students to follow changes in the chromosome‬
‭number and DNA content of cells during the mitotic phase. While‬‭Figure 13.9‬‭contains‬
‭substantial detail, moving through it step-by-step and carefully reading the figure‬
‭description should help clarify these concepts.‬

‭13.4 Cell Cycle Control‬

‭ very elaborate and precise regulation system controls how cells proceed from one‬
A
‭phase to the next in the cell cycle and begin mitosis. The control system involves molecules‬
‭within the cell and external signals, which function as “triggers” to regulate movement‬
‭(both stopping and advancing) through the different stages of the cycle. Precise regulation‬
‭o f the cell cycle is critical for maintaining the health of an organism.‬
‭ s the cell proceeds through its cycle, each phase involves specific processes that must be‬
A
‭completed before it advances to the next phase. A‬‭c heckpoint‬‭is a point in the cell cycle at‬
‭which the cycle can be signaled to move forward or stop.‬

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‭ igure 13.9‬ ‭A summary of the stages of the mitotic‬‭phase (mitosis and cytokinesis) that‬
F
‭shows changes in the associated chromosome number and DNA content. A diploid (2n)‬
‭cell begins interphase with its complement of unduplicated chromosomes (4 in the‬
‭example above; 2 pairs of homologous chromosomes) and a total DNA content designated‬
‭as 2c. When the chromosomes duplicate during the S phase, the number of chromosomes‬
‭remains the same (2n) because sister chromatids are joined at the centromere. In other‬
‭words, a duplicated chromosome is still counted as one chromosome. However, the DNA‬
‭content is doubled from 2c to 4c. At anaphase and telophase, sister chromatids separate.‬
‭However, both sets of chromosomes are still in one cell, so the DNA content is represented‬
‭as 2c x 2 and the chromosome number as 2n x 2. After cytokinesis, each daughter cell has‬
‭a DNA content 2c and a chromosome number 2n. (credit:‬‭Mitosis diagram.jpg‬‭; Marek‬
‭Kultys;‬‭CC BY-SA 3.0‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

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‭Chapter 13: Cell Cycle and Cell Division‬

‭ igure 13.10‬‭shows the G1, G2, and M checkpoints. Different types of intracellular‬
F
‭molecules provide the stop or go signals at each one, depending on certain conditions‬
‭within the cell. At the‬‭G‬‭1‬ ‭c heckpoint‬‭, the cell must‬‭be ready for DNA synthesis. At the‬‭G‬‭2‬
‭c heckpoint,‬‭the cell must be fully prepared for mitosis, including proper replication of the‬
‭chromosomes. Even during mitosis, a crucial stop-and-go checkpoint in metaphase‬
‭ensures the cell is fully prepared to complete cell division. The‬‭metaphase checkpoint‬
‭ensures that all sister chromatids are correctly attached to their respective microtubules‬
‭and lined up at the metaphase plate before the signal is given to separate them during‬
‭anaphase.‬

‭ igure 13.10‬ ‭The cell cycle is controlled at three‬‭checkpoints. The integrity of the DNA is‬
F
‭assessed at the G‬‭1‬ ‭checkpoint. Proper chromosome duplication‬‭is verified at the G‬‭2‬
‭checkpoint. The attachment of each kinetochore to a spindle fiber is assessed at the M‬
‭checkpoint. The mitotic phase consists of mitosis followed by cytokinesis. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 13: Cell Cycle and Cell Division‬

‭13.5 Sexual Reproduction and Meiosis Overview‬

S‭ exual reproduction was likely an early evolutionary innovation after the appearance of‬
‭eukaryotic cells. It appears to have been very successful because most eukaryotes can‬
‭reproduce sexually; in many animals, it is the only mode of reproduction. And yet,‬
‭scientists also recognize some real disadvantages to sexual reproduction. On the surface,‬
‭creating offspring that are genetic clones of the parent is a better system. If the parent‬
‭o rganism successfully occupies a habitat, offspring with the same traits should be similarly‬
‭successful. There is also the obvious benefit to an organism that can produce offspring‬
‭whenever circumstances are favorable by asexual budding, fragmentation, or by creating‬
‭eggs asexually. These methods of reproduction do not require another organism of the‬
‭o pposite sex. Indeed, some organisms that lead a solitary lifestyle have retained the ability‬
‭to reproduce asexually. In addition, in asexual populations, every individual is capable of‬
‭reproduction. In sexual populations, the males are not producing the offspring themselves,‬
‭so hypothetically, an asexual population could grow twice as fast.‬
‭ owever, multicellular organisms that exclusively depend on asexual reproduction are‬
H
‭exceedingly rare. Why are meiosis and sexual reproductive strategies so common? These‬
‭are important (and as yet not wholly answered) questions in biology, even though they‬
‭have been the focus of much research beginning in the latter half of the 20th century.‬
‭There are several possible explanations, one of which is that the variation that sexual‬
‭reproduction creates among offspring is significant to the survival and reproduction of the‬
‭population. Thus, on average, a sexually reproducing population will leave more‬
‭descendants than an otherwise similar asexually reproducing population. The only source‬
‭o f variation in asexual organisms is mutation. Mutations that occur during the formation‬
‭o f cells destined to become gametes are also the ultimate source of variation in sexually‬
‭reproducing organisms.‬
‭ owever, in contrast to mutations during asexual reproduction, mutations during sexual‬
H
‭reproduction can be continually reshuffled from one generation to the next when‬
‭different parents combine their unique genomes. The genes are mixed into different‬
‭combinations by two mechanisms that will be further discussed: crossovers during‬
‭prophase I and random assortment at metaphase I.‬

‭Life Cycle of Sexually Reproducing Organisms‬

‭ ertilization and meiosis alternate in sexual life cycles. The process of‬‭meiosis‬‭reduces the‬
F
‭chromosome number by half, from 2n to n. Fertilization, the joining of two haploid (n)‬
‭gametes, restores the diploid (2n) condition. Nearly all animals employ a‬
‭diploid-dominant life-cycle strategy‬‭in which the‬‭o nly haploid cells produced by the‬
‭o rganism are the‬‭gametes‬‭(sperm and egg). Early in‬‭the embryo's development,‬

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s‭ pecialized diploid cells, called‬‭germ cells‬‭, are made within the‬‭gonads‬‭(such as the‬‭testes‬
‭and‬‭ovaries‬‭). Germ cells are capable of mitosis to‬‭perpetuate the germ cell line and‬
‭meiosis to produce haploid gametes. Once the haploid gametes are formed, they lose the‬
‭ability to divide again. Fertilization occurs with the fusion of two gametes, usually from‬
‭different individuals, restoring the diploid state (‬‭Figure 13.11‬‭).‬

‭Meiosis Overview‬
S‭ exual reproduction requires the union of two specialized haploid cells called gametes,‬
‭each containing one set of chromosomes (n). When gametes unite, they form a zygote, or‬
‭fertilized egg, that includes two sets of chromosomes (diploid; 2n). Each member of a‬
‭chromosome set has a similar member in the other set, and the pair is referred to as‬
‭homologous. In other words, diploid organisms inherit one copy of each homologous‬
‭chromosome from each parent.‬

‭ igure 13.11‬ ‭In animals, sexually reproducing adults‬‭form haploid (n) gametes from‬
F
‭diploid (2n) germ cells located in the gonads (testes or ovaries). Fusion of the gametes‬
‭gives rise to a fertilized egg cell or zygote. The zygote will undergo many rounds of mitosis‬
‭to produce multicellular offspring. The germ cells are generated early in the development‬
‭o f the zygote. (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 13: Cell Cycle and Cell Division‬

I‭ f the reproductive cycle is to continue for any sexually reproducing species, then the‬
‭diploid (2n) cell must somehow reduce its number of chromosome sets to produce‬
‭haploid (n) gametes; otherwise, the number of chromosome sets will double with every‬
‭future round of fertilization. Therefore, sexual reproduction requires a nuclear division‬
‭that reduces the number of chromosome sets by half. The type of nuclear division that‬
‭accomplishes this outcome is called meiosis.‬
‭ s with mitosis, DNA replication occurs before meiosis during the S phase of the cell cycle‬
A
‭so that each chromosome becomes a pair of sister chromatids. In meiosis, there are two‬
‭rounds of nuclear division resulting in four nuclei and usually four daughter cells, each‬
‭with half the number of chromosomes as the parent cell. The first division separates‬
‭homologs (homologous chromosomes) and is also referred to as‬‭reductional division‬
‭because it reduces the number of chromosome sets from two (2n) to one (n). The second‬
‭division—like mitosis—separates sister chromatids into individual, unduplicated‬
‭chromosomes. As it doesn’t further reduce the number of sets of chromosomes, it is also‬
‭referred to as‬‭equational division‬‭.‬
‭ eiosis and mitosis share similar processes but have distinct outcomes. Mitotic divisions‬
M
‭are single nuclear divisions that produce genetically identical daughter nuclei (i.e., each‬
‭daughter nucleus has the same number of chromosome sets as the original cell). In‬
‭contrast, meiotic divisions include two nuclear divisions that produce genetically different‬
‭daughter nuclei with only one chromosome set (n) instead of the two sets (2n) of‬
‭chromosomes in the parent cell.‬

‭13.6 The Process of Meiosis‬

I‭ n meiosis, the starting nucleus is always diploid (2n), and the resulting daughter nuclei‬
‭are haploid (n). Meiosis consists of one round of chromosome replication followed by two‬
‭rounds of nuclear division to achieve this reduction in chromosome number. Because the‬
‭events that occur during each division stage are analogous to the events of mitosis, the‬
‭same stage names are assigned. However, because there are two division rounds, the‬
‭major process and the stages are designated with an “I” or a “II.” Thus,‬‭meiosis I‬‭is the‬
‭first round of meiotic division and consists of prophase I, prometaphase I, and so on.‬
‭Likewise, meiosis II (during which the second round of meiotic division occurs) includes‬
‭prophase II, prometaphase II, and so on.‬

‭Meiosis I‬
‭ eiosis is preceded by an interphase consisting of G‬‭1‭,‬ S, and G‬‭2‬ ‭phases, nearly identical to‬
M
‭the stages preceding mitosis. The G‬‭1‬ ‭phase is focused‬‭o n cell growth. During the S phase,‬
‭the cell replicates the DNA of the chromosomes to produce sister chromatids attached at‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 13: Cell Cycle and Cell Division‬

t‭ he centromere. Finally, in the G‬‭2‬ ‭phase, the cell undergoes the final preparations for‬
‭meiosis.‬

‭Prophase I‬
‭ arly in‬‭prophase I‬‭, before the chromosomes can be‬‭seen clearly with a microscope, the‬
E
‭homologous chromosomes of each pair are moved closer together so that they can join.‬
‭Recall that in mitosis, homologous chromosomes do not pair together. A lattice of proteins‬
‭between the homologous chromosomes first forms at specific locations and then spreads‬
‭o utward to cover the entire length of the chromosomes. The tight pairing of the‬
‭homologous chromosomes is called‬‭synapsis‬‭(‬‭Figure‬‭13.12‬‭). These pairs are called‬
‭tetrads‬‭because each pair of homologous chromosomes'‬‭four chromatids are now visible.‬
‭In humans, even though the X and Y sex chromosomes are not completely homologous‬
‭(that is, most of their genes differ), a small region of homology allows the X and Y‬
‭chromosomes to pair up during prophase I.‬

‭ igure 13.12‬ ‭Early in prophase I, homologous chromosomes‬‭come together (synapse).‬

F
‭The chromosomes are bound tightly together and perfectly aligned by a protein lattice‬
‭called a synaptonemal complex. (credit:‬‭OpenStax‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

I‭ n synapsis, the genes on the chromatids of the homologous chromosomes are aligned‬
‭precisely with each other. This arrangement supports the exchange of chromosomal‬
‭segments between homologous, non-sister chromatids—a process called‬‭c rossing over‬
‭(‬‭Figure 13.13‬‭). If crossing over occurs between non-sister‬‭chromatids with different‬
‭alleles of a gene, then‬‭genetic recombination‬‭can‬‭o ccur. The recombinant chromatid has‬

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‭Chapter 13: Cell Cycle and Cell Division‬

a‭ combination of the alleles of maternal and paternal genes that did not exist before the‬
‭crossover.‬

‭ igure 13.13‬ ‭The result is an exchange of genetic‬‭material between homologous‬

F
‭chromosomes. Assume there are three genes on each homolog, and the first one has‬
‭dominant alleles at all three (A, B, and C), while the second one has recessive alleles at all‬
‭three (a, b, and c). Crossover occurs between non-sister chromatids of homologous‬
‭chromosomes‬‭.‬‭In the figure above, crossing over occurred‬‭o ne time between the B/b and‬
‭C/c genes. Because there were allelic differences between the homologs, crossing over‬
‭produced two recombinant chromatids with allelic content ABc and abC. The two‬
‭non-sister chromatids that didn’t participate in crossing over are classified as‬
‭nonrecombinant and have the same allelic collection for the three genes as the parental‬
‭homologs (ABC and abc). (credit:‬‭OpenStax‬‭). A link‬‭to a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭ rossover events can occur almost anywhere along the length of the synapsed‬
C
‭chromosomes. Therefore, different cells undergoing meiosis will produce separate‬

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‭Chapter 13: Cell Cycle and Cell Division‬

r‭ ecombinant chromatids with varying combinations of maternal and parental genes.‬

‭Multiple crossovers in an arm of the chromosome have the same effect, exchanging‬
‭segments of DNA to produce genetically recombined chromosomes.‬

‭Metaphase I‬
J‭ ust before‬‭metaphase I‬‭, the spindle fiber microtubules attach to the kinetochore‬
‭proteins at the centromeres. During this phase, the homologous chromosomes are‬
‭arranged at the‬‭metaphase plate‬‭—roughly in the cell's‬‭midline, with the kinetochores‬
‭facing opposite poles. The homologous pairs orient themselves randomly at the equator.‬
‭For example, if the two homologous members of chromosome 1 are labeled‬‭a‬‭and‬‭b‬‭, then‬
‭the chromosomes could line up a-b or b-a. This is important in determining the genes‬
‭carried by a gamete, as each will only receive one of the two homologous chromosomes.‬
‭Recall that after crossing over takes place, homologous chromosomes are not identical as‬
‭long as the two homologs contain different alleles for the same genes. They contain slight‬
‭differences in their genetic information, causing each gamete to have a unique genetic‬
‭makeup.‬
‭ he randomness in the alignment of recombined chromosomes at the metaphase plate,‬
T
‭coupled with the crossing-over events between non-sister chromatids, is responsible for‬
‭much of the genetic variation in the offspring. To clarify this further, remember that the‬
‭homologous chromosomes of a sexually reproducing organism are initially inherited as‬
‭two separate sets, one from each parent. Using humans as an example, one set of 23‬
‭chromosomes is present in the egg donated by the mother. The father provides the other‬
‭set of 23 chromosomes in the sperm that fertilizes the egg. Every cell of the multicellular‬
‭o ffspring has copies of the original two sets of homologous chromosomes. In prophase I‬
‭o f meiosis, the homologous chromosomes form the tetrads. In metaphase I, these pairs‬
‭line up at the midway point between the two poles of the cell to create the metaphase‬
‭plate. Because there is an equal chance that a microtubule fiber will encounter a‬
‭maternally or paternally inherited chromosome, the arrangement of the tetrads at the‬
‭metaphase plate is random. Thus, any maternally inherited chromosome may face either‬
‭pole. Likewise, any paternally inherited chromosome may also face either pole. The‬
‭o rientation of each tetrad is independent of the orientation of the other 22 tetrads.‬
‭ his event—the random or‬‭independent assortment‬‭o f‬‭homologous chromosomes at‬
T
‭the metaphase plate—is the second mechanism that introduces variation into the gametes‬
‭(Crossing over is the first mechanism). In each cell that undergoes meiosis, the‬
‭arrangement of the tetrads is different. Variations depend on the number of‬
‭chromosomes making up a set. There are two possibilities for orientation at the‬
‭metaphase plate; the possible number of alignments equals 2‬‭n‬ ‭in a diploid cell, where‬‭n‬‭is‬
‭the number of chromosomes per haploid set (‬‭Figure‬‭13.14‬‭). Humans have 23‬

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c‭ hromosome pairs, resulting in over eight million (2‬‭23‬‭) possible genetically distinct‬
‭gametes from the random alignment of chromosomes at the metaphase plate. This‬
‭number does not include the variability previously produced by crossing over between‬
‭the non-sister chromatids. Given these two mechanisms, it is improbable that any two‬
‭haploid cells resulting from meiosis will have the same genetic‬
‭composition.‬

‭ igure 13.14‬ ‭Random, independent‬‭assortment during metaphase I can be‬

F
‭demonstrated by considering a cell with a set of two chromosomes (2n = 4;‬‭n‬‭= 2). In this‬
‭case, metaphase I has two possible arrangements at the equator. The total possible‬
‭number of different gametes is 2‬‭n‬‭, where‬‭n‬‭equals the number of chromosomes in a set. In‬
‭this example, there are four possible genetic combinations for the gametes. With‬‭n‬‭= 23 in‬
‭human cells, there are over eight million possible paternal and maternal chromosome‬
‭combinations. (credit:‬‭OpenStax‬‭). A link to a video explanation of this figure is available at‬
‭Biology411.com‬‭.‬

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‭Chapter 13: Cell Cycle and Cell Division‬

‭ o summarize, meiosis I creates genetically diverse gametes in two ways. First, during‬
T
‭prophase I, crossover events between the non-sister chromatids of each homologous pair‬
‭o f chromosomes generate recombinant chromatids with new combinations of alleles of‬
‭maternal and paternal genes. Second, the random assortment of tetrads on the metaphase‬
‭plate produces unique combinations of maternal and paternal chromosomes that will‬
‭make their way into the gametes.‬

‭Anaphase I‬
I‭ n‬‭anaphase I‬‭, the microtubules pull the duplicated‬‭homologous chromosomes apart. The‬
‭sister chromatids of each duplicated homolog remain tightly bound together at the‬
‭centromere.‬

‭Telophase I and Cytokinesis‬

I‭ n‬‭telophase I‬‭, the separated homologous chromosomes‬‭arrive at opposite poles. During‬
‭cytokinesis, the cleavage furrow deepens, as was discussed in mitosis, and the cytoplasm is‬
‭divided between the two daughter cells. Two haploid cells result from the first meiotic‬
‭division of a diploid cell. The cells are haploid because, at each pole, there is just one of‬
‭each pair of homologous chromosomes. Therefore, only one complete set of‬
‭chromosomes is present. This is why the cells are considered haploid—there is only one‬
‭chromosome set, even though each chromosome still consists of two sister chromatids‬‭.‬
‭Recall that sister chromatids are duplicates of one of the two homologous chromosomes‬
‭(except for changes that occurred during crossing over). In meiosis II, these two sister‬
‭chromatids will separate, creating four haploid daughter cells.‬

‭Meiosis II‬
‭ he DNA isn’t replicated before the cell begins meiosis II, so chromosomes are not‬
T
‭duplicated. The two cells produced during meiosis I go through the events of meiosis II in‬
‭synchrony. During‬‭meiosis II‬‭, the sister chromatids‬‭within the two daughter cells separate,‬
‭forming four new haploid gametes. The mechanics of meiosis II are similar to mitosis,‬
‭except that each dividing cell has only one set of homologous chromosomes, each with‬
‭two chromatids. Therefore, each cell has half the number of sister chromatids to separate‬
‭as a diploid cell undergoing mitosis.‬
‭ he events of meiosis II (prophase II, metaphase II, anaphase II, telophase II, and‬
T
‭cytokinesis) in which sister chromatids separate are similar to those discussed in mitosis.‬
‭Figure 13.15‬‭illustrates the differences between meiosis‬‭I (homologous chromosomes‬
‭separate) and meiosis II (sister chromatids separate).‬

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‭Chapter 13: Cell Cycle and Cell Division‬

‭Telophase II and Cytokinesis‬

I‭ n‬‭telophase II‬‭, the chromosomes arrive at opposite‬‭poles and begin to decondense.‬
‭Nuclear envelopes form around the chromosomes, and cytokinesis separates the two into‬
‭four unique haploid cells. The cells produced are genetically unique because of the‬
‭random assortment of paternal and maternal homologs and the recombination of‬
‭maternal and paternal segments of chromosomes (with their sets of genes) that occurs‬
‭during crossing over. The entire process of meiosis is outlined in‬‭Figure 13.16‬‭.‬

‭ igure 13.15‬ ‭The process of chromosome alignment‬‭differs between meiosis I and‬

F
‭meiosis II. In prometaphase I, microtubules attach to the fused kinetochores of‬
‭homologous chromosomes, and the homologous chromosomes are arranged at the‬
‭midline of the cell (the metaphase plate) in metaphase I. In anaphase I, the homologous‬
‭chromosomes separate. In prometaphase II, microtubules attach to the kinetochores of‬
‭sister chromatids, and the sister chromatids are arranged at the midpoint of the cells in‬
‭metaphase II. In anaphase II, the sister chromatids separate. (credit:‬‭OpenStax‬‭). A link to‬
‭a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

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‭ igure 13.16‬ ‭A summary of meiosis in an animal‬‭cell with a diploid number of four (2‬‭n‬
F
‭= 4). The original diploid germ cell proceeds through the stages of meiosis to form four‬
‭haploid (n = 2) daughter cells. (credit:‬‭OpenStax‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

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‭13.7 Comparing Meiosis and Mitosis‬

‭ itosis and meiosis are both forms of nuclear division in eukaryotic cells. They share‬
M
‭some similarities but exhibit significant differences, leading to different outcomes (‬‭Figure‬
‭13.17‬‭). Mitosis is a single nuclear division resulting‬‭in two nuclei usually partitioned into‬
‭two new cells. The nuclei resulting from a mitotic division are genetically identical to the‬
‭o riginal nucleus. They have the same number of sets of chromosomes: one set in the case‬
‭o f haploid cells and two sets in the case of diploid cells.‬
I‭ n contrast, meiosis consists of two nuclear divisions, resulting in four nuclei that are‬
‭usually partitioned into four new, genetically distinct cells. The four nuclei produced‬
‭during meiosis are not genetically identical and contain only one chromosome set, half the‬
‭number of chromosome sets in the original diploid cell.‬
‭ he main differences between mitosis and meiosis occur during meiosis I, a very different‬
T
‭nuclear division from mitosis. In meiosis I, the homologous chromosome pairs physically‬
‭meet and are bound together with the synaptonemal complex. Following this, the‬
‭chromosomes undergo crossover between non-sister chromatids. In the end, the‬
‭chromosomes line up along the metaphase plate as tetrads—with kinetochore fibers from‬
‭o pposite spindle poles attached to each kinetochore of a homolog to form a tetrad. All of‬
‭these events occur only in meiosis I.‬
‭ hen the tetrad is broken up with the homologs moving to one pole or another, the‬
W
‭ploidy level—the number of sets of chromosomes in each future nucleus—has been‬
‭reduced from two to one. For this reason, as mentioned previously, meiosis I is referred to‬
‭as a reductional division. There is no such reduction in ploidy level during mitosis.‬
‭ eiosis II is analogous to a mitotic division. In this case, the duplicated chromosomes‬
M
‭(only one set) line up on the metaphase plate with divided kinetochores attached to‬
‭kinetochore fibers from opposite poles. During anaphase II, as in mitotic anaphase, the‬
‭kinetochores divide. One sister chromatid—now referred to as a chromosome—is pulled‬
‭to one pole, and the other sister chromatid is pulled to the other pole. If it were not for the‬
‭fact that there had been a crossover, the two products of each meiosis II division would be‬
‭identical (as in mitosis). Instead, they are different because there has always been at least‬
‭o ne crossover per chromosome.‬
‭ eiosis II doesn’t further reduce the number of copies of the genome in its daughter cells,‬
M
‭as only the sister chromatids are separated. Therefore, as stated previously, it is referred‬
‭to as equational division.‬

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‭ igure 13.17‬ ‭Meiosis and mitosis are preceded by‬‭o ne DNA replication cycle; however,‬
F
‭meiosis includes two nuclear divisions. The four daughter cells resulting from meiosis are‬
‭haploid (n) and genetically distinct due to crossing over and independent assortment. The‬
‭daughter cells resulting from mitosis are diploid (2n) and genetically identical to the‬
‭parent cell. (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

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‭13.8 Disorders Associated with Cell Division‬

‭Cancer‬

‭ ancer‬‭is a collective name for many diseases caused‬‭by a common mechanism:‬

C
‭uncontrolled cell division. Despite the redundancy and overlapping levels of cell-cycle‬
‭control, errors occur. One of the critical processes monitored by the cell-cycle checkpoint‬
‭surveillance mechanism is the proper replication of DNA during the S phase. Even when‬
‭all cell-cycle controls are fully functional, a small percentage of replication errors‬
‭(mutations) will be passed on to the daughter cells. If one of these changes to the DNA‬
‭nucleotide sequence occurs within a gene, a gene mutation results.‬

‭ ll cancers begin when a gene mutation gives rise to a faulty protein that participates in‬
A
‭cell reproduction. The change in the cell that results from the malformed protein may be‬
‭minor. Even minor mistakes, however, may allow subsequent mistakes to occur more‬
‭readily. Over and over, minor, uncorrected errors are passed from parent cells to daughter‬
‭cells and accumulate as each generation of cells produces more non-functional proteins‬
‭from uncorrected DNA damage. Eventually, the pace of the cell cycle speeds up as the‬
‭effectiveness of the control and repair mechanisms decreases. Uncontrolled growth of the‬
‭mutated cells outpaces the growth of normal cells in the area, and a tumor can result.‬
‭ or additional information about how cancerous cells differ from normal cells, click the‬
F
‭link below or scan the QR code to watch the TED-Ed video by‬‭George Zaidan titled “How‬
‭do cancer cells behave differently from healthy ones?”.‬

How do cancer cells behave differently from healthy ones? - Geor…

‭Proto-oncogenes‬

‭ he genes that code for the positive cell-cycle regulators are called proto-oncogenes.‬
T
‭Proto-oncogenes‬‭are normal genes that, when mutated,‬‭become oncogenes—genes that‬
‭cause a cell to become cancerous. Consider what might happen to the cell cycle in a cell‬
‭with a recently acquired oncogene. In most instances, altering the DNA sequence will‬
‭result in a less functional (or non-functional) protein. The result is detrimental to the cell‬
‭and will likely prevent the cell from completing the cell cycle; however, the organism is not‬

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‭ armed because the mutation will not be passed on. If a cell cannot reproduce, the‬
h
‭mutation is not propagated, and the damage is minimal. Occasionally, however, a gene‬
‭mutation causes a change that increases the activity of a positive regulator. For example, a‬
‭mutation that allows Cdk, a protein involved in cell-cycle regulation, to be activated before‬
‭it should be could push the cell cycle past a checkpoint before all of the required‬
‭conditions are met. If the resulting daughter cells are too damaged to undertake further‬
‭cell divisions, the mutation would not be propagated, and no harm comes to the organism.‬
‭However, suppose the atypical daughter cells can divide further. In that case, the‬
‭subsequent generation of cells will likely accumulate even more mutations, some possibly‬
‭in additional genes that regulate the cell cycle.‬

‭ he Cdk example is only one of many genes that are considered proto-oncogenes. In‬
T
‭addition to the cell-cycle regulatory proteins, any protein that influences the cycle can be‬
‭altered to override cell-cycle checkpoints. Once a proto-oncogene has been changed to‬
‭increase the cell cycle rate, it is called an‬‭oncogene‬‭.‬

‭Tumor Suppressor Genes‬

‭ ike proto-oncogenes, many of the negative cell-cycle regulatory proteins were discovered‬
L
‭in cells that had become cancerous.‬‭Tumor suppressor‬‭genes‬‭code for the negative‬
‭regulator proteins, the type of regulator that—when activated—can prevent the cell from‬
‭undergoing uncontrolled division. The collective function of the best-understood tumor‬
‭suppressor gene proteins (e.g., retinoblastoma protein (RB1), p53, BRCA1, BRCA2, and‬
‭p2) is to put up a roadblock to cell-cycle progress until certain events are completed. A cell‬
‭that carries a mutated form of a negative regulator might be unable to halt the cell cycle if‬
‭there is a problem.‬

‭ or additional information about the BRCA1 gene and its association with cancer, click the‬
F
‭link below or scan the QR code to watch the TED-Ed video by Michael Windelspecht titled‬
‭“The cancer gene we all have.”‬

The cancer gene we all have - Michael Windelspecht

‭ utated p53 genes have been identified in over half of all human tumor cells. This‬
M
‭discovery is not surprising in light of the multiple roles that the p53 protein plays at the‬
‭G1 checkpoint. The p53 protein activates other genes whose products halt the cell cycle‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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(‭ allowing time for DNA repair), activates genes whose products participate in DNA repair,‬
‭o r activates genes that initiate cell death when DNA damage cannot be repaired. A‬
‭damaged p53 gene can result in the cell behaving as if there are no mutations (‬‭Figure‬
‭13.18‬‭). This allows cells to divide, propagating the‬‭mutation in daughter cells and allowing‬
‭the accumulation of new mutations. In addition, the damaged version of p53 found in‬
‭cancer cells cannot trigger cell death.‬

‭ igure 13.18‬ ‭(a) The role of p53 is to monitor DNA.‬‭If damage is detected, p53 triggers‬
F
‭repair mechanisms. If repairs are unsuccessful, p53 signals apoptosis. (b) A cell with an‬
‭abnormal p53 protein cannot repair damaged DNA and cannot signal apoptosis. Cells with‬
‭abnormal p53 can become cancerous. (credit: modification of work by Thierry Soussi;‬
‭OpenStax‬‭).‬ ‭A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ ost people understand that cancer or tumors are caused by abnormal cells that multiply‬
M
‭continuously, resulting in a tumor or leukemia (blood cancer). For additional information‬
‭about leukemia, click the link below or scan the QR code to watch the TED-Ed video by‬
‭Danilo Allegra and Dania Puggioni titled “What is leukemia?”.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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What is leukemia? - Danilo Allegra and Dania Puggioni

I‭ f the abnormal cells continue to divide unstopped, they can damage the tissues around‬
‭them, spread to other body parts (‬‭metastasis‬‭), and‬‭eventually result in death. For‬
‭additional information about how cancer can metastasize, click the link below or scan the‬
‭QR code to watch the TED-Ed video by Ivan Seah Yu Jun titled “How does cancer spread‬
‭through the body?”.‬

How does cancer spread through the body? - Ivan Seah Yu J…

I‭ n healthy cells, the tight regulation mechanisms of the cell cycle prevent this from‬
‭happening, while failures of cell cycle control can cause unwanted and excessive cell‬
‭division. Failures of control may be caused by inherited genetic abnormalities that‬
‭compromise the function of specific “stop” and “go” signals. Environmental insults that‬
‭damage DNA can also cause dysfunction in those signals. A combination of genetic‬
‭predisposition and environmental factors often leads to cancer.‬
‭ cell escaping its regular control system and becoming cancerous may frequently happen‬
A
‭throughout the body. Fortunately, specific immune system cells can recognize and destroy‬
‭cancerous cells. However, in some instances, cancerous cells remain undetected and‬
‭continue to increase. If the resulting tumor does not threaten surrounding tissues, it is‬
‭considered benign and can usually be easily removed. If capable of damage, the tumor is‬
‭considered malignant, and the patient is diagnosed with cancer.‬

‭ here are various traditional cancer treatments, such as chemotherapy and radiation‬
T
‭therapy. For additional information about chemotherapy, click the link below or scan the‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ R code to watch the TED-Ed video by Hyunsoo Joshua No titled “How does‬
Q
‭chemotherapy work?”‬

How does chemotherapy work? - Hyunsoo Joshua No

‭ relatively new cancer treatment involves manipulating (i.e., “biohacking”) the DNA of‬
A
‭bacterial cells. For additional information, click the link on the next page or scan the QR‬
‭code to watch the TED-Ed video by Tal Danino titled “Hacking bacteria to fight cancer.”‬

Hacking bacteria to fight cancer - Tal Danino

‭ arly detection of cancer is vital to the success of any treatment method. One novel‬
E
‭approach involves using a breathalyzer to detect chemicals associated with the disease.‬
‭For additional details, click the link below or scan the QR code to watch the TED-Ed video‬
‭by Julian Berschka titled “Could a breathalyzer detect cancer?”.‬

Could a breathalyzer detect cancer? - Julian Burschka

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ or more information about the challenges of treating cancer, click the link below or scan‬
F
‭the QR code to watch the TED-Ed video by Kyuson Yun titled “Why is it so hard to cure‬
‭cancer?”.‬

Why is it so hard to cure cancer? - Kyuson Yun

‭Chromosome Number Disorders‬

‭ f all chromosomal disorders, chromosome number abnormalities are the most obviously‬
O
‭identifiable from a karyogram/karyotype. Chromosome number disorders include‬
‭duplicating or losing entire chromosomes and changes in the number of complete sets of‬
‭chromosomes. They are caused by‬‭nondisjunction‬‭, which‬‭o ccurs when homologous‬
‭chromosome pairs or sister chromatids fail to separate during meiosis. Misaligned or‬
‭incomplete synapsis, or a spindle apparatus dysfunction that facilitates chromosome‬
‭migration, can cause nondisjunction. The risk of nondisjunction occurring increases with‬
‭the parents' age.‬

‭ ondisjunction can occur during either meiosis I or II, with differing results (‬‭Figure‬
N
‭13.19‬‭). If homologous chromosomes fail to separate‬‭during meiosis I, the result is two‬
‭gametes that lack that particular chromosome and two gametes with two chromosome‬
‭copies. Suppose sister chromatids fail to separate during meiosis II. In that case, the result‬
‭is one gamete that lacks that chromosome, two normal gametes with one chromosome‬
‭copy, and one gamete with two chromosome copies.‬

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‭ igure 13.19‬ ‭Nondisjunction occurs when homologous‬‭chromosomes or sister‬

F
‭chromatids fail to separate during meiosis, resulting in an abnormal chromosome number.‬
‭Nondisjunction may occur during meiosis I or meiosis II. (credit: Rao, A. and Tag, A.‬
‭Department of Biology, Texas A&M University;‬‭OpenStax‬‭).‬ ‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭ neuploidy‬
A
‭Scientists call individuals with the appropriate number of chromosomes for their species‬
‭euploid‬‭. In humans, euploidy corresponds to 22 pairs‬‭o f autosomes and one pair of sex‬
‭chromosomes. An individual with an error in chromosome number is described as‬
‭aneuploid‬‭, a term that includes‬‭monosomy‬‭(losing one‬‭chromosome) or‬‭trisomy‬
‭(gaining an extra chromosome). Monosomic human zygotes missing one copy of an‬
‭autosome fail to develop because they lack essential genes. This underscores the‬
‭importance of “gene dosage” in humans. Most autosomal trisomies also fail to develop to‬
‭birth; however, duplications of some smaller chromosomes (13, 15, 18, 21, or 22) can‬
‭result in offspring that survive for several weeks to many years. Trisomic individuals‬

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s‭ uffer from a different genetic imbalance: an excess gene dose. Individuals with an extra‬
‭chromosome may synthesize an abundance of the gene products which that chromosome‬
‭encodes. This extra dose (150 percent) of specific genes can lead to several functional‬
‭challenges and often precludes development. The most common trisomy among viable‬
‭births is chromosome 21, which corresponds to‬‭Down‬‭Syndrome‬‭. Short stature and‬
‭stunted digits, facial distinctions that include a broad skull and large tongue, and‬
‭significant developmental delays characterize individuals with this inherited disorder. We‬
‭can correlate the incidence of Down syndrome with maternal age. Older people are more‬
‭likely to become pregnant with fetuses carrying the‬‭trisomy 21‬‭genotype (‬‭Figure 13.20‬‭).‬

‭ igure 13.20‬ ‭The incidence of having a fetus with‬‭trisomy 21 increases with maternal age.‬
F
‭(credit:‬‭OpenStax‬‭)‬

‭Sex Chromosome Nondisjunction in Humans‬

‭ umans display dramatic deleterious effects with autosomal trisomies and monosomies.‬
H
‭Therefore, it may seem counterintuitive that human females and males can function‬
‭normally despite carrying different numbers of the X chromosome. Rather than a gain or‬
‭loss of autosomes, variations in the number of sex chromosomes occur with relatively‬
‭mild effects. In part, this happens because of the molecular process called‬‭X inactivation‬‭.‬

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‭ arly in development, when female mammalian embryos consist of just a few thousand‬
E
‭cells (relative to trillions in the newborn), one X chromosome in each cell inactivates by‬
‭tightly condensing into a quiescent (dormant) structure called a‬‭Barr body‬‭. The chance‬
‭that an X chromosome (maternally or paternally derived) inactivates in each cell is‬
‭random. Still, once this occurs, all cells derived from that one will have the same inactive X‬
‭chromosome or Barr body. Through this process, females compensate for their double‬
‭genetic dose of the X chromosome. In so-called “tortoiseshell” cats, we observe embryonic‬
‭X inactivation as color variegation (‬‭Figure 13.21‬‭).‬‭Females heterozygous for an X-linked‬
‭coat color gene will express one of two different coat colors over different body regions,‬
‭corresponding to whichever X chromosome inactivates in that region's embryonic cell‬
‭progenitor.‬

‭ igure 13.21‬‭In cats, the gene for coat color is located‬‭o n the X chromosome. In female‬
F
‭cats' embryonic development, one of the two X chromosomes randomly inactivates in each‬
‭cell, resulting in a tortoiseshell pattern if the cat has two different alleles for coat color.‬
‭Male cats, having only one X chromosome, never exhibit a tortoiseshell coat color. (credit:‬
‭Michael Bodega;‬‭OpenStax‬‭).‬ ‭A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

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‭ n individual carrying an abnormal number of X chromosomes will inactivate all but one X‬
A
‭chromosome in each of her cells. However, even inactivated X chromosomes continue to‬
‭express a few genes, and X chromosomes must reactivate for the proper maturation of‬
‭female ovaries. As a result, X-chromosomal abnormalities typically occur with mild‬
‭intellectual and physical disorders or disabilities and sterility. The individual will not‬
‭develop in utero if the X chromosome is absent.‬

S‭ cientists have identified and characterized several errors in sex chromosome numbers.‬
‭Individuals with three X chromosomes,‬‭triple-X‬‭, are‬‭phenotypically female but express‬
‭developmental delays and reduced fertility. The XXY genotype, corresponding to one type‬
‭o f‬‭Klinefelter syndrome‬‭, corresponds to phenotypically‬‭male individuals with small‬
‭testes, enlarged breasts, and reduced body hair. More complex types of Klinefelter‬
‭syndrome exist in which the individual has as many as five X chromosomes. In all types,‬
‭every X chromosome except one undergoes inactivation to compensate for the excess‬
‭genetic dosage. We see this as several Barr bodies in each cell nucleus.‬‭Turner syndrome‬‭,‬
‭characterized as an X0 genotype (i.e., only a single sex chromosome), corresponds to a‬
‭phenotypically female individual with short stature, webbed skin in the neck region,‬
‭hearing and cardiac impairments, and sterility.‬

‭13.9 Career Connection - Cytogenetic Technologist‬

‭ lthough we refer to Mendel as the “father of modern genetics,” he performed his‬

A
‭experiments with none of the tools today's geneticists routinely employ. One such‬
‭powerful cytological technique is karyotyping, a method in which geneticists can identify‬
‭traits characterized by chromosomal abnormalities from a single cell. A cytogenetic‬
‭technologist first collects a person’s cells (like white blood cells) from a blood sample or‬
‭o ther tissue to observe an individual's karyotype. In the laboratory, the technologist‬
‭stimulates the isolated cells to begin actively dividing and then applies the chemical‬
‭colchicine to cells to arrest condensed chromosomes in metaphase. The technologist then‬
‭induces swelling in the cells using a hypotonic solution, spreading the chromosomes, and‬
‭finally preserves the sample in a fixative and applies it to a slide.‬

‭ he technologist then stains chromosomes with one of several dyes to better visualize‬
T
‭each chromosome pair's distinct and reproducible banding patterns. Following staining,‬
‭the technologist views the chromosomes using bright-field microscopy. A common stain‬
‭choice is the Giemsa stain. Giemsa staining results in approximately 400–800 bands (of‬
‭tightly coiled DNA and condensed proteins) arranged along all 23 chromosome pairs. An‬
‭experienced technologist can identify each band. In addition to the banding patterns,‬
‭geneticists further identify chromosomes based on size and centromere location.‬

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‭ o obtain the classic depiction of the karyotype in which homologous chromosome pairs‬
T
‭align in numerical order from longest to shortest, the technologist obtains a digital image,‬
‭identifies each chromosome, and manually arranges the chromosomes into this pattern‬
‭(‬‭Figure 13.22‬‭).‬

‭ igure 13.22‬ ‭This is a human female karyotype. Notice‬‭that homologous chromosomes‬

F
‭are the same size and have the same centromere positions and banding patterns. A human‬
‭male would have an XY chromosome pair instead of the XX pair. (credit: Andreas Blozer et‬
‭al.;‬‭OpenStax‬‭).‬ ‭A link to a video explanation of‬‭this figure is available at‬‭Biology411.com‬‭.‬

‭ t its most basic, the karyotype may reveal genetic abnormalities in which an individual‬
A
‭has too many or too few chromosomes per cell. Examples of this are Down Syndrome,‬
‭identified by a third copy of chromosome 21, and Turner Syndrome, characterized by the‬
‭presence of only one X chromosome in females instead of the usual two. Technologists can‬
‭also identify large DNA deletions or insertions. For instance, geneticists can identify‬
‭Jacobsen Syndrome—which involves distinctive facial features and heart and bleeding‬
‭defects—by a deletion on chromosome 11. Finally, the karyotype can pinpoint‬
‭translocations, which occur when a segment of genetic material breaks from one‬
‭chromosome and reattaches to another one or to a different part of the same‬
‭chromosome. Translocations are implicated in certain cancers, including chronic‬
‭myelogenous leukemia.‬

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P
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‭ uring Mendel’s lifetime, inheritance was an abstract concept that one could only infer by‬
D
‭performing crosses and observing offspring's traits. By studying a karyotype, cytogenetic‬
‭technologists can visualize an individual's chromosomal composition to confirm or predict‬
‭genetic abnormalities in offspring, even before birth.‬

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‭ igure 14.1‬ ‭A 10 to 12-week human fetus. (credit:‬‭Fetus 10-12 Weeks‬‭; Lunar Caustic;‬
F
‭Flickr;‬‭CC BY 2.0‬‭)‬

‭14.1 Introduction‬
‭ eproduction is necessary for the survival of a species. In humans, which reproduce‬
R
‭sexually, the genetic material of two individuals combines via the fusion of the gametes to‬
‭produce the fertilized egg, also called a zygote. This single cell divides via mitosis to‬
‭produce other cells, forming the embryo. Cell division continues, and the cells become‬
‭o rganized into tissues, organs, and organ systems that function in an integrated manner.‬
‭The human embryo becomes the fetus after approximately ten weeks of gestation (‬‭Figure‬
‭14.1‬‭); the processes of gestational growth and development‬‭then continue until birth.‬
‭In this chapter, you will learn about human reproductive anatomy, physiology, and the‬
a‭ ssociated events that ultimately gave rise to you as a newborn! Other related topics, such‬
‭as reproductive technologies, contraceptives, and eugenics, are included.‬

‭14.2 Male Reproductive Anatomy‬

‭ ur discussion of male reproductive anatomy begins with the male gonad, or‬‭testis‬
O
‭(plural: testes;‬‭Figure 14.2‬‭). A testis is approximately‬‭1.5 by 1.0 inches, and it functions to‬

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‭ roduce both gametes (sperm) and hormones. During the seventh month of male fetal‬
p
‭development, the testes move through the muscle of the abdominal cavity and descend‬
‭into a sac-like structure called the‬‭scrotum‬‭. This event is called “descent of the testes,”‬‭and‬
‭if it doesn’t happen to one or both testes, the resulting condition is “‬‭c ryptorchidism‬‭”‬
‭(unilateral or bilateral, respectively). A male affected with bilateral cryptorchidism will‬
‭experience a more significant fertility reduction than a male with unilateral‬
‭cryptorchidism.‬

‭ igure 14.2‬ ‭A depiction of male reproductive anatomy.‬ ‭Organs of other systems (e.g.,‬
F
‭rectum and bladder) are provided as reference points. (credit:‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ aving the testes outside the body cavity permits them to remain at a slightly cooler‬
H
‭temperature, which is necessary for the survival of sperm cells. The scrotum also includes‬
‭a passage for blood vessels, nerves, and muscles associated with the testes' function.‬
‭ he‬‭epididymis‬‭is a highly coiled tube attached to‬‭each testis on one end and the vas‬
T
‭deferens on the other. Sperm cells produced in the testis move through the epididymis for‬
‭several weeks, where they undergo a maturation process that results in the ability to move‬

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(‭ mobility) and fertilize an egg. During ejaculation, sperm cells stored in the last part of the‬
‭epididymis move into the‬‭vas deferens‬‭, a tube that transports them‬‭back into the‬
‭abdominal cavity towards the‬‭urethra.‬‭Glandular tissue in the vas deferens contributes‬‭a‬
‭relatively small percentage (10%) of the fluid secretions that combine with the sperm to‬
‭produce‬‭semen‬‭.‬
‭ ecause the vas deferens are physically accessible within the scrotum, surgical‬
B
‭sterilization to interrupt sperm delivery can be performed by cutting and sealing a small‬
‭section of them. This procedure, called a‬‭vasectomy‬‭,‬ ‭is an effective form of male birth‬
‭control. Although it may be possible to reverse a vasectomy, the procedure is considered‬
‭permanent. Therefore, men are advised to undergo it only if they no longer want to have‬
‭children. A male with a vasectomy will still ejaculate, but if the procedure is successful, no‬
‭sperm cells will be present in the semen.‬
‭ he bulk of the fluids in semen comes from the three types of accessory glands: the‬
T
‭seminal vesicles, the prostate gland, and the bulbourethral glands (‬‭Figure 14.3‬‭). The‬
‭seminal vesicles‬‭are a pair of glands that lie along‬‭the back of the urinary bladder,‬
‭contributing approximately 60% of the semen volume. These glands make an alkaline‬
‭solution necessary because sperm are only motile in such an environment. Also, a basic‬
‭pH (greater than 7) is essential to neutralize the acidity of the vaginal environment.‬
‭Seminal vesicle secretions also contain large amounts of fructose (a monosaccharide‬
‭sugar) used by the sperm mitochondria to generate ATP necessary for movement through‬
‭the female reproductive tract.‬
‭ he walnut-shaped‬‭prostate gland‬‭surrounds the urethra‬‭and connects to the urinary‬
T
‭bladder. It has a series of short ducts that directly connect to the urethra. The gland is a‬
‭mixture of smooth muscle and glandular tissue. The muscle provides much of the force‬
‭needed for ejaculation. The glandular tissue makes a thin, milky fluid that contains buffers‬
‭and enzymes such as prostate-specific antigen (PSA). Prostate gland secretions account‬
‭for about 30% of the bulk of semen.‬
I‭ n mid to late 20-year-old males, the prostate gland enlarges gradually. This enlargement‬
‭does not usually cause problems; however, abnormal growth of the prostate can cause the‬
‭narrowing of the urethra because it passes through the middle of the gland. This‬
‭enlargement can lead to several lower urinary tract symptoms, such as a frequent and‬
‭intense urge to urinate, a weak urine stream, and a sensation that the bladder has not‬
‭emptied. By age 60, approximately 40% of men have some enlargement. By age 80, the‬
‭number of affected individuals increases to as many as 80%. Treatments for this condition‬
‭attempt to relieve the pressure on the urethra so that urine can flow more normally. Mild‬
‭to moderate symptoms are treated with medication, whereas severe enlargement of the‬
‭prostate is treated by surgery in which a portion of the prostate tissue is removed.‬

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P
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‭ igure 14.3‬ ‭Another depiction of male reproductive‬‭anatomy. (credit:‬‭Sperm release‬

F
‭pathway.svg‬‭; K.D. Schroeder (‬‭KDS4444‬‭);‬‭CC BY-SA 4.0‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ nother common disorder involving the prostate is‬‭prostate cancer‬‭. According to the‬
A
‭Centers for Disease Control and Prevention (CDC), prostate cancer is men's second most‬
‭common cancer. However, some forms of prostate cancer grow very slowly and thus may‬
‭never require treatment. In contrast, aggressive forms of prostate cancer involve‬
‭metastasis to vulnerable organs like the lungs and brain.‬
‭ octors check the prostate gland for enlargement and cancerous growths via a digital‬
D
‭rectal exam (‬‭Figure 14.4‬‭). During this procedure,‬‭the clinician accesses the gland via the‬
‭rectum and feels for changes in size, tenderness, and lumps.‬
‭ he‬‭bulbourethral gland‬‭releases its secretion before‬‭releasing the bulk of the semen. It‬
T
‭lubricates and cleans the end of the urethra before sperm cells move through. These‬
‭secretions account for a relatively minor percentage of fluids in the total ejaculate. It is‬
‭important to note that bulbourethral fluid can pick up sperm already present in the‬
‭urethra, and therefore, it may be able to cause pregnancy.‬
‭ he‬‭penis‬‭is an organ that transports urine from the bladder out of the body‬‭and‬
T
‭functions as a copulatory organ during intercourse. The penis contains three regions of‬
‭erectile tissue‬‭running through the organ's length. During sexual arousal, blood‬‭vessels in‬
‭this tissue will dilate (open), allowing increased blood flow, and the penis will become‬

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e‭ rect and hard in preparation for intercourse. During intercourse, the smooth muscle‬
‭sphincters at the opening of the bladder close and prevent urine from entering the penis.‬
‭After ejaculation, the blood drains from the erectile tissue, the penis becomes flaccid, the‬
‭sphincters relax, and urination can resume.‬

‭ igure 14.4‬ ‭A digital rectal examination to inspect‬‭the prostate gland for abnormalities.‬
F
‭(credit:‬‭Digital rectal exam‬‭;‬‭NCI‬‭;‬‭CC0 1.0‬‭)‬

‭ rectile dysfunction‬‭(‬‭ED‬‭) is when a man has difficulty‬‭initiating or maintaining an‬

E
‭erection. The combined prevalence of minimal, moderate, and complete ED is‬
‭approximately 40% in men at age 40 and reaches nearly 70% by 70 years of age. In‬
‭addition to aging, the risk of ED increases with diabetes, vascular disease, psychiatric‬
‭disorders, prostate disorders, the use of some drugs such as certain antidepressants, and‬
‭problems with the testes resulting in low testosterone concentrations. These physical and‬
‭emotional conditions can lead to interruptions in the vasodilation pathway and result in‬
‭an inability to achieve an erection.‬

‭ he release of‬‭nitric oxide‬‭(NO) causes relaxation of the smooth muscles surrounding‬‭the‬

T
‭arteries in the penis, leading to the vasodilation (increased diameter) necessary to achieve‬
‭an erection. To reverse the process of vasodilation, an enzyme called phosphodiesterase‬
‭(PDE) degrades a vital component of the NO signaling pathway. There are several different‬

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f‭ orms of this enzyme, and PDE-5 is the type found in the tissues of the penis. Scientists‬
‭discovered that inhibiting PDE-5, which is the mechanism of action of ED medications such‬
‭as Viagra, increases blood flow and permits vasodilation of the blood vessels in the‬
‭erectile tissue of the penis.‬

‭14.3 Spermatogenesis‬
‭ permatogenesis‬‭is the process of sperm production‬‭in the testes. This process begins at‬
S
‭puberty, after which time sperm is produced constantly throughout a man’s life.‬
‭Spermatogenesis occurs in structures called‬‭seminiferous tubules‬‭, which are coiled‬
‭inside the testes, as illustrated in‬‭Figure 14.5a‬‭.‬ ‭The walls of the seminiferous tubules are‬
‭made up of the developing sperm cells, with the least developed sperm (spermatogonia)‬
‭at the outer edges of the tubule and the fully developed sperm (spermatozoa) in the‬
‭lumen (‬‭Figure 14.5b‬‭). As the cells move from the outer‬‭edge towards the lumen of the‬
‭tubules, which takes approximately 60 days, they undergo the process of meiosis to‬
‭become haploid gametes.‬
‭ ithin the seminiferous tubules, the various forms of sperm cells are mixed with‬
W
‭“nursemaid” cells called‬‭Sertoli cells‬‭. Other cells‬‭located in spaces between the tubules‬
‭are called the‬‭interstitial‬‭(or Leydig)‬‭cells‬‭, which‬‭produce high levels of testosterone‬
‭o nce the male reaches puberty. When the sperm has developed flagella (tails) and is‬
‭nearly mature, it leaves the testicles and enters the epididymis to complete maturation.‬
S‭ perm cells are smaller than most cells in the body; in fact, the volume of a sperm cell is‬
‭85,000 times less than that of the female egg cell. Approximately 100 to 300 million‬
‭sperm are produced each day. As is true for most cells in the body, the structure of sperm‬
‭cells speaks to their function. Sperm have a distinctive head, mid-piece, and tail region‬
‭(‬‭Figure 14.6‬‭). The head of the sperm contains the‬‭genetic material, which is a haploid‬
‭nucleus with minimal cytoplasm. A structure called the‬‭acrosome‬‭covers most of the head‬
‭o f the sperm cell as a “cap” filled with enzymes that can digest the protective coverings‬
‭surrounding the egg to help the sperm penetrate and fertilize the egg.‬
‭ n average ejaculate contains from two to five milliliters of fluid with approximately‬
A
‭50–120 million sperm per milliliter. Sperm cells make up only five percent of the final‬
‭volume of semen; the majority of the remaining volume is contributed by the accessory‬
‭sex glands (seminal vesicles, prostate, and bulbourethral). Tightly packed‬‭mitochondria‬
‭fill the mid-piece of the sperm. ATP produced by these mitochondria will power the‬
‭flagellum‬‭, which extends from the neck and the mid-piece through the sperm's tail,‬
‭enabling it to move the entire sperm cell.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭(a)‬ ‭(b)‬

‭ igure 14.5‬ ‭(a) Diagram of a cross-section of the‬‭testis showing the seminiferous‬

F
‭(sperm-producing) tubules. (b) A stained section of the testis shows a seminiferous‬
‭tubule. The white portion in the center of the tubule is the lumen, which contains the‬
‭flagella (tails) attached to the small spherical heads of the spermatozoa. (credit: (a)‬‭The‬
‭testis & a magnified seminiferous tubule.jpg‬‭;‬ ‭Sunshineconnelly‬‭;‬‭CC BY 3.0‬ ‭(b)‬
‭Photomicrograph of Rabbit Testicular Tissue‬‭; Majid‬‭Shokoohi;‬‭CC BY 4.0‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ or additional information about the physical and chemical challenges sperm encounter‬
F
‭when moving through the female reproductive tract, click the link below or scan the QR‬
‭code to watch the TED-Ed video by Aatish Bhatia titled “Human sperm vs. the sperm‬
‭whale.”‬

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Human sperm vs. the sperm whale - Aatish Bhatia

‭ igure 14.6‬ ‭Human sperm cells, visualized using‬‭scanning electron microscopy, have a‬
F
‭flagellum, neck, and head. (credit b: modification of work by Mariana Ruiz Villareal;‬
‭scale-bar data from Matt Russell;‬‭OpenStax‬‭). A link‬‭to a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭14.4 Male Reproductive Hormones‬

‭ t the onset of puberty, the hypothalamus releases‬‭gonadotropin-releasing hormone‬
A
‭(GnRH),‬‭which causes the release of‬‭follicle-stimulating‬‭hormone (FSH)‬‭and‬
‭luteinizing hormone (LH)‬‭from the‬‭anterior pituitary‬‭gland‬‭for the first time. FSH‬
‭enters the testes and stimulates the Sertoli cells to begin facilitating spermatogenesis, as‬
‭illustrated in‬‭Figure 14.7‬‭. LH also enters the testes and stimulates the interstitial‬‭(Leydig)‬
‭cells‬‭to make and release testosterone into the testes and the blood.‬
‭ estosterone‬‭stimulates spermatogenesis and is also responsible for‬‭the secondary sexual‬
T
‭characteristics that develop in the male during adolescence and stimulating‬

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s‭ permatogenesis. These secondary sex characteristics include a deepening of the voice,‬

‭the growth of facial and pubic hair, and initiation of the sex drive.‬
‭ ‬‭negative feedback control system‬‭o ccurs with rising testosterone levels acting on the‬
A
‭hypothalamus and anterior pituitary to inhibit the release of GnRH, FSH, and LH. When‬
‭the sperm count is too high, the Sertoli cells produce and secrete the hormone inhibin.‬
‭This hormone inhibits the release of GnRH and FSH, which will cause spermatogenesis to‬
‭slow down. If the sperm count drops below threshold concentrations, the Sertoli cells‬
‭cease releasing inhibin, which removes the blocked release of the other hormones, and‬
‭the sperm count increases.‬

‭ igure 14.7‬ ‭Hormones control sperm production in‬‭a negative feedback system. (credit:‬
F
‭OpenStax‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭14.5 Female Reproductive Anatomy‬

‭ he female reproductive system also functions to produce gametes and reproductive‬
T
‭hormones; however, it also has the additional task of supporting the developing fetus and‬
‭delivering it. Unlike its male counterpart, the female reproductive system is primarily‬
‭inside the pelvic cavity (‬‭Figure 14.8‬‭). Recall that‬‭the‬‭ovaries‬‭are the female gonads, and‬
‭the gamete they produce is called an‬‭oocyte‬‭(egg). We’ll discuss the production of oocytes,‬
‭called‬‭oogenesis‬‭, in detail shortly. First, let’s‬‭look at some of the structures of the female‬
‭reproductive system.‬
S‭ everal reproductive structures in the figures above are exterior to the female’s body.‬
‭These include the clitoris and the labia (majora and minora). The‬‭c litoris‬‭is a structure‬
‭with erectile tissue that contains many sensory nerves and is a source of stimulation‬
‭during intercourse. The‬‭labia majora‬‭are a pair of‬‭elongated folds of tissue derived from‬
‭the same tissue that produces the scrotum in a male. The‬‭labia minora‬‭are thin folds of‬
‭tissue located within the labia majora. These labia protect the openings to the vagina and‬
‭urethra.‬

‭ igure 14.8‬ ‭The reproductive structures of the human‬‭female, which are located inside‬
F
‭the pelvic cavity, are shown above. (credit a: modification of work by Gray's Anatomy;‬
‭credit b: modification of work by CDC;‬‭OpenStax‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

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I‭ nternal female reproductive structures include the ovaries, oviducts/fallopian tubes, the‬
‭uterus, and the vagina. The pair of ovaries is held in place in the abdominal cavity by a‬
‭ligament system, a type of connective tissue. Like the kidneys, ovaries consist of an inner‬
‭region called the medulla and an outer region called the cortex. The cortex primarily‬
‭consists of‬‭follicles‬‭, sac-like structures that contain‬‭o ocytes (‬‭Figure 14.9‬‭). Each ovary is‬
‭estimated to have 300,000 to 400,000 follicles at puberty. During each menstrual cycle, 10‬
‭to 12 follicles develop and prepare their oocytes for release. However, the majority of‬
‭these developing follicles and their oocytes will degrade. Typically, at‬‭ovulation‬‭, one large,‬
‭developing follicle, called a‬‭Graafian‬‭o r‬‭pre-ovulatory‬‭follicle‬‭, ruptures and releases its‬
‭o ocyte, as illustrated in‬‭Figure 14.9‬‭.‬

‭ igure 14.9‬ ‭Oocytes develop in (a) follicles located‬‭in the ovary. At the beginning of the‬
F
‭menstrual cycle, the follicle matures. At ovulation, the follicle ruptures, releasing the egg.‬
‭The follicle becomes a corpus luteum (CL), which eventually degenerates. The (b) follicle‬
‭in this light micrograph has an oocyte at its center. (credit: modification of work by NIH;‬
‭scale-bar data from Matt Russell;‬‭OpenStax‬‭). A link‬‭to a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

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‭ he‬‭oviducts‬‭, or‬‭fallopian tubes‬‭, extend from the ovaries to the‬‭uterus‬‭but are not in‬
T
‭direct contact with the ovaries. The ends of the oviducts adjacent to the ovaries flare out‬
‭into a trumpet-like structure and have a fringe of finger-like projections called‬‭fimbriae‬‭,‬
‭illustrated in‬‭Figure 14.8b‬‭. When an oocyte is released‬‭at ovulation, the fimbriae help‬
‭guide it into the oviduct, the passage to the uterus. The oviducts' walls are ciliated and‬
‭mostly made up of smooth muscle. The cilia beat toward the middle, and the smooth‬
‭muscle contracts in the same direction, moving the egg toward the uterus.‬‭Fertilization‬
‭usually takes place within the oviducts, and the developing embryo moves toward the‬
‭uterus, where‬‭implantation‬‭will occur. It usually‬‭takes the egg or embryo approximately a‬
‭week to travel through the oviduct to the uterus.‬
‭ he developing embryo may implant in other locations instead of the endometrium of the‬
T
‭uterus. If this happens, an‬‭ectopic pregnancy‬‭results (‬‭Figure 14.10‬‭). The most‬‭common‬
‭location for an ectopic pregnancy is in the fallopian tube, which results in a tubal‬
‭pregnancy. In cases of an ectopic pregnancy, the developing embryo won’t survive. Also,‬
‭the associated growth could cause the tube to rupture and, if untreated, result in‬
‭life-threatening internal bleeding in the mother.‬

‭ igure 14.10‬ ‭In a normal pregnancy, the embryo implants‬‭in the endometrium of the‬
F
‭uterus. In an ectopic pregnancy, implantation happens at another location (e.g., the‬
‭fallopian tube) that can’t sustain the pregnancy. Ectopic pregnancies are a serious medical‬
‭condition that can be potentially life-threatening if untreated. (credit:‬‭Ectopic‬
‭Pregnancy.png‬‭;‬‭BruceBlaus‬‭;‬‭CC BY-SA 4.0‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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S‭ terilization in women is called a‬‭tubal ligation‬‭; it is analogous to a vasectomy in males in‬

‭that the oviducts are cut and sealed. Although it may be possible to reverse a tubal‬
‭ligation, the procedure is considered permanent. Therefore, women are advised to‬
‭undergo it only if they no longer want to have children. A female with a tubal ligation will‬
‭still ovulate, but if the procedure is successful, neither the egg nor sperm can move‬
‭through the oviduct.‬
‭ he uterus is a structure about the size of a woman’s fist, and it supports the developing‬
T
‭embryo and fetus during gestation. The inner layer, the‬‭endometrium‬‭, is rich in blood‬
‭vessels and mucus glands. A portion of the inner layer sloughs off during the menstrual‬
‭cycle's menses (bleeding) portion. Then, it builds up again in preparation for possible‬
‭embryo implantation during the next cycle.‬
‭ ndometrial tissue is sometimes found outside the uterus in the pelvic or abdominal‬
E
‭cavity. These locations are problematic because the tissue responds to menstrual cycle‬
‭hormones and becomes inflamed and painful. This condition, known as‬‭endometriosis‬‭,‬
‭can negatively impact a female’s menstrual cycle and fertility if left untreated (‬‭Figure‬
‭14.11‬‭).‬
‭ he thickest portion of the uterus wall, the‬‭myometrium‬‭,‬‭is made of smooth muscle.‬
T
‭Contractions of this smooth muscle aid in moving sperm cells to the oviduct during‬
‭o rgasm and the baby through the birth canal during labor.‬

‭ igure 14.11‬ ‭Endometriosis is a condition in which‬‭tissue that normally lines the uterus‬
F
‭(endometrium) is found in other regions of the abdomen or pelvis. This tissue will‬
‭respond to hormones during the menstrual cycle and become inflamed and painful. It can‬
‭adversely affect a woman’s menstrual cycles and fertility if untreated. (credit:‬
‭Endometriosis-it.png‬‭;‬‭BruceBlaus‬‭;‬‭CC BY-SA 4.0‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ he lower end of the uterus that joins the‬‭vagina‬‭is called the‬‭cervix,‬‭and one of its‬
T
‭functions is to serve as the birth canal during delivery. The cervix must dilate (open) to‬
‭allow the fetus to pass from the uterus into the vagina. During birthing, the cervix will‬
‭dilate to a maximum diameter of 10 centimeters, which is necessary for a vaginal birth.‬
‭Also, ligaments and bones in the pelvic region relax, providing some additional “wiggle‬
‭room” to accommodate the baby's birth. You might wonder how the baby’s head fits‬
‭through the birth canal. Fortunately, the five bone plates that make up the baby’s skull‬
‭haven’t fused at birth, so they can shift and overlap to reduce the diameter.‬
‭ he‬‭vagina‬‭is a muscular tube, approximately four inches long, that serves several‬
T
‭functions. It allows the menstrual flow to leave the body, it is the receptacle for the penis‬
‭during intercourse, and it is the opening through which a baby will pass to be delivered.‬
‭ he vagina is home to a resident population of microorganisms that help to protect‬
T
‭against infection by pathogenic bacteria, yeast, or other organisms that enter the vagina.‬
‭In a healthy woman, the most predominant type of vaginal bacteria is‬‭Lactobacillus‬‭. This‬
‭family of beneficial bacteria secretes lactic acid and thus protects the vagina by‬
‭maintaining an acidic pH (below 4.5). Potential disease-causing organisms are less likely to‬
‭survive in these acidic conditions. In combination with other vaginal secretions, lactic acid‬
‭makes the vagina a self-cleansing organ. However, douching—or washing out the vagina‬
‭with fluid—can disrupt healthy microorganisms' normal balance and increase a woman’s‬
‭risk for infections and irritation.‬
‭ or additional information about yeast infections, click the link below or scan the QR code‬
F
‭to watch the TED-Ed video by Liesbeth Demuyser titled “What causes yeast infections, and‬
‭how do you get rid of them?”.‬

What causes yeast infections, and how do you get rid of them? …

‭14.6 Oogenesis‬
‭ ogenesis, which produces the female gamete (oocyte; egg), is illustrated in‬‭Figure 14.12‬‭.‬
O
‭This process occurs in the outermost layers of the ovaries. As with sperm production,‬
‭o ogenesis starts with a diploid (2n) germ cell called an oogonium (plural: oogonia); this‬
‭cell undergoes mitosis to increase in number.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 14.12‬ ‭The process of oogenesis results in‬‭haploid (n) gametes. Upon fertilization‬
F
‭by a haploid sperm cell, the fertilized egg (zygote) is diploid (2n). Polar bodies, which are‬
‭nonviable products of meiosis) are produced in meiosis I and II. For every oogonium that‬
‭begins meiosis, one via gamete is produced. The polar bodies degrade. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ he cell type that begins meiosis is called a‬‭primary oocyte‬‭. This cell will start the‬‭first‬
T
‭meiotic division and then temporarily stop its progress in prophase I (Chapter 13). At the‬
‭time of a female’s birth, all future eggs are in the prophase I stage of meiosis. The‬
‭initiation of ovulation—the release of an oocyte from the ovary—marks women's‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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t‭ ransition from puberty into reproductive maturity. From this point on, ovulation occurs‬
‭approximately once every 28 days throughout a woman's reproductive years. Just before‬
‭ovulation, a‬‭luteinizing hormone (LH) surge‬‭triggers the resumption of meiosis I in a‬
‭primary oocyte. This surge initiates the transition from primary to secondary oocyte.‬
‭However, as shown in‬‭Figure 14.12‬‭, this cell division does not result in two identical‬
‭haploid cells. The primary oocyte divides unequally, with most of the cellular material and‬
‭o rganelles going to one cell, called a‬‭secondary oocyte‬‭, and only one set of chromosomes‬
‭and a small amount of cytoplasm going to the other cell. This second cell is called a‬‭polar‬
‭body‬‭and usually dies. At this point, the cell stops meiosis at the metaphase II stage. This‬
‭secondary oocyte will be released at ovulation and travel toward the uterus through the‬
‭oviduct. If the secondary oocyte is fertilized, the cell continues through meiosis II,‬
‭producing a second polar body and a fertilized egg containing all 46 chromosomes of a‬
‭human being, half of them coming from the sperm. Only one viable gamete (secondary‬
‭o ocyte) is produced for every oogonium that begins meiosis.‬

‭14.7 The Ovarian Cycle, Menstrual Cycle, and Female Hormonal Regulation‬
‭ he‬‭ovarian cycle‬‭governs the preparation of endocrine‬‭tissues and the release of eggs.‬
T
‭In contrast, the‬‭menstrual cycle‬‭governs the preparation‬‭and maintenance of the uterine‬
‭lining (endometrium). These cycles coincide and are coordinated over a 22–32 day cycle,‬
‭with an average length of 28 days. The previously discussed hormones GnRH, FSH, and LH‬
‭play an essential role in the ovarian cycle. In contrast, hormones produced by the follicles‬
‭and corpus luteum (estrogen and progesterone) play a direct role in the ovarian cycle (via‬
‭negative feedback control) and the menstrual cycle, as discussed later in this section.‬
‭ uring the first half of the ovarian cycle, called the‬‭follicular phase,‬‭a small group of‬
D
‭follicles on the surface of the ovary grows in response to FSH and LH secreted from the‬
‭anterior pituitary gland (‬‭Figure 14.13; steps 2 and‬‭3)‬‭. As the follicles grow, they begin‬
‭releasing estrogens and a low level of progesterone, which will impact the development of‬
‭the uterine endometrium in preparation for the possible implantation of a fertilized egg.‬
‭Just before the middle of the menstrual cycle (approximately day 14), the high estrogen‬
‭level causes FSH and LH to rise rapidly and then fall. The spike in LH causes ovulation‬
‭(‬‭Figure 14.13; step 4‬‭). During this event, the most mature follicle, called the Graafian‬
‭follicle, ruptures and releases its egg. The fimbriae then help guide the ovulated egg into‬
‭the fallopian tube. The follicles that did not rupture degenerate, along with their eggs. The‬
‭level of estrogen decreases when the extra follicles degenerate.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ ollowing ovulation, the ovarian cycle enters the second phase, called the‬‭luteal phase‬‭.‬
F
‭During this time, the cells of the ruptured follicle undergo physical changes and produce a‬
‭structure called a‬‭corpus luteum (CL),‬‭as shown in‬‭Figure 14.13, step 5,‬‭and‬‭Figure‬
‭14.14a‬‭. The corpus luteum produces estrogen and progesterone. These hormones‬
‭facilitate the regrowth of the endometrium and inhibit, via negative feedback control, the‬
‭release of further FSH and LH in a similar manner to the effects of testosterone.‬
‭Therefore, while estrogen and progesterone levels are elevated, additional eggs and‬
‭follicles won’t develop. This outcome hopefully makes sense, as the female’s body is‬
‭“waiting” to determine if the previously ovulated egg was fertilized and implanted.‬
‭Estrogen is also an essential initiator of the development of female secondary sex‬
‭characteristics, such as breast development, hip changes, increased fat deposition, and‬
‭initiation of menstrual cycles.‬

‭ igure 14.13‬ ‭Diagram of the various stages of the‬‭ovarian cycle: 1) Primordial follicles,‬
F
‭2) Developing follicle, 3) Mature/Graafian follicle, 4) Ovulation, 5) Corpus luteum (CL),‬
‭and 6) Deterioration of the CL. (credit:‬ ‭Order of changes in ovary.svg‬‭;‬‭Shazz‬‭;‬‭CC BY-SA‬
‭4.0‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 14.14‬ ‭(a) A fully-developed corpus luteum‬‭(CL) on the ovary. Note the relatively‬
F
‭large size of the CL relative to the size of the entire ovary. (b) An example of degrading‬
‭CL, called a corpus albicans (CA), from a previous ovarian cycle. Several CA of earlier‬
‭cycles are visible in the ovary. (credit: Modified from‬‭Human ovary.jpg‬‭;‬‭Euthman‬‭;‬‭CC‬
‭BY-SA 4.0‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ fter ovulation, the oocyte’s trip through the fallopian tube takes about a week. If the egg‬
A
‭isn’t fertilized, implantation in the endometrium does not occur, and the lining is sloughed‬
‭o ff (menses). If a fertilized egg doesn’t implant into the uterus, the corpus luteum‬
‭degenerates (‬‭Figure 14.13; step 6; Figure 14.14b‬‭),‬‭and the levels of estrogen and‬
‭progesterone decrease. As these levels decrease, the endometrium begins to degenerate,‬
‭which initiates menses/menstruation and the start of the next menstrual cycle. The decline‬
‭in progesterone also allows the hypothalamus to send GnRH to the anterior pituitary,‬
‭releasing the block on FSH and LH secretion and starting the ovarian and uterine cycles‬
‭again.‬‭Figure 14.15‬‭visually compares the ovarian‬‭and uterine cycles, body temperature,‬
‭and relevant hormones (FSH, LH, estrogen/estradiol, and progesterone).‬
‭ or additional information about the ovarian and menstrual cycles, click the link below or‬
F
‭scan the QR code to watch the TED-Ed video by Emma Bryce titled “How menstruation‬
‭works.”‬

How menstruation works - Emma Bryce

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ or additional information about the history and evolution of menstruation, click the link‬
F
‭below or scan the QR code to watch the TED-Ed video titled “Why do women have‬
‭periods?”.‬

Why do women have periods?

‭ s women approach their mid-40s to mid-50s, their ovaries begin to lose their sensitivity‬
A
‭to FSH and LH. Menstrual periods become less frequent and finally cease, referred to as‬
‭menopause‬‭. There are still eggs and potential follicles‬‭o n the ovaries, but without the‬
‭stimulation of FSH and LH, they will not produce a viable egg to be released. The outcome‬
‭o f this situation is the inability to have children.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 14.15‬ ‭The ovarian and uterine cycles, with additional information‬‭about body‬
F
‭temperature (useful in monitoring when ovulation occurs) and relevant hormones (FSH,‬
‭LH, estrogen/estradiol, and progesterone). (credit:‬ ‭MenstrualCycle2.png‬‭;‬‭Lyrl‬‭;‬‭CC BY-SA‬
‭3.0‬‭). A link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ he side effects of menopause include hot flashes, heavy sweating (especially at night),‬
T
‭headaches, hair loss, muscle pain, vaginal dryness, insomnia, depression, weight gain, and‬
‭mood swings. Estrogen is involved in calcium metabolism, and without it, blood levels of‬
‭calcium decrease. Calcium is removed from the bone to replenish these levels, which may‬
‭decrease bone density and lead to‬‭osteoporosis‬‭(‬‭Figure 14.16‬‭). Supplementation of‬
‭estrogen as‬‭hormone replacement therapy‬‭(‬‭HRT‬‭) can‬‭prevent bone loss, but the‬
‭treatment can have adverse side effects. While doctors think that HRT gives some‬
‭protection from colon cancer, osteoporosis, heart disease, macular degeneration, and‬
‭possibly depression, its adverse side effects include increased risk of stroke or heart‬
‭attack, blood clots, breast cancer, ovarian cancer, endometrial cancer, and gallbladder‬
‭disease.‬

‭ igure 14.16‬ ‭A visual comparison of normal bone and bone affected by osteoporosis.‬
F
‭Two common locations of this disorder are in the vertebrae and hip. (credit:‬‭Osteoporosis‬
‭Locations.png‬‭;‬‭BruceBlaus‬‭;‬‭CC BY-SA 4.0‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭14.8 Fertilization and Twins‬

‭ ertilization is a numbers game. During ejaculation, hundreds of millions of sperm are‬
F
‭released into the vagina. Almost immediately, millions of these sperm are overcome by the‬
‭acidity of the vagina (approximately pH 3.8), and millions more may be blocked from‬
‭entering the uterus by thick cervical mucus. Thousands of those that do enter are‬
‭destroyed by phagocytic uterine white blood cells. Thus, the race into the fallopian tubes,‬
‭the most typical site for sperm to encounter the oocyte, is reduced to a few thousand‬
‭contenders. Their journey—thought to be facilitated by uterine contractions—usually‬
‭takes 30 minutes to 2 hours. If the sperm do not encounter an oocyte immediately, they‬
‭can survive in the uterine tubes for another 3–5 days. Thus, fertilization can still occur if‬
‭intercourse occurs a few days before ovulation. An oocyte can survive independently for‬
‭o nly approximately 24 hours following ovulation. Intercourse, more than a day after‬
‭ovulation, will usually not result in fertilization.‬
‭ hen the first sperm fuses with the oocyte, the oocyte deploys two mechanisms to block‬
W
‭polyspermy‬‭, which is penetration by more than one‬‭sperm. This block is critical because if‬
‭another sperm was to fertilize the oocyte, the resulting zygote would be a triploid‬
‭o rganism with three sets of chromosomes, which is incompatible with life.‬
‭ ost of the time, a woman releases a single egg during an ovulation cycle. However, two‬
M
‭eggs are released and fertilized in approximately one percent of ovulation cycles. Two‬
‭zygotes form, implant, and develop, resulting in the birth of‬‭dizygotic‬‭(or fraternal) twins.‬
‭Because dizygotic twins develop from two eggs fertilized by two sperm, they are no more‬
‭identical than siblings born at different times.‬
‭ ess commonly, a zygote can divide into two offspring during early development, resulting‬
L
‭in the birth of‬‭monozygotic‬‭(or identical) twins.‬‭Although the zygote can split as early as‬
‭the two-cell stage, splitting occurs most commonly during a later stage, with roughly‬
‭70–100 cells present. These two scenarios are distinct because twin embryos separated at‬
‭the two-cell stage will have individual placentas. In contrast, twin embryos that form from‬
‭separation at the later stage will share a placenta.‬

‭14.9 Development of the Fetal Male and Female Reproductive Systems‬

‭ he reproductive systems of humans begin to develop soon after conception. A gene on‬
T
‭the male’s Y chromosome called‬‭SRY‬‭(‬‭S‬‭ex Determining‬‭R‬‭egion of the‬‭Y‬‭Chromosome)is‬
‭critical in stimulating a cascade of events that simultaneously stimulate testis development‬
‭and repress the development of female structures. Testosterone produced by Leydig cells‬
‭in the embryonic testis stimulates the development of male sexual organs. If testosterone‬
‭is not present, female sexual organs will develop.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ urther development of the reproductive system occurs at puberty. The changes that‬
F
‭o ccur in puberty result from a decrease in sensitivity to negative feedback in the‬
‭hypothalamus and pituitary gland and an increase in sensitivity of the gonads to FSH and‬
‭LH stimulation. These changes increase estrogen or testosterone in female and male‬
‭adolescents, respectively. The increase in sex steroid hormones leads to the maturation of‬
‭the gonads and other reproductive organs. The initiation of spermatogenesis begins in‬
‭boys, and girls begin ovulating and menstruating.‬
‭ echanisms of sex determination differ across nature. For additional information about‬
M
‭the various mechanisms, click the link on the next page or scan the QR code to watch the‬
‭TED-Ed video titled “Sex determination: more complicated than you thought.”‬

Sex Determination: More Complicated Than You Thought

‭14.10 Pregnancy, Gestation, and Birth‬

‭ uman pregnancy begins with the fertilization of an egg and proceeds through the three‬
H
‭trimesters of‬‭gestation‬‭(‬‭Figure 14.17‬‭). A full-term‬‭pregnancy lasts approximately 270‬
‭days (approximately 38.5 weeks) from conception to birth. Because it is easier to‬
‭remember the first day of the last menstrual period (LMP) than to estimate the date of‬
‭conception, obstetricians set the due date as 284 days (approximately 40.5 weeks) from‬
‭the LMP. This estimate assumes conception occurred on day 14 of the woman’s cycle,‬
‭which is usually a good approximation. The 40 weeks of an average pregnancy are‬
‭typically discussed in terms of three trimesters, each approximately 13 weeks.‬

‭ igure 14.17‬ ‭A timeline of information about human pregnancy‬‭,‬‭including (from top to‬
F
‭bottom): Trimesters, embryo/fetus development, gestational age in weeks and months,‬
‭viability, and maturity stages. (credit:‬‭Pregnancy‬‭timeline.png‬‭;‬ ‭Mikael Hä ggströ m‬‭;‬‭CC0‬
‭1.0‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ or additional information about how a woman’s body is affected by a pregnancy, click the‬
F
‭link or scan the QR code to watch the TED-Ed video titled “The surprising effects of‬
‭pregnancy.”‬

The surprising effects of pregnancy

‭ he developing embryo must implant into the uterus wall within seven days, or it will‬
T
‭deteriorate and die. The outer layers of the developing zygote grow into the endometrium‬
‭by digesting the endometrial cells, and wound healing of the endometrium closes up the‬
‭blastocyst into the tissue. Another layer of the developing zygote, the chorion, begins‬
‭releasing a hormone called‬‭human beta chorionic gonadotropin‬‭(‬‭β‭-‬ HCG),‬‭which makes‬
‭its way to the corpus luteum and keeps that structure active. This result ensures adequate‬
‭levels of progesterone that will maintain the endometrium of the uterus for the support of‬
‭the developing embryo. Pregnancy tests determine the level of‬‭β‭-‬ HCG in urine or serum. If‬
‭the hormone is present, the test is positive.‬

‭ or additional information about pregnancy tests, click the link below or scan the QR code‬
F
‭to watch the TED-Ed video by Tien Nguyen titled “How do pregnancy tests work?”.‬

How do pregnancy tests work? - Tien Nguyen

‭ he endometrial lining handles nutrition and waste through diffusion during the first two‬
T
‭to four weeks of the first trimester. As the trimester progresses, the embryo's outer layer‬
‭merges with the endometrium, and the‬‭placenta‬‭forms.‬‭This organ takes over the nutrient‬
‭and waste requirements of the embryo and fetus, with the mother’s blood passing‬
‭nutrients to the placenta and removing waste. Some of the mother’s immunoglobulins‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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(‭ antibodies) will pass through the placenta, providing passive immunity against some‬
‭potential infections.‬
I‭ nternal organs and body structures begin to develop during the first trimester. By five‬
‭weeks, limb buds, eyes, the heart, and the liver have been formed. The term‬‭fetus‬‭applies‬
‭by eight weeks, and the body is essentially formed, as shown in‬‭Figure 14.18‬‭. The‬
‭individual is about two inches long, and many organs, such as the lungs and liver, are not‬
‭yet functioning. Exposure to toxins is hazardous during the first trimester, as all of the‬
‭body’s organs and structures undergo initial development. Anything that affects that‬
‭development can severely affect the fetus’ survival.‬

‭Figure 14.18‬ ‭Fetal development at nine weeks gestation.‬‭(credit: Ed Uthman;‬‭OpenStax‬‭)‬

‭ he fetus grows to about 12 inches during the second trimester, as shown in‬‭Figure‬
T
‭14.19‬‭. It becomes active, and the mother usually‬‭feels the first movements. All organs‬
‭and structures continue to develop. The placenta has taken over the functions of nutrition‬
‭and waste and the production of estrogen and progesterone from the corpus luteum,‬
‭which has degenerated. The placenta will continue functioning up through the delivery of‬
‭the baby.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ uring the third trimester, the fetus grows to 6 ½ -8 ½ lbs. and is about 19-20 inches long,‬
D
‭as illustrated in‬‭Figure 14.20‬‭. This trimester is‬‭the period of the most rapid growth during‬
‭the pregnancy. Organ development continues until birth, and some systems, such as the‬
‭nervous system, continue to develop after birth.‬

‭ igure 14.19‬ ‭This fetus is just entering the second‬‭trimester, when the placenta, located‬
F
‭above the fetus, takes over more functions performed during development. (credit:‬
‭National Museum of Health and Medicine;‬‭OpenStax‬‭)‬

‭ fter the newborn is successfully delivered, the placenta and associated membranes,‬
A
‭commonly called‬‭afterbirth‬‭, are delivered. Contractions‬‭o f the myometrium shear the‬
‭placenta from the back of the uterine wall, and it is delivered through the vagina.‬
‭Continued uterine contractions then reduce blood loss from the site of the placenta.‬
‭Delivery of the placenta marks the beginning of the‬‭postpartum period‬‭—approximately‬
‭six weeks immediately following childbirth, during which the mother’s body gradually‬
‭returns to a non-pregnant state. If the placenta does not birth spontaneously within about‬
‭30 minutes, it is classified as retained, and the doctor may attempt manual removal. If this‬
‭is not successful, surgery may be required.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ he doctor must examine the expelled placenta and fetal membranes to ensure they are‬
T
‭intact. If fragments of the placenta remain in the uterus, they can cause postpartum‬
‭hemorrhage. Uterine contractions continue for several hours after birth to return the‬
‭uterus to its pre-pregnancy size in a process called involution, allowing the mother’s‬
‭abdominal organs to return to their pre-pregnancy locations.‬

‭ igure 14.20‬ ‭There is rapid fetal growth during‬‭the third trimester. (credit: modification‬
F
‭o f work by Gray’s Anatomy;‬‭OpenStax‬‭)‬

‭14.11 Contraception and Birth Control‬

‭ he prevention of pregnancy comes under the terms‬‭contraception‬‭o r‬‭birth control‬‭.‬
T
‭Strictly speaking, contraception refers to preventing the sperm and egg from joining. Both‬
‭terms are, however, frequently used interchangeably.‬
‭ igure 14.21‬‭lists some common methods of contraception.‬‭The failure rates listed are not‬
F
‭the ideal ones that could be realized but the typical ones that occur. A failure rate is the‬
‭number of pregnancies resulting from the method’s use over twelve months.‬‭Barrier‬
‭methods,‬‭such as condoms, cervical caps, and diaphragms, block sperm from entering the‬
‭uterus, preventing fertilization.‬‭Spermicides‬‭are‬‭chemicals that are placed in the vagina‬
‭that kill sperm. Sponges, which are saturated with spermicides, are placed in the vagina at‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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t‭ he cervical opening. Combinations of spermicidal chemicals and barrier methods achieve‬

‭lower failure rates than the methods when used separately.‬
‭ early a quarter of the couples using barrier methods, natural family planning, or‬
N
‭withdrawal can expect a failure of the method. Natural family planning is based on‬
‭monitoring the menstrual cycle and having intercourse only when the egg is unavailable. A‬
‭woman’s body temperature may rise a degree Celsius at ovulation, and the cervical mucus‬
‭may increase in volume and become more pliable. These changes give a general indication‬
‭o f when intercourse is more or less likely to result in fertilization.‬

‭Failure Rate in Typical‬

‭Method‬ ‭Examples‬ ‭Use Over 12 Months‬
‭Barrier‬ ‭ ale condom, female‬
m ‭15 to 24%‬
‭condom, sponge,‬
‭cervical cap, diaphragm,‬
‭spermicides‬
‭Hormonal‬ ‭o ral, patch, vaginal ring‬ ‭8%‬
‭injection‬ ‭3%‬

‭implant‬ ‭less than 1%‬

‭Other‬ ‭natural family planning‬ ‭12 to 25%‬

‭withdrawal‬ ‭27%‬

‭sterilization‬ ‭less than 1%‬

‭ igure 14.21‬ ‭Information regarding various contraceptive‬‭methods and their associated‬

F
‭failure rates.‬

‭ ithdrawal involves the removal of the penis from the vagina during intercourse before‬
W
‭ejaculation. This method has a high failure rate due to the possible presence of sperm in‬
‭the bulbourethral gland’s secretion, which may enter the vagina before removing the‬
‭penis.‬
‭ ormonal methods‬‭use synthetic progesterone (sometimes in combination with‬
H
‭estrogen) to inhibit the hypothalamus from releasing FSH or LH and thus prevent an egg‬
‭from being available for fertilization. The method of administering the hormone affects the‬
‭failure rate. The most reliable method, with a failure rate of less than one percent, is the‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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i‭mplantation of the hormone under the skin. The same rate can be achieved through the‬
‭sterilization procedures of vasectomy in the man, tubal ligation in the woman, or by using‬
‭an‬‭intrauterine device (IUD)‬‭. IUDs are inserted into‬‭the uterus and establish an‬
‭inflammatory condition that prevents fertilized eggs from implanting into the uterine wall.‬
‭ irth control pills provide constant levels of estrogen and progesterone, which negatively‬
B
‭feedback on the hypothalamus and pituitary and suppress the release of FSH and LH,‬
‭which inhibits ovulation and prevents pregnancy.‬
‭ ompliance with the contraceptive method strongly contributes to any method's success‬
C
‭o r failure rate. The only method that is entirely effective at preventing conception is‬
‭abstinence. The choice of contraceptive method depends on the woman's or couple's‬
‭goals. Tubal ligation and vasectomy are considered permanent prevention, while other‬
‭methods are reversible and provide short-term contraception.‬
‭ or additional information about the physical basis of different birth control methods,‬
F
‭click the link below or scan the QR code to watch the TED-Ed video by NW Hunter titled‬
‭“How do contraceptives work?”.‬

How do contraceptives work? - NWHunter

‭14.12 Infertility, IVF, and “Designer Babies”‬

I‭ nfertility‬‭is the inability to conceive or carry‬‭a child to birth. About 75% of causes of‬
‭infertility include diseases, such as sexually transmitted diseases that can cause scarring of‬
‭the reproductive tubes in either men or women or developmental problems frequently‬
‭related to abnormal hormone levels in one of the individuals. Inadequate nutrition,‬
‭especially starvation, can delay menstruation. Stress can also lead to infertility. Short-term‬
‭stress can affect hormone levels, while long-term stress can delay puberty and cause less‬
‭frequent menstrual cycles. Other factors that affect fertility include toxins, tobacco‬
‭smoking, marijuana use, gonadal injuries, and aging.‬
I‭ f infertility is identified, several assisted reproductive technologies (ART) are available to‬
‭aid conception. A common type of ART is‬‭in vitro‬‭fertilization (IVF)‬‭.‬‭In vitro‬‭, which in‬
‭Latin translates to “in glass,” refers to a procedure outside the body. Many different‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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r‭ easons support using IVF as a solution. For example, a woman may produce normal eggs,‬
‭but they cannot reach the uterus because the uterine tubes are blocked or otherwise‬
‭compromised. A man may have a low sperm count, low sperm motility, sperm with an‬
‭unusually high percentage of morphological abnormalities, or sperm incapable of‬
‭penetrating an egg's outermost layer.‬
‭ he IVF procedure (‬‭Figure 14.22‬‭) begins with obtaining eggs from the woman‬‭after‬
T
‭extensive hormonal treatments that prepare mature eggs for fertilization and the uterus‬
‭for implantation of the fertilized egg. Sperm cells, obtained from a designated male or a‬
‭sperm bank, are combined in a glass (Petrie) dish with the eggs; the ideal ratio is 75,000‬
‭sperm to one egg. The embryos are incubated upon successful fertilization until they‬
‭reach the eight-cell or blastocyst stages. In the United States, fertilized eggs are typically‬
‭cultured to the blastocyst stage because this results in a higher pregnancy rate. Finally, the‬
‭embryos are transferred to a woman’s uterus using a plastic catheter (tube).‬
‭ or additional information about IVF, click the link or scan the QR code to watch the‬
F
‭TED-Ed video by Nassim Assefi and Brian A. Levine titled “How‬‭in vitro‬‭fertilization works.”‬

How in vitro fertilization (IVF) works - Nassim Assefi and Brian A. L…

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 14.22‬ ‭In vitro fertilization involves egg collection from the ovaries, fertilization‬‭in a‬
F
‭petri dish, and the transfer of embryos into the uterus. (credit:‬‭OpenStax‬‭). A link‬‭to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Everyday Connection‬

‭Designer Babies and Eugenics‬

‭ igure 14.23‬ ‭This logo from the Second International‬‭Eugenics Conference in New York‬
F
‭City in September of 1921 shows how eugenics attempted to merge several fields of study‬
‭to produce a genetically superior human race. (credit:‬‭Cold Spring Harbor Laboratory‬‭)‬

I‭ f you could prevent your child from getting a devastating genetic disease, would you do‬
‭it? Would you select the sex of your child or choose their attractiveness, strength, or‬
‭intelligence? How far would you go to maximize the possibility of disease resistance? The‬
‭genetic engineering of a human child, producing "designer babies" with desirable‬
‭phenotypic characteristics, was once a topic restricted to science fiction. This is the case no‬
‭longer: science fiction now overlaps with science facts. Many phenotypic choices for‬
‭o ffspring are already available, with many more likely to be possible in the not-too-distant‬
‭future. Which traits should be selected and how they should be selected are topics of‬
‭much debate within the worldwide medical community. The ethical and moral line is not‬
‭always clear or agreed upon, and some fear that modern reproductive technologies could‬
‭be detrimental.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ ugenics‬‭is the use of information and technology from various sources to “improve” the‬
E
‭genetic makeup of the human race. The conceptual basis for improvement was founded‬
‭o n similar principles used in livestock breeding. Progeny with economically desired‬
‭phenotypes (e.g., higher growth rate, higher milk production, greater numbers of‬
‭o ffspring produced, etc.) result from matings between parents with superior phenotypes‬
‭compared to others. This is because some portion of the progeny phenotypes for these‬
‭traits have an inherited (genetic) basis.‬

‭ he goal of creating genetically superior humans was prevalent in several countries,‬

T
‭including the United States, during the early 20‬‭th‬ ‭century. Eugenics classes were‬
‭commonly offered in colleges, and county and state fairs included informational displays‬
‭(‬‭Figure 14.24; p. 378‬‭).‬

I‭ n 1927, the Supreme Court accepted the case‬‭Carrie‬‭Buck v. John Hendren Bell,‬
‭Superintendent of State Colony for Epileptics and Feeble-Minded‬‭. It focused on a new‬
‭Virginia State law (‬‭The Virginia Sterilization Act‬‭)‬‭that gave the state government the legal‬
‭right to forcibly sterilize individuals deemed “mentally defective.” Carrie Buck, chosen as‬
‭the defendant in the test case to determine the legality of the new law, was an unwed‬
‭woman who became pregnant at the age of 17 years. Her pregnancy was later‬
‭determined to have occurred as a result of rape. Carrie was relocated to the Virginia‬
‭Colony for Epileptics and Feeble-Minded, where her birth mother was also housed. The‬
‭superintendent of this colony, Dr. Albert Priddy, submitted the following paragraph‬
‭(‬‭Figure 14.25; p. 379‬‭) in his petition to the Supreme‬‭Court to uphold the law:‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 14.24‬ ‭A Eugenics Society exhibit from the‬‭1930s. Note the inclusion of‬
F
‭information about Mendel’s Laws, pedigrees, and the traits associated with them: “Two‬
‭Well-To-Do Families,” “Inheritance of Mental Deficiency and Insanity,” and “Dramatic‬
‭Ability: Music and Art.” (credit:‬‭Eugenics Society‬‭Exhibit (1930s).jpg‬‭; Wellcome Library;‬
‭Wikimedia Commons)‬

‭A link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 14.25‬ ‭Statement by Dr. Albert Priddy from‬‭his petition to the Supreme Court in‬
F
‭the case of‬‭Carrie Buck v. John Hendren Bell, Superintendent‬‭of State Colony for Epileptics and‬
‭Feeble-Minded‬‭. John Bell, who was the colony superintendent‬‭at the time the case was‬
‭initially filed, was replaced by Dr. Priddy. (credit:‬ ‭National Archives Education Updates‬‭)‬

‭ he Supreme Court, in an 8-1 vote, ruled the‬‭Virginia‬‭Sterilization Act‬‭was constitutional‬

T
‭and that forced sterilization of “feeble-minded” individuals was permissible. Writing for‬
‭the majority opinion, Chief Justice Oliver Wendell Holmes stated that “Three generations of‬
‭imbeciles are enough.” (‬‭Figure 14.26‬‭)‬

‭ lthough the‬‭Buck v. Bell‬‭decision was never formally‬‭overturned, the Supreme Court‬

A
‭rendered a decision in another case in 1942 that outlawed sterilization as a punitive‬
‭action. The Virginia Sterilization Act was repealed by the state legislature in 1974. In‬
‭2002, 50 years after the initial ruling, the governor of Virginia renounced the state’s‬
‭sterilization legislation and apologized for the past eugenics-related actions.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 14.26‬ ‭An excerpt of the opinion by Oliver‬‭Wendell Holmes in the case of‬‭Carrie‬
F
‭Buck v. John Hendren Bell, Superintendent of State Colony for Epileptics and Feeble-Minded.‬
‭The Supreme Court ruled that the‬‭Virginia Sterilization‬‭Act‬‭was constitutional.‬ ‭The‬
‭infamous phrase from the opinion, “Three generations of imbeciles are enough,” is‬
‭underlined. (credit:‬ ‭National Archives Education‬‭Updates‬‭)‬

‭ azi Germany developed an extensive eugenics program in the 1930s and 40s using‬
N
‭America's eugenics policies as a foundation. As part of their program, the Nazis forcibly‬
‭sterilized hundreds of thousands of the so-called "unfit." They killed thousands of‬
‭institutionally disabled people as part of a systematic program to develop a genetically‬
‭superior race of Germans known as Aryans. Since then, eugenic ideas have not been as‬
‭publicly expressed, but some still promote them.‬

‭ or additional details regarding the history of eugenics in the United States, click the link‬
F
‭below or scan the QR code to watch the TED-Ed video by Alexandra M. Stern and Natalie‬
‭Lira titled “The movement that inspired the Holocaust.”‬

The movement that inspired the Holocaust - Alexandra Min…

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ fforts have been made to control traits in human children using donated sperm from‬
E
‭men with desired characteristics. Eugenicist Robert Klark Graham established a sperm‬
‭bank in 1980 that included samples exclusively from donors with high IQs. The "genius"‬
‭sperm bank failed to capture the public's imagination, and the operation closed in 1999.‬

‭ he procedure known as‬‭prenatal genetic diagnosis‬‭(PGD)‬‭has recently been‬

T
‭developed. PGD involves screening human embryos as part of the process of IVF when‬
‭embryos are conceived and grown outside the mother's body for some time before they‬
‭are implanted. PGD usually refers to the selected embryos' diagnosis, selection, and‬
‭implantation of the selected embryos.‬

I‭ n the least controversial use of PGD, embryos are tested for the presence of alleles that‬
‭cause genetic diseases such as sickle cell disease, muscular dystrophy, and hemophilia, in‬
‭which a single disease-causing allele or pair of alleles has been identified. The disease is‬
‭prevented by excluding embryos containing these alleles from implantation into the‬
‭mother, and the unused embryos are either donated to science or discarded. There is also‬
‭considerable controversy about what happens to unused embryos.‬

‭ here are relatively few in the worldwide medical community that question the ethics of‬
T
‭this type of procedure, which allows individuals scared to have children because of the‬
‭alleles they carry to do so successfully. The major limitation of this procedure is its‬
‭expense. Not usually covered by medical insurance and thus out of reach financially for‬
‭most couples, only a tiny percentage of all live births use such complicated methodologies.‬
‭Yet, even in cases where the ethical issues may seem clear-cut, not everyone agrees with‬
‭the morality of these types of procedures. For example, to those who believe that human‬
‭life begins at conception, discarding unused embryos, a necessary result of PGD, is‬
‭unacceptable under any circumstances.‬

‭ murkier ethical situation is found in selecting a child's sex, which PGD quickly performs.‬
A
‭Countries such as Great Britain have banned the selection of a child's sex for reasons‬
‭o ther than preventing sex-linked diseases. Other countries allow the procedure for "family‬
‭balancing" based on the desire of some parents to have at least one child of each sex. Still,‬
‭o thers, including the United States, have taken a scattershot approach to regulating these‬
‭practices, essentially leaving it to the individual practicing physician to decide which‬
‭practices are acceptable and which are not.‬

‭ ven murkier are rare instances of disabled parents, such as those with deafness or‬
E
‭dwarfism, who select embryos via PGD to ensure they share their disability. These parents‬
‭usually cite many positive aspects of their disabilities and associated culture as reasons for‬
‭their choice, which they see as their moral right. To others, to purposely cause a disability‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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i‭n a child violates the basic medical principle of‬‭Primum non nocere,‬‭"first, do no harm."‬
‭Although not illegal in most countries, this procedure demonstrates the complexity of‬
‭ethical issues associated with choosing genetic traits in offspring.‬

‭ or additional information about designer babies and ethical concerns, click the link‬
F
‭below or scan the QR code to watch the TED-Ed video by Paul Knepfler titled “The ethical‬
‭dilemma of designer babies.”‬

The ethical dilemma of designer babies | Paul Knoep…

‭ here could these technologies lead? Will these processes become more affordable, and‬
W
‭how should they be used? With the ability of technology to progress rapidly and‬
‭unpredictably, a lack of definitive guidelines for the use of reproductive technologies‬
‭before they arise might make it easier for legislators to keep pace once they are realized,‬
‭assuming the process needs any government regulation. Other bioethicists argue that we‬
‭should only deal with current technologies, not in some uncertain future. They say that‬
‭these procedures will always be expensive and rare, so the fears of eugenics and “master”‬
‭races are unfounded and overstated. The debate continues.‬

‭14.13 Career Connection - Reproductive Endocrinologist‬

‭ reproductive endocrinologist is a physician who treats a variety of hormonal disorders‬

A
‭related to reproduction and infertility in people of any gender. The disorders include‬
‭menstrual problems, infertility, pregnancy loss, sexual dysfunction, and menopause.‬
‭Doctors may use fertility drugs, surgery, or assisted reproductive techniques (ART) in‬
‭their therapy. ART involves using procedures to manipulate the egg or sperm to facilitate‬
‭reproduction, such as in vitro fertilization.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ eproductive endocrinologists undergo extensive medical training, first in a four-year‬

R
‭residency in obstetrics and gynecology, then in a three-year fellowship in reproductive‬
‭endocrinology. To be board-certified in this area, the physician must pass written and oral‬
‭exams in both areas.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 15: Skeletal System‬

‭ igure 15.1‬ ‭Different types of bones found in the‬‭human body (credit:‬ ‭Bones of the‬
F
‭Skeletal System‬‭;‬‭Scientific Animations‬‭;‬‭CC BY-SA 4.0‬‭)‬

‭15.1 Introduction‬
‭ hen most students think of the skeletal system, images of bones, such as those shown in‬
W
‭Figure 15.1,‬‭understandably come to mind. Your skeleton‬‭comprises living tissue that‬
‭grows, repairs, and renews itself. The bones associated with this system are dynamic and‬
‭complex organs that serve several essential functions: supporting the body, storing‬
‭minerals and lipids, producing blood cells, protecting internal organs, and allowing‬
‭movement.‬
‭ he human skeleton, which consists of 206 bones in the adult, is divided into the axial‬
T
‭skeleton (which consists of the skull, vertebral column, and rib cage) and the appendicular‬
‭skeleton (which consists of the shoulders, limb bones, the pectoral girdle, and the pelvic‬
‭girdle).‬

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‭15.2 Human Axial Skeleton‬

‭ he‬‭axial skeleton‬‭forms the body's central axis and‬‭includes the skull, vertebral column,‬
T
‭ribcage, auditory ossicles, and hyoid bone (‬‭Figure 15.2‬‭). The function of the axial skeleton‬
‭is to provide support and protection for the brain, the spinal cord, and other organs. It‬
‭provides a surface for the attachment of muscles that move the head, neck, and trunk,‬
‭perform respiratory movements, and stabilize parts of the appendicular skeleton.‬

‭ igure 15.2‬ ‭The axial skeleton consists of the bones‬‭o f the skull, ossicles of the middle‬
F
‭ear, hyoid bone, vertebral column, and rib cage. The hyoid bone and ossicles are shown in‬
‭Figure 15.7. (credit: modification of work by Mariana Ruiz Villareal;‬‭OpenStax‬‭)‬

‭The Skull‬
‭ he bones of the‬‭skull‬‭support the structures of the‬‭face and protect the brain. The skull‬
T
‭consists of 22 bones divided into two categories: cranial bones and facial bones. The‬
‭c ranial bones‬‭are eight bones that form the cranial‬‭cavity, which encloses the brain and‬
‭serves as an attachment site for the head and neck muscles. The eight cranial bones are‬
‭the frontal bone, two parietal bones, two temporal bones, the occipital bone, the sphenoid‬
‭bone, and the ethmoid bone. Although the bones develop separately in the embryo and‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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f‭ etus, they are tightly fused with connective tissue in the adult, and adjoining bones do not‬
‭move (‬‭Figure 15.3‬‭).‬

‭ igure 15.3‬ ‭The bones of the skull support the structures‬‭o f the face and protect the‬
F
‭brain. (credit: modification of work by Mariana Ruiz Villareal;‬‭OpenStax‬‭)‬

‭ ourteen‬‭facial bones‬‭form the face, provide cavities‬‭for the sense organs (eyes, mouth,‬
F
‭and nose), protect the entrances to the digestive and respiratory tracts, and serve as‬
‭attachment points for facial muscles. The facial bones include the nasal bones, the‬
‭maxillary bones, the zygomatic bones, the palatine, vomer, lacrimal bones, the inferior‬
‭nasal conchae, and the mandible. These bones occur in pairs except for the mandible and‬
‭the vomer (‬‭Figure 15.4‬‭).‬

‭The Rib Cage‬

‭ he‬‭rib cage‬‭consists of the ribs, sternum, thoracic‬‭vertebrae, and costal cartilages‬
T
‭(‬‭Figure 15.5‬‭). It encloses and protects the organs‬‭o f the thoracic cavity, including the‬
‭heart and lungs. It also provides support for the shoulder girdles and upper limbs and‬
‭serves as an attachment point for the diaphragm and muscles of the back, chest, neck, and‬
‭shoulders.‬

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‭ igure 15.4‬ ‭The cranial bones, including the frontal,‬‭parietal, and sphenoid bones, cover‬
F
‭the top of the head. The facial bones of the skull form the face and provide cavities for the‬
‭eyes, nose, and mouth. (credit:‬‭OpenStax‬‭)‬

‭ igure 15.5‬ ‭The thoracic cage, or rib cage, protects‬‭the heart and the lungs. (credit:‬
F
‭modification of work by NCI, NIH;‬‭OpenStax‬‭). A link‬‭to a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

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‭ he‬‭sternum‬‭, or breastbone, is a long, flat bone at the front of the chest. The‬‭ribs‬‭are 12‬
T
‭pairs of long, curved bones that attach to the thoracic vertebrae and curve toward the‬
‭front of the body, forming the ribcage. Cartilage connects one end of the ribs to the‬
‭sternum, except for rib pairs 11 and 12, which are free-floating ribs.‬

‭The Vertebral Column‬

‭ he‬‭vertebral column‬‭, or‬‭spinal column‬‭, surrounds‬‭and protects the spinal cord,‬
T
‭supports the head, and acts as an attachment point for the ribs and muscles of the back‬
‭and neck. The adult vertebral column comprises 26 bones: the 24 vertebrae, the sacrum,‬
‭and the coccyx bones. In adults, the‬‭sacrum‬‭is typically‬‭composed of five vertebrae that‬
‭fuse into one; the‬‭coccyx‬‭generally is three to four‬‭vertebrae that fuse into one. We begin‬
‭life with approximately 33 vertebrae, but as we grow, several vertebrae fuse. The adult‬
‭vertebrae are divided into seven cervical vertebrae, 12 thoracic vertebrae, and five lumbar‬
‭vertebrae (‬‭Figure 15.6‬‭).‬

‭The Auditory Ossicles and the Hyoid Bone‬

‭ he‬‭auditory ossicles‬‭o f the middle ear transmit sounds‬‭from the air as vibrations to a‬
T
‭fluid-filled structure called the cochlea. The auditory ossicles consist of three bones: the‬
‭malleus, incus, and stapes (‬‭Figure 15.7a‬‭). These are‬‭the smallest bones in the body and‬
‭are unique to mammals.‬

‭ lthough it is not found in the skull, the hyoid bone is considered a component of the axial‬
A
‭skeleton. It lies below the mandible in the front of the neck (‬‭Figure 15.7b‬‭), and it acts as a‬
‭movable base for the tongue and is connected to the jaw, larynx, and tongue muscles. The‬
‭mandible articulates with the base of the skull and controls the opening to the airway and‬
‭gut.‬

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‭ igure 15.6‬ ‭(a) The vertebral column consists of‬‭seven cervical vertebrae (C1–7), twelve‬
F
‭thoracic vertebrae (Th1–12), five lumbar vertebrae (L1–5), the os sacrum, and the coccyx.‬
‭(b) Spinal curves increase the strength and flexibility of the spine. (credit a: modification‬
‭o f work by Uwe Gille;‬‭OpenStax‬‭)‬

‭ igure 15.7‬ ‭a) The ossicles of the human ear: malleus,‬‭incus, and stapes. b) The position‬
F
‭o f the hyoid bone above the larynx. credit: (a)‬‭auditory ossicles.jpg‬‭; US Government;‬‭CC0‬
‭1.0‬ ‭(b)‬‭Hyoid in throat.jpg‬‭; Openstax College;‬‭CC‬‭BY 3.0‬‭)‬

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P
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‭15.3 Human Appendicular Skeleton‬

‭ he‬‭appendicular skeleton‬‭comprises the bones of the‬‭upper limbs (which function to‬
T
‭grasp and manipulate objects) and the lower limbs (which permit locomotion). It also‬
‭includes the pectoral girdle, or shoulder girdle, which attaches the upper limbs to the‬
‭body, and the pelvic girdle, which connects the lower limbs to the body (‬‭Figure 15.8‬‭).‬

‭ igure 15.8‬ ‭The appendicular skeleton comprises‬‭the bones of the pectoral limbs (arm,‬
F
‭forearm, hand), the pelvic limbs (thigh, leg, foot), the pectoral girdle, and the pelvic girdle.‬
‭(credit: modification of work by Mariana Ruiz Villareal;‬‭OpenStax‬‭)‬

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P
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‭The Pectoral (Upper) Limbs‬

‭ he upper limbs contain 30 bones in three regions: the arm (shoulder to elbow), the‬
T
‭forearm (ulna and radius), and the wrist and hand (‬‭Figure 15.9‬‭). An‬‭articulation‬‭is any‬
‭place at which two bones are joined. The humerus is the largest and longest bone of the‬
‭upper limbs and the only arm bone. It articulates with the shoulder scapula and the‬
‭forearm at the elbow. The forearm extends from the elbow to the wrist and consists of‬
‭two bones: the ulna and the radius. The radius is located along the thumb side of the‬
‭forearm and articulates with the humerus at the elbow. The ulna is located on the pinky‬
‭finger side of the forearm and articulates with the humerus at the elbow. The radius and‬
‭ulna also articulate with the carpal bones and each other. The hand includes the carpals‬
‭(wrist; 8 bones), the metacarpals (palm; 5 bones), and the phalanges (digits; 14 bones).‬
‭Each digit consists of three phalanges, except for the thumb, which has two.‬

‭ igure 15.9‬ ‭The pectoral limbs consist of the humerus‬‭o f the upper arm, the radius and‬
F
‭ulna of the forearm, the carpals (8 bones), the metacarpals (5 bones), and the phalanges‬
‭(14 bones). (credit:‬ ‭OpenStax‬‭)‬

‭ any students are familiar with “cracking” or “popping” the hand joints. However, what is‬
M
‭the cause of this noise? For an explanation, click the link below or scan the QR code to‬
‭watch the TED-Ed video by Eleanor Nelsen titled “Why do your knuckles pop?”.‬

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Why do your knuckles pop? - Eleanor Nelsen

‭The Pelvic (Lower) Limbs‬

‭ he lower limbs consist of the thigh, the leg, and the foot; the bones associated with these‬
T
‭regions are the femur (thigh bone), patella (kneecap), tibia, and fibula (bones of the leg),‬
‭tarsals (bones of the ankle), and metatarsals and phalanges (bones of the foot) (‬‭Figure‬
‭15.10‬‭). The bones of the lower limbs are thicker and‬‭stronger than the bones of the upper‬
‭limbs because of the need to support the entire weight of the body and the resulting‬
‭forces from movement.‬
‭ he femur, or thighbone, is the body's longest, heaviest, and strongest bone. The femur‬
T
‭and pelvis form the hip joint at the proximal end. The femur, tibia, and patella form the‬
‭knee joint at the distal end. The patella, or kneecap, is a triangular bone in front of the‬
‭knee joint. It improves knee extension by reducing friction. The tibia, or shinbone, is a‬
‭large leg bone directly below the knee. The tibia articulates with the femur at its proximal‬
‭end, with the fibula and the tarsal bones at its distal end. It is the second largest bone in‬
‭the human body and is responsible for transmitting the body's weight from the femur to‬
‭the foot. The fibula, or calf bone, parallels and articulates with the tibia. It does not‬
‭articulate with the femur and does not bear weight. The fibula acts as a site for muscle‬
‭attachment and forms part of the ankle joint.‬
‭ he tarsals are the seven bones of the ankle, which transmit the body's weight from the‬
T
‭tibia and fibula to the foot. The metatarsals are the five bones of the foot. The phalanges‬
‭are the 14 bones of the toes. Each toe consists of three phalanges, except for the big toe,‬
‭which has only two (‬‭Figure 15.11‬‭).‬

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‭ igure 15.10‬ ‭The lower limbs consist of the thigh‬‭(femur), the kneecap (patella), the leg‬
F
‭(tibia and fibula), the ankle (tarsals), and the foot (metatarsals and phalanges) bones.‬
‭(credit:‬ ‭OpenStax‬‭)‬

‭ igure 15.11‬ ‭The human foot and ankle bones, including‬‭the tarsals, metatarsals, and‬
F
‭phalanges. (credit:‬ ‭OpenStax‬‭)‬

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‭The Pectoral Girdle‬

‭ he‬‭pectoral girdle‬‭bones provide the points of attachment‬‭o f the upper limbs to the‬
T
‭axial skeleton. The human pectoral girdle consists of the clavicle (or collarbone) towards‬
‭the front side of the body and the scapula (or shoulder blades) towards the back side‬
‭(‬‭Figure 15.12‬‭).‬

‭ igure 15.12‬‭(a) The pectoral girdle in primates consists‬‭o f the clavicles and scapulae. (b)‬
F
‭The posterior view reveals the spine of the scapula to which muscle attaches. (credit:‬
‭OpenStax‬‭)‬

‭ he clavicles are S-shaped bones that position the arms on the body. The clavicles lie‬
T
‭horizontally across the front of the thorax (chest) just above the first rib. These bones are‬
‭relatively fragile and are susceptible to fractures. For example, a fall with the arms‬
‭o utstretched causes the force to be transmitted to the clavicles, which can break if the‬
‭force is excessive. The clavicles articulate with the sternum and the scapulae.‬

‭ he scapulae (plural of scapula) are flat, triangular-shaped bones that function in the‬
T
‭shoulder joint and participate in diverse shoulder movements. They also protect the back‬
‭o f the chest and the lungs in the thoracic cavity. The scapulae articulate with the humeri‬
‭(plural of humerus) and the clavicles.‬

‭The Pelvic Girdle‬

‭ he‬‭pelvic girdle‬‭attaches to the lower limbs of the‬‭axial skeleton via strong ligaments.‬
T
‭This is necessary because it bears the body's weight and movement.‬

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I‭ t also has deep sockets with strong ligaments to securely attach the femur to the body.‬
‭Two large hip bones further strengthen the pelvic girdle.‬
‭ he female pelvis is slightly different from the male pelvis. Over generations of evolution,‬
T
‭females with a wider pubic angle and larger diameter pelvic canal reproduced more‬
‭successfully. Therefore, their female offspring also had these characteristics that facilitate‬
‭successful childbirth (‬‭Figure 15.13‬‭).‬

‭ igure 15.13‬ ‭To improve reproductive fitness, the‬‭female pelvis is lighter, wider, and‬
F
‭shallower and has a broader angle between the pubic bones than the male pelvis. (credit:‬
‭OpenStax‬‭)‬

‭15.4 Bone Tissue‬

‭ ones are considered organs because they contain various types of tissues, such as blood,‬
B
‭connective tissue, nerves, and bone. The two types of bone tissue are compact and spongy.‬

‭Compact Bone Tissue‬

‭ ompact bone‬‭forms the hard external layer of all‬‭bones, providing protection and‬
C
‭strength. Compact bone tissue consists of units called‬‭osteons‬‭, cylindrical structures‬
‭containing a mineral matrix (consisting of calcium and phosphorus), and living cells‬
‭(called‬‭osteocytes‬‭). Each osteon consists of‬‭lamellae‬‭,‬‭which are layers of compact matrix‬
‭surrounding a central canal called the Haversian canal. The‬‭Haversian canal‬‭contains the‬
‭bone’s blood vessels and nerve fibers (‬‭Figure 15.14‬‭). Osteons in compact bone tissue are‬
‭aligned in the same direction along the lines of stress and help the bone resist bending or‬
‭fracturing.‬

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‭ he osteocytes are located inside spaces called‬‭lacunae‬‭(singular = lacuna), found at the‬

T
‭borders of adjacent lamellae. The cells communicate with each other and receive nutrients‬
‭via long cytoplasmic processes that extend through channels within the bone matrix called‬
‭c analiculi‬‭(singular = canaliculus). The canaliculi‬‭eventually merge with the Haversian‬
‭canal, which allows nutrients to be transported to the osteocytes and wastes to be‬
‭removed from them.‬

‭(a) (b)‬

‭ igure 15.14‬ ‭Compact bone tissue consists of osteons‬‭aligned parallel to the long axis of‬
F
‭the bone and the Haversian canal containing the bone’s blood vessels and nerve fibers.‬
‭The inner layer of bones consists of spongy bone tissue. The small dark ovals in the‬
‭o steon represent the living osteocytes. (credit: (a)‬‭U.S. National Cancer Institute's SEER‬
‭Program‬‭;‬‭CC0 1.0‬‭); (b) Provided by Willy Cushwa).‬ ‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭Spongy Bone Tissue‬

‭ pongy bone‬‭, also called‬‭c ancellous bone‬‭, contains‬‭o steocytes housed in lacunae, but‬
S
‭they are not arranged in concentric circles. Instead, the lacunae and osteocytes are found‬

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i‭n a lattice-like network of matrix spikes called‬‭trabeculae‬‭(singular = trabecula) (‬‭Figure‬

‭15.15‬‭). The trabeculae may appear to be a random network, but each trabecula forms‬
‭along the stress lines to strengthen the bone. The spaces of the trabeculated network‬
‭provide balance to the dense and heavy compact bone by making bones lighter so muscles‬
‭can move them more easily. In addition, the spaces in some spongy bones contain red‬
‭marrow, protected by the trabeculae, where the production of blood cells occurs.‬

‭(a) (b)‬
‭ igure 15.15‬ ‭(a) Diagram of spongy (cancellous)‬‭bone. (b) Histology of spongy bone,‬
F
‭showing the trabeculae and red bone marrow. (credit: (a)‬ ‭Servier Laboratories;‬
‭Wikimedia Commons;‬ ‭CC BY-SA 3.0‬‭;‬‭(b) Provided by‬‭Willy Cushwa)‬

‭15.5 Bone Classification and Gross Anatomy‬

‭ ones are classified according to their shapes. Long bones, such as the femur, are longer‬
B
‭than wide. Short bones, such as the carpals, are approximately equal in length, width, and‬
‭thickness. Flat bones are thin but are often curved, such as the ribs. Irregular bones, such‬
‭as those of the face, have no characteristic shape. Sesamoid bones, such as the patellae,‬
‭are small and round.‬
‭ he structure of a long bone allows for the best visualization of all of the parts of a bone‬
T
‭(‬‭Figure 15.16‬‭).‬‭Long bones‬‭have a shaft and two ends.‬‭The‬‭diaphysis‬‭, or central shaft,‬
‭has a hollow region called the‬‭medullary cavity‬‭and‬‭contains the‬‭yellow marrow‬‭, a‬
‭concentrated source of fat and energy potential. The walls of the diaphysis are composed‬
‭o f dense and hard compact bone.‬

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‭ igure 15.16‬ ‭The gross anatomy of the femur, classified‬‭as a long bone. (credit:‬
F
‭OpenStax‬‭)‬

‭ he rounded ends, or epiphyses, contain spongy bone, and its spaces are filled with red‬
T
‭marrow (‬‭Figure 15.17‬‭). Each epiphysis meets the diaphysis‬‭at the metaphysis, the narrow‬
‭area that contains the‬‭epiphyseal plate‬‭(growth plate),‬‭a layer of hyaline cartilage in a‬
‭growing bone. When the bone stops growing in early adulthood (approximately 18–21‬
‭years), the cartilage is replaced by bone, and the epiphyseal plate becomes an‬‭epiphyseal‬
‭line‬‭.‬

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‭ he medullary cavity has a delicate membranous lining called the‬‭endosteum‬‭(end- =‬

T
‭“inside”; oste- = “bone”), where bone growth, repair, and remodeling occur. The outer‬
‭surface of the bone is covered with a fibrous membrane called the‬‭periosteum‬‭(peri‬‭- =‬
‭“around” or “surrounding”). The periosteum contains blood vessels, nerves, and lymphatic‬
‭vessels that nourish compact bone. Tendons and ligaments also attach to bones at the‬
‭periosteum. The periosteum covers the outer surface except where the epiphyses meet‬
‭o ther bones to form joints. In these regions, the epiphyses are covered with a thin layer of‬
‭cartilage that reduces friction and acts as a shock absorber.‬

‭ igure 15.17‬ ‭The head of the femur contains both‬‭yellow and red marrow. Yellow‬
F
‭marrow stores fat (an energy reserve), and red marrow is responsible for the production‬
‭o f blood cells. (credit:‬‭OpenStax‬‭)‬

‭ or additional information about red bone marrow and the production of blood cells, click‬
F
‭the link below or scan the QR code to watch the TED-Ed video by Melody Smith‬‭titled‬
‭“How bones make blood.”‬

How bones make blood - Melody Smith

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‭15.6 Cell Types in Bones‬

‭ lthough bone cells compose a small amount of bone volume, they are crucial to bone‬
A
‭function. Four types of cells are found within bone tissue: osteoblasts, osteocytes,‬
‭o steogenic cells, and osteoclasts (‬‭Figure 15.18‬‭).‬

‭ steoblasts‬‭are bone cells responsible for bone formation.‬‭They synthesize and secrete‬
O
‭the organic and inorganic parts of the extracellular matrix of bone tissue and collagen‬
‭fibers. Osteoblasts become trapped in these secretions and differentiate into less active‬
‭osteocytes‬‭. These mature bone cells, located in lacunae,‬‭are surrounded by a matrix.‬
‭Osteocytes are the main ones in bone connective tissue, and they maintain typical bone‬
‭structure by recycling mineral salts.‬

‭ igure 15.18‬ ‭The four types of cells found within‬‭bone tissue and their respective‬
F
‭functions. (credit:‬‭OpenStax‬‭)‬

‭ steogenic cells‬‭are stem cells that divide by mitosis‬‭to produce daughter cells that‬
O
‭differentiate into osteoblasts. This is important because osteoblasts and osteocytes are‬
‭incapable of mitosis.‬

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‭ steoclasts‬‭are large bone cells that originate from two types of white blood cells‬
O
‭(monocytes and macrophages). They remove bone structure by releasing enzymes and‬
‭acids that dissolve the bony matrix. These minerals, released from bones into the blood,‬
‭help regulate calcium concentrations in body fluids. The bone may also be resorbed for‬
‭remodeling if the applied stresses have changed.‬
‭ steoclasts continually break down old bone, while osteoblasts continually form new‬
O
‭bone. The dynamic nature of bone means that new tissue is constantly formed, and old,‬
‭injured, or unnecessary bone is dissolved for repair or calcium release. The ongoing‬
‭balance between osteoblasts and osteoclasts is responsible for the constant but subtle‬
‭reshaping of bone.‬

‭15.7 Development and Growth of Bone‬

‭Development of Bone‬
‭ ssification‬‭, or‬‭osteogenesis‬‭, is the process of bone‬‭formation by osteoblasts.‬
O
‭Ossification is distinct from the process of calcification; whereas calcification occurs‬
‭during the ossification of bones, it can also occur in other tissues. Ossification begins‬
‭approximately six weeks after fertilization in an embryo. Before this time, the embryonic‬
‭skeleton consists entirely of fibrous membranes and hyaline cartilage. The development of‬
‭bone from fibrous membranes is called intramembranous ossification; development from‬
‭hyaline cartilage is called endochondral ossification. Bone growth continues until‬
‭approximately age 25. Bones can grow in thickness throughout life, but after age 25,‬
‭o ssification functions primarily in bone remodeling and repair.‬
‭ or additional information about bone formation and development, click the link below or‬
F
‭scan the QR code to watch the TED-Ed video by Nina Tandon‬‭titled “How to grow a bone.”‬

How to grow a bone - Nina Tandon

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Lengthening of Long Bones‬

‭ hondrocytes on the epiphyseal side of the epiphyseal plate divide; one daughter cell‬
C
‭remains undifferentiated near the epiphysis and the other moves toward the diaphysis.‬
‭The cells pushed from the epiphysis mature and are destroyed by calcification. This‬
‭process replaces cartilage with bone on the diaphyseal side of the plate, resulting in a‬
‭lengthening of the bone.‬
‭ ong bones stop growing at around the age of 18 in females and 21 in males in a process‬
L
‭called epiphyseal plate closure. During this process, cartilage cells stop dividing, and all of‬
‭the cartilage is replaced by bone. The epiphyseal plate fades, leaving a structure called the‬
‭epiphyseal line, and the epiphysis and diaphysis fuse.‬

‭Thickening of Long Bones‬

‭ ones increase in diameter by adding bony tissue at the surface of bones. Osteoblasts at‬
B
‭the bone surface secrete bone matrix, and osteoclasts on the inner surface break down‬
‭bone. The osteoblasts differentiate into osteocytes. A balance between these two‬
‭processes allows the bone to thicken without becoming too heavy.‬

‭15.8 Bone Remodeling and Repair‬

‭ one renewal continues after birth into adulthood.‬‭Bone remodeling‬‭is the replacement‬
B
‭o f old bone tissue with new tissue. It involves the processes of bone deposition by‬
‭o steoblasts and bone resorption by osteoclasts. Normal bone growth requires vitamins D,‬
‭C, and A, plus minerals such as calcium, phosphorus, and magnesium. Hormones such as‬
‭parathyroid hormone (PTH), growth hormone (GH), and calcitonin are also required for‬
‭proper bone growth and maintenance.‬
‭ one turnover rates are high, with five to seven percent of bone mass recycled weekly.‬
B
‭Differences in turnover rate exist in different areas of the skeleton and other regions of a‬
‭bone. For example, the bone in the head of the femur may be entirely replaced every six‬
‭months, whereas the bone along the shaft is altered much more slowly.‬
‭ one remodeling allows bones to adapt to stresses by becoming thicker and stronger‬
B
‭when subjected to stress. Bones that are not subject to everyday stress, for example, when‬
‭a limb is in a cast, will begin to lose mass.‬
‭ one fractures‬‭are classified by complexity, location,‬‭and other features (‬‭Figures 15.19‬
B
‭and‬‭15.20‬‭). Common types of fractures are transverse,‬‭o blique, spiral, comminuted,‬
‭impacted, greenstick, open (or compound), and closed (or simple). Some fractures may be‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ escribed using more than one term because they may have features of more than one‬
d
‭type (e.g., an open transverse fracture).‬

‭Type of fracture‬ ‭ escription‬

D
‭Transverse‬ ‭Occurs straight across the long axis of the‬
‭bone‬
‭Oblique‬ ‭Occurs at an angle that is not 90 degrees‬
‭Spiral‬ ‭ one segments are pulled apart as a‬
B
‭result of a twisting motion‬
‭Comminuted‬ S‭ everal breaks result in many small pieces‬
‭between two large segments‬
‭Impacted‬ ‭ ne fragment is driven into the other,‬
O
‭usually as a result of compression‬
‭Greenstick‬ ‭ partial fracture in which only one side‬
A
‭o f the bone is broken‬
‭Open (or compound)‬ ‭ fracture in which at least one end of the‬
A
‭broken bone tears through the skin;‬
‭carries a high risk of infection‬
‭Closed (or simple)‬ ‭ fracture in which the skin remains‬
A
‭intact‬

‭Figure 15.19‬ ‭A comparison of different types of‬‭bone fractures.‬

‭ hen a bone breaks, blood flows from any vessel torn by the fracture. These vessels‬
W
‭could be in the periosteum, osteons, and medullary cavity. The blood begins to clot, and‬
‭about six to eight hours after the fracture, the clotting blood has formed a fracture‬
‭hematoma (‬‭Figure 15.21a‬‭). The disruption of blood‬‭flow to the bone results in the death‬
‭o f bone cells around the fracture.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 15.20‬ ‭A comparison of different types of‬‭fractures: (a) closed fracture, (b) open‬
F
‭fracture, (c) transverse fracture, (d) spiral fracture, (e) comminuted fracture, (f) impacted‬
‭fracture, (g) greenstick fracture, and (h) oblique fracture. (credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 15.21‬ ‭The healing of a bone fracture follows‬‭a series of progressive steps: (a) A‬
F
‭fracture hematoma forms. (b) Internal and external calli form. (c) The cartilage of the calli‬
‭is replaced by trabecular bone. (d) Remodeling occurs. (credit:‬‭OpenStax‬‭)‬

‭ ithin about 48 hours after the fracture, chondrocytes have created an internal callus‬
W
‭(plural = calli) by secreting a fibrocartilaginous matrix between the two ends of the‬
‭broken bone, while other chondrocytes and osteoblasts create an external callus of‬
‭hyaline cartilage and bone, respectively, around the outside of the break (Figure‬‭15.21b‬‭).‬
‭This stabilizes the fracture.‬
‭ ver the next several weeks, osteoclasts resorb the dead bone; osteogenic cells become‬
O
‭active, divide, and differentiate into osteoblasts. The cartilage in the calli is replaced by‬
‭trabecular bone via ossification (‬‭Figure 15.21c‬‭).‬
‭ ventually, the internal and external calli unite, compact bone replaces spongy bone at the‬
E
‭o uter margins of the fracture, and healing is complete. A slight swelling may remain on the‬
‭o uter surface of the bone, but quite often, that region undergoes remodeling (‬‭Figure‬
‭15.21d‬‭), and no external evidence of the fracture‬‭remains.‬

‭ 5.9 Calcium Homeostasis: Interactions of the Skeletal System and Other Organ‬
1
‭Systems‬
‭ alcium‬‭(Ca) is a chemical element that cannot be‬‭produced by biological processes and‬
C
‭can only be obtained via the diet. It is the most abundant mineral in bone and the most‬
‭abundant mineral in the human body. Calcium ions are needed not only for bone‬

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‭ ineralization but for tooth health, regulation of the heart rate and strength of‬
m
‭contraction, blood coagulation, contraction of smooth and skeletal muscle cells, and‬
‭regulation of nerve impulse conduction. When the body cannot maintain an appropriate‬
‭concentration, a person will experience hypo- or hypercalcemia.‬
‭ ypocalcemia‬‭, characterized by abnormally low calcium‬‭levels, can adversely affect‬
H
‭several body systems, including circulation, muscles, nerves, and bone. Without adequate‬
‭calcium, blood has difficulty coagulating, the heart may skip beats or stop beating‬
‭altogether, muscles may have difficulty contracting, nerves may have trouble functioning,‬
‭and bones may become brittle. The causes of hypocalcemia can range from hormonal‬
‭imbalances to an improper diet. Treatments vary according to the cause, but prognoses‬
‭are generally good.‬
‭ onversely, in‬‭hypercalcemia‬‭, a condition characterized‬‭by abnormally high calcium‬
C
‭levels, the nervous system is underactive, resulting in lethargy, sluggish reflexes,‬
‭constipation and loss of appetite, confusion, and, in severe cases, coma.‬
‭ he bones act as a storage site for calcium: The body deposits calcium in the bones when‬
T
‭blood levels get too high, and it releases calcium when blood levels drop too low. The‬
‭skeletal, endocrine, and digestive systems and the kidneys work together to maintain‬
‭calcium homeostasis in the body (‬‭Figure 15.22‬‭).‬
‭ he cells of the‬‭parathyroid glands‬‭(‬‭Figure 15.23‬‭)‬‭have plasma membrane receptors for‬
T
‭calcium. When calcium does not bind to these receptors, the cells release‬‭parathyroid‬
‭hormone‬‭(‬‭PTH)‬‭, which stimulates an increasing number‬‭o f osteoclasts and bone‬
‭resorption. This demineralization process releases calcium into the blood. PTH promotes‬
‭the kidneys' calcium reabsorption from the urine so that the calcium returns to the blood.‬
‭Finally, PTH stimulates vitamin D synthesis, which promotes calcium absorption from any‬
‭digested food in the small intestine.‬
‭ hen all these processes return blood calcium levels to normal, there is enough calcium‬
W
‭to bind with the receptors on the surface of the cells of the parathyroid glands, and this‬
‭cycle of events is turned off.‬
‭ hen blood calcium levels get too high, the thyroid gland is stimulated to release‬
W
‭c alcitonin‬‭, which inhibits osteoclast activity, stimulates‬‭calcium uptake by the bones, and‬
‭decreases calcium reabsorption by the kidneys. These actions lower blood calcium levels.‬
‭When blood calcium levels return to normal, the thyroid gland stops secreting calcitonin.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 15.22‬ ‭The body regulates calcium homeostasis‬‭with two pathways; one is signaled‬
F
‭to turn on when blood calcium levels drop below average, and one is the pathway that is‬
‭signaled to turn on when blood calcium levels are elevated. (credit:‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

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‭ igure 15.23‬‭: The locations of the parathyroid glands‬‭(yellow) within the thyroid gland.‬
F
‭(credit: modification of‬‭parathyroid glands‬‭;‬‭Scientific‬‭Animations‬‭;‬‭CC BY-SA 4.0‬‭)‬

‭15.10 Nutrition and Bone Tissue‬

‭ he vitamins and minerals in our food are essential for the proper functioning of all organ‬
T
‭systems. However, certain nutrients specifically affect bone health.‬ ‭You already know that‬
‭calcium is a critical component of bone and must be obtained from the diet. However,‬
‭calcium cannot be absorbed from the small intestine without vitamin D. Therefore, vitamin‬
‭D intake is also critical to bone health. In addition to vitamin D’s role in calcium‬
‭absorption, it also plays a role in bone remodeling, though not as clearly understood.‬
‭ ilk and other dairy foods are not the only sources of calcium. This vital nutrient is also‬
M
‭found in green leafy vegetables, broccoli, intact salmon, and canned sardines with their‬
‭soft bones. Nuts, beans, seeds, and shellfish provide calcium in smaller quantities.‬
‭ itamin D is not found naturally in many foods besides fatty fish like salmon and tuna or‬
V
‭fortified milk or cereal. Sunlight on the skin triggers the body to produce vitamin D‬
‭(‬‭Figure 15.24‬‭). Still, many people, especially those‬‭o f darker complexions and those living‬
‭in northern latitudes where the sun’s rays are not as strong, are deficient in vitamin D. In‬
‭cases of deficiency, a doctor can prescribe a vitamin D supplement.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 15.24‬ ‭Vitamin D synthesis. (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of this‬

F
‭figure is available at‬‭Biology411.com‬‭.‬

‭ or additional information about bone health, click the link on the next page or scan the‬
F
‭QR code to watch the TED Talk by Jen Gunter‬‭titled‬‭“Why healthy bones are about so much‬
‭more than milk.”‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Jen Gunter: Why healthy bones are about so much‬‭more than milk |‬
‭TED Talk‬

‭15.11 Diseases and the Skeletal System‬

‭Osteoporosis‬

‭ steoporosis‬‭is a disease characterized by a decrease‬‭in bone mass that occurs when the‬
O
‭bone resorption rate exceeds the bone formation rate, a common occurrence as the body‬
‭ages. While osteoporosis can involve any bone, it most commonly affects the proximal‬
‭ends of the femur, vertebrae, and wrist. As a result of the loss of bone density, the osseous‬
‭tissue may not provide adequate support for everyday functions, and something as simple‬
‭as a sneeze can cause a vertebral fracture. When an older adult falls and breaks a hip‬
‭(really, the femur), it is very likely that the femur broke first, resulting in the fall.‬
‭Histologically, osteoporosis is characterized by a reduction in the thickness of compact‬
‭bone and the number and size of trabeculae in cancellous bone.‬

‭ igure 15.25‬‭shows that women lose bone mass more‬‭quickly than men starting at about‬
F
‭50 years of age. This occurs because 50 is the approximate age at which women go‬
‭through menopause. Not only do their menstrual periods lessen and eventually cease, but‬
‭their ovaries reduce in size and then cease the production of estrogen, a hormone that‬
‭promotes osteoblastic activity and the production of bone matrix. Thus, osteoporosis is‬
‭more common in women than men, but men can also develop it. Anyone with a family‬
‭history of osteoporosis has a greater risk of developing the disease, so the best treatment‬
‭is prevention, which should start with a childhood diet that includes adequate calcium and‬
‭vitamin D intake and a lifestyle that includes weight-bearing exercise. These actions, as‬
‭discussed above, are essential in building bone mass. Promoting proper nutrition and‬
‭weight-bearing exercise early in life can maximize bone mass before age 30, thus reducing‬
‭the risk of osteoporosis.‬

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‭ igure 15.25‬ ‭A graph showing the relationship between‬‭age and bone mass. Bone density‬
F
‭peaks at about 30 years of age in both sexes. However, as time progresses, women lose‬
‭bone mass more rapidly than men. (credit:‬‭OpenStax‬‭)‬

‭ or many older adults, a hip fracture can be life-threatening. The fracture itself may not be‬
F
‭serious, but the immobility that comes during the healing process can lead to a variety of‬
‭health consequences: formation of blood clots that can lodge in the capillaries of the‬
‭lungs, resulting in respiratory failure; pneumonia due to the lack of poor air exchange that‬
‭accompanies immobility; pressure sores (bed sores) that allow pathogens to enter the‬
‭body and cause infections; and urinary tract infections from catheterization.‬

‭ urrent treatments for osteoporosis include bisphosphonates, calcitonin, and estrogen‬

C
‭(for women only). Minimizing the risk of falls, for example, by removing tripping hazards,‬
‭is also an important step in managing the disease's potential outcomes.‬

‭Osteogenesis Imperfecta‬

‭ steogenesis imperfecta‬‭(‬‭OI‬‭) is a genetic disease‬‭in which bones do not form properly,‬

O
‭becoming fragile and breaking easily. It is also called brittle bone disease. The disease is‬
‭present from birth and affects a person throughout life.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ he genetic mutation that causes OI affects the body’s production of collagen, one of the‬
T
‭critical components of the bone matrix. The severity of the disease can range from mild to‬
‭severe. Those with the most severe forms of the disease sustain many more fractures than‬
‭those with a mild form. Frequent and multiple fractures typically lead to bone deformities‬
‭and short stature. Bowing of the long bones and curvature of the spine are also common‬
‭in people afflicted with OI. The curvature of the spine makes breathing difficult because‬
‭the lungs are compressed.‬

‭ ecause collagen is an essential structural protein in many body parts, people with OI may‬
B
‭also experience fragile skin, weak muscles, loose joints, easy bruising, frequent‬
‭nosebleeds, brittle teeth, blue sclera, and hearing loss. There is no known cure for OI, so‬
‭treatment focuses on helping the person retain as much independence as possible while‬
‭minimizing fractures and maximizing mobility. Safe exercises (e.g., swimming), where the‬
‭body is less likely to experience collisions or compressive forces, are recommended.‬
‭Braces to support legs, ankles, knees, and wrists are used as needed. Canes, walkers, or‬
‭wheelchairs can also help compensate for weaknesses.‬

‭15.12 Career Connection - Rheumatologist‬

‭ heumatologists are medical doctors who specialize in diagnosing and treating disorders‬
R
‭o f the joints, muscles, and bones. They diagnose and treat arthritis, musculoskeletal‬
‭disorders, osteoporosis, and autoimmune diseases such as ankylosing spondylitis and‬
‭rheumatoid arthritis.‬

‭ heumatoid arthritis (RA) is an inflammatory disorder that primarily affects the synovial‬
R
‭joints of the hands, feet, and cervical spine. Affected joints become swollen, stiff, and‬
‭painful. Although it is known that RA is an autoimmune disease in which the body’s‬
‭immune system mistakenly attacks healthy tissue, the cause of RA remains unknown.‬
‭Immune cells from the blood enter joints and the synovium, causing cartilage breakdown,‬
‭swelling, and joint lining inflammation. The breakdown of cartilage causes bones to rub‬
‭against each other, causing pain. RA is more common in women than men, and the age of‬
‭o nset is usually 40–50.‬

‭ heumatologists can diagnose RA based on symptoms such as joint inflammation and‬

R
‭pain, X-ray and MRI imaging, and blood tests. Arthrography is a type of medical imaging of‬
‭joints that uses a contrast agent, such as a dye, that is opaque to X-rays. This allows the‬
‭soft tissue structures of joints—such as cartilage, tendons, and ligaments—to be‬
‭visualized. An arthrogram differs from a regular X-ray by showing the surface of soft‬
‭tissues lining the joint in addition to joint bones. An arthrogram allows early degenerative‬
‭changes in joint cartilage to be detected before bones become affected.‬

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‭ here is currently no cure for RA; however, rheumatologists have several treatment‬
T
‭o ptions. Early stages can be treated with the rest of the affected joints using a cane or joint‬
‭splints that minimize inflammation. When inflammation has decreased, exercise can be‬
‭used to strengthen the muscles that surround the joint and to maintain joint flexibility. If‬
‭joint damage is more extensive, medications such as aspirin, topical pain relievers, and‬
‭corticosteroid injections may relieve pain and decrease inflammation. Surgery may be‬
‭required in cases in which joint damage is severe.‬

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‭Chapter 16: Muscular System and Movement‬

‭ igure 16.1‬ ‭Athletes like this tennis player rely‬‭o n toned skeletal muscles that supply the‬
F
‭force required for movement. (credit: Emmanuel Huybrechts; Flickr;‬‭OpenStax‬‭)‬

‭16.1 Introduction‬
‭ he muscular and skeletal systems support the body and allow for a wide range of‬
T
‭movement. The bones of the skeletal system protect the body’s internal organs and‬
‭support the body's weight. The muscles of the muscular system contract and pull on the‬
‭bones, allowing for movements as diverse as standing, walking, running, and grasping‬
‭items.‬
‭ s discussed in Chapter 4,‬‭muscle‬‭is one of the four‬‭primary tissue types of the body‬
A
‭(muscle, epithelial, connective, and neural), and the body contains three types of muscle‬
‭tissue: cardiac, smooth, and skeletal.‬ ‭Cardiac muscle‬‭tissue in the heart functions to‬
‭pump blood through the circulatory system.‬‭Smooth‬‭muscle‬‭tissue is responsible for‬
‭various involuntary movements, such as moving food through the digestive system (e.g.,‬
‭peristalsis). However, when most people think of muscles, they think of the‬‭skeletal‬
‭muscles‬‭because most of them move the skeleton. In‬‭this chapter, we will focus on skeletal‬
‭muscle structure and function.‬
‭ or an introduction to various aspects of this system, click the link on the next page or‬
F
‭scan the QR code to watch the TED-Ed video by Emma Bryce titled “How your muscular‬
‭system works.”‬

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How your muscular system works - Emma Bry…

‭16.2 Skeletal Muscle Function and Structure‬

S‭ keletal muscle tissue forms skeletal muscles, which attach to bones and control‬
‭locomotion. Because it can be controlled by thought, skeletal muscle is also called‬
‭voluntary muscle‬‭. Skeletal muscles depend on signaling‬‭from the nervous system to‬
‭function correctly; unlike cardiac and smooth muscles, the only way to functionally‬
‭contract a skeletal muscle is through signaling from the nervous system. Related aspects‬
‭will be discussed in this chapter and Chapter 17.‬
S‭ keletal muscles act to produce movement and stop movement, such as resisting gravity to‬
‭maintain posture. Minor, constant adjustments of the skeletal muscles are needed to hold‬
‭a body upright or balanced in any position. Muscles also prevent excess movement of the‬
‭bones and joints, maintaining skeletal stability and preventing skeletal structure damage‬
‭o r deformation. Joints can become misaligned or dislocated entirely by pulling on the‬
‭associated bones; muscles work to keep these joints stable. Skeletal muscles are located‬
‭throughout the body at the openings of internal tracts to control the movement of various‬
‭substances. These muscles allow functions, such as swallowing, urination, and defecation,‬
‭to be under voluntary control. Skeletal muscles also protect internal organs (particularly‬
‭abdominal and pelvic organs) by acting as a shield against internal trauma and by‬
‭supporting the weight of the organs.‬
S‭ keletal muscles generate heat to maintain body temperature homeostasis. Muscle‬
‭contraction requires energy; heat is released when ATP is broken down. This heat is‬
‭noticeable during exercise when sustained muscle movement causes body temperature to‬
‭rise. Shivering produces random skeletal muscle contractions in extreme cold to generate‬
‭heat.‬
‭ ach skeletal muscle is an organ that consists of various integrated tissues. These tissues‬
E
‭include the skeletal muscle fibers, blood vessels, nerve fibers, and connective tissue. Each‬
‭skeletal muscle has three layers of connective tissue that enclose it, provide structure to‬
‭the muscle as a whole, and compartmentalize the muscle fibers within the muscle (‬‭Figure‬
‭16.2‬‭). Each muscle is wrapped in a sheath of dense,‬‭irregular connective tissue called the‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ pimysium‬‭, which allows a muscle to contract and move powerfully while maintaining its‬
e
‭structural integrity. The epimysium also separates muscle from other tissues and organs in‬
‭the area, allowing the muscle to move independently.‬
I‭ nside each skeletal muscle, muscle fibers are organized into individual bundles, called‬
‭fascicles‬‭, by a middle layer of connective tissue called the perimysium. This fascicular‬
‭o rganization is typical in muscles of the limbs; it allows the nervous system to trigger a‬
‭specific movement of a muscle by activating a subset of muscle fibers within a bundle or‬
‭fascicle of the muscle. Each muscle fiber is encased inside a fascicle in a thin connective‬
‭tissue layer called the‬‭endomysium‬‭. The endomysium‬‭contains extracellular fluid and‬
‭nutrients to support the muscle fibers. These nutrients are supplied via blood to the‬
‭muscle tissue.‬

‭ igure 16.2‬ ‭Skeletal muscles comprise bundles of‬‭muscle fibers called fascicles, covered‬
F
‭by the perimysium. The endomysium covers muscle fibers. (credit:‬‭OpenStax‬‭)‬

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‭ lood vessels also supply every skeletal muscle richly for nourishment, oxygen delivery,‬
B
‭and waste removal. In addition, every muscle fiber in a skeletal muscle is in very close‬
‭proximity to a neuron, which signals the fiber to contract.‬

‭16.3 Skeletal Muscle Fiber Structure‬

‭ ach‬‭skeletal muscle fiber‬‭is a skeletal muscle cell. These cells are incredibly large, with‬
E
‭diameters of up to 0.004 inches and lengths of up to approximately 12 inches. The plasma‬
‭membrane of a skeletal muscle fiber is called the‬‭sarcolemma‬‭. The sarcolemma is the site‬
‭o f action potential (i.e., an electrical current) conduction, which triggers muscle‬
‭contraction. Within each muscle fiber are‬‭myofibrils‬‭—long‬‭cylindrical structures that lie‬
‭parallel to the fiber. Myofibrils run the entire length of the muscle fiber, and because they‬
‭are only approximately 0.00005 inches in diameter, hundreds to thousands can be found‬
‭inside one muscle fiber. They attach to the sarcolemma at their ends so that as myofibrils‬
‭shorten, the entire muscle cell contracts (‬‭Figure‬‭16.3‬‭).‬

‭ igure 16.3‬ ‭A plasma membrane surrounds a skeletal‬‭muscle cell called the sarcolemma‬
F
‭with a cytoplasm called the sarcoplasm. A muscle fiber is composed of many fibrils‬
‭packaged into orderly units. (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of this figure‬
‭is available at‬‭Biology411.com‬‭.‬

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‭ he‬‭sarcomere,‬‭the space between two consecutive‬‭Z lines‬‭(‬‭Figure 16.4‬‭), is the most‬

T
‭basic unit associated with contraction. The sarcomere is bundled within the myofibril that‬
‭runs the entire muscle fiber length and attaches to the sarcolemma at its end. A myofibril‬
‭is composed of many sarcomeres running along its length, and as the sarcomeres‬
‭individually contract, the myofibrils and muscle cells shorten.‬
‭ yofibrils are composed of smaller structures called‬‭myofilaments‬‭. These filaments have‬
M
‭two main types:‬‭thick filaments‬‭(composed mainly of‬‭the‬‭myosin‬‭protein) and‬‭thin‬
‭filaments‬‭(composed primarily of the‬‭actin‬‭protein); each has different compositions and‬
‭locations (‬‭Figure 16.5‬‭).‬

‭(a)‬ ‭(b)‬

‭ igure 16.4‬ ‭(a) A sarcomere is the region from one‬‭Z line to the following Z line. Many‬
F
‭sarcomeres are present in a myofibril, resulting in the striation pattern characteristic of‬
‭skeletal muscle. (b) A magnified sample of skeletal muscle showing striations. The fibers‬
‭are horizontally oriented, and the striations are vertically oriented. (credit: (a)‬‭OpenStax‬‭;‬
‭(b) Willy Cushwa). A link to a video explanation of this figure is available at‬
‭Biology411.com‬‭.‬

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‭16.4 Skeletal Muscle Contraction and Relaxation‬

‭ ll living cells have membrane potentials or electrical gradients across their membranes.‬
A
‭The inside of the membrane is usually around -60 to -90 millivolts (mV) relative to the‬
‭o utside (This concept will be explained in detail in Chapter 17.). This is referred to as a‬
‭cell’s‬‭membrane potential‬‭. Neurons and muscle cells‬‭can use their membrane potentials‬
‭to generate electrical signals. They do this by controlling the movement of‬‭ions‬‭(Chapter‬
‭2) across their membranes to create electrical currents. This is achieved by opening and‬
‭closing specialized proteins in the membrane called‬‭ion channels‬‭(Chapter 3). Although‬
‭the currents generated by ions moving through these channel proteins are minimal, they‬
‭are the basis for neural signaling and muscle contraction.‬

‭ igure 16.5‬ ‭A second simplified diagram of a sarcomere‬‭(the first one in Figure 16.4),‬
F
‭defined as the region between two Z lines. It is composed largely of regularly-spaced actin‬
‭and myosin contractile myofilaments. (credit: modification of‬ ‭Normal Sarcomere‬‭;‬
‭MecciaC0410‬‭;‬‭CC BY-SA 4.0‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ oth neurons and skeletal muscle cells are electrically excitable, meaning they can‬
B
‭generate action potentials. An‬‭action potential‬‭is‬‭an electrical signal that can travel along‬
‭a cell membrane as a wave. This allows a signal to be transmitted quickly and consistently‬
‭over long distances.‬

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‭ owever, how is an action potential generated and transmitted along a muscle fiber? The‬
H
‭process responsible for these events is‬‭excitation-contraction‬‭coupling‬‭, a potentially‬
‭intimidating term to some students! However, it comes down to this: for a skeletal muscle‬
‭fiber to contract, its membrane must first be “excited.” In other words, it must be‬
‭stimulated to “fire” (initiate) an action potential. The muscle fiber action potential, which‬
‭sweeps along the sarcolemma as a wave, is “coupled” to the actual contraction via the‬
‭release of calcium ions (Ca‬‭++‬‭) from the‬‭sarcoplasmic‬‭reticulum (SR)‬‭, which is a‬
‭specialized form of the endoplasmic reticulum discussed in Chapter 3. The effect of the‬
‭calcium ions will be discussed in more detail later in this section.‬
I‭ n skeletal muscle, the initiation of this sequence of events begins with input from the‬
‭nervous system at points called‬‭neuromuscular junctions‬‭(NMJ)‬‭, where neurons‬
‭associate or come into close physical contact with the membranes of skeletal muscles. In‬
‭o ther words, the “excitation” step in skeletal muscles is always triggered by signaling from‬
‭the nervous system to the skeletal muscle at these junctions.‬
‭ he neurons that tell the skeletal muscle fibers to contract originate in the spinal cord,‬
T
‭with a smaller number located in the brainstem for activating the skeletal muscles of the‬
‭face, head, and neck. These neurons have long processes called‬‭axons‬‭, which are‬
‭specialized to transmit action potentials long distances—in this case, from the spinal cord‬
‭to the muscle (which may be up to three feet away). The axons of multiple neurons bundle‬
‭together to form nerves, like wires bundled together in a cable.‬
S‭ ignaling begins when an action potential travels along the axon of a type of neuron and‬
‭then along the individual branches to terminate at the NMJ (‬‭Figure 16.6‬‭). At the NMJ, the‬
‭axon terminal releases a chemical messenger, or‬‭neurotransmitter‬‭,‬‭called‬‭acetylcholine‬
‭(‬‭ACh‬‭) via‬‭exocytosis‬‭(Chapter 3). The ACh molecules‬‭diffuse‬‭across a tiny space called the‬
‭synaptic cleft‬‭and bind to ACh receptors located on‬‭sarcolemma on the other side of the‬
‭synapse. Once ACh binds, a channel in the ACh receptor opens, and positively charged ions‬
‭can pass through into the muscle fiber, causing it to‬‭depolarize‬‭, meaning that the‬
‭membrane potential of the muscle fiber becomes less negative (i.e., closer to zero.)‬
‭ s the membrane depolarizes, another set of ion channels,‬‭voltage-gated sodium‬
A
‭c hannels‬‭, are triggered to open. Sodium ions enter‬‭the muscle fiber, and an action‬
‭potential rapidly spreads (or “fires”) along the entire membrane to initiate‬
‭excitation-contraction coupling.‬

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‭ igure 16.6‬ ‭At the neuromuscular junction, vesicles‬‭(Chapter 3) that contain a chemical‬
F
‭called a neurotransmitter (e.g., acetylcholine; ACh) are released from the end of the axon.‬
‭(credit:‬‭OpenStax‬‭); A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

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‭ hings happen very quickly in the world of excitable membranes; just think about how‬
T
‭quickly you can snap your fingers when you decide to do it. Immediately after‬
‭depolarization, the membrane‬‭repolarizes‬‭(meaning‬‭the ions are moved to their original‬
‭positions), re-establishing the negative membrane potential. Meanwhile, the ACh in the‬
‭synaptic cleft is degraded by the enzyme‬‭acetylcholinesterase‬‭(AChE)‬‭so that the ACh‬
‭cannot rebind to a receptor and reopen its channel, which would cause unwanted‬
‭extended muscle excitation and contraction.‬
‭ s previously stated, the movement of an action potential along the sarcolemma triggers‬
A
‭the release of calcium ions (Ca‬‭+2‬‭) from their storage‬‭in the cell’s SR. For the action‬
‭potential to reach the membrane of the SR, there are periodic infoldings in the‬
‭sarcolemma called‬‭T-tubules‬‭(“T” stands for “transverse”).‬‭These T-tubules ensure the‬
‭membrane can get close to the SR in the sarcoplasm. The arrangement of a T-tubule with‬
‭the membranes of SR on either side ensures that the action potential will trigger the‬
‭release of calcium ions in the myofibrils, which contain actin and myosin filaments (‬‭Figure‬
‭16.7‬‭).‬

‭ igure 16.7‬ ‭Narrow T-tubules (transverse tubules)‬‭permit the conduction of electrical‬

F
‭impulses (action potentials) to the sarcoplasmic reticulum (SR), which causes the release‬
‭o f the calcium ions needed to bring about muscle contraction. (credit:‬‭OpenStax‬‭). A link‬
‭to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

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‭ he T-tubules carry the action potential into the cell's interior, which triggers the opening‬
T
‭o f calcium channels in the membrane of the adjacent SR. This causes Ca‬‭++‬ ‭to diffuse from‬
‭the SR and into the sarcoplasm. The arrival of Ca‬‭++‬ ‭in the sarcoplasm initiates the‬
‭contraction of the muscle fiber by its contractile units (i.e., sarcomeres).‬
‭ efore continuing this discussion, it is advantageous to stop and think for a moment about‬
B
‭the structure of the sarcomere, the relative position of actin (thin) and myosin (thick)‬
‭myofilaments, the Z lines, and how these change during muscle contraction. The name of‬
‭the contraction mechanism, the‬‭sliding filament model‬‭,‬‭explains these changes. The‬
‭sarcomere must shorten for a muscle cell to contract, meaning the distance between Z‬
‭lines must decrease. However, thick and thin filaments—the components of‬
‭sarcomeres—do not shorten. Instead, they slide by one another, by a mechanism‬
‭discussed later in the chapter, causing the sarcomere to shorten while the filaments‬
‭remain the same length (‬‭Figure 16.8‬‭).‬
‭ reasonable question is, what causes the actin filaments to slide over the myosin‬
A
‭filaments during contraction? The answer is that the structural features of myosin fiber,‬
‭called myosin heads, bind to unique sites on the actin fibers to form‬‭c ross bridges‬‭. The‬
‭movement of the myosin heads, discussed later in the section, is responsible for sliding the‬
‭actin filaments and the subsequent shortening of the sarcomeres (‬‭Figure 16.9‬‭).‬

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‭ igure 16.8‬ ‭When (a) a sarcomere (b) contracts,‬‭the Z lines move closer together, and‬
F
‭the sarcomere length decreases due to the actin filaments sliding along the myosin‬
‭filaments. The mechanism of this movement will be explained in the text and other figures.‬
‭(credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

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‭ igure 16.9‬ ‭(a) A sarcomere is the distance between‬‭two Z lines. (a) A relaxed‬
F
‭sarcomere shows the relative positions of the thin (actin) and thick (myosin) filaments.‬
‭(b) A contracted sarcomere showing the Z lines moved closer together because the‬
‭myosin heads create cross bridges with the actin filaments and pull them towards the‬
‭center. At full contraction, the thin and thick filaments overlap entirely. (credit:‬
‭modification of an‬‭OpenStax‬‭figure)‬

‭ n important question to ask at this point is, “How does the release of calcium ions bring‬
A
‭about muscle contraction?”. When a muscle fiber rests, actin and myosin are separated‬
‭because of two regulatory proteins, tropomyosin and troponin.‬ ‭Tropomyosin‬‭blocks the‬
‭myosin binding sites on actin filaments, so cross bridges can’t form. To enable muscle‬
‭contraction, tropomyosin must change conformation, uncovering the myosin-binding site‬
‭o n an actin molecule. This can only happen in the presence of calcium ions, which are kept‬
‭at extremely low concentrations in the sarcoplasm. If present, calcium ions bind to‬
‭troponin‬‭, causing conformational changes in troponin‬‭that allow tropomyosin to move‬
‭away from the myosin binding sites on actin. Once the position of tropomyosin shifts,‬
‭cross bridges can form between actin and myosin, triggering contraction by the sliding‬
‭filament model (‬‭Figures 16.10‬‭and‬‭16.12‬‭).‬

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‭ igure 16.10‬ ‭During muscle fiber contraction, cross-bridges‬‭form between actin‬

F
‭filaments and the myosin heads. As long as Ca‬‭+2‬ ‭ions‬‭remain in the sarcoplasm to bind to‬
‭troponin and ATP is available, the muscle fiber will continue to shorten. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ uscle relaxation‬‭results when excitation from the‬‭neuron ceases and calcium ions are‬
M
‭pumped back into the SR, thus reducing the concentration in the sarcoplasm. Troponin‬
‭changes conformation and causes tropomyosin to cover the binding sites on actin again‬
‭and prevent the formation of cross bridges. These actions result in the actin filaments‬

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s‭ liding back over the myosin filaments and increasing the length of the sarcomere (‬‭Figure‬
‭16.11‬‭).‬

‭ igure 16.11‬ ‭During muscle fiber relaxation, Ca‬‭+2‬ ‭ions are pumped back into the SR,‬
F
‭which causes the tropomyosin to shield the binding sites on the actin strands. A muscle‬
‭may also stop contracting when it runs out of ATP and becomes fatigued. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ uring contraction, the myosin heads require ATP to form crossing bridges and move the‬
D
‭actin filaments in what is referred to as the‬‭power‬‭stroke‬‭event. Full contraction is not‬
‭achieved in one step but via a series of repeated steps called the‬‭c ross-bridge muscle‬
‭contraction cycle‬‭. The myosin head can only pull‬‭the actin filament a very short distance‬

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‭ efore it has reached its limit and must be reset before it can pull again, a step that‬
b
‭requires ATP. A visual analogy might be helpful. The motion of the myosin heads is‬
‭similar to the oars when an individual rows a boat: The paddle of the oars (the myosin‬
‭heads) pull, are lifted from the water (detach), repositioned (re-cocked) and then‬
‭immersed again to pull. Each cycle requires energy, and the action of the myosin heads in‬
‭the sarcomeres repetitively pulling on the thin filaments also requires energy, which ATP‬
‭provides.‬ ‭Figure 16.12‬‭shows the associated steps‬‭o f the cycle and how ATP is needed to‬
‭continue the power strokes and move the actin filaments.‬

‭ igure 16.12‬ ‭The cross-bridge muscle contraction‬‭cycle is triggered by Ca‬‭2+‬ ‭binding to‬
F
‭the actin active site. With each contraction cycle, actin moves relative to myosin. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

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‭ s these are complex concepts, it will be helpful to “put the pieces together” in a brief‬
A
‭review of skeletal muscle contraction.‬‭Figure 16.13‬‭summarizes the steps of initiating an‬
‭action potential on the sarcolemma, propagating the action potential, releasing calcium‬
‭ions from the SR, contracting, and relaxing.‬
‭ ach thick filament of roughly 300 myosin molecules has multiple myosin heads, and many‬
E
‭cross-bridges form and break continuously during muscle contraction. Multiply this by all‬
‭o f the sarcomeres in one myofibril, all the myofibrils in one muscle fiber, and all of the‬
‭muscle fibers in one skeletal muscle, and you can understand why so much energy (ATP)‬
‭is needed to keep skeletal muscles working. In fact, the loss of ATP results in the rigor‬
‭mortis observed soon after someone dies. With no further ATP production possible, there‬
‭is no ATP available for myosin heads to detach from the actin-binding sites, so the‬
‭cross-bridges stay in place, causing rigidity in the skeletal muscles.‬
‭ uscle contraction can generate differing amounts of force. For example, the force‬
M
‭required to pick up a pencil using the biceps muscle is substantially less than that needed‬
‭for a textbook. The primary variable determining force production is the number of‬
‭myofibers within the muscle that receive an action potential from the neuron that controls‬
‭that fiber. When lifting a light object (e.g., a pencil), the brain's motor cortex only signals a‬
‭few neurons of the biceps, and only a few myofibers respond. In vertebrates, each‬
‭myofiber responds fully if stimulated. When picking up a heavier object (e.g., a textbook),‬
‭the motor cortex signals more neurons in the biceps, and more myofibers participate. If‬
‭the maximum force required to lift an object (e.g., a car) exceeds the force generated by‬
‭the muscle fully contracting, the object won’t move.‬
‭ he number of skeletal muscle fibers in a given muscle is genetically determined and does‬
T
‭not change. Muscle strength is directly related to the amount of myofibrils and sarcomeres‬
‭within each fiber. Factors such as hormones and stress (and artificial anabolic steroids)‬
‭acting on the muscle can increase the production of sarcomeres and myofibrils within the‬
‭muscle fibers, a change called‬‭hypertrophy‬‭, which‬‭results in increased mass and bulk in a‬
‭skeletal muscle. Likewise, decreased use of skeletal muscle results in‬‭atrophy‬‭, where the‬
‭number of sarcomeres and myofibrils disappear (but not the number of muscle fibers). It‬
‭is common for a limb in a cast to show atrophied muscles when the cast is removed, and‬
‭certain diseases, such as polio, show atrophied muscles.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 16.13‬ ‭Summary of the steps of muscle contraction‬‭and relaxation. (credit:‬

F
‭OpenStax‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ or information about these topics, click the link below or scan the QR code to watch the‬
F
‭TED-Ed video by Jeffrey Seigel titled “What makes muscles grow?”‬

What makes muscles grow? - Jeffrey Siegel

‭16.5 Sources of ATP for Muscle Contraction‬

‭ TP supplies the energy for muscle contraction. In addition to its direct role in the‬
A
‭cross-bridge cycle, ATP provides energy for the active-transport Ca‬‭++‬ ‭pumps in the SR. The‬
‭amount of ATP stored in muscle is very low, only enough to power a few seconds of‬
‭contractions. As ATP is broken down, it must be regenerated and replaced quickly for‬
‭sustained contraction. There are three mechanisms by which ATP can be regenerated:‬
‭creatine phosphate metabolism, anaerobic metabolism (lactic acid fermentation), and‬
‭aerobic/cellular respiration.‬

‭ reatine phosphate‬‭is a molecule that can store energy‬‭in its phosphate bonds. Excess‬
C
‭ATP in a resting muscle transfers energy to creatine, producing ADP and creatine‬
‭phosphate. This acts as an energy reserve that can quickly create more ATP. When the‬
‭muscle starts to contract and needs energy, creatine phosphate transfers its phosphate‬
‭back to ADP to form ATP and creatine. The enzyme creatine kinase catalyzes this reaction‬
‭and occurs quickly (‬‭Figure 16.14a‬‭); thus, creatine‬‭phosphate-derived ATP powers the‬
‭first few seconds of muscle contraction. However, creatine phosphate can only provide‬
‭approximately 15 seconds of energy, at which point another energy source must be used.‬
‭ s the ATP produced by creatine phosphate is depleted, muscles turn to glycolysis as an‬
A
‭ATP source.‬‭Glycolysis‬‭(Chapter 6) is an anaerobic‬‭(non-oxygen-dependent) process that‬
‭breaks down glucose to produce ATP; however, glycolysis cannot generate ATP as quickly‬
‭as creatine phosphate. Thus, the switch to glycolysis results in a slower rate of ATP‬
‭availability to the muscle. The glucose used in glycolysis can be provided by blood glucose‬
‭o r metabolizing glycogen stored in the muscle. The glycolytic breakdown of one glucose‬
‭molecule produces two ATP and two molecules of pyruvic acid/pyruvate (‬‭Figure 16.14b‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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I‭ f oxygen is available, pyruvate is used in‬‭aerobic/cellular respiration‬‭. However, if‬

‭oxygen is unavailable, pyruvate is converted to lactic acid via‬‭fermentation‬‭(anaerobic‬
‭metabolism). This conversion allows the recycling of the enzyme NAD‬‭+‬ ‭from NADH, which‬
‭is needed for glycolysis to continue. This occurs during strenuous exercise when high‬
‭energy is required, but oxygen cannot be sufficiently delivered to muscles. Glycolysis‬
‭cannot be sustained for very long (approximately 1 minute of muscle activity), but it helps‬
‭facilitate short bursts of high-intensity output. This is because glycolysis does not utilize‬
‭glucose very efficiently, producing a net gain of only two ATPs per glucose molecule; also,‬
‭the end product of lactic acid may contribute to muscle fatigue as it accumulates.‬
‭ erobic/cellular respiration is the breakdown of glucose or other nutrients in the‬
A
‭presence of oxygen to produce carbon dioxide, water, and ATP (Chapter 6). Approximately‬
‭95 percent of the ATP required for resting or moderately active muscles is provided by‬
‭aerobic respiration, which takes place in mitochondria. The inputs for aerobic respiration‬
‭include glucose circulating in the bloodstream, pyruvic acid, and fatty acids. Aerobic‬
‭respiration is much more efficient than anaerobic glycolysis, producing a net yield of‬
‭approximately 36 ATPs per glucose molecule. However, aerobic respiration cannot be‬
‭sustained without a steady oxygen supply to the skeletal muscle and is much slower‬
‭(‬‭Figure 16.14c‬‭). To compensate, muscles store a small‬‭amount of excess oxygen in‬
‭proteins called‬‭myoglobin‬‭, allowing for more efficient‬‭muscle contractions and less‬
‭fatigue. Aerobic training also increases the efficiency of the circulatory system so that‬
‭oxygen can be supplied to the muscles for extended periods.‬
‭ or more information about muscle structure, energy considerations, and fatigue, click the‬
F
‭link below or scan the QR code to watch the TED-Ed video by Christian Morro titled “The‬
‭surprising reason our muscles get tired.”‬

The surprising reason our muscles get tired - Christian Moro

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 16.14‬ ‭A summary of muscle metabolism (a)‬‭Some ATP is stored in a resting‬

F
‭muscle. As contraction starts, it is used up in seconds. More ATP is generated from‬
‭creatine phosphate for about 15 seconds. (b) Each glucose molecule produces two ATP‬
‭and two molecules of pyruvic acid, which can be used in aerobic respiration or converted‬
‭to lactic acid. If oxygen is unavailable, pyruvic acid is converted to lactic acid, which may‬
‭contribute to muscle fatigue. This occurs during strenuous exercise when large amounts‬
‭o f energy are needed, but oxygen cannot be sufficiently delivered to muscle. (c) Aerobic‬
‭respiration is the breakdown of glucose in the presence of oxygen (O‬‭2‭)‬ to produce carbon‬
‭dioxide, water, and ATP. Approximately 95 percent of the ATP required for resting or‬
‭moderately active muscles is provided by aerobic respiration, which takes place in‬
‭mitochondria. (credit:‬‭OpenStax‬‭); A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭16.6 Performance-Enhancing Substances‬

S‭ ome athletes attempt to boost their performance by using various agents that may‬
‭enhance muscle performance. Anabolic steroids are one of the more widely known agents‬
‭used to boost muscle mass and increase power output.‬‭Anabolic steroids‬‭are a form of‬
‭testosterone‬‭, a male sex hormone that stimulates muscle‬‭formation and increases muscle‬
‭mass. For more information about steroids and muscle mass, click the link below or scan‬
‭the QR code to watch the TED-Ed video by Anees Bahji titled “How do steroids affect your‬
‭muscles - and the rest of your body?”.‬

How do steroids affect your muscles— and the rest of your bod…

‭ ndurance athletes may also try to boost oxygen availability to muscles to increase‬
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‭aerobic respiration by using substances such as‬‭erythropoietin‬‭(‬‭EPO‬‭), a hormone usually‬
‭produced in the kidneys, which triggers the production of red blood cells. Muscles can‬
‭then use the extra oxygen these blood cells carry for aerobic respiration.‬‭Human growth‬
‭hormone‬‭(‬‭hGH‬‭) is another supplement. Although it can‬‭facilitate building muscle mass, its‬
‭primary role is to promote the healing of muscle and other tissues after strenuous‬
‭exercise. Increased hGH may allow for faster recovery after muscle damage, reducing the‬
‭rest required after exercise and allowing for more sustained high-level performance.‬
‭ lthough performance-enhancing substances often improve performance, most are‬
A
‭banned by governing bodies in sports and are illegal for non-medical purposes. Their use‬
‭to enhance performance raises ethical cheating issues because they give users an unfair‬
‭advantage over non-users. A more significant concern, however, is that their use carries‬
‭serious health risks. The side effects of these substances are often substantial,‬
‭nonreversible, and sometimes fatal. The physiological strain caused by these substances is‬
‭o ften greater than what the body can handle, leading to unpredictable and dangerous‬
‭effects. Anabolic steroid use has been linked to infertility, aggressive behavior,‬
‭cardiovascular disease, and brain cancer.‬
S‭ imilarly, some athletes have used creatine to increase power output. Creatine phosphate‬
‭provides quick bursts of ATP to muscles in the initial stages of contraction. Increasing the‬
‭amount of creatine available to cells is thought to produce more ATP and increase‬
‭explosive power output, although its effectiveness as a supplement has been questioned.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭16.7 Disorder of the Muscular System - Duchenne Muscular Dystrophy‬

‭ uchenne muscular dystrophy‬‭(‬‭DMD‬‭) is a progressive‬‭weakening of the skeletal‬

D
‭muscles, and is one of several diseases collectively referred to as “muscular dystrophy.”‬
‭DMD is caused by a lack of the protein‬‭dystrophin‬‭,‬‭which helps the thin filaments of‬
‭myofibrils bind to the sarcolemma. Without sufficient dystrophin, muscle contractions‬
‭cause the sarcolemma to tear, resulting in an influx of Ca‬‭+2‬‭, which leads to cellular damage‬
‭and muscle fiber degradation. Over time, as muscle damage accumulates, muscle mass is‬
‭lost, and more significant functional impairments develop.‬

‭ MD is an inherited disorder caused by an abnormal gene on the X chromosome. It‬

D
‭primarily affects males, and it is usually diagnosed in early childhood. DMD usually first‬
‭appears as difficulty with balance and motion and then progresses to an inability to walk.‬
‭It continues progressing upward in the body from the lower extremities to the upper‬
‭body, where it affects the muscles responsible for breathing and circulation. It ultimately‬
‭causes death due to respiratory failure, and those afflicted do not usually live past their‬
‭20s.‬

‭ ecause DMD is caused by a mutation in the gene that codes for dystrophin, it was‬
B
‭thought that introducing healthy myoblasts into patients might be an effective treatment.‬
‭Myoblasts are the embryonic cells responsible for muscle development, and ideally, they‬
‭carry healthy genes that could produce the dystrophin needed for normal muscle‬
‭contraction. This approach has been largely unsuccessful in humans. A recent strategy has‬
‭involved attempting to boost the muscle’s production of utrophin, a protein similar to‬
‭dystrophin that may be able to assume the role of dystrophin and prevent cellular damage‬
‭from occurring.‬

‭16.8 Career Connection - Physical Therapist‬

‭ s muscle cells die, they are not regenerated but instead are replaced by connective tissue‬
A
‭and adipose tissue, which do not possess contractile abilities. Muscles atrophy when not‬
‭used, and muscle cells die over time if atrophy is prolonged. Therefore, those susceptible‬
‭to muscle atrophy must exercise to maintain muscle function and prevent the complete‬
‭loss of muscle tissue. In extreme cases, electrical stimulation can be introduced to a‬
‭muscle from an external source when movement is impossible. This is a substitute for‬
‭endogenous neural stimulation, stimulating the muscle to contract and preventing the loss‬
‭o f proteins that occurs with a lack of use.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ hysical therapists work with patients to maintain muscles. They are trained to target‬
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‭muscles susceptible to atrophy and to prescribe and monitor exercises designed to‬
‭stimulate those muscles. Various causes of atrophy include mechanical injury, disease, and‬
‭age. Muscle use is impaired after breaking a limb or undergoing surgery and can lead to‬
‭disuse atrophy. If the muscles are not exercised, this atrophy can lead to long-term muscle‬
‭weakness. A stroke can also cause muscle impairment by interrupting neural stimulation‬
‭to specific muscles. Without neural inputs, these muscles do not contract and thus begin to‬
‭lose structural integrity. Exercising these muscles can help to restore muscle function and‬
‭minimize functional impairments. Age-related muscle loss is also a target of physical‬
‭therapy, as exercise can reduce the effects of age-related atrophy and improve muscle‬
‭function.‬

‭ he goal of a physical therapist is to improve physical functioning and reduce functional‬

T
‭impairments; this is achieved by understanding the cause of muscle impairment and‬
‭assessing a patient's capabilities, after which a program to enhance these capabilities is‬
‭designed. Some assessed factors include strength, balance, and endurance, which are‬
‭continually monitored as exercises are introduced to track improvements in muscle‬
‭function. Physical therapists can also instruct patients on properly using equipment, such‬
‭as crutches, and assess whether someone has sufficient strength to use the equipment‬
‭and when they can function without it.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 17.1‬ ‭An athlete’s nervous system is hard‬‭at work during the planning and‬
F
‭execution of a movement as precise as a high jump. Parts of the nervous system are‬
‭involved in determining how hard to push off and when to turn and controlling the‬
‭muscles throughout the body that make this complicated movement possible without‬
‭knocking the bar down—all in just a few seconds. (credit:‬‭Jumping at the Military World‬
‭Games.jpg‬‭; Shane T. McCoy; Wikimedia Commons;‬‭CC0‬‭1.0‬‭)‬

‭17.1 Introduction‬
‭ he nervous system is a very complex organ system. In Peter D. Kramer’s book‬‭Listening‬
T
‭to Prozac‬‭, a pharmaceutical researcher says, “If the‬‭human brain were simple enough for‬
‭us to understand, we would be too simple to understand it” (1994). That quote is from the‬
‭early 1990s; in the decades since progress has continued at an amazing rate within the‬
‭scientific disciplines of neuroscience. It is interesting to consider that the complexity of the‬
‭nervous system may be too complex for it (that is, for us) to unravel completely. But our‬
‭current level of understanding is probably nowhere close to that limit.‬
‭ he picture you have in your mind of the nervous system likely includes the‬‭brain‬‭, the‬
T
‭nervous tissue within the skull, and the‬‭spinal cord‬‭,‬‭the extension of nervous tissue‬
‭within the vertebral column. That suggests it is made of two organs—and you may not‬
‭even think of the spinal cord as an organ—but the nervous system is very complex. Many‬
‭different and separate regions are responsible for many distinct and individual functions‬
‭within the brain. It is as if the nervous system is composed of many organs that appear‬
‭similar and can only be differentiated using tools such as the microscope or‬
‭electrophysiology. In comparison, it is easy to see that the stomach differs from the‬

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‭Chapter 17: Nervous System‬

e‭ sophagus or the liver, so you can imagine the digestive system as a collection of specific‬
‭o rgans.‬

‭17.2 Nervous System Categorization‬

‭ he nervous system can be categorized in various ways, depending on the basis used (e.g.,‬
T
‭anatomy, physiology/function, the direction of transport of the nerve messages, and the‬
‭basis for control of the body). However, there is some overlap between the different‬
‭methods. The anatomical divisions are the central and peripheral nervous systems‬
‭(‬‭Figure 17.2‬‭). The‬‭central nervous system‬‭(‬‭CNS‬‭) consists‬‭o f the brain and spinal cord,‬
‭protected by the skull and vertebral column. The CNS receives sensory information,‬
‭integrates this information, and initiates events that lead to a response. The brain is the‬
‭control center for processing sensory information and directing responses.‬
‭ he‬‭peripheral nervous system‬‭(‬‭PNS‬‭) contains all other‬‭elements of this organ system‬
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‭and is so named because it is on the periphery—meaning beyond the brain and spinal‬
‭cord.‬

‭Figure 17.2‬ ‭Anatomical categorization of the nervous‬‭system. (credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ he structures of the PNS are referred to as ganglia (a collection of nerve cell bodies) and‬
T
‭nerves (cells that conduct electrical impulses), which can be seen as distinct structures.‬
‭The equivalent structures in the CNS are not apparent from this overall perspective and‬
‭are best examined in prepared tissue under the microscope.‬
‭ he CNS is the power plant of the nervous system. It creates signals that control the‬
T
‭functions of the body. Without those wires, the CNS could not control the body (and the‬
‭CNS would not be able to receive sensory information from the body.‬
‭ he PNS is the connection between the central nervous system and the rest of the body‬
T
‭and it can be further categorized based on the direction of the impulse conduction. The‬
‭sensory division‬‭conducts impulses from receptors‬‭to the CNS. In contrast, the‬‭motor‬
‭division‬‭conducts impulses from the CNS to the effectors,‬‭which are the muscles or glands‬
‭that will respond.‬
‭ he first physiological (or functional) division of the nervous system is associated with the‬
T
‭tasks of receiving information about the environment around us (‬‭sensation‬‭), integrating‬
‭and processing the information (‬‭integration‬‭), and‬‭generating responses to that‬
‭information (‬‭motor responses‬‭):‬
‭ ensation‬‭: The first primary function of the nervous‬‭system is sensation—receiving‬
S
‭information about the environment to gain input about what is happening outside the‬
‭body or, sometimes, within the body. The sensory functions of the nervous system register‬
‭the presence of a change from homeostasis or a particular event in the environment,‬
‭known as a stimulus. The senses we think of most are the “big five”: taste, smell, touch,‬
‭sight, and hearing. The stimuli for taste and smell are both chemical substances‬
‭(molecules, compounds, ions, etc.), touch is physical or mechanical stimuli that interact‬
‭with the skin, sight is light stimuli, and hearing is the perception of sound, which is a‬
‭physical stimulus similar to some aspects of touch. There are more senses than just those;‬
‭that list represents the primary senses. Those five are all senses that receive stimuli from‬
‭the outside world, of which conscious perception exists. Additional sensory stimuli might‬
‭be from the internal environment (inside the body), such as the stretch of an organ wall or‬
‭the concentration of specific ions in the blood.‬
I‭ ntegration:‬ ‭Stimuli received by sensory structures‬‭are communicated to the nervous‬
‭system, where that information is processed. This is called integration. Stimuli are‬
‭compared with, or integrated with, other stimuli, memories of previous stimuli, or the‬
‭state of a person at a particular time. This process leads to the specific response that will‬
‭be generated. Seeing a baseball pitched to a batter will not automatically cause the batter‬
‭to swing. The trajectory of the ball and its speed will need to be considered. The count‬
‭may be three balls and one strike, and the batter wants to let this pitch go by in the hope‬
‭o f getting a walk to first base. Or maybe the batter’s team is so far ahead it would be fun‬
‭just to swing away.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ otor Response:‬ ‭The nervous system produces a response based on the stimuli‬
M
‭perceived by sensory structures. An obvious response would be the movement of‬
‭muscles, such as withdrawing a hand from a hot stove, but the term has broader uses. The‬
‭nervous system can cause the contraction of all three types of muscle tissue. For example,‬
‭skeletal muscle contracts to move the skeleton, cardiac muscle is influenced as heart rate‬
‭increases during exercise, and smooth muscle contracts as the digestive system moves‬
‭food along the GI tract. Responses also include the neural control of glands in the body,‬
‭such as the production and secretion of sweat by glands found in the skin to lower body‬
‭temperature.‬
‭ he second physiological division of the nervous system is based on control of the body,‬
T
‭which can be somatic or autonomic. The‬‭somatic nervous‬‭system‬‭(‬‭SNS‬‭) is responsible for‬
‭conscious perception and voluntary motor responses.‬‭Voluntary motor response‬‭means‬
‭the contraction of skeletal muscle, but those contractions are not always voluntary‬
‭because you just want to perform them. Some somatic motor responses are‬‭reflexes‬‭and‬
‭o ften happen without a conscious decision to complete them. If your friend jumps out‬
‭from behind a corner and yells, “Boo!” you will be startled, and you might scream or leap‬
‭back. You didn’t decide to do that, but it is a reflex involving skeletal muscle contractions.‬
‭ he‬‭autonomic nervous system‬‭(‬‭ANS‬‭) is responsible‬‭for involuntary body control,‬
T
‭usually for homeostasis (regulation of the internal environment). Sensory input for‬
‭autonomic functions can be from sensory structures tuned to external or internal‬
‭environmental stimuli. The motor output extends to smooth and cardiac muscle and‬
‭glands. The role of the autonomic system is to regulate the body's organ systems, which‬
‭usually means to control homeostasis. Sweat glands, for example, are governed by the‬
‭autonomic system. When you are hot, sweating helps cool your body down. That is a‬
‭homeostatic mechanism. But when you are nervous, you might start sweating, too. That is‬
‭not homeostatic; it is the physiological response to an emotional state.‬
‭ he autonomic nervous system can be divided into the‬‭parasympathetic‬‭and‬
T
‭sympathetic‬‭pathways‬‭(Figure 17.3‬‭). The parasympathetic‬‭division activates “rest and‬
‭digest” responses, whereas the sympathetic division activates “‬‭fight or flight” responses‬‭.‬
‭In other words, these two systems often have opposing effects on target organs; for‬
‭example, activation of the parasympathetic system slows heart rate, whereas activation of‬
‭the sympathetic system increases heart rate.‬
‭ or many students, it is helpful to stop at this point and review the various divisions of the‬
F
‭nervous system.‬ ‭Figure 17.4‬‭provides a useful summary‬‭o f this information.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 17.3‬ ‭In the ANS, a preganglionic neuron of‬‭the CNS synapses with a postganglionic‬
F
‭neuron of the PNS. The postganglionic neuron, in turn, acts on a target organ. Autonomic‬
‭responses are mediated by the sympathetic and the parasympathetic systems, which are‬
‭antagonistic to one another. The sympathetic system activates the “fight or flight”‬
‭response, while the parasympathetic system activates the “rest and digest” response.‬
‭(credit:‬‭OpenStax‬‭); A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 17.4‬ ‭Divisions of the nervous system and‬‭their associated structure(s) and‬
F
‭function(s). (credit:‬‭Nervous System Diagram‬‭;‬‭Fuzzform‬‭at‬‭English Wikipedia‬‭;‬‭CC BY-SA‬
‭3.0‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭17.3 Nervous Tissue‬

‭ ervous tissue, present in both the CNS and PNS, comprises two basic types of cells: glial‬
N
‭cells and neurons.‬‭Glial cells‬‭o utnumber neurons and‬‭are supporting cells that assist the‬
‭development of neurons, maintain the chemical environment around them, and provide‬
‭some structural elements (i.e., myelin sheath).‬

‭ eurons‬‭are the more functionally important of the‬‭two cell types in terms of the‬
N
‭communicative function of the nervous system. They are responsible for the electrical‬
‭signals communicating information about sensations, producing movements in response‬
‭to those stimuli, and inducing thought processes within the brain. There are many‬
‭neurons in the nervous system, a number in the trillions, and many different types of‬
‭neurons among them. They can be classified by many criteria, such as the number of‬
‭processes attached to the cell body, where they are found, what they do, and what‬
‭chemicals they use to communicate.‬
‭ n essential part of the function of neurons is their structure or shape. The‬
A
‭three-dimensional shape of these cells makes the immense number of connections within‬
‭the nervous system possible.‬ ‭The central part of‬‭a neuron is the‬‭cell body‬‭(‬‭Figure 17.5‬‭),‬
‭which contains the nucleus and most of the major organelles. But what makes neurons‬
‭unique is that they have many extensions of their cell membranes, generally called‬
‭processes. Neurons are usually described as having one, and only one,‬‭axon‬‭—a fiber that‬
‭emerges from the cell body and projects to target cells. That single axon can repeatedly‬
‭branch to communicate with many target cells. The axon propagates the nerve impulse,‬
‭which is transmitted to one or more cells. The other neuron processes are‬‭dendrites‬‭,‬
‭which receive information from other neurons. The dendrites are usually highly branched‬
‭processes, providing locations for other neurons to communicate with the cell body.‬
‭Information flows through a neuron from the dendrites, across the cell body, and down‬
‭the axon. This gives the neuron a‬‭polarity‬‭—meaning‬‭that information flows in this one‬
‭direction.‬
‭ any axons are wrapped by an insulating substance called myelin, made from a glial cell‬
M
‭type called‬‭Schwann cells‬‭.‬‭Myelin‬‭acts as insulation,‬‭much like the plastic or rubber used‬
‭to insulate electrical wires. The myelin sheath's appearance can be considered similar to‬
‭the pastry wrapped around a hot dog for “pigs in a blanket” or similar food. The Schwann‬
‭cell is wrapped around the axon several times with little to no cytoplasm between the cell‬
‭layers.‬

‭ critical difference between myelin and the insulation on a wire is that there are gaps in‬
A
‭the myelin covering of an axon. Each gap is called a‬‭node of Ranvier‬‭and is vital to how‬
‭electrical signals travel down the axon. At the end of the axon is the‬‭axon terminal‬‭, where‬
‭there are usually several branches extending toward the target cell, each of which ends in‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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a‭ n enlargement called a‬‭synaptic end bulb‬‭. As discussed in the previous chapter, these‬
‭bulbs connect with the target cell at the‬‭synapse‬‭.‬

‭ igure 17.5‬ ‭A simplified diagram of a neuron. (credit:‬ ‭Neuron.svg‬‭;‬‭Dhp1080‬‭;‬‭CC BY-SA‬

F
‭3.0‬‭). A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ ervous tissue consists of some regions predominantly containing cell bodies and others‬
N
‭primarily composed of axons. These two regions within nervous system structures are‬
‭o ften called‬‭gray matter‬‭(the regions with many cell‬‭bodies and dendrites) or‬‭white‬
‭matter‬‭(the regions with many axons).‬‭Figure 17.6‬‭shows the appearance of these‬
‭regions in the brain and spinal cord.‬
‭ he colors ascribed to these regions would be seen in “fresh,” or unstained, nervous‬
T
‭tissue. Gray matter is not necessarily gray. Depending on how long the tissue has been‬
‭preserved, it can be pinkish because of blood content or even slightly tan. However, the‬
‭white matter is white because of the lipid-rich myelin that insulates the axons. Lipids can‬
‭appear as white (“fatty”) material, much like the fat on a raw piece of chicken or beef. The‬
‭gray matter may have that color ascribed to it because next to the white matter, it is just‬
‭darker—hence, gray.‬

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‭ igure 17.6‬ ‭A brain obtained from an autopsy, with‬‭a partial section removed, shows‬
F
‭white matter surrounded by gray matter. Gray matter makes up the outer cortex of the‬
‭brain. (credit: modification of work by “Suseno”; Wikimedia Commons;‬‭OpenStax‬‭)‬

‭17.4 The General Function of Nervous Tissue‬

S‭ ensation starts with activating a sensory ending, such as the thermoreceptor in the skin‬
‭sensing the temperature of the water. The sensory endings in the skin initiate an electrical‬
‭signal that travels along the sensory axon within a nerve into the spinal cord, where it‬
‭synapses with a neuron in the gray matter of the spinal cord. The temperature‬
‭information represented in that electrical signal is passed to the next neuron by a‬
‭chemical signal that diffuses across the small gap of the synapse and initiates a new‬
‭electrical signal in the target cell. That signal travels through the sensory pathway to the‬
‭brain, where conscious perception of the water temperature is made possible by a region‬
‭called the cerebral cortex. After integrating that information with other cognitive‬
‭processes and sensory input, the brain sends a command back to the spinal cord to‬
‭initiate a motor response by controlling a skeletal muscle. A motor neuron will ultimately‬
‭synapse with a skeletal muscle in a neuromuscular junction and initiate an action potential‬
‭resulting in contraction.‬

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P
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‭17.5 The Action Potential‬

‭ he functions of the nervous system—sensation, integration, and response—depend on‬
T
‭the functions of the neurons underlying these pathways. To understand how neurons can‬
‭communicate, one must describe the role of an excitable membrane in generating these‬
‭signals. The basis of this communication is the‬‭action‬‭potential‬‭, which demonstrates how‬
‭changes in the membrane can constitute a signal. Looking at how these signals work in‬
‭more variable circumstances involves a look at graded potentials, which will be covered in‬
‭the next section.‬

‭Electrically Active Cell Membranes‬

‭ ost cells in the body use charged particles, called ions, to build up a charge across the‬
M
‭cell membrane. In the previous chapter, this was shown to be a part of how muscle cells‬
‭work. For skeletal muscles to contract, input must be obtained from one or more neurons.‬
‭Both muscle and neural cells use the cell membrane to regulate ion movement between‬
‭the extracellular fluid and cytosol.‬
‭ s you learned in Chapter 3, the‬‭cell membrane‬‭regulates‬‭what can cross the membrane‬
A
‭and what stays on only one side. The cell membrane is a‬‭phospholipid bilayer‬‭, so only‬
‭substances passing directly through the hydrophobic core can diffuse unaided. Charged‬
‭particles, which are hydrophilic by definition, cannot pass through the cell membrane‬
‭without assistance (‬‭Figure 17.7‬‭). Transmembrane proteins,‬‭specifically‬‭c hannel proteins‬‭,‬
‭make this possible. Several‬‭passive transport channels‬‭,‬‭as well as‬‭active transport‬
‭pumps‬‭, are necessary to generate a transmembrane potential‬‭and an action potential. Of‬
‭particular interest is the carrier protein referred to as the‬‭sodium-potassium pump‬‭that‬
‭moves‬‭sodium ions‬‭(‬‭Na‬‭+‭)‬ out of a cell and‬‭potassium‬‭ions‬‭(‬‭K‭+‬ ‭)‬ into a cell, thus regulating‬
‭ion concentration on both sides of the cell membrane.‬
‭ he sodium/potassium pump requires adenosine triphosphate (‬‭ATP‬‭) energy, which was‬
T
‭discussed extensively in Chapter 6. The concentration of Na‬‭+‬ ‭is higher outside the cell‬
‭than inside, and the concentration of K‬‭+‬ ‭is higher‬‭inside the cell than outside. That means‬
‭that this pump is moving the ions against the concentration gradients for sodium and‬
‭potassium, which is why it requires energy. The pump maintains those concentration‬
‭gradients.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 17.7‬ ‭The cell membrane comprises a phospholipid‬‭bilayer with many‬

F
‭transmembrane proteins, including different channel proteins that serve as ion channels.‬
‭(credit:‬‭OpenStax‬‭); A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

I‭ on channels‬‭are pores that allow specific charged‬‭particles to cross the membrane in‬
‭response to an existing‬‭concentration gradient‬‭. Proteins‬‭can span the cell membrane,‬
‭including its hydrophobic core. They can interact with the charge of ions because of the‬
‭amino acids' various properties within specific regions of the protein channel. Ion‬
‭channels do not always freely allow ions to diffuse across the membrane. Some are‬
‭o pened by certain events, meaning the channels are gated. Another way that channels can‬
‭be categorized is based on how they are gated.‬
‭ ‬‭ligand-gated channel‬‭o pens because a signaling molecule,‬‭a ligand, binds to the‬
A
‭extracellular region of the channel. When the‬‭ligand‬‭,‬‭known as a‬‭neurotransmitter‬‭in the‬
‭nervous system, binds to the protein, ions cross the membrane, changing its electrical‬
‭potential (‬‭Figure 17.8‬‭).‬
‭ ‬‭voltage-gated channel‬‭responds to changes in the‬‭electrical properties of the‬
A
‭membrane in which it is embedded. Usually, the inner portion of the membrane has a‬
‭negative voltage. When that voltage becomes less negative, the channel allows ions to‬
‭cross the membrane (‬‭Figure 17.9‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 17.8‬ ‭When the neurotransmitter acetylcholine‬‭binds to a specific location on the‬

F
‭extracellular surface of the channel protein, the pore opens to allow select ions through.‬
‭In this case, the ions are sodium, calcium, and potassium cations. (credit:‬‭OpenStax‬‭); A link‬
‭to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ igure 17.9‬ ‭Voltage-gated channels open when the‬‭transmembrane voltage changes‬

F
‭around them. Amino acids in the protein structure are sensitive to charge and cause the‬
‭pore to open to the selected ion. (credit:‬‭OpenStax‬‭);‬ ‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ ‬‭leakage channel‬‭is randomly gated, meaning it opens and closes randomly, hence the‬
A
‭reference to leaking. No actual event opens the channel; instead, it has an intrinsic rate of‬
‭switching between the open and closed states. Leakage channels contribute to the resting‬
‭transmembrane voltage of an excitable membrane (‬‭Figure‬‭17.10‬‭).‬

‭The Membrane Potential‬

‭ he electrical state of the cell membrane can have several variations. These are all‬
T
‭variations in the‬‭membrane potential‬‭. A potential‬‭is a charge distribution across the cell‬
‭membrane, measured in‬‭millivolts‬‭(‬‭mV‬‭). The standard‬‭is to compare the inside of the cell‬
‭relative to the outside, so the membrane potential represents the charge on the‬
‭intracellular side of the membrane based on the outside being zero, relatively speaking‬
‭(‬‭Figure 17.11‬‭).‬

‭ igure 17.10‬ ‭In certain situations, ions must randomly‬‭move across the membrane via‬
F
‭leakage channels. This type of channel modifies the particular electrical properties of‬
‭specific cells. (credit:‬‭OpenStax‬‭)‬

‭ he concentration of ions in extracellular and intracellular fluids is mainly balanced, with a‬

T
‭net neutral charge. However, a slight difference in charge occurs right at the membrane‬
‭surface, both internally and externally. It is the difference in this minimal region that has‬
‭all the power in neurons (and muscle cells) to generate electrical signals, including action‬
‭potentials.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 17.11‬ ‭When measuring charge across a membrane‬‭with a voltmeter, a recording‬

F
‭electrode is inserted into the cell, and a reference electrode is outside the cell. The‬
‭transmembrane voltage is determined by comparing the charge measured by these two‬
‭electrodes. Expressing that value for the cytosol relative to the outside is conventional.‬
‭(credit:‬‭OpenStax‬‭); A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ efore these electrical signals can be described, the‬‭resting state‬‭o f the membrane must‬
B
‭be explained. When the cell is at rest and the ion channels are closed (except for leakage‬
‭channels, which are randomly open), ions are distributed across the membrane in a very‬
‭predictable way. The concentration of Na‬‭+‬ ‭o utside‬‭the cell is ten times greater than the‬
‭concentration inside. Also, the concentration of K‬‭+‬ ‭inside the cell is greater than outside.‬
‭The cytosol contains a high concentration of anions in the form of phosphate ions and‬
‭negatively charged proteins. Large anions are a component of the inner cell membrane,‬
‭including specialized phospholipids and proteins.‬
‭ ith the ions distributed across the membrane at these concentrations, the difference in‬
W
‭charge is measured at‬‭-70 mV‬‭, the value described‬‭as the‬‭resting membrane potential‬‭.‬
‭The exact value measured for the resting membrane potential varies between cells, but‬
‭-70 mV is most commonly used as this value. This voltage would be much lower except for‬
‭the contributions of some important proteins in the membrane. Leakage channels allow‬
‭Na‬‭+‬ ‭to slowly move into the cell or K‬‭+‬ ‭to move out‬‭slowly, and the Na‬‭+‭/‬ K‬‭+‬ ‭pump restores‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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t‭ hem. This may appear to be a waste of energy, but each has a role in maintaining the‬
‭membrane potential.‬

‭The Action Potential‬

‭ esting membrane potential describes the steady state of the cell, which is a dynamic‬
R
‭process balanced by ion leakage and ion pumping. It will not change without any outside‬
‭influence. To initiate an electrical signal, the membrane potential has to change.‬
‭ his event begins with a channel opening for Na‬‭+‬ ‭in the membrane (‬‭Figure 17.12‬‭).‬
T
‭Because the concentration of Na‬‭+‬ ‭is higher outside‬‭the cell than inside the cell by a factor‬
‭o f 10, ions will rush into the cell, primarily driven by the concentration gradient. Because‬
‭sodium is a positively charged ion, it will change the relative voltage immediately inside the‬
‭cell relative to the outside. The resting potential is the state of the membrane at a voltage‬
‭o f -70 mV, so the sodium cation entering the cell will cause it to become less negative. This‬
‭is known as‬‭depolarization‬‭, meaning the membrane potential‬‭moves toward zero.‬
‭ he concentration gradient for Na‬‭+‬ ‭is so strong that‬‭it will continue to enter the cell even‬
T
‭after the membrane potential has become zero so that the voltage immediately around the‬
‭pore begins to become positive. The electrical gradient also plays a role, as negative‬
‭proteins below the membrane attract the sodium ion. The membrane potential will reach‬
‭+30 mV‬‭when sodium has entered the cell.‬
‭ s the membrane potential reaches +30 mV, other voltage-gated channels open in the‬
A
‭membrane. These channels are specific for the potassium ion. A concentration gradient‬
‭also acts on K‬‭+‭.‬ As K‬‭+‬ ‭leaves the cell, taking a positive‬‭charge with it, the membrane‬
‭potential moves back toward its resting voltage. This is called‬‭repolarization‬‭, meaning‬
‭that the membrane voltage moves back toward the -70 mV value of the resting membrane‬
‭potential.‬
‭ epolarization returns the membrane potential to the -70 mV value, which indicates the‬
R
‭resting potential, but it overshoots that value. Potassium ions reach equilibrium when the‬
‭membrane voltage is below -70 mV, so hyperpolarization occurs while the K‬‭+‬ ‭channels are‬
‭o pen. Those K‬‭+‬ ‭channels are slightly delayed in closing,‬‭accounting for this short‬
‭overshoot.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 17.12‬ ‭The (a) resting membrane potential‬‭results from different concentrations‬
F
‭o f Na‬‭+‬ ‭and K‬‭+‬ ‭ions inside and outside the cell. A‬‭nerve impulse causes Na‬‭+‬ ‭to enter the cell,‬
‭depolarizing (b). At the peak action potential, K‬‭+‬ ‭channels open, and the cell becomes (c)‬
‭hyperpolarized. (credit:‬‭OpenStax‬‭); A link to a‬‭video explanation of this figure is available‬
‭at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ he action potential is presented as a voltage over time graph in‬‭Figure 17.13‬‭. It is the‬
T
‭electrical signal that nervous tissue generates for communication. The change in the‬
‭membrane voltage from -70 mV at rest to +30 mV at the end of depolarization is a 100 mV‬
‭change. That can also be written as a 0.1 V change. To put that value in perspective, think‬
‭about a battery. An AA battery that you might find in a television remote has a voltage of‬
‭1.5 V, or a 9 V battery (the rectangular battery with two posts on one end) is, obviously, 9‬
‭V. The change seen in the action potential is one or two orders of magnitude less than the‬
‭charge in these batteries. The membrane potential can be described as a battery. A charge‬
‭is stored across the membrane that can be released under the right conditions. A battery‬
‭in your remote has stored a charge that is “released” when you push a button.‬

‭ igure 17.13‬ ‭A graph of an action potential is obtained‬‭by plotting voltage measured‬

F
‭across the cell membrane (Y-axis) against time (X-axis). The action potential begins with‬
‭depolarization, followed by repolarization, which goes past the resting potential into‬
‭hyperpolarization, and finally, the membrane returns to a resting state at -70 mV. (credit:‬
‭OpenStax‬‭); A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭ he question is, now, what initiates the action potential? The description above‬
T
‭conveniently glosses over that point, but it is vital to understand what is happening. The‬
‭membrane potential will stay at the resting voltage until something changes. The‬
‭description above just says that a Na‬‭+‬ ‭channel opens.‬‭Saying “a channel opens” does not‬
‭mean one individual transmembrane protein changes. Instead, it means that one kind of‬
‭channel opens. A few different types of channels allow Na‬‭+‬ ‭to cross the membrane.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ ‬‭ligand-gated Na‬‭+‬ ‭c hannel‬‭will open when a neurotransmitter binds to it, and some‬
A
‭stimulus gets the process started. Sodium starts to enter the cell, and the membrane‬
‭becomes less negative.‬
‭ third channel important for depolarization in the action potential is the‬‭voltage-gated‬
A
‭Na‬‭+‬ ‭c hannel‬‭. The channels that start depolarizing‬‭the membrane because of a stimulus‬
‭help the cell to depolarize from -70 mV to -55 mV. Once the membrane reaches that‬
‭voltage, the voltage-gated Na‬‭+‬ ‭channels open. This‬‭is what is known as the‬‭threshold‬‭. Any‬
‭depolarization that does not change the membrane potential to‬‭-55 mV‬‭o r higher will not‬
‭reach the threshold and, thus, will not result in an action potential. Also, any stimulus that‬
‭depolarizes the membrane to -55 mV or beyond will cause many channels to open, and an‬
‭action potential will be initiated.‬
‭ ecause of the threshold, the action potential can be likened to a digital event—it either‬
B
‭happens or does not. If the threshold is not reached, then no action potential occurs. If‬
‭depolarization reaches -55 mV, the action potential continues and runs to +30 mV, at‬
‭which K‬‭+‬ ‭causes repolarization, including the hyperpolarizing‬‭overshoot. Also, those‬
‭changes are the same for every action potential, which means that the same thing happens‬
‭o nce the threshold is reached. A more substantial stimulus, which might depolarize the‬
‭membrane well past the threshold, will not make a “bigger” action potential. Action‬
‭potentials are‬‭“all or none.”‬‭Either the membrane‬‭reaches the threshold, everything‬
‭o ccurs as described above, or the membrane does not, and nothing else happens. All‬
‭action potentials peak at the same voltage (+30 mV), so one action potential is not bigger‬
‭than another. More potent stimuli will initiate multiple action potentials more quickly, but‬
‭the individual signals are not bigger. Thus, for example, you will not feel a greater pain‬
‭sensation or have a stronger muscle contraction because the action potentials are not‬
‭different sizes.‬
‭ ll of this takes place within approximately two milliseconds (‬‭Figure 17.14‬‭). While an‬
A
‭action potential is in progress, another cannot be initiated, which is referred to as the‬
‭refractory period‬‭.‬

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‭ igure 17.14‬ ‭The action potential events can be‬‭related to specific changes in the‬
F
‭membrane voltage. (1) At rest, the membrane voltage is -70 mV. (2) The membrane‬
‭depolarizes when an external stimulus is applied. (3) The membrane voltage begins a‬
‭rapid rise toward +30 mV. (4) The membrane voltage starts to return to a negative value.‬
‭(5) Repolarization continues past the resting membrane voltage, resulting in‬
‭hyperpolarization. (6) The membrane voltage returns to the resting value shortly after‬
‭hyperpolarization. (credit:‬‭OpenStax‬‭); A link to‬‭a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

‭Propagation of the Action Potential‬

‭ he action potential is initiated at the beginning of the axon. There is a high density of‬
T
‭voltage-gated Na‬‭+‬ ‭channels, so that rapid depolarization‬‭can occur here. The action‬
‭potential is‬‭propagated‬‭along the length of the axon‬‭because more voltage-gated Na‬‭+‬
‭channels open as the depolarization spreads. This spreading happens because Na‬‭+‬ ‭enters‬
‭through the channel and moves along the inside of the cell membrane. As the Na‬‭+‬ ‭moves,‬
‭o r flows, a short distance along the cell membrane, its positive charge depolarizes a little‬
‭more of the cell membrane. As that depolarization spreads, new voltage-gated Na‬‭+‬
‭channels open, and more ions rush into the cell, spreading the depolarization a little‬
‭farther (‬‭Figure 17.15‬‭).‬

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‭ igure 17.15‬ ‭The action potential is conducted down‬‭the axon as the axon membrane‬
F
‭depolarizes, then repolarizes. (credit:‬‭OpenStax‬‭);‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ ropagation, as described above, applies to unmyelinated axons. When myelination is‬

P
‭present, the action potential propagates differently (‬‭Figure 17.16‬‭). Sodium ions that enter‬
‭the cell at the initial segment start to spread along the length of the axon segment, but‬
‭there are no voltage-gated Na‬‭+‬ ‭channels until the‬‭first node of Ranvier. The depolarization‬
‭spreads at an optimal speed because these channels aren’t constantly opened along the‬
‭axon segment. The distance between nodes is optimal to keep the membrane depolarized‬
‭above the next node's threshold. As Na‬‭+‬ ‭spreads along‬‭the inside of the membrane of the‬

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a‭ xon segment, the charge starts to dissipate. If the node were any farther down the axon,‬
‭that depolarization would have fallen off too much for voltage-gated Na‬‭+‬ ‭channels to be‬
‭activated at the next node of Ranvier.‬

‭ igure 17.16‬ ‭Nodes of Ranvier are gaps in myelin‬‭coverage along axons. Nodes contain‬
F
‭voltage-gated K‬‭+‬ ‭and Na‬‭+‬ ‭channels. Action potentials‬‭travel down the axon by jumping‬
‭from one node to the next. (credit:‬‭OpenStax‬‭); A‬‭link to a video explanation of this figure‬
‭is available at‬‭Biology411.com‬‭.‬

‭ ropagation along an unmyelinated axon is called‬‭continuous‬‭conduction‬‭; along the‬

P
‭length of a myelinated axon, it is‬‭saltatory conduction‬‭.‬‭Continuous conduction is slow‬
‭because there are always voltage-gated Na‬‭+‬ ‭channels‬‭o pening, and more and more Na‬‭+‬ ‭is‬
‭rushing into the cell. Saltatory conduction is faster because the action potential jumps‬
‭from one node to the next (saltare = “to leap”), and the new influx of Na‬‭+‬ ‭renews the‬
‭depolarized membrane. If nodes of Ranvier were not present along an axon, the action‬
‭potential would propagate very slowly since Na‬‭+‬ ‭and‬‭K‭+‬ ‬ ‭channels would have to‬
‭continuously regenerate action potentials at every point along the axon instead of at‬
‭specific points. Nodes of Ranvier also save energy for the neuron since the channels only‬
‭need to be present at the nodes and not along the entire axon.‬
‭ or additional information about how nerves function, click the link or scan the QR code‬
F
‭to watch the TED-Ed video by Elliot Krane titled “How do nerves work?”.‬

How do nerves work? - Elliot Krane

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‭17.6 Communication Between Neurons‬

‭ s described in the previous section, the electrical changes within a neuron are similar to‬
A
‭a light switch being turned on. A stimulus starts depolarization, but the action potential‬
‭runs independently once a threshold is achieved. The question is now, “What flips the light‬
‭switch on?” Temporary changes to the cell membrane voltage can result from neurons‬
‭receiving information from the environment or the action of one neuron on another.‬
‭These special types of potentials influence a neuron and determine whether an action‬
‭potential will occur. Many of these transient signals originate at the synapse.‬

‭Chemical Synapses‬
S‭ ynapses are the contacts between neurons, which can be chemical or electrical, but‬
‭c hemical synapses‬‭are far more common. An example‬‭o f a chemical synapse is the‬
‭neuromuscular junction (NMJ) described in the previous chapter, which deals with skeletal‬
‭muscle tissue and contraction. In the nervous system, many more synapses are the same‬
‭as the NMJ. All synapses have common characteristics, which can be summarized in this‬
‭list:‬
•‭ ‬ ‭ resynaptic element‬
p
‭•‬ ‭neurotransmitter (packaged in vesicles)‬
‭•‬ ‭synaptic cleft‬
‭•‬ ‭receptor proteins‬
‭•‬ ‭postsynaptic element‬
‭•‬ ‭neurotransmitter elimination or re-uptake‬
‭ or the NMJ, these characteristics are as follows: the presynaptic element is the motor‬
F
‭neuron's axon terminals, the neurotransmitter is acetylcholine, the synaptic cleft is the‬
‭space between the cells where the neurotransmitter diffuses, the receptor protein is the‬
‭nicotinic acetylcholine receptor, the postsynaptic element is the sarcolemma of the muscle‬
‭cell, and the neurotransmitter is eliminated by acetylcholinesterase. Other synapses are‬
‭similar to this, and the specifics are different, but they all contain the same characteristics.‬
‭ t a chemical synapse,‬‭synaptic vesicles‬‭(‬‭Figure 17.17‬‭)‬‭at the axon terminal of the‬
A
‭presynaptic neuron are stimulated, via increasing calcium ion concentrations, to migrate to‬
‭and fuse with the membrane. Chemical messengers, called‬‭neurotransmitters,‬‭are‬
‭released into the junction via‬‭exocytosis‬‭. The neurotransmitters‬‭diffuse across the‬
‭synaptic cleft‬‭, bind to‬‭receptors‬‭embedded in the‬‭membrane of the postsynaptic neuron,‬
‭and cause a change in the postsynaptic membrane. Receptors are specific for‬
‭neurotransmitters; the two fit together like a key and lock. One neurotransmitter binds to‬

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i‭ts receptor and will not attach to receptors for other neurotransmitters, making the‬
‭binding a specific chemical event.‬
‭ eurotransmitters may be either‬‭excitatory‬‭(such as‬‭acetylcholine or epinephrine) or‬
N
‭inhibitory‬‭(such as serotonin) as they either increase‬‭o r decrease the change of an action‬
‭potential in the postsynaptic neuron. Many drugs can induce changes in synaptic‬
‭transmission; for example, tetrahydrocannabinol (more commonly known as THC) in‬
‭marijuana binds to a naturally occurring neurotransmitter important to short-term‬
‭memory.‬
‭ nce neurotransmission has occurred, the neurotransmitter must be removed from the‬
O
‭synaptic cleft so the postsynaptic membrane can “reset” and be ready to receive another‬
‭signal. This removal can be accomplished in three ways: the neurotransmitter can diffuse‬
‭away from the synaptic cleft, be degraded by enzymes in the synaptic cleft, or be recycled‬
‭(sometimes called reuptake) by the presynaptic neuron. Several drugs act at this step of‬
‭neurotransmission. For example, some medications given to Alzheimer’s patients work by‬
‭inhibiting acetylcholinesterase, the enzyme that degrades acetylcholine. This inhibition of‬
‭the enzyme essentially increases neurotransmission at these synapses.‬

‭ igure 17.17‬ ‭This pseudo-colored image, taken with‬‭a scanning electron microscope,‬
F
‭shows an axon terminal broken open to reveal synaptic vesicles (blue and orange) inside‬
‭the neuron. (credit: modification of work by Tina Carvalho, NIH-NIGMS; scale-bar data‬
‭from Matt Russell;‬‭OpenStax‬‭)‬

‭ hen an action potential reaches the axon terminals, voltage-gated Ca‬‭+2‬ ‭channels in the‬
W
‭membrane of the synaptic end bulb open. The concentration of Ca‬‭+2‬ ‭increases inside the‬

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e‭ nd bulb and the Ca‬‭+2‬ ‭ion associates with proteins in the outer surface of‬
‭neurotransmitter vesicles. The Ca‬‭+2‬ ‭facilitates the‬‭merging of the vesicle with the‬
‭presynaptic membrane so that the neurotransmitter is released through‬‭exocytosis‬‭into‬
‭the small gap between the cells, known as the synaptic cleft‬‭(‬‭Figure 17.18‬‭). Once in the‬
‭synaptic cleft, the neurotransmitter diffuses a short distance to the postsynaptic‬
‭membrane and can interact with neurotransmitter receptors.‬

‭Signal Summation‬
S‭ ometimes, a single excitatory input signal is strong enough to induce an action potential‬
‭in the postsynaptic neuron. Still, multiple excitatory inputs are often required around the‬
‭same time for the postsynaptic neuron to be sufficiently depolarized to fire an action‬
‭potential. This process is called‬‭summation‬‭and occurs‬‭at the connection point of the‬
‭axon to the cell body (‬‭Figure 17.19)‬‭.‬
‭ dditionally, one neuron often has inputs from many presynaptic neurons—some‬
A
‭excitatory and some inhibitory—so inhibitory input signals can cancel out excitatory‬
‭signals and vice versa. The net change in postsynaptic membrane voltage determines‬
‭whether the postsynaptic cell has reached its threshold of excitation needed to fire an‬
‭action potential. Together, synaptic summation and the threshold for excitation act as a‬
‭filter so that random “noise” in the system is not transmitted as necessary information.‬

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‭ igure 17.18‬ ‭Communication at chemical synapses requires‬‭the release of‬

F
‭neurotransmitters. When the presynaptic membrane is depolarized, voltage-gated Ca‬‭2+‬
‭channels open, releasing Ca‬‭2+‬‭. The calcium entry causes‬‭synaptic vesicles to fuse with the‬
‭membrane and release neurotransmitters into the synaptic cleft. The neurotransmitter‬
‭diffuses across the gap and binds to ligand-gated ion channels in the postsynaptic‬
‭membrane, resulting in the postsynaptic neuron's localized depolarization or‬
‭hyperpolarization. Neurotransmitters are removed from the gap by one of several‬
‭possible mechanisms. (credit:‬‭OpenStax‬‭); A link to‬‭a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

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‭ igure 17.19‬ ‭A single neuron can receive excitatory‬‭and inhibitory inputs from multiple‬
F
‭neurons, resulting in local membrane depolarization (EPSP input) and hyperpolarization‬
‭(IPSP input). All these inputs are added together at the axon hillock. The neuron will fire if‬
‭the EPSPs are strong enough to overcome the IPSPs and reach the excitation threshold.‬
‭(credit:‬‭OpenStax‬‭); A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭17.7 Nervous System Disorders‬

‭ nervous system that functions correctly is a complex and well-oiled machine—synapses‬
A
‭fire appropriately, muscles move when needed, memories are formed and stored, and‬
‭emotions are well-regulated. You can now appreciate what it takes for you to be able to‬
‭read and comprehend the information in this textbook. Unfortunately, each year, millions‬
‭o f people in the United States deal with some sort of disorder involving the nervous‬
‭system. Neurodegenerative disorders are characterized by loss of nervous system‬
‭functioning, usually caused by the death of neurons; examples include multiple sclerosis,‬
‭Alzheimer’s disease, Parkinson’s disease, and amyotrophic lateral sclerosis (ALS).‬
‭Neurodevelopmental disorders occur when the development of the nervous system is‬
‭disturbed; examples include autism spectrum disorder (ASD), attention deficit‬
‭hyperactivity disorder (ADHD), schizophrenia, and major depression. Epilepsy and stroke‬
‭also have neurological origins. Although scientists have discovered potential causes of‬
‭many of these diseases and effective treatments for some, ongoing research into the‬
‭prevention and treatment of these disorders continues.‬

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‭Neurodegenerative Disorders‬
‭ eurodegenerative disorders‬‭are illnesses characterized‬‭by a loss of nervous system‬
N
‭functioning, usually caused by neuronal death. These diseases generally worsen over time‬
‭as more and more neurons die. The symptoms of a particular neurodegenerative disease‬
‭are related to where neurons are killed in the nervous system. The death of these neurons‬
‭causes problems in balance and walking. Neurodegenerative disorders include multiple‬
‭sclerosis, Alzheimer’s disease, Parkinson’s disease, and amyotrophic lateral sclerosis (Lou‬
‭Gehrig’s disease), which are discussed in more detail.‬

‭Multiple Sclerosis‬

‭ ultiple sclerosis‬‭(MS) is an example of an autoimmune‬‭disease. The antibodies‬

M
‭produced by lymphocytes (a type of white blood cell) mark myelin as something that‬
‭should not be in the body. This response causes inflammation and the destruction of the‬
‭myelin in the central nervous system. As the disease destroys the insulation around the‬
‭axons, scarring becomes obvious. This is where the name of the disease comes from;‬
‭sclerosis means hardening of tissue, which is what a scar is. Multiple scars are found in the‬
‭white matter of the brain and spinal cord. The symptoms of MS include both somatic and‬
‭autonomic deficits. Control of the musculature is compromised, as is control of organs‬
‭such as the bladder.‬

‭Alzheimer’s Disease‬
‭ lzheimer’s disease‬‭is the most common cause of dementia‬‭in the elderly. In 2012, an‬
A
‭estimated 5.4 million Americans had Alzheimer’s disease, and payments for their care are‬
‭estimated at $200 billion. Roughly one in every eight people age 65 or older has the‬
‭disease. Due to the aging of the baby-boomer generation, there are projected to be as‬
‭many as 13 million Alzheimer’s patients in the United States in the year 2050.‬
S‭ ymptoms of Alzheimer’s disease include disruptive memory loss, confusion about time or‬
‭place, difficulty planning or executing tasks, poor judgment, and personality changes.‬
‭Problems smelling certain scents can also indicate Alzheimer’s disease and may be an‬
‭early warning sign. Many of these symptoms are also common in people who are aging‬
‭normally, so the severity and longevity of the symptoms determine whether a person has‬
‭Alzheimer’s.‬
‭ lzheimer’s disease is named for Alois Alzheimer, a German psychiatrist who published a‬
A
‭report in 1911 about a woman who showed severe dementia symptoms. He and his‬
‭colleagues examined the woman’s brain following her death. He reported the presence of‬

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a‭ bnormal clumps, which are now called amyloid plaques, along with tangled brain fibers‬
‭called neurofibrillary tangles. Amyloid plaques, neurofibrillary tangles, and an overall‬
‭shrinking of brain volume are commonly seen in the brains of Alzheimer’s patients. Loss‬
‭o f neurons in the hippocampus is especially severe in advanced Alzheimer’s patients.‬
‭Figure 17.20‬‭compares a normal brain to the brain‬‭o f an Alzheimer’s patient. Many‬
‭research groups are examining the causes of these hallmarks of the disease.‬

‭ igure 17.20‬ ‭Compared to a normal brain (left),‬‭the brain of a patient with Alzheimer’s‬
F
‭disease (right) shows dramatic neurodegeneration, particularly within the ventricles and‬
‭hippocampus. (credit: modification of work by “Garrando”/Wikimedia Commons based on‬
‭o riginal images by ADEAR: "Alzheimer's Disease Education and Referral Center, a service‬
‭o f the National Institute on Aging”;‬‭OpenStax‬‭)‬

‭ ne form of the disease is usually caused by mutations in one of three known genes. This‬
O
‭rare form of early-onset Alzheimer’s disease affects fewer than five percent of patients‬
‭with the disease and causes dementia beginning between the ages of 30 and 60. The‬
‭disease's more prevalent, late-onset form likely has a genetic component. One particular‬
‭gene, apolipoprotein E (APOE), has a variant (E4) that increases a carrier’s (an individual‬
‭with one normal allele and one abnormal one) likelihood of getting the disease. Many‬
‭o ther genes that might be involved in the pathology have been identified.‬
‭ nfortunately, there is no cure for Alzheimer’s disease. Current treatments focus on‬
U
‭managing the symptoms of the disease. Because the decrease in the activity of cholinergic‬
‭neurons (neurons that use the neurotransmitter acetylcholine) is common in Alzheimer’s‬
‭disease, several drugs used to treat the disease work by increasing acetylcholine‬

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‭ eurotransmission, often by inhibiting the enzyme that breaks down acetylcholine in the‬
n
‭synaptic cleft. Other clinical interventions focus on behavioral therapies like‬
‭psychotherapy, sensory therapy, and cognitive exercises. Because Alzheimer’s disease‬
‭appears to hijack the normal aging process, prevention research is prevalent. Smoking,‬
‭o besity, and cardiovascular problems may be risk factors for the disease, so treatments‬
‭for those may also help to prevent Alzheimer’s disease. Some studies have shown that‬
‭people who remain intellectually active by playing games, reading, playing musical‬
‭instruments, and being socially active in later life have a reduced risk of developing the‬
‭disease.‬
‭ or more information about this disease, click the link below or scan the QR code to watch‬
F
‭the TED-Ed video by Ivan Seah Yun Jun titled “What is Alzheimer’s disease?”.‬

What is Alzheimer's disease? - Ivan Seah Yu Jun

‭Parkinson’s Disease‬
‭ ike Alzheimer’s disease,‬‭Parkinson’s disease‬‭is neurodegenerative.‬‭It was first‬
L
‭characterized by James Parkinson in 1817. Each year, 50,000-60,000 people in the United‬
‭States are diagnosed with the disease. Parkinson’s disease causes the loss of dopamine‬
‭neurons in the substantia nigra, a midbrain structure that regulates movement. Loss of‬
‭these neurons causes many symptoms, including tremors (shaking of fingers or a limb),‬
‭slowed movement, speech changes, balance and posture problems, and rigid muscles. The‬
‭combination of these symptoms often causes a characteristic slow, hunched, shuffling‬
‭walk, illustrated in‬‭Figure 17.21‬‭. Patients with Parkinson’s‬‭disease can also exhibit‬
‭psychological symptoms, such as dementia or emotional problems.‬
‭ lthough some patients have a form of the disease known to be caused by a single‬
A
‭mutation, the exact causes of Parkinson’s disease remain unknown: the condition likely‬
‭results from a combination of genetic and environmental factors (similar to Alzheimer’s‬
‭disease). Post-mortem analysis of the brains of Parkinson’s patients shows the presence of‬
‭Lewy bodies—abnormal protein clumps—in specific neurons. The prevalence of these‬
‭Lewy bodies often correlates with the severity of the disease.‬
‭ here is no cure for Parkinson’s disease, and treatment is focused on easing symptoms.‬
T
‭One of the most commonly prescribed drugs for Parkinson’s is L-DOPA, a chemical‬
‭converted into dopamine by neurons in the brain. This conversion increases the overall‬

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l‭evel of dopamine neurotransmission and can help compensate for the loss of‬
‭dopaminergic neurons in the substantia nigra. Other drugs work by inhibiting the enzyme‬
‭that breaks down dopamine.‬
‭ or additional information about this disease, click the link below or scan the QR code to‬
F
‭watch the Nature video “Understanding Parkinson’s disease.”‬

‭Understanding Parkinson's Disease‬

‭ igure 17.21‬ ‭Parkinson’s patients often have a characteristic‬‭hunched walk. (credit:‬

F
‭OpenStax‬‭)‬

‭Amyotrophic Lateral Sclerosis (Lou Gehrig’s disease)‬

‭ he roots of the term‬‭amyotrophic lateral sclerosis‬‭(‬‭ALS‬‭) provide clues as to what‬

T
‭happens to an individual with this disease. “Amyotrophic” means “no muscle nourishment,”‬
‭“lateral” refers to the part of the spine that tells muscles what to do, and “sclerosis”‬
‭indicates that the lateral portion of the spine that controls muscle movement hardens. ALS‬
‭causes a decline in voluntary muscle control over time via two mechanisms:‬

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‭1.‬ A
‭ LS degenerates sensory neurons that control voluntary muscle movement. As the‬
‭lateral portion of the spine that controls muscle movement hardens, signals are no‬
‭longer sent to muscles. Muscles weaken, but coordination is not affected, and‬
‭eventually, paralysis occurs.‬

‭2.‬ A
‭ LS degenerates motor neurons that control voluntary muscle movement. As the‬
‭lateral portion of the spine that controls muscle movement hardens, signals are no‬
‭longer sent to muscles. Initially, muscles strengthen, coordination is affected, and‬
‭eventually, paralysis occurs.‬

‭ or additional information about this disease, click the link below or scan the QR code to‬
F
‭watch the TED-Ed video by Fernando G. Vieira titled “Why is it so hard to cure ALS?”.‬

Why is it so hard to cure ALS? - Fernando G. Vieira

‭Everyday Connection‬

‭ rain-computer interface‬
B
‭Amyotrophic lateral sclerosis (ALS, also called Lou Gehrig’s Disease) is a neurological‬
‭disease characterized by the degeneration of the motor neurons that control voluntary‬
‭movements. The disease begins with muscle weakening and lack of coordination and‬
‭eventually destroys the neurons that control speech, breathing, and swallowing; in the‬
‭end, the condition can lead to paralysis. At that point, patients require assistance from‬
‭machines to breathe and communicate. Several technologies have been developed to allow‬
‭“locked-in” patients to communicate with the rest of the world. One technology, for‬
‭example, will enable patients to type sentences by twitching their cheeks. These sentences‬
‭can then be read aloud by a computer.‬

‭ relatively new line of research for helping paralyzed patients, including those with ALS,‬
A
‭to communicate and retain a degree of self-sufficiency is called brain-computer interface‬
‭(BCI) technology and is illustrated in‬‭Figure 17.22‬‭.‬‭This technology sounds like something‬
‭o ut of science fiction: it allows paralyzed patients to control a computer using only their‬
‭thoughts. There are several forms of BCI. Some forms use EEG recordings from electrodes‬
‭taped onto the skull. These recordings contain information from large populations of‬
‭neurons that a computer can decode. Other forms of BCI require implanting an array of‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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e‭ lectrodes smaller than a postage stamp in the arm and hand area of the motor cortex.‬
‭This form of BCI, while more invasive, is potent as each electrode can record actual action‬
‭potentials from one or more neurons. These signals are then sent to a computer, which‬
‭has been trained to decode the signal and feed it to a tool—such as a cursor on a‬
‭computer screen. This means a patient with ALS can use e-mail, read the Internet, and‬
‭communicate with others by thinking of moving their hand or arm (even though the‬
‭paralyzed patient cannot make that bodily movement). Recent advances have allowed a‬
‭paralyzed locked-in patient who suffered a stroke 15 years ago to control a robotic arm‬
‭and drink coffee using BCI technology.‬

‭ espite the fantastic advancements in BCI technology, it also has limitations. The‬
D
‭technology can require many hours of training and long periods of intense concentration‬
‭for the patient, and implanting the devices can require brain surgery.‬

‭ igure 17.22‬ ‭With brain-computer interface technology,‬‭neural signals from a paralyzed‬

F
‭patient are collected, decoded, and then fed to a tool, such as a computer, a wheelchair, or‬
‭a robotic arm. (credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Neurodevelopmental Disorders‬
‭ eurodevelopmental disorders‬‭o ccur when the nervous system's development is‬
N
‭disturbed. There are several different classes of neurodevelopmental disorders. Some, like‬
‭Down Syndrome, cause intellectual deficits. Others specifically affect communication,‬
‭learning, or the motor system. Some disorders, like autism spectrum disorder and‬
‭attention-deficit/hyperactivity disorder, have complex symptoms.‬

‭Autism‬
‭ utism spectrum disorder (ASD)‬‭is a neurodevelopmental‬‭disorder. Its severity differs‬
A
‭from person to person. Estimates for the prevalence of the disorder have changed rapidly‬
‭in the past few decades. Current estimates suggest that one in 88 children will develop the‬
‭disorder; ASD is four times more prevalent in males than females.‬
‭ characteristic symptom of ASD is impaired social skills. Children with autism may have‬
A
‭difficulty making and maintaining eye contact and reading social cues. They also may have‬
‭problems feeling empathy for others. Other symptoms of ASD include repetitive motor‬
‭behaviors (such as rocking back and forth), preoccupation with specific subjects, strict‬
‭adherence to certain rituals, and unusual language use. Up to 30 percent of patients with‬
‭ASD develop epilepsy, and patients with some forms of the disorder (like Fragile X) also‬
‭have an intellectual disability. Because it is a spectrum disorder, other ASD patients are‬
‭very functional and have good-to-excellent language skills. Many of these patients do not‬
‭feel they suffer from a disorder; instead, they think their brains process information‬
‭differently.‬
‭ xcept for some well-characterized, clearly genetic forms of autism (like Fragile X and‬
E
‭Rett’s Syndrome), the causes of ASD are largely unknown. Variants of several genes‬
‭correlate with ASD, but for any given patient, many different mutations in different genes‬
‭may be required for the disease to develop. Generally, ASD is considered a disease of‬
‭“incorrect” wiring. Accordingly, the brains of some ASD patients lack the same level of‬
‭synaptic pruning that occurs in non-affected people. In the 1990s, a research paper linked‬
‭autism to a common vaccine given to children. This paper was retracted when it was‬
‭discovered that the author falsified data, and follow-up studies showed no connection‬
‭between vaccines and autism.‬
‭ reatment for autism usually combines behavioral therapies, interventions, and‬
T
‭medications to treat other disorders common to people with autism (depression, anxiety,‬
‭o bsessive-compulsive disorder). Although early interventions can help mitigate the effects‬
‭o f the disease, there is currently no cure for ASD.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Attention Deficit Hyperactivity Disorder (ADHD)‬

‭ pproximately three to five percent of children and adults are affected by‬
A
‭attention-deficit/hyperactivity disorder (ADHD)‬‭. Like‬‭ASD, ADHD is more prevalent in‬
‭males than females. Symptoms of the disorder include inattention (lack of focus), executive‬
‭functioning difficulties, impulsivity, and hyperactivity beyond what is characteristic of the‬
‭normal developmental stage. Some patients do not have the hyperactive component of‬
‭symptoms and are diagnosed with a subtype of ADHD: attention deficit disorder (ADD).‬
‭Many people with ADHD also show comorbidity in that they develop secondary disorders‬
‭in addition to ADHD. Examples include depression or obsessive-compulsive disorder‬
‭(OCD).‬‭Figure 17.23‬‭provides some statistics concerning‬‭comorbidity with ADHD.‬
‭ he cause of ADHD is unknown, although research points to a delay and dysfunction in‬
T
‭the development of the prefrontal cortex and disturbances in neurotransmission.‬
‭According to studies of twins, the disorder has a substantial genetic component. Several‬
‭candidate genes may contribute to the disorder, but no definitive links have been‬
‭discovered. Environmental factors, including exposure to certain pesticides, may also‬
‭contribute to the development of ADHD in some patients. Treatment for ADHD often‬
‭involves behavioral therapies and prescription stimulant medications, which paradoxically‬
‭cause a calming effect in these patients.‬

‭ igure 17.23‬ ‭Many people with ADHD have one or more‬‭o ther neurological disorders.‬
F
‭(credit “chart design and illustration”: modification of work by Leigh Coriale; credit “data”:‬
‭Drs. Biederman and Faraone, Massachusetts General Hospital;‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Mental Illnesses‬
‭ ental illnesses‬‭are nervous system disorders that result in problems with thinking,‬
M
‭mood, or relating to others. These disorders are severe enough to affect a person’s quality‬
‭o f life and often make it difficult for people to perform the routine tasks of daily living.‬
‭Debilitating mental disorders plague approximately 12.5 million Americans (about 1 in 17‬
‭people) at an annual cost of more than $300 billion. Several types of mental disorders‬
‭include schizophrenia, major depression, bipolar disorder, anxiety disorders and phobias,‬
‭post-traumatic stress disorders, and obsessive-compulsive disorder (OCD), among others.‬
‭The American Psychiatric Association publishes the Diagnostic and Statistical Manual of‬
‭Mental Disorders (DSM), which describes the symptoms required for a patient to be‬
‭diagnosed with a particular mental disorder. Each newly released version of the DSM‬
‭contains different symptoms and classifications as scientists learn more about these‬
‭disorders, their causes, and how they relate. A more detailed discussion of two mental‬
‭illnesses—schizophrenia and major depression—is given below.‬

‭Schizophrenia‬
‭ chizophrenia‬‭is a severe and often debilitating mental‬‭illness affecting one percent of‬
S
‭people in the United States. Symptoms of the disease include the inability to differentiate‬
‭between reality and imagination, inappropriate and unregulated emotional responses,‬
‭difficulty thinking, and problems with social situations. People with schizophrenia can‬
‭suffer from hallucinations and hear voices; they may also suffer from delusions. Patients‬
‭also have so-called “negative” symptoms like a flattened emotional state, loss of pleasure,‬
‭and loss of fundamental drives. Many schizophrenic patients are diagnosed in their late‬
‭adolescence or early 20s. The development of schizophrenia is thought to involve‬
‭malfunctioning dopaminergic neurons and may also include problems with glutamate‬
‭signaling. Treatment for the disease usually requires antipsychotic medications that block‬
‭dopamine receptors and decrease dopamine neurotransmission in the brain. This‬
‭decrease in dopamine can cause Parkinson’s disease-like symptoms in some patients.‬
‭While some classes of antipsychotics can be quite effective at treating the disease, they are‬
‭not a cure, and most patients must remain medicated for the rest of their lives.‬
‭ or additional information about this disease, click the link below or scan the QR code to‬
F
‭watch the TED-Ed video by Anees Bahji titled “What is schizophrenia?”.‬

What is schizophrenia? - Anees Bahji

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Depression‬
‭ ajor depression‬‭affects approximately 6.7 percent‬‭o f adults in the United States each‬
M
‭year and is one of the most common mental disorders. To be diagnosed with major‬
‭depressive disorder, a person must have experienced a severely depressed mood lasting‬
‭longer than two weeks along with other symptoms, including a loss of enjoyment in‬
‭activities that were previously enjoyed, changes in appetite and sleep schedules, difficulty‬
‭concentrating, feelings of worthlessness, and suicidal thoughts. For additional information‬
‭about this disorder, click the link below or scan the QR code to watch the TED-Ed video by‬
‭Helen M. Farrell titled “What is depression?”.‬

What is depression? - Helen M. Farrell

‭ he exact causes of major depression are unknown and likely include genetic and‬
T
‭environmental risk factors. Some research supports the “classic monoamine hypothesis,”‬
‭which suggests that a decrease in norepinephrine and serotonin neurotransmission‬
‭causes depression. One argument against this hypothesis is that some antidepressant‬
‭medications cause an increase in norepinephrine and serotonin release within a few‬
‭hours of beginning treatment—but clinical results of these medications are not seen until‬
‭weeks later. This observation has led to alternative hypotheses: for example, dopamine‬
‭may also be decreased in depressed patients, or it may be an increase in norepinephrine‬
‭and serotonin that causes the disease, and antidepressants force a feedback loop that‬
‭reduces this release. Treatments for depression include psychotherapy, electroconvulsive‬
‭therapy, deep-brain stimulation, and prescription medications.‬
‭ here are several classes of‬‭antidepressant‬‭drugs‬‭that work through different‬
T
‭mechanisms. For example, monoamine oxidase inhibitors (MAO inhibitors) block the‬
‭enzyme degrading many neurotransmitters (including dopamine, serotonin, and‬
‭norepinephrine), increasing neurotransmitters in the synaptic cleft. Selective serotonin‬
‭reuptake inhibitors (SSRIs) block the reuptake of serotonin into the presynaptic neuron.‬
‭This blockage results in an increase in serotonin in the synaptic cleft. Other drugs, such as‬
‭norepinephrine-dopamine reuptake inhibitors and norepinephrine-serotonin reuptake‬
‭inhibitors, are also used to treat depression.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ or additional details about the mechanisms of action of antidepressants, click the link or‬
F
‭scan the QR code below to watch the TED-Ed video by Neil R. Jeyasingam titled “How do‬
‭antidepressants work?”.‬

How do antidepressants work? - Neil R. Jeyasingam

‭Other Neurological Disorders‬

S‭ everal other neurological disorders cannot be easily placed in the above categories.‬
‭These include chronic pain conditions, nervous system cancers, epilepsy disorders, and‬
‭stroke. For additional details about pain and brain function, click the link or scan the QR‬
‭code to watch the TED-Ed video by Karen D. Davis titled “How does your brain respond to‬
‭pain?”‬

How does your brain respond to pain? - Karen D. Davis

‭Epilepsy and stroke are discussed in more detail in the following sections.‬

‭Epilepsy‬
‭ stimates suggest that up to three percent of people in the United States will be diagnosed‬
E
‭with‬‭epilepsy‬‭in their lifetime. Recurrent seizures‬‭characterize all types of epilepsy.‬
‭Epilepsy can be a symptom of a brain injury, disease, or other illness. For example, people‬
‭with an intellectual disability or ASD can experience seizures, presumably because the‬
‭developmental wiring malfunctions that caused their disorders also put them at risk for‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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e‭ pilepsy. For many patients, however, the cause of their epilepsy is never identified and is‬
‭likely to be a combination of genetic and environmental factors. Often, seizures are‬
‭controlled with anticonvulsant medications. However, for very severe cases, patients may‬
‭undergo brain surgery to remove the brain area where seizures originate.‬
‭ or more information about this disease and identification of the seizure's associated‬
F
‭brain region, click the link below or scan the QR code to watch the TED-Ed video by Moran‬
‭Cerf titled “What if we could look inside human brains?”.‬

What if we could look inside human brains? - Moran Cerf

‭Stroke‬
‭ ‬‭stroke‬‭results when blood fails to reach a portion‬‭o f the brain long enough to cause‬
A
‭damage. Without the oxygen supplied by blood flow, neurons in this brain region die. This‬
‭neuronal death can cause many different symptoms—depending on the brain area‬
‭affected— including headache, muscle weakness or paralysis, speech disturbances,‬
‭sensory problems, memory loss, and confusion. Blood clots often cause strokes and can‬
‭also be caused by the bursting of a weak blood vessel. Strokes are prevalent and are the‬
‭third most common cause of death in the United States. On average, one person‬
‭experiences a stroke every 40 seconds in the United States. Approximately 75 percent of‬
‭strokes occur in people older than 65. Risk factors for stroke include high blood pressure,‬
‭diabetes, high cholesterol, and a family history of stroke. Because a stroke is a medical‬
‭emergency, patients with symptoms of a stroke should immediately go to the emergency‬
‭room, where they can receive drugs that will dissolve any clot that may have formed.‬
‭These drugs will not work if the stroke was caused by a burst blood vessel or if the stroke‬
‭o ccurred more than three hours before arriving at the hospital. Treatment following a‬
‭stroke can include blood pressure medication (to prevent future strokes) and (sometimes‬
‭intense) physical therapy.‬
‭ or additional information about strokes, click the link below or scan the QR code to‬
F
‭watch the TED-Ed video by Vaibhav Goswami titled “What happens during a stroke?”.‬

What happens during a stroke? - Vaibhav Goswami

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭17.8 Career Connection - Neurologist‬

‭ eurologists are physicians who specialize in disorders of the nervous system. They‬
N
‭diagnose and treat disorders such as epilepsy, stroke, dementia, nervous system injuries,‬
‭Parkinson’s disease, sleep disorders, and multiple sclerosis. Neurologists are medical‬
‭doctors who have attended college and medical school and completed three to four years‬
‭o f neurology residency.‬

‭ hen examining a new patient, a neurologist takes an entire medical history and‬
W
‭performs a complete physical exam. The physical exam contains specific tasks to‬
‭determine what brain, spinal cord, or peripheral nervous system areas may be damaged.‬
‭For example, to check whether the hypoglossal nerve is functioning correctly, the‬
‭neurologist will ask the patient to move their tongue in different ways. Suppose the patient‬
‭does not have complete control over tongue movements. In that case, the hypoglossal‬
‭nerve may be damaged, or there may be a lesion in the brainstem where the cell bodies of‬
‭these neurons reside (or there could be damage to the tongue muscle itself).‬

‭ eurologists have other tools besides a physical exam they can use to diagnose particular‬
N
‭problems in the nervous system. Suppose the patient has had a seizure, for example. In‬
‭that case, the neurologist can use electroencephalography (EEG), which involves taping‬
‭electrodes to the scalp to record brain activity to determine which brain regions are‬
‭involved in the seizure. In suspected stroke patients, a neurologist can use a computerized‬
‭tomography (CT) scan, a type of X-ray, to look for bleeding in the brain or other health‬
‭conditions.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ hapter 18: Sensation, Somatosensation, and‬
C
‭Special Senses‬

‭ igure 18.1‬ ‭This shark uses its senses of sight,‬‭vibration, and smell to hunt, but it also‬
F
‭relies on its ability to sense the electric fields of prey, a sense not present in most land‬
‭animals. (credit: modification of work by Hermanus Backpackers Hostel, South Africa;‬
‭OpenStax‬‭)‬

‭18.1 Introduction‬
‭ ll animals have a sensory system, the development of which has been driven by natural‬
A
‭selection, which is the survival of organisms with favorable traits in a particular‬
‭environment and more successful reproduction. Thus, sensory systems differ among‬
‭species according to the demands of their environments. Animals’ senses are constantly at‬
‭work, making them aware of stimuli, such as light, sound, or the presence of a chemical‬
‭substance in the external environment. They also monitor information about the‬
‭o rganism’s internal environment. The shark pictured above (‬‭Figure 18.1‬‭) can perceive‬
‭natural electrical stimuli produced by other animals in its environment, a sense called‬
‭electroreception. This enhanced ability to sense prey gives the shark an evolutionary‬
‭advantage over other fish. While it is helpful to this underwater predator, electroreception‬
‭is a sense not found in most land animals.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭.‬

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‭18.2 Sensory Processes‬

S‭ enses provide information about the body and its environment via‬‭sensation‬‭. Humans‬
‭possess general senses called somatosensation, which respond to stimuli like temperature,‬
‭pain, pressure, and vibration. We also have several commonly known special senses: taste,‬
‭smell, hearing, and vision. Although the sensory systems associated with these senses are‬
‭very different, all share a common function: to convert a sensation (such as light, sound,‬
‭o r the body's position) into an electrical signal in the nervous system via‬‭sensory‬
‭transduction‬‭.‬‭Two broad types of cellular systems‬‭perform sensory transduction. In one,‬
‭a neuron works with a‬‭sensory receptor‬‭, cell, or specialized‬‭cell process to engage with‬
‭and detect a specific stimulus. Stimulation of the sensory receptor activates the associated‬
‭afferent neuron, which carries information about the stimulus to the central nervous‬
‭system. In the second type of sensory transduction, a‬‭sensory nerve ending‬‭responds to a‬
‭stimulus in the internal or external environment: this neuron constitutes the sensory‬
‭receptor. Several stimuli can stimulate free nerve endings, thus showing little receptor‬
‭specificity. For example, temperature changes, chemical stimulation, or pressure may‬
‭stimulate pain receptors in your gums and teeth.‬

‭Reception‬
‭ he first step in sensation is‬‭reception‬‭, which is‬‭the activation of sensory receptors by‬
T
‭mechanical stimuli (being bent or squished, for example), chemicals, or temperature. The‬
‭receptor can then respond to the stimuli. The region in space in which a given sensory‬
‭receptor can respond to a stimulus, be it far away or in contact with the body, is that‬
‭receptor’s‬‭receptive field‬‭. Think briefly about the‬‭differences in receptive fields for the‬
‭different senses. For the sense of touch, a stimulus must come into contact with the body.‬
‭For the sense of hearing, a stimulus can be a moderate distance away (some baleen whale‬
‭sounds can propagate for many kilometers). For vision, a stimulus can be very far away;‬
‭for example, the visual system perceives light from stars at enormous distances.‬

‭Transduction‬
‭ he second step in sensation is‬‭transduction‬‭, which‬‭is translating a sensory signal into an‬
T
‭electrical signal in the nervous system. This occurs at the sensory receptor, and the change‬
‭in electrical potential is called the‬‭receptor potential‬‭.‬‭How is sensory input, such as‬
‭pressure on the skin, changed to a receptor potential? In this example, a type of receptor‬
‭called a mechanoreceptor (‬‭Figure 18.2‬‭) possesses specialized‬‭membranes that respond‬
‭to pressure. Disturbance of these dendrites by compressing or bending them opens gated‬
‭ion channels in the plasma membrane of the sensory neuron, changing its electrical‬
‭potential. Recall that in the nervous system, a positive change in a neuron’s electrical‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ otential (also called the membrane potential) depolarizes the neuron. Receptor‬
p
‭potentials are graded potentials: the magnitude of these graded (receptor) potentials‬
‭varies with the strength of the stimulus. If the magnitude of depolarization is sufficient‬
‭(that is, if membrane potential reaches a threshold), the neuron will fire‬
‭an action potential.‬

‭ igure 18.2‬ ‭(a) Mechanosensitive ion channels are‬‭gated ion channels that respond to‬
F
‭mechanical deformation of the plasma membrane. Hair-like tethers connect A‬
‭mechanosensitive channel to the plasma membrane and the cytoskeleton. When pressure‬
‭causes the extracellular matrix to move, the channel opens, allowing ions to enter or exit‬
‭the cell. (b) Stereocilia in the human ear are connected to mechanosensitive ion channels.‬
‭When a sound causes the stereocilia to move, mechanosensitive ion channels transduce‬
‭the signal to the cochlear nerve. (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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S‭ ensory receptors for different senses are very different from each other, and they are‬
‭specialized according to the type of stimulus they sense. For example, touch receptors,‬
‭light receptors, and sound receptors are each activated by different stimuli. Touch‬
‭receptors are not sensitive to light or sound but only to touch or pressure. However,‬
‭stimuli may be combined at higher levels in the brain, as happens with our sense of smell‬
‭contributing to our sense of taste.‬
S‭ ensory systems encode four aspects of sensory information: the type of stimulus, the‬
‭location of the stimulus in the receptive field, the duration of the stimulus, and the relative‬
‭intensity of the stimulus. Thus, action potentials transmitted over a sensory receptor’s‬
‭afferent axons encode one type of stimulus, and this segregation of the senses is‬
‭preserved in other sensory circuits. For example, auditory receptors transmit signals over‬
‭a dedicated system, and the brain interprets electrical activity in the axons of the auditory‬
‭receptors as an auditory stimulus—a sound.‬
‭ he intensity of a stimulus is often encoded in the rate of action potentials produced by‬
T
‭the sensory receptor. Thus, an intense stimulus will create a more rapid train of action‬
‭potentials, and reducing the stimulus will likewise slow the production rate of action‬
‭potentials. A second way in which intensity is encoded is by the number of receptors‬
‭activated. An intense stimulus might initiate action potentials in many adjacent receptors,‬
‭while a less intense stimulus might stimulate fewer receptors. Integration of sensory‬
‭information begins as soon as the information is received in the CNS, and the brain will‬
‭further process incoming signals.‬

‭Perception‬
‭ he third step in sensation is‬‭perception‬‭, an individual’s‬‭interpretation of a sensation.‬
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‭Although perception relies on the activation of sensory receptors, perception happens not‬
‭at the sensory receptor level but at higher levels in the central nervous system. The brain‬
‭distinguishes sensory stimuli through a sensory pathway: action potentials from sensory‬
‭receptors travel along neurons dedicated to a particular stimulus. These neurons are‬
‭dedicated to that specific stimulus and synapse with certain neurons in the brain or spinal‬
‭cord.‬
‭ ll sensory signals, except those from the olfactory system, are transmitted through the‬
A
‭central nervous system and routed to the thalamus and the appropriate cortex region. The‬
‭thalamus‬‭is a structure in the forebrain that serves‬‭as a clearinghouse and relay station‬
‭for sensory (as well as motor) signals. When the sensory signal exits the thalamus, it is‬
‭conducted to the specific area of the‬‭cortex‬‭(‬‭Figure‬‭18.3‬‭) dedicated to processing that‬
‭particular sense.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ ow are neural signals interpreted? The interpretation of sensory signals between‬

H
‭individuals of the same species is mainly similar, owing to the inherited similarity of their‬
‭nervous systems; however, there are some individual differences. An excellent example is‬
‭individual tolerance to a painful stimulus, such as dental pain, which certainly differs.‬

‭ igure 18.3‬ ‭In humans, except for olfaction, all‬‭sensory signals are routed from the (a)‬
F
‭thalamus to (b) final processing regions in the brain's cortex. (credit b: modification of‬
‭work by Polina Tishina;‬‭OpenStax‬‭)‬

‭18.3 Somatosensation‬
‭ omatosensation‬‭is a mixed sensory category and includes‬‭all sensations received from‬
S
‭the skin and mucous membranes, as well as from the limbs and joints. Somatosensation‬
‭o ccurs all over the body's exterior and at some interior locations. Various receptor‬
‭types—embedded in the skin, mucous membranes, muscles, joints, internal organs, and‬
‭cardiovascular system—play a role.‬
‭ or an introduction to somatosensation and our general senses, as well as an interesting‬
F
‭story, click the link below or scan the QR code to watch the TED-Ed video by Antonio‬
‭Cataldo titled “The man who lost his sense of touch.”‬

The man who lost his sense of touch - Antonio Cataldo

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ he‬‭skin‬‭, the largest organ in the human body, is a vital component of the‬‭integumentary‬
T
‭system‬‭. Its complex structure includes different types of receptors‬‭that function in‬
‭somatosensation. For an introduction to this organ and its role in somatosensation, click‬
‭the link on the next page or scan the QR code to watch the TED-Ed video by Emma Bryce‬
‭titled “The science of skin.”‬

The science of skin - Emma Bryce

‭ ne skin-related phenomenon that is relevant to everyone is itching. But why do we itch‬

O
‭o ur skin? For an answer to this question, click the link below or scan the QR code to‬
‭watch the TED-Ed video by Emma Bryce titled “Why do we itch?”‬

Why do we itch? - Emma Bryce

S‭ omatosensory receptors are classified into five categories: chemoreceptors,‬

‭o smoreceptors, mechanoreceptors, thermoreceptors, and pain receptors. These‬
‭categories are based on the nature of stimuli each receptor class transduces.‬
‭Chemoreceptors‬‭interpret chemical stimuli, such as‬‭an object’s taste or smell.‬
‭Osmoreceptors‬‭respond to solute concentrations of‬‭body fluids. Physical stimuli, such as‬
‭pressure and vibration, as well as the sensation of sound and body position (balance), are‬
‭interpreted through‬‭mechanoreceptors‬‭. Another physical‬‭stimulus with its own receptor‬
‭type is temperature, which is sensed through‬‭thermoreceptors‬‭that are either sensitive to‬
‭temperatures above (heat) or below (cold) normal body temperature.‬
‭ ain receptors‬‭function to detect injurious stimuli‬‭in response to tissue damage. Pain is‬
P
‭caused by actual sources of injury, such as contact with a heat source that causes a‬
‭thermal burn or contact with a corrosive chemical. But pain also can be caused by‬
‭harmless stimuli that mimic the action of damaging stimuli, such as contact with‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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c‭ apsaicinoids, the compounds that cause peppers to taste hot and which are used in‬
‭self-defense pepper sprays and certain topical medications. Peppers taste “hot” because‬
‭the protein receptors that bind capsaicin open the same calcium channels activated by‬
‭warm receptors.‬

‭ or more information about pain and how our brains perceive pain, click the links or scan‬
F
‭the QR codes to watch two TED-Ed videos: “The mysterious science of pain,” by Joshua W.‬
‭Pate and “How does your brain respond to pain?” by Karen D. Davis.‬

The mysterious science of pain - Joshua W. Pate

How does your brain respond to pain? - Karen D. Davis

‭18.4 Smell and Taste‬

S‭ mell, also called‬‭olfaction‬‭, and taste, also called‬‭gustation‬‭, are the most interconnected‬
‭senses in that both involve molecules of the stimulus entering the body and binding to‬
‭receptors. Smell lets an animal sense the presence of food or other animals—whether‬
‭potential mates, predators, or prey—or other environmental chemicals that can impact‬
‭their survival. Similarly, the sense of taste allows animals to discriminate between types of‬
‭foods. While the value of a sense of smell is obvious, what is the value of a sense of taste?‬
‭Different tasting foods have various attributes, both helpful and harmful. For example,‬
‭sweet-tasting substances tend to be highly caloric, which could be necessary for survival in‬
‭lean times. Bitterness is associated with toxicity, and sourness is associated with spoiled‬
‭food. Salty foods are valuable in maintaining homeostasis by helping the body retain water‬
‭and providing ions necessary for cells to function.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Odors and Tastes‬

‭ ll odors that we perceive are molecules in the air we breathe. Therefore, if a substance‬
A
‭o r organism doesn’t release molecules into the air, it has no smell. Also, if a human does‬
‭not have a receptor that recognizes a specific molecule, then that molecule elicits no smell.‬
‭Humans have about 350 olfactory receptor subtypes that work in various combinations,‬
‭allowing us to sense about 10,000 different odors. Compare that to mice, which have‬
‭about 1,300 olfactory receptor types and probably detect more odors.‬
‭ astes are generated in response to ingested molecules, and the primary tastes detected‬
T
‭by humans are sweet, sour, bitter, salty, and umami. The first four tastes need little‬
‭explanation. The identification of‬‭umami‬‭as a fundamental‬‭taste occurred fairly recently; it‬
‭was identified in 1908 by Japanese scientist Kikunae Ikeda while he worked with seaweed‬
‭broth. However, it was only widely accepted as a taste that could be physiologically‬
‭distinguished many years later. The taste of umami, also known as savoriness, is‬
‭attributable to the taste of the amino acid L-glutamate. Monosodium glutamate, or MSG, is‬
‭o ften used in cooking to enhance the savory taste of certain foods. What is the adaptive‬
‭value of being able to distinguish umami? Savory substances tend to be high in protein, a‬
‭necessary nutrient.‬
‭ oth odors and tastes involve molecules that stimulate specific chemoreceptors. Although‬
B
‭humans commonly distinguish smell as one sense and taste as another, they work‬
‭together to create the perception of flavor. A person’s perception of flavor is reduced if‬
‭they have congested nasal passages.‬

‭Odor Reception and Transduction‬

‭ dorants‬‭(odor molecules) enter the nose and dissolve‬‭in the olfactory epithelium, the‬
O
‭mucosa at the back of the nasal cavity, as illustrated in‬‭Figure 18.4‬‭. The‬‭olfactory‬
‭epithelium‬‭is a collection of specialized olfactory‬‭receptors in this region that spans an‬
‭area of about 5 cm‬‭2‬ ‭(approximately 0.75 in‬‭2‭)‬ in humans.‬‭Recall that sensory cells are‬
‭neurons. An‬‭olfactory receptor‬‭, a dendrite of a specialized‬‭neuron, responds when it‬
‭binds specific molecules inhaled from the environment and sends impulses directly to the‬
‭brain's olfactory bulb. Humans have about 12 million olfactory receptors, distributed‬
‭among hundreds of receptor types that respond to different odors. Twelve million seems‬
‭like a large number of receptors, but compare that to other animals: rabbits have about‬
‭100 million, most dogs have about 1 billion, and bloodhounds—dogs selectively bred for‬
‭their sense of smell—have about 4 billion. The overall size of the olfactory epithelium also‬
‭differs between species, with that of bloodhounds, for example, being many times larger‬
‭than that of humans.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ lfactory neurons are bipolar neurons, meaning they have two processes extending from‬
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‭the cell body. Each neuron has a single dendrite buried in the olfactory epithelium, and‬
‭extending from this dendrite are five to twenty receptor-laden, hair-like cilia that trap‬
‭o dorant molecules. The sensory receptors on the cilia are proteins, and the variations in‬
‭their amino acid chains make the receptors sensitive to different odorants. Each olfactory‬
‭sensory neuron has only one type of receptor on its cilia, and the receptors are‬
‭specialized to detect specific odorants, so the bipolar neurons themselves are specialized.‬
‭When an odorant binds with a receptor that recognizes it, the sensory neuron associated‬
‭with the receptor is stimulated, generating an action potential. Olfactory stimulation is the‬
‭o nly sensory information directly reaching the brain’s cerebral cortex, whereas other‬
‭sensations are relayed through the thalamus.‬

‭ igure 18.4‬ ‭In the human olfactory system, (a) bipolar‬‭o lfactory neurons extend from (b)‬
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‭the olfactory epithelium, where olfactory receptors are located, to the olfactory bulb.‬
‭(credit: modification of work by Patrick J. Lynch, medical illustrator; C. Carl Jaffe, MD,‬
‭cardiologist;‬‭OpenStax‬‭)‬

‭Evolution Connection‬

‭ heromones‬
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‭A‬‭pheromone‬‭is a chemical released by an animal that‬‭affects the behavior or physiology‬
‭o f animals of the same species. Pheromonal signals can profoundly affect animals that‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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i‭nhale them, but pheromones apparently are not consciously perceived in the same way as‬
‭o ther odors. There are several different types of pheromones, which are released in urine‬
‭o r as glandular secretions. Specific pheromones are attractants to potential mates, others‬
‭are repellants to potential competitors of the same sex, and still others play roles in‬
‭mother-infant attachment. Some pheromones can also influence the timing of puberty,‬
‭modify reproductive cycles, and even prevent embryonic implantation. While the roles of‬
‭pheromones in many nonhuman species are essential, pheromones have become less‬
‭important in human behavior over evolutionary time compared to their importance to‬
‭o rganisms with more limited behavioral repertoires.‬

‭ he vomeronasal organ (VNO) is a tubular, fluid-filled, olfactory organ in many vertebrate‬

T
‭animals that sits adjacent to the nasal cavity. It is very sensitive to pheromones and is‬
‭connected to the nasal cavity by a duct. When molecules dissolve in the mucosa of the‬
‭nasal cavity, they then enter the VNO, where the pheromone molecules among them bind‬
‭with specialized pheromone receptors. Upon exposure to pheromones from their species‬
‭o r others, many animals, including cats, may display the flehmen response, shown in‬
‭Figure 18.5‬‭, which is a curling of the upper lip that‬‭helps pheromone molecules enter the‬
‭VNO.‬

‭ hile some scientists assert that the VNO is apparently functionally vestigial in humans,‬
W
‭even though a similar structure is located near human nasal cavities, others are‬
‭researching it as a possible functional system that may, for example, contribute to the‬
‭synchronization of menstrual cycles in women living in close proximity.‬

‭ igure 18.5‬‭The flehmen response in this tiger results‬‭in the curling of the upper lip and‬
F
‭helps airborne pheromone molecules enter the vomeronasal organ. (credit: modification‬
‭o f work by "chadh"/Flickr;‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ or additional information about our sense of smell, click the link below or scan the QR‬
F
‭code to watch the TED-Ed video by Rose Eveleth‬‭titled‬‭“How do we smell?”‬

How do we smell? - Rose Eveleth

‭Taste Reception and Transduction‬

‭ he primary organ of taste is the‬‭taste bud‬‭, a cluster‬‭o f gustatory receptors (taste cells)‬
T
‭located within the bumps on the tongue called‬‭papillae‬‭(singular: papilla), which are‬
‭illustrated in‬‭Figure 18.6‬‭. There are several structurally‬‭distinct papillae. Filiform papillae,‬
‭which are located across the tongue, are tactile, providing friction that helps the tongue‬
‭move substances and contain no taste cells. In contrast, fungiform papillae, located mainly‬
‭o n the anterior two-thirds of the tongue, contain one to eight taste buds and have‬
‭receptors for pressure and temperature. The large circumvallate papillae contain up to‬
‭250 taste buds and form a V near the posterior edge of the tongue.‬

‭(a) (b)‬

‭ igure 18.6‬ ‭(a) Foliate, circumvallate, and fungiform‬‭papillae are located on different‬
F
‭tongue regions. (b) Foliate papillae are prominent protrusions on this light micrograph.‬
‭(credit a: modification of work by NCI; scale-bar data from Matt Russell;‬‭OpenStax‬‭)‬
I‭ n addition to those two types of chemically and mechanically sensitive papillae, foliate‬
‭papillae—leaf-like papillae located in parallel folds along the edges and toward the back of‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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t‭ he tongue. Foliate papillae contain about 1,300 taste buds within their folds. Lastly, there‬
‭are circumvallate papillae, wall-like papillae in the shape of an inverted “V” at the back of‬
‭the tongue. A groove surrounds each of these papillae and contains about 250 taste buds.‬
‭ ach taste bud’s taste cells are replaced every 10 to 14 days. These are elongated cells‬
E
‭with hair-like processes called microvilli at the tips that extend into the taste bud pore‬
‭(‬‭Figure 18.7‬‭). Food molecules (‬‭tastants‬‭) are dissolved‬‭in saliva and bind with and‬
‭stimulate the receptors on the microvilli. The receptors for tastants are located across the‬
‭o uter portion and front of the tongue, outside of the middle area where the filiform‬
‭papillae are most prominent.‬

‭ igure 18.7‬ ‭Pores in the tongue allow tastants to‬‭enter. (credit: modification of work by‬
F
‭Vincenzo Rizzo;‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

I‭ n humans, there are five primary tastes, each with only one corresponding receptor type.‬
‭Thus, like olfaction, each receptor is specific to its stimulus (tastant). Transduction of the‬
‭five tastes happens through different mechanisms that reflect the molecular composition‬
‭o f the tastant. For example, salty tastants (containing NaCl) provide the sodium ions (Na‬‭+‭)‬ ‬
‭that enter the taste neurons and excite them directly. Sour tastants are acids and belong to‬
‭the thermoreceptor protein family. The binding of an acid or other sour-tasting molecule‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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t‭ riggers a change in the ion channel, increasing hydrogen ion (H‬‭+‭)‬ concentrations in the‬
‭taste neurons, thus depolarizing them.‬
‭ oth tasting abilities and sense of smell change with age. In humans, the senses decline‬
B
‭dramatically by age 50 and continue to decline. A child may find a specific food too spicy,‬
‭whereas an older person may find the same food bland and unappetizing.‬
‭ or additional information about our sense of taste, click the link below or scan the QR‬
F
‭code to watch the Neuro Transmissions video titled “How do we taste?”‬

How Do We Taste?

‭ or additional information about the interrelationship between smell and taste, click the‬
F
‭link below or scan the QR code to watch the TED-Ed video by Jen Gunter‬‭titled “How your‬
‭sense of smell helps you savor flavor.”‬

How Your Sense of Smell Helps You Savor Flavor | Body Stuff …

‭18.5 Hearing‬
‭ udition‬‭, or hearing, is essential for many human‬‭and other animal interactions. It‬
A
‭enables an organism to detect and receive information about danger, such as an‬
‭approaching predator, and to participate in communal exchanges like those concerning‬
‭territories or mating.‬

‭Sound‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ uditory stimuli are sound waves, which are mechanical pressure waves that move‬
A
‭through a medium, such as air or water. There are no sound waves in a vacuum since‬
‭there are no air molecules to move in waves. The speed of sound waves differs based on‬
‭altitude, temperature, and medium, but at sea level and a temperature of 68 F, sound‬
‭waves travel in the air at about 1,125 feet per second.‬
‭ s is true for all waves, a sound wave has four primary characteristics: frequency,‬
A
‭wavelength, period, and amplitude. Frequency is the number of waves per unit of time,‬
‭and, in sound, is heard as pitch. High-frequency (≥15.000Hz) sounds are higher-pitched‬
‭(short wavelength) than low-frequency (long wavelengths; ≤100Hz) sounds. Frequency is‬
‭measured in cycles per second, and for sound, the most commonly used unit is hertz (Hz),‬
‭o r cycles per second. Most humans can perceive sounds with frequencies between 30 and‬
‭20,000 Hz. Women are typically better at hearing high frequencies, but everyone’s ability‬
‭to hear high frequencies decreases with age. Dogs detect up to about 40,000 Hz; cats,‬
‭60,000 Hz; bats, 100,000 Hz; and dolphins, 150,000 Hz. Those frequencies above the‬
‭human range are called‬‭ultrasound‬‭.‬
‭ mplitude, or the dimension of a wave from peak to trough, in sound is heard as volume‬
A
‭and is illustrated in‬‭Figure 18.8‬‭. The sound waves‬‭o f louder sounds have greater‬
‭amplitude than softer sounds. For sound, volume is measured in decibels (dB). The softest‬
‭sound that a human can hear is the zero point. Humans usually speak at approximately 60‬
‭decibels.‬

‭ igure 18.8‬ ‭For sound waves, wavelength corresponds‬‭to pitch. The amplitude of the‬
F
‭wave corresponds to volume. The sound wave shown with a dashed line is softer in‬
‭volume than the sound wave shown with a solid line. (credit: NIH;‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Sound Reception‬
I‭ n mammals, sound waves are collected by the external, cartilaginous part of the ear called‬
‭the‬‭pinna‬‭, then travel through the auditory canal‬‭and cause vibrations of the thin‬
‭diaphragm (or membrane) called the‬‭tympanum,‬‭o r ear‬‭drum, which is the innermost‬
‭part of the‬‭outer ear‬‭(‬‭Figure 18.9‬‭). Interior to the‬‭tympanum is the‬‭middle ear‬‭. The‬
‭middle ear holds three tiny bones called the‬‭ossicles‬‭,‬‭which transfer energy from the‬
‭moving tympanum to the inner ear. The three ossicles are the‬‭malleus‬‭(also known as the‬
‭hammer), the‬‭incus‬‭(the anvil), and the‬‭stapes‬‭(the‬‭stirrup). The aptly named stapes looks‬
‭very much like a stirrup on a saddle. The three ossicles are unique to mammals, and each‬
‭plays a role in hearing.‬
‭ he malleus attaches at three points to the interior surface of the tympanic membrane.‬
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‭The incus attaches the malleus to the stapes. In humans, the stapes isn’t long enough to‬
‭reach the tympanum. If we did not have the malleus and the incus, the tympanum‬
‭vibrations would never reach the inner ear. These bones also function to collect force and‬
‭amplify sounds. The ear ossicles are homologous to bones in a fish mouth: the bones that‬
‭support gills in fish are thought to be adapted for use in the vertebrate ear over‬
‭evolutionary time. Many animals (frogs, reptiles, and birds, for example) use the stapes of‬
‭the middle ear to transmit vibrations to the middle ear.‬

‭ igure 18.9‬ ‭Sound travels through the outer ear to‬‭the middle ear, which is bounded on‬
F
‭its exterior by the tympanic membrane. The middle ear contains three bones called‬
‭o ssicles that transfer the sound wave to the oval window, the exterior boundary of the‬
‭inner ear. The organ of Corti, which is the organ of sound transduction, lies inside the‬
‭cochlea. (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Sound Transduction‬
‭ ibrating objects, such as vocal cords, create sound waves or pressure waves in the air.‬
V
‭When these pressure waves reach the ear, the ear transduces, or changes, this mechanical‬
‭stimulus (pressure wave) into a nerve impulse (electrical signal) that the brain perceives‬
‭as sound. The pressure waves strike the tympanum, causing it to vibrate. The mechanical‬
‭energy from the moving tympanum transmits the vibrations to the three bones of the‬
‭middle ear. The stapes transmits the vibrations to a thin diaphragm called the‬‭oval‬
‭window‬‭, which is the outermost structure of the‬‭inner‬‭ear‬‭. The inner ear structures are‬
‭found in the labyrinth, a bony, hollow structure that is the most interior portion of the ear.‬
‭Here, the energy from the sound wave is transferred from the stapes through the flexible‬
‭oval window and to the cochlea's fluid. The vibrations of the oval window create pressure‬
‭waves in the fluid inside the cochlea. The‬‭cochlea‬‭is a whorled structure, like a snail's‬
‭shell, and it contains receptors for the transduction of the mechanical wave into an‬
‭electrical signal (‬‭Figure 18.10‬‭). Inside the cochlea,‬‭the‬‭basilar membrane‬‭is a mechanical‬
‭analyzer that runs the length of the cochlea, curling toward the cochlea’s center.‬
‭ he mechanical properties of the basilar membrane change along its length, such that it is‬
T
‭thicker, tauter, and narrower at the outside of the whorl (where the cochlea is largest) and‬
‭thinner, floppier, and broader toward the apex, or center, of the whorl (where the cochlea‬
‭is smallest). Different regions of the basilar membrane vibrate according to the sound‬
‭wave frequency conducted through the fluid in the cochlea. For these reasons, the‬
‭fluid-filled cochlea detects different wave frequencies (pitches) at different membrane‬
‭regions. When the sound waves in the cochlear fluid contact the basilar membrane, it‬
‭flexes back and forth in a wave-like fashion.‬
‭ he site of transduction, or changing the signal from mechanical to electrical, is in the‬
T
‭organ of Corti‬‭(spiral organ). It is composed of hair‬‭cells held in place above the basilar‬
‭membrane like flowers projecting up from the soil, with their exposed short, hair-like‬
‭stereocilia‬‭contacting or embedded in the‬‭tectorial‬‭membrane‬‭above them. The inner‬
‭hair cells are the primary auditory receptors and exist in a single row, numbering‬
‭approximately 3,500. The stereocilia from inner hair cells extend into small dimples on the‬
‭tectorial membrane’s lower surface. The outer hair cells are arranged in three or four‬
‭rows. They number approximately 12,000, and they function to fine-tune incoming sound‬
‭waves. The longer stereocilia that project from the outer hair cells attach to the tectorial‬
‭membrane. All of the stereocilia are mechanoreceptors, and when bent by vibrations, they‬
‭respond by opening a gated ion channel (‬‭Figure 18.2‬‭).‬‭As a result, the hair cell membrane‬
‭is depolarized, and a signal is transmitted to the cochlear nerve. The intensity (volume) of‬
‭sound is determined by how many hair cells at a particular location are stimulated.‬
‭ ochlear implants can restore hearing in people who have a nonfunctional cochlea. The‬
C
‭implant consists of a microphone that picks up sound. A speech processor selects sounds‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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i‭n the range of human speech, and a transmitter converts these sounds‬
‭to electrical impulses, which are then sent to the auditory nerve.‬

‭ igure 18.10‬ ‭In the human ear, sound waves cause‬‭the stapes to press against the oval‬
F
‭window. Vibrations travel up the fluid-filled interior of the cochlea. The basilar membrane‬
‭that lines the cochlea gets continuously thinner toward the apex of the cochlea. Different‬
‭thicknesses of membrane vibrate in response to varying frequencies of sound. Sound‬
‭waves then exit through the round window. In the cross-section of the cochlea (top right‬
‭figure), note that the cochlea also has a middle canal in addition to the upper canal and‬
‭lower canal. The organ of Corti (bottom image) is the site of sound transduction.‬
‭Movement of stereocilia on hair cells results in an action potential that travels along the‬
‭auditory (cochlear) nerve. (credit:‬‭Opentextbc.ca‬‭).‬‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭ or additional information about hearing, click the link below or scan the QR code to‬
F
‭watch the TED-Ed video by Douglas L. Oliver‬‭titled‬‭“The science of hearing.”‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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The science of hearing - Douglas L. Oliver

‭ n ear-related medical condition that affects an estimated 10% to 25% of the US‬
A
‭population is tinnitus, or ringing in the ears. To learn more about this condition, click the‬
‭link below or scan the QR code to watch the TED-Ed video by Marc Fagelson titled “What’s‬
‭that ringing in your ears?”‬

What’s that ringing in your ears? - Marc Fagelson

‭18.6 Vision‬
‭ ision‬‭, the ability to detect light patterns from‬‭the outside environment and interpret‬
V
‭them into images, is the only photo-responsive sense. Animals are bombarded with‬
‭sensory information, and the sheer volume of visual information can be problematic.‬
‭Fortunately, the visual systems of species have evolved to attend to the most important‬
‭stimuli. The importance of vision to humans is further substantiated by the fact that about‬
‭o ne-third of the human’s cerebral cortex is dedicated to analyzing and perceiving visual‬
‭information.‬

‭Light‬
‭ s with auditory stimuli, light travels in waves. The compression waves that compose‬
A
‭sound must travel in a medium—a gas, a liquid, or a solid. In contrast, light is composed of‬
‭electromagnetic waves and needs no medium; light can travel in a vacuum. The behavior‬
‭o f light can be discussed in terms of the behavior of waves and the behavior of the‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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f‭ undamental unit of light—a packet of electromagnetic radiation called a photon. A glance‬

‭at the‬‭electromagnetic spectrum‬‭(‬‭Figure 18.11‬‭) shows‬‭that visible light for humans is‬
‭just a tiny slice of the entire spectrum (i.e., the‬‭visible region‬‭), which includes radiation‬
‭that we cannot see as light because it is below the frequency of visible red light and above‬
‭the frequency of visible violet light.‬
‭ ertain variables are important when discussing the perception of light. Wavelength‬
C
‭(which varies inversely with frequency) manifests itself as hue. Light at the red end of the‬
‭visible spectrum has longer wavelengths (and a lower frequency), while light at the violet‬
‭end has shorter wavelengths (and a higher frequency). The wavelength of light is‬
‭expressed in nanometers (nm); one nanometer is one billionth of a meter. Humans‬
‭perceive light that ranges between approximately 380 nm and 740 nm. Some other‬
‭animals, though, can detect wavelengths outside the human range. For example, bees see‬
‭near-ultraviolet light to locate nectar guides on flowers, and some non-avian reptiles‬
‭sense infrared light (heat that prey gives off).‬

‭ igure 18.11‬ ‭In the electromagnetic spectrum, visible‬‭light lies between 380 nm‬
F
‭(nanometers) and 740 nm. (credit: modification of work by NASA;‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬
‭ ave amplitude is perceived as luminous intensity or brightness. The standard unit of‬
W
‭light intensity is the‬‭c andela‬‭, which is approximately‬‭the luminous intensity of one‬
‭common candle.‬
‭ ight waves travel at approximately 984,000 feet per second in a vacuum (and somewhat‬
L
‭slower in various media such as air and water), and those waves arrive at the eye as long‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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(‭ red), medium (green), and short (blue) waves. What is termed “white light” is light that is‬
‭perceived as white by the human eye. This effect is produced by light that stimulates‬
‭equally the color receptors in the human eye. The apparent color of an object is the color‬
‭(or colors) that the object reflects. Thus, a red object reflects the red wavelengths in mixed‬
‭(white) light and absorbs all other wavelengths of light.‬

‭Light Reception‬
‭ he eye's photoreceptor cells, where light transduction to nervous impulses occurs, are‬
T
‭located in the‬‭retina‬‭(‬‭Figure 18.12‬‭) on the inner‬‭surface of the back of the eye. But light‬
‭does not impact the retina unaltered. Instead, it passes through other layers that process‬
‭it so the retina can interpret it. The‬‭cornea‬‭, the‬‭front transparent layer of the eye, and the‬
‭crystalline‬‭lens‬‭, a transparent convex structure behind‬‭the cornea, refract (bend) light to‬
‭focus the image on the retina. The‬‭iris‬‭, which is‬‭conspicuous as the colored part of the eye,‬
‭is a circular muscular ring between the lens and cornea that regulates the amount of light‬
‭entering the eye. In high ambient light conditions, the iris contracts, reducing the size of‬
‭the pupil at its center. In conditions of low light, the iris relaxes, and the pupil enlarges.‬
‭ he primary function of the lens is to focus light on the retina and a region called the‬
T
‭fovea centralis. The lens is dynamic, focusing and refocusing light as the eye rests on near‬
‭and far objects in the visual field. The lens is operated by muscles that stretch it flat or‬
‭allow it to thicken, changing the focal length of light coming through it to focus it sharply‬
‭o n the retina. With age comes the loss of the lens's flexibility, and a form of farsightedness‬
‭called‬‭presbyopia‬‭results. Presbyopia occurs because‬‭the image focuses behind the retina.‬
‭Presbyopia is a deficit similar to a type of farsightedness called‬‭hyperopia‬‭caused by an‬
‭eyeball that is too short. Images in the distance are clear for both defects, but those‬
‭nearby are blurry.‬‭Myopia‬‭(nearsightedness) occurs‬‭when an eyeball is elongated and the‬
‭image focus falls in front of the retina. In this case, images in the distance are blurry, but‬
‭images nearby are clear.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 18.12‬ ‭(a) A cross-section of the human eye.‬ ‭(b) An enlargement shows the layers‬
F
‭o f the retina. (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ or additional information about eye-focusing issues and how they can be corrected with‬
F
‭glasses, click the link below or scan the QR code to watch the TED-Ed video by Andrew‬
‭Bastawrous and Clare Gilbert‬‭titled “How do glasses‬‭help us see?”‬

How do glasses help us see? - Andrew Bastawrous and Clare Gilbert

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ or additional information about how laser eye surgery corrects focusing issues, click the‬
F
‭link on the next page or scan the QR code to watch the TED-Ed video by Dan Reinstein‬
‭titled “How does laser eye surgery work?”‬

How does laser eye surgery work? - Dan Reinstein

‭ he retina has two types of photoreceptors:‬‭rods‬‭and‬‭cones‬‭, named for their general‬

T
‭appearance (‬‭Figure 18.13‬‭). Rods are strongly photosensitive‬‭and located on the outer‬
‭edges of the retina. They detect dim light and are used primarily for peripheral and‬
‭nighttime vision. Cones are weakly photosensitive and located near the center of the‬
‭retina. They respond to bright light, and their primary role is in daytime color vision.‬

‭ igure 18.13‬ ‭Rods and cones are photoreceptors in‬‭the retina. Rods respond in low light‬
F
‭and can detect only shades of gray. Cones respond in intense light and are responsible for‬
‭color vision. (credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ he‬‭fovea‬‭is the region in the center back of the eye responsible for acute vision. The‬
T
‭fovea has a high density of cones. When you bring your gaze to an object to examine it‬
‭intently in bright light, the eyes orient so that the object’s image falls on the fovea.‬
‭However, when looking at a star in the night sky or another object in dim light, the object‬
‭can be better viewed by the peripheral vision because the rods at the edges of the retina,‬
‭rather than the cones at the center, operate better in low light. In humans, cones far‬
‭o utnumber rods in the fovea.‬

‭ or additional information about color vision, click the link on the next page or scan the‬
F
‭QR code to watch the TED-Ed video by Colm Kelleher‬‭titled “How do we see color?”‬

How we see color - Colm Kelleher

‭ or additional information about the diversity of eye structure and function in animals,‬
F
‭click the link below or scan the QR code to watch the TED-Ed video by Thomas W. Cronin‬
‭titled “Which animal has the best eyesight?”‬

Which animal has the best eyesight? - Thomas W. Cronin

‭Light Transduction‬
‭ he rods and cones are the sites of light transduction to a neural signal. Both rods and‬
T
‭cones contain photopigments. In vertebrates, the main photopigment,‬‭rhodopsin‬‭, has two‬
‭main parts‬‭Figure 27.19‬‭): an opsin, which is a membrane‬‭protein, and a retinal, a molecule‬
‭that absorbs light. When light hits a photoreceptor, it causes a shape change in the retinal,‬
‭altering its structure from a bent (‬‭cis‬‭) form of the‬‭molecule to its linear (‬‭trans‬‭) form. This‬
‭change in the retina activates rhodopsin and starts a cascade of events that end with the‬
‭closing of Na‬‭+‬ ‭channels in the photoreceptor membrane.‬‭Thus, unlike most other sensory‬
‭neurons (which become depolarized by exposure to a stimulus), visual receptors become‬
‭hyperpolarized and, therefore, driven away from the threshold (‬‭Figure 18.14‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭18.7 Career Connections - Audiologist, Orthoptist, and Optometrists‬

‭Audiologist‬

‭ udiologists are healthcare professionals who diagnose, manage, and treat hearing,‬
A
‭balance, and ear problems. They work in audiology, the science of hearing and balance.‬
‭They determine the severity and type of hearing loss a patient has and develop a‬
‭treatment plan.‬

‭ udiologists counsel patients, manage hearing loss prevention programs, assist patients‬
A
‭with ringing in the ears (tinnitus), and design educational plans for children. They‬
‭specialize in hearing aids, inner ear implants, and assistive listening devices.‬

‭ udiologists work with doctors, speech-language pathologists, physical therapists,‬

A
‭classroom teachers, social workers, and psychologists — and treat patients of all ages.‬

‭Typical roles and responsibilities of an audiologist include:‬

‭‬
● I‭ nterpret hearing test results.‬
‭●‬ ‭Develop treatment plans with other healthcare professionals.‬
‭●‬ ‭Train and counsel patients in the use of various listening devices.‬
‭●‬ ‭Select and fit hearing aids and cochlear implants.‬
‭●‬ ‭Conduct research to enhance knowledge about hearing and balance function.‬
‭●‬ ‭Manage hearing conservation and hearing loss prevention programs.‬

‭ hrough these daily tasks, they work to prevent, diagnose, and manage their patients'‬
T
‭hearing and balance disorders through audiometers, computers, and other testing‬
‭devices, as well as hearing aids and cochlear implants.‬

‭To become an audiologist, you must:‬

‭‬
● ‭ omplete a bachelor’s degree in any field.‬
C
‭●‬ ‭Complete a doctoral degree in audiology.‬
‭●‬ ‭Earn a Certificate of Clinical Competence in Audiology (CCC-A).‬
‭●‬ ‭Obtain a license to practice as an audiologist in your state.‬

‭(‬‭Credit: https://college.mayo.edu/academics/explore-health-care-careers/careers-a-z/audiologist‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Orthoptist and Optometrists‬

‭ n orthoptist is an advanced healthcare provider within the ophthalmic field who helps‬
A
‭people with specific eye conditions.‬‭Orthoptists specialize in evaluating and treating eye‬
‭movement problems and binocular vision and are uniquely trained to treat a variety of‬
‭eye-related disorders.‬

‭ n orthoptist is specifically trained to evaluate how the eye functions and interpret test‬
A
‭results. They diagnose eye movement disorders and work with the ophthalmologist to‬
‭formulate and manage individual treatment plans.‬

‭In a typical day, orthoptists:‬

‭●‬ ‭ erform patient examinations and tests to evaluate vision and look for eye issues‬
P
‭and disorders.‬
‭●‬ ‭Evaluate, detect, and diagnose amblyopia, diplopia, genetic disorders of the eye,‬
‭complex pediatric and adult strabismus, and other eye movement conditions.‬
‭●‬ ‭Create treatment plans for patients with eye movement problems and binocular‬
‭vision.‬
‭●‬ ‭Work with ophthalmologists with surgical planning of some eye movement‬
‭disorders.‬
‭●‬ ‭Help patients and their family members understand their eye disorders and‬
‭treatment plans.‬

‭ ptometrists diagnose and treat various eye and vision disorders, injuries, diseases, and‬
O
‭o ther issues. After obtaining a bachelor’s degree, they must complete a Doctor of‬
‭Optometry (O.D.) degree (typically four years) and get a license to practice.‬

‭ rthoptists are more specialized in their work, specifically with eye movement problems‬
O
‭and binocular vision. They are uniquely skilled in diagnostic techniques. After obtaining a‬
‭bachelor’s degree, orthoptists typically complete a two-year fellowship certification and‬
‭o btain a license to practice.‬

‭(‬‭Credit:‬‭https://college.mayo.edu/academics/explore-health-care-careers/careers-a-z/orthoptist/‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 19: Immune and Lymphatic Systems‬

‭ igure 19.1‬ ‭In this microscope view, the purple-stained‬‭neutrophil (upper left) and‬
F
‭eosinophil (lower right) are white blood cells that float among red blood cells in this blood‬
‭smear. Neutrophils provide an early, rapid, and nonspecific defense against invading‬
‭pathogens. Eosinophils play a variety of roles in the immune response. Red blood cells are‬
‭about 7–8 µm (micrometers) in diameter, and a neutrophil is approximately 10–12µm.‬
‭(credit: modification of work by Dr. David Csaba;‬‭OpenStax‬‭)‬

‭19.1 Introduction‬
‭ he environment consists of numerous‬‭pathogens‬‭, which‬‭are agents, usually‬
T
‭microorganisms, that cause diseases in their hosts. A‬‭host‬‭is an organism invaded and‬
‭o ften harmed by a pathogen. Pathogens include bacteria, protists, fungi, and other‬
‭infectious organisms. We are constantly exposed to pathogens on surfaces and in food,‬
‭water, and air. Mammalian immune systems evolved for protection from such pathogens;‬
‭they are composed of a highly diverse array of specialized cells and molecules that‬
‭coordinate a rapid and flexible defense system capable of protecting from most of these‬
‭disease agents.‬
‭ accines were developed to reduce the chance of infection of a particular disease, such as‬
V
‭measles, mumps, polio, or chickenpox, by assisting the body to create immunity. However,‬
‭many diseases still do not have a vaccine, such as the deadly disease caused by the Ebola‬
‭virus. Data from the World Health Organization indicates that more than 11,000 people‬
‭died out of over 27,000 cases reported during the 2014–2015 outbreak. Though most of‬

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‭Chapter 19: Immune and Lymphatic Systems‬

t‭ he cases were in Africa, Ebola spread to other countries and prompted researchers to try‬
‭to find a treatment.‬
‭ or a general introduction to the topic, click the link below or scan the QR code to watch‬
F
‭the TED-Ed video by Shannon Stile titled “Cell vs. virus - a battle for health.”‬

Cell vs. virus: A battle for health - Shannon Stiles

‭19.2 The Innate Immune Response‬

‭ he immune system can be divided into two overlapping mechanisms to destroy‬
T
‭pathogens: the‬‭innate immune response‬‭, which is relatively‬‭rapid but nonspecific and‬
‭thus not consistently effective, and the‬‭adaptive‬‭immune response‬‭, which is slower in its‬
‭development during an initial infection with a pathogen, but is highly specific and effective‬
‭at attacking a wide variety of pathogens (‬‭Figure 19.2‬‭).‬ ‭There is some functional overlap‬
‭between the two responses, as the innate responses enhance the adaptive responses, so‬
‭they are more effective. Before reading further in the chapter, it will be helpful to click the‬
‭link below or scan the QR code to watch the TED-Ed video by Emma Bryce titled “How‬
‭does your immune system work?”‬

How does your immune system work? - Emma Bryce

I‭ nnate immunity occurs naturally because of genetic factors or physiology; it is not‬

‭induced by infection or vaccination but works to reduce the workload for the adaptive‬
‭immune response. Innate immunity has a limited number of specific targets: any‬
‭pathogenic threat triggers a consistent sequence of events that can identify the type of‬
‭pathogen and either clear the infection independently or mobilize a highly specialized‬
‭adaptive immune response.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ he innate immune system is the first responder to pathogenic threats that bypass the‬
T
‭body's natural physical and chemical barriers. Using a combination of cellular and‬
‭molecular attacks, the innate immune system identifies the nature of a pathogen and‬
‭responds with inflammation, phagocytosis, cytokine release, destruction by NK cells, and a‬
‭complement system. When innate mechanisms are insufficient to clear an infection, the‬
‭adaptive immune response is informed and mobilized.‬

‭Barrier Defenses (Physical and Chemical)‬

‭ ny discussion of the innate immune response usually begins with the physical barriers‬
A
‭that prevent pathogens from entering the body, destroying them after they enter, or‬
‭flushing them out before they can establish themselves in the hospitable environment of‬
‭the body’s soft tissues.‬‭Barrier defenses‬‭are part‬‭o f the body’s most basic defense‬
‭mechanisms. They are not a response to infections, but they continuously work to protect‬
‭against a broad range of pathogens.‬
‭ he different modes of barrier defenses are associated with the external surfaces of the‬
T
‭body, where pathogens may try to enter. The skin is the primary barrier to‬
‭microorganisms' entrance into the body. Not only is the skin covered with a layer of dead,‬
‭keratinized epithelium that is too dry for bacteria to grow, but as these cells are‬
‭continuously sloughed off from the skin, they carry bacteria and other pathogens with‬
‭them. Additionally, sweat and other skin secretions may lower pH, contain antimicrobial‬
‭components, and physically wash microbes away.‬
‭ nother barrier is the saliva in the mouth, which is rich in‬‭lysozyme‬‭—an enzyme that‬
A
‭destroys bacteria by digesting the protective cell walls surrounding them. The stomach's‬
‭acidic environment, which is fatal to many pathogens, is also a barrier. Additionally, the‬
‭mucus layer of the gastrointestinal tract, respiratory tract, reproductive tract, eyes, ears,‬
‭and nose traps microbes and facilitates removal. In the case of the upper respiratory tract,‬
‭ciliated epithelial cells move potentially contaminated mucus upwards to the mouth, where‬
‭it is then swallowed into the digestive tract, ending up in the stomach's harsh, acidic‬
‭environment. Considering how often you breathe compared to how frequently you eat or‬
‭perform other activities that expose you to pathogens, it is not surprising that multiple‬
‭barrier mechanisms have evolved to work in concert to protect this vital area.‬

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‭ igure 19.2‬ ‭Cooperation between the innate and adaptive‬‭immune responses enhances‬
F
‭the adaptive responses to be more effective. (credit:‬‭OpenStax‬‭). A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

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‭Cells of the Innate Immune Response‬

‭ s stated previously, various cell types are associated with the innate immune response,‬
A
‭such as macrophages, natural killer (NK) cells, monocytes, dendrites, and neutrophils.‬
‭Macrophages‬‭are irregularly shaped phagocytes that‬‭are the most versatile type in the‬
‭body (‬‭Figure 19.3‬‭).‬ ‭Phagocytes‬‭are cells that can‬‭surround and engulf a particle or cell, a‬
‭process called‬‭phagocytosis‬‭. The phagocyte takes the‬‭o rganism inside and fuses with a‬
‭lysosome and its digestive enzymes, effectively killing many pathogens. The immune‬
‭system's phagocytes engulf other particles or cells to clean an area of debris or old cells‬
‭o r destroy pathogenic organisms such as bacteria. They participate in innate immune‬
‭responses and have evolved to cooperate with lymphocytes, a type of white blood cell, as‬
‭part of the adaptive immune response. Macrophages exist in many body tissues, freely‬
‭roaming through connective tissues or within specific tissues such as lymph nodes.‬
‭Macrophages move through tissues and squeeze through capillary walls. When pathogens‬
‭breach the body’s barrier defenses, macrophages are the first line of defense.‬
‭ atural killer‬‭cells are a type of lymphocyte that‬‭can induce‬‭apoptosis‬‭, that is,‬
N
‭programmed cell death, in cells infected with pathogens such as bacteria and viruses. NK‬
‭cells recognize these cells by mechanisms that are still poorly understood, presumably‬
‭involving their surface receptors. If apoptosis is induced before the virus can synthesize‬
‭and assemble all its components, no infectious virus will be released from the cell, thus‬
‭preventing further infection.‬
‭ onocytes‬‭are circulating precursor cells that differentiate‬‭into either macrophage or‬
M
‭dendritic cells. Signal molecules of inflammation can rapidly attract monocytes to areas of‬
‭infection.‬
‭ endritic cells‬‭and‬‭neutrophils‬‭also function as phagocytes‬‭that are attracted via signal‬
D
‭molecules from the bloodstream to infected tissues. These cells contain a variety of‬
‭vasoactive mediators, such as histamine. Whereas macrophages act like sentries, always‬
‭o n guard against infection, dendritic cells and neutrophils can be considered military‬
‭reinforcements that are called into a battle to hasten the destruction of the enemy.‬

‭Inflammatory Response‬
‭ he hallmark of the innate immune response is‬‭inflammation‬‭,‬‭which everyone has‬
T
‭experienced. Stub a toe, cut a finger, or do any activity that causes tissue damage, and‬
‭inflammation will result, with its four characteristics: heat, redness, pain, and swelling‬
‭(“loss of function” is sometimes mentioned as a fifth characteristic).‬

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‭ igure 19.3‬ ‭The characteristics and location of‬‭cells involved in the innate immune‬
F
‭system are described. (credit: modification of work by NIH;‬‭OpenStax‬‭)‬

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I‭ t is important to note that inflammation does not have to be initiated by an infection but‬
‭can also be caused by tissue injuries. The release of damaged cellular contents into the‬
‭injury site is enough to stimulate the response, even without breaks in physical barriers‬
‭that would allow pathogens to enter (e.g., by hitting your thumb with a hammer). The‬
‭inflammatory reaction brings phagocytic cells to the damaged area to clear cellular debris‬
‭and set the stage for wound repair (‬‭Figures 19.4 and‬‭19.5‬‭).‬
I‭ nflammation not only brings fluid and cells into the site to destroy the pathogen and‬
‭remove it and debris from the site but also helps isolate the site, limiting the pathogen's‬
‭spread. Acute inflammation is a short-term inflammatory response to an insult to the‬
‭body. If the cause of the inflammation is not resolved, however, it can lead to chronic‬
‭inflammation, an ongoing condition associated with significant tissue destruction and‬
‭fibrosis. Foreign bodies, persistent pathogens, and autoimmune diseases such as‬
‭rheumatoid arthritis can cause it.‬
‭There are four important parts to the inflammatory response:‬
‭•‬ ‭ issue Injury.‬‭The released contents of injured cells‬‭stimulate the release of various‬
T
‭substances, including histamines and prostaglandins. Histamine increases the‬
‭diameter of local blood vessels (vasodilation), causing an increase in blood flow.‬
‭Histamine also increases the permeability of local capillaries, causing plasma to leak‬
‭o ut and form interstitial fluid. This causes swelling associated with inflammation.‬
‭Additionally, injured cells and phagocytes are sources of inflammatory mediators,‬
‭including prostaglandins, which cause vasodilation by relaxing the vascular smooth‬
‭muscle and majorly cause inflammation-related pain. Nonsteroidal anti-inflammatory‬
‭drugs such as aspirin and ibuprofen relieve pain by inhibiting prostaglandin‬
‭production.‬

‭•‬ ‭ asodilation.‬‭Many inflammatory mediators, such as‬‭histamine, are vasodilators that‬

V
‭increase the diameters of local capillaries. This causes increased blood flow and is‬
‭responsible for the heat and redness of inflamed tissue. It allows blood greater access‬
‭to the site of inflammation.‬

‭•‬ I‭ ncreased Vascular Permeability.‬‭At the same time,‬‭inflammatory mediators increase‬

‭the permeability of the local capillaries, causing fluid leakage into the interstitial‬
‭space, resulting in swelling, or edema, associated with inflammation.‬

‭•‬ ‭ ecruitment of Phagocytes.‬‭Macrophages are recruited‬‭via cytokines to clean up the‬

R
‭debris left at the site. When local infections are severe, neutrophils are attracted to‬
‭the areas of infections in large numbers, and as they phagocytose the pathogens and‬
‭subsequently die, their accumulated cellular remains are visible as pus at the infection‬
‭site.‬

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‭Figure 19.4‬ ‭The inflammatory response. (credit:‬‭OpenStax‬‭)‬

‭ igure 19.5‬ ‭The inflammatory response. (credit:‬‭Inflammatory response.png‬‭;‬‭Alan Sved‬‭;‬

F
‭CC BY-SA 4.0‬‭)‬

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‭ verall, inflammation is valuable for many reasons. Not only are the pathogens killed and‬
O
‭debris removed, but the increase in vascular permeability encourages the entry of clotting‬
‭factors, the first step toward wound repair. Inflammation also facilitates the transport of‬
‭antigens to lymph nodes by dendritic cells to develop the adaptive immune response.‬

‭Soluble Mediators of the Innate Immune Response‬

‭ he previous discussions have alluded to chemical signals that can induce cells to change‬
T
‭various physiological characteristics. These soluble factors, such as cytokines, C-reactive‬
‭proteins, and complement proteins, are secreted during innate immune responses and‬
‭may also play a subsequent role in adaptive immune responses.‬

‭Cytokines‬
‭ ‬‭c ytokine‬‭is a chemical messenger that allows cells‬‭to communicate with each other over‬
A
‭short distances. Cytokines are secreted into the intercellular space, and the cytokine's‬
‭action induces the receiving cell to change its physiology. These messengers regulate cell‬
‭differentiation, proliferation, and gene expression to affect immune responses. At least 40‬
‭types of cytokines exist in humans that differ in terms of the cell type that produces them,‬
‭the cell type that responds to them, and the changes they make. Cytokines also send‬
‭feedback to nervous system cells to bring about the symptoms of feeling sick, including‬
‭lethargy, muscle pain, and nausea. These effects may have evolved because the symptoms‬
‭encourage the individual to rest and prevent them from spreading the infection to others.‬
‭Cytokines also increase the core body temperature, causing a fever, which causes the liver‬
‭to withhold iron from the blood. Without iron, certain pathogens, such as some bacteria,‬
‭are unable to replicate; this is called nutritional immunity.‬
I‭ nterferons‬‭are a type of cytokine that is secreted‬‭by cells infected with viruses and then‬
‭travel to adjacent cells (‬‭Figure 19.6‬‭). In response‬‭to these interferons, uninfected cells‬
‭alter gene expression to increase their resistance to infection. Thus, even though the initial‬
‭cell is sacrificed, the surrounding cells are protected.‬
‭ -reactive protein‬‭is another example of a pro-inflammatory‬‭cytokine specific for‬
C
‭polysaccharide bacterial cell walls. Phagocytes such as macrophages have receptors for‬
‭these proteins, and they are thus able to recognize them as they are bound to the bacteria.‬
‭This brings the phagocyte and bacterium into close proximity and enhances the‬
‭phagocytosis of the bacterium.‬

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‭ igure 19.6‬ ‭Interferons are cytokines released by‬‭a cell infected with a virus. Response‬
F
‭o f neighboring cells to interferon helps stem the infection. (credit:‬‭OpenStax‬‭); A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Complement System‬
‭ he‬‭complement system‬‭is a collection of approximately‬‭20 types of soluble proteins that‬
T
‭function to destroy extracellular pathogens. Cells of the liver and macrophages synthesize‬
‭complement proteins continuously, so these proteins are abundant in the blood and can‬
‭respond immediately to infecting microorganisms. The complement system is so named‬
‭because it is complementary to the antibody response of the adaptive immune system.‬
‭Complement proteins bind to the surfaces of microorganisms and are particularly‬
‭attracted to pathogens already bound by antibodies. The binding of complement proteins‬
‭o ccurs in a specific and highly regulated sequence, with each successive protein activated‬
‭by cleavage or structural changes induced upon binding the preceding protein(s). After‬
‭the first few complement proteins bind, a cascade of sequential binding events follows in‬
‭which the pathogen rapidly becomes coated in complement proteins.‬

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‭Complement proteins perform several functions:‬

•‭ ‬ ‭ ind to the pathogen's cell membrane that activates it, labeling it for phagocytosis.‬
B
‭•‬ ‭Diffuse away from the pathogen and act as chemotactic agents to attract phagocytic‬
‭cells to the site of inflammation.‬
‭•‬ ‭Form damaging pores in the pathogen's plasma membrane.‬

‭ igure 19.7‬‭summarizes key points from the prior discussion‬‭o f the innate immune‬
F
‭system.‬

‭19.3 The Adaptive Immune Response‬

‭ daptive immune responses‬‭take days or weeks to establish,‬‭much longer than the‬
A
‭innate response. This type of immunity occurs after exposure to an antigen, either from a‬
‭pathogen or through vaccination, and is activated when the innate immune response is‬
‭insufficient to control the infection. This immune response also involves molecular‬
‭memory, which means re-exposure to the same pathogen will elicit an efficient and quick‬
‭response. This gives long-term protection from reinfection.‬
‭ here are two types of adaptive responses: the‬‭cell-mediated‬‭immune response‬‭, carried‬
T
‭o ut by T-cell lymphocytes, and the‬‭humoral (or antibody-mediated)‬‭immune response‬‭,‬
‭governed by activated B-cell lymphocytes and their production of antibodies.‬

‭Antigen-presenting Cells‬
‭ n‬‭antigen‬‭is a foreign or “non-self” macromolecule‬‭that reacts with immune system cells.‬
A
‭Not all antigens will provoke a response. For instance, individuals produce innumerable‬
‭“self” antigens and are constantly exposed to harmless foreign antigens, such as food‬
‭proteins, pollen, or dust components. The suppression of immune responses to harmless‬
‭macromolecules is highly regulated and typically prevents processes that could damage‬
‭the host, known as tolerance.‬
‭ he innate immune system contains cells that detect potentially harmful antigens and then‬
T
‭inform the adaptive immune response about their presence. An‬‭antigen-presenting cell‬
‭(APC)‬‭is an immune cell that detects, engulfs, and‬‭informs the adaptive immune response‬
‭about an infection (‬‭Figure 19.8‬‭).‬

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‭ igure 19.7‬ ‭Components of the innate immune response‬‭and their associated function(s).‬
F
‭(credit:‬‭Innate defense system‬‭; Cenevo; pressbooksccconline.org).‬

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‭ hen a pathogen is detected, these APCs consume it via phagocytosis and digest it to form‬
W
‭many different antigen fragments. Antigen fragments are then transported to the surface‬
‭o f the APC, form complexes with MHC II molecules, and serve as an indicator to other‬
‭immune cells of a “non-self” invader. Dendritic cells and macrophages are examples of‬
‭APCs.‬

‭ igure 19.8‬ ‭An APC, such as a macrophage, engulfs‬‭and digests a foreign bacterium. An‬
F
‭antigen from the bacterium is presented on the cell surface in conjunction with an MHC II‬
‭molecule. (credit:‬‭OpenStax‬‭); A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭T and B Lymphocytes‬
‭ ymphocytes‬‭are a class of white blood cells in human‬‭circulating blood consisting of‬
L
‭approximately 80 to 90 percent T cells (‬‭Figure 19.9‬‭).‬ ‭T cells‬‭are a vital component in the‬
‭cell-mediated response, which is the specific immune response that destroys cells infected‬
‭with viruses and certain bacteria. There are three types of T cells: cytotoxic, helper, and‬
‭suppressor T cells.‬‭Cytotoxic T cells‬‭destroy virus-infected‬‭cells in the cell-mediated‬

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i‭mmune response.‬‭Helper T cells‬‭activate both the cell-mediated and antibody-mediated‬

‭immune responses.‬ ‭Suppressor T cells‬‭deactivate T‬‭and B cells when needed and thus‬
‭prevent the immune response from becoming too intense.‬
‭ he remaining 10 to 20 percent of the lymphocytes are‬‭B cells‬‭, which function as part of‬
T
‭the humoral or antibody-mediated immune response.‬

‭ igure 19.9‬ ‭This image from a scanning electron‬‭microscope shows a T lymphocyte,‬

F
‭which is responsible for the cell-mediated immune response. The value shown for scale is‬
‭1 micrometer, which is 1/1,000,000 of a meter. (credit: modification of work by NCI;‬
‭scale-bar data from Matt Russell;‬‭OpenStax‬‭)‬

‭ and B cells are similar in that each cell only expresses one type of antigen receptor.‬
T
‭Individuals may possess a population of T and B cells expressing a near-limitless variety of‬
‭antigen receptors capable of recognizing virtually any infecting pathogen.‬
‭ and B cells are activated when they recognize small components of antigens, called‬
T
‭epitopes‬‭, presented by APCs (‬‭Figure 19.10‬‭). Recognition‬‭o ccurs at a specific epitope‬
‭rather than on the entire antigen, so epitopes are known as antigenic determinants.‬
‭Without information from APCs, T, and B cells remain inactive or naïve and cannot prepare‬
‭an immune response.‬

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‭ igure 19.10‬ ‭An antigen is a macromolecule that‬‭reacts with immune system‬

F
‭components. A given antigen may contain several regions, called epitopes, that immune‬
‭cells recognize. In this figure, the entire structure is an antigen, and the orange, salmon,‬
‭and green components projecting from it represent potential epitopes. (credit:‬‭OpenStax‬‭)‬

‭ aïve T cells can express one of two molecules, CD4 or CD8, on their surface, as shown in‬
N
‭Figure‬‭19.11‬‭, and are accordingly classified as CD4‬‭+‬ ‭o r CD8‬‭+‬ ‭cells. These molecules are‬
‭important because they regulate how a T cell interacts with and responds to an APC. Naïve‬
‭CD4‬‭+‬ ‭cells bind APCs via their antigen-embedded MHC‬‭II molecules. They are stimulated to‬
‭become helper T cells that go on to activate B cells or cytotoxic T cells; the activated helper‬
‭T cells also secrete cytokines to inform more cells about the pathogenic threat.‬
I‭ n contrast, CD8‬‭+‬ ‭cells engage antigen-embedded MHC‬‭I molecules on APCs. They are‬
‭stimulated to become cytotoxic T cells, which directly kill infected cells by apoptosis and‬
‭emit cytokines to amplify the immune response.‬
‭ aive B cells require two interactions to become activated. The first is with the intact‬
N
‭antigen, and the second is with the activated helper T cell. The B cells must be able to bind‬
‭intact antigens because they secrete antibodies that must recognize the pathogen directly‬
‭rather than digested remnants of the pathogen.‬
‭ hen the helper T cell with the correct MHC II-epitope combination detects that a B cell is‬
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‭bound to its relevant antigen, the T cell secretes specific cytokines that induce the B cell to‬
‭differentiate into plasma cells, which secrete antibodies. The cytokines also stimulate‬
‭accelerated cell division of the activated B cell, in a process known as‬‭c lonal selection‬‭, to‬
‭make thousands of identical (clonal) copies of it. This is beneficial because the resulting‬
‭population of plasma cells produces the specific antibody for the particular antigen.‬

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‭ igure 19.11‬ ‭Naïve CD4‬‭+‬ ‭T cells engage MHC II molecules‬‭o n antigen-presenting cells‬
F
‭(APCs) and become activated. Clones of the activated helper T cell, in turn, activate B cells‬
‭and CD8‬‭+‬ ‭T cells, which become cytotoxic T cells.‬‭Cytotoxic T cells kill infected cells. (credit:‬
‭OpenStax‬‭); A link to a video explanation of this figure‬‭is available at‬‭Biology411.com‬‭.‬

‭Immunological Memory‬
‭ he adaptive immune system possesses a memory component that allows for an efficient‬
T
‭and dramatic response upon a second infection by the same pathogen. Memory is handled‬
‭by the adaptive immune system with little reliance on cues from the innate response.‬
‭During the adaptive immune response to a pathogen that has not been encountered‬
‭before, called a‬‭primary response‬‭, plasma cells secreting‬‭antibodies and differentiated T‬

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c‭ ells increase, then plateau over time. As naive B and T cells transition into activated forms‬
‭(i.e.,‬‭effector cells‬‭), a subset of the naïve populations‬‭differentiates into B and T memory‬
‭cells with the same antigen specificities.‬
‭ herefore, a‬‭memory cell‬‭is an antigen-specific B‬‭o r T cell that does not differentiate into‬
T
‭effector cells during the primary immune response but can immediately become effector‬
‭cells upon re-exposure to the same pathogen. During the primary immune response,‬
‭memory cells do not respond to antigens and do not contribute to host defenses. As the‬
‭infection is cleared and pathogenic stimuli subside, the effectors are no longer needed and‬
‭undergo apoptosis. In contrast, memory cells persist in the circulation.‬
S‭ uppose the pathogen is never reencountered during the individual’s lifetime. In that case,‬
‭B and T memory cells will circulate for a few years or even several decades and will‬
‭gradually die off, having never functioned as effector cells. However, if the host is‬
‭re-exposed to the same pathogen type, circulating memory cells will immediately‬
‭differentiate into plasma cells without input from APCs or T‬‭H‬ ‭cells. One reason the adaptive‬
‭immune response is delayed is that it takes time for naïve B and T cells to identify and‬
‭activate the appropriate antigen specificities. Upon reinfection, this step is skipped, and the‬
‭result, designated as the‬‭secondary response‬‭, is a‬‭more rapid production of immune‬
‭defenses (‬‭Figure 19.12‬‭).‬
‭ emory B cells that differentiate into plasma cells output tens to hundreds-fold greater‬
M
‭antibody amounts than were secreted during the primary response, as the graph in‬
‭Figure 19.13‬‭illustrates. This rapid and dramatic‬‭antibody response may stop the infection‬
‭before it can even become established, and the individual may not realize they have been‬
‭exposed.‬
‭ accinations are based on the knowledge that exposure to noninfectious antigens from‬
V
‭known pathogens generates a mild primary immune response. The host may not perceive‬
‭the immune response to vaccination as an illness but still confers immune memory. The‬
‭reaction is similar to secondary exposure when exposed to the corresponding pathogen‬
‭to which an individual was vaccinated. Because each reinfection generates more memory‬
‭cells and increased resistance to the pathogen, and some memory cells die, specific‬
‭vaccine courses involve one or more booster vaccinations to mimic repeat exposures. For‬
‭instance, tetanus boosters are necessary every ten years because the memory cells only‬
‭live that long.‬

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‭ igure 19.12‬ ‭In the primary immune response of B‬‭cells, an activated B cell will undergo‬
F
‭clonal selection to produce memory and effector (i.e., plasma) cells. In a subsequent‬
‭second response, the memory B cells will be activated to form clones of memory and‬
‭effector cells. (credit:‬‭OpenStax‬‭); A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 19.13‬ ‭In the primary response to infection,‬‭antibodies are secreted first from‬
F
‭plasma cells. Upon re-exposure to the same pathogen, memory cells differentiate into‬
‭antibody-secreting plasma cells that output more antibodies for longer. (credit:‬‭OpenStax‬‭)‬

‭Antibodies‬
‭ ntibodies‬‭, also known as‬‭immunoglobulins‬‭, are the‬‭molecules secreted from effector B‬
A
‭cells, called‬‭plasma cells‬‭, that mediate the humoral‬‭immune response. Antibodies‬
‭circulate freely and act independently of plasma cells. Antibodies affect pathogens via one‬
‭o f three general mechanisms: neutralization, opsonization, or complement activation‬
‭(‬‭Figure 19.14‬‭).‬

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‭ igure 19.14‬ ‭Antibodies may inhibit infection by‬‭(a) preventing the antigen from binding‬
F
‭its target, (b) tagging a pathogen for destruction by macrophages or neutrophils, or (c)‬
‭activating the complement cascade. (credit:‬‭OpenStax‬‭);‬‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭Mechanisms of Acquiring Immunity‬

‭ hile the general concept of immunity has been discussed extensively, it is helpful to‬
W
‭categorize the various mechanisms by which it is obtained formally. The two types of‬
‭immunity are classified as passive and active, which reflects the role of an individual’s‬
‭immune system in generating the response.‬ ‭Passive‬‭immunity‬‭results when antibodies‬

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a‭ re passed from one individual to another, while‬‭active immunity‬‭results from an‬

‭individual’s immune system response. Each category has two options for how the‬
‭immunity was acquired -‬‭naturally‬‭o r‬‭artificially‬‭.‬ ‭Therefore, there are four immunity‬
‭acquisition mechanisms: passive natural, passive artificial, active natural, and active‬
‭artificial (‬‭Figure 19.15‬‭). For example, a person‬‭who has recently produced a successful‬
‭immune response against a particular disease agent can donate blood to a nonimmune‬
‭recipient and confer temporary immunity through antibodies in the donor’s blood serum.‬
‭This phenomenon is called passive, artificial-acquired immunity; it also occurs naturally‬
‭during breastfeeding, which makes breastfed infants highly resistant to infections during‬
‭the first few months of life. The immunity that results from vaccinations is termed active,‬
‭artificially acquired.‬

‭ igure 19.15‬ ‭The four mechanisms for acquiring immunity:‬ ‭Passive, natural; passive‬
F
‭artificial; active, natural; active artificial. (credit:‬‭OpenStax‬‭); A link to a video explanation‬
‭o f this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ or additional information about‬‭vaccines‬‭, an example of active, artificial-acquired‬

F
‭immunity, click the link below or scan the QR code to watch the TED-Ed video by‬‭Kelwalin‬
‭Dhanasarnsombut titled “How do vaccines work?”‬

How do vaccines work? - Kelwalin Dhanasarnsombut

S‭ mallpox is a highly contagious and deadly disease responsible for more than 300 million‬
‭deaths during the 20th century alone. In 1796, physician Edward Jenner showed that‬
‭individuals who were purposefully exposed to cowpox, a less pathogenic member in the‬
‭same viral family, were immune to smallpox. This was the first demonstration of a vaccine‬
‭developed specifically for a contagious disease. For more information about this disease‬
‭and the associated vaccine, click the link below or scan the QR code to view the TED-Ed‬
‭video by Simona Zompi titled “How we conquered the deadly smallpox virus.”‬

‭How we conquered the deadly smallpox virus - Simona‬‭Zompi‬

‭ uring the‬‭COVID‬‭pandemic, there were multiple initiatives‬‭to quickly develop effective‬

D
‭vaccines against the‬‭SARs-CoV-2 coronavirus‬‭causing‬‭the disease. For more information,‬
‭click the links below to watch the associated TED-Ed videos:‬

What is a coronavirus? - Elizabeth Cox

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How the COVID-19 vaccines were created so quickly - …

‭19.4 Lymphatic System‬

‭ he lymphatic system is a network of vessels, cells, and organs that carries excess fluids to‬
T
‭the bloodstream and filters pathogens from the blood. For most people, this system is‬
‭associated with the immune system to such a degree that the two systems are virtually‬
‭indistinguishable. For example, the swelling of lymph nodes during an infection and the‬
‭transport of lymphocytes via the lymphatic vessels are but two examples of the many‬
‭connections between these critical organ systems.‬

‭Functions of the Lymphatic System‬

‭ significant function of the lymphatic system is to drain body fluids and return them to‬
A
‭the bloodstream. Blood pressure causes fluid leakage from the capillaries, resulting in‬
‭fluid accumulation in the interstitial space—that is, spaces between individual cells in the‬
‭tissues. In humans, approximately 5 gallons of plasma is released into the interstitial space‬
‭o f the tissues each day due to capillary filtration. Once this filtrate is out of the‬
‭bloodstream and in the tissue spaces, it is called interstitial fluid. Of this, the blood vessels‬
‭reabsorbed 4.5 gallons (or 90%). But what happens to the remaining fluid? This is where‬
‭the lymphatic system comes into play. It drains this excess fluid and empties it into the‬
‭bloodstream via a series of vessels, trunks, and ducts.‬
‭ ymph‬‭is the term used to describe interstitial fluid‬‭o nce it has entered the lymphatic‬
L
‭system. When the lymphatic system is damaged, such as by being blocked by cancer cells‬
‭o r destroyed by injury, protein-rich interstitial fluid accumulates (sometimes “backs up”‬
‭from the lymph vessels) in the tissue spaces. This inappropriate accumulation of fluid,‬
‭referred to as lymphedema, may lead to serious medical consequences.‬
‭ s the vertebrate immune system evolved, the network of lymphatic vessels became‬
A
‭convenient avenues for transporting the cells of the immune system. Additionally, the‬
‭transport of dietary lipids and fat-soluble vitamins absorbed in the gut uses this system.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ he immune system's cells use lymphatic vessels to move from interstitial spaces back into‬
T
‭the circulation. Lymph nodes, which are small, bean-shaped organs located throughout‬
‭the lymphatic system, also serve as major staging areas for the development of critical‬
‭immune responses.‬

‭Structure of the Lymphatic System‬

‭ he lymphatic vessels begin as blind-ending or closed-at-one-end capillaries, which feed‬
T
‭into larger and larger lymphatic vessels and eventually empty into the bloodstream by a‬
‭series of ducts. Along the way, the lymph travels through the‬‭lymph nodes‬‭commonly‬
‭found near the groin, armpits, neck, chest, and abdomen. Humans have about 500–600‬
‭lymph nodes throughout the body (‬‭Figure 19.16‬‭).‬

‭ significant distinction between the lymphatic and cardiovascular systems in humans is‬
A
‭that lymph is not actively pumped by the heart but is forced through the vessels by the‬
‭body's movements, the contraction of skeletal muscles during body movements, and‬
‭breathing. One-way valves (semilunar-type valves) in lymphatic vessels keep the lymph‬
‭moving toward the heart. Lymph flows from the lymphatic capillaries through lymphatic‬
‭vessels and is then returned into the circulatory system via the lymphatic ducts located at‬
‭the junction of the jugular and subclavian veins in the neck.‬

‭Lymphatic Capillaries‬
‭ ymphatic capillaries, also called terminal lymphatics, are vessels where interstitial fluid‬
L
‭enters the lymphatic system to become lymph fluid. Located in almost every tissue in the‬
‭body, these vessels are interlaced among the arterioles and venules of the circulatory‬
‭system in the soft connective tissues (‬‭Figure 19.17‬‭).‬‭Exceptions are the central nervous‬
‭system, bone marrow, bones, teeth, and the cornea of the eye, which do not contain lymph‬
‭vessels.‬

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P
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‭ igure 19.16‬ ‭The anatomy of the lymphatic system.‬ ‭Lymphatic vessels in the arms and‬
F
‭legs transport lymph to the larger lymphatic vessels in the torso. Ultimately, this fluid is‬
‭returned to the circulatory system just before venous blood enters the superior vena cava.‬
‭(credit:‬‭OpenStax‬‭)‬

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‭ igure 19.17‬ ‭Lymphatic Capillaries Lymphatic capillaries‬‭are interlaced with the‬

F
‭arterioles and venules of the cardiovascular system. Collagen fibers anchor a lymphatic‬
‭capillary in the tissue (inset). Interstitial fluid slips through spaces between the‬
‭overlapping endothelial cells that compose the lymphatic capillary. (credit:‬‭OpenStax‬‭)‬

I‭ n the small intestine, lymphatic capillaries called lacteals are critical for transporting‬
‭dietary lipids and lipid-soluble vitamins to the bloodstream. In the small intestine, dietary‬
‭triglycerides combine with other lipids and proteins and enter the lacteals to form a milky‬
‭fluid called chyle. The chyle then travels through the lymphatic system, eventually entering‬
‭the bloodstream.‬

‭Larger Lymphatic Vessels, Trunks, and Ducts‬

‭ he lymphatic capillaries empty into larger lymphatic vessels, which are similar to veins in‬
T
‭terms of their structure and the presence of valves. These one-way valves are located‬

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r‭ elatively close to one another, and each one causes a bulge in the lymphatic vessel, giving‬
‭the vessels a beaded appearance (‬‭Figure 19.17‬‭).‬
‭ n the right side of the body, the right sides of the head, thorax, and right upper limb‬
O
‭drain lymph fluid into the right subclavian vein via the right lymphatic duct (‬‭Figure 19.18‬‭).‬
‭On the left side of the body, the remaining portions drain into the larger thoracic duct,‬
‭which empties into the left subclavian vein. The thoracic duct begins just beneath the‬
‭diaphragm in the cisterna chyli, a sac-like chamber receiving lymph from the lower‬
‭abdomen, pelvis, and lower limbs through the left and right lumbar trunks and the‬
‭intestinal trunk.‬

‭19.5 Primary Centers of the Immune System‬

‭ lthough the immune system is characterized by circulating cells throughout the body, the‬
A
‭regulation, maturation, and intercommunication of immune factors occur at specific sites.‬
‭The blood circulates immune cells, proteins, and other factors throughout the body.‬
‭Approximately 0.1 percent of all cells in the blood are leukocytes, which encompass‬
‭monocytes (the precursor of macrophages) and lymphocytes. Most cells in the blood are‬
‭erythrocytes (red blood cells). As discussed previously, lymph is a watery fluid that bathes‬
‭tissues and organs with protective white blood cells and does not contain erythrocytes.‬
‭Cells of the immune system can travel between the distinct lymphatic and blood‬
‭circulatory systems by passing through to surrounding tissue.‬
‭ he immune system cells originate from hematopoietic stem cells in the bone marrow.‬
T
‭Cytokines stimulate these stem cells to differentiate into various immune cell types. B cell‬
‭maturation occurs in the‬‭bone marrow‬‭, whereas naïve‬‭T cells move from the bone‬
‭marrow to the‬‭thymus‬‭for maturation (‬‭Figure 19.19‬‭).‬

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P
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‭ igure 19.18‬ ‭The major trunks and ducts of the lymphatic system are shown. The‬
F
‭thoracic duct drains a much more significant portion of the body than the right lymphatic‬
‭duct. (Credit:‬‭OpenStax‬‭)‬

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‭ igure 19.19‬ ‭The location of the thymus gland relative to the lungs and thyroid gland.‬
F
‭(credit:‬‭Position of the thymus gland‬‭; Cancer Research‬‭UK;‬‭CC BY-SA 4.0‬‭)‬

‭ n maturation, T and B lymphocytes circulate to various destinations. Lymph nodes are‬

O
‭scattered throughout the body, as illustrated in‬‭Figure 19.20‬‭, and house large populations‬
‭o f T and B cells, dendritic cells, and macrophages. The lymph gathers antigens as it drains‬
‭from tissues, and these antigens are filtered through lymph nodes before the lymph is‬
‭returned to blood circulation. APCs in the lymph nodes capture and process antigens and‬
‭inform nearby lymphocytes about potential pathogens.‬
‭ he spleen, an organ associated with the circulatory system, is similar to a lymph node but‬
T
‭is much larger and filters blood instead of lymph (‬‭Figure 19.21‬‭). Blood enters the spleen‬
‭through arteries and exits through veins. The spleen contains two types of tissue: red pulp‬
‭and white pulp. Red pulp consists of cavities that store blood. Within the red pulp,‬
‭damaged red blood cells are removed and replaced by new ones. White pulp is rich in B‬
‭and T cells, macrophage, dendritic cells, and NK cells, so antigen-coated bacterial cells in‬
‭the blood are removed at this point.‬

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‭ igure 19.20‬ ‭(a) Lymphatic vessels carry a clear fluid called lymph throughout the body.‬
F
‭The liquid enters (b) lymph nodes through afferent vessels. Lymph nodes are filled with‬
‭lymphocytes that purge infecting cells. The lymph then exits through efferent vessels.‬
‭(credit: modification of work by NIH, NCI;‬‭OpenStax‬‭)‬

‭ igure 19.21‬ ‭The location and structure of the spleen. (credit: modification of work by‬
F
‭NCI;‬‭OpenStax‬‭)‬

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‭19.6 Disruptions of the Immune System‬

‭ athogens can sometimes defeat immune systems. For example, some bacteria, including‬
P
‭Streccus pneumoniae‬‭, surround themselves with a capsule‬‭that inhibits phagocytes from‬
‭engulfing them and displaying antigens to the adaptive immune system.‬‭Human‬
‭immunodeficiency virus‬‭(‬‭HIV‬‭), the virus that causes‬‭AIDS, infects helper T-cells via their‬
‭CD4 surface molecules, gradually depleting the number of these cells in the body; this‬
‭inhibits the adaptive immune system’s capacity to sufficiently respond to infection or‬
‭tumors that persons with healthy immune systems can defend against. Allergies to pollen‬
‭o r pet dander occur when the immune system attacks the body’s cells or tissues.‬
I‭ mmune disruptions may involve insufficient immune responses or inappropriate immune‬
‭targets. Immunodeficiency increases an individual's susceptibility to infections and‬
‭cancers. Hypersensitivities are misdirected responses to harmless foreign particles, such‬
‭as allergies, or host factors, such as autoimmunity.‬

‭Immunodeficiency‬
‭ ailures, insufficiencies, or delays at any level of the immune response can allow‬
F
‭pathogens or tumor cells to gain a foothold and replicate or increase to high enough‬
‭levels that the immune system becomes overwhelmed.‬‭Immunodeficiency‬‭is the failure,‬
‭insufficiency, or delay in the immune system's response, which may be acquired or‬
‭inherited. Immunodeficiency can be acquired due to infection with certain pathogens‬
‭(such as HIV), chemical exposure (including certain medical treatments), malnutrition, or‬
‭possibly extreme stress. For instance, radiation exposure can destroy populations of‬
‭lymphocytes and elevate an individual’s susceptibility to infections and cancer. Dozens of‬
‭genetic disorders result in immunodeficiencies, including Severe Combined‬
‭Immunodeficiency (SCID) and MHC II deficiencies. Rarely, primary immunodeficiencies‬
‭that are present from birth may occur. Neutropenia is one form in which the immune‬
‭system produces a below-average number of neutrophils, the body’s most abundant‬
‭phagocytes. As a result, bacterial infections may go unrestricted in the blood, causing‬
‭severe complications.‬

‭Hypersensitivities‬
‭ aladaptive immune responses toward harmless foreign substances or self-antigens that‬
M
‭o ccur after tissue sensitization are termed‬‭hypersensitivities‬‭.‬‭The types of‬
‭hypersensitivities include immediate, delayed, and autoimmunity. A large proportion of the‬
‭population is affected by one or more types of hypersensitivity.‬

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‭Allergies‬
‭ n allergy is an immune reaction resulting from immediate hypersensitivities in which an‬
A
‭antibody-mediated immune response occurs within minutes of exposure to a harmless‬
‭antigen. In the United States, 20 percent of the population exhibits allergy or asthma‬
‭symptoms, whereas 55 percent test positive for one or more allergens. Upon initial‬
‭exposure to a potential allergen, an allergic individual synthesizes antibodies of the IgE‬
‭class via the typical process of APCs presenting processed antigens to helper T cells that‬
‭stimulate B cells to produce IgE. Some IgE molecules interact with mast cells embedded in‬
‭connective tissues, which primes or sensitizes the tissue. Upon subsequent exposure to‬
‭the same allergen, IgE molecules on mast cells bind the antigen via another region and‬
‭stimulate the mast cell to release substances such as histamine. This substance recruits a‬
‭type of white blood cell that mediates allergic responses.‬
‭ igure 19.22‬‭shows an example of an allergic response to ragweed pollen. The effects of‬
F
‭an allergic reaction range from mild symptoms like sneezing and itchy, watery eyes to‬
‭more severe or even life-threatening reactions involving intensely itchy welts or hives,‬
‭airway contraction with severe respiratory distress, and plummeting blood pressure. This‬
‭extreme reaction is known as‬‭anaphylactic shock‬‭. This‬‭condition can be fatal if not‬
‭treated with epinephrine to counter the blood pressure and breathing effects.‬

‭ igure 19.22‬ ‭On first exposure to an allergen, an IgE antibody is synthesized by plasma‬
F
‭cells in response to a harmless antigen. The IgE molecules bind to mast cells, and on‬
‭secondary exposure, the mast cells release histamines and other modulators that affect‬
‭allergy symptoms. (credit: modification of work by NIH;‬‭OpenStax‬‭)‬

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‭ elayed hypersensitivity is a cell-mediated immune response that takes approximately one‬

D
‭to two days after secondary exposure for a maximal reaction to be observed. This type of‬
‭hypersensitivity may manifest as local tissue lesions or contact dermatitis (rash or skin‬
‭irritation). Delayed hypersensitivity occurs in some individuals in response to contact with‬
‭certain jewelry or cosmetics. Delayed hypersensitivity facilitates the immune response to‬
‭poison ivy. This is also why the skin test for tuberculosis results in a small region of‬
‭inflammation in individuals who have previously been exposed to‬‭Mycobacterium‬
‭tuberculosis‬‭. That is also why cortisone treats such‬‭responses, as it inhibits cytokine‬
‭production.‬

‭Autoimmunity‬
‭ utoimmunity‬‭is hypersensitivity to self-antigens‬‭that affects approximately five percent‬
A
‭o f the population. Most types of autoimmunity involve the humoral immune response.‬
‭Antibodies that inappropriately mark self-components as foreign are termed‬
‭autoantibodies‬‭. In patients with the autoimmune disease‬‭myasthenia gravis‬‭, antibodies‬
‭target muscle cell receptors that induce contraction in response to acetylcholine. The‬
‭result is muscle weakness that may include considerable difficulty with fine and gross‬
‭motor functions.‬
‭ utoimmunity can develop with time, and its causes may be rooted in molecular mimicry,‬
A
‭which is the structural similarity between epitopes of self-molecules in the host and those‬
‭o f invading pathogens. Antibodies may bind self-antigens structurally similar to pathogen‬
‭antigens, which the immune receptors first raise. For example, infection with‬‭Streptococcus‬
‭pyogenes‬‭(the bacterium that causes strep throat)‬‭may generate antibodies or T cells that‬
‭react with heart muscle, which has a similar structure to the surface of‬‭S. pyogenes‬‭. These‬
‭antibodies can damage heart muscle with autoimmune attacks, leading to‬‭rheumatic‬
‭fever‬‭. This situation is also the cause of‬‭Insulin-dependent‬‭(Type 1) diabetes mellitus‬‭,‬
‭in which a destructive inflammatory response arises against insulin-producing pancreas‬
‭cells. Patients with this autoimmunity must be injected with insulin that originates from‬
‭o ther sources.‬

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‭Everyday Connection‬

‭Immune Rejection of Medical Devices‬

I‭ t makes sense that our body's immune system will attack bacteria and viruses, as those‬
‭could harm the body. However, what about implanted devices (e.g., pacemakers, insulin‬
‭pumps, or replacement joints)? These devices improve our overall health, but the immune‬
‭system still regards them as foreign, affecting the device's longevity and functioning. For‬
‭more information, click the link below or scan the QR code to watch the TED-Ed video by‬
‭Kaitlyn Sadtler titled “Your body vs. implants.”‬

Your body vs. implants - Kaitlyn Sadtler

‭19.7 Career Connection -Vaccinologist‬

‭ accination (or immunization) involves the delivery, usually by injection of noninfectious‬

V
‭antigen(s) derived from known pathogens (‬‭Figure 19.23‬‭). Other components, called‬
‭adjuvants, are delivered in parallel to help stimulate the immune response. Immunological‬
‭memory is the reason vaccines work. Ideally, vaccination elicits immunological memory‬
‭and, thus, resistance to specific pathogens without the individual having to experience an‬
‭infection.‬

‭ accinologists are involved in the process of vaccine development, from the initial idea to‬
V
‭the availability of the completed vaccine. This process can take decades, can cost millions‬
‭o f dollars, and can involve many obstacles along the way. For instance, injected vaccines‬
‭stimulate the systemic immune system, eliciting humoral and cell-mediated immunity, but‬
‭have little effect on the mucosal response. This presents a challenge because many‬
‭pathogens are deposited and replicate in mucosal compartments. The injection does not‬
‭provide the most efficient immune memory for these disease agents. For this reason,‬
‭vaccinologists are actively involved in developing new vaccines that are applied via‬
‭intranasal, aerosol, oral, or transcutaneous (absorbed through the skin) delivery methods.‬
‭Importantly, mucosal-administered vaccines elicit both mucosal and systemic immunity‬
‭and produce the same level of disease resistance as injected vaccines.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭538‬
‭Chapter 19: Immune and Lymphatic Systems‬

‭ igure 19.23‬ ‭Vaccines are often delivered by injection into the arm. (credit: U.S. Navy‬
F
‭Photographer's Mate Airman Apprentice Christopher D. Blachly;‬‭WikiMedia Commons‬‭)‬

‭ version of the intranasal influenza vaccine is available, and polio and typhoid vaccines‬
A
‭can be administered orally (‬‭Figure 19.24‬‭). Similarly, the measles and rubella vaccines are‬
‭being adapted to aerosol delivery using inhalation devices. Eventually, transgenic plants‬
‭may be engineered to produce vaccine antigens that can be eaten to confer disease‬
‭resistance. Other vaccines may be adapted to rectal or vaginal applications to elicit‬
‭immune responses in rectal, urinary, or reproductive mucosa. Finally, vaccine antigens‬
‭may be adapted to transdermal application in which the skin is lightly scraped, and‬
‭microneedles are used to pierce the outermost layer. In addition to mobilizing the mucosal‬
‭immune response, this new generation of vaccines may end the anxiety associated with‬
‭injections and, in turn, improve patient participation.‬

‭ igure 19.24‬ ‭The polio vaccine can be administered orally. (credit: modification of work‬
F
‭by UNICEF Sverige;‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭539‬
‭ hapter 20: Mendelian Genetics and Associated‬
C
‭Topics‬

‭ igure 20.1‬ ‭Experimenting with thousands of garden‬‭peas, Mendel uncovered the‬

F
‭fundamentals of genetics. (credit: modification of work by Jerry Kirkhart;‬‭OpenStax‬‭)‬

‭20.1 Introduction‬
‭ enetics can be defined as the study of heredity or the inheritance of genetic information‬
G
‭from generation to generation. Johann Gregor Mendel set the framework for genetics long‬
‭before chromosomes or genes were identified and when meiosis was not well‬
‭understood. Mendel selected a simple biological system and conducted methodical,‬
‭quantitative analyses using large sample sizes. Because of Mendel’s work, the fundamental‬
‭principles of heredity were revealed. We now know that genes, carried on chromosomes,‬
‭are the basic functional units of heredity that can be replicated, expressed, or mutated.‬
‭Today, the postulates put forth by Mendel form the basis of classical, or Mendelian,‬
‭genetics. Not all genes are transmitted from parents to offspring according to Mendel’s‬
‭genetic principles, but Mendel’s experiments serve as an excellent starting point for‬
‭thinking about inheritance. In Chapter 7 (blood types), the concepts of genes, alleles,‬
‭genotypes, phenotypes, and allelic interactions (dominant, recessive, and codominant)‬
‭were introduced as part of the discussion of ABO blood types. In this chapter, we will‬
‭engage in a more in-depth discussion of Mendelian genetics, Punnett squares, pedigree‬
‭analysis, and extensions of Mendelian inheritance.‬

‭540‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

‭20.2 Mendelian Genetics‬

‭Gregor Mendel‬
J‭ ohann Gregor Mendel‬‭(1822–1884) (‬‭Figure 20.2‬‭) was‬‭a lifelong learner, teacher,‬
‭scientist, and man of faith. As a young adult, he joined the Augustinian Abbey of St.‬
‭Thomas in Brno in what is now the Czech Republic. Supported by the monastery, he taught‬
‭physics, botany, and natural science courses at the secondary and university levels. In‬
‭1856, he began a decade-long research project involving inheritance patterns in‬
‭honeybees and plants, ultimately settling on pea plants as his primary model system (a‬
‭system with convenient characteristics used to study a specific biological phenomenon to‬
‭be applied to other systems). In 1865, Mendel presented the results of his experiments‬
‭with nearly 30,000 pea plants to the local Natural History Society. He demonstrated that‬
‭traits are transmitted from parents to offspring independently of other traits and in‬
‭dominant and recessive patterns. In 1866, he published his work,‬‭Experiments in Plant‬
‭Hybridization‬‭(‬‭1‬‭), in the Natural History Society‬‭o f Brü nn proceedings.‬

‭ igure 20.2‬ ‭Johann Gregor Mendel set the framework‬‭for the study of genetics.‬
F
‭(credit:‬ ‭OpenStax‬‭)‬

‭ endel’s work went virtually unnoticed by the scientific community, which believed,‬
M
‭incorrectly, that the inheritance process involved a blending of parental traits that‬
‭produced an intermediate physical appearance in offspring. The‬‭blending theory of‬
‭inheritance‬‭asserted that the original parental traits‬‭were lost or absorbed by the‬

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‭541‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

‭ lending in the offspring, but we now know this is not the case. This hypothetical process‬
b
‭appeared correct because of what we now know as continuous variation.‬‭Continuous‬
‭variation‬‭results from the action of many genes to determine a characteristic like human‬
‭height. Offspring appear to be a “blend” of their parents’ traits.‬
I‭ nstead of continuous characteristics, Mendel worked with traits inherited in distinct‬
‭classes (for example, violet versus white flowers), referred to as‬‭discontinuous‬
‭variation‬‭. Mendel’s choice of these traits allowed‬‭him to experimentally determine that‬
‭the traits were not blended in the offspring, nor were they absorbed, but rather that they‬
‭kept their distinctness and could be passed on. In 1868, Mendel became abbot of the‬
‭monastery and exchanged his scientific pursuits for his pastoral duties. He was not‬
‭recognized for his extraordinary scientific contributions during his lifetime. In fact, it was‬
‭not until 1900 that his work was rediscovered, reproduced, and extended by other‬
‭scientists on the brink of discovering the chromosomal basis of heredity.‬

‭Mendel’s Crosses‬
‭ endel’s seminal work was accomplished using the garden pea,‬‭Pisum sativum‬‭, to study‬
M
‭inheritance. This species naturally self-fertilizes, meaning pollen (the male gamete)‬
‭encounters ova within the same flower. Because every pea plant has both male and female‬
‭reproductive organs, each plant produces both types of gametes required for‬
‭reproduction—pollen and ova. In plants, just as in animals, reproductive organs are‬
‭classified by the size of the gametes produced. The organs producing the smaller pollen‬
‭are called male reproductive organs, while the organs producing the larger ova are called‬
‭female reproductive organs.‬
I‭ n garden peas, the flower petals remain tightly sealed until pollination is complete to‬
‭prevent the pollination of other plants. The result is highly‬‭inbred‬‭, or “‬‭true-breeding,‬‭”‬
‭pea plants that are homozygous at all loci. Therefore, these plants always produce‬
‭o ffspring that look like the parent. By experimenting with true-breeding pea plants,‬
‭Mendel avoided the appearance of unexpected traits in offspring that might occur if the‬
‭plants were not true-breeding. The garden pea also matures within one season, meaning‬
‭several generations could be evaluated relatively quickly. Finally, large quantities of garden‬
‭peas could be cultivated simultaneously, allowing Mendel to conclude that his results did‬
‭not come about simply by chance.‬
‭ endel performed‬‭hybridizations‬‭, which involve mating‬‭two true-breeding individuals‬
M
‭with different traits or phenotypes for a particular characteristic. Pea plants are naturally‬
‭self-pollinating, so pollen is transferred from the anther of a mature pea plant of one‬
‭variety to the stigma of a separate mature pea plant of the second variety.‬
‭ lants used in first-generation crosses were called‬‭P‬‭, or‬‭parental generation‬‭plants‬
P
‭(‬‭Figure 20.3‬‭). Mendel collected the seeds produced‬‭by the P plants that resulted from‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭542‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

e‭ ach cross and grew them the following season. These offspring were called the‬‭F‬‭1‭,‬ or the‬
‭first filial‬‭(filial = daughter or son) generation.‬‭Once Mendel examined the characteristics‬
‭o f the F‬‭1‬ ‭generation of plants, he allowed them to‬‭self-fertilize naturally. He then collected‬
‭and grew the seeds from the F‬‭1‬ ‭plants to produce the‬‭F‭2‬ ‭,‬ or‬‭second filial‬‭, generation.‬
‭Mendel’s experiments extended beyond the F‬‭2‬ ‭generation‬‭to the F‬‭3‬ ‭generation, F‬‭4‬
‭generation, etc. Still, the ratios of characteristics in the P, F‬‭1‭,‬ and F‬‭2‬ ‭generations were the‬
‭most intriguing and became the basis of Mendel’s postulates.‬
I‭ n his 1865 publication, Mendel reported the results of his crosses involving seven‬
‭different characteristics, each with two contrasting traits. A trait is defined as a variation in‬
‭the physical appearance of a heritable characteristic. The characteristics included plant‬
‭height, seed texture, seed color, flower color, pod size, pod color, and flower position. For‬
‭the characteristic of flower color, for example, the two contrasting traits were white versus‬
‭violet. Mendel generated large numbers of F‬‭1‬ ‭and F‬‭2‬ ‭plants to examine each characteristic‬
‭thoroughly and reported results from thousands of F‬‭2‬ ‭plants.‬
‭ hat results did Mendel find in his crosses for flower color? First, Mendel confirmed that‬
W
‭he was using plants that bred true for white or violet flower color. Irrespective of the‬
‭number of generations that Mendel examined, all self-crossed offspring of parents with‬
‭white flowers had white flowers, and all self-crossed offspring of parents with violet‬
‭flowers had violet flowers. In addition, Mendel confirmed that the pea plants were‬
‭physically identical other than flower color. This was an important check to ensure that the‬
‭two varieties of pea plants only differed regarding flower color.‬
‭ nce these validations were complete, Mendel applied the pollen from a plant with violet‬
O
‭flowers to the stigma of a plant with white flowers. After gathering and sowing the seeds‬
‭from this cross, Mendel found that 100 percent of the F‬‭1‬ ‭hybrid generation had violet‬
‭flowers. Conventional wisdom at that time would have predicted the hybrid flowers to be‬
‭pale violet or for hybrid plants to have equal numbers of white and violet flowers. In‬
‭o ther words, the contrasting parental traits were expected to blend in the offspring.‬
‭Instead, Mendel’s results demonstrated that the white flower trait had disappeared‬
‭entirely in the F‬‭1‬ ‭generation.‬

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‭Chapter 20: Mendelian Genetics and Associated Topics‬

‭ igure 20.3‬ ‭Mendel’s process for performing crosses‬‭included examining flower color.‬
F
‭For the seven characters that Mendel examined, the F‬‭1‬ ‭progeny had the phenotype‬
‭associated with the dominant allele. By intercrossing the F‬‭1‬ ‭generation (heterozygous)‬
‭individuals, the F‬‭2‬ ‭generation of progeny had an approximately‬‭3:1 phenotype ratio of the‬
‭dominant to the recessive form. (credit:‬‭OpenStax‬‭)‬

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‭544‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

I‭ mportantly, Mendel did not stop his experimentation there. He allowed the F‬‭1‬ ‭plants to‬
‭self-fertilize and found that 705 plants in the F‬‭2‬ ‭generation had violet flowers and 224 had‬
‭white flowers. This was a ratio of 3.15 violet flowers to one white flower, or approximately‬
‭3:1. Mendel performed an additional experiment to ascertain differences in the‬
‭inheritance of traits carried in the pollen versus the ovum. When Mendel transferred‬
‭pollen from a plant with violet flowers to fertilize the ova of a plant with white flowers and‬
‭vice versa, he obtained approximately the same ratio irrespective of which gamete‬
‭contributed which trait. This is called a‬‭reciprocal‬‭c ross‬‭—a paired cross in which the‬
‭traits of the male and female in one cross become the traits of the female and male in the‬
‭o ther cross. For the other six characteristics that Mendel examined, the F‬‭1‬ ‭and F‬‭2‬
‭generations behaved in the same way that they behaved for flower color. One of the two‬
‭traits would disappear entirely from the F‬‭1‬ ‭generation,‬‭o nly to reappear in the F‬‭2‬
‭generation at a ratio of roughly 3:1 (‬‭Figure 20.4‬‭).‬

‭ igure 20.4‬ ‭Mendel identified seven pea plant characteristics‬‭associated with‬‭pea plants'‬
F
‭seeds, flowers, pods, and stems, with each‬‭characteristic‬‭expressed as one of two versions‬
‭o r traits.‬‭He used these observable traits as the‬‭basis for his breeding experiments, noting‬
‭which traits were expressed (or dominant) and unexpressed (or recessive) in offspring.‬

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‭545‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

‭ pon compiling his results for thousands of plants, Mendel concluded that the‬
U
‭characteristics could be divided into expressed and latent traits. He called these dominant‬
‭and recessive traits, respectively.‬‭Dominant traits‬‭are those that are inherited unchanged‬
‭in a hybridization.‬‭Recessive traits‬‭become latent‬‭o r disappear in the offspring of a‬
‭hybridization. The recessive trait does, however, reappear in the progeny of the hybrid‬
‭o ffspring. An example of a dominant trait is the violet-colored flower trait. White-colored‬
‭flowers are a recessive trait for this same characteristic (flower color). The fact that the‬
‭recessive trait reappeared in the F‬‭2‬ ‭generation meant that the traits remained separate‬
‭(and were not blended) in the plants of the F‬‭1‬ ‭generation.‬‭Mendel proposed that this was‬
‭because the plants possessed two copies of the trait for the flower-color characteristic‬
‭and that each parent transmitted one of their two copies to their offspring, where they‬
‭came together. Moreover, the physical observation of a dominant trait could mean that the‬
‭o rganism's genetic composition included two dominant versions of the characteristic or‬
‭that it included one dominant and one recessive version. Conversely, observing a recessive‬
‭trait meant that the organism lacked any dominant versions of this characteristic.‬

‭A Modern Interpretation of Mendel’s Crosses‬

‭ endel deduced from his results that each individual had two discrete copies of the‬
M
‭characteristic that were passed individually to offspring. We now call those two copies‬
‭genes‬‭, which are carried on‬‭c hromosomes‬‭. We have two‬‭copies of each gene because we‬
‭inherit one from each parent. In fact, the two copies of each gene are located on paired‬
‭homologous chromosomes‬‭. Recall that in meiosis, these‬‭chromosomes are separated‬
‭during meiosis I. This separation, or segregation, of the homologous chromosomes, also‬
‭means that only one of the copies of the gene gets moved into a gamete. The offspring‬
‭form when that gamete unites with one from another parent (i.e., fertilization) and the two‬
‭copies of each gene (and chromosome) are restored.‬
‭ or cases in which a single gene controls a single characteristic, a diploid organism has‬
F
‭two genetic copies that may or may not encode the same version of that characteristic. For‬
‭example, one individual may carry a gene that determines white flower color and a gene‬
‭that determines violet flower color. Gene variants arising from mutation and existing at‬
‭the same relative locations on homologous chromosomes are called‬‭alleles‬‭. Mendel‬
‭examined the inheritance of genes with just two allele forms, but it is common to‬
‭encounter more than two alleles for any given gene in a natural population.‬
‭ wo alleles for a given gene in a diploid organism are expressed and interact to produce‬
T
‭physical characteristics. The observable traits expressed by an organism are referred to as‬
‭its‬‭phenotype‬‭. An organism’s genetic makeup, consisting‬‭o f physically visible and‬
‭non-expressed alleles, is called its‬‭genotype‬‭. Mendel’s‬‭hybridization experiments‬
‭demonstrate the difference between phenotype and genotype. For example, the‬

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‭546‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

‭ henotypes that Mendel observed in his crosses between pea plants with differing traits‬
p
‭are connected to the diploid genotypes of the plants in the P, F‬‭1‭,‬ and F‬‭2‬ ‭generations.‬
‭ e will use a second trait that Mendel investigated, seed color, as an example. A single‬
W
‭gene with two alleles governs seed color. The yellow-seed allele is‬‭dominant,‬‭and the‬
‭green-seed allele is‬‭recessive‬‭. When true-breeding‬‭plants were cross-fertilized, in which‬
‭o ne parent had yellow seeds and one had green seeds, all F‬‭1‬ ‭hybrid offspring had yellow‬
‭seeds. The hybrid offspring were phenotypically identical to the true-breeding parent with‬
‭yellow seeds. However, we know that the allele donated by the parent with green seeds‬
‭was not simply lost because it reappeared in some of the F‬‭2‬ ‭o ffspring (‬‭Figure 20.5‬‭).‬
‭Therefore, the F‬‭1‬ ‭plants must have been genotypically‬‭different from the parent with‬
‭yellow seeds.‬
‭ he P plants that Mendel used in his experiments were each‬‭homozygous‬‭for the trait he‬
T
‭was studying. Diploid organisms that are homozygous for a gene have two identical alleles,‬
‭o ne on each of their homologous chromosomes. The genotype is often written as‬‭YY‬‭o r‬‭yy‬‭,‬
‭with each letter representing one of the two alleles in the genotype. The dominant allele is‬
‭capitalized, and the recessive allele is lowercase. The letter used for the gene (seed color,‬
‭in this case) is usually related to the dominant trait (yellow allele, in this case, or “‬‭Y‭”‬ ).‬
‭Mendel’s parental pea plants always bred true because both produced gametes carried‬
‭the same allele. When P plants with contrasting traits were cross-fertilized, all of the‬
‭o ffspring were‬‭heterozygous‬‭for the contrasting trait,‬‭meaning their genotype had‬
‭different alleles for the gene being examined. For example, the F‬‭1‬ ‭yellow plants that‬
‭received a‬‭Y‬‭allele from their yellow parent and a‬‭y‬‭allele from their green parent had the‬
‭genotype‬‭Yy‬‭.‬

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‭547‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

‭ igure 20.5‬ ‭Phenotypes are physical expressions‬‭o f traits transmitted by alleles. Capital‬
F
‭letters represent dominant alleles, and lowercase letters represent recessive alleles. The‬
‭phenotypic ratios are the ratios of visible characteristics. Genotypic ratios are the ratio of‬
‭gene combinations in the offspring, which are not always distinguishable in the‬
‭phenotypes (credit:‬‭OpenStax‬‭). A link to a video‬‭explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭Law of Dominance‬
‭ ur discussion of homozygous and heterozygous organisms brings us to why the F‬‭1‬
O
‭heterozygous offspring were identical to one of the parents rather than expressing both‬
‭alleles. One of the two contrasting alleles was dominant in all seven pea-plant‬
‭characteristics, and the other was recessive. Mendel called the dominant allele the‬
‭expressed unit factor‬‭; the recessive allele was referred to as the‬‭latent unit factor‬‭. We‬
‭now know these so-called unit factors are genes on homologous chromosomes. For a‬
‭gene expressed in a dominant and recessive pattern, homozygous dominant and‬
‭heterozygous organisms will look identical (that is, they will have different genotypes but‬
‭the same phenotype). The traits of the recessive allele will only be observed in‬
‭homozygous recessive individuals (‬‭Figure 20.6‬‭).‬

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‭548‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

‭ omozygous‬
H ‭ eterozygous‬
H ‭ omozygous‬
H
‭Genotype‬ ‭YY‬ ‭Yy‬ ‭yy‬
‭Phenotype‬ ‭yellow‬ ‭yellow‬ ‭green‬

‭ igure 20.6‬ ‭Correspondence between genotype and‬‭phenotype for a dominant-recessive‬

F
‭characteristic. (credit:‬‭OpenStax‬‭)‬

‭ endel’s law of dominance‬‭states that in a heterozygote,‬‭o ne trait will conceal the‬

M
‭presence of another trait for the same characteristic. For example, when crossing‬
‭true-breeding (homozygous) violet-flowered plants with true-breeding white-flowered‬
‭plants, all of the offspring were violet-flowered, even though they all had one allele for‬
‭violet and one allele for white. Rather than both alleles contributing to a phenotype, the‬
‭dominant allele will be expressed exclusively. The recessive allele will remain latent but‬
‭will be transmitted to offspring in the same manner as that by which the dominant allele is‬
‭transmitted. The recessive trait will only be expressed by offspring with two copies of this‬
‭allele (‬‭Figure 20.7‬‭), and these offspring will breed‬‭true when self-crossed.‬

‭ igure 20.7‬ ‭The allele for albinism, expressed here in humans, is recessive. Each of this‬
F
‭child’s parents carried the recessive allele, meaning they were heterozygous for the‬
‭associated gene. The presence of a copy of the dominant allele meant that their‬
‭phenotype for skin color was “normal” or unaffected. (credit:‬‭OpenStax‬‭)‬

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‭549‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

S‭ everal conventions exist for referring to genes and alleles. In this chapter, we will‬
‭abbreviate genes using the first letter of the gene’s corresponding dominant trait. For‬
‭example, violet is the dominant trait for a pea plant’s flower color, so the flower-color‬
‭gene would be abbreviated as‬‭V‬‭(note that it is customary‬‭to italicize gene designations).‬
‭Furthermore, we will use uppercase and lowercase letters to represent dominant and‬
‭recessive alleles. Therefore, we would refer to the genotype of a homozygous dominant‬
‭pea plant with violet flowers as‬‭VV‬‭, a homozygous‬‭recessive pea plant with white flowers‬
‭as‬‭vv‬‭, and a heterozygous pea plant with violet flowers‬‭as‬‭Vv‬‭.‬

‭Using a Punnett Square with Monohybrid Crosses‬

‭ hen fertilization occurs between two true-breeding parents that differ in only one‬
W
‭characteristic, the process is called a‬‭monohybrid‬‭c ross‬‭, and the resulting offspring are‬
‭monohybrids. Mendel performed seven monohybrid crosses involving contrasting traits‬
‭for each characteristic (‬‭Figure 20.4‬‭). Based on his‬‭results in F‬‭1‬ ‭and F‬‭2‬ ‭generations,‬
‭Mendel postulated that each parent in the monohybrid cross contributed one of two‬
‭paired unit factors to each offspring, and every possible combination of unit factors was‬
‭equally likely.‬
‭ he results of Mendel’s research can be explained in terms of‬‭probabilities‬‭, which are‬
T
‭mathematical measures of likelihood. The probability of an event is calculated by the‬
‭number of times the event occurs divided by the total number of opportunities for the‬
‭event to occur. A probability of one (100 percent) for some event indicates that it is‬
‭guaranteed to occur. In contrast, a probability of zero (0 percent) suggests that it is‬
‭guaranteed not to occur, and a probability of 0.5 (50 percent) means it has an equal‬
‭chance of occurring or not occurring.‬
‭ o demonstrate a monohybrid cross, consider the case of true-breeding pea plants with‬
T
‭yellow versus green pea seeds. The dominant seed color is yellow; therefore, the parental‬
‭genotypes were‬‭YY‬‭for the plants with yellow seeds‬‭and‬‭yy‬‭for the plants with green seeds,‬
‭respectively. A‬‭Punnett square‬‭, devised by the British‬‭geneticist Reginald Punnett, can be‬
‭drawn that applies the rules of probability to predict the possible outcomes of a genetic‬
‭cross or mating and their expected frequencies. When preparing a Punnett square, all‬
‭possible combinations of the parental alleles are listed along the top (for one parent) and‬
‭side (for the other parent) of a grid, representing their meiotic segregation into haploid‬
‭gametes. Then, the combinations of egg and sperm are made in the boxes on the table to‬
‭show which alleles are combining. Each box represents the diploid genotype of a zygote,‬
‭o r fertilized egg, that could result from this mating. Because each possibility is equally‬
‭likely, genotypic ratios can be determined from a Punnett square. The phenotypic ratios‬
‭can also be inferred if the inheritance pattern (dominant or recessive) is known. Each‬
‭parent contributes one type of allele for a monohybrid cross of two true-breeding‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ arents. In this case, only one genotype is possible. All offspring are‬‭Yy‬‭and have yellow‬
p
‭seeds (‬‭Figure 20.8‬‭).‬

‭ igure 20.8‬ ‭In the P generation, pea plants that‬‭are true-breeding for the dominant‬
F
‭yellow phenotype are crossed with recessive green phenotype plants. This cross produces‬
‭F‭1‬ ‬ ‭heterozygotes with a yellow phenotype. Punnett‬‭square analysis can be used to predict‬
‭the genotypes of the F‬‭2‬ ‭generation. (credit:‬‭OpenStax‬‭)‬

‭ hen the F‬‭1‬ ‭o ffspring are crossed with each other,‬‭each has an equal probability of‬
W
‭contributing either a‬‭Y‬‭o r a‬‭y‬‭to the F‬‭2‬ ‭o ffspring.‬‭The result is a 1 in 4 (25 percent)‬
‭probability of both parents contributing a‬‭Y‬‭, resulting‬‭in an offspring with a yellow‬

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‭ henotype; a 25 percent probability of parent 1 contributing a‬‭Y‬‭and parent 2 a‬‭y‭,‬‬

p
‭resulting in offspring with a yellow phenotype; a 25 percent probability of parent 1‬
‭contributing a‬‭y‬‭and parent 2 a‬‭Y‬‭, also resulting‬‭in a yellow phenotype; and a (25 percent)‬
‭probability of both parents contributing a‬‭y‬‭, resulting‬‭in a green phenotype. When‬
‭counting all four possible outcomes, there is a 3 in 4 likelihood of offspring having the‬
‭yellow phenotype and a 1 in 4 probability of offspring having the green phenotype. This‬
‭explains why Mendel’s F‬‭2‬ ‭generation results occurred‬‭in a‬‭3:1 phenotypic ratio‬‭. Using‬
‭large numbers of crosses, Mendel calculated probabilities, found that they fit the‬
‭inheritance model, and used these to predict the outcomes of other crosses (‬‭Figure 20.9‬‭).‬

‭ ‬‭1‬
F
‭ ontrasting‬
C ‭Offspring‬ ‭ ‬‭2‬ ‭Trait‬
F
‭Characteristic‬ ‭P‬‭0‬ ‭Traits‬ ‭Traits‬ ‭ ‬‭2‬ ‭Offspring Traits‬
F ‭Ratios‬
‭ lower color‬
F ‭Violet vs.‬ ‭100 percent‬ ‭•‬ ‭705 violet‬ ‭3.15:1‬
‭white‬ ‭violet‬ ‭•‬ ‭224 white‬
‭ lower‬
F ‭ xial vs.‬
A ‭ 00 percent‬ •‭ ‬
1 ‭ 51 axial‬
6 ‭3.14:1‬
‭position‬ ‭terminal‬ ‭axial‬ ‭•‬ ‭207 terminal‬
‭Plant height‬ ‭Tall vs. dwarf‬ 1
‭ 00 percent‬ •‭ ‬ ‭ 87 tall‬
7 ‭2.84:1‬
‭tall‬ ‭•‬ ‭277 dwarf‬
‭Seed texture‬ ‭ ound vs.‬
R ‭ 00 percent‬ •‭ ‬
1 ‭ ,474 round‬
5 ‭2.96:1‬
‭wrinkled‬ ‭round‬ ‭•‬ ‭1,850 wrinkled‬
‭Seed color‬ ‭ ellow vs.‬
Y ‭ 00 percent‬ •‭ ‬
1 ‭ ,022 yellow‬
6 ‭3.01:1‬
‭green‬ ‭yellow‬ ‭•‬ ‭2,001 green‬
‭ ea pod‬
P I‭ nflated vs.‬ ‭ 00 percent‬ •‭ ‬
1 ‭ 82 inflated‬
8 ‭2.95:1‬
‭texture‬ ‭constricted‬ ‭inflated‬ ‭•‬ ‭299 constricted‬
‭Pea pod color‬ ‭ reen vs.‬
G ‭ 00 percent‬ •‭ ‬
1 ‭ 28 green‬
4 ‭2.82:1‬
‭yellow‬ ‭green‬ ‭•‬ ‭152 yellow‬

‭Figure 20.9‬ ‭The results of Mendel’s garden pea hybridizations.‬ ‭(credit:‬‭OpenStax‬‭)‬

‭Law of Segregation‬
‭ bserving that true-breeding pea plants with contrasting traits gave rise to F‬‭1‬ ‭generations‬
O
‭that all expressed the dominant trait and F‬‭2‬ ‭generations‬‭that expressed the dominant and‬
‭recessive traits in a 3:1 ratio, Mendel proposed‬‭the‬‭law of segregation‬‭. This law states‬
‭that paired unit factors (genes) must segregate equally into gametes so that offspring have‬
‭an equal likelihood of inheriting either factor. For the F‬‭2‬ ‭generation of a monohybrid‬

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c‭ ross, the following three possible combinations of genotypes result:‬‭homozygous‬

‭dominant‬‭,‬‭heterozygous‬‭, or‬‭homozygous recessive‬‭. Because‬‭heterozygotes could arise‬
‭from two different pathways (receiving one dominant and one recessive allele from either‬
‭parent), and because heterozygous and homozygous dominant individuals are‬
‭phenotypically identical, the law supports Mendel’s observed 3:1 phenotypic ratio. The‬
‭equal segregation of alleles is why we can apply the Punnett square to predict the‬
‭o ffspring of parents with known genotypes accurately. The physical basis of Mendel’s law‬
‭o f segregation is the first division of meiosis in which the homologous chromosomes with‬
‭their different versions of each gene segregate into daughter nuclei. The scientific‬
‭community did not understand this process during Mendel’s lifetime (‬‭Figure 20.10‬‭).‬

‭Figure 20.10‬ ‭The first division in meiosis is shown.‬ ‭(credit:‬‭OpenStax‬‭)‬

‭ or additional information about monohybrid crosses and Punnett squares, click the link‬
F
‭o r scan the QR code to watch the Amoeba Sisters video titled “Monohybrids and the‬
‭Punnett Square Guinea Pigs.”‬

Monohybrids and the Punnett Square Guinea Pigs

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‭Test Cross‬
‭ eyond predicting the offspring of a cross between known homozygous or heterozygous‬
B
‭parents, Mendel also developed a way to determine whether an organism that expressed a‬
‭dominant trait was a heterozygote or a homozygote. A technique called a‬‭test cross‬‭is still‬
‭used by plant and animal breeders. In a test cross, the dominant-expressing organism is‬
‭crossed with a homozygous recessive organism for the same characteristic. If the‬
‭dominant-expressing organism is a homozygote, all F‬‭1‬ ‭o ffspring will be heterozygotes‬
‭expressing the dominant trait (‬‭Figure 20.11‬‭). Alternatively, if the dominant-expressing‬
‭o rganism is a heterozygote, the F‬‭1‬ ‭o ffspring will exhibit a 1:1 ratio of heterozygous and‬
‭recessive homozygotes (‬‭Figure 20.11‬‭). The test cross further validates Mendel’s postulate‬
‭that pairs of unit factors segregate equally.‬

‭ igure 20.11‬ ‭A test cross determines whether an‬‭o rganism expressing a dominant trait‬
F
‭is a homozygote or a heterozygote (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of‬
‭this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Chapter 20: Mendelian Genetics and Associated Topics‬

‭Law of Independent Assortment‬

‭ endel also established crosses to examine the inheritance of different traits for two‬
M
‭characteristics. Consider seed color and texture characteristics for two pea plants, one‬
‭with wrinkled, green seeds (‬‭rryy‬‭) and another with‬‭round, yellow seeds (‬‭RRYY‬‭). Because‬
‭each parent is homozygous, the law of segregation indicates that the gametes for the‬
‭wrinkled–green plant are all‬‭ry‬‭, and the gametes for‬‭the round–yellow plant are all‬‭RY‬‭.‬
‭Therefore, the F‬‭1‬ ‭o ffspring generation is all‬‭RrYy‬‭(‬‭Figure 20.12‬‭).‬

‭ igure 20.12‬ ‭A dihybrid cross in pea plants involves‬‭seed color and texture genes. The P‬
F
‭cross produces F‬‭1‬ ‭o ffspring that are all heterozygous‬‭for both characteristics. The‬
‭resulting 9:3:3:1 F‬‭2‬ ‭phenotypic ratio is obtained‬‭using a Punnett square. (credit:‬
‭OpenStax‬‭). A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ he gametes produced by the F‬‭1‬ ‭individuals must have‬‭o ne allele from each of the two‬
T
‭genes. For example, a gamete could get an‬‭R‬‭allele‬‭for the seed shape gene and either a‬‭Y‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭555‬
‭Chapter 20: Mendelian Genetics and Associated Topics‬

‭ r a‬‭y‬‭allele for the seed color gene. It cannot get an‬‭R‬‭and an‬‭r‬‭allele; each gamete has‬
o
‭o nly one allele per gene. The‬‭law of independent assortment‬‭states that a gamete into‬
‭which an‬‭r‬‭allele is sorted would be equally likely‬‭to contain either a‬‭Y‬‭o r a‬‭y‬‭allele. Thus,‬
‭four equally likely gametes can be formed when the‬‭RrYy‬‭heterozygote is self-crossed:‬‭RY‬‭,‬
‭rY‬‭,‬‭Ry‬‭, and‬‭ry‬‭. Arranging these gametes along the‬‭top and left of a 4 × 4 Punnett square‬
‭(‬‭Figure 20.12‬‭) gives us 16 equally likely genotypic‬‭combinations. These genotypes show a‬
‭phenotypic ratio of 9 round–yellow:3 round–green:3 wrinkled–yellow:1 wrinkled–green‬
‭(‬‭9:3:3:1‬‭;‬‭Figure 20.12‬‭). In other words, 9/16 of the‬‭progeny have the dominant‬
‭phenotype for both genes, 3/16 have the dominant phenotype for the first gene and the‬
‭recessive phenotype for the second one, 3/16 have the recessive phenotype for the first‬
‭gene and the dominant phenotype for the second one, and 1/16 have the recessive‬
‭phenotypes for both genes. These are the expected offspring ratios, assuming we‬
‭performed the crosses with a large enough sample size.‬
‭ endel’s‬‭law of independent assortment‬‭states that genes do not influence each other‬
M
‭concerning the sorting of alleles into gametes, and every possible combination of alleles‬
‭for every gene is equally likely to occur. The physical basis for the law of independent‬
‭assortment also lies in meiosis I, in which the different homologous pairs line up in‬
‭random orientations. Each gamete can contain any combination of paternal and maternal‬
‭chromosomes (and, therefore, the genes on them) because the orientation of tetrads on‬
‭the metaphase plane is random (‬‭Figure 20.13‬‭).‬
‭ or additional information about Gregor Mendel and the significance of his research, click‬
F
‭the link or scan the QR code to watch the TED-Ed video by Hortensia Jimenez Diaz titled‬
‭“How Mendel’s pea plants helped us understand genetics.”‬

How Mendel's pea plants helped us understand genetics - Hort…

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‭ igure 20.13‬ ‭The random segregation into daughter‬‭nuclei during the first division in‬
F
‭meiosis can lead to various possible genetic arrangements.‬
‭(credit:‬‭OpenStax‬‭)‬

‭ or additional information about dihybrid crosses and Punnett squares, click the link or‬
F
‭scan the QR code to watch the Amoeba Sisters video titled “Dihybrid and two-trait‬
‭crosses.”‬

Dihybrid and Two-Trait Crosses

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‭20.3 Probability Basics‬

‭ s stated earlier, probabilities are mathematical measures of likelihood. The empirical‬
A
‭probability of an event is calculated by dividing the number of times the event occurs by‬
‭the total number of opportunities for the event to occur. It is also possible to calculate‬
‭theoretical probabilities by dividing the number of times an event is‬‭expected‬‭to occur by‬
‭the number of times it could occur. Empirical probabilities come from observations like‬
‭those of Mendel. On the other hand, theoretical probabilities come from knowing how the‬
‭events are produced and assuming that the probabilities of individual outcomes are equal.‬
‭A probability of one for some event indicates that it is guaranteed to occur, whereas a‬
‭probability of zero indicates that it is guaranteed not to occur. An example of a genetic‬
‭event is a round seed produced by a pea plant.‬
I‭ n one experiment, Mendel demonstrated that the probability of the event “round seed”‬
‭o ccurring was one in the F‬‭1‬ ‭o ffspring of true-breeding‬‭parents, one of which has round‬
‭seeds and one of which has wrinkled seeds. When the F‬‭1‬ ‭plants were subsequently‬
‭self-crossed, the probability of any given F‬‭2‬ ‭o ffspring‬‭having round seeds was three out of‬
‭four. In other words, in a large population of F‬‭2‬ ‭o ffspring chosen at random, 75 percent‬
‭were expected to have round seeds, whereas 25 percent were expected to have wrinkled‬
‭seeds. Using large numbers of crosses, Mendel was able to calculate probabilities and use‬
‭these to predict the outcomes of other crosses.‬

‭The Product Rule‬

‭ endel demonstrated that pea plants transmit characteristics as discrete units from‬
M
‭parent to offspring. As will be discussed, Mendel also determined that different‬
‭characteristics, like seed color and seed texture, were transmitted independently of one‬
‭another and could be considered in separate probability analyses. For instance,‬
‭performing a cross between a plant with green, wrinkled seeds and a plant with yellow,‬
‭round seeds still produced offspring with a 3:1 ratio of yellow: green seeds (ignoring seed‬
‭texture) and a 3:1 ratio of wrinkled: round seeds (ignoring seed color). The characteristics‬
‭o f color and texture did not influence each other.‬
‭ he product rule of probability can be applied to this phenomenon of the independent‬
T
‭transmission of characteristics. The‬‭product rule‬‭states that the probability of two‬
‭independent events occurring together can be calculated by multiplying the individual‬
‭probabilities of each event occurring alone. To demonstrate the product rule, imagine‬
‭rolling a six-sided die (D) and flipping a penny (P) simultaneously. The die may roll any‬
‭number from 1–6 (D‬‭#‭)‬ , whereas the penny may turn up‬‭heads (P‬‭H‬‭) or tails (P‬‭T‬‭). Rolling the‬
‭die does not affect the outcome of flipping the penny and vice versa. There are 12 possible‬
‭o utcomes of this action (‬‭Figure 20.14‬‭), and each event‬‭is expected to occur with equal‬
‭probability.‬

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‭ olling Die‬
R ‭ lipping Penny‬
F
‭D‬‭1‬ ‭P‬‭H‬
‭D‬‭1‬ ‭P‬‭T‬
‭D‬‭2‬ ‭P‬‭H‬
‭D‬‭2‬ ‭P‬‭T‬
‭D‬‭3‬ ‭P‬‭H‬
‭D‬‭3‬ ‭P‬‭T‬
‭D‬‭4‬ ‭P‬‭H‬
‭D‬‭4‬ ‭P‬‭T‬
‭D‬‭5‬ ‭P‬‭H‬
‭D‬‭5‬ ‭P‬‭T‬
‭D‬‭6‬ ‭P‬‭H‬
‭D‬‭6‬ ‭P‬‭T‬

‭ igure 20.14‬ ‭Twelve Equally Likely Outcomes of Rolling‬‭a Die and Flipping a Penny‬
F
‭(credit:‬‭OpenStax‬‭)‬

‭ f the 12 possible outcomes, the die has a 2/12 (or 1/6) probability of rolling a two, and‬
O
‭the penny has a 6/12 (or 1/2) probability of coming up heads. By the product rule, the‬
‭likelihood that you will obtain the combined outcome a 2 and heads is (D‬‭2‭)‬ x (P‬‭H‬‭) = (1/6)‬
‭x (1/2) or 1/12. Notice the word “and” in the description of the probability. The “and” is a‬
‭signal to apply the product rule. For example, consider how the product rule is applied to‬
‭the dihybrid cross: the probability of having both dominant traits (for example, yellow and‬
‭round) in the F‬‭2‬ ‭progeny is the product of the probabilities‬‭o f having the dominant trait‬
‭for each characteristic, as shown here:‬
‭‬
3 ‭3‬ ‭9‬
‭4‬
‭‬ × ‭‬ ‭4‬ ‭‬ = ‭‬ ‭16‬

‭The Sum Rule‬

‭ he sum rule of probability is applied when considering two mutually exclusive outcomes‬
T
‭that can come about by more than one pathway. The‬‭sum rule‬‭states that the likelihood of‬
‭o ne event or the other event of two mutually exclusive events is the sum of their individual‬
‭probabilities. Notice the word “or” in the description of the likelihood. The “or” indicates‬

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t‭ hat you should apply the sum rule. Let’s imagine you are flipping a penny (P) and a‬
‭quarter (Q). What is the probability of one coin coming up heads and one coin coming up‬
‭tails? This outcome can occur in two ways: the penny lands on heads (P‬‭H‬‭), and the quarter‬
‭lands on tails (Q‬‭T‭)‬ , or the quarter lands on heads‬‭(Q‬‭H‬‭), and the penny lands on tails (P‬‭T‭)‬ .‬
‭Either case fulfills the outcome. By the sum rule, we calculate the probability of obtaining‬
‭o ne head and one tail as [(P‬‭H‬‭) × (Q‬‭T‭)‬ ] + [(Q‬‭H‭)‬ × (P‬‭T‭)‬ ]‬‭= [(1/2) × (1/2)] + [(1/2) × (1/2)] =‬
‭1/2.‬
‭ ou should also notice that we used the product rule to calculate the probability of P‬‭H‬ ‭and‬
Y
‭Q‭T‬ ‬ ‭and the probability of P‬‭T‬ ‭and Q‬‭H‬ ‭before we summed‬‭them. Again, the sum rule can be‬
‭applied to show the likelihood of having at least one dominant trait in the F‬‭2‬ ‭generation of‬
‭a dihybrid cross:‬

( ‭‬
1
‭4‬
×
‭‬
3
‭4‬ )+ ( ‭‬
3
‭4‬
×
‭‬
1
‭4‬ )= ‭3‬
‭16‬
+
‭3‬
‭16‬
=
‭6‬
‭16‬
=
‭‬
3
‭8‬

‭ o use probability laws in practice, we must work with large sample sizes because small‬
T
‭sample sizes are prone to deviations caused by chance. The large quantities of pea plants‬
‭that Mendel examined allowed him to calculate the probabilities of the traits appearing in‬
‭his F‬‭2‬ ‭generation. As you will learn, this discovery‬‭meant that when parental‬
‭characteristics are known, the offspring’s traits can predicted accurately even before‬
‭fertilization.‬
‭ or additional information about the product and sum rules, click the link or scan the QR‬
F
‭code to watch the Bozeman Science video titled “Probability in Genetics: Multiplication and‬
‭Addition Rules.”‬

Probability in Genetics: Multiplication and Addition Rules

‭20.4 Extensions of Mendel’s Laws‬

‭ endel’s experiments with pea plants suggested that (1) two “units” or alleles exist for‬
M
‭every gene; (2) alleles maintain their integrity in each generation (no blending); and (3) in‬
‭the presence of the dominant allele, the recessive allele is hidden and makes no‬
‭contribution to the phenotype. Therefore, recessive alleles can be “carried” and not‬
‭expressed by individuals. Such heterozygous individuals are known as “‬‭c arriers‬‭.” Further‬
‭genetic studies in other plants and animals have shown that much more complexity exists‬
‭but that the fundamental principles of Mendelian genetics still hold. In the following‬
‭sections, we consider some of the‬‭extensions‬‭o f Mendelism.‬‭If Mendel had chosen an‬

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e‭ xperimental system that exhibited these genetic complexities, he likely couldn’t have‬
‭explained the results.‬

‭Incomplete Dominance‬
‭ endel’s results, demonstrating that traits are inherited as dominant and recessive pairs,‬
M
‭contradicted the view at that time that offspring exhibited a blend of their parents’ traits.‬
‭However, the heterozygote phenotype occasionally does appear to be intermediate‬
‭between the two parents. For example, in the snapdragon,‬‭Antirrhinum majus‬‭(‬‭Figure‬
‭20.15‬‭), a cross between a homozygous parent with white‬‭flowers (‬‭C‬‭W‬‭C‭W ‬ ‬‭) and a‬
‭homozygous parent with red flowers (‬‭C‬‭R‭C ‬ ‭R‬
‬ ‭) will produce‬‭o ffspring with pink flowers‬
‭R‬ ‭W‬
‭(‬‭C‬ ‭C‬ ‭). Note that different genotypic abbreviations‬‭are used for Mendelian extensions to‬
‭distinguish these patterns from simple dominance and recessiveness. This inheritance‬
‭pattern is described as‬‭incomplete dominance‬‭, meaning‬‭that one of the alleles appears‬
‭in the phenotype in the heterozygote but not to the exclusion of the other, which can also‬
‭be seen. The allele for red flowers is incompletely dominant over the allele for white‬
‭flowers. However, the results of a heterozygote self-cross can still be predicted, just as‬
‭with Mendelian dominant and recessive crosses. In this case, the genotypic ratio would be‬
‭1‬‭C‬‭R‭C‬ ‭R‬
‬ ‭:2‬‭C‭R‬ ‭C
‬ ‭W‬
‬ ‭:1‬‭C‬‭W‭C
‬ ‭W‬
‬ ‭, and the phenotypic ratio would‬‭be 1:2:1 for red:pink:white. The‬
‭basis for the intermediate color in the heterozygote is that the pigment produced by the‬
‭red allele (anthocyanin) is diluted in the heterozygote and, therefore, appears pink‬
‭because of the white background of the flower petals.‬

‭ igure 20.15‬ ‭The pink flowers of a heterozygote snapdragon‬‭result from incomplete‬

F
‭dominance of the red color allele to the white color allele (credit:‬‭Bennilover‬‭;‬‭Snapdragon‬
‭Dragon Weather‬‭; Flickr;‬‭CC BY-ND 2.0‬‭)‬

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‭Codominance‬
‭ variation on incomplete dominance is‬‭codominance‬‭, in which both alleles for the same‬
A
‭characteristic are simultaneously expressed in the heterozygote. An example of‬
‭codominance occurs in the ABO blood groups of humans. The A and B alleles are‬
‭expressed as A or B molecules on the surface of red blood cells. Homozygotes (‬‭I‭A‬ ‬‭I‭A‬ ‬ ‭and‬
‭I‭B‬ ‭I‬ ‭B‬ ‬‭) express either the A or the B phenotype, and‬‭heterozygotes (‬‭I‭A‬ ‬‭I‭B‬ ‭)‬ express both‬
‭phenotypes equally. The‬‭I‭A‬ ‬‭I‭B‬ ‬ ‭individual has blood‬‭type AB. The three possible offspring‬
‭genotypes are phenotypically distinct in a self-cross between heterozygotes expressing a‬
‭codominant trait. However, a Mendelian monohybrid cross's 1:2:1 genotypic ratio‬
‭characteristic still applies (‬‭Figure 20.16‬‭).‬

‭ igure 20.16‬ ‭This Punnett square shows an AB/AB‬‭blood type cross. Note the‬
F
‭phenotype ratio in the progeny is 1:2:1, not the typical 3:1 observed in Mendel’s crosses.‬
‭(credit:‬‭OpenStax‬‭)‬

‭Multiple Alleles‬
‭ endel implied that only two alleles, one dominant and one recessive, could exist for a‬
M
‭given gene. We now know that this is an oversimplification. Although individual humans‬
‭(and all diploid organisms) can only have two alleles for a given gene, multiple alleles may‬
‭exist at the population level, such that many combinations of two alleles are observed.‬
‭Note that when many alleles exist for the same gene, the convention is to denote the most‬

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c‭ ommon phenotype or genotype in the natural population as the wild type (often‬
‭abbreviated “+”). All other phenotypes or genotypes are considered variants (mutants) of‬
‭this typical form, meaning they deviate from the wild type. The variant may be recessive or‬
‭dominant to the wild-type allele.‬
‭ n example of multiple alleles is the ABO blood-type system in humans was discussed in‬
A
‭Chapter 7. In this case, there are three alleles in the population. The‬‭I‬‭A‬ ‭allele codes for A‬
‭molecules on the red blood cells, the‬‭I‭B‬ ‬ ‭allele codes‬‭for B molecules on the surface of red‬
‭blood cells, and the‬‭i‬‭allele codes for no molecules‬‭o n the red blood cells. In this case, the‬
‭I‭A‬ ‬ ‭and‬‭I‬‭B‬ ‭alleles are codominant with each other and‬‭are both dominant over the‬‭i‬‭allele.‬
‭Although three alleles are present in a population, each individual only gets two from their‬
‭parents. This produces the genotypes and phenotypes shown in‬‭Figure 20.17‬‭. Notice that‬
‭instead of three genotypes, there are six different genotypes when there are three alleles.‬
‭The number of possible phenotypes depends on the dominance relationships between the‬
‭three alleles.‬

‭ igure 20.17‬ ‭Inheritance of the ABO blood system‬‭in humans is shown.‬

F
‭(credit:‬‭OpenStax‬‭)‬

‭ or additional information about multiple alleles and Punnett squares, click the link or‬
F
‭scan the QR code to watch the Amoeba Sisters video titled “Multiple alleles (ABO blood‬
‭types) and Punnett squares.”‬

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Multiple Alleles (ABO Blood Types) and Punnett Squares

‭Evolution Connection‬

‭ ultiple Alleles Confer Drug Resistance in the Malaria Parasite‬

M
‭Malaria is a parasitic disease in humans transmitted by infected female mosquitoes,‬
‭including‬‭Anopheles gambiae‬‭(‬‭Figure 20.18a‬‭), and is‬‭characterized by cyclic high fevers,‬
‭chills, flu-like symptoms, and severe anemia.‬‭Plasmodium‬‭falciparum‬‭and‬‭P. vivax‬‭are‬
‭malaria's most common causative agents, and‬‭P. falciparum‬‭is the most deadly (‬‭Figure‬
‭20.18b‬‭)‬‭.‬‭When promptly and correctly treated,‬‭P. falciparum‬‭malaria has a mortality rate‬
‭o f 0.1 percent. However, in some parts of the world, the parasite has evolved to be‬
‭resistant to commonly used malaria treatments, so the most effective malaria treatments‬
‭can vary by geographic region.‬

‭ igure 20.18‬ ‭The (a)‬‭Anopheles gambiae‬‭, or African‬‭malaria mosquito, acts as a vector in‬
F
‭the transmission to humans of the malaria-causing parasite (b)‬‭Plasmodium falciparum‬‭,‬
‭here visualized using false-color transmission electron microscopy. (credit a: James D.‬
‭Gathany; credit b: Ute Frevert; false color by Margaret Shear; scale-bar data from Matt‬
‭Russell;‬‭OpenStax‬‭)‬

I‭ n Southeast Asia, Africa, and South America,‬‭P. falciparum‬‭has developed resistance to the‬
‭anti-malarial drugs chloroquine, mefloquine, and sulfadoxine-pyrimethamine.‬‭P.‬

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f‭ alciparum‬‭, which is haploid during the life stage in which it is infectious to humans, has‬
‭evolved multiple drug-resistant mutant alleles of the‬‭dhps‬‭gene. Varying degrees of‬
‭sulfadoxine resistance are associated with each of these alleles. Being haploid,‬‭P.‬
‭falciparum‬‭needs only one drug-resistant allele to‬‭express this trait.‬

I‭ n Southeast Asia, different sulfadoxine-resistant alleles of the‬‭dhps‬‭gene are localized to‬

‭various geographic regions. This is a common evolutionary phenomenon because‬
‭drug-resistant mutants arise in a population and interbreed with other‬‭P. falciparum‬
‭isolates nearby. Sulfadoxine-resistant parasites cause considerable human hardship in‬
‭regions where this drug is widely used as an over-the-counter malaria remedy. As is‬
‭typical with pathogens that multiply to large numbers within an infection cycle,‬‭P.‬
‭falciparum‬‭evolves relatively rapidly (over a decade‬‭o r so) in response to the selective‬
‭pressure of commonly used anti-malarial drugs. For this reason, scientists must constantly‬
‭work to develop new drugs or drug combinations to combat the worldwide malaria‬
‭burden (‬‭2‬‭)‬

I‭ n late 2021, R21/Matrix-M became the first vaccine recommended for widespread use by‬
‭the World Health Organization. At least ten other candidate vaccines are in development.‬
‭The effort is a multinational one involving governments, universities, nonprofits,‬
‭philanthropists, and pharmaceutical companies. Much of the recent progress can be‬
‭credited to organizations within the most affected countries, such as the Malaria Research‬
‭and Training Center in Mali. Founded by Ogobara Duombo and Yeya Touré in the 1990s,‬
‭the center has emerged as a primary front-line research driver, including running many‬
‭critical clinical trials important to vaccine development and approval.‬

‭Sex-Linked Traits‬
I‭ n humans, as well as in many other animals and some plants, the sex of the individual is‬
‭determined by‬‭sex chromosomes‬‭—one pair of non-homologous‬‭chromosomes. Until now,‬
‭we have only considered inheritance patterns among non-sex chromosomes or‬
‭autosomes. In addition to 22 homologous pairs of autosomes, human females have a‬
‭homologous pair of X chromosomes, whereas human males have an XY chromosome pair.‬
‭Although the Y chromosome contains a small region of similarity to the X chromosome so‬
‭that they can pair during meiosis, the Y chromosome is much shorter and contains fewer‬
‭genes. When a gene being examined is present on the X chromosome but not the Y, it is‬
‭X-linked‬‭.‬
‭ ye color in‬‭Drosophila‬‭, the common fruit fly, was‬‭the first X-linked trait to be identified.‬
E
‭Thomas Hunt Morgan mapped this trait to the X chromosome in 1910.‬‭Drosophila‬‭males‬
‭have an XY chromosome pair like humans, and females are XX. In flies, the wild-type eye‬
‭color is red (X‬‭W‭)‬ and is dominant to white eye color‬‭(X‬‭w‭)‬ (‬‭Figure 20.19‬‭). Because of the‬

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l‭ocation of the eye-color gene, reciprocal crosses do not produce the same offspring‬
‭ratios. Males are said to be‬‭hemizygous‬‭because they‬‭have only one allele for any X-linked‬
‭characteristic. Hemizygosity makes descriptions of dominance and recessiveness‬
‭irrelevant for XY males.‬‭Drosophila‬‭males lack the‬‭white gene on the Y chromosome; their‬
‭genotype can only be X‬‭W‭Y ‬
or X‬‭w‬‭Y. In contrast, females‬‭have two allele copies of this gene,‬
‭and can be X‬‭W‬‭X‭W
‬ ‭,‬ X‬‭W‬‭X‭w‬ ‬‭, or X‬‭w‬‭X‭w‬ ‭.‬ ‬

‭ igure 20.19‬ ‭In‬‭Drosophila‬‭, the gene for eye color‬‭is located on the X chromosome. Red‬
F
‭eye color is wild-type and is dominant to white eye color. (credit:‬‭OpenStax‬‭)‬

I‭ n an X-linked cross, the genotypes of F‬‭1‬ ‭and F‬‭2‬ ‭o ffspring‬‭depend on whether the male or‬
‭the female in the P generation expressed the recessive trait. Concerning‬‭Drosophila‬‭eye‬
‭color, when the P male expresses the white-eye phenotype and the female is homozygous‬
‭red-eyed, all members of the F‬‭1‬ ‭generation exhibit‬‭red eyes (‬‭Figure 20.20‬‭). The F‬‭1‬ ‭females‬
‭are heterozygous (X‬‭W‭X ‬ ‭w‬
‬ ‭), and the males are all X‬‭W‭Y
‬
,‬‭having received their X chromosome‬
‭from the homozygous dominant P female and their Y chromosome from the P male. A‬
‭subsequent cross between the X‬‭W‬‭X‭w‬ ‬ ‭female and the X‬‭W‭Y ‬
‬ ‭male would produce only‬
‭W‬ ‭W‬ ‭W‬ ‭w‬
‭red-eyed females (with X‬ ‭X‬ ‭o r X‬ ‭X‬ ‭genotypes) and‬‭both red- and white-eyed males‬
‭(with X‬‭W‬‭Y or X‬‭w‭Y ‬
genotypes). Consider a cross between‬‭a homozygous white-eyed female‬
‭and a male with red eyes. The F‬‭1‬ ‭generation would‬‭exhibit only heterozygous red-eyed‬
‭females (X‬‭W‭X
‬ ‭w‬
‬ ‭) and only white-eyed males (X‬‭w‬‭Y). Half‬‭o f the F‬‭2‬ ‭females would be red-eyed‬

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(‭ X‬‭W‭X
‬ ‭w‬
‬ ‭), and half would be white-eyed (X‬‭w‬‭X‭w‬ ‬‭). Similarly, half of the F‬‭2‬ ‭males would be‬
‭red-eyed (X‬‭W‭Y ‬
), and half would be white-eyed (X‬‭w‭Y ‬
).‬

‭ igure 20.20‬ ‭Crosses involving sex-linked traits‬‭o ften give rise to different phenotypes‬
F
‭for the different sexes of offspring, as is the case for this cross involving red and white eye‬
‭color in Drosophila. In the diagram, w is the white-eye mutant allele, and W is the‬
‭wild-type, red-eye allele. (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation of this figure‬
‭is available at‬‭Biology411.com‬‭.‬

S‭ ex-linkage studies in Morgan’s laboratory provided the fundamentals for understanding‬

‭X-linked recessive disorders in humans, which include red-green color blindness and‬
‭Types A and B hemophilia. Because human males need to inherit only one recessive‬
‭mutant X allele to be affected, X-linked disorders are disproportionately observed in‬
‭males. Females must inherit recessive X-linked alleles from both parents to express the‬
‭trait. When they inherit one recessive X-linked mutant allele and one dominant X-linked‬

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‭ ild-type allele, they are carriers of the trait and are typically unaffected. Carrier females‬
w
‭can manifest mild forms of the trait due to the inactivation of the dominant allele located‬
‭o n one of the X chromosomes. However, female carriers can contribute the trait to their‬
‭sons, resulting in the son exhibiting the trait, or they can contribute the recessive allele to‬
‭their daughters, resulting in the daughters being carriers of the trait (‬‭Figure 20.21‬‭).‬
‭Although some Y-linked recessive disorders exist, typically, they are associated with‬
‭infertility in males and are, therefore, not transmitted to subsequent generations.‬

‭ igure 20.21‬ ‭The son of a woman who is a carrier‬‭o f a recessive X-linked disorder will‬
F
‭have a 50 percent chance of being affected. A daughter will not be affected, but she will‬
‭have a 50 percent chance of being a carrier like her mother. (credit:‬‭OpenStax‬‭). A link to‬
‭a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Epistasis‬
‭ endel’s studies in pea plants implied that the sum of an individual’s phenotype was‬
M
‭controlled by genes (or, as he called them, unit factors), such that a single gene distinctly‬

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a‭ nd completely controlled every characteristic. In fact, single observable characteristics‬

‭are almost always influenced by multiple genes (each with two or more alleles) acting in‬
‭unison. For example, at least eight genes contribute to eye color in humans.‬
I‭ n some cases, several genes can contribute to aspects of a common phenotype without‬
‭their gene products directly interacting. In the case of organ development, for instance,‬
‭genes may be expressed sequentially, with each gene adding to the complexity and‬
‭specificity of the organ. Genes may function in complementary or synergistic fashions,‬
‭such that two or more genes expressed simultaneously affect a phenotype. An apparent‬
‭example of this occurs with human skin color, which involves the action of at least three‬
‭(and probably more) genes. Polygenic inheritance is a case in which a characteristic like‬
‭skin color or human height depends on the combined effects of numerous genes.‬
‭ enes may also oppose each other, with one gene suppressing the expression of another.‬
G
‭In‬‭epistasis‬‭, the interaction between genes is antagonistic,‬‭such that one gene masks or‬
‭interferes with the expression of another. “Epistasis” is a Greek word that means “standing‬
‭upon.” The alleles being masked or silenced are said to be hypostatic to the epistatic alleles‬
‭doing the masking. Often, the biochemical basis of epistasis is a gene pathway in which the‬
‭expression of one gene is dependent on the function of a gene that precedes or follows it‬
‭in the pathway.‬
‭ n example of epistasis is pigmentation in mice. The wild-type coat color, agouti (AA), is‬
A
‭dominant to the solid-colored fur (aa). However, a separate gene C, when present as the‬
‭recessive homozygote (cc), negates any expression of pigment from the A gene and results‬
‭in an albino mouse (‬‭Figure 20.22‬‭). Therefore, the‬‭genotypes‬‭AAcc‬‭,‬‭Aacc‬‭, and‬‭aacc‬‭all‬
‭produce the same albino phenotype. A cross between heterozygotes for both genes (‬‭AaCc‬
‭x‬‭AaCc‬‭) would generate offspring with a phenotypic‬‭ratio of 9 agouti:3 black:4 albino‬
‭(‬‭Figure 20.22‬‭). In this case, the‬‭C‬‭gene is epistatic‬‭to the‬‭A‬‭gene.‬

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‭ igure 20.22‬ ‭In this example of epistasis, one gene‬‭(‬‭C‭)‬ masks the expression of another‬
F
‭(‬‭A‬‭) for coat color. When the‬‭C‬‭allele is present,‬‭coat color is expressed; no coat color is‬
‭expressed when it is absent (‬‭cc‬‭). Coat color depends‬‭o n the‬‭A‬‭gene, which shows‬
‭dominance, with the recessive homozygote showing a different phenotype than the‬
‭heterozygote or dominant homozygote. (credit:‬‭OpenStax‬‭).‬ ‭A link to a video explanation‬
‭o f this figure is available at‬‭Biology411.com‬‭.‬

‭ or additional information about extensions to Mendelian inheritance, click the link or‬
F
‭scan the QR code to watch the Amoeba Sisters video titled “Incomplete dominance,‬
‭codominance, polygenic traits, and epistasis.”‬

Incomplete Dominance, Codominance, Polygenic Traits, and Epist…

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‭Linked Genes Violate the Law of Independent Assortment‬

‭ lthough all of Mendel’s pea plant characteristics behaved according to the law of‬
A
‭independent assortment, we now know that some allele combinations are not inherited‬
‭independently of each other. Genes that are located on separate, non-homologous‬
‭chromosomes will always sort independently. However, each chromosome contains‬
‭hundreds or thousands of genes, organized linearly on chromosomes like beads on a‬
‭string. The segregation of alleles into gametes can be influenced by‬‭linkage‬‭, in which‬
‭genes located physically close to each other on the same chromosome are more likely to‬
‭be inherited as a pair because they are less likely to be separated by crossing over.‬
‭However, because of the process of‬‭recombination‬‭,‬‭o r “‬‭c rossover‬‭,” two genes on the‬
‭same chromosome can behave independently or as if they are not linked. To understand‬
‭this, let us consider the biological basis of gene linkage and recombination.‬
‭ omologous chromosomes possess the same genes in the same order, though the specific‬
H
‭alleles of the gene can be different on each of the two chromosomes. Recall that during‬
‭interphase and prophase I of meiosis, homologous chromosomes first replicate and then‬
‭synapse, with like genes on the homologs aligning with each other. At this stage, segments‬
‭o f homologous chromosomes exchange linear segments of genetic material (‬‭Figure‬
‭20.23‬‭). This process is called recombination or crossover‬‭and is a common genetic‬
‭process. Because the genes are aligned during recombination, the gene order is not‬
‭altered. Instead, recombination results in maternal and paternal alleles being combined‬
‭o nto the same chromosome. Across a given chromosome, several recombination events‬
‭may occur, causing extensive shuffling of alleles.‬

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‭ igure 20.23‬ ‭Crossover, or recombination, occurs‬‭when two homologous chromosomes‬

F
‭align and exchange a segment of genetic material. (credit:‬‭OpenStax‬‭)‬

‭ hen two genes are located on the same chromosome, they are considered linked, and‬
W
‭their alleles tend to be transmitted through meiosis together. To exemplify this, imagine a‬
‭dihybrid cross involving flower color and plant height in which the genes are next to each‬
‭o ther on the chromosome. Suppose one homologous chromosome has alleles for tall‬
‭plants and red flowers, and the other has genes for short plants and yellow flowers. In‬
‭that case, when the gametes are formed, the tall and red alleles will tend to go together‬
‭into a gamete, and the short and yellow alleles will go into other gametes. These are called‬
‭the parental genotypes because they have been inherited intact from the parents of the‬
‭individual-producing gametes. But unlike if the genes were on different chromosomes,‬
‭there would be no gametes with tall and yellow alleles and no gametes with short and red‬
‭alleles. If you create a Punnett square with these gametes, you will see that the classical‬
‭Mendelian prediction of a 9:3:3:1 outcome of a dihybrid cross would not apply. As the‬
‭distance between two genes increases, the probability of one or more crossovers between‬
‭them increases, and the genes behave more like they are on separate chromosomes.‬
‭Geneticists have used the proportion of recombinant gametes (the ones not like the‬
‭parents) as a measure of how far apart genes are on a chromosome. Using this‬
‭information, they have constructed‬‭linkage maps‬‭o f‬‭genes on chromosomes for‬
‭well-studied organisms, including humans.‬

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‭ or additional information about genetic linkage and its impact on dihybrid cross results,‬
F
‭click the link or scan the QR code to watch the Bozeman Science video titled “Linked‬
‭Genes.”‬

Linked Genes

‭ endel’s seminal publication does not mention linkage, and many researchers have‬
M
‭questioned whether he encountered linkage but chose not to publish those crosses out of‬
‭concern that they would invalidate his independent assortment postulate. The garden pea‬
‭has seven pairs of chromosomes, and some have suggested that his choice of seven‬
‭characteristics was not a coincidence. However, even if the genes he examined were not‬
‭located on separate chromosomes, it is possible that he did not observe linkage because‬
‭o f the extensive shuffling effects of recombination.‬

‭20.5 Pedigree Analysis‬

‭ any human diseases are genetically inherited. A healthy person in a family in which‬
M
‭some members suffer from a recessive genetic disorder may want to know if they have the‬
‭disease-causing gene and what risk exists of passing the disorder on to their offspring. Of‬
‭course, doing a test cross in humans is unethical and impractical. Instead, geneticists use‬
‭pedigree analysis‬‭to study the inheritance pattern‬‭o f human genetic diseases.‬
‭ or an introduction to pedigrees and their analysis, click the link or scan the QR code to‬
F
‭watch the Amoeba Sisters video titled “Pedigrees.”‬

Pedigrees

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‭Autosomal Recessive Inheritance‬

‭ hen a genetic disorder is inherited in an‬‭autosomal‬‭recessive‬‭pattern, the disorder‬

W
‭corresponds to the recessive phenotype. Heterozygous individuals will not display‬
‭symptoms of this disorder because their unaffected gene will compensate. Such an‬
‭individual, as mentioned previously, is called a carrier. Carriers for an autosomal recessive‬
‭disorder may never know their genotype unless they have a child with the disorder.‬

‭ n example of an autosomal recessive disorder is cystic fibrosis (CF). The chronic‬

A
‭accumulation of thick, tenacious mucus in the lungs and digestive tract characterizes CF.‬
‭Decades ago, children with CF rarely lived to adulthood. With advances in medical‬
‭technology, the average lifespan in developed countries has increased into middle‬
‭adulthood. CF is a relatively common disorder that occurs in approximately 30,000 people‬
‭in the United States. A child born to two CF carriers would have a 25 percent chance of‬
‭inheriting the disease. This is the same 3:1 dominant:recessive ratio that Mendel observed‬
‭in his pea plants would apply here. The pattern is shown in‬‭Figure 20.24‬‭, using a diagram‬
‭that tracks the likely incidence of an autosomal recessive disorder based on parental‬
‭genotypes.‬

‭ igure 20.24‬ ‭The inheritance pattern of an autosomal‬‭recessive disorder with two‬

F
‭carrier parents reflects a 3:1 probability of expression among offspring. (credit: U.S.‬
‭National Library of Medicine;‬‭OpenStax‬‭)‬

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‭ n the other hand, a child born to a CF carrier and someone with two unaffected alleles‬
O
‭would have a 0 percent probability of inheriting CF but a 50 percent chance of being a‬
‭carrier. Other examples of autosomal recessive genetic illnesses include the blood‬
‭disorder sickle-cell anemia, the fatal neurological disorder Tay–Sachs disease, and the‬
‭metabolic disorders phenylketonuria (PKU) and alkaptonuria. A human pedigree‬
‭demonstrating the inheritance of alkaptonuria is shown in‬‭Figure 20.25‬‭.‬

‭ igure 20.25‬ ‭Alkaptonuria is a recessive genetic‬‭disorder in which two amino acids,‬

F
‭phenylalanine and tyrosine, are not properly metabolized. Affected individuals may have‬
‭darkened skin and brown urine and may suffer joint damage and other complications. In‬
‭this pedigree, individuals with the disorder are indicated in blue and have the genotype aa.‬
‭Unaffected individuals are indicated in yellow and have the genotype AA or Aa. Note that it‬
‭is often possible to determine a person’s genotype from the genotype of their offspring.‬
‭For example, if neither parent has the disorder but their child does, they must be‬
‭heterozygous. Two individuals on the pedigree have an unaffected phenotype but an‬
‭unknown genotype. Because they do not have the disorder, they must have at least one‬
‭normal allele, so their genotype gets the “A?” designation. (credit:‬‭OpenStax‬‭). A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Autosomal Dominant Inheritance‬

I‭ n the case of cystic fibrosis, the disorder is recessive to the normal phenotype. However, a‬
‭genetic abnormality may be dominant to the normal phenotype. When the dominant allele‬
‭is located on one of the 22 pairs of autosomes (non-sex chromosomes), we refer to its‬

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i‭nheritance pattern as‬‭autosomal dominant‬‭. An example of an autosomal dominant‬

‭disorder is neurofibromatosis type I, a disease that induces tumor formation within the‬
‭nervous system that leads to skin and skeletal deformities. Consider a couple in which one‬
‭parent is heterozygous for this disorder (and therefore has neurofibromatosis),‬‭Nn‬‭, and‬
‭o ne parent is homozygous for the normal gene,‬‭nn‬‭.‬‭The heterozygous parent would have a‬
‭50 percent chance of passing the dominant allele for this disorder to their offspring, and‬
‭the homozygous parent would always pass the normal allele. Therefore, four possible‬
‭o ffspring genotypes are equally likely to occur:‬‭Nn‬‭,‬‭Nn‬‭,‬‭nn‬‭, and‬‭nn‬‭. That is, every child of‬
‭this couple would have a 50 percent chance of inheriting neurofibromatosis. This‬
‭inheritance pattern is shown in Figure‬‭20.26‬‭in a‬‭Punnett square.‬

‭ igure 20.26‬‭The Inheritance pattern of an autosomal‬‭dominant disorder, such as‬

F
‭neurofibromatosis, is shown in a Punnett square. (credit:‬‭OpenStax‬‭)‬

‭ ther genetic diseases inherited in this pattern are achondroplastic dwarfism, Marfan‬
O
‭syndrome, and Huntington’s disease. Because autosomal dominant disorders are‬
‭expressed by the presence of just one gene, an individual with the disorder will know that‬
‭they have at least one faulty gene. The expression of the disease may manifest later in life,‬
‭after the childbearing years, which is the case in Huntington’s disease, a neurological‬
‭condition.‬

‭X-linked Dominant or Recessive Inheritance‬

‭ n‬‭X-linked‬‭transmission pattern involves genes located‬‭o n the X chromosome of the‬

A
‭23rd pair (‬‭Figure 20.27‬‭). Recall that a male has one‬‭X and one Y chromosome. When a‬
‭male transmits a Y chromosome, the child is male, and when a male parent transmits an X‬
‭chromosome, the child is female. A female can transmit only an X chromosome, as both‬
‭her sex chromosomes are X.‬

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‭ hen an abnormal allele for a gene that occurs on the X chromosome is dominant over‬
W
‭the normal allele, the pattern is described as‬‭X-linked‬‭dominant‬‭. This is the case with‬
‭vitamin D-resistant rickets: an affected male would pass the disease gene to all of the‬
‭female offspring but none of the male offspring because the male transmits only the Y‬
‭chromosome to male offspring (see‬‭Figure 20.27a‬‭).‬‭If the female parent is affected, all of‬
‭the progeny—male or female—would have a 50 percent chance of inheriting the disorder‬
‭because the female parent can only pass an X chromosome on to children (see‬‭Figure‬
‭20.27b‬‭). For an affected female, the inheritance pattern‬‭would be identical to an‬
‭autosomal dominant inheritance pattern in which one parent is heterozygous, and the‬
‭o ther is homozygous for the normal gene.‬

‭ -linked recessive‬‭inheritance is much more common‬‭because females can be carriers of‬

X
‭the disease yet still have a normal phenotype. Diseases transmitted by X-linked recessive‬
‭inheritance include color blindness, the blood-clotting disorder hemophilia, and some‬
‭forms of muscular dystrophy. For an example of X-linked recessive inheritance, consider‬
‭parents in which the female is an unaffected carrier, and the male is normal. None of the‬
‭female offspring would have the disease because they receive a normal gene from their‬
‭male parent. However, they have a 50 percent chance of receiving the disease gene from‬
‭their female parent and becoming a carrier. In contrast, 50 percent of the male offspring‬
‭would be affected (‬‭Figure 20.28‬‭).‬

‭ ith X-linked recessive diseases, males either have the disease or are genotypically‬
W
‭normal—they cannot be carriers. Females, however, can be genotypically normal, a carrier‬
‭who is phenotypically normal, or affected by the disease. A female can inherit the gene for‬
‭an X-linked recessive illness when the female parent is a carrier or affected or the male‬
‭parent is affected. The disease will only affect female offspring if they inherit an X-linked‬
‭recessive gene from both parents. As you can imagine, X-linked recessive disorders affect‬
‭many more males than females. For example, color blindness affects at least 1 in 20 males‬
‭but only about 1 in 400 females.‬

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‭ igure 20.27‬ ‭A chart of X-linked dominant inheritance‬‭patterns differs depending on‬

F
‭which parent is affected with the disease. (a) The affected parent is the father. (b) The‬
‭affected parent is the mother. (credit: U.S. National Library of Medicine;‬‭OpenStax‬‭)‬

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‭ igure 28.28‬ ‭Given two parents in which the male‬‭is normal and the female is a carrier‬
F
‭o f an X-linked recessive disorder, a male offspring would have a 50 percent probability of‬
‭being affected with the disorder. In contrast, female offspring would either be carriers or‬
‭entirely unaffected. (credit: U.S. National Library of Medicine;‬‭OpenStax‬‭).‬ ‭A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ or information about sex-linked traits and their analysis using Punnett squares, click the‬
F
‭link or scan the QR code to watch the Amoeba Sisters video titled “Punnett Squares and‬
‭Sex-Linked Traits.”‬

Punnett Squares and Sex-Linked Traits (UPDATED)

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‭Chapter 21: Evolution and Population Genetics‬

‭ igure 21.1‬ ‭Living things may be single-celled or‬‭complex, multicellular organisms. They‬
F
‭may be plants, animals, fungi, bacteria, or archaea. This diversity results from evolution.‬
‭(credit "wolf": modification of work by Gary Kramer; credit "coral": modification of work‬
‭by William Harrigan, NOAA; credit "river": modification of work by Vojtě ch Dostá l; credit‬
‭"fish" modification of work by Christian Mehlfü hrer; credit "mushroom": modification of‬
‭work by Cory Zanker; credit "tree": modification of work by Joseph Kranak; credit "bee":‬
‭modification of work by Cory Zanker;‬ ‭OpenStax‬‭)‬

‭21.1 Introduction‬
‭ volutionary theory states that all organisms share a common ancestor but that millions‬
E
‭o f years of‬‭evolution‬‭(i.e., genetic changes to populations‬‭over time) have resulted in the‬
‭incredible diversity of organisms we see today.‬
‭ wo factors are fundamentally responsible for much of the observed variation in‬
T
‭o rganisms. The first is‬‭natural selection‬‭, which‬‭promotes traits and behaviors that‬
‭increase an organism’s chances of survival and reproduction while eliminating those traits‬
‭and behaviors that are detrimental to the organism. However, as its name implies, natural‬
‭selection can only select—it cannot create. Novel traits and behaviors can be attributed to‬
‭o ther factors, such as‬‭mutations‬‭, which are heritable‬‭changes in the DNA. Mutation and‬
‭o ther sources of variation among individuals and the evolutionary forces that act upon‬
‭them alter species (individuals that can interbreed to produce offspring) and populations‬

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(‭ members of the same species that live in the same area simultaneously). This‬
‭combination of processes has led to the world of life we see today.‬
I‭ n this chapter, you will learn about the history of evolution theory, the factors associated‬
‭with it, and a brief introduction to human evolution. Also, the impact of evolution on the‬
‭genetics of populations is discussed.‬

‭21.2 A Brief History of the Theory of Evolution by Natural Selection‬

‭ volution by natural selection‬‭describes a mechanism‬‭for how species change over time.‬
E
‭Scientists, philosophers, researchers, and others had made suggestions and debated this‬
‭topic well before Charles Darwin and others began to explore this idea. Classical Greek‬
‭philosopher Plato emphasized in his writings that species were static and unchanging, yet‬
‭there were also ancient Greeks who expressed evolutionary ideas. In the eighteenth‬
‭century, naturalist Georges-Louis Leclerc Comte de Buffon reintroduced ideas about the‬
‭evolution of animals and observed that various geographic regions have different plant‬
‭and animal populations, even when the environments are similar. Some at this time also‬
‭accepted that there were extinct species.‬
‭ lso during the eighteenth century, James Hutton, a Scottish geologist and naturalist,‬
A
‭proposed that geological change occurred gradually by accumulating small changes from‬
‭processes operating like they are today over long periods of time. This contrasted with the‬
‭predominant view that the planet's geology resulted from catastrophic events occurring‬
‭during a relatively brief past. Nineteenth-century geologist Charles Lyell popularized‬
‭Hutton's view and promoted the idea of the greater age of Earth, which gave more time‬
‭for gradual change in species.‬
‭ he mechanism for evolution was independently conceived of and described by two‬
T
‭naturalists,‬‭Charles Darwin‬‭and‬‭Alfred Russell Wallace‬‭,‬‭in the mid-nineteenth century.‬
‭Importantly, each explored the natural world on expeditions to the tropics. From 1831 to‬
‭1836, Darwin traveled the world on‬‭H.M.S. Beagle‬‭,‬‭visiting South America, Australia, and‬
‭the southern tip of Africa. Wallace traveled to Brazil to collect insects in the Amazon‬
‭rainforest from 1848 to 1852 and the Malay Archipelago from 1854 to 1862. Darwin’s‬
‭journey, like Wallace’s later journeys in the Malay Archipelago, included stops at several‬
‭island chains, the last being the Galá pagos Islands (west of Ecuador). Darwin observed‬
‭species of organisms on these islands that were similar yet had distinct differences. For‬
‭example, the ground finches inhabiting the Galá pagos Islands comprised several species‬
‭that each had a unique beak shape (‬‭Figure 21.2‬‭). He‬‭o bserved that these finches closely‬
‭resembled other finch species on the mainland of South America and that the species in‬
‭the Galá pagos formed a graded series of beak sizes and shapes, with minimal differences‬

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‭ etween the most similar. Darwin imagined the island species might be modified from one‬
b
‭o riginal mainland species.‬

‭ igure 21.2‬ ‭Darwin observed that beak shape varies‬‭among finch species. He postulated‬
F
‭that the beak of an ancestral species had adapted over time to equip the finches to acquire‬
‭different food sources. This illustration shows the beak shapes of four ground finch‬
‭species: 1.‬‭Geospiza magnirostris‬‭(the large ground‬‭finch), 2.‬‭G. fortis‬‭(the medium ground‬
‭finch), 3.‬‭G. parvula‬‭(the small tree finch), and‬‭4.‬‭Certhidea olivacea‬‭(the green-warbler‬
‭finch). (credit:‬‭OpenStax‬‭)‬

‭ allace and Darwin observed similar patterns in other organisms and independently‬
W
‭conceived a mechanism to explain how and why such changes could occur. Darwin called‬
‭this mechanism‬‭natural selection‬‭. Natural selection,‬‭Darwin argued, was an inevitable‬
‭o utcome of three principles that operated in nature. First, the characteristics of organisms‬
‭are inherited or passed from parent to offspring. Second, more offspring are produced‬
‭than can survive; in other words, resources for survival and reproduction are limited. The‬
‭capacity for reproduction in all organisms outstrips the availability of resources to‬
‭support their numbers. Thus, there is competition for those resources in each generation.‬
‭Darwin and Wallace’s understanding of this principle came from reading an essay by the‬
‭economist Thomas Malthus, who discussed this principle concerning human populations.‬
‭Third, offspring vary among each other in regard to their characteristics, and those‬

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‭ ariations are inherited. Out of these three principles, Darwin and Wallace reasoned that‬
v
‭o ffspring with inherited characteristics that allow them to best compete for limited‬
‭resources will survive and have more offspring than those individuals with variations that‬
‭are less able to compete. Because characteristics are inherited, these traits will be better‬
‭represented in the next generation. This will lead to population changes over generations,‬
‭which Darwin called “descent with modification.”‬
‭ apers by Darwin and Wallace (‬‭Figure 21.3‬‭) presenting‬‭the idea of natural selection were‬
P
‭read together in 1858 before the Linnaean Society in London. The following year,‬
‭Darwin’s book,‬‭On the Origin of Species,‬‭was published, outlining his arguments for‬
‭evolution by natural selection in considerable detail.‬

‭ igure 21.3‬ ‭(a) Charles Darwin and (b) Alfred Wallace‬‭wrote scientific papers on‬
F
‭natural selection that were presented together before the Linnean Society in 1858.‬
‭(credit:‬‭OpenStax‬‭)‬

‭21.3 Understanding Darwin’s Theory of Natural Selection‬

‭The theory of natural selection has five main components:‬
‭1.‬ ‭ ll organisms are capable of producing offspring faster than the food supply‬
A
‭increases.‬
‭2.‬ ‭All organisms show variation.‬

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‭3.‬ ‭ here is a fierce struggle for existence, and those with the most suitable variations‬
T
‭are most likely to survive and reproduce.‬
‭ .‬
4 ‭Variations, or traits, are passed on to offspring (inherited).‬
‭5.‬ ‭Small changes in every generation lead to significant changes over long periods of‬
‭time.‬
‭ popular but often misunderstood concept related to natural selection is the term‬
A
‭survival of the fittest‬‭. Survival of the fittest does‬‭not necessarily mean that the biggest‬
‭and fastest survive; instead, it refers to the most‬‭evolutionarily‬‭fit. This means that an‬
‭o rganism has traits that are sufficient for survival and will be passed on to future‬
‭generations. Darwin did not introduce the term survival of the fittest; it was first used by‬
‭English philosopher, anthropologist, and sociologist Herbert Spencer.‬
‭ xamples of Darwin’s theory of natural selection can be found throughout the natural‬
E
‭world. Perhaps one of the best-known is the color change observed in peppered moths in‬
‭England during the 19th century (‬‭Figure 21.4‬‭). Before the Industrial Revolution,‬
‭peppered moths in England were a light gray color, well camouflaged on tree branches,‬
‭and less likely to be eaten by birds. Occasionally, through the process of mutation, black‬
‭moths would appear in the population, but these were usually quickly eaten because they‬
‭were more visible against light-colored bark.‬

‭ igure 21.4‬ ‭This peppered moth is well camouflaged on the trunk of this tree. A‬
F
‭darker-colored moth would more easily be seen and eaten, thus less likely to pass on its‬
‭genes to offspring. Natural selection relies upon the ability of natural variations to‬
‭increase an individual’s chances of reproduction. (credit: Ben Sale/Wikimedia Commons;‬
‭CC BY 2.0‬‭;‬ ‭OpenStax‬‭)‬

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‭ hen soot from coal factories began to cover the bark of the trees, the black moths‬
W
‭became better camouflaged, and the white moths were now more visible. Consequently,‬
‭the black moths survived to reproduce while the white ones were eaten. In a few decades,‬
‭all the peppered moths in the cities were black. The process was termed industrial‬
‭melanism. As coal usage decreased and the bark of the trees once again became lighter in‬
‭color, white moths again dominated the urban areas.‬
‭ here are numerous examples of natural selection in modern times. One of the best‬
T
‭demonstrations has been in the very birds that helped to inspire the theory: the‬
‭Galá pagos finches. Peter and Rosemary Grant and their colleagues have studied Galá pagos‬
‭finch populations every year since 1976 and have provided important demonstrations of‬
‭the operation of natural selection. The Grants found changes from generation to‬
‭generation in the beak shapes of the medium ground finches on the island of Daphne‬
‭Major.‬
‭ he medium-ground finch feeds on seeds. The birds have inherited variation in the bill‬
T
‭shape, with some individuals having wide, deep bills and others having thinner bills.‬
‭Large-billed birds feed more efficiently on large, hard seeds, whereas smaller-billed birds‬
‭feed more efficiently on small, soft seeds. In 1977, a drought period altered vegetation on‬
‭the island. After this period, the number of seeds declined dramatically: the decline in‬
‭small, soft seeds was more significant than in large, hard seeds. The large-billed birds‬
‭survived better than the small-billed birds the following year. The year following the‬
‭drought, when the Grants measured beak sizes in the much-reduced population, they‬
‭found that the average bill size was larger (‬‭Figure 21.5‬‭). This was clear evidence for‬
‭natural selection (differences in survival) of bill size caused by the availability of seeds.‬
‭The Grants had studied the inheritance of bill sizes and knew that the surviving‬
‭large-billed birds would tend to produce offspring with larger bills, so the selection would‬
‭lead to the evolution of bill size. Subsequent studies by the Grants have demonstrated the‬
‭selection on and evolution of bill size in this species in response to changing conditions on‬
‭the island. The evolution has occurred to larger bills, as in this case, and smaller bills when‬
‭large seeds became rare.‬
‭ nother example of natural selection in modern times includes pesticide resistance in‬
A
‭insects. Pesticide resistance refers to the decreasing susceptibility of a pest population to a‬
‭pesticide that previously was effective at controlling it. Pest species evolve pesticide‬
‭resistance via natural selection, with the most resistant individuals surviving to pass on‬
‭their ability to resist the pesticide to their offspring.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 21.5‬ ‭A drought on the Galá pagos island of Daphne Major in 1977 reduced the‬
F
‭number of small seeds available to finches, causing many small-beaked finches to die. This‬
‭caused an increase in the finches’ average beak size between 1976 and 1978. (credit:‬
‭OpenStax‬‭)‬

‭21.4 The Processes of Evolution‬

‭ atural selection can only occur if there is variation or differences among individuals in a‬
N
‭population. Importantly, these differences must have some genetic basis; otherwise,‬
‭selection will not lead to change in the next generation. This is critical because variation‬
‭among individuals can be caused by non-genetic reasons, such as an individual being taller‬
‭because of better nutrition rather than different genes.‬
‭ n adaptation is a heritable trait that aids an organism's survival and reproduction in its‬
A
‭present environment. It is the organism's “match” to the environment. Adaptation to an‬
‭environment occurs when a change in the range of genetic variation occurs over time that‬
‭increases or maintains the population's match with its environment. The variations in‬
‭finch beaks shifted from generation to generation, providing adaptation to food‬
‭availability.‬
‭ enetic diversity in a population can arise from sources such as mutation, meiosis, genetic‬
G
‭drift, and gene flow.‬

‭Mutation‬
‭ utation is the creative force of evolution and represents the first stage of the‬
M
‭evolutionary process. As stated previously, Mutations are defined as alterations in genetic‬
‭sequences that result in variant forms. For a mutation to have evolutionary significance, it‬

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‭ ust occur in the‬‭gametes‬‭(sperm and ova). This is because only genetic information in‬
m
‭the gametes is passed on from generation to generation. Mutations in non-sex cells will‬
‭not be passed on from generation to generation. Whereas other evolutionary forces can‬
‭modify existing genetic material, only mutation can produce new genetic material.‬
‭ ne of the most interesting things about mutations is that they are mainly random. There‬
O
‭is no way of predicting when a specific mutation will occur; all scientists can do is estimate‬
‭the probability of a mutation occurring. Mutations do not necessarily appear when they‬
‭are needed.‬
‭ he conventional view is that mutations are harmful, but this is not always true. Some‬
T
‭mutations are harmful, some are advantageous, and some are neutral.‬ ‭Advantageous‬
‭mutations‬‭lead to changes that improve an individual’s‬‭survival and/or chances of‬
‭reproduction. The mutation that confers resistance to insecticide in mosquitos led to‬
‭changes that improved their survival. Likewise, the mutation for black coloration in‬
‭peppered moths led to increased survival during the Industrial Revolution.‬‭Neutral‬
‭mutations‬‭do not affect survival or reproduction.‬ ‭Deleterious mutation‬‭s are very‬
‭harmful and negatively impact individuals’ survival and reproduction.‬
‭ utations generally occur spontaneously in response to body or environment conditions.‬
M
‭The exact cause of a mutation cannot usually be determined, and the mutation rate is‬
‭challenging to determine. This is because mutations that are neutral or do not lead to‬
‭apparent changes often go unnoticed. The probability of a mutation at any given gene is‬
‭between 1 in 10,000 and 1 in 100,000. While the probability that a specific point in an‬
‭individual’s genetic material will have a mutation is very low, the probability that the‬
‭totality of an individual’s genetic material will have at least one mutation is much higher.‬
‭The point is that while rare, mutation is not uncommon. For example, although many‬
‭mosquitoes have adapted to insecticides through a mutation that confers some resistance‬
‭to the chemicals if the mutation had not already been present in the population, the‬
‭mosquitoes would have died out. The need for a specific mutation did not affect whether‬
‭the mutation appeared.‬
‭ lorida has a controversial pilot program to deal with mosquitoes against which‬
F
‭insecticide sprays have increasingly become ineffective. The first genetically modified‬
‭mosquitoes were released in the Florida Keys in May 2021 (‬‭Figure 21.6‬‭). The genetically‬
‭altered mosquitoes produce female offspring that die in the larval stage, preventing them‬
‭from growing to adulthood, in which they can then bite and spread disease. Genetic‬
‭science currently can use mutations to control or even wipe out an entire species. Genetic‬
‭engineering has the potential to benefit humanity, but it will undoubtedly also raise ethical‬
‭questions and controversy.‬

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‭ igure 21.6‬ ‭Genetically modified mosquitoes that‬‭will die in the larval stage are currently‬
F
‭being bred, thus greatly reducing the mosquito population. (attribution: Rice University,‬
‭OpenStax‬‭; CC BY 4.0). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭Meiosis‬
‭ eiosis‬‭is the type of cell division that results in gametes, or haploid cells. As discussed in‬
M
‭Chapter 13, two events associated with meiosis can result in genetic diversity: crossing‬
‭over, which occurs during prophase I, and random alignment of the homologous‬
‭chromosome pairs on the metaphase I plate.‬
‭ rossing over‬‭is the exchange of chromosomal segments‬‭between homologous, non-sister‬
C
‭chromatids (‬‭Figure 21.7‬‭). As long as the segments‬‭contain different alleles of the genes,‬
‭the gametes will contain new combinations.‬
‭ he second mechanism that introduces variation in the gametes is the random or‬
T
‭independent assortment‬‭o f homologous chromosomes at‬‭the metaphase plate (‬‭Figure‬
‭21.8‬‭). There are two possibilities for the orientation‬‭o f the tetrad at the metaphase plate‬
‭(the maternal homolog to the left or the right). Therefore, the possible number of‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ ifferent alignments equals 2‬‭n‬ ‭in a diploid cell, where‬‭n‬‭is the number of chromosomes‬
d
‭per haploid set.‬

‭ igure 21.7‬ ‭The result of crossing over is an exchange‬‭o f genetic material between‬
F
‭non-sister chromatids of a pair of homologous chromosomes. Assume there are three‬
‭genes on each homolog, and the first one has dominant alleles at all three (A, B, and C),‬
‭while the second one has recessive alleles at all three (a, b, and c). Crossover occurs‬
‭between non-sister chromatids of homologous chromosomes‬‭.‬‭In the figure above,‬
‭crossing over occurred one time between the B/b and C/c genes. Because there were‬
‭allelic differences between the homologs, crossing over produced two recombinant‬
‭chromatids with allelic content ABc and abC. The two non-sister chromatids that didn’t‬
‭participate in crossing over are classified as nonrecombinant and have the same allelic‬
‭collection for the three genes as the parental homologs (ABC and abc). (credit:‬‭OpenStax‬‭).‬
‭A link to a video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 21.8‬ ‭Random, independent assortment during metaphase I can be demonstrated‬

F
‭by considering a cell with a set of two chromosomes (2n = 4;‬‭n‬‭= 2). In this case,‬
‭metaphase I has two possible arrangements at the equator. The total possible number of‬
‭different gametes is 2‬‭n‭,‬ where‬‭n‬‭equals the number‬‭o f chromosomes in a set. In this‬
‭example, there are four possible genetic combinations for the gametes. With‬‭n‬‭= 23 in‬
‭human cells, there are over eight million possible paternal and maternal chromosome‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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c‭ ombinations. (credit:‬‭OpenStax‬‭). A link to a video explanation of this figure is available at‬

‭Biology411.com‬‭.‬
‭ umans have 23 chromosome pairs, resulting in over eight million (2‬‭23‬‭) possible‬
H
‭genetically distinct gametes from the random alignment of chromosomes at the metaphase‬
‭plate. This number does not include the variability previously produced by crossing over‬
‭between the non-sister chromatids. Given these two mechanisms, it is improbable that any‬
‭two haploid cells resulting from meiosis will have the same genetic composition.‬
‭Therefore, when two parents reproduce, unique combinations of alleles assemble to‬
‭produce unique genotypes and, thus, phenotypes in each offspring.‬

‭Genetic Drift‬
‭ enetic drift‬‭is defined as the effect of random chance‬‭o n a population, notably how it‬
G
‭determines whether an individual survives and reproduces or dies. Imagine that you stick‬
‭your hand into a bucket filled with different brands of candy bars. What is the probability‬
‭you will withdraw a Snickers bar? The composition of candy bars in your bucket will be‬
‭affected by the proportion of Snickers bars compared to other brands. If each bucket of‬
‭candy bars were a population, then one could say that genetic drift—random‬
‭chance—affected the candy composition in your bucket.‬
‭ s discussed previously (Chapter 7),‬‭alleles‬‭are different‬‭forms of a gene.‬ ‭Allele‬
A
‭frequency‬‭is the number of copies of a particular‬‭allele divided by the total number of‬
‭alleles in a population. Remember, each individual possesses two alleles for each gene, so‬
‭the total number of alleles is calculated by doubling the number of individuals in the‬
‭population.‬ ‭Figure 21.9‬‭shows genetic drift in a‬‭small, hypothetical rabbit population.‬
‭The brown color allele (‬‭B‭)‬ is dominant to the white‬‭allele (‬‭b‬‭), so individuals with‬‭BB‬‭o r‬‭Bb‬
‭genotypes are brown, and individuals with a‬‭bb‬‭genotype‬‭are white. Notice how the allele‬
‭frequencies change over successive generations simply because of which individuals mate‬
‭and how many progeny they produce.‬
‭ n important point about genetic drift is that it is directly and inversely related to‬
A
‭population size. Small populations are more susceptible to the forces of genetic drift.‬
‭Large populations, alternatively, are buffered against the effects of chance. If one‬
‭individual of a population of 10 individuals happens to die at a young age before it leaves‬
‭any offspring to the next generation, all of its genes—1/10 of the population’s gene‬
‭pool—will be suddenly lost. However, in a population of 100, that’s only 1 percent of the‬
‭overall gene pool; therefore, it is much less impactful on the population’s genetic‬
‭structure. However, in early human evolution, population sizes were small, so the effect of‬
‭genetic drift may have been substantial.‬

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‭ igure 21.9‬ ‭Genetic drift occurs when the gene frequency‬‭o f a population shifts by‬
F
‭random chance (i.e., without selective pressure). Over time, genetic drift can eliminate an‬
‭allele from the population. For example, the two alleles B and b occur with equal‬
‭frequency in the first generation, so p = q = 0.5. If only half the individuals reproduce, and‬
‭by chance, most of the reproducing alleles are of B, then the second generation results in‬
‭p = 0.7 and q = 0.3. In the second generation, only two individuals, both of whom are‬
‭homozygous for the B allele, reproduce. This leads to a loss of b from the third‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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g‭ eneration. (credit:‬ ‭OpenStax‬‭). A link to a video explanation of this figure is available at‬
‭Biology411.com‬‭.‬

‭Gene Flow‬
‭ ene flow‬‭is another important evolutionary force‬‭involving genetic material exchange‬
G
‭between populations and geographic regions. Without gene flow, there would be‬
‭diminished diversity—and without diversity, a species is at higher risk of extinction. Gene‬
‭flow can be seen in the process of pollination, in which bees or butterflies carry and‬
‭transfer pollen from one area to another (‬‭Figure 21.10‬‭).‬ ‭Gene flow also results from the‬
‭movement of individuals of the same species between populations.‬

‭ igure 21.10‬ ‭The process of pollination is an excellent‬‭example of gene flow. In this case,‬
F
‭bees and butterflies transfer genetic material, pollen, from one flower to another. (credit:‬
‭“‬‭Honey Bee on a Dandelion, Sandy, Bedfordshire”‬‭by‬‭Orangeaurochs/flickr, CC BY 2.0)‬

‭21.5 Evidence of Evolution‬

‭ he evidence for evolution is compelling and extensive. Biologists see the signature of past‬
T
‭and present evolution in every organization in living systems. Darwin dedicated a large‬
‭portion of his book,‬‭On the‬‭Origin of Species‬‭, to‬‭identifying patterns in nature that were‬
‭consistent with evolution. Since Darwin's time, our understanding has become clearer and‬
‭broader.‬

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‭Fossils‬
‭ ossils provide solid evidence that organisms from the past are not the same as those‬
F
‭today, and fossils show a progression of evolution. Scientists determine the age of fossils‬
‭and categorize them worldwide to determine when the organisms lived relative to each‬
‭o ther. The resulting fossil record tells the past story and shows the evolution of form over‬
‭millions of years. For example, scientists have recovered highly detailed records showing‬
‭the evolution of humans and horses (‬‭Figure 21.11‬‭).‬

‭ igure 21.11‬ ‭In this (a) display, fossil hominids‬‭are arranged from oldest (bottom) to‬
F
‭newest (top). As hominids evolved, the skull's shape changed. An artist’s rendition of (b)‬
‭extinct species of the genus‬‭Equus‬‭reveals that these‬‭ancient species resembled the‬
‭modern horse (‬‭Equus ferus‬‭) but varied in size. (credit:‬‭OpenStax‬‭)‬

‭Anatomy‬
‭ nother type of evidence for evolution is the presence of structures in organisms that‬
A
‭share the same basic form. For example, human, dog, bird, and whale appendages' bones‬
‭share the same overall construction (‬‭Figure 21.12‬‭),‬‭resulting from their origin in a‬
‭common ancestor's appendages. Over time, evolution led to changes in the bones' shapes‬
‭and sizes of different species, but they have maintained the same overall layout. Scientists‬
‭call these synonymous parts‬‭homologous structures‬‭.‬ ‭The wings of a bat and a bird are‬
‭another example of homologous structures. They did not evolve independently in each‬
‭lineage; instead, they descended from a common ancestor with wings.‬

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‭ igure 21.12‬ ‭The similar construction of these appendages‬‭indicates that these‬

F
‭o rganisms share a common ancestor. (credit:‬‭OpenStax‬‭)‬

I‭ t should be mentioned that similar phenotypes sometimes evolve independently in‬

‭distantly related species. For example, flight has evolved in both bats and insects, and they‬
‭both have structures we refer to as wings, which are adaptations to flight. However, the‬
‭wings of bats and insects evolved from very different original structures and are called‬
‭analogous structures‬‭; they are similar in function‬‭and appearance but do not share an‬
‭o rigin in a common ancestor. Instead, they evolved independently in the two lineages‬
‭(Figure 21.13)‬‭.‬
S‭ ome structures exist in organisms with no apparent function and appear to be residual‬
‭parts from a past common ancestor. These unused structures without function are called‬
‭vestigial structures‬‭. Other vestigial structures include‬‭wings on flightless birds, leaves on‬
‭some cacti, and hind leg bones in whales.‬
‭ nother example of evidence of evolution is the convergence of form in organisms that‬
A
‭share similar environments. For example, species of unrelated animals, such as the arctic‬
‭fox and ptarmigan, living in the Arctic region have been selected for seasonal white‬
‭phenotypes during winter to blend with the snow and ice (‬‭Figure 21.14‬‭). These‬
‭similarities occur not because of common ancestry but because of similar selection‬
‭pressures—the benefits of predators not seeing them.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 21.13‬ ‭A honey bee's wing is similar in shape‬‭to a bird wing and a bat wing and‬
F
‭serves the same function (flight). The bird and bat wings are homologous structures.‬
‭However, the honey bee wing has a different structure (it is made of a chitinous‬
‭exoskeleton, not a bony endoskeleton) and embryonic origin. The bee and bird or bat‬
‭wing types illustrate an analogy—similar structures that do not share an evolutionary‬
‭history. (credit a photo: modification of work by U.S. BLM; credit b: modification of work‬
‭by Steve Hillebrand, USFWS; credit c: modification of work by Jon Sullivan;‬ ‭OpenStax‬‭)‬

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‭ igure 21.14‬ ‭The white winter coat of the (a) arctic‬‭fox and the (b) ptarmigan’s plumage‬
F
‭are adaptations to their environments. (credit a: modification of work by Keith‬
‭Morehouse;‬ ‭OpenStax‬‭)‬

‭Embryology‬
‭ mbryology‬‭, the study of the anatomy of an organism's‬‭development to its adult form,‬
E
‭also provides evidence of relatedness between now widely divergent groups of organisms‬
‭(‬‭Figure 21.15‬‭). Mutational tweaking in the embryo‬‭can have magnified consequences in‬
‭the adult, which tends to conserve embryo formation. As a result, absent structures in‬
‭some groups often appear in their embryonic forms and disappear when they reach the‬
‭adult or juvenile form. For example, all vertebrate embryos, including humans, exhibit gill‬
‭slits and tails at some point in their early development. These disappear in the adults of‬
‭terrestrial groups, but adult forms of aquatic groups, such as fish and some amphibians,‬
‭maintain them. Great ape embryos, including humans, have a tail structure during their‬
‭development that they lose when they are born.‬

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‭ igure 21.15‬ ‭Figure demonstrating physical similarities‬‭between different species during‬

F
‭early embryonic development. (credit:‬ ‭Romanes,‬‭G. J.‬ ‭Haeckel Drawings‬‭; Wikimedia‬
‭Commons;‬‭CC0 1.0‬‭)‬

‭Biogeography‬
‭ he geographic distribution of organisms on the planet follows patterns that we can‬
T
‭explain best by evolution in conjunction with tectonic plate movement over geological‬
‭time. Broad groups that evolved before the supercontinent Pangaea broke up (about 200‬
‭million years ago) are distributed worldwide. Groups that evolved since the breakup‬
‭appear uniquely in regions of the planet, such as the unique plants and animals of the‬

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‭ orthern continents that formed from the supercontinent Laurasia and of the southern‬
n
‭continents that formed from the supercontinent Gondwana. The presence of the plant‬
‭family Proteaceae members in Australia, southern Africa, and South America was most‬
‭predominant prior to the southern supercontinent Gondwana breaking up.‬
‭ arsupial diversification in Australia and the absence of other mammals reflect Australia’s‬
M
‭long isolation. Australia has an abundance of endemic species—species found nowhere‬
‭else—which is typical of islands whose isolation by expanses of water prevents species‬
‭from migrating. Over time, these species diverge evolutionarily into new species that look‬
‭very different from their ancestors who may have existed on the mainland. Australia's‬
‭marsupials, the Galá pagos' finches, and many species on the Hawaiian Islands are unique‬
‭to their one point of origin, yet they display distant relationships to ancestral species on‬
‭mainlands.‬

‭Molecular Biology‬
‭ ike anatomical structures, the molecular structures of life reflect descent with‬
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‭modification. DNA's universality reflects evidence of a common ancestor for all of life.‬
‭Fundamental divisions in life between the genetic code, DNA replication, and expression‬
‭are reflected in major structural differences in otherwise conservative structures such as‬
‭ribosome components and membrane structures. Generally, the relatedness of groups of‬
‭o rganisms is reflected in the similarity of their DNA sequences—exactly the pattern we‬
‭would expect from descent and diversification from a common ancestor. Organisms with‬
‭similar physical features‬‭and‬‭genetic sequences tend‬‭to be more closely related than those‬
‭without.‬
‭ NA sequences have also shed light on some of the mechanisms of evolution. For‬
D
‭example, it is clear that the evolution of new functions for proteins commonly occurs after‬
‭gene duplication events that allow freely modifying one copy by mutation, selection, or‬
‭drift (changes in a population’s gene pool resulting from chance). In contrast, the second‬
‭copy continues to produce a functional protein.‬

‭21.6 Misconceptions of Evolution‬

‭ lthough the theory of evolution generated some controversy when Darwin first‬
A
‭proposed it, biologists widely accepted it within 20 years after the publication of‬‭On the‬
‭Origin of Species‬‭. Nevertheless, the theory of evolution‬‭is a difficult concept, and‬
‭misconceptions about how it works abound. Here are several common misconceptions:‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Evolution Is Just a Theory‬

‭ ritics of the theory of evolution dismiss its importance by purposefully confounding the‬
C
‭everyday usage of the word “theory” with how scientists use it. In science, we understand‬
‭a “theory” to be a body of thoroughly tested and verified explanations for a set of‬
‭o bservations of the natural world. Scientists have a theory of the atom, a theory of gravity,‬
‭and the theory of relativity, each of which describes understood facts about the world. In‬
‭the same way, the theory of evolution describes facts about the living world. As such, a‬
‭theory in science has survived significant efforts to discredit it by scientists. In contrast, a‬
‭“theory” in common vernacular means a guess or suggested explanation. This meaning is‬
‭more akin to the scientific concept of “hypothesis.” When critics of evolution say it is “just‬
‭a theory,” they are implying that there is little evidence supporting it and that it is still in‬
‭the process of rigorous testing. This is a mischaracterization.‬

‭Individuals Evolve‬
‭ volution is the change in a population's genetic composition over time, specifically over‬
E
‭generations, resulting from the differential reproduction of individuals with certain alleles.‬
‭Individuals do change over their lifetime, but this is development and involves changes‬
‭programmed by the genes the individual acquired at birth in coordination with the‬
‭individual’s environment. When thinking about the evolution of a characteristic, it is‬
‭probably best to think about the change in the average value of the characteristic in the‬
‭population over time. For example, when natural selection leads to bill-size changes in‬
‭medium-ground finches in the Galá pagos, this does not mean that individual bills on the‬
‭finches are changing. If one measures the average bill size among all individuals in the‬
‭population at one time and then measures them in the population several years later, this‬
‭average value will be different due to evolution. Although some individuals may survive‬
‭from the first time to the second, they will still have the same bill size; however, many new‬
‭individuals will contribute to the shift in average bill size.‬

‭Evolution Explains the Origin of Life‬

I‭ t is a common misunderstanding that evolution includes an explanation of life’s origins.‬
‭Conversely, some of the theory’s critics believe it cannot explain the origin of life. The‬
‭theory does not try to explain the origin of life. The theory of evolution explains how‬
‭populations change over time and how life diversifies the origin of species. It does not‬
‭shed light on life's beginnings, including the origins of the first cells that define life.‬
‭Importantly, biologists believe that life on Earth precludes the possibility that the events‬
‭that led to life on Earth can repeat themselves because the intermediate stages would‬
‭immediately become food for existing living things.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ owever, once a mechanism of inheritance was in place in the form of a molecule like DNA‬
H
‭either within a cell or pre-cell, these entities would be subject to the principle of natural‬
‭selection. More effective reproducers would increase in frequency at the expense of‬
‭inefficient reproducers. While evolution does not explain the origin of life, it may have‬
‭something to say about some of the processes operating once pre-living entities acquired‬
‭certain properties.‬

‭Organisms Evolve on Purpose‬

S‭ tatements such as “organisms evolve in response to a change in an environment” are‬
‭quite common, but such statements can lead to two types of misunderstandings. First, do‬
‭not interpret the statement to mean that individual organisms evolve. The statement is‬
‭shorthand for “a population evolves in response to a changing environment.” However, a‬
‭second misunderstanding may arise when interpreting the statement to mean that the‬
‭evolution is somehow intentional. A changed environment results in some individuals in‬
‭the population, those with particular phenotypes, benefiting and producing‬
‭proportionately more offspring than other phenotypes. This results in changes in the‬
‭population if the characteristics are genetically determined.‬
I‭ t is also important to understand that the variation that natural selection works on is‬
‭already in a population and does not arise in response to an environmental change. For‬
‭example, applying antibiotics to a population of bacteria will, over time, select a population‬
‭resistant to antibiotics. The resistance a gene causes did not arise from mutation because‬
‭o f the application of the antibiotic. The gene for resistance was already present in the‬
‭bacteria's gene pool, likely at a low frequency. The antibiotic, which kills the bacterial cells‬
‭without the resistance gene, strongly selects resistant individuals since these would be the‬
‭o nly ones that survive and divide. Experiments have demonstrated that mutations for‬
‭antibiotic resistance do not arise due to antibiotic use.‬
I‭ n a larger sense, evolution is not goal-directed. Species do not become “better” over time.‬
‭They simply track their changing environment with adaptations that maximize their‬
‭reproduction in a particular environment at a particular time. Evolution has no goal of‬
‭making faster, bigger, more complex, or even smarter species, despite the commonness of‬
‭this kind of language in popular discourse. The characteristics of a species are a function‬
‭o f the variation present and the environment, which constantly changes non-directionally.‬
‭A trait that fits in one environment at one time may be fatal at some point.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ or additional information about evolution and misconceptions, click the links or scan the‬
F
‭QR codes to watch the TED-Ed videos by Prosanta Chakrabarty (“Four billion years of‬
‭evolution in six minutes”) and Alex Gendler (Myths and misconceptions about evolution).‬

Four billion years of evolution in six minutes | Prosanta Ch…

Myths and misconceptions about evolution - Alex Gendler

‭21.7 Taxonomy, Phylogeny, and Speciation‬

‭ efore we examine human evolution briefly, it will be helpful to discuss three associated‬
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‭foundational concepts: classification (taxonomy), phylogenetic trees, and speciation.‬

‭Taxonomy‬
‭ axonomy‬‭(which literally means “arrangement law”)‬‭is the science of naming and‬
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‭grouping species to construct an internationally shared classification system. The‬
‭taxonomic classification system (also called the Linnaean system after its inventor, Carl‬
‭Linnaeus, a Swedish naturalist) uses a hierarchical model. A hierarchical system has levels,‬
‭and each group at one of the levels includes groups at the next lowest level, so each‬
‭member belongs to a series of nested groups at the lowest level. An analogy is the nested‬
‭series of directories on the main disk drive of a computer. For example, in the most‬
‭inclusive grouping, scientists divide organisms into three‬‭domains‬‭: Bacteria, Archaea, and‬
‭Eukarya. Within each domain is a second level called a‬‭kingdom‬‭. Each domain contains‬
‭several kingdoms. Within kingdoms, the subsequent categories of increasing specificity are‬
‭phylum‬‭,‬‭c lass‬‭,‬‭order‬‭,‬‭family‬‭,‬‭genus‬‭, and‬‭species‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ or example, the domestic dog's classification levels are shown in‬‭Figure 21.16‬‭. The group‬
F
‭at each level is called a taxon (plural: taxa). In other words, for the dog, Carnivora is the‬
‭taxon at the order level, Canidae is the taxon at the family level, and so forth. Organisms‬
‭also have a common name that people typically use, such as domestic dogs or wolves.‬
‭Each taxon name is capitalized except for species; the genus and species names are‬
‭italicized. Scientists refer to an organism by its genus and species names, commonly called‬
‭a‬‭scientific‬‭o r Latin name. This two-name system is‬‭called‬‭binomial nomenclature‬‭. The‬
‭scientific name of the wolf is therefore‬‭Canis lupus‬‭.‬‭Recent studies of the DNA of domestic‬
‭dogs and wolves suggest that the domestic dog is a subspecies of the wolf, not its own‬
‭species; thus, it is given an extra name to indicate its subspecies status,‬‭Canis lupus‬
‭familiaris‬‭.‬
‭ igure 21.16‬‭also shows how taxonomic levels move‬‭toward specificity. Notice how, within‬
F
‭the domain, the dog is grouped with the broadest diversity of organisms. These include‬
‭plants and other organisms not pictured, such as fungi and protists. The organisms‬
‭become more similar at each sublevel because they are more closely related. Before‬
‭Darwin’s theory of evolution was developed, naturalists sometimes classified organisms‬
‭using arbitrary similarities. Still, since the theory of evolution was proposed in the 19‬‭th‬
‭century, biologists have worked to make the classification system reflect evolutionary‬
‭relationships. This means that all of the members of a taxon should have a common‬
‭ancestor and be more closely related to each other than to members of other taxa.‬
‭ ecent genetic analysis and other advancements have found that some earlier taxonomic‬
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‭classifications do not reflect actual evolutionary relationships. Therefore, changes and‬
‭updates must be made as new discoveries take place. One dramatic and recent example‬
‭was the breaking apart of prokaryotic species, which, until the 1970s, were all classified as‬
‭bacteria. Their division into Archaea and Bacteria came about after recognizing that their‬
‭significant genetic differences warranted their separation into two of three fundamental‬
‭branches of life.‬

‭Phylogeny and Phylogenetic Trees‬

‭ hylogeny‬‭is the evolutionary history and the relationships‬‭among a species or group of‬
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‭species. Each organism shares relatedness with others, and based on morphologic and‬
‭genetic evidence, scientists attempt to map the evolutionary pathways of all life on Earth.‬
‭Historically, organisms were organized into a taxonomic classification system. However,‬
‭today, many scientists build phylogenetic trees to illustrate evolutionary relationships, and‬
‭the taxonomic classification system is expected to reflect evolutionary relationships.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 21.16‬ ‭At each sublevel in the taxonomic classification‬‭system, organisms become‬
F
‭more similar. Dogs and wolves are the same species because they can breed and produce‬
‭viable offspring, but they are different enough to be classified as distinct subspecies.‬
‭(credit “plant”: modification of work by "berduchwal"/Flickr; credit “insect”: modification‬
‭o f work by Jon Sullivan; credit “fish”: modification of work by Christian Mehlfü hrer; credit‬
‭“rabbit”: modification of work by Aidan Wojtas; credit “cat”: modification of work by‬
‭Jonathan Lidbeck; credit “fox”: modification of work by Kevin Bacher, NPS; credit “jackal”:‬
‭modification of work by Thomas A. Hermann, NBII, USGS; credit “wolf” modification of‬
‭work by Robert Dewar; credit “dog”: modification of work by "digital_image_fan"/Flickr;‬
‭OpenStax‬‭)‬

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‭ ‬‭phylogenetic tree‬‭is a diagram that reflects evolutionary relationships among‬

A
‭o rganisms or groups of organisms. In other words, a “tree of life” can be constructed to‬
‭illustrate when different organisms evolved and to show the relationships among various‬
‭o rganisms, as shown in‬‭Figure 21.17‬‭. The hierarchical‬‭classification of groups nested‬
‭within more inclusive groups is reflected in these diagrams. The point where a split‬
‭o ccurs in a tree is called a‬‭branch point‬‭, and it‬‭represents where a single lineage evolved‬
‭into distinct new ones. Notice that from a single point, the three domains of Archaea,‬
‭Bacteria, and Eukarya diverge and then branch repeatedly. Note the position of animals,‬
‭including humans, on this tree.‬
S‭ cientists consider phylogenetic trees to be a hypothesis of the evolutionary past because‬
‭o ne cannot go back through time to confirm the proposed relationships.‬

‭ igure 21.17‬ ‭In the evolution of life on Earth,‬‭the three domains of life—Archaea,‬
F
‭Bacteria, and Eukarya—branch from a single point. (credit: modification of work by Eric‬
‭Gaba;‬ ‭OpenStax‬‭)‬

‭ any phylogenetic trees have a single branch point at the base, representing a common‬
M
‭ancestor of all the branches in the tree. Scientists call such trees rooted, meaning there is a‬
‭single ancestral taxon at the base of a phylogenetic tree from which all organisms‬
‭represented in the diagram descend (‬‭Figure 21.18‬‭).‬‭When two lineages stem from the‬

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s‭ ame branch point, they are called sister taxa, for example, the two species of orangutans.‬
‭A branch point with more than two groups illustrates a situation where scientists have not‬
‭definitively determined relationships. An example is illustrated by the three branches‬
‭leading to the gorilla subspecies; their exact relationships still need to be understood. It is‬
‭important to note that sister taxa share an ancestor, which does not mean that one taxon‬
‭evolved from the other. The branch point, or split, represents a common ancestor that‬
‭existed in the past but no longer exists.‬
‭ umans did not evolve from chimpanzees (nor did chimpanzees evolve from humans),‬
H
‭although they are our closest living relatives. Both humans and chimpanzees evolved from‬
‭a common ancestor that lived, scientists believe, approximately six million years ago and‬
‭looked different from both modern chimpanzees and modern humans.‬

‭ igure 21.18‬ ‭A phylogenetic tree is rooted and shows‬‭how different organisms, in this‬
F
‭case, the species and subspecies of living apes, evolved from a common ancestor. (credit:‬
‭OpenStax‬‭)‬

‭ he branch points and the branches in the phylogenetic tree structure also imply‬
T
‭evolutionary change. Sometimes, significant character changes are identified on a branch‬
‭o r branch point. For example, in‬‭Figure 21.19‬‭, the‬‭branch point that gives rise to the‬
‭mammal and reptile lineage from the frog lineage shows the origin of the amniotic egg‬
‭character (i.e., the embryo is surrounded by fluid-filled membranes inside the shell). Also,‬
‭the branch point that gives rise to organisms with legs is indicated as the common‬
‭ancestor of mammals, reptiles, amphibians, and jawed fishes.‬
I‭ t is easy to assume that more closely related organisms look more alike, and while this is‬
‭o ften the case, it is not always true. Suppose two closely related lineages evolved under‬
‭significantly different surroundings or after the evolution of a major new adaptation. In‬
‭that case, they might look quite different from each other, even more so than other groups‬
‭that are not as closely related. For example, the phylogenetic tree in‬‭Figure 21.19‬‭shows‬

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t‭ hat lizards and rabbits both have amniotic eggs, whereas salamanders (within the frog‬
‭lineage) do not; yet, on the surface, lizards and salamanders appear more similar than the‬
‭lizards and rabbits.‬

‭ igure 21.19‬ ‭This phylogenetic tree is rooted in‬‭an organism that lacked a vertebral‬
F
‭column. At each branch point, organisms with different characters are placed in different‬
‭groups. (credit:‬‭OpenStax‬‭)‬

‭ nother aspect of phylogenetic trees is that, unless otherwise indicated, the branches do‬
A
‭not show length of time; they show only the order in time of evolutionary events. In other‬
‭words, a long branch does not necessarily mean more time passed, nor does a short‬
‭branch mean less time passed— unless specified on the diagram. For example, in‬‭Figure‬
‭21.19‬‭, the tree does not indicate how much time has‬‭passed between the evolution of‬
‭amniotic eggs and hair. What the tree does show is the order in which things took place.‬
‭In this same figure, the tree shows that the oldest trait is the vertebral column, followed by‬
‭hinged jaws, and so forth. Remember that any phylogenetic tree is a part of the greater‬
‭whole, and similar to a real tree, it does not grow in only one direction after a new branch‬
‭develops. Therefore, just because a vertebral column evolved does not mean that‬
‭invertebrate evolution ceased; it only means that a new branch formed. Also, groups that‬
‭are not closely related but evolve under similar conditions may appear more similar to‬
‭each other than to a close relative.‬

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‭Speciation‬
‭ he biological definition of‬‭species‬‭, which works for sexually reproducing organisms, is a‬
T
‭group of actual or potential interbreeding individuals. There are exceptions to this rule.‬
‭Many species are similar enough that hybrid offspring are possible and may often occur in‬
‭nature, but this rule generally holds for most species. The presence in nature of hybrids‬
‭between similar species suggests that they may have descended from a single‬
‭interbreeding species, and the speciation process may still need to be completed.‬
‭ iven the extraordinary diversity of life on the planet, there must be mechanisms for‬
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‭speciation‬‭, such as the formation of two species from‬‭o ne original species. Darwin‬
‭envisioned this process as a branching event and diagrammed the process in the only‬
‭illustration in‬‭On the‬‭Origin of Species‬‭(‬‭Figure 21.20a‬‭).‬‭Compare this illustration to the‬
‭diagram of elephant evolution (‬‭Figure 21.20b‬‭), which‬‭shows that as one species changes‬
‭over time, it branches to form more than one new species, repeatedly, as long as the‬
‭population survives or until the organism becomes extinct.‬

‭ igure 21.20‬ ‭The only illustration in Darwin's‬‭On‬‭the Origin of Species‬‭is (a) a diagram‬
F
‭showing speciation events leading to biological diversity. The diagram shows similarities to‬
‭phylogenetic charts that today illustrate the relationships of species. (b) Modern elephants‬
‭evolved from the‬‭Palaeomastodon‬‭, a species in Egypt‬‭35–50 million years ago. (credit:‬
‭OpenStax‬‭)‬

‭ or speciation to occur, two new populations must form from one original population, and‬
F
‭they must evolve in such a way that it becomes impossible for individuals from the two‬

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‭ ew populations to interbreed. Biologists have proposed mechanisms by which this could‬

n
‭o ccur that fall into two broad categories.‬‭Allopatric‬‭speciation‬‭(allo- = "other"; -patric =‬
‭"homeland") involves the geographic separation of populations from a parent species and‬
‭subsequent evolution.‬‭Sympatric speciation‬‭(sym- =‬‭"same"; -patric = "homeland")‬
‭involves speciation occurring within a parent species remaining in one location.‬
‭ iologists think of speciation events as splitting one ancestral species into two descendant‬
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‭species. There is no reason why more than two species might not form simultaneously‬
‭except that it is less likely, and we can conceptualize multiple events as single splits‬
‭o ccurring close in time.‬

‭21.8 A Brief Look at Human Evolution‬

‭The Evolution of‬‭Homo sapiens‬

‭ ur evolutionary story is associated with the rise of the genus‬‭Homo‬‭from the genus‬
O
‭Australopithecus,‬‭an event thought to have occurred‬‭approximately two to three million‬
‭years ago. At one time, the genus‬‭Homo‬‭represented‬‭at least eight different species in our‬
‭human lineage:‬‭Homo habilis‬‭,‬‭Homo rudolfensis‬‭,‬‭Homo‬‭erectus‬‭,‬‭Homo antecessor‬‭,‬‭Homo‬
‭heidelbergensis‬‭,‬‭Homo floresiensis‬‭,‬‭Homo neanderthalensis‬‭(the Neanderthals); however,‬
‭o nly‬‭Homo sapiens‬‭survived.‬
‭ odern‬‭Homo sapiens‬‭first appeared about 200,000 years‬‭ago in Africa. Anthropologists‬
M
‭generally classify these people as “anatomically modern‬‭H. sapiens‬‭,” which is a way of‬
‭noting that while their bodies are the same as modern humans, they had not yet‬
‭developed the cultural traditions, symbolic behaviors, and technologies that are seen‬
‭among later‬‭H. sapiens,‬‭including people of today.‬‭Probably the most defining feature of‬
‭anatomically modern‬‭H. sapiens‬‭is their chin. Modern‬‭H. sapiens‬‭is the first hominin (i.e.,‬
‭bipedal ancestor) to exhibit a projecting chin. One of the most common explanations for‬
‭this anatomical feature is that the chin evolved in response to human speech and protects‬
‭the jaw against stresses produced by the contraction of certain tongue muscles.‬
‭ round 40,000 years ago, there was an abrupt change in tool technology, subsistence‬
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‭patterns, and symbolic expression among‬‭H. sapiens.‬‭These changes seem to have‬
‭o ccurred almost simultaneously in Africa, Asia, Europe, and Australia. While there is‬
‭evidence of some creative artistic activity in earlier groups like the Neanderthal, they were‬
‭not on the same scale as that seen during this time, also referred to as “the human‬
‭revolution.” The level of cultural changes associated with this period has been compared‬
‭to the level of change that occurred during the Industrial Revolution of the 19th century.‬

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‭ or a more in-depth but concise review of human evolution, click the link or scan the QR‬
F
‭code to watch the American Museum of Natural History video titled “Seven Million Years‬
‭o f Human Evolution.”‬

Seven Million Years of Human Evolution

‭Are Humans Still Evolving?‬

‭ umans evolved for one million years as hunter-gatherers. Today, human bodies are still‬
H
‭trying to adapt to the largely grain-based diet brought about by agriculture, a diet‬
‭characterized by less diversity and lower nutrition levels than that of a typical‬
‭hunter-gatherer. Incomplete adaptation to this change has made people susceptible to‬
‭several diseases and nutritional deficiencies. Lactose intolerance is a prime example. The‬
‭domestication of cattle and the drinking of cow’s milk began during the agricultural age,‬
‭not very long ago in evolutionary history. Currently, 65 percent of humans are unable to‬
‭digest cow’s milk. Dental caries (cavities) are another problem linked to the change in diet‬
‭associated with agriculture. The grain-based and high-sugar diets associated with‬
‭agriculture are very different from the diet of hunter-gatherers. Neither our bodies nor‬
‭the bacteria in our mouths have had time to adapt to this change fully.‬
‭ nother adaptation that occurred during the Neolithic era (approximately 12,000 years‬
A
‭ago) is related to variation in skin pigmentation. Humans who left Africa and settled in‬
‭Europe about 40,000 years ago most likely had dark skin with high levels of melanin,‬
‭which protects against ultraviolet radiation. New data confirms that about 8,500 years‬
‭ago, early hunter-gatherers in Spain, Luxembourg, and Hungary also had darker skin. Skin‬
‭pigmentation is an adaptation to ultraviolet radiation, with different tones offering‬
‭different advantages depending on one’s distance from the equator. As humans migrated‬
‭to the Northern Hemisphere, they were exposed to less ultraviolet radiation, which meant‬
‭less absorption of the Vitamin D needed for strong bones and other essential immune‬
‭functions. Skin pigmentation became lighter to compensate for this loss and allow for‬
‭greater exposure to ultraviolet radiation.‬
‭ nother example of human variation due to environmental adaptation can be seen in‬
A
‭Indigenous populations in the Andes, Tibet, and the Ethiopian highlands. Each of these‬
‭three groups faces the same environmental challenge, living in a low-oxygen environment,‬
‭and they have responded with unique adaptations. Tibetans compensate for low oxygen‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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l‭evels by taking more breaths per minute than people who live at sea level. Those living at‬
‭high altitudes in the Andes have been found to have higher concentrations of hemoglobin‬
‭in their blood than other people. Ethiopians living at altitudes of 9,800 to 11,580 feet have‬
‭neither of these adaptations. The explanation of how Ethiopian highlanders thrive in their‬
‭environment is still a mystery.‬
‭ or more information about the rate of human evolution, click on the link or scan the QR‬
F
‭code to watch the Ted-Ed video by Laurence Hurst titled “Is human evolution speeding up‬
‭o r slowing down?”‬

Is human evolution speeding up or slowing down? - Laurence Hu…

‭21.9 Genetic Changes in Populations‬

‭ he modern synthesis of evolutionary theory grew from the reconciliation of Darwin’s,‬
T
‭Wallace’s, and Mendel’s thoughts on evolution and heredity.‬‭Population genetics‬‭is a‬
‭theoretical framework for describing evolutionary change in populations through the‬
‭change in allele frequencies over generations. Without evolutionary forces, allele‬
‭frequencies will not change in a population; this is known as the Hardy-Weinberg‬
‭equilibrium principle. However, in most populations, factors such as mutation, natural‬
‭selection, genetic drift, and gene flow act to change allele frequencies.‬
‭ or more information about factors that change allele frequency, click on the link or scan‬
F
‭the QR code to watch the Ted Ed video by Paul Anderson titled “Five fingers of evolution.”‬

Five fingers of evolution - Paul Andersen

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Allele Frequencies‬
‭ ecall that a gene for a particular character may have several alleles or variants that code‬
R
‭for different traits associated with that character. For example, in the ABO blood type‬
‭system in humans, three alleles determine the particular blood-type carbohydrate on the‬
‭surface of red blood cells. Each individual in a population of diploid organisms can only‬
‭carry two alleles for a specific gene. Still, more than two may be present in the‬
‭population's individuals. Mendel followed alleles as they were inherited from parent to‬
‭o ffspring. In the early twentieth century, biologists in population genetics began to study‬
‭how selective forces change a population through changes in allele and genotypic‬
‭frequencies.‬
‭ llele frequency‬‭is the rate at which a specific allele‬‭appears within a population. Until‬
A
‭now, we have discussed evolution as a change in the characteristics of a population of‬
‭o rganisms, but behind that, phenotypic change is genetic change. In population genetics,‬
‭scientists define evolution as a change in the allele's frequency in a population. Using the‬
‭ABO blood type system as an example, the frequency of one of the alleles,‬‭I‭A‬ ‬‭, is the‬
‭number of copies of that allele divided by all the copies of the ABO gene in the population.‬
‭For example, a study in Jordan (‬‭1‬‭) found that I‬‭A‬ ‭frequency‬‭was 26.1 percent. The‬‭I‬‭B‬ ‭and‬‭I‬‭0‬
‭alleles comprise 13.4 percent and 60.5 percent, respectively, and all of the frequencies‬
‭added up to 100 percent. A change in this frequency over time would constitute evolution‬
‭in the population.‬
‭ he allele frequency within a given population can change depending on environmental‬
T
‭factors; therefore, certain alleles become more widespread than others during the natural‬
‭selection process. Natural selection can alter the population’s genetic makeup. An example‬
‭is if a given allele confers a phenotype that allows an individual to survive better or have‬
‭more offspring. Because many of those offspring will also carry the beneficial allele, and‬
‭o ften the corresponding phenotype, they will have more offspring that also carry the‬
‭allele, thus perpetuating the cycle. Over time, the allele will spread throughout the‬
‭population. Some alleles will quickly become‬‭fixed‬‭in this way, meaning that every‬
‭individual in the population will carry the allele. At the same time, detrimental mutations‬
‭may be swiftly eliminated if derived from a dominant allele from the gene pool. The‬‭gene‬
‭pool‬‭is the sum of all the alleles in a population.‬

‭Hardy-Weinberg Principle of Equilibrium‬

I‭ n the early twentieth century, English mathematician Godfrey Hardy and German‬
‭physician Wilhelm Weinberg stated the principle of equilibrium to describe the‬
‭population's genetic makeup. The theory, later known as the‬‭Hardy-Weinberg principle‬
‭of equilibrium‬‭, states that a population’s allele‬‭and genotype frequencies are inherently‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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s‭ table. In other words, unless some evolutionary force is acting upon the population,‬
‭neither the allele nor the genotypic frequencies would change. The Hardy-Weinberg‬
‭principle assumes conditions with no mutations, gene flow, or selective pressure for or‬
‭against genotype, plus an infinitely large population size. While no population can satisfy‬
‭those conditions, the principle offers a useful model against which to compare real‬
‭population changes.‬
‭ orking under this theory, population geneticists represent different alleles as different‬
W
‭variables in their mathematical models. The variable‬‭p‬‭, for example, often represents the‬
‭frequency of a particular allele, say‬‭Y,‬‭for the trait‬‭o f yellow in Mendel’s peas. In contrast,‬
‭the variable‬‭q‬‭represents the frequency of‬‭y‬‭alleles‬‭that confer the color green. If these are‬
‭the only two possible alleles for a given locus in the population, then‬‭p + q = 1‬‭. In other‬
‭words, all the p and q alleles comprise all of the alleles for that gene in the population.‬
‭ owever, what ultimately interests most biologists is not the frequencies of different‬
H
‭alleles but the frequencies of the resulting genotypes, known as the population’s genetic‬
‭structure, from which scientists can surmise phenotype distribution. If we observe the‬
‭phenotype, we can know only the homozygous recessive allele's genotype. The‬
‭calculations provide an estimate of the remaining genotypes. Since each individual carries‬
‭two alleles per gene if we know the allele frequencies (p and q), predicting the genotypes'‬
‭frequencies is a simple mathematical calculation to determine the probability of obtaining‬
‭these genotypes if we randomly draw two alleles from the gene pool. In the above‬
‭scenario, an individual pea plant could be pp (‬‭YY‬‭)‬‭and thus produce yellow peas; pq (‬‭Yy‬‭),‬
‭also yellow; or qq (‬‭yy‬‭), and thus produce green peas‬‭(‬‭Figure 21.21‬‭). In other words, the‬
‭frequency of pp individuals is simply‬‭p‭2‬ ‭;‬ the frequency‬‭o f pq individuals is‬‭2pq‬‭, and the‬
‭frequency of qq individuals is‬‭q‭2‬ ‭.‬ Again, if p and‬‭q are the only two possible alleles for a‬
‭trait in the population, these genotype frequencies will sum to one:‬‭p‭2‬ ‬ ‭+ 2pq + q‬‭2‬ ‭= 1‬‭.‬
I‭ f a population is genuinely at equilibrium—that is, no evolutionary forces are acting upon‬
‭it—generation after generation would have the same gene pool and genetic structure.‬
‭These equations would all hold true all of the time. Of course, even Hardy and Weinberg‬
‭recognized that no natural population is immune to evolution. Populations in nature are‬
‭constantly changing in genetic makeup due to drift, mutation, possibly migration, and‬
‭selection. As a result, the only way to determine the exact distribution of phenotypes in a‬
‭population is to go out and count them. However, the Hardy-Weinberg principle gives‬
‭scientists a mathematical baseline of a non-evolving population to which they can compare‬
‭evolving populations and thereby infer what evolutionary forces might be at play. If the‬
‭frequencies of alleles or genotypes deviate from the value expected from the‬
‭Hardy-Weinberg equation, then the population is evolving.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ or additional information about the Hardy-Weinberg equation and its uses, click the link‬
F
‭o r scan the QR code to watch the Amoeba Sisters video titled “Hardy-Weinberg‬
‭Equilibrium.”‬

Hardy-Weinberg Equilibrium

‭Adaptive Evolution‬
‭ atural selection acts on the population’s heritable traits: selecting for beneficial alleles‬
N
‭that allow for environmental adaptation, thus increasing their frequency in the population,‬
‭while selecting against deleterious alleles and decreasing their frequency. Scientists call‬
‭this process‬‭adaptive evolution‬‭. Natural selection‬‭acts on entire organisms, not on an‬
‭individual allele within the organism. An individual may carry a very beneficial genotype‬
‭with a resulting phenotype that, for example, increases the ability to reproduce, but if that‬
‭same individual also carries an allele that results in a fatal childhood disease, that‬
‭reproductive success phenotype will not pass to the next generation because the‬
‭individual will not live to reach reproductive age. Natural selection acts at the individual's‬
‭level. It selects individuals with greater contributions to the next generation's gene pool.‬
‭Scientists call this an organism’s evolutionary (Darwinian) fitness.‬
‭ itness is often quantifiable and measured by scientists in the field. However, it is not an‬
F
‭individual's absolute fitness that counts but rather how it compares to the other‬
‭o rganisms in the population. Scientists call this concept‬‭relative fitness‬‭, which allows‬
‭researchers to determine which individuals contribute additional offspring to the next‬
‭generation and, thus, how the population might evolve.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 21.21‬ ‭When populations are in the Hardy-Weinberg‬‭equilibrium, the allelic‬

F
‭frequency is stable from generation to generation, and we can determine the allele‬
‭distribution from the Hardy-Weinberg equation. If the allelic frequency measured in the‬
‭field differs from the predicted value, scientists can infer what evolutionary forces are at‬
‭play. (credit:‬‭OpenStax‬‭). A link to a video explanation‬‭o f this figure is available at‬
‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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S‭ election can affect population variation in several ways, including stabilizing selection,‬
‭directional selection, and diversifying selection. As natural selection influences the allele‬
‭frequencies in a population, individuals can become more or less genetically similar, and‬
‭the phenotypes can become more similar or more disparate.‬

‭Stabilizing Selection‬
I‭ f natural selection favors an average phenotype, selecting against extreme variation, the‬
‭population will undergo‬‭stabilizing selection‬‭(‬‭Figure‬‭21.22a‬‭). In a mouse population‬
‭that lives in the woods, for example, natural selection is likely to favor mice that best blend‬
‭in with the forest floor and are less likely for predators to spot. Assuming the ground is a‬
‭relatively consistent shade of brown, those mice whose fur is most closely matched to that‬
‭color will most likely survive and reproduce, passing on their genes for their brown coat.‬
‭Mice that carry alleles that make them a bit lighter or darker will stand out against the‬
‭ground and be more likely to fall victim to predation. As a result of this selection, the‬
‭population’s genetic variance (or variation) will decrease.‬

‭Directional Selection‬
‭ hen the environment changes, populations often undergo‬‭directional selection‬‭(‬‭Figure‬
W
‭21.22b‬‭), which selects for phenotypes at one end of‬‭the spectrum of existing variation. A‬
‭classic example of this type of selection is the evolution of the peppered moth in‬
‭eighteenth- and nineteenth-century England. Before the Industrial Revolution, the moths‬
‭were predominately light in color, which allowed them to blend in with the light-colored‬
‭trees and lichens in their environment. However, as soot began spewing from factories,‬
‭the trees darkened, and the light-colored moths became easier for predatory birds to‬
‭spot. Over time, the frequency of the moth's melanic form increased because they had a‬
‭higher survival rate in habitats affected by air pollution because their darker coloration‬
‭blended with the sooty trees. Similarly, the hypothetical mouse population may evolve to‬
‭take on a different coloration if something were to cause the forest floor where they live‬
‭to change color. This type of selection results in a shift in the population’s genetic variance‬
‭toward the new, fit phenotype.‬

‭Diversifying Selection‬
S‭ ometimes, two or more distinct phenotypes can each have their advantages for natural‬
‭selection, while the intermediate phenotypes are, on average, less fit. Scientists call this‬
‭diversifying selection‬‭(‬‭Figure 21.22c‬‭). We see this‬‭in many animal populations that have‬
‭multiple male forms. Large, dominant alpha males use brute force to obtain mates, while‬
‭small males can sneak in for furtive copulations with the females in an alpha male’s‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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t‭ erritory. In this case, both the alpha males and the “sneaking” males will be selected for,‬
‭but medium-sized males, who can’t overtake the alpha males and are too big to sneak‬
‭copulations, are selected against. Diversifying selection can also occur when‬
‭environmental changes favor individuals on either end of the phenotypic spectrum.‬
‭Imagine a mouse population living at the beach with light-colored sand interspersed with‬
‭patches of tall grass. In this scenario, light-colored mice that blend in with the sand and‬
‭dark-colored mice that can hide in the grass would be favored. Medium-colored mice,‬
‭alternatively, would not blend in with either the grass or the sand, and thus predators‬
‭would most likely eat them. This type of selection results in increased genetic variance as‬
‭the population becomes more diverse.‬

‭No Perfect Organism‬

‭ atural selection is a driving force in evolution and can generate populations better‬
N
‭adapted to survive and successfully reproduce in their environments. However, natural‬
‭selection cannot produce the perfect organism. Natural selection can only be based on‬
‭existing variations in the population. It does not create anything from scratch. Thus, it is‬
‭limited by a population’s existing genetic variance and whatever new alleles arise through‬
‭mutation and gene flow.‬
‭ inally, it is important to understand that not all evolution is adaptive. While natural‬
F
‭selection selects the fittest individuals and often results in a more fit population overall,‬
‭o ther forces of evolution, including genetic drift and gene flow, usually do the opposite:‬
‭introducing deleterious alleles to the population’s gene pool. Evolution has no purpose—it‬
‭is not changing a population into a preconceived ideal. It is simply the sum of the various‬
‭forces described in this chapter and how they influence the population's genetic and‬
‭phenotypic variance.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 21.22‬ ‭Different types of natural selection‬‭can impact the distribution of‬
F
‭phenotypes within a population. In (a) stabilizing selection, an average phenotype is‬
‭favored. In (b) directional selection, the environment changes the spectrum of observed‬
‭phenotypes. In (c) diversifying selection, two or more extreme phenotypes are selected‬
‭for, while the average phenotype is selected against. (credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭21.10 Career Connection - Field Biologist‬

‭ any people hike, explore caves, scuba dive, or climb mountains for recreation. People‬
M
‭o ften participate in these activities, hoping to see wildlife. Experiencing the outdoors can‬
‭be incredibly enjoyable and invigorating. What if your job entailed working in the‬
‭wilderness? Field biologists, by definition, work outdoors in the “field.” In this case, the‬
‭term field refers to any location outdoors, even underwater. A field biologist typically‬
‭focuses research on a specific species, group of organisms, or a single habitat (‬‭Figure‬
‭21.23‬‭).‬

‭ igure 21.23‬ ‭A field biologist tranquilizes a polar‬‭bear for study. (credit: Karen Rhode;‬
F
‭OpenStax‬‭)‬

‭ ne objective of many field biologists includes discovering new, unrecorded species. Not‬
O
‭o nly do such findings expand our understanding of the natural world, but they also lead‬
‭to important innovations in fields such as medicine and agriculture. Plant and microbial‬
‭species, in particular, can reveal new medicinal and nutritional knowledge. Other‬
‭o rganisms can play key roles in ecosystems or, if rare, require protection. When‬
‭discovered, researchers can use these important species as evidence for environmental‬
‭regulations and laws.‬

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‭Chapter 22: Molecular Biology and Biotechnology‬

‭ igure 22.1‬ ‭Laboratory scientists use various molecular‬‭biology techniques and‬

F
‭protocols as part of biotechnology research and development. This research scientist is‬
‭using a micropipette to transfer small volumes of liquid. (Credit: David Baras; Armed‬
‭Forces Medical Examiner;‬‭Nara‬‭Public Domain Archives‬‭;‬‭CC0 1.0‬‭)‬

‭22.1 Introduction‬
‭ iotechnology‬‭uses molecular biology techniques to‬‭modify the genetic material of living‬
B
‭o rganisms or cells to produce desired compounds or perform new functions. Since the‬
‭discovery of DNA’s structure in 1953, biotechnology has grown rapidly through academic‬
‭research and private companies. The primary applications are in medicine (production of‬
‭vaccines and antibiotics) and agriculture (genetic modification of crops to increase yields).‬
‭Biotechnology also has many industrial applications, such as fermentation, the treatment‬
‭o f oil spills, and the production of biofuels.‬‭The‬‭rate of discovery and the development of‬
‭new applications in medicine, agriculture, and energy is expected to accelerate, bringing‬
‭considerable benefits to humankind and perhaps also significant risks. Many of these‬
‭developments are also likely to raise important ethical and social questions that human‬
‭societies have not yet considered.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭22.2 Molecular Biology Techniques‬

‭ iotechnology commonly utilizes various molecular biology techniques to accomplish the‬
B
‭desired outcomes. It will be helpful to discuss these techniques before learning about their‬
‭application.‬

‭Basic Techniques to Manipulate Genetic Material (DNA and RNA)‬

‭ o understand the basic techniques used to work with nucleic acids, remember that‬
T
‭nucleic acids‬‭are macromolecules made of‬‭nucleotides‬‭(a sugar, a phosphate, and a‬
‭nitrogenous base) linked by‬‭phosphodiester bonds‬‭.‬‭The phosphate groups on these‬
‭molecules each have a net negative charge. DNA has two complementary strands linked by‬
‭hydrogen bonds‬‭between the paired bases. Exposure‬‭to high temperatures can separate‬
‭the two DNA strands by breaking the hydrogen bonds (i.e.,‬‭denaturation‬‭), and cooling‬
‭can‬‭reanneal‬‭them. The DNA polymerase enzyme replicates‬‭DNA, using available‬
‭nucleotides, to produce strands with complementary sequences. Unlike DNA, which is‬
‭located in the eukaryotic cells' nucleus, RNA molecules leave the nucleus. The most‬
‭common type of RNA researchers analyze is‬‭messenger‬‭RNA‬‭(mRNA) because it‬
‭represents the protein-coding genes actively expressed. However, RNA molecules present‬
‭some other challenges to analysis, as they are often less stable than DNA.‬

‭DNA and RNA Extraction‬

‭ ne must first isolate or‬‭extract‬‭the DNA or RNA from‬‭the cells to study or manipulate‬
O
‭nucleic acids. Most extraction techniques use a lysis buffer containing detergent to break‬
‭o pen cells’ plasma and nuclear membranes made of phospholipids (‬‭lysis‬‭means "to split").‬
‭After steps to remove proteins and other undesired components are completed, alcohol is‬
‭added to‬‭precipitate‬‭the nucleic acid (i.e., to change‬‭DNA and RNA from a soluble to an‬
‭insoluble form) so it can be isolated (‬‭Figure 22.2‬‭).‬‭Human genomic DNA is usually visible‬
‭as a gelatinous, white mass.‬

‭Gel Electrophoresis‬
‭ ecause nucleic acids are negatively charged ions at neutral or basic pH in an aqueous‬
B
‭environment, an electric field can mobilize them.‬‭Gel electrophoresis‬‭is a technique that‬
‭scientists use to separate molecules based on size using this charge. The isolated,‬
‭dissolved nucleic acid samples are loaded, using a micropipette (‬‭Figure 22.1‬‭), into a slot‬
‭(called a “well”) near the semisolid, porous gel matrix's‬‭negative electrode‬‭. They are then‬
‭pulled toward the‬‭positive electrode‬‭at the gel's‬‭o pposite end. Smaller molecules move‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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t‭ hrough the gel's pores faster than larger molecules, and this difference in the migration‬
‭rate separates the fragments based on size.‬

‭(a)‬ ‭(b)‬

‭ igure 22.2‬ ‭(a) This diagram shows the basic DNA‬‭extraction method. (credit:‬‭OpenStax‬‭)‬
F
‭(b) DNA extracted from an orange. (credit:‬ ‭Mariano‬‭Avagliano‬‭;‬‭Estrazione DNA‬
‭(cropped).jpg‬‭; Wikimedia Commons;‬‭CC BY 4.0‬‭)‬

‭ e can observe nucleic acids in an electrophoresis gel using various fluorescent or‬
W
‭colored dyes. Distinct nucleic acid fragments appear as‬‭bands‬‭at specific distances from‬
‭the gel's top (the negative electrode end) based on their size (base pairs; bp). A mixture of‬
‭genomic DNA fragments of varying sizes appears as a long smear; uncut genomic DNA is‬
‭usually too large to run through the gel and forms a single large band at the gel's top.‬
‭Molecular weight standard samples, which are collections of fragments of known size (bp),‬
‭can be included when loading the gel and run alongside other nucleic acid molecules to‬
‭provide a size comparison (‬‭Figure 22.3‬‭).‬
‭ or additional information about electrophoresis, click the link or scan the QR code to‬
F
‭watch the Amoeba Sisters video titled “Gel Electrophoresis.”‬

Gel Electrophoresis

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 22.3‬ ‭a) Shown are DNA fragments from seven‬‭samples run on a gel, stained with‬
F
‭a fluorescent dye, and viewed under UV light; the first and last wells contained molecular‬
‭weight size markers, and the wells between them contained fragments of unknown sizes;‬
‭b) A researcher from International Rice Research Institute, reviewing gel electrophoresis‬
‭results using UV light. (credit: a: James Jacob, Tompkins Cortland Community College; b:‬
‭International Rice Research Institute;‬‭OpenStax‬‭);‬ ‭A link to a video explanation of this‬
‭figure is available at‬‭Biology411.com‬‭.‬

‭Hybridization, Southern Blotting, and Northern Blotting‬

S‭ cientists can‬‭probe (i.e., search) nucleic acid samples,‬‭such as fragmented genomic DNA‬
‭and RNA extracts, for the presence of specific sequences of bases. Gel electrophoresis‬
‭separates the nucleic acid fragments according to their size. Scientists then transfer the‬
‭fragments in the gel to a nylon membrane using a procedure called‬‭blotting‬‭(‬‭Figure‬
‭22.4‬‭). When‬‭DNA‬‭is transferred to a nylon membrane,‬‭the technique is called‬‭Southern‬
‭blotting‬‭. When‬‭RNA‬‭is transferred to a nylon membrane,‬‭it is called‬‭Northern blotting‬‭.‬
‭Scientists use Southern blots to detect the presence of specific DNA sequences in a given‬
‭genome and Northern blots to detect gene expression.‬

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‭ igure 22.4‬ ‭Scientists use Southern blotting to‬‭find a particular sequence in a DNA‬
F
‭sample. Scientists separate DNA fragments on a gel, denature them via pH changes,‬
‭transfer them to a nylon membrane, and incubate them with a single-stranded DNA probe‬
‭complementary to the sequence of interest. Northern blotting is similar to Southern‬
‭blotting, but scientists run RNA on the gel instead of DNA. In Western blotting, scientists‬
‭run proteins on a gel and detect them using antibodies. (credit:‬‭OpenStax‬‭); A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ nce the nylon membrane is prepared, scientists can‬‭probe‬‭the nucleic acid fragments‬
O
‭bound to its surface.‬ ‭Probes‬‭are single-stranded‬‭DNA fragments joined to radioactive or‬
‭fluorescent dyes, which aid in the detection of complementary fragments on the nylon‬
‭membranes. In a Southern blot, a DNA probe will bind to DNA fragments with‬
‭complementary base sequences (i.e., A to T and G to C). In a Northern blot, a DNA probe‬
‭will bind to RNA fragments with complementary base sequences (i.e., A to U and G to C).‬

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‭ or example, a specific probe could be used to determine if a gene in mice is also present‬
F
‭in humans. If so, mice could be used as a model organism to study the gene and its effects.‬

‭ or additional information about blotting methods, click the link or scan the QR code to‬
F
‭watch the Cleavaforce video titled “Introduction to blotting.”‬

Introduction to Blotting

‭Cloning‬
I‭ n general, cloning means the creation of a perfect replica. Typically, the word describes‬
‭the creation of a genetically identical copy. Long before attempts were made to clone an‬
‭entire organism (i.e., reproductive cloning), researchers learned how to copy short‬
‭stretches of DNA using a process called bacterial cloning. These methods are discussed in‬
‭the following sections.‬

‭In-vivo (Bacterial) Cloning‬

‭ acterial cloning‬‭allows for the creation of multiple‬‭copies of genes and the expression‬
B
‭o f genes using bacteria. This method is called‬‭in-vivo‬‭cloning, as it occurs within a living‬
‭o rganism. The fragment is first inserted into a plasmid to get the DNA fragment of interest‬
‭into a bacterial cell in a form that will be copied or expressed. A‬‭plasmid‬‭(also called a‬
‭vector‬‭in this context) is a small circular DNA molecule‬‭that replicates independently of‬
‭the bacterial chromosomal DNA. In cloning, plasmids provide a "vehicle" to insert a‬
‭desired DNA fragment that will be moved into a bacterial host cell for replication or‬
‭expression. As the bacteria divide, they copy their DNA and also the plasmids. Any DNA‬
‭fragment inserted into a plasmid and the rest of the bacterial DNA are copied.‬
‭ lasmids occur naturally in bacterial populations (such as‬‭E. coli‬‭) and have genes that can‬
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‭contribute favorable traits to the organism, such as antibiotic resistance (the ability to be‬
‭unaffected by antibiotics). Plasmids have been highly engineered as vectors for molecular‬
‭cloning and the subsequent large-scale production of important molecules via gene‬
‭expression for proteins such as insulin.‬

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‭ valuable characteristic of plasmid vectors is the ease with which a foreign DNA fragment‬
A
‭can be introduced. Plasmid vectors contain many short DNA sequences that can be cut‬
‭(i.e., the phosphodiester bonds between bases are broken) with different available‬
‭restriction enzymes.‬‭Restriction enzymes‬‭(also called‬‭restriction endonucleases) are‬
‭enzymes produced by bacterial cells that recognize specific DNA sequences and‬
‭predictably cut them; bacteria naturally produce them as a defense mechanism against‬
‭foreign DNA from viruses. The sequence that is recognized by the restriction enzyme is a‬
‭four- to eight-nucleotide sequence that is a‬‭palindrome‬‭.‬‭This means the sequence reads‬
‭the same forward and backward (e.g., radar, level, and noon). In the case of DNA, a‬
‭palindromic sequence reads the same forward on one strand and backward on the‬
‭complementary strand. (e.g., 5’GGATCC3’/5’CCTAGG3’). Many restriction enzymes make‬
‭staggered cuts in the two strands of DNA, such that the cut ends have two- to‬
‭four-nucleotide, single-stranded overhangs. When a staggered cut is made in a sequence‬
‭like this, the overhangs are complementary (‬‭Figure‬‭22.5‬‭).‬

‭ igure 22.5‬ ‭In this (a) six-nucleotide restriction‬‭enzyme recognition site, notice that the‬
F
‭sequence of six nucleotides reads the same in the 5' to 3' direction on one strand as in the‬
‭5' to 3' direction on the complementary strand. This is known as a palindrome. (b) The‬
‭restriction enzyme makes breaks in the DNA strands, and (c) the cuts in the DNA result in‬
‭“sticky ends”. Another piece of DNA cut on either end by the same restriction enzyme‬
‭could attach to these sticky ends and be inserted into the gap made by this cut. (credit:‬
‭OpenStax‬‭); A link to a video explanation of this‬‭figure is available at‬‭Biology411.com‬‭.‬

‭ ecause these overhangs are capable of coming back together by hydrogen bonding‬
B
‭between complementary bases on pieces of DNA cut with the same restriction enzyme,‬
‭they are called “‬‭sticky ends‬‭.” The process of forming‬‭hydrogen bonds between‬
‭complementary sequences on single strands to form double-stranded DNA is called‬

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‭ nnealing‬‭. Addition of an enzyme called‬‭DNA ligase‬‭, which takes part in DNA replication‬
a
‭in cells, permanently joins the DNA fragments, via phosphodiester bonds, when the sticky‬
‭ends come together. In this way, any DNA fragment can be spliced between the two ends‬
‭o f a plasmid DNA cut with the same restriction enzyme (‬‭Figure 22.6‬‭). Plasmids with‬
‭foreign DNA inserted into them are called‬‭recombinant‬‭because the addition has altered‬
‭their original DNA sequence.‬
‭ or additional information about bacterial cloning and blue-white screening, click the link‬
F
‭o r scan the QR code to watch the Henrick’s Lab video titled “Blue-white screening and‬
‭transformation.”‬

Blue-White Screen & Transformation

‭In-vitro Cloning: Polymerase Chain Reaction (PCR)‬

‭ NA analysis often requires focusing on one or more specific genome regions. It also‬
D
‭frequently involves situations in which only one or a few copies of a DNA molecule are‬
‭available for further analysis. These amounts are insufficient for most procedures, such as‬
‭gel electrophoresis.‬‭Polymerase chain reaction‬‭(‬‭PCR‬‭)‬‭is a technique used to rapidly‬
‭increase the number of copies of specific DNA regions for further analyses (‬‭Figure 22.7‬‭).‬
‭PCR uses a particular form of‬‭DNA polymerase‬‭, the‬‭enzyme that replicates DNA, and‬
‭short nucleotide sequences called‬‭primers‬‭that base‬‭pair to a specific portion of the DNA‬
‭being copied, in a similar manner that probes used in blotting protocols hybridize with‬
‭complementary sequences on nylon membranes. Heat (approximately 95°C or 203°F) is‬
‭used to denature the double-stranded DNA. When the reaction mixture is cooled to‬
‭approximately 50°C, the primers will find and bind with complementary sequences faster‬
‭than the original DNA strands will renature. Then, the mixture is heated again (72°C) to‬
‭allow DNA polymerase to replicate the DNA from the points of the primers flanking the‬
‭region to be copied. The combination of these three temperature profiles make up one‬
‭PCR cycle (i.e., denaturation, primer annealing, and primer extension), and the process is‬
‭repeated over and over, resulting in an exponential increase in the number of copies of‬
‭the segment defined by the primers. For example, after 30 cycles of PCR, one template‬
‭copy will be amplified to produce more than 1 billion copies!‬

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‭ igure 22.6‬ ‭This diagram shows the steps involved‬‭in molecular cloning using blue and‬
F
‭white screening. (credit:‬‭OpenStax‬‭); A link to‬‭a video explanation of this figure is‬
‭available at‬‭Biology411.com‬‭.‬

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‭ igure 22.7‬ ‭Scientists use polymerase chain reaction,‬‭o r PCR, to amplify (i.e., make‬
F
‭copies of) a specific DNA sequence. Primers, which are short, single-stranded pieces of‬
‭DNA, are complementary to each end of the target sequence. The two primers are‬
‭combined with genomic DNA, Taq DNA polymerase, and deoxynucleotides (dNTPs). The‬
‭tubes containing this reaction mixture are then heated and cooled in a specific manner,‬
‭usually using a thermal cycler. The result is an exponential increase in the number of‬
‭copies of the target double-stranded DNA sequence. Taq polymerase is a DNA polymerase‬
‭isolated from the thermostable bacterium‬‭Thermus aquaticus‬‭that can withstand the high‬
‭temperatures (e.g., 94 to 97℃) that scientists use in PCR.‬‭Thermus aquaticus‬‭grows in the‬
‭Lower Geyser Basin of Yellowstone National Park. (credit:‬‭OpenStax‬‭); A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ CR is used in laboratories for many purposes. These include 1) the identification of the‬
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‭owner of a DNA sample left at a crime scene; 2) paternity analysis (i.e., determination of a‬
‭child’s father); 3) the comparison of small amounts of ancient DNA with modern‬
‭o rganisms; and 4) determining the sequence of nucleotides in a specific region (i.e., DNA‬
‭sequencing).‬
‭ or additional information about PCR, click the link or scan the QR code to watch the‬
F
‭Amoeba Sisters video titled “PCR (Polymerase Chain Reaction).”‬

PCR (Polymerase Chain Reaction)

‭ igure 22.8‬ ‭An example of a thermal cycler used‬‭in the PCR technique. On the top is an‬
F
‭aluminum block with spaces for tubes containing the PCR mixture that can be heated and‬
‭cooled to the preprogrammed temperatures for specific intervals. Lids cover the tubes to‬
‭make sure they achieve uniform temperatures. (credit:‬ ‭Double-block PCR thermocycler‬
‭MJ Research PTC-200‬‭;‬ ‭Cygaretka‬‭;‬‭Wikimedia Commons;‬‭CC BY-SA 1.0 DEED‬‭); A link to a‬
‭video explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭Reproductive Cloning‬

‭ eproductive cloning‬‭is a method used to make a clone‬‭o r an identical copy of an entire‬

R
‭multicellular organism. Most multicellular organisms undergo reproduction by sexual‬
‭means, which involves the union, during fertilization, of a sperm and an egg. Each of these‬
‭gametes is haploid, meaning they contain one set of chromosomes in their nuclei. The‬
‭resulting cell, or zygote, is then diploid and includes two sets of chromosomes. This cell‬
‭divides mitotically to produce a multicellular organism. However, the union of just any two‬
‭cells cannot make a viable zygote; there are components in the cytoplasm of the egg cell‬
‭that are essential for the early development of the embryo during its first few cell‬
‭divisions. These provisions are necessary for subsequent development. Therefore, a‬
‭diploid genetic complement and an egg’s cytoplasm are required to produce a new‬
‭individual. The approach to making an artificially cloned individual is to take an egg cell of‬
‭o ne individual and remove the haploid nucleus via a process called‬‭enucleation‬‭. Then a‬
‭diploid nucleus from a somatic cell of a second individual, the donor, is put into the egg‬
‭cell. The egg is then stimulated to divide so that development proceeds. This sounds‬
‭simple, but it takes many attempts before each of the steps is completed successfully.‬
‭ he first cloned agricultural animal was‬‭Dolly‬‭, a‬‭sheep born in 1996 (‬‭Figure 22.9‬‭). The‬
T
‭success rate of reproductive cloning at the time was very low. Dolly lived for six years and‬
‭died of a lung tumor. There was speculation that because the cell DNA that gave rise to‬
‭Dolly came from an older individual, the age of the DNA may have affected her life‬
‭expectancy. Since Dolly, several species of domestic animals (e.g., horses, cattle, pigs, and‬
‭goats) have been successfully cloned.‬
‭ here have been attempts at producing cloned human embryos as sources of embryonic‬
T
‭stem cells. In the procedure, the DNA from an adult human somatic cell is introduced into‬
‭a human egg cell, which is then stimulated to divide. The technology is similar to the‬
‭technology used to produce Dolly, but the embryo is never implanted into a surrogate‬
‭carrier. The cells produced are called‬‭embryonic stem‬‭cells (ESCs)‬‭because they can‬
‭develop into many different kinds of cells, such as muscle or nerve cells. The stem cells‬
‭could be used for research and ultimately provide therapeutic applications, such as‬
‭replacing damaged tissues. The benefit of cloning in this instance is that the cells used to‬
‭regenerate new tissues would be a perfect match to the donor of the original DNA. For‬
‭example, a leukemia patient would not require a sibling with a tissue match for a bone‬
‭marrow transplant. However, producing embryos as a source of ESCs is controversial‬
‭because the embryo could develop into a fetus if it were implanted into a surrogate‬
‭female.‬

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‭ igure 22.9‬ ‭Dolly the sheep was the first agricultural‬‭animal to be cloned. To create‬
F
‭Dolly, the nucleus was removed from a donor egg cell. The enucleated egg was placed next‬
‭to the other cell, and they were shocked to fuse. They were shocked again to start division.‬
‭The cells were allowed to divide for several days until an early embryonic stage was‬
‭reached before being implanted in a surrogate mother. (credit:‬‭OpenStax‬‭); a link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ or additional information about Dolly the sheep and the process used to produce her,‬
F
‭click the link or scan the QR code to watch the Henrick’s Lab video titled “The story of‬
‭Dolly the cloned sheep - animal cloning”.‬

The Story of Dolly the cloned Sheep - Animal Cloning

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‭DNA Sequencing‬
‭ NA sequencing‬‭is a process used to determine the‬‭specific order of nucleotide bases‬
D
‭(i.e., A, T, G, and C) in a segment of DNA. The basic sequencing technique used in all‬
‭modern-day sequencing projects is the‬‭c hain termination‬‭method‬‭(also known as the‬
‭dideoxy method), which Fred Sanger developed in the 1970s. This method involves DNA‬
‭replication of a single-stranded template using a primer and regular‬‭deoxynucleotides‬
‭(dNTPs), the monomer units of DNA. There are four types of dNTPs, with N representing‬
‭o ne of the four bases in DNA (i.e., dATP, dTTP, dGTP, and dCTP). The primer and dNTPs‬
‭are mixed with a small proportion of fluorescently labeled‬‭dideoxynucleotides‬‭(ddNTPs;‬
‭ddATP, ddTTP, ddGTP, and ddCTP). The ddNTPs are monomers missing a hydroxyl group‬
‭(–OH) on the 3’ carbon, which is the site at which another nucleotide usually attaches to‬
‭form a chain (‬‭Figure 22.10‬‭).‬
‭ very time a ddNTP incorporates into the growing complementary strands, it terminates‬
E
‭the DNA replication process, which results in multiple short strands of replicated DNA that‬
‭each terminates at a different point. Electrophoresis is then used to separate the‬
‭numerous newly replicated DNA strands by size. Because the ddNTPs are fluorescently‬
‭labeled with other colors, the specific one that terminated the reaction can be determined.‬
‭A computer analyzes the colors and generates a pattern that reflects the order of bases‬
‭(‬‭Figure 22.11‬‭). This DNA sequence is complementary‬‭to the template strand’s sequence.‬
I‭ n 1989, scientists formally began to discuss the application of DNA sequencing to the‬
‭human genome. In 1991, the Human Genome Project (HGP) began. This international‬
‭collaborative research effort included groups from twenty separate research centers and‬
‭universities in six countries: the United States, the United Kingdom, China, Germany,‬
‭France, and Japan. The project took 13 years to complete and cost approximately three‬
‭billion dollars. In 2000, the first draft of the sequence results, estimated to include 90‬
‭percent of the genome, was published. Three years later, the complete genome sequence‬
‭was published.‬

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‭ igure 22.10‬ ‭A dideoxynucleotide is similar in structure‬‭to a deoxynucleotide, but is‬

F
‭missing the 3' hydroxyl group (indicated by the box). When a dideoxynucleotide is‬
‭incorporated into a DNA strand, DNA synthesis stops. (credit:‬‭OpenStax‬‭); A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭ igure 22.11‬ ‭This figure illustrates Frederick‬‭Sanger's dideoxy chain termination‬

F
‭method. Using dideoxynucleotides, the synthesized DNA fragments will terminate at‬
‭different points/lengths. The DNA is separated based on size (bp), and the colors‬
‭associated with the ddNTPs are interpreted to determine the DNA sequence. In the figure‬
‭above, 120 and 130 represent the nucleotide number in the DNA sequence; only a small‬
‭portion of the entire sequence is shown. (credit:‬‭OpenStax‬‭); A link to a video explanation‬
‭o f this figure is available at‬‭Biology411.com‬‭.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ or additional information about the HGP, click the links or scan the QR codes below to‬
F
‭watch Ted-Ed videos by Mark J. Keil and Tien Nguyen, titled “How to sequence the human‬
‭genome,” and “The race to sequence the human genome,” respectively.‬

How to sequence the human genome - Mark J. Kiel

The race to sequence the human genome - Tien Nguyen

‭ o review many of the molecular biology techniques discussed thus far, click the link or‬
T
‭scan the QR code to watch the Bozeman Science video titled “Molecular Biology.”‬

Molecular Biology

‭22.3 Biotechnology in Medicine and Agriculture‬

S‭ ince the discovery of the structure of DNA in 1953, and particularly since the‬
‭development of tools and methods to manipulate DNA in the 1970s, biotechnology has‬
‭become synonymous with manipulating organisms’ DNA at the molecular level. Adding‬
‭foreign DNA in the form of recombinant DNA vectors generated by bacterial cloning is the‬
‭most common method of manipulation. Specific applications of biotechnology are‬
‭discussed in the following sections.‬

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‭Genetic Diagnosis and Gene Therapy‬

‭ he process of testing for suspected genetic defects before administering treatment is‬
T
‭called‬‭genetic diagnosis by genetic testing‬‭. In cases‬‭where a genetic disease is present in‬
‭an individual’s family, members may be advised to undergo genetic testing. For example,‬
‭mutations in the‬‭BRCA‬‭genes may increase the likelihood‬‭o f developing breast, ovarian,‬
‭and some other cancers. A person with breast cancer can be screened for these‬
‭mutations. If one of the high-risk mutations is found, relatives (males and females) may‬
‭also wish to be screened for that particular mutation or simply be more vigilant for the‬
‭o ccurrence of cancers. Genetic testing is also offered for fetuses (or embryos with in vitro‬
‭fertilization) to determine the presence or absence of disease-causing genes in families‬
‭with specific debilitating diseases.‬
‭ ene therapy‬‭is a genetic engineering technique that‬‭may one day be commonly used to‬
G
‭cure various genetic diseases. In its simplest form, gene therapy involves the introduction‬
‭o f a non-mutated gene at a random location in the genome to cure a disease by replacing‬
‭a protein that may be absent in these individuals because of a genetic mutation. The‬
‭non-mutated gene is usually introduced into diseased cells as part of a vector transmitted‬
‭by a‬‭virus‬‭, such as an adenovirus, that can infect‬‭the host cell and deliver the foreign DNA‬
‭into the genome of the targeted cells (‬‭Figure 22.12‬‭).‬‭To date, gene therapies have been‬
‭primarily experimental procedures in humans. A few of these experimental treatments‬
‭have been successful, but the methods may be necessary in the future as the factors‬
‭limiting their success are resolved.‬

‭Production of Vaccines, Antibiotics, and Hormones‬

‭ raditional vaccination strategies use weakened or inactive forms of microorganisms or‬
T
‭viruses to stimulate the immune system. Modern techniques use specific genes of‬
‭microorganisms cloned into vectors and mass-produced in bacteria to make large‬
‭quantities of particular substances to stimulate the immune system. The substance is then‬
‭used as a vaccine. In some cases, such as the H1N1 flu vaccine, genes cloned from the‬
‭virus have been used to combat the constantly changing strains of this virus.‬
‭ ntibiotics kill bacteria and are naturally produced by microorganisms such as fungi;‬
A
‭penicillin is perhaps the most well-known example. Antibiotics are made on a large scale‬
‭by cultivating and manipulating fungal cells. The fungal cells have typically been genetically‬
‭modified to improve the yields of the antibiotic compound.‬
‭ ecombinant DNA technology was used to produce large-scale quantities of the human‬
R
‭hormone insulin in‬‭E. coli‬‭as early as 1978. Previously,‬‭it was only possible to treat‬
‭diabetes with pig insulin, which caused allergic reactions in many humans because of‬
‭differences in the insulin molecule. In addition, human growth hormone (HGH) is used to‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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t‭ reat growth disorders in children. The HGH gene was cloned and inserted into‬‭E. coli‬‭cells‬
‭via a bacterial vector.‬

‭ igure 22.12‬ ‭This diagram shows the steps in curing‬‭disease with gene therapy using an‬
F
‭adenovirus vector. (credit: modification of work by NIH;‬‭OpenStax‬‭); A link to a video‬
‭explanation of this figure is available at‬‭Biology411.com‬‭.‬

‭Transgenic Animals‬
I‭ n biotechnology, the organism that receives the recombinant DNA is called a‬‭genetically‬
‭modified organism‬‭(‬‭GMO‬‭). The host organism is transgenic‬‭if the foreign DNA comes‬
‭from a different species. Therefore, not all GMOs are transgenic. However, all transgenics‬
‭are GMOs.‬
‭ lthough several proteins used in medicine are successfully produced in bacteria, some‬
A
‭proteins need a eukaryotic animal host for proper processing. For this reason, genes have‬
‭been cloned and expressed in animals such as sheep, goats, chickens, and mice (‬‭Figure‬
‭22.13‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 22.13‬‭Two of these mice are transgenic because‬‭they have a gene that causes‬
F
‭them to fluoresce under UV light. The non-transgenic mouse does not have the gene that‬
‭causes fluorescence. (credit: Ingrid Moen et al.;‬‭OpenStax‬‭)‬

S‭ everal human proteins are expressed in the milk of transgenic sheep and goats. In one‬
‭commercial example, the FDA has approved a blood anticoagulant protein produced in the‬
‭milk of transgenic goats for use in humans.‬

‭Transgenic Plants‬
‭ anipulating the DNA of plants, such as corn, wheat, potatoes, and tomatoes, has helped‬
M
‭create desirable traits such as disease resistance, herbicide resistance, better nutritional‬
‭value, and better shelf life (‬‭Figure 22.14‬‭). These‬‭changes have been valuable because‬
‭plants are the most important food source for the human population.‬
‭ ecause transgenic plants contain unique combinations of genes and are not restricted to‬
B
‭the laboratory, they, along with other GMOs, are closely monitored by government‬
‭agencies to ensure that they are fit for human consumption and do not endanger other‬
‭plant and animal life. Because foreign genes can spread to other species in the‬
‭environment, particularly in the pollen and seeds of plants, extensive testing is required to‬
‭ensure ecological stability.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 22.14‬ ‭Corn, a major agricultural crop used‬‭to create products for various‬
F
‭industries, is often modified through plant biotechnology. (credit: Keith Weller, USDA;‬
‭OpenStax‬‭)‬

‭ or information about the controversy surrounding GMOs, click the link or scan the QR‬
F
‭code below to watch Kurzgesagt's video‬‭In a Nutshell‬‭,‬‭titled “Are GMOs Good or Bad?‬
‭Genetic Engineering and Our Food.”‬

Are GMOs Good or Bad? Genetic Engineering & Our Food

‭ or additional information about biotechnology and applications, click the link below or‬
F
‭scan the QR code to watch the Amoeba Sisters’ video titled “Genetic Engineering.”‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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Genetic Engineering

‭22.4 Genomics and Its Applications‬

‭ he study of nucleic acids began with the discovery of DNA, progressed to the study of‬
T
‭genes and small fragments, and has now exploded into the field of genomics.‬‭Genomics‬‭is‬
‭the study of entire genomes, including the complete set of genes, their nucleotide‬
‭sequence and organization, and their interactions within a species and with other species.‬
‭The advances in genomics have been made possible by DNA sequencing technology. Just‬
‭as information technology has led to Google Maps, which enables us to get detailed‬
‭information about locations around the globe, genomic data is used to create similar maps‬
‭o f the DNA of different organisms. These resources are now being used in various fields,‬
‭as discussed in the following sections.‬

‭Predicting Disease Risk at the Individual Level‬

‭ redicting disease risk involves screening currently healthy individuals by genome‬
P
‭analysis at the individual level. Health care professionals can recommend interventions,‬
‭such as lifestyle changes and drugs, before disease onset. However, this approach is most‬
‭applicable when the problem resides within a single gene defect. Such defects only‬
‭account for approximately 5 percent of diseases in developed countries. Most common‬
‭diseases, such as heart disease, are multi-factored or‬‭polygenic‬‭, a phenotypic‬
‭characteristic involving two or more genes, as well as environmental factors such as diet.‬
‭In April 2010, scientists at Stanford University published the genome analysis of a healthy‬
‭individual (Stephen Quake, a scientist at Stanford University) who had his genome‬
‭sequenced. The study predicted his propensity to acquire various diseases. The medical‬
‭team performed a risk assessment to analyze Quake’s risk percentage for 55 different‬
‭medical conditions. The team found a rare genetic mutation, which showed him to be at‬
‭risk for sudden heart attack. The results also predicted that Quake had a 23 percent risk of‬
‭developing prostate cancer and a 1.4 percent risk of developing Alzheimer’s. The scientists‬
‭used databases and several publications to analyze the genomic data. Even though‬
‭genomic sequencing is becoming more affordable and analytical tools are becoming more‬
‭reliable, researchers still must address ethical issues surrounding genomic analysis at a‬
‭population level.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Gene Editing‬
‭ or thousands of years, humans have engaged in some level of control over genes and‬
F
‭heredity regarding the plants and animals we rely on. The technology now exists to exert‬
‭that control more directly by altering organisms' DNA. The technique is usually called‬
‭CRISPR‬‭, for the portions of DNA it targets: "‬‭C‬‭lustered‬‭R‬‭egularly‬‭I‬‭nterspaced‬‭S‬‭hort‬
‭P‬‭alindromic‬‭R‬‭epeats." In essence, DNA contains repetitive‬‭sequences with "spacers"‬
‭between them. CRISPR-associated nucleases (known as "Cas") are enzymes that can‬
‭identify, attach to, and cut DNA strands at precise locations (‬‭Figure 22.15‬‭). In 2012,‬
‭Jennifer Doudna and Emmanuelle Charpentier developed a method to combine the Cas‬
‭nuclease with a synthetically produced "guide RNA" that leads the nuclease to selected‬
‭locations on the DNA strand. The discovery revolutionized gene editing. Researchers‬
‭worldwide have used CRISPR to manipulate the DNA of plants, animals, laboratory cell‬
‭lines, and (in trials) human patients. In 2020, Doudna and Charpentier were awarded the‬
‭Nobel Prize for their work (‬‭Figure 22.16‬‭).‬

‭ igure 22.15‬ ‭A general overview of how an organism’s‬‭genome can be edited using the‬
F
‭CRISPR/Cas-9 method. (credit: Bartz/Stockmar,‬‭Agrifood‬‭Atlas‬‭, Wikimedia Commons;‬‭CC‬
‭BY-SA 4.0‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 22.16‬ ‭Emmanuelle Charpentier and Jennifer‬‭Doudna, awardees of the 2020‬

F
‭Nobel Prize in Chemistry for their work on the CRISPR-Cas 9 DNA editing system. (credit:‬
‭Bianca Fioretti of Hallbauer & Fioretti;‬‭Duncan.Hull‬‭and The Royal Society;‬‭Wikimedia‬
‭Commons‬‭;‬‭CC BY-SA 4.0‬‭)‬

‭ ene editing is so promising because it can be used experimentally to understand disease‬

G
‭o r organismal limitations and then applied to overcome those issues. For example, in a‬
‭human trial, cancerous cells were removed from a person, edited to remove their‬
‭cancerous properties at the DNA level, and reintroduced into the patient so that those‬
‭now edited cells could multiply and replace the cancerous ones. Using the person's own‬
‭cells increases the likelihood of acceptance and success.‬
‭ ike other genomics applications, the prospect of directly editing genes raises ethical‬
L
‭issues. Doudna and Chapentier, as well as many other genetic engineers, support only‬
‭certain CRISPR applications. Many governments and other entities have placed strict‬
‭guidelines on the use of this powerful technology.‬
‭ or additional information about CRISPR and its uses, click the link or scan the QR code‬
F
‭below to watch the Ted-Ed video by Andrea M. Henle titled “How CRISPR lets you edit‬
‭DNA.”‬

How CRISPR lets you edit DNA - Andrea M. Henle

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Pharmacogenomics‬
‭ harmacogenomic‬‭s involves evaluating drug effectiveness‬‭and safety based on‬
P
‭information from an individual's genomic sequence. We can study genomic responses to‬
‭drugs using experimental animals (such as laboratory rats or mice) or live cells in the‬
‭laboratory before embarking on studies with humans. Studying changes in gene‬
‭expression could provide information about the transcription profile in the drug's‬
‭presence, which can be used as an early indicator of the potential for toxic effects. For‬
‭example, when disturbed, genes involved in cellular growth and controlled cell death‬
‭could lead to cancerous cell growth. Genome-wide studies can also help to find new genes‬
‭involved in drug toxicity. Medical professionals can use personal genome sequence‬
‭information to prescribe medications that will be most effective and least toxic based on‬
‭the individual patient’s genotype. The gene signatures may not be completely accurate, but‬
‭medical professionals can test them further before pathology symptoms arise.‬
‭ or additional information about pharmacogenomics, click the link or scan the QR code‬
F
‭below to watch the Mayo Clinic video titled “Pharmacogenomics: The right drug, for the‬
‭right patient, at the right dose.”‬

Pharmacogenomics: The Right Drug,…

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭Chapter 23: Ecology, Populations, and Biodiversity‬

‭Chapter 23: Ecology, Populations, and Biodiversity‬

‭ igure 23.1‬ ‭An Australian Institute of Marine Sciences‬‭(AIMS) researcher inspects a‬

F
‭bleached coral on the Great Barrier Reef. AIMS is part of an international research‬
‭collaboration to help predict how reef-building corals will respond to climate change.‬
‭(credit: Eric Matson; AIMS; “‬‭Can Corals Survive Climate‬‭Change?‬‭”)‬

‭23.1 Introduction‬
‭ ife in an ecosystem is often about competition for limited resources, a characteristic of‬
L
‭the theory of natural selection. Competition in‬‭communities‬‭(all living things within‬
‭specific habitats) is observed both within species and among species. The resources for‬
‭which organisms compete include organic material, sunlight, and mineral nutrients, which‬
‭provide the energy for living processes and the matter that makeup organisms’ physical‬
‭structures. Other critical factors influencing community dynamics are the components of‬
‭its physical and geographic environment: a habitat’s latitude, amount of rainfall,‬
‭topography (elevation), and available species. These are all critical environmental‬

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P
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‭ ariables determining which organisms can exist within a particular area. Understanding‬
v
‭the tremendous variety of living species will help us better understand how to conserve‬
‭the diversity of life on Earth.‬

‭23.2 Ecosystems and Energy‬

‭ n‬‭ecosystem‬‭is a community of living organisms and‬‭their abiotic (non-living)‬
A
‭environment;‬‭ecology‬‭can be defined as the study of‬‭ecosystems. Ecosystems are‬
‭tremendously diverse. They can be small, such as the tide pools found near the rocky‬
‭shores of many oceans, or large, such as those found in the tropical rainforest of the‬
‭Amazon in Brazil (‬‭Figure 23.2‬‭).‬

‭ igure 23.2‬ ‭A (a) tidal pool ecosystem in Matinicus‬‭Island, Maine, is a small ecosystem,‬
F
‭while the (b) Amazon rainforest in Brazil is a large ecosystem. (credit a: modification of‬
‭work by Jim Kuhn; credit b: modification of work by Ivan Mlinaric;‬‭OpenStax‬‭)‬

‭ here are three broad categories of ecosystems based on their general environment:‬
T
‭freshwater, marine, and terrestrial. Within these three categories are individual ecosystem‬
‭types based on the environmental habitat and organisms present:‬
‭ reshwater ecosystems‬‭are the least common, occurring‬‭o n only 1.8 percent of Earth's‬
F
‭surface. These systems comprise lakes, rivers, streams, and springs. They are quite diverse‬
‭and support a variety of animals, plants, fungi, protists, and prokaryotes.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ arine ecosystems‬‭are the most common, comprising 75 percent of Earth's surface and‬
M
‭consisting of three basic types: shallow ocean, deep ocean water, and deep ocean bottom.‬
‭Shallow ocean ecosystems include extremely biodiverse coral reef ecosystems, yet the‬
‭deep ocean water is known for the large numbers of plankton and krill (small‬
‭crustaceans) that support it. These two environments are critical to aerobic respirators‬
‭worldwide, as phytoplankton perform 40 percent of all photosynthesis on Earth. Although‬
‭not as diverse as the other two, deep ocean bottom ecosystems still contain a variety of‬
‭marine organisms. Such ecosystems exist even at depths where light cannot penetrate the‬
‭water.‬
‭ errestrial ecosystems‬‭, also known for their diversity,‬‭are grouped into large categories‬
T
‭called biomes. A‬‭biome‬‭is a large-scale community‬‭o f organisms primarily defined on land‬
‭by the dominant plant types that exist in geographic regions of the planet with similar‬
‭climatic conditions. Examples of biomes include tropical rainforests, savannas, deserts,‬
‭grasslands, temperate forests, and tundras. Grouping these ecosystems into just a few‬
‭biome categories obscures the great diversity of the individual ecosystems within them.‬
‭For example, the saguaro cacti (‬‭Carnegiea gigantean‬‭)‬‭and other plant life in the Sonoran‬
‭Desert in the United States are relatively diverse compared with the desolate rocky desert‬
‭o f Boa Vista, an island off the coast of Western Africa (‬‭Figure 23.3‬‭).‬

‭ igure 23.3‬ ‭Desert ecosystems, like all ecosystems,‬‭can vary greatly. The desert in (a)‬
F
‭Saguaro National Park, Arizona, has abundant plant life, while the rocky desert of (b) Boa‬
‭Vista island, Cape Verde, Africa, is devoid of plant life. (credit a: modification of work by‬
‭Jay Galvin; credit b: modification of work by Ingo Wö lbern;‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Ecosystems and Disturbance‬

‭ cosystems are complex, with many interacting parts. They are routinely exposed to‬
E
‭various environmental changes that affect their compositions, such as yearly variations in‬
‭rainfall and temperature. Many disturbances are a result of natural processes. For‬
‭example, when lightning causes a forest fire and destroys part of a forest ecosystem, the‬
‭ground is eventually populated with grasses, followed by bushes and shrubs, and later‬
‭mature trees; thus, the forest is restored to its former state. This process is so universal‬
‭that ecologists have named it‬‭succession‬‭. The impact‬‭o f environmental disturbances‬
‭caused by human activities is now as significant as the changes wrought by natural‬
‭processes. Human agricultural practices, air pollution, acid rain, global deforestation,‬
‭overfishing, oil spills, and illegal dumping on land and into the ocean all impact‬
‭ecosystems.‬
‭ quilibrium is a dynamic state of an ecosystem in which biodiversity remains somewhat‬
E
‭constant despite changes in species numbers and occurrence. In ecology, two parameters‬
‭measure ecosystem changes: resistance and resilience.‬ ‭Resistance‬‭is the ability of an‬
‭ecosystem to remain at equilibrium despite disturbances.‬ ‭Resiliency‬‭is the speed at which‬
‭an ecosystem recovers equilibrium after being disturbed. Ecosystem resistance and‬
‭resilience are especially important when considering human impact. The nature of an‬
‭ecosystem may change to such a degree that it can lose its resilience entirely. This process‬
‭can lead to the irreversible altering of an ecosystem.‬

‭Food Chains and Food Webs‬

‭ ‬‭food chain‬‭is a linear sequence of organisms through‬‭which nutrients and energy pass‬
A
‭as one organism eats another; the levels in the food chain are producers, primary‬
‭consumers, higher-level consumers, and decomposers. There is a single path through a‬
‭food chain. Each organism in a food chain occupies a specific‬‭trophic level‬‭(energy level),‬
‭which is its position in the food chain or food web.‬
‭ ood chains' base or foundation in many ecosystems consists of photosynthetic organisms‬
F
‭(plants or phytoplankton) called‬‭producers‬‭. The organisms‬‭that consume the producers‬
‭are‬‭herbivores‬‭(i.e., the‬‭primary consumers)‬‭.‬‭Secondary‬‭consumers‬‭are usually‬
‭c arnivores‬‭that eat the primary consumers.‬‭Tertiary‬‭consumers‬‭are carnivores that eat‬
‭o ther carnivores. Higher-level consumers feed on the next lower trophic levels, and so on,‬
‭up to the organisms at the top of the food chain, containing the‬‭apex consumers‬‭. In the‬
‭Lake Ontario food chain, shown in‬‭Figure 23.4‬‭, the‬‭Chinook salmon is the apex consumer‬
‭at the top of this food chain.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 23.4‬ ‭These are the trophic levels of a food‬‭chain in Lake Ontario at the United‬
F
‭States–Canada border. Energy and nutrients flow from photosynthetic green algae at the‬
‭base to the top of the food chain: the Chinook salmon. (credit: modification of work by‬
‭National Oceanic and Atmospheric Administration/NOAA;‬ ‭OpenStax‬‭)‬

‭ nergy is one major factor limiting the number of steps in a food chain. Energy is lost at‬
E
‭each trophic level and between trophic levels as heat and in the transfer to decomposers‬
‭(Chapter 6; Second Law of Thermodynamics;‬‭Figure 23.5‬‭).‬‭Thus, after a limited number of‬
‭trophic energy transfers, the amount of energy remaining in the food chain may need to‬
‭be greater to support viable populations at a higher trophic level.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
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‭ igure 23.5‬ ‭The relative energy in trophic levels‬‭in a Silver Springs, Florida, ecosystem is‬
F
‭shown. Each trophic level has less energy available and usually, but only sometimes,‬
‭supports a smaller mass of organisms at the next level. (credit:‬‭OpenStax‬‭)‬

‭ here is one problem when using food chains to describe most ecosystems. Even when all‬
T
‭o rganisms are grouped into appropriate trophic levels, some can feed on more than one‬
‭trophic level; likewise, some can also be fed on from multiple trophic levels. In addition,‬
‭species feed on and are eaten by more than one species. In other words, the linear model‬
‭o f ecosystems, the food chain, is a hypothetical, overly simplistic representation of‬
‭ecosystem structure. A holistic model—including all the interactions between different‬
‭species and their complex interconnected relationships with each other and the‬
‭environment—is a more accurate and descriptive model for ecosystems. A‬‭food web‬‭is a‬
‭concept that accounts for the multiple trophic (feeding) interactions between each species‬
‭and the many species it may feed on or that feed on it. In a food web, the several trophic‬
‭connections between each species and the other species that interact with it may cross‬
‭multiple trophic levels. Food webs describe virtually all ecosystems' matter and energy‬
‭movements more accurately (‬‭Figure 23.6‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
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‭ igure 23.6‬ ‭This food web shows the interactions‬‭between organisms across trophic‬
F
‭levels. Arrows point from an organism that is consumed to the organism that consumes it.‬
‭All the producers and consumers eventually become nourishment for the decomposers‬
‭(fungi, mold, earthworms, and bacteria in the soil). (credit "fox": modification of work by‬
‭Kevin Bacher, NPS; credit "owl": modification of work by John and Karen Hollingsworth,‬
‭USFWS; credit "snake": modification of work by Steve Jurvetson; credit "robin":‬
‭modification of work by Alan Vernon; credit "frog": modification of work by Alessandro‬
‭Catenazzi; credit "spider": modification of work by "Sanba38"/Wikimedia Commons; credit‬
‭"centipede": modification of work by “Bauerph”/Wikimedia Commons; credit "squirrel":‬
‭modification of work by Dawn Huczek; credit "mouse": modification of work by NIGMS,‬
‭NIH; credit "sparrow": modification of work by David Friel; credit "beetle": modification of‬
‭work by Scott Bauer, USDA Agricultural Research Service; credit "mushrooms":‬
‭modification of work by Chris Wee; credit "mold": modification of work by Dr. Lucille‬
‭Georg, CDC; credit "earthworm": modification of work by Rob Hille; credit "bacteria":‬
‭modification of work by Don Stalons, CDC;‬ ‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ wo general types of food webs are often shown interacting within a single ecosystem. A‬
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‭grazing food web‬‭has plants or other photosynthetic‬‭o rganisms at its base, followed by‬
‭herbivores and various carnivores. A‬‭detrital food‬‭web‬‭consists of a base of organisms‬
‭that feed on decaying organic matter (dead organisms), including decomposers (which‬
‭break down dead and decaying organisms) and detritivores (which consume organic‬
‭detritus). These organisms are usually bacteria, fungi, and invertebrate animals that‬
‭recycle organic material back into the biotic part of the ecosystem as other organisms‬
‭consume them. As ecosystems require a method to recycle material from dead organisms,‬
‭grazing food webs have an associated detrital food web. For example, in a meadow‬
‭ecosystem, plants may support a grazing food web of different organisms, primary and‬
‭o ther levels of consumers, while at the same time supporting a detrital food web of‬
‭bacteria and fungi feeding off dead plants and animals. Simultaneously, a detrital food web‬
‭can contribute energy to a grazing food web, as when a robin eats an earthworm.‬

‭ or additional information about the importance of detrital food webs, click the link or‬
F
‭scan the QR code to watch the TED-Ed video by John C. Moore titled “The secret ingredient‬
‭in our food chain.”‬

Dead stuff: The secret ingredient in our food chain - John C. Mo…

‭How Organisms Acquire Energy in a Food Web‬

‭ ll living things require energy in one form or another. Energy is used by most complex‬
A
‭metabolic pathways (usually in the form of‬‭ATP‬‭), especially‬‭those responsible for building‬
‭large molecules from smaller compounds. Without a constant energy input, living‬
‭o rganisms could not assemble macromolecules (proteins, lipids, nucleic acids, and‬
‭complex carbohydrates) from their monomers.‬
‭ ood web diagrams illustrate how energy flows directionally through ecosystems. They‬
F
‭can also indicate how efficiently organisms acquire and use energy and how much‬
‭remains for use by other organisms of the food web. Energy is acquired by living things in‬
‭two ways: autotrophs harness light or chemical energy, and heterotrophs acquire energy‬
‭through the consumption and digestion of other living or previously living organisms.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ hotosynthetic and chemosynthetic organisms are‬‭autotrophs‬‭capable of synthesizing‬

P
‭their food (specifically, capable of using inorganic carbon as a carbon source).‬
‭Photosynthetic autotrophs (‬‭photoautotrophs‬‭) use sunlight‬‭as an energy source, and‬
‭chemosynthetic autotrophs (‬‭c hemoautotrophs‬‭) use inorganic‬‭molecules as an energy‬
‭source. Autotrophs are critical for most ecosystems: they are the producers trophic level.‬
‭Without these organisms, energy would not be available to other living organisms, and life‬
‭itself would not be possible.‬
‭ hotoautotrophs, such as plants, algae, and photosynthetic bacteria, are the energy source‬
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‭for most of the world’s ecosystems. These ecosystems are often described by grazing and‬
‭detrital food webs. Photoautotrophs harness the Sun’s solar energy by converting it to‬
‭chemical energy as ATP. The energy stored in ATP is used to synthesize complex organic‬
‭molecules, such as glucose. The rate at which photosynthetic producers incorporate‬
‭energy from the Sun is called‬‭gross primary productivity‬‭.‬‭However, not all the energy‬
‭producers incorporate is available to other organisms on the food web because‬
‭producers must also grow and reproduce, which consumes energy.‬‭Net primary‬
‭productivity‬‭is the energy that remains in the producers‬‭after accounting for these‬
‭o rganisms’ respiration and heat loss. The net productivity is then available to the primary‬
‭consumers at the next trophic level.‬
‭ hemoautotrophs are primarily bacteria and archaea found in rare ecosystems where‬
C
‭sunlight is unavailable, such as those associated with dark caves or hydrothermal vents at‬
‭the bottom of the ocean (‬‭Figure 23.7‬‭). Many chemoautotrophs‬‭in hydrothermal vents use‬
‭hydrogen sulfide (H‬‭2‭S‬ ), which is released from the‬‭vents, as a source of chemical energy;‬
‭this allows them to synthesize complex organic molecules, such as glucose, for their‬
‭energy and, in turn, supplies energy to the rest of the ecosystem.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.7‬ ‭Swimming shrimp, a few squat lobsters,‬‭and hundreds of vent mussels are‬
F
‭seen at a hydrothermal vent at the bottom of the ocean. As no sunlight penetrates this‬
‭depth, the ecosystem is supported by chemoautotrophic bacteria and organic material‬
‭that sinks from the ocean’s surface. This picture was taken in 2006 at the submerged NW‬
‭Eifuku volcano off the coast of Japan by the National Oceanic and Atmospheric‬
‭Administration (NOAA). The summit of this highly active volcano lies 1535 m below the‬
‭surface. (credit:‬‭OpenStax‬‭)‬

‭23.3 Biogeochemical Cycles‬

‭ nergy flows directionally through ecosystems, entering as sunlight (or inorganic‬
E
‭molecules for chemoautotrophs) and leaving as heat during the transfers between trophic‬
‭levels. Rather than flowing through an ecosystem, the matter that makes up living‬
‭o rganisms is conserved and recycled. The six most common elements associated with‬
‭o rganic molecules—carbon, nitrogen, hydrogen, oxygen, phosphorus, and sulfur—take‬
‭various chemical forms and may exist for long periods in the atmosphere, on land, in‬
‭water, or beneath Earth’s surface. Geologic processes, such as weathering, erosion, water‬
‭drainage, and the subduction of the continental plates, all play a role in the cycling of‬
‭elements on Earth. Because geology and chemistry have significant roles in the study of‬
‭this process, the recycling of inorganic matter between living organisms and their‬
‭nonliving environment is called a‬‭biogeochemical cycle‬‭.‬ ‭Three examples of these cycles‬
‭(water, carbon, and nitrogen) are discussed in the following sections.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭The Water Cycle‬

‭ ater, which contains hydrogen and oxygen, is essential to all living processes. The‬
W
‭hydrosphere‬‭is the area of Earth where water movement‬‭and storage occur as liquid‬
‭water on the surface (rivers, lakes, oceans), beneath the surface (groundwater), as ice‬
‭(polar ice caps and glaciers), and as water vapor in the atmosphere.‬
‭ he human body is more than one-half water, and human cells are more than 70 percent‬
T
‭water. Thus, most land animals need a supply of freshwater to survive. Of the stores of‬
‭water on Earth, 97.5 percent is‬‭saltwater‬‭(‬‭Figure‬‭23.8‬‭). Of the remaining water, 99‬
‭percent is locked as underground water or ice. Thus, less than one percent of‬‭freshwater‬
‭is present in lakes and rivers. Many living things depend on this small amount of surface‬
‭fresh water supply, a lack of which can significantly affect ecosystem dynamics. Humans‬
‭have developed technologies to increase water availability, such as digging wells to harvest‬
‭groundwater, storing rainwater, and using desalination to obtain drinkable water from the‬
‭o cean. Although this pursuit of drinkable water has been ongoing throughout human‬
‭history, freshwater supply remains a significant issue.‬

‭ igure 23.8‬ ‭Only 2.5 percent of water on Earth‬‭is freshwater, and less than 1 percent of‬
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‭freshwater is easily accessible to living things. (credit:‬‭OpenStax‬‭)‬

‭ or additional information about the hydrosphere, click the link or scan the QR code to‬
F
‭watch the TED-Ed video by Christiana Z. Peppard titled “Why is biodiversity so‬
‭important?”.‬

Where we get our fresh water - Christiana Z. Peppard

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ he various processes that occur during the cycling of water are illustrated in‬‭Figure 23.9‬‭.‬
T
‭The processes include the following:‬
•‭ ‬ e‭ vaporation and sublimation‬
‭•‬ ‭condensation and precipitation‬
‭•‬ ‭subsurface water flow‬
‭•‬ ‭surface runoff and snowmelt‬
‭•‬ ‭streamflow‬
‭ he Sun’s energy drives the water cycle as it warms the oceans and other surface waters.‬
T
‭This leads to‬‭evaporation‬‭(water to water vapor) of‬‭liquid surface water and‬‭sublimation‬
‭(ice to water vapor) of frozen water, thus moving large amounts of water into the‬
‭atmosphere as water vapor. Over time, this water vapor condenses (i.e.,‬‭condensation‬‭)‬
‭into clouds as liquid or frozen droplets and eventually leads to‬‭precipitation‬‭(rain or‬
‭snow), which returns water to the Earth’s surface. Rain reaching Earth’s surface may‬
‭evaporate again, flow over the surface, or percolate into the ground. Most easily observed‬
‭is surface runoff: the flow of fresh water either from rain or melting ice. Runoff can make‬
‭its way through streams and lakes to the oceans or flow directly to the oceans themselves.‬
I‭ n most natural terrestrial environments, rain encounters vegetation before it reaches the‬
‭soil surface. A significant percentage of water evaporates immediately from the surfaces of‬
‭plants. What is left reaches the soil and begins to move down. Surface runoff will only‬
‭o ccur if the soil becomes saturated with water in heavy rainfall. Plant roots will take up‬
‭most of the water in the soil. The plant will use some of this water for its metabolism, and‬
‭some of that will find its way into animals that eat the plants, but much of it will be lost‬
‭back to the atmosphere through a process known as‬‭evapotranspiration‬‭. Water enters‬
‭the plant's vascular system through the roots and evaporates, or transpires, through‬
‭o penings in the leaves. Water in the soil not taken up by a plant and that does not‬
‭evaporate can percolate into the subsoil and bedrock to groundwater.‬
‭ roundwater‬‭is a significant reservoir of fresh water.‬‭It exists in the pores between sand‬
G
‭and gravel particles or in rocks' fissures. Shallow groundwater flows slowly through these‬
‭pores and fissures and eventually finds its way to a stream or lake, where it becomes a‬
‭part of the surface water again. Streams do not flow because they are replenished from‬
‭rainwater directly; they flow because there is a constant inflow from groundwater below.‬
‭Some groundwater is deep in the bedrock and can persist there for millennia. Most‬
‭groundwater reservoirs, or aquifers, are the source of drinking or irrigation water drawn‬
‭up through wells. In many cases, these aquifers are being depleted faster than being‬
‭replenished by water percolating down from above.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.9‬ ‭Water from the land and oceans enters‬‭the atmosphere by evaporation or‬
F
‭sublimation, condensing into clouds and falling as rain or snow. Precipitated water may‬
‭enter freshwater bodies or infiltrate the soil. The cycle is complete when surface or‬
‭groundwater reenters the ocean. (credit: modification of work by John M. Evans and‬
‭Howard Perlman, USGS;‬‭OpenStax‬‭)‬

‭The Carbon Cycle‬

‭ arbon is the fourth most abundant element in living organisms. Carbon is present in all‬
C
‭o rganic molecules, and its role in the structure of macromolecules is of primary‬
‭importance to living organisms. Carbon compounds contain energy, and many of these‬
‭compounds from plants and algae have remained stored as fossilized carbon, which‬
‭humans use as fuel. Since the 1800s, the use of fossil fuels has accelerated. As global‬
‭demand for Earth’s limited fossil fuel supplies has risen since the beginning of the‬
‭Industrial Revolution, the amount of carbon dioxide in our atmosphere has increased as‬
‭the fuels are burned. This increase in carbon dioxide has been associated with climate‬
‭change and is a primary environmental concern worldwide.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ he carbon cycle is most easily studied as two interconnected subcycles: one dealing with‬
T
‭rapid carbon exchange among living organisms and the other dealing with the long-term‬
‭cycling of carbon through geologic processes. The carbon cycle is shown in‬‭Figure‬‭23.10‬‭.‬

‭ igure 23.10‬ ‭Carbon dioxide gas exists in the atmosphere‬‭and is dissolved in water.‬
F
‭Photosynthesis converts carbon dioxide gas to organic carbon, and respiration cycles the‬
‭o rganic carbon back into carbon dioxide gas. Long-term organic carbon storage occurs‬
‭when matter from living organisms is buried deep underground and fossilized. Volcanic‬
‭activity and human emissions bring this stored carbon back into the carbon cycle. (credit:‬
‭modification of work by John M. Evans and Howard Perlman, USGS;‬‭OpenStax‬‭)‬

‭The Biological Carbon Cycle‬

‭ iving organisms are connected in many ways, even between ecosystems. An excellent‬
L
‭example of this connection is the carbon exchange between heterotrophs and autotrophs‬
‭within and between ecosystems through atmospheric carbon dioxide. Carbon dioxide is‬
‭the basic building block autotrophs use to build multi-carbon, high-energy compounds like‬
‭glucose. The energy harnessed from the Sun is used by these organisms in‬
‭photosynthesis‬‭to form the covalent bonds that link‬‭carbon atoms.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ hese chemical bonds store this energy for later use in metabolism to produce ATP. Most‬
T
‭terrestrial autotrophs obtain their carbon dioxide directly from the atmosphere, while‬
‭marine autotrophs acquire it in the dissolved form (carbonic acid, HCO‬‭3‭–‬ ‬‭). However,‬
‭oxygen is a byproduct of fixing carbon in organic compounds when carbon dioxide is‬
‭acquired. Photosynthetic organisms maintain approximately 21 percent of the‬
‭atmosphere's oxygen content that we observe today.‬
‭ he partners in biological carbon exchange are the heterotrophs (especially the primary‬
T
‭consumers, largely herbivores). Heterotrophs acquire the high-energy carbon compounds‬
‭from the autotrophs by consuming them and breaking them down by cellular respiration‬
‭to obtain cellular energy (ATP). The most efficient type of respiration,‬‭aerobic‬
‭respiration‬‭, requires oxygen from the atmosphere or‬‭dissolved in water. Thus, oxygen‬
‭and carbon dioxide are constantly exchanged between the autotrophs (which need the‬
‭carbon) and the heterotrophs (which require the oxygen). Autotrophs also respire and‬
‭consume the organic molecules they form using oxygen and releasing carbon dioxide.‬
‭They release more oxygen gas as a waste product of photosynthesis than they use for‬
‭their respiration; therefore, there is excess available for the respiration of other aerobic‬
‭o rganisms. Gas exchange through the atmosphere and water is one way the carbon cycle‬
‭connects all living organisms on Earth.‬

‭The Biogeochemical Carbon Cycle‬

‭ he movement of carbon through land, water, and air is complex, and, in many cases, it‬
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‭o ccurs much more slowly geologically than the movement between living organisms.‬
‭Carbon is stored for long periods in what are known as carbon reservoirs, which include‬
‭the atmosphere, bodies of liquid water (mostly oceans), ocean sediment, soil, rocks‬
‭(including fossil fuels), and Earth’s interior.‬
‭ s stated, the atmosphere is a significant carbon reservoir in the form of carbon dioxide,‬
A
‭which is essential to photosynthesis. The level of carbon dioxide in the atmosphere is‬
‭greatly influenced by the reservoir of carbon in the oceans. The exchange of carbon‬
‭between the atmosphere and water reservoirs influences how much carbon is found in‬
‭each, and each one affects the other reciprocally. Carbon dioxide from the atmosphere‬
‭dissolves in water and, unlike oxygen and nitrogen gas, reacts with water molecules to‬
‭form ionic compounds. Some of these ions combine with calcium ions in the seawater to‬
‭form calcium carbonate (CaCO‬‭3‭)‬ , a major component‬‭o f the shells of marine organisms.‬
‭These organisms eventually form sediments on the ocean floor. Over geologic time, the‬
‭calcium carbonate forms limestone, which comprises the largest carbon reservoir on‬
‭Earth.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ n land, carbon is stored in the soil as organic carbon due to the decomposition of living‬
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‭o rganisms or the weathering of terrestrial rock and minerals. Deeper under the ground,‬
‭at land, and sea are fossil fuels, the anaerobically decomposed remains of plants that take‬
‭millions of years to form. Fossil fuels are considered nonrenewable because their use far‬
‭exceeds their formation rate. A non-renewable resource is either regenerated very slowly‬
‭o r not at all.‬
‭ nother way for carbon to enter the atmosphere is from land (including land beneath the‬
A
‭o cean's surface) by the eruption of volcanoes and other geothermal systems. Carbon is‬
‭released as carbon dioxide when a volcano erupts or from volcanic hydrothermal vents.‬
‭ arbon dioxide is also added to the atmosphere by animal husbandry practices. The large‬
C
‭number of land animals raised to feed Earth’s growing human population results in‬
‭increased atmospheric carbon dioxide levels caused by their respiration. This is another‬
‭example of how human activity indirectly affects biogeochemical cycles in a significant way.‬
‭Although much of the debate about the future effects of increasing atmospheric carbon on‬
‭climate change focuses on fossil fuels, scientists take natural processes, such as volcanoes,‬
‭plant growth, soil carbon levels, and respiration, into account as they model and predict‬
‭the future impact of this increase.‬
‭ or additional information about the carbon cycle and climate change, click the link or‬
F
‭scan the QR code to watch the TED-Ed video by Nathaniel Manning titled “The carbon‬
‭cycle.”‬

The carbon cycle - Nathaniel Manning

‭The Nitrogen Cycle‬

‭ etting nitrogen into the living world is complicated. Plants and phytoplankton are not‬
G
‭equipped to incorporate nitrogen from the atmosphere (which is tightly bonded, triple‬
‭covalent N‬‭2‭)‬ even though this molecule comprises approximately‬‭78 percent of the‬
‭atmosphere. Nitrogen enters the living world via free-living and symbiotic bacteria, which‬
‭incorporate nitrogen into their macromolecules through‬‭nitrogen fixation‬‭(conversion of‬
‭N‬‭2‬‭). Cyanobacteria live in most aquatic ecosystems‬‭where sunlight is present; they play a‬
‭crucial role in nitrogen fixation. Cyanobacteria can use inorganic sources of nitrogen to‬
‭“fix” nitrogen.‬‭Rhizobium‬‭bacteria live symbiotically‬‭in the root nodules of legumes (such‬
‭as peas, beans, and peanuts) and provide them with the organic nitrogen they need.‬
‭Free-living bacteria, such as‬‭Azotobacter‬‭, are also‬‭essential nitrogen fixers.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ rganic nitrogen is vital to studying ecosystem dynamics since the available nitrogen‬
O
‭supply limits many ecosystem processes, such as primary production and decomposition.‬
‭As shown in‬‭Figure 23.11‬‭, the nitrogen that enters‬‭living systems by nitrogen fixation is‬
‭eventually converted from organic nitrogen into nitrogen gas by bacteria. This process‬
‭o ccurs in three steps in terrestrial systems: ammonification, nitrification, and‬
‭denitrification. First, the ammonification process converts nitrogenous waste from living‬
‭animals or the remains of dead animals into ammonium (NH‬‭4‭+‬ ‬ ‭) via certain bacteria and‬
‭fungi. Second, this ammonium is then converted to nitrites (NO‬‭2‭−‬ ‭)‬ by nitrifying bacteria,‬
‭such as‬‭Nitrosomonas‬‭, through nitrification. Subsequently,‬‭similar organisms convert‬
‭nitrites to nitrates (NO‬‭3‭−‬ ‭)‬ . Lastly, denitrification‬‭o ccurs, whereby bacteria, such as‬
‭Pseudomonas‬‭and‬‭Clostridium‬‭, convert the nitrates‬‭into nitrogen gas, thus allowing it to‬
‭re-enter the atmosphere.‬

‭ igure 23.11‬ ‭Nitrogen enters the living world from‬‭the atmosphere through‬
F
‭nitrogen-fixing bacteria. This nitrogen and nitrogenous waste from animals is then‬
‭processed back into gaseous nitrogen by soil bacteria, which also supply terrestrial food‬
‭webs with the organic nitrogen they need. (credit: modification of work by John M. Evans;‬
‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ uman activity can release nitrogen into the environment by two primary means: the‬
H
‭combustion of fossil fuels, which releases different nitrogen oxides, and the use of‬
‭artificial fertilizers (which contain nitrogen and phosphorus compounds) in agriculture,‬
‭which are then washed into lakes, streams, and rivers by surface runoff. Atmospheric‬
‭nitrogen (other than N‬‭2‭)‬ is associated with several‬‭effects on Earth’s ecosystems, including‬
‭the production of acid rain (as nitric acid, HNO‬‭3‭)‬ ‬‭and greenhouse gas effects (as nitrous‬
‭oxide, N‬‭2‬‭O), potentially causing climate change. A‬‭significant impact of fertilizer runoff is‬
‭saltwater and freshwater‬‭eutrophication‬‭, a process‬‭whereby nutrient runoff causes the‬
‭overgrowth of algae and several consequential problems.‬

‭ similar process occurs in the marine nitrogen cycle, where marine bacteria and archaea‬
A
‭perform the ammonification, nitrification, and denitrification processes. Some of this‬
‭nitrogen falls to the ocean floor as sediment, which can then be moved to land in geologic‬
‭time by uplift of Earth’s surface, and thereby incorporated into terrestrial rock. Although‬
‭the movement of nitrogen from rock directly into living systems has been traditionally‬
‭seen as insignificant compared with nitrogen fixed from the atmosphere, a recent study‬
‭showed that this process may be significant and should be included in any analysis of the‬
‭global nitrogen cycle.‬

‭23.4 Population Demographics‬

‭ opulations in ecosystems are dynamic entities. Their size and composition fluctuate due‬
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‭to numerous factors, including seasonal and yearly environmental changes, natural‬
‭disasters such as forest fires and volcanic eruptions, and competition for resources‬
‭between and within species. The statistical study of populations is called demography, a‬
‭set of mathematical tools designed to describe populations and investigate how they‬
‭change. Many of these tools were designed to study human populations. For example, life‬
‭tables, which detail the life expectancy of individuals within a population, were initially‬
‭developed by life insurance companies to set insurance rates. While “demographics” is‬
‭sometimes assumed to mean a study of human populations, all living populations can be‬
‭studied using this approach.‬

‭Population Size and Density‬

‭ opulations are characterized by‬‭population size‬‭(total‬‭number of individuals) and‬
P
‭population density‬‭(number of individuals per unit‬‭area). A population may have many‬
‭individuals that are distributed densely or sparsely. There are also populations with small‬
‭numbers of individuals that may be dense or very sparsely distributed in a local area.‬
‭Population size can affect the potential for adaptation because it affects the population's‬
‭genetic variation. Density can affect interactions within a population, such as competition‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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f‭ or food and the ability of individuals to find a mate. Smaller organisms tend to be more‬
‭densely distributed than larger organisms (‬‭Figure‬‭23.12‬‭).‬

‭ igure 23.12‬ ‭Australian mammals show a typical‬‭inverse relationship between‬

F
‭population density and body size. (credit:‬‭OpenStax‬‭)‬

‭Estimating Population Size‬

‭ he most accurate way to determine population size is to count all individuals within the‬
T
‭area. However, this method is usually not logistically or economically feasible, especially‬
‭when studying large areas. Thus, scientists usually study populations by sampling a‬
‭representative portion of each habitat and use this sample to make inferences about the‬
‭population as a whole. The methods used to sample populations to determine their size‬
‭and density are typically tailored to the characteristics of the organism being studied. A‬
‭quadrat may be utilized for immobile organisms such as plants or small and slow-moving‬
‭o rganisms. A‬‭quadrat‬‭is a wood, plastic, or metal‬‭square randomly located on the ground‬
‭and used to count the number of individuals within its boundaries. To obtain an accurate‬
‭count using this method, the square must be placed at random locations within the habitat‬
‭enough times to produce an accurate estimate. This counting method will provide an‬
‭estimate of both population size and density. The number and size of quadrat samples‬
‭depend on the type of organisms and the nature of their distribution.‬
‭ arking and recapture are often used for smaller mobile organisms like mammals. This‬
M
‭method involves marking a sample of captured animals in some way and releasing them‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ ack into the environment to mix with the rest of the population; then, a new sample is‬
b
‭captured, and scientists determine how many of the marked animals are in the new‬
‭sample. This method assumes that the larger the population, the lower the percentage of‬
‭marked organisms that will be recaptured since they will have mixed with more unmarked‬
‭individuals. For example, if 80 field mice are captured, marked, and released into the‬
‭forest, then a second trapping 100 field mice are captured, and 20 of them are marked,‬
‭the population size (‬‭N‬‭) can be determined using the following equation:‬

‭𝑛𝑢𝑚𝑏𝑒𝑟‬‭‬‭𝑚𝑎𝑟𝑘𝑒𝑑‬‭‬‭𝑓𝑖𝑟𝑠𝑡‬‭‬‭𝑐𝑎𝑡𝑐ℎ‬‭‬×‭‬‭𝑡𝑜𝑡𝑎𝑙‬‭‬‭𝑛𝑢𝑚𝑏𝑒𝑟‬‭‭𝑠‬ 𝑒𝑐𝑜𝑛𝑑‬‭‬‭𝑐𝑎𝑡𝑐ℎ‬
‭𝑛𝑢𝑚𝑏𝑒𝑟‬‭‬‭𝑚𝑎𝑟𝑘𝑒𝑑‬‭‭𝑠‬ 𝑒𝑐𝑜𝑛𝑑‬‭‭𝑐‬ 𝑎𝑡𝑐ℎ‬
‭‬ = ‭‬‭𝑁‬

‭Using our example, the population size would be 400.‬

‭80 ×
‬ ‭100‬
‭20‬
‭‬ = ‭400‬

‭ hese results give us an estimate of 400 total individuals in the original population. The‬
T
‭actual number usually will be different from this because of chance errors and possible‬
‭bias caused by the sampling methods.‬

‭Species Distribution‬
I‭ n addition to measuring density, further information about a population can be obtained‬
‭by looking at the distribution of the individuals throughout their range. A species‬
‭distribution pattern is the distribution of individuals within a habitat at a particular‬
‭time—broad categories of patterns are used to describe them.‬
I‭ ndividuals within a population can be distributed randomly, in groups, or equally spaced‬
‭apart (more or less). These are known as random, clumped, and uniform distribution‬
‭patterns (‬‭Figure 23.13‬‭). Different distributions reflect‬‭important aspects of the species'‬
‭biology and affect the mathematical methods required to estimate population sizes. An‬
‭example of random distribution occurs with dandelion and other plants with‬
‭wind-dispersed seeds germinating wherever they fall in favorable environments. A‬
‭clumped distribution may be seen in plants that drop their seeds straight to the ground,‬
‭such as oak trees; it can also be seen in animals that live in social groups (schools of fish‬
‭o r herds of elephants). Uniform distribution is observed in plants that secrete substances‬
‭inhibiting the growth of nearby individuals (such as the release of toxic chemicals by sage‬
‭plants). It is also seen in territorial animal species, such as penguins, that maintain a‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ efined territory for nesting. Each individual's territorial defensive behaviors create a‬
d
‭regular distribution pattern of similar-sized territories and individuals within those‬
‭territories. Thus, the distribution of the individuals within a population provides more‬
‭information about how they interact with each other than a simple density measurement.‬
‭Just as lower-density species might have more difficulty finding a mate, solitary species‬
‭with a random distribution might have a similar difficulty when compared to social species‬
‭clumped together in groups.‬

‭ igure 23.13‬ ‭Species may have a random, clumped,‬‭o r uniform distribution. Plants such‬
F
‭as (a) dandelions with wind-dispersed seeds tend to be randomly distributed. Animals‬
‭such as (b) elephants that travel in groups exhibit a clumped distribution. Territorial birds‬
‭such as (c) penguins tend to have a uniform distribution. (credit a: modification of work‬
‭by Rosendahl; credit b: modification of work by Rebecca Wood; credit c: modification of‬
‭work by Ben Tubby;‬ ‭OpenStax‬‭)‬

‭Demography‬
‭ hile population size and density describe a population at one particular point in time,‬
W
‭scientists must use demography to study the dynamics of a population.‬‭Demography‬‭is‬
‭the statistical study of population changes over time: birth rates, death rates, and life‬
‭expectancies. These population characteristics are often displayed in a life table.‬
‭ ife tables‬‭provide important information about the‬‭life history of an organism and the‬
L
‭life expectancy of individuals at each age. They are modeled after actuarial tables used by‬
‭the insurance industry for estimating human life expectancy. Life tables may include the‬
‭probability of each age group dying before their next birthday as well as the percentage‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ f surviving individuals dying at a particular age interval (their mortality rate, and their life‬
o
‭expectancy at each interval). An example of a life table is shown in‬‭Figure 23.14‬‭from a‬
‭study of Dall mountain sheep, a species native to northwestern North America.‬
‭Notice that the population is divided into age intervals (column A). The mortality rate (per‬
‭1000) shown in column D is based on the number of individuals dying during the age‬
‭interval (column B), divided by the number of individuals surviving at the beginning of the‬
‭interval (Column C) multiplied by 1000.‬

‭𝑛𝑢𝑚𝑏𝑒𝑟‬‭‭𝑜
‬ 𝑓‬‭‬‭𝑖𝑛𝑑𝑖𝑣𝑖𝑑𝑢𝑎𝑙𝑠‬‭‬‭𝑑𝑦𝑖𝑛𝑔‬
‭𝑚𝑜𝑟𝑡𝑎𝑙𝑖𝑡𝑦‬‭𝑟𝑎𝑡𝑒‬ = ‭‬ ‭𝑛𝑢𝑚𝑏𝑒𝑟‬‭‬‭𝑜𝑓‬‭‬‭𝑖𝑛𝑑𝑖𝑣𝑖𝑑𝑢𝑎𝑙𝑠‬‭‬‭𝑠𝑢𝑟𝑣𝑖𝑣𝑖𝑛𝑔‬ ‭‬ × ‭1000‬

‭ or example, 12 individuals died out of the 776 remaining from the original 1000 sheep‬
F
‭between ages three and four. This number is multiplied by 1000 to give the mortality rate‬
‭per thousand.‬

‭12‬
‭𝑚𝑜𝑟𝑡𝑎𝑙𝑖𝑡𝑦‬‭𝑟𝑎𝑡𝑒‬ = ‭‬ ‭776‬ ‭‬ × ‭1000 ‬ ≈ ‭15‬. ‭5‬

‭ s can be seen from the mortality rate data (column D), a high death rate occurred when‬
A
‭the sheep were between six months and a year old and then increased even more from 8‬
‭to 9 years old, after which there were few survivors. The data indicate that if a sheep in‬
‭this population survived to age one, it could be expected to live another 7.7 years on‬
‭average, as shown by the life-expectancy numbers in column E.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭A‬ ‭B‬ ‭C‬ ‭D‬ ‭E‬

‭ ge interval‬
A ‭ umber dying‬
N ‭ umber‬
N ‭ ortality rate per‬
M ‭ ife expectancy‬
L
‭(years)‬ ‭in age interval‬ ‭surviving at the‬ ‭1000 alive at the‬ ‭or mean lifetime‬
‭out of 1000‬ ‭beginning of age‬ ‭beginning of the‬ ‭remaining to‬
‭born‬ ‭interval out of‬ ‭age interval‬ ‭those attaining‬
‭1000 born‬ ‭age interval‬
‭0–0.5‬ ‭54‬ ‭1000‬ ‭54.0‬ ‭7.06‬
‭0.5–1‬ ‭145‬ ‭946‬ ‭153.3‬ ‭—‬
‭1–2‬ ‭12‬ ‭801‬ ‭15.0‬ ‭7.7‬
‭2–3‬ ‭13‬ ‭789‬ ‭16.5‬ ‭6.8‬
‭3–4‬ ‭12‬ ‭776‬ ‭15.5‬ ‭5.9‬
‭4–5‬ ‭30‬ ‭764‬ ‭39.3‬ ‭5.0‬
‭5–6‬ ‭46‬ ‭734‬ ‭62.7‬ ‭4.2‬
‭6–7‬ ‭48‬ ‭688‬ ‭69.8‬ ‭3.4‬
‭7–8‬ ‭69‬ ‭640‬ ‭107.8‬ ‭2.6‬
‭8–9‬ ‭132‬ ‭571‬ ‭231.2‬ ‭1.9‬

‭ igure 23.14‬ ‭This life table of‬‭Ovis dalli‬‭shows‬‭the number of deaths, survivors, mortality‬
F
‭rate, and life expectancy at each age interval for Dall mountain sheep. (credit:‬‭OpenStax‬‭)‬

‭23.5 Population Growth and Carrying Capacity‬

‭ he two simplest models of population growth use deterministic equations (equations‬
T
‭that do not account for random events) to describe the rate of change in the size of a‬
‭population over time. The first of these models,‬‭exponential‬‭growth‬‭, describes‬
‭theoretical populations that increase in numbers without limits to their growth. The‬
‭second model,‬‭logistic growth‬‭, limits reproductive‬‭growth, which becomes more intense‬
‭as the population increases. Neither model adequately describes natural populations, but‬
‭they provide points of comparison.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Exponential Growth‬
I‭ n his theory of natural selection, Charles Darwin was greatly influenced by the English‬
‭clergyman Thomas Malthus. Malthus published a book in 1798 stating that populations‬
‭with unlimited natural resources multiply, representing exponential growth, and then‬
‭population growth decreases as resources become depleted, indicating a logistic growth.‬
‭ he best example of exponential growth in organisms is seen in bacteria. Bacteria are‬
T
‭prokaryotes that reproduce primarily by binary fission. This division takes about an hour‬
‭for many bacterial species. If 1000 bacteria are placed in a large flask with an abundant‬
‭supply of nutrients (so the nutrients will not become quickly depleted), the number of‬
‭bacteria will have doubled from 1000 to 2000 after just an hour. Each of the 2000‬
‭bacteria will divide in another hour, producing 4000 bacteria. After the third hour, there‬
‭should be 8000 bacteria in the flask. The critical concept of exponential growth is that the‬
‭growth rate—the number of organisms added in each reproductive generation—is itself‬
‭increasing; that is, the population size is growing at a greater and greater rate. After 24‬
‭cycles, the population would have increased from 1000 to more than 16 billion bacteria. A‬
‭J-shaped growth curve is produced when the population size,‬‭N‭,‬ is plotted over time‬
‭(‬‭Figure 23.15a‬‭).‬
‭ he bacteria-in-a-flask example does not represent the real world, where resources are‬
T
‭usually limited. However, when a species is introduced into a new habitat that it finds‬
‭suitable, it may show exponential growth for a while. In the case of the bacteria in the‬
‭flask, some bacteria will die during the experiment and thus not reproduce; therefore, the‬
‭growth rate is lowered from a maximal rate in which there is no mortality. The‬‭growth‬
‭rate of a population‬‭is primarily determined by subtracting‬‭the‬‭death rate‬‭,‬‭D‬‭(the‬
‭number of organisms that die during an interval) from the‬‭birth rate‬‭, and‬‭B‬‭(the number‬
‭o f organisms that are born during an interval). The growth rate can be expressed in a‬
‭simple equation combining birth and death rates into a single factor:‬‭r‬‭. This is shown in‬
‭the following formula:‬

‭𝑃𝑜𝑝𝑢𝑙𝑎𝑡𝑖𝑜𝑛‬‭𝑔𝑟𝑜𝑤𝑡ℎ‬ = ‭‭𝑟‬ 𝑁‬

‭ he value of‬‭r‬‭can be positive, meaning the population‬‭is increasing in size (the rate of‬
T
‭change is positive); or negative, meaning the population is decreasing in size; or zero, in‬
‭which case the population size is unchanging, a condition known as zero population‬
‭growth.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Logistic Growth‬
‭ xtended exponential growth is possible only when infinite natural resources are‬
E
‭available; this is not true in the real world. Charles Darwin recognized this fact in his‬
‭description of the “struggle for existence,” which states that individuals will compete (with‬
‭members of their own or other species) for limited resources. The successful ones are‬
‭more likely to survive and pass on the traits that made them successful to the next‬
‭generation at a greater rate (natural selection). Population ecologists developed the‬
‭logistic growth model to model the reality of limited resources.‬

‭Carrying Capacity and the Logistic Model‬

‭ ith limited resources, exponential growth can only continue for a while in the real world.‬
W
‭Exponential growth may occur in environments with few individuals and plentiful‬
‭resources. However, when the number of individuals gets large enough, resources will be‬
‭depleted, and the growth rate will slow down. Eventually, the growth rate will plateau or‬
‭level off (‬‭Figure 23.15b‬‭). This population size, determined‬‭by the maximum population‬
‭size that a particular environment can sustain, is called the‬‭c arrying capacity‬‭, or‬‭K‭.‬ In‬
‭natural populations, a growing population often overshoots its carrying capacity, and the‬
‭death rate increases beyond the birth rate, causing the population size to decline back to‬
‭the carrying capacity or below it. Most populations usually fluctuate around the carrying‬
‭capacity in an undulating fashion rather than existing right at it.‬
‭ he formula used to calculate logistic growth adds the carrying capacity as a moderating‬
T
‭force in the growth rate. The expression “‬‭K‬‭–‬‭N‬‭” is‬‭equal to the number of individuals that‬
‭may be added to a population at a given time, and “‬‭K‬‭–‬‭N‬‭” divided by “‬‭K‬‭” is the fraction of‬
‭the carrying capacity available for further growth. Thus, the exponential growth model is‬
‭restricted by this factor to generate the logistic growth equation:‬

‭𝐾‬−‭𝑁‬
‭𝑃𝑜𝑝𝑢𝑙𝑎𝑡𝑖𝑜𝑛‬‭𝑔𝑟𝑜𝑤𝑡ℎ‬ = ‭‭𝑟‬ 𝑁‬‭⎡‬ ⎤
⎣ ‭𝐾‬ ⎦

‭ otice that when‬‭N‬‭is almost zero, the quantity in‬‭brackets is nearly equal to 1 (or‬‭K‬‭/‬‭K‬‭),‬
N
‭and growth is close to exponential. When the population size equals the carrying capacity,‬
‭o r‬‭N‬‭=‬‭K‬‭, the quantity in brackets equals zero, and‬‭growth equals zero.‬
‭ graph of this equation (logistic growth) yields the‬‭S-shaped growth curve‬‭(‬‭Figure‬
A
‭23.15b‬‭). It is a more realistic model of population‬‭growth than exponential growth. An‬
‭S-shaped curve has three different sections. Initially, growth is exponential because few‬
‭individuals and ample resources are available. Then, as resources begin to become limited,‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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t‭ he growth rate decreases. Finally, the growth rate levels off at the environment's carrying‬
‭capacity, with little change in population number over time.‬

‭ igure 23.15‬ ‭When resources are unlimited, populations‬‭exhibit (a) exponential growth,‬
F
‭shown in a J-shaped curve. When resources are limited, populations exhibit (b) logistic‬
‭growth. In logistic growth, population expansion decreases as resources become scarce‬
‭and levels off when the environment's carrying capacity is reached. The logistic growth‬
‭curve is S-shaped. (credit:‬‭OpenStax‬‭)‬

‭Examples of Logistic Growth‬

‭ east, a microscopic fungus used to make bread and alcoholic beverages, exhibits the‬
Y
‭classical S-shaped curve when grown in a test tube (‬‭Figure 23.16a‬‭). Its growth levels off‬
‭as the population depletes the nutrients necessary. In the real world, however, there are‬
‭variations to this idealized curve. Examples in wild populations include sheep and harbor‬
‭seals (‬‭Figure 23.16b‬‭). In both examples, the population‬‭size exceeds the carrying capacity‬
‭for short periods and then falls below the carrying capacity afterward. This fluctuation in‬
‭population size continues to occur as the population oscillates around its carrying‬
‭capacity. Still, even with this oscillation, the logistic model is confirmed.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.16‬ ‭(a) Yeast grown in ideal conditions‬‭in a test tube shows a classical S-shaped‬
F
‭logistic growth curve, whereas (b) a natural population of seals shows real-world‬
‭fluctuation. The yeast is visualized using differential interference contrast light‬
‭micrography. (credit a: scale-bar data from Matt Russell;‬ ‭OpenStax‬‭)‬

‭23.6 The Human Population‬

‭ oncepts of animal population dynamics can be applied to human population growth.‬
C
‭Humans are not unique in their ability to alter their environment. For example, beaver‬
‭dams alter the stream environment where they are built. Humans, however, can change‬
‭their environment to increase its carrying capacity, sometimes to the detriment of other‬
‭species. Earth’s human population and its use of resources are growing rapidly, to the‬
‭extent that some worry about the ability of Earth’s environment to sustain its human‬
‭population. Long-term exponential growth carries with it the potential risks of famine,‬
‭disease, and large-scale death, as well as social consequences of crowding, such as‬
‭increased crime.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Chapter 23: Ecology, Populations, and Biodiversity‬

‭ lthough humans have increased the carrying capacity of their environment, the‬
A
‭technologies used to achieve this transformation have caused unprecedented changes to‬
‭Earth’s environment, altering ecosystems to the point where some may be in danger of‬
‭collapse. The depletion of the ozone layer, erosion due to acid rain, and damage from‬
‭global climate change are caused by human activities. The ultimate effect of these changes‬
‭o n our carrying capacity is unknown. As some point out, it is likely that the adverse effects‬
‭o f increasing carrying capacity will outweigh the positive ones—the world’s carrying‬
‭capacity for human beings might decrease.‬
‭ he world’s human population is presently growing exponentially (‬‭Figure 23.17‬‭).‬
T
‭However, the growth appears to be slowing in more economically-developed countries. A‬
‭consequence of the exponential growth rate is that the time it takes to add a particular‬
‭number of humans to the population is becoming shorter. However, only 126 years were‬
‭necessary to add 1 billion humans between 1804 and 1930, and only 24 years to add the‬
‭two billion people between 1975 and 1999 (‬‭Figure 23.18‬‭).‬ ‭This acceleration in growth‬
‭rate will likely begin to decrease in the coming decades. Despite this, the population will‬
‭continue to increase, and the threat of overpopulation remains, mainly because the‬
‭damage caused to ecosystems and biodiversity is lowering the planet's human carrying‬
‭capacity.‬

‭ igure 23.17‬ ‭Human population growth since 1000‬‭AD is exponential (dark blue line).‬
F
‭Notice that while the population in Asia (yellow line), which has many economically‬
‭underdeveloped countries, is increasing exponentially, the population in Europe (light blue‬
‭line), where most countries are economically developed, is growing much more slowly.‬
‭(credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.18‬ ‭The time between the addition of each‬‭billion human beings to Earth‬
F
‭decreases over time. (credit: modification of work by Ryan T. Cragun;‬‭OpenStax‬‭)‬

‭Age Structure, Population Growth, and Economic Development‬

‭ he age structure of a population is an essential factor in population dynamics. Age‬
T
‭structure is the proportion of a population at different age ranges. Age structure allows‬
‭better prediction of population growth, plus the ability to associate this growth with the‬
‭region's economic development level. Countries with rapid growth have a pyramidal shape‬
‭in their‬‭age structure diagrams‬‭, showing a preponderance‬‭o f younger individuals, many‬
‭o f whom are of reproductive age or will be soon (‬‭Figure‬‭23.19‬‭). This pattern is often‬
‭o bserved in underdeveloped countries where individuals do not live to old age because of‬
‭less-than-optimal living conditions. Age structures of areas with slow growth, including‬
‭developed countries such as the United States, still have a pyramidal structure but with‬
‭fewer young and reproductive-aged individuals and a more significant proportion of older‬
‭individuals. Other developed countries, such as Italy, have zero population growth. The age‬
‭structure of these populations is more conical, with an even greater percentage of‬
‭middle-aged and older individuals. The actual growth rates in different countries are‬
‭shown in‬‭Figure 23.20‬‭, with the highest rates tending‬‭to be in Africa and Asia's less‬
‭economically developed countries.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.19‬ ‭Typical age structure diagrams are‬‭shown. The rapid growth diagram‬
F
‭narrows to a point, indicating that the number of individuals decreases rapidly with age. In‬
‭the slow growth model, the number of individuals decreases steadily with age. Stable‬
‭population diagrams are rounded on the top, showing that the number of individuals per‬
‭age group decreases gradually and then increases for the older part of the population.‬
‭(credit:‬‭OpenStax‬‭)‬

‭ igure 23.20‬ ‭The percent growth rate of populations‬‭in different countries is shown.‬
F
‭Notice that the highest growth is occurring in less economically developed countries in‬
‭Africa and Asia. (credit:‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ or additional information about the age structure diagrams, also known as population‬
F
‭pyramids, click the link or scan the QR code to watch the TED-Ed video by Kim Preshoff‬
‭titled “Population pyramids: Powerful predictors of the future.”‬

Population pyramids: Powerful predictors of the future - Kim Pr…

‭Long-Term Consequences of Exponential Human Population Growth‬

‭ any dire predictions have been made about the world’s population, leading to a‬
M
‭significant crisis called the “population explosion.” In the 1968 book‬‭The Population Bomb‬‭,‬
‭biologist Dr. Paul R. Ehrlich wrote, “The battle to feed all of humanity is over. In the 1970s,‬
‭hundreds of millions of people would starve to death despite any crash programs‬
‭embarked upon. Nothing can prevent a substantial increase in the world death rate at this‬
‭late date.” While many critics view this statement as an exaggeration, the laws of‬
‭exponential population growth are still in effect, and unchecked human population growth‬
‭cannot continue indefinitely.‬
‭ fforts to moderate population control led to the one-child policy in China, which imposes‬
E
‭fines on urban couples who have more than one child. Because some couples wish to have‬
‭a male heir, many Chinese couples continue to have more than one child. The effectiveness‬
‭o f the policy in limiting overall population growth is controversial, as is the policy itself.‬
‭Moreover, there are stories of female infanticide having occurred in some of the more‬
‭rural areas of the country. Family planning education programs in other countries have‬
‭had highly positive effects on limiting population growth rates and increasing living‬
‭standards. Despite population control policies, the human population continues to grow.‬
‭Because of the subsequent need to produce more and more food to feed our population,‬
‭inequalities in access to food and other resources will continue to widen. The United‬
‭Nations estimates the future world population size could vary from 6 billion (a decrease)‬
‭to 16 billion people by 2100. There is no way to know whether human population growth‬
‭will moderate to the point where the crisis described by Dr. Ehrlich will be averted.‬
‭ nother consequence of population growth is the change and degradation of the natural‬
A
‭environment. Many countries have attempted to reduce the human impact on climate‬
‭change by limiting their emission of greenhouse gases. However, a global climate change‬
‭treaty remains elusive, and many underdeveloped countries trying to improve their‬
‭economic condition may be less likely to agree with such provisions without compensation‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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i‭f it means slowing their economic development. Furthermore, the role of human activity‬
‭in causing climate change has become a hotly debated socio-political issue in some‬
‭developed countries, including the United States. Thus, we enter the future with‬
‭considerable uncertainty about our ability to curb human population growth and protect‬
‭o ur environment to maintain the carrying capacity of the human species.‬
‭ or additional information about the history and challenges of human population growth,‬
F
‭click the link or scan the QR code to watch the‬‭Kurzgesagt‬‭– In a Nuts‬‭deo titled‬
‭“Overpopulation - The Human Explosion Explained.”‬

Overpopulation – The Human Explosion Explained

‭23.7 What is Biodiversity, and Why is it Important?‬

‭ iodiversity‬‭is a broad term for biological variety,‬‭and it can be measured at various‬
B
‭o rganizational levels. Traditionally, ecologists have measured biodiversity by considering‬
‭both the number of species and the number of individuals in each species. However,‬
‭biologists are using measures of biodiversity at several levels of biological organization‬
‭(including genes, populations, and ecosystems) to help focus efforts to preserve the‬
‭biologically and technologically important elements of biodiversity.‬
‭ hen biodiversity loss through extinction is considered the loss of the passenger pigeon,‬
W
‭the dodo, or even the wooly mammoth, there seems to be no reason to care about it‬
‭because these events happened long ago. How is this loss practically important for the‬
‭welfare of the human species? Would these species have made our lives any better? From‬
‭the perspective of evolution and ecology, losing a particular species may seem‬
‭unimportant, with some exceptions. Still, the accelerated extinction rate means the loss of‬
‭tens of thousands of species within our lifetimes. Much of this loss occurs in tropical‬
‭rainforests, especially high-diversity ecosystems that are being cleared for timber and‬
‭agriculture. This is likely to affect human welfare dramatically through the collapse of‬
‭ecosystems and add costs to maintaining food production, clean air and water, and‬
‭improving human health.‬
‭ iologists recognize that human populations are embedded in ecosystems and depend on‬
B
‭them, just as is every other species on the planet. Agriculture began after early‬
‭hunter-gatherer societies settled in one place and heavily modified their immediate‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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e‭ nvironment: the ecosystem in which they existed. This cultural transition has made it‬
‭difficult for humans to recognize their dependence on living things other than crops and‬
‭domesticated animals. Today, our technology smoothes out the extremes of existence. It‬
‭allows many of us to live longer, more comfortable lives, but ultimately, the human species‬
‭cannot exist without its surrounding ecosystems. Our ecosystems provide our food. This‬
‭includes living plants that grow in soil ecosystems and the animals that eat these plants (or‬
‭o ther animals), as well as photosynthetic organisms in the oceans and the different‬
‭o rganisms that eat them. Our ecosystems have provided and will provide many‬
‭medications that maintain our health, commonly made from compounds found in living‬
‭o rganisms. Ecosystems provide our clean water, which is held in lake and river ecosystems‬
‭o r passes through terrestrial ecosystems on its way into groundwater.‬

‭Types of Biodiversity‬
‭ common meaning of biodiversity is simply the number of species in a location or on‬
A
‭Earth; for example, the American Ornithologists’ Union lists 2078 species of birds in North‬
‭and Central America. This is one measure of the bird biodiversity on the continent. More‬
‭sophisticated measures of diversity take into account the relative abundances of species.‬
‭For example, a forest with ten equally common species of trees is more diverse than a‬
‭forest with ten species of trees wherein just one of those species makes up 95 percent of‬
‭the trees rather than being equally distributed. Biologists have also identified alternate‬
‭measures of biodiversity, some of which are important in planning how to preserve‬
‭biodiversity.‬

‭Genetic and Chemical Diversity‬

‭ enetic diversity is one alternate concept of biodiversity.‬‭Genetic diversity‬‭(or variation)‬
G
‭is the raw material for a species. A species’ future potential for adaptation depends on the‬
‭genetic diversity held in the genomes of the individuals in populations that make up the‬
‭species. The same is true for higher taxonomic categories. A genus with very different‬
‭types of species will have more genetic diversity than a genus with species that look alike‬
‭and have similar ecologies. The genus with the most significant potential for subsequent‬
‭evolution is the most genetically diverse one.‬
‭ ost genes code for proteins, which in turn carry out the metabolic processes that keep‬
M
‭o rganisms alive and reproducing. Genetic diversity can also be considered as‬‭c hemical‬
‭diversity‬‭in that species with different genetic makeups‬‭produce different assortments of‬
‭chemicals in their cells (proteins as well as the products and byproducts of metabolism).‬
‭This chemical diversity is vital for humans because of the potential uses for these‬
‭chemicals, such as medications. For example, the drug eptifibatide is derived from‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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r‭ attlesnake venom and is used to prevent heart attacks in individuals with certain heart‬
‭conditions.‬
‭ t present, it is far cheaper to discover compounds made by an organism than to imagine‬
A
‭them and then synthesize them in a laboratory. Chemical diversity is one way to measure‬
‭diversity that is important to human health and welfare. Through selective breeding,‬
‭humans have domesticated animals, plants, and fungi, but even this diversity is suffering‬
‭losses because of market forces and increasing globalism in human agriculture and‬
‭migration. For example, international seed companies produce only a few varieties of a‬
‭given crop and provide incentives worldwide for farmers to buy these few varieties while‬
‭abandoning their traditional varieties, which are far more diverse. The human population‬
‭depends on crop diversity directly as a stable food source, and its decline is troubling to‬
‭biologists and agricultural scientists.‬

‭Ecosystems Diversity‬
I‭ t is also helpful to define‬‭ecosystem diversity‬‭,‬‭which is the number of different‬
‭ecosystems on Earth or in a geographical area. Whole ecosystems can disappear even if‬
‭some species survive by adapting to other ecosystems. The loss of an ecosystem means‬
‭the loss of the interactions between species, the loss of unique features of coadaptation,‬
‭and the loss of biological productivity that an ecosystem can create. The prairie ecosystem‬
‭is an example of a largely extinct ecosystem in North America (‬‭Figure 23.21‬‭). Prairies‬
‭o nce spanned central North America from the boreal forest in northern Canada down into‬
‭Mexico. They are now all but gone, replaced by crop fields, pasture lands, and suburban‬
‭sprawl. Many species survive, but the hugely productive ecosystem responsible for‬
‭creating our most productive agricultural soils is gone. Consequently, their soils are now‬
‭being depleted unless maintained artificially at a more significant expense. The decline in‬
‭soil productivity occurs because the interactions in the original ecosystem have been lost;‬
‭this was a far more critical loss than the relatively few species that were driven extinct‬
‭when the prairie ecosystem was destroyed.‬

‭Current Species Diversity‬

‭ espite considerable effort, knowledge of the‬‭species‬‭diversity‬‭that inhabits the planet is‬
D
‭limited. A recent estimate suggests that the eukaryote species for which science has‬
‭names, about 1.5 million species, account for less than 20 percent of the total number of‬
‭eukaryote species present on the planet (8.7 million species, by one estimate). Estimates of‬
‭numbers of prokaryotic species are largely guesses, but biologists agree that science has‬
‭o nly just begun to catalog their diversity. Even with what is known, there is no centralized‬
‭repository of names or samples of the described species; therefore, there is no way to be‬
‭sure that the 1.5 million descriptions are accurate. It is a best guess based on experts'‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ pinions on different taxonomic groups. Since Earth is losing species at an accelerating‬

o
‭pace, science needs to learn more about what is being lost.‬

‭ igure 23.21‬ ‭The variety of ecosystems on Earth—from‬‭coral reefs to prairies—enables‬

F
‭a great diversity of species to exist. (Credit “coral reef” and “prairie”: modification of work‬
‭by Jim Maragos, USFWS).‬

‭ arious initiatives are underway to catalog described species in accessible and more‬
V
‭o rganized ways, and the internet is facilitating that effort. Naming and counting species‬
‭may seem unimportant given the other needs of humanity, but it is more than just an‬
‭accounting. Describing species is a complex process by which biologists determine an‬
‭o rganism’s unique characteristics and whether or not that organism belongs to any other‬
‭described species. It allows biologists to find and recognize the species after the initial‬
‭discovery to follow up on questions about its biology. That subsequent research will‬
‭produce discoveries that make the species valuable to us and our ecosystems. Without a‬
‭name and description, a species cannot be studied in depth and in a coordinated way by‬
‭multiple scientists.‬

‭Importance of Biodiversity‬
‭ oss of biodiversity eventually threatens other species we do not impact directly because‬
L
‭o f their interconnectedness; as species disappear from an ecosystem, other species are‬
‭threatened by the changes in available resources. Biodiversity is essential to the survival‬
‭and welfare of human populations because it impacts our health and ability to feed‬
‭o urselves through agriculture and harvesting populations of wild animals.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Human Health‬
‭ any medications are derived from natural chemicals made by diverse organisms. For‬
M
‭example, many plants produce secondary plant compounds, which are toxins used to‬
‭protect the plant from insects and other animals that eat them. Some of these secondary‬
‭plant compounds also work as human medicines. Contemporary societies that live close to‬
‭the land often have a broad knowledge of the medicinal uses of plants growing in their‬
‭area. For centuries in Europe, older knowledge about the medical uses of plants was‬
‭compiled in herbals—books that identified the plants and their uses.‬
‭ odern pharmaceutical science also recognizes the importance of these plant compounds.‬
M
‭Significant medicines derived from plant compounds include aspirin, codeine, digoxin,‬
‭atropine, and vincristine (‬‭Figure 23.22‬‭). Many medications‬‭were once derived from plant‬
‭extracts but are now synthesized. It is estimated that, at one time, 25 percent of modern‬
‭drugs contained at least one plant extract. That number has probably decreased to about‬
‭10 percent as synthetic versions of the plant compounds replace natural plant ingredients.‬
‭Antibiotics, which are responsible for extraordinary improvements in health and lifespans‬
‭in developed countries, are compounds derived mainly from fungi and bacteria.‬

I‭ n recent years, animal venoms and poisons have excited intense research for their‬
‭medicinal potential. By 2007, the FDA had approved five drugs based on animal toxins to‬
‭treat diseases such as hypertension, chronic pain, and diabetes. Another five drugs are‬
‭undergoing clinical trials, and at least six are being used in other countries. Other toxins‬
‭under investigation come from mammals, snakes, lizards, various amphibians, fish, snails,‬
‭o ctopuses, and scorpions.‬

‭ side from representing billions of dollars in profits, these medications improve people’s‬
A
‭lives. Pharmaceutical companies are looking for new natural compounds that can function‬
‭as medicines. It is estimated that one-third of pharmaceutical research and development is‬
‭spent on natural compounds, and about 35 percent of new drugs brought to market‬
‭between 1981 and 2002 were from natural compounds.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.22‬ ‭Catharanthus roseus‬‭, the Madagascar‬‭periwinkle, has various medicinal‬

F
‭properties. Among other uses, it is a source of vincristine, a drug used to treat lymphomas.‬
‭(credit: Forest and Kim Starr;‬ ‭OpenStax‬‭)‬

‭Agriculture‬
S‭ ince the beginning of human agriculture, more than 10,000 years ago, human groups‬
‭have been breeding and selecting crop varieties. This crop diversity matched the cultural‬
‭diversity of highly subdivided populations of humans. For example, potatoes were‬
‭domesticated beginning around 7,000 years ago in the central Andes of Peru and Bolivia.‬
‭The people in this region traditionally lived in relatively isolated settlements separated by‬
‭mountains. The potatoes grown in that region belong to seven species, and the number of‬
‭varieties is likely in the thousands. Each variety has been bred to thrive at particular‬
‭elevations and soil and climate conditions. The diversity is driven by the diverse demands‬
‭o f the dramatic elevation changes, the limited movement of people, and the demands‬
‭created by crop rotation for different varieties that will do well in other fields.‬
‭ otatoes are only one example of agricultural diversity. Every plant, animal, and fungus‬
P
‭cultivated by humans has been bred from original wild ancestor species into diverse‬
‭varieties arising from the demands for food value, adaptation to growing conditions, and‬
‭resistance to pests. The potato demonstrates a well-known example of the risks of low‬
‭crop diversity: during the tragic Irish potato famine (1845–1852 AD), the single potato‬
‭variety grown in Ireland became susceptible to a potato blight—wiping out the crop. The‬
‭crop loss led to famine, death, and mass emigration. Disease resistance is a chief benefit to‬
‭maintaining crop biodiversity, and the lack of diversity in contemporary crop species‬
‭carries similar risks. Seed companies, which are the source of most crop varieties in‬
‭developed countries, must continually breed new varieties to keep up with evolving pest‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ rganisms. These same seed companies, however, have participated in the decline of the‬
o
‭number of varieties available as they focus on selling fewer varieties in more areas of the‬
‭world, replacing traditional local varieties.‬
‭ he ability to create new crop varieties relies on the diversity of varieties available and the‬
T
‭availability of wild forms related to the crop plant. These wild forms are often the source‬
‭o f new gene variants that can be bred with existing varieties to create varieties with new‬
‭attributes. Loss of wild species related to a crop will mean the loss of potential for crop‬
‭improvement. Maintaining the genetic diversity of wild species related to domesticated‬
‭species ensures our continued food supply.‬
S‭ ince the 1920s, government agriculture departments have maintained seed banks of crop‬
‭varieties to preserve crop diversity. This system has flaws because, over time, seed‬
‭varieties are lost through accidents, and there is no way to replace them. In 2008, the‬
‭Svalbard Global Seed Vault, located on Spitsbergen Island, Norway (‬‭Figure 23.23‬‭), began‬
‭storing seeds from around the world as a backup system for the regional seed banks. If a‬
‭regional seed bank stores varieties in Svalbard, losses can be replaced from Svalbard‬
‭should something happen to the regional seeds. The Svalbard seed vault is deep into the‬
‭rock of the Arctic island. Conditions within the vault are maintained at ideal temperature‬
‭and humidity for seed survival. Still, the deep underground location of the vault in the‬
‭Arctic means that failure of the vault’s systems will not compromise the climatic conditions‬
‭inside the vault.‬
‭ or additional information about the importance of biodiversity, click the link or scan the‬
F
‭QR code to watch the TED-Ed video by Kim Preshoff titled “Why is biodiversity so‬
‭important?”‬

Why is biodiversity so important? - Kim Preshoff

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.23‬ ‭The Svalbard Global Seed Vault is‬‭a storage facility for seeds of Earth’s‬
F
‭diverse crops. (credit: Mari Tefre, Svalbard Global Seed Vault;‬‭OpenStax‬‭)‬

‭Wild Food Sources‬

I‭ n addition to growing crops and raising animals for food, humans obtain food resources‬
‭from wild populations, primarily fish populations. For approximately 1 billion people,‬
‭aquatic resources provide the primary source of animal protein. But since 1990, global‬
‭fish production has declined. Despite considerable effort, few fisheries on the planet are‬
‭managed for sustainability.‬

‭ ishery extinctions rarely lead to the complete extinction of the harvested species but‬
F
‭rather to a radical restructuring of the marine ecosystem in which a dominant species is‬
‭so over-harvested that it becomes a minor player, ecologically. In addition to humans‬
‭losing the food source, these alterations affect many other species in ways that are difficult‬
‭o r impossible to predict. The collapse of fisheries has dramatic and long-lasting effects on‬
‭local populations working there. In addition, losing an inexpensive protein source to‬
‭populations that cannot afford to replace it will increase the cost of living and limit‬
‭societies in other ways. In general, the fish taken from fisheries have shifted to smaller‬
‭species as larger species are fished to extinction. The ultimate outcome could be the loss‬
‭o f aquatic systems as food sources.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Psychological and Moral Value‬

‭ inally, it has been argued that humans benefit psychologically from living in a biodiverse‬
F
‭world. A chief proponent of this idea is entomologist E. O. Wilson. He argues that human‬
‭evolutionary history has adapted us to live in a natural environment and that built‬
‭environments generate stressors that affect human health and well-being. There is‬
‭considerable research into the psychological regenerative benefits of natural landscapes,‬
‭suggesting the hypothesis may hold some truth. In addition, there is a moral argument‬
‭that humans have a responsibility to inflict as little harm as possible on other species.‬

‭23.8 Threats to Biodiversity‬

‭ he core threat to biodiversity on the planet, and therefore to human welfare, is the‬
T
‭combination of human population growth and the resources used by that population. The‬
‭human population requires resources to survive and grow, and those resources are being‬
‭removed unsustainably from the environment. The three greatest proximate threats to‬
‭biodiversity are habitat loss, overharvesting, and introduction of exotic species. The first‬
‭two directly result from human population growth and resource use. The third one results‬
‭from increased mobility and trade.‬
‭ lobal climate change is also a consequence of the human population’s need for energy‬
G
‭and the use of fossil fuels to meet those needs (‬‭Figure‬‭23.24‬‭). Environmental issues, such‬
‭as toxic pollution, have specific targeted effects on species but are not generally seen as‬
‭threats at the magnitude of the others.‬

‭Habitat Loss‬
‭ umans rely on technology to modify their environment and replace certain functions‬
H
‭o nce performed by the natural ecosystem. Other species cannot do this. Eliminating their‬
‭habitat—whether it is a forest, coral reef, grassland, or flowing river—will kill the‬
‭individuals in the species. Remove the entire habitat within the range of a species; unless‬
‭they are one of the few species that do well in human-built environments, the species will‬
‭become extinct. Human destruction of habitats (habitats generally refer to the part of the‬
‭ecosystem required by a particular species) accelerated in the latter half of the twentieth‬
‭century. Consider the exceptional biodiversity of Sumatra: it is home to one species of‬
‭o rangutan, a species of critically endangered elephant, and the Sumatran tiger, but half of‬
‭Sumatra’s forest is now gone. The neighboring island of Borneo, home to the other‬
‭o rangutan species, has lost a similar forest area. Forest loss continues in protected areas‬
‭o f Borneo.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.24‬ ‭Atmospheric carbon dioxide levels‬‭fluctuate cyclically. However, the‬

F
‭burning of fossil fuels in recent history has caused a dramatic increase in the levels of‬
‭carbon dioxide in the Earth’s atmosphere, which have now reached levels never before‬
‭seen on Earth. Scientists predict that the addition of this “greenhouse gas” to the‬
‭atmosphere will result in climate change that will significantly impact biodiversity in the‬
‭coming century. (credit:‬‭OpenStax‬‭)‬

‭ he orangutan in Borneo is listed as endangered by the International Union for‬

T
‭Conservation of Nature (IUCN). Still, it is simply the most visible of thousands of species‬
‭that will not survive the disappearance of the forests of Borneo. The forests are removed‬
‭for timber and to plant palm oil plantations (‬‭Figure‬‭23.25‬‭). Palm oil is used in many‬
‭European products, including food products, cosmetics, and biodiesel. A 5-year estimate of‬
‭global forest cover loss from 2000 to 2005 was 3.1 percent. Much of the loss (2.4 percent)‬
‭o ccurred in the humid tropics, where forest loss is primarily from timber extraction. These‬
‭losses also represent the extinction of species unique to those areas.‬
‭ abitat destruction can affect ecosystems other than forests. Rivers and streams are‬
H
‭important ecosystems and are frequently the target of habitat modification through‬
‭building and from damming or water removal. Damming of rivers affects flows and access‬
‭to all parts of a river. Altering a flow regime can reduce or eliminate populations adapted‬
‭to seasonal flow changes. For example, an estimated 91 percent of river lengths in the‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ nited States have been modified with damming or bank modifications. Many fish species‬
U
‭in the United States, especially rare species or species with restricted distributions, have‬
‭seen declines caused by river damming and habitat loss.‬

‭ igure 23.25‬ ‭An oil palm plantation in Sabah Province‬‭Borneo, Malaysia, replaces the‬
F
‭native forest habitat that a variety of species depended on to live. (credit: Lian Pin Koh;‬
‭OpenStax‬‭)‬

‭Overharvesting‬
‭ verharvesting is a serious threat to many species, particularly aquatic ones. Many‬
O
‭examples of regulated fisheries (including hunting marine mammals and harvesting‬
‭crustaceans and other species) monitored by fisheries scientists have nevertheless‬
‭collapsed. The western Atlantic cod fishery is the most spectacular recent collapse. While‬
‭it was a hugely productive fishery for 400 years, the introduction of modern factory‬
‭trawlers in the 1980s and the pressure on the fishery made it unsustainable. The causes‬
‭o f fishery collapse are both economic and political. Most fisheries are managed as a‬
‭shared resource, available to anyone willing to fish, even when the fishing territory lies‬
‭within a country’s territorial waters. Common resources are subject to an economic‬
‭pressure known as the tragedy of the commons, in which fishers have little motivation to‬
‭exercise restraint in harvesting a fishery when they do not own the fishery. The general‬
‭o utcome of harvests of resources held in common is their overexploitation. While large‬
‭fisheries are regulated to attempt to avoid this pressure, it still exists in the background.‬
‭This overexploitation is exacerbated when access to the fishery is open and unregulated‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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a‭ nd when technology gives fishers the ability to overfish. In a few fisheries, the biological‬
‭growth of the resource is less than the potential growth of the profits made from fishing if‬
‭that time and money were invested elsewhere. In these cases—whales are an‬
‭example—economic forces will drive fishing the population to extinction.‬

‭Exotic Species‬
‭ xotic species‬‭are species that humans have intentionally‬‭o r unintentionally introduced‬
E
‭into an ecosystem in which they did not evolve. Human transportation of people and‬
‭goods, including the intentional transport of organisms for trade, has dramatically‬
‭increased the introduction of species into new ecosystems. These new introductions are‬
‭sometimes at distances well beyond the species' capacity to ever travel itself and outside‬
‭the range of the species’ natural predators.‬
‭ ost exotic species introductions fail because of the low number of individuals introduced‬
M
‭o r poor adaptation to the ecosystem they enter. Some species, however, have‬
‭characteristics that can make them especially successful in a new ecosystem. These exotic‬
‭species often undergo dramatic population increases in their new habitat and reset the‬
‭ecological conditions in the new environment, threatening the species that exist there.‬
‭When this happens, the exotic species also becomes an‬‭invasive species‬‭. Invasive species‬
‭can threaten other species through resource competition, predation, or disease.‬
‭ akes and islands are particularly vulnerable to extinction threats from introduced‬
L
‭species. In Lake Victoria, the intentional introduction of the Nile perch was primarily‬
‭responsible for the extinction of about 200 species of cichlids. The accidental introduction‬
‭o f the brown tree snake via aircraft (‬‭Figure 23.26‬‭)‬‭from the Solomon Islands to Guam in‬
‭1950 has led to the extinction of three species of birds and three to five species of reptiles‬
‭endemic to the island. Several other species are still threatened. The brown tree snake is‬
‭adept at exploiting human transportation as a means to migrate; one was even found on‬
‭an aircraft arriving in Corpus Christi, Texas. Constant vigilance on the part of airport,‬
‭military, and commercial aircraft personnel is required to prevent the snake from moving‬
‭from Guam to other islands in the Pacific, especially Hawaii. Islands do not comprise a‬
‭large area of land on the globe, but they do contain a disproportionate number of‬
‭endemic species because of their isolation from mainland ancestors.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.26‬ ‭The brown tree snake,‬‭Boiga irregularis‬‭,‬‭is an exotic species that has‬
F
‭caused numerous extinctions on the island of Guam since its accidental introduction in‬
‭1950. (credit: NPS;‬ ‭OpenStax‬‭)‬

‭ any introductions of marine and freshwater aquatic species have occurred when ships‬
M
‭have dumped ballast water taken on at a port of origin into waters at a destination port.‬
‭Water from the port of origin is pumped into tanks on a ship empty of cargo to increase‬
‭stability. The water is drawn from the ocean or estuary of the port and typically contains‬
‭living organisms such as plant parts, microorganisms, eggs, larvae, or aquatic animals. The‬
‭water is pumped out before the ship takes on cargo at the destination port, which may be‬
‭o n a different continent. The zebra mussel was introduced to the Great Lakes from Europe‬
‭before 1988 in ship ballast. The zebra mussels in the Great Lakes have cost the industry‬
‭millions of dollars in clean-up costs to maintain water intakes and other facilities. The‬
‭mussels have also altered the ecology of the lakes dramatically. They threaten native‬
‭mollusk populations but have benefited some species, such as smallmouth bass. The‬
‭mussels are filter feeders and have significantly improved water clarity, which has allowed‬
‭aquatic plants to grow along shorelines, providing shelter for young fish where it did not‬
‭exist before.‬
‭ or additional information about invasive species and their impact, click the link or scan‬
F
‭the QR code to watch the TED-Ed video by Jennifer Klos titled “The threat of invasive‬
‭species.”‬
The threat of invasive species - Jennifer Klos

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭Climate Change‬
‭ limate change is recognized as a major extinction threat, particularly when combined‬
C
‭with other threats such as habitat loss. Warming of the planet has been observed and is‬
‭hypothesized to continue due to past and ongoing emissions of greenhouse gases,‬
‭primarily carbon dioxide and methane, into the atmosphere caused by burning fossil fuels‬
‭and deforestation. These gases decrease the degree to which Earth can radiate heat‬
‭energy created by the sunlight that enters the atmosphere. The climate and energy‬
‭balance changes caused by increasing greenhouse gases are complex, and our‬
‭understanding of them depends on predictions generated from detailed computer models.‬
‭Scientists generally agree that humans cause the present warming trend, and some of the‬
‭likely effects include dramatic and dangerous climate changes in the coming decades.‬
‭However, there is still debate and a need for more understanding about specific outcomes.‬
‭Scientists disagree about the likely magnitude of the effects on extinction rates, with‬
‭estimates ranging from 15 to 40 percent of species committed to extinction by 2050.‬
S‭ cientists agree that climate change will alter regional climates, including rainfall and‬
‭snowfall patterns, making habitats less hospitable to their species. The warming trend will‬
‭shift colder climates toward the north and south poles, forcing species to move with their‬
‭adapted climate norms and face habitat gaps along the way. The shifting ranges will‬
‭impose new competitive regimes on species as they are in contact with other species not‬
‭present in their historic range. One such unexpected species contact is between polar‬
‭bears and grizzly bears. Previously, these two species had separate ranges. Their ranges‬
‭overlap, and there are documented cases of these two species mating and producing‬
‭viable offspring. Changing climates also disrupt species' delicate timing adaptations to‬
‭seasonal food resources and breeding times. Scientists have already documented many‬
‭contemporary mismatches to resource availability and timing shifts.‬
‭ limate gradients will also move up mountains, eventually crowding species higher in‬
C
‭altitude and eliminating the habitat for those species adapted to the highest elevations.‬
‭Some climates will completely disappear. The rate of warming appears to be accelerated in‬
‭the Arctic, which is recognized as a serious threat to polar bear populations that require‬
‭sea ice to hunt seals during the winter months: seals are the only source of protein‬
‭available to polar bears. A trend of decreasing sea ice coverage has occurred since‬
‭o bservations began in the mid-twentieth century. The rate of decline observed in recent‬
‭years is far greater than previously predicted by climate models (‬‭Figure 23.27‬‭).‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.27‬ ‭The effect of global warming can be‬‭seen in the continuing retreat of‬
F
‭Grinnell Glacier. The mean annual temperature in Glacier National Park has increased by‬
‭1.33°C since 1900. The loss of a glacier results in the loss of summer meltwaters, sharply‬
‭reducing seasonal water supplies and severely affecting local ecosystems. (credit: USGS,‬
‭GNP Archives;‬ ‭OpenStax‬‭)‬

‭ inally, global warming will raise ocean levels due to meltwater from glaciers and the‬
F
‭greater volume occupied by warmer water. Shorelines will be inundated, reducing island‬
‭size, which will affect some species, and some islands will disappear entirely. Additionally,‬
‭the gradual melting and subsequent refreezing of the poles, glaciers, and higher-elevation‬
‭mountains—a cycle that has provided freshwater to environments for centuries—will be‬
‭altered. This could result in an overabundance of salt water and a shortage of fresh water.‬
‭ or additional information about climate change, its impact on wildlife, and how they‬
F
‭respond, click the link or scan the QR code to watch the TED-Ed video by Erin Eastwood‬
‭titled “Can wildlife adapt to climate change?”‬

Can wildlife adapt to climate change? - Erin Eastwood

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭23.9 Preserving Biodiversity‬

‭ he threats to biodiversity at the genetic, species, and ecosystem levels have been‬
T
‭recognized for some time. In the United States, the first national park with land set aside to‬
‭remain in a wilderness state was Yellowstone Park in 1890. However, attempts to preserve‬
‭nature for various reasons have occurred for centuries. Primary efforts to conserve‬
‭biodiversity today involve legislative approaches to regulate human and corporate‬
‭behavior, set aside protected areas, and restore habitats.‬

‭Changing Human Behavior‬

‭ egislation has been enacted to protect species throughout the world. The legislation‬
L
‭includes international treaties as well as national and state laws. The Convention on‬
‭International Trade in Endangered Species of Wild Fauna and Flora (CITES) treaty came‬
‭into force in 1975. The treaty and the national legislation that supports it provide a legal‬
‭framework for preventing “listed” species from being transported across nations’ borders,‬
‭thus protecting them from being caught or killed in the first place when the purpose‬
‭involves international trade. The listed species that are protected to one degree or‬
‭another by the treaty number some 33,000. The treaty is limited in its reach because it‬
‭o nly deals with the international movement of organisms or their parts. It is also limited‬
‭by various countries’ ability or willingness to enforce the treaty and supporting legislation.‬
‭The illegal trade in organisms and their parts is probably a hundreds of millions of dollars‬
‭market.‬
I‭ n many countries, laws protect endangered species and regulate hunting and fishing. In‬
‭the United States, the Endangered Species Act was enacted in 1973. When the Act lists an‬
‭at-risk species, the U.S. Fish & Wildlife Service must develop a management plan to protect‬
‭the species and bring it back to sustainable numbers. The Act, and others like it in other‬
‭countries, is a helpful tool, but it suffers because it is often difficult to get a species listed‬
‭o r an effective management plan once a species is listed. Additionally, species may be‬
‭controversially taken off the list without necessarily having had a change in their situation.‬
‭More fundamentally, the approach to protecting individual species rather than entire‬
‭ecosystems (although the management plans commonly involve the protection of the‬
‭particular species’ habitat) is both inefficient and focuses efforts on a few highly visible‬
‭and often charismatic species, perhaps at the expense of other species that go‬
‭unprotected.‬
‭ he Migratory Bird Treaty Act (MBTA) is an agreement between the United States and‬
T
‭Canada signed into law in 1918 in response to declines in North American bird species‬
‭caused by hunting. The Act now lists over 800 protected species. It makes it illegal to‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ isturb or kill the protected species or distribute their parts (much of the hunting of birds‬
d
‭in the past was for their feathers). Examples of protected species include northern‬
‭cardinals, the red-tailed hawk, and the American black vulture.‬
‭ lobal warming is expected to be a major driver of biodiversity loss. Many governments‬
G
‭are concerned about the effects of global warming, primarily on their economies and food‬
‭resources. Since greenhouse gas emissions do not respect national boundaries, the effort‬
‭to curb them is international. The international response to global warming has been‬
‭mixed. The Kyoto Protocol, an international agreement from the United Nations‬
‭Framework Convention on Climate Change that committed countries to reducing‬
‭greenhouse gas emissions by 2012, was ratified by some countries but rejected by others.‬
‭Two countries that were especially important in terms of their potential impact that did‬
‭not ratify the Kyoto Protocol were the United States and China. Some goals for reduction‬
‭in greenhouse gasses were met and exceeded by individual countries, but worldwide, the‬
‭effort to limit greenhouse gas production is failing. The intended replacement for the‬
‭Kyoto Protocol has yet to materialize because governments cannot agree on timelines and‬
‭benchmarks. Meanwhile, as predicted by most climate scientists, the resulting costs to‬
‭human societies and biodiversity will be high.‬
‭ s mentioned, the non-profit, non-governmental sector plays a significant role in‬
A
‭conservation efforts in North America and worldwide. The approaches range from‬
‭species-specific organizations to the broadly focused IUCN and Trade Records Analysis of‬
‭Flora and Fauna in Commerce (TRAFFIC). The Nature Conservancy takes a novel‬
‭approach. It purchases land and protects it in an attempt to set up preserves for‬
‭ecosystems. Ultimately, human behavior will change when human values change. The‬
‭growing urbanization of the human population is a force that mitigates against valuing‬
‭biodiversity because many people no longer come in contact with natural environments‬
‭and the species that inhabit them.‬

‭Conservation in Preserves‬
‭ stablishing wildlife and ecosystem preserves is one essential tool in conservation efforts.‬
E
‭A‬‭preserve‬‭is an area of land set aside with varying‬‭degrees of protection for the‬
‭o rganisms within its boundaries. Preserves can effectively protect species and ecosystems,‬
‭but they have serious drawbacks.‬
‭ simple measure of success in setting aside preserves for biodiversity protection is to set‬
A
‭a target percentage of land or marine habitat to protect. However, a more detailed‬
‭preserve design and choice of location is usually necessary because of the way protected‬
‭lands are allocated and how biodiversity is distributed: protected lands tend to contain‬
‭less economically valuable resources rather than being set aside expressly for the species‬
‭o r ecosystems at risk. In 2003, the IUCN World Parks Congress estimated that preserves‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ f various kinds covered 11.5 percent of Earth’s land surface. This area is greater than‬
o
‭previous goals; however, it only represents 9 out of 14 recognized major biomes, and‬
‭research has shown that 12 percent of all species live outside preserves; these‬
‭percentages are much higher when threatened species are considered and when only‬
‭high-quality preserves are considered. For example, high-quality preserves include only‬
‭about 50 percent of threatened amphibian species. The conclusion must be that the‬
‭percentage of area protected must be increased, the percentage of high-quality preserves‬
‭must be increased, or preserves must be targeted with greater attention to biodiversity‬
‭protection. Researchers argue that more attention is required to the latter solution.‬
‭ ‬‭biodiversity hotspot‬‭is a conservation concept developed‬‭by Norman Myers in 1988.‬
A
‭Hotspots are geographical areas that contain high numbers of endemic species—the idea‬
‭aimed to identify essential locations on the planet for conservation efforts, a conservation‬
‭triage. By protecting hotspots, governments can protect a larger number of species. The‬
‭o riginal criteria for a hotspot included 1500 or more species of endemic plants and 70‬
‭percent of the area disturbed by human activity. There are now 34 biodiversity hotspots‬
‭(‬‭Figure 23.28‬‭) that contain many endemic species,‬‭including half of Earth’s endemic‬
‭plants.‬
‭ here has been extensive research into optimal preserve designs for maintaining‬
T
‭biodiversity. The fundamental principles behind much of the research come from the‬
‭seminal theoretical work of Robert H. MacArthur and Edward O. Wilson, published in‬
‭1967 on island biogeography. This work sought to understand the factors affecting‬
‭biodiversity on islands. Conservation preserves are “islands” of habitat within “an ocean”‬
‭o f non-habitat. In general, large preserves are better because they support more species,‬
‭including species with large home ranges; they have more core areas of optimal habitat‬
‭for individual species; they have more niches to support more species; and they attract‬
‭more species because they can be found and reached more quickly.‬
‭ reserves perform better when there are partially protected buffer zones around them of‬
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‭suboptimal habitat. The buffer allows organisms to exit the preserve's boundaries without‬
‭immediate negative consequences from hunting or lack of resources. One large preserve is‬
‭better than the same area of several smaller preserves because there is more core habitat‬
‭unaffected by less hospitable ecosystems outside the preserve boundary. For this same‬
‭reason, preserves in a square or circle shape will be better than a preserve with many thin‬
‭“arms.” If preserves must be smaller, then providing wildlife corridors between them so‬
‭species and their genes can move between them; for example, preserves along rivers and‬
‭streams will make the smaller preserves behave more like large ones. All of these factors‬
‭are considered when planning the nature of a preserve before the land is set aside.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ igure 23.28‬ ‭Conservation International has identified‬‭34 biodiversity hotspots.‬

F
‭Although these cover only 2.3 percent of the Earth’s surface, 42 percent of the terrestrial‬
‭vertebrate species and 50 percent of the world’s plants are endemic to those hotspots.‬
‭(credit:‬‭OpenStax‬‭)‬

I‭ n addition to the physical specifications of a preserve, there are a variety of regulations‬

‭related to the use of a preserve. These can include anything from timber extraction,‬
‭mineral extraction, regulated hunting, human habitation, and non-destructive human‬
‭recreation. Many of the decisions to include these other uses are made based on political‬
‭pressures rather than conservation considerations. On the other hand, in some cases,‬
‭wildlife protection policies have been so strict that subsistence-living indigenous‬
‭populations have been forced from ancestral lands that fell within a preserve. In other‬
‭cases, even if a preserve is designed to protect wildlife, if the protections are not or cannot‬
‭be enforced, the preserve status will have little meaning in the face of illegal poaching and‬
‭timber extraction. This is a widespread problem with preserves in the tropics.‬
‭ he discussion of preserve design reveals some limitations of preserves as conservation‬
T
‭tools. Political and economic pressures typically make preserves smaller, never larger, so‬
‭setting aside areas that are large enough is difficult. Enforcement of protections is also a‬
‭significant issue in countries without the resources or political will to prevent poaching‬
‭and illegal resource extraction.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

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‭ limate change will create inevitable problems with the location of preserves as the‬
C
‭species within them migrate to higher latitudes as the preserve's habitat becomes less‬
‭favorable. Planning for the effects of global warming on future preserves or adding new‬
‭preserves to accommodate the changes expected from global warming is in progress but‬
‭will only be as effective as the accuracy of the predictions of the effects of global warming‬
‭o n future habitats.‬
‭ inally, an argument can be made that conservation preserves reinforce the cultural‬
F
‭perception that humans are separate from nature, can exist outside of it, and can only‬
‭o perate in ways that damage biodiversity. Creating preserves reduces the pressure on‬
‭human activities outside the preserves to be sustainable and non-damaging to biodiversity.‬
‭Ultimately, the political, economic, and human demographic pressures will degrade and‬
‭reduce the size of conservation preserves if the activities outside them are not altered to‬
‭be less damaging to biodiversity.‬

‭Habitat Restoration‬
‭ abitat restoration holds considerable promise as a mechanism for maintaining or‬
H
‭restoring biodiversity. Of course, once a species has become extinct, restoration is‬
‭impossible. However, restoration can improve the biodiversity of degraded ecosystems.‬
‭Reintroducing wolves, a top predator, to Yellowstone National Park in 1995 led to dramatic‬
‭changes in the ecosystem that increased biodiversity. The wolves (‬‭Figure 23.29‬‭) function‬
‭to suppress elk and coyote populations and provide more abundant resources to the guild‬
‭o f carrion eaters. Reducing elk populations has allowed revegetation of riparian (the‬
‭areas along the banks of a stream or river) areas, which has increased the diversity of‬
‭species in that habitat. The suppression of coyotes has increased the species previously‬
‭suppressed by this predator. The number of species of carrion eaters has increased‬
‭because of the predatory activities of the wolves. In this habitat, the wolf is a‬‭keystone‬
‭species‬‭, meaning a species that is instrumental in‬‭maintaining diversity within an‬
‭ecosystem. Removing a keystone species from an ecological community causes a collapse‬
‭in diversity. The results from the Yellowstone experiment suggest that restoring a keystone‬
‭species effectively can restore biodiversity in the community. Ecologists have argued for‬
‭identifying keystone species where possible and focusing protection efforts on these‬
‭species. It makes sense to return the keystone species to the ecosystems where they have‬
‭been removed.‬
‭ ther large-scale restoration experiments underway involve dam removal. In the United‬
O
‭States, since the mid-1980s, many aging dams have been considered for removal rather‬
‭than replacement because of shifting beliefs about the ecological value of free-flowing‬
‭rivers.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭694‬
‭Chapter 23: Ecology, Populations, and Biodiversity‬

‭ igure 23.29‬ ‭This photograph shows the Gibbon wolf‬‭pack in Yellowstone National Park‬
F
‭o n March 1, 2007. Wolves have been identified as a keystone species. (credit: Doug Smith,‬
‭NPS;‬‭OpenStax‬‭)‬

‭ he measured benefits of dam removal include restoration of naturally fluctuating water‬

T
‭levels (often, the purpose of dams is to reduce variation in river flows), which leads to‬
‭increased fish diversity and improved water quality. In the Pacific Northwest, dam removal‬
‭projects are expected to increase populations of salmon, which is considered a keystone‬
‭species because it transports nutrients to inland ecosystems during its annual spawning‬
‭migrations. In other regions, such as the Atlantic coast, dam removal has allowed the‬
‭return of other spawning anadromous fish species (species born in freshwater, live most‬
‭o f their lives in saltwater, and return to freshwater to spawn). Some of the largest dam‬
‭removal projects have yet to occur or have happened too recently for the consequences to‬
‭be measured. The large-scale ecological experiments these removal projects constitute will‬
‭provide valuable data for other dam projects slated for removal or construction.‬
‭ or additional information about approaches to restoring ecosystems and their species,‬
F
‭click the link or scan the QR code to watch the TED-Ed video by George Monobiot titled‬
‭“From the top of the food chain down: Rewilding our world.”‬

From the top of the food chain down: Rewilding our world …

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭695‬
‭Chapter 23: Ecology, Populations, and Biodiversity‬

‭The Role of Zoos and Captive Breeding‬

‭ oos have sought to participate in conservation efforts through captive breeding‬
Z
‭programs and education (‬‭Figure 23.30‬‭). The transformation‬‭o f the missions of zoos from‬
‭collection and exhibition facilities to organizations dedicated to conservation is ongoing. In‬
‭general, it has been recognized that, except in some specifically targeted cases, captive‬
‭breeding programs for endangered species are inefficient and often prone to failure when‬
‭the species are reintroduced to the wild. Zoo facilities are far too limited to contemplate‬
‭captive breeding programs for the number of species now at risk. On the other hand,‬
‭education is a potential positive impact of zoos on conservation efforts, particularly given‬
‭the global trend to urbanization and the consequent reduction in contact between people‬
‭and wildlife. Several studies have been performed to look at the effectiveness of zoos on‬
‭people’s attitudes and actions regarding conservation; at present, the results are mixed.‬
‭ or additional information about approaches to saving endangered species (the Northern‬
F
‭White Rhino), click the link or scan the QR code to watch the TED-Ed video by Jan Stejskal‬
‭titled “The last living members of an extinct species.”‬

The last living members of an extinct species - Jan Stejskal

‭ igure 23.30‬ ‭Zoos and captive breeding programs‬‭help preserve many endangered‬
F
‭species, such as this golden lion tamarin. (credit: Garrett Ziegler;‬‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭696‬
‭Chapter 23: Ecology, Populations, and Biodiversity‬

‭23.10 Career Connections - Biogeographer and Ecologist‬

‭Biogeographer‬

‭ iogeography is the study of the distribution of the world’s species in the past and‬
B
‭present. Biogeographers' work is critical to understanding our physical environment, how‬
‭the environment affects species, and how environmental changes impact species‬
‭distribution; it has also been critical to developing modern evolutionary theory.‬
‭Biogeographers need to understand both biology and ecology. They must also be‬
‭well-versed in evolutionary studies, soil science, and climatology.‬

‭ here are three main fields of study under the heading of biogeography: ecological‬
T
‭biogeography, historical biogeography (called paleobiogeography), and conservation‬
‭biogeography. Ecological biogeography studies the current factors affecting the‬
‭distribution of plants and animals. Historical biogeography, as the name implies, studies‬
‭the past distribution of species. On the other hand, conservation biogeography is focused‬
‭o n protecting and restoring species based on known historical and current ecological‬
‭information. Each of these fields considers both zoogeography and phytogeography—the‬
‭past and present distribution of animals and plants.‬

‭Ecologist‬

‭ career in ecology contributes to many facets of human society. Understanding ecological‬

A
‭issues can help society meet the basic human needs of food, shelter, and health care.‬
‭Ecologists can conduct their research in the laboratory and outside in natural‬
‭environments (‬‭Figure 23.31‬‭). These natural environments‬‭can be as close to home as the‬
‭stream running through your campus or as far away as the hydrothermal vents at the‬
‭bottom of the Pacific Ocean. Ecologists manage natural resources such as white-tailed deer‬
‭populations (Odocoileus virginianus) for hunting or aspen (Populus spp.) timber stands‬
‭for paper production. Ecologists also work as educators who teach children and adults at‬
‭various institutions, including universities, high schools, museums, and nature centers.‬
‭Ecologists may also work in advisory positions assisting local, state, and federal‬
‭policymakers to develop ecologically sound laws, or they may develop those policies and‬
‭legislation themselves. To become an ecologist requires at least an undergraduate degree,‬
‭usually in a natural science. The undergraduate degree is often followed by specialized‬
‭training or an advanced degree, depending on the area of ecology selected. Ecologists‬
‭should also have a broad background in the physical sciences and a solid foundation in‬
‭mathematics and statistics.‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭697‬
‭Chapter 23: Ecology, Populations, and Biodiversity‬

‭ igure 23.31‬ ‭This landscape ecologist is releasing‬‭a black-footed ferret into its native‬
F
‭habitat as part of a study. (credit: USFWS Mountain Prairie Region, NPS;‬ ‭OpenStax‬‭)‬

‭ rincipally derived from‬‭Anatomy and Physiology‬‭and‬‭Biology 2e,‬‭both available for free at‬
P
‭https://openstax.org‬‭. © William Cushwa;‬ ‭CC BY-NC-ND‬

‭698‬
Index
Symbols & Equations ADH 282
3' polyA tail 112 Adipose tissue 125
5' cap 112 ADP 156
3:1 F2 phenotypic ratio 552 Adrenal cortex 286
9:3:3:1 F2 phenotypic ratio 556 Adrenal glands 286
negative 55 mV 458 Adrenal medulla 287
Adrenocorticotrophic hormone
negative 70 mV 454 (ACTH) 283
P1V1 = P2V2 256 Advantageous mutations 587
positive 30 mV 455 Afferent arteriole 308
positive delta G 150 Afterbirth 374
p 613 Age structure diagrams 672
p+q=1 613 Agglutination 201
p2 613 Albumin 196
q 613 Alcohol fermentation 170
q2 613 Aldosterone 283
p2 + 2pq + q2 = 1 613 Alfred Russell Wallace 581
2pq 613 Allele fixation 612
Allele frequency 591
Alleles 198
A Allergy 536
Accessory organs 127 Allopatric speciation 609
Acclimatization 271 All-or-none principle 458
Acetyl CoA 162 Alveolar sacs 252
Acetyl group 162 Alzheimer's disease 467
Acetylcholine (Ach) 424 Amino acid-derived hormones 277
Acetylcholinesterase (AChE) 426 Amino acids 56
Acid reflux 130 Amino group 47
Acidic 45 Ammonia 174
Acrosome 354 Amplitude 493
Actin 422 Amylase 138
Action potential 423 Amyotrophic lateral sclerosis (ALS) 470
Activation energy 153 Anabolic reactions 151
Active immunity 525 Anabolic steroids 293
Active sites 61 Anabolism 12
Active transport 80 Analogous structures 595
Active transport pumps 450 Anaphase 321
Acute mountain sickness 271 Anaphase I 133
Adaptation 20 Anaphylactic shock 536
Adaptation 586 Anatomy 6
Adaptive evolution 614 Anemia 189
Adaptive immune response 50 Aneuploid 344
Addison's disease 311 Angina 211
Adenine 63 Angina pectoris 228
Adenosine triphosphate (ATP) 66 Anion 39
Annealing 627 Autosomal dominant 576

699
Index
Anterior pituitary gland 281 Autosomal recessive 574
Antibiotics 74 Autotrophs 652
Antibodies 201 Axial skeleton 389
Antidepressant 476 Axon 424
Antidiuretic hormone (ADH) 282 Axon terminal 447
Antigen-presenting cell (APC) 515
Antigens 515 B
Aorta 210 B cells 518
Aortic semilunar valve 211 Bacterial cloning 625
Apex consumer 647 Balloon angioplasty 245
Aplastic anemia 192 Bands 622
Apoptosis 509 Barometer 254
Appendicular skeleton 394 Barr body 346
Appendix 134 Barrier defenses 507
Aquaporin channel proteins 307 Barrier methods 375
Arteries 209 Basic/Alkaline 45
Arterioles 234 Basilar membrane 495
Artery 234 Bicarbonate 136
Articulation 395 Bicarbonate buffer system 262
Artificially-acquired immunity 525 Bicarbonate ions 262
Asexual reproduction 314 Biconcave disks 185
Asthma 265 Bicuspid valve 211
Atherosclerosis 211 Bile 135
Atmospheric pressure 254 Binomial nomenclature 603
Atom 29 Biodiversity 675
Atomic mass unit (amu) 30 Biodiversity hotspot 691
Atomic number 32 Biogeochemical carbon cycle 658
ATP 66 Biogeochemical cycles 653
ATP synthase 165 Biological carbon cycle 657
ATP-ADP cycle 156 Biology 6
Atria 209 Biome 646
Atrioventricular (AV) node 220 Biotechnology 620
Atrioventricular valve (AV) 213 Bladder 301
Atrophy 432 Blending theory of inheritance 541
Attention-deficit/hyperactivity
disorder (ADHD) 474 Blood 125
Audition 492 Blood pressure (BP) 229
Auditory ossicles 392 Blood transfusion 202
Auscultation 219 Bolus 128
Autism spectrum disorder (ASD) 473 Bone fractures 406
Autoantibodies 537 Bone marrow 196
Autoimmunity 537 Bone marrow transplant 197
Autonomic nervous system (ANS) 444 Bone remodeling 406
Autorhythmicity 220 Bones 125

700
Index
Brain 441 Cell cycle 317
Branch point 605 Cell cycle checkpoint 338
Brightness 499 Cell membrane 450
Bronchioles 252 Cell-mediated immune response 515
Buck v. Bell 381 Cellular portion 181
Buffy coat 184 Cellular respiration 159
Bulbourethral gland 352 Cellulose 51
Bypass surgery 243 Central canal 399
Central dogma 109
C Central nervous system (CNS) 442
Calcitonin 410 Centrifuge 184
Calcium 409 Cervix 362
Calcium ions 424 Chain termination method 633
Calorie 175 Channel proteins 81
cAMP 279 Chargaff's rules 106
Canaliculi 400 Charles Darwin 581
Cancellous bone 400 Chemical bond 7
Cancer 338 Chemical digestion 137
Capillaries 185 Chemical diversity 676
Capsule 304 Chemical energy 147
Carbaminohemoglobin 187 Chemical symbols 28
Carbohydrate 49 Chemical synapses 462
Carbon cycle 256 Chemoautotrophs 652
Carbon dioxide 658 Cholecystokinin (CCK) 143
Carbon monoxide poisoning 268 Cholesterol 55
Carbonic anhydrase 262 Chordae tendineae 216
Carboxyl group 47 Chromatin 317
Cardiac cycle 217 Chromosome compaction 317
Cardiac muscle 220 Chromosomes 102
Cardiac sphincter 130 Chyme 131
Cardiac veins 241 Class 602
Cardiopulmonary resuscitation 225 Cleavage furrow 323
Carnivores 647 Clitoris 358
Carrier proteins 81 Clonal selection 519
Carriers 560 Clot 195
Carrying capacity 668 Clotting factors 195
Cas nuclease 641 Coccyx 392
Catabolic reactions 151 Cochlea 495
Catabolism 12 Cochlear nerve 495
Catalyst 153 Codominance 562
Cation 39 Codominant 199
Cecum 134 Codon 112
Cell 7 Coefficients 159
Cell body 447 Coenzyme A 12

701
Index
Coenzymes 158 D
Collecting ducts 304 Decompression sickness 270
Colon 134 Degenerate 115
Dehydration synthesis
Community 645 (Condensation) 48
Compact bone 399 Deleterious mutations 587
Complement system 514 Deletions 348
Complementary 105 Demography 661
Compound 28 Denaturation 59
Concentration gradient 80 Dendrites 447
Conclusion 24 Deoxyhemoglobin 186
Condensation 655 Deoxynucleotides (dNTPs) 633
Cones 501 Deoxyribonucleic Acid (DNA) 7
Connective tissue 125 Deoxyribose 50
Constipation 145 Dephosphorylation 156
Continuous conduction 461 Depolarization 455
Continuous variation 542 Depolarize 424
Contraception 375 Detrital food web 651
Control center 16 Development 14
Control group 23 Diabetes insipidus 298
Cornea 499 Diabetes mellitus 298
Coronary arteries 211 Dialysis 312
Coronary circuit 209 Diaphragm 251
Coronary veins 211 Diaphysis 401
Corpus luteum (CL) 365 Diarrhea 145
Cortex 483 Diastole 217
Covalent bond 40 Diastolic pressure 229
COVID 526 Dideoxynucleotides (ddNTPs) 633
CPAP machine 269 Diffusion 80
Cranial bones 389 Diploid 314
C-reactive protein 513 Diploid number (2n) 315
Creatine phosphate 435 Diploid-dominant life-cycle strategy 326
CRISPR 641 Directional selection 616
Cross-bridge contraction cycle 431 Disaccharidases 140
Cross-bridges 433 Disaccharide 49
Crossing over 329 Discontinuous variation 542
Cryptorchidism 350 Discussion 24
Cytokines 513 Distal convoluted tubule (DCT) 307
Cytokinesis 319 Diuretics 311
Cytoplasm 92 Diversifying selection 616
Cytoplasmic receptors 277 Dizygotic twins 370
Cytosine 63 DNA ligase 627
Cytoskeleton 92 DNA polymerase 108
Cytosol 91 DNA replication 108
Cytotoxic T cells 517 DNA sequencing 633

702
Index
Dolly 631 Enzymatic digestion 137
Domain 602 Enzymes 61
Dominant 198 Enzyme-substrate complex 61
Dominant traits 546 Epididymis 351
Double circulation 211 Epiglottis 128
Double covalent bond 41 Epilepsy 477
Double helix 63 Epimysium 419
Down syndrome 345 Epinephrine 275
Duchenne muscular dystrophy 439 Epiphyseal line 402
Duodenum 132 Epiphyseal plate 402
Dup 217 Epistasis 569
Dynamic equilibrium 122 Epithelial tissue 123
Dystrophin 439 Epitopes 518
Equational division 328
E Erectile dysfunction (ED) 353
Ecology 645 Erectile tissue 353
Ecosystem 645 Erythrocytes 181
Ecosystem diversity 677 Erythropoietin (EPO) 188
Ectopic pregnancy 360 Esophagus 130
Effector 16 Essential amino acids 58
Effector cells 521 Estrogen 283
Efferent arteriole 308 Eugenics 381
Electrocardiogram (EKG) 222 Eukaryotes 73
Electromagnetic spectrum 498 Eukaryotic cell 75
Electron shells 35 Euploid 344
Electron transport chain 164 Eutrophication 661
Electrons 30 Evaporation 655
Elements 28 Evolution 580
Elimination 145 Evolution by natural selection 581
Elongation 108 Excitation-contraction coupling 424
Embryology 597 Excitatory neurotransmitter 463
Embryonic stem cells 631 Exergonic reactions 149
Emulsification 138 Exhalation 248
Endergonic reactions 150 Exocrine system 277
Endocrine gland 276 Exocytosis 88
Endocrine system 274 Exon 112
Endocytosis 87 Exotic species 685
Endomembrane system 95 Experiments 23
Endometriosis 361 Exponential growth 666
Endometrium 361 Expressed unit factor 548
Endomysium 420 External respiration 260
Endoplasmic reticulum (ER) 95 Extract 621
Endosteum 403
Energy 147

703
Index
F Gallbladder 135
Facilitated diffusion 82 Gametes 283
FAD 158 Gastrin 143
FADH2 158 Gastrointestinal tract 127
Falsifiable 23 Gel electrophoresis 621
Family 602 Gene editing 641
Farsighted 499 Gene expression 109
Fascicle 420 Gene flow 593
Fatty acids 53 Gene pool 612
Fermentation 169 Gene therapy 636
Fertilization 314 Genes 109
Fetus 373 Genetic diagnosis 636
Fibrin 195 Genetic diversity 676
Fibrinogen 195 Genetic drift 591
Fimbriae 360 Genetic recombination 329
First filial generation (F1) 543 Genetic testing 636
First law of thermodynamics 12 Genome 315
First messenger 277 Genomics 640
Flagellum 354 Genotype 198
Flatulence 143 Genus 602
Flight-or-fight response 287 Germ cells 327
Fluid mosaic model 77 Gestation 371
Follicles 283 Gibbs free energy 149
Follicle-stimulating hormone (FSH) 283 Glial cells 447
Follicular phase 365 Globin 186
Food chain 647 Globulin 196
Food web 649 Glomerular filtration 309
Formed elements 181 Glomeruli 296
Fovea 502 Glomerulus 305
Frameshift mutation 118 Glottis 128
Frequency 493 Glucagon 288
Freshwater 654 Glucocorticoid hormones 286
Freshwater ecosystems 645 Glucose 50
Fructose 50 Glycerol 53
FSH 283 Glycogen 51
Functional group 47 Glycolysis 160
GMOs 637
G GnRH 283
G0 phase 320 Goiter 290
G1 checkpoint 325 Golgi apparatus 96
Gonadotropin-releasing hormone
G1 phase 319 (GnRH) 283
G2 checkpoint 325 Gonads 289
G2 phase 320 Graafian follicle 359
Galactose 50 Graves' disease 291

704
Index
Gray matter 448 Human biology 6
Grazing food web 651 Human Genome Project (HGP) 633
Groundwater 655 Human growth hormone (hGH) 438
Human immunodeficiency virus
Growth 15 (HIV) 535
Growth hormone (GH) 282 Humoral immune response 523
Guanine 63 Hybridize 542
Gustation 486 Hydrochloric acid 138
Hydrogen bond 43
H Hydrogen ions 45
Half-life 35 Hydrolysis 48
Haploid 314 Hydrophilic 44
Haploid number (n) 315 Hydrophilic heads 78
Hardy-Weinberg equilibrium 612 Hydrophobic 44
Haversian canal 399 Hydrophobic tails 78
Heart attack 211 Hydroxyl group 48
Heat energy 149 Hyoid bone 392
Heimlich maneuver 128 Hyperbaric chamber 270
Helper T cells 518 Hypercalcemia 410
Hematocrit 184 Hypersensitivities 537
Heme 186 Hypertension 231
Hemizygous 566 Hyperthyroidism 291
Hemoglobin 186 Hypertonic 84
Hemoglobin saturation 261 Hypertrophy 433
Hemolysis 201 Hypocalcemia 410
Hemolytic disease of the newborn 205 Hypothalamus 281
Hemophilia 195 Hypothesis 22
Hemorrhagic stroke 246 Hypothyroidism 290
Herbivores 647 Hypotonic 86
Heterozygous 198 Hypoxemia 187
Histone proteins 102
Homeostasis 15 I
Homo sapiens 609 Ileocecal sphincter 132
Homologous chromosomes 316 Immunodeficiency 535
Homologous structures 594 Immunoglobulins 523
Homozygous 198 Implantation 360
Homozygous dominant 553 in vitro fertilization (IVF) 377
Homozygous recessive 553 Inbred 542
Hormonal methods 376 Incomplete dominance 561
Hormone replacement therapy
(HRT) 369 Incus 494
Hormones 274 Independent assortment 331
Host 505 Induced fit 61
Hue 498 Induced mutations 109
Human beta chorionic gonadotropin 372 Inferior vena cava 209

705
Index
Infertility 377 Kinetochore 321
Inflammation 509 Kingdom 602
Inhalation 249 Klinefelter syndrome 347
Inhibin 357 Korotkoff sounds 229
Inhibitory neurotransmitter 463 Krebs/citric acid cycle 163
Initiation 108
Innate immune response 506 L
Inner ear 495 Labia majora 358
Insertion 118 Labia minora 358
Insulin 288 Lactase 140
Integrated functioning 120 Lactic acid (lactate) 169
Integration 443 Lactose 50
Integumentary system 485 Lactose intolerance 142
Interferons 513 Lacunae 400
Internal respiration 260 Lamellae 399
Interphase 318 Large intestine 134
Interstitial cells 354 Larynx 249
Intrauterine device (IUD) 377 Latent unit factor 548
Intrinsic factor 139 Law of conservation of energy 149
Introduction 24 Law of dominance 549
Intron 112 Law of independent assortment 556
Invasive species 685 Law of segregation 552
Iodine 283 Leakage channel 453
Ion 39 Left atrium 211
Ion channels 423 Left ventricle 211
Ionic bond 39 Lens 499
Ions 39 Leukemia 193
Iris 499 Leukocytes 181
Iron 186 Leukopenia 194
Iron deficiency anemia 190 LH 283
Ischemic stroke 246 LH surge 364
Isotonic 84 Life tables 664
Isotope 34 Ligaments 125
Ligand 451
J Ligand-gated channel 451
Johann Gregor Mendel 541 Ligand-gated sodium channel 458
J-shaped growth curve 667 Lingual lipase 138
Linkage 571
K Linkage maps 572
Karyotype 315 Lipid bilayer 78
Ketones 299 Lipid-derived hormones 277
Keystone species 694 Liver 135
Kidneys 299 Loci 316
Kinetic energy 149 Logistic growth 666

706
Index
Long bones 401 Middle ear 494
Loop of Henle 306 Millimeters of mercury (mm Hg) 254
Lumen 144 Millivolts (mV) 453
Lup 217 Mineralocorticoid 283
Luteal phase 365 Missense mutations 118
Lymph 527 Mitochondria 94
Lymph fluid 240 Mitosis 319
Lymphatic vessels 240 Mitotic phases 321
Lymphocytes 517 Mitral valve 211
Lymphoma 193 Molecules 38
Lysis 621 Monocyte 509
Lysozyme 97 Monohybrid cross 550
Monomers 7
M Monosaccharide 49
Macromolecules 7 Monosomy 344
Macrophage 189 Monounsaturated fatty acids 53
Major depression 476 Monozygotic twins 370
Malleus 494 Motor division 443
Maltase 140 Motor responses 443
Maltose 50 mRNA 621
Marine ecosystems 645 Multiple sclerosis (MS) 467
Mark and recapture 663 Muscle relaxation 430
Mass number 32 Muscle tissue 126
Materials and methods 24 Mutation 580
Mechanical digestion 137 Mutations 109
Medulla 304 Myelin 447
Medullary cavity 401 Myocardial infarction (MI) 211
Meiosis 326 Myocardium 211
Meiosis I 328 Myofibrils 421
Meiosis II 333 Myofilaments 422
Membrane potential 423 Myoglobin 436
Membrane-bound receptors 277 Myometrium 362
Memory cells 521 Myosin 422
Menopause 367
Menstrual cycle 365 N
Mental Illnesses 475 NAD+ 158
Messenger RNA (mRNA) 93 NADH 158
Metabolism 147 Nasal cavity 249
Metaphase 321 Natural killer (NK) cells 509
Metaphase checkpoint 325 Natural selection 582
Metaphase I 331 Naturally-acquired immunity 525
Metaphase plate 331 Nearsighted 499
Metastasis 341 Negative delta G 149
Microvilli 78 Negative electrode 621

707
Index
Negative feedback 183 Olfaction 486
Negative feedback control system 357 Olfactory receptors 487
Negative feedback loops 122 Oncogene 338
Nephrons 296 One-way valves 238
Nervous tissue 127 Oocyte 358
Net diffusion 84 Oogenesis 358
Neurodegenerative disorders 467 Optic nerve 500
Neurodevelopmental disorders 473 Oral cavity 127
Neuromuscular junction (NMJ) 424 Order 602
Neurons 127 Organ 9
Neurotransmitter 424 Organ of Corti 495
Neutral 45 Organ system 9
Neutral mutations 587 Organelle 91
Neutrons 7 Organic compounds 47
Neutrophil 509 Organism 9
Nitric oxide 354 Organs 9
Nitrogen cycle 659 Osmosis 84
Nitrogen fixation 659 Ossification (Osteogenesis) 405
Node of Ranvier 447 Osteoblasts 404
Noncellular portion (matrix) 181 Osteoclasts 405
Non-coding regions 113 Osteocytes 399
Nondisjunction 343 Osteogenesis imperfecta (OI) 415
Nonpolar covalent bond 41 Osteogenic cells 404
Nonpolar molecule 41 Osteons 399
Nonsense mutations 118 Osteoporosis 414
Norepinephrine 275 Outer ear 494
Normal range 15 Ova 358
Northern blotting 623 Oval window 495
Nostrils 249 Ovarian cycle 365
Nuclear envelope 92 Ovaries 358
Nuclear pore 102 Oviducts 360
Nucleases 140 Ovulation 365
Nucleic acids 63 Oxidation 157
Nucleolus 92 Oxidative phosphorylation 157
Nucleoplasm 92 Oxyhemoglobin 186
Nucleosome 102 Oxytocin (OT) 275
Nucleotides 621
Nucleus 7 P
Nylon membrane 623 P wave 222
Pain receptors 485
O Palindromic sequence 626
Observation 21 Pancreas 136
Octet rule 37 Pancreatic islets 288
Odorants 487 Pancreatic lipase 140

708
Index
Papillae 490 Photoreceptors 501
Parasympathetic pathway 444 Photosynthesis 147
Parathyroid glands 410 Phylogenetic tree 605
Parathyroid hormone (PTH) 410 Phylogeny 603
Parental generation (P1) 543 Phylum 602
Parkinson's disease 469 Physical digestion 137
Partial pressure 254 Physiology 6
Passive immunity 524 Pinna 494
Passive transport 80 Pitch 493
Passive transport channels 450 Pituitary gland 281
Pathogens 505 Placenta 373
Pectoral girdle 398 Plasma 181
Pedigree analysis 573 Plasma cells 523
Peer-reviewed 23 Plasma membrane 76
Pelvic girdle 398 Plasmid 625
Penis 353 Platelets 181
Pepsin 138 Pneumonia 267
Pepsinogen 139 Point mutations 118
Peptide bond 56 Polar body 364
Peptide-derived hormones 277 Polar covalent bonds 41
Percent saturation 187 Polar molecule 41
Perception 483 Polarity 447
Periodic table 32 Polycythemia 192
Periosteum 403 Polycythemia vera 192
Peripheral nervous system (PNS) 442 Polygenic inheritance 569
Peristalsis 130 Polymer 7
Peritubular capillary network 308 Polymerase Chain Reaction (PCR) 627
pH 45 Polysaccharide 49
pH scale 45 Polysome or Polyribosome 115
Phagocyte 509 Polyspermy 370
Phagocytosis 509 Polyunsaturated fatty acids 53
Pharmacogenomics 643 Population density 661
Pharynx 128 Population genetics 611
Phenotype 198 Population size 661
Phenylalanine 178 Positive electrode 621
Phenylketonuria 178 Positive feedback 18
Pheromone 488 Positive feedback loops 18
Phosphate group 47 Posterior pituitary gland 282
Phosphodiester bonds 621 Postpartum period 374
Phospholipid bilayer 79 Potassium ions (K+1) 450
Phospholipids 55 Potential energy 149
Phosphorylate 66 Prader-Willi syndrome 178
Phosphorylation 157 Precipitate 621
Photoautotrophs 652 Precipitation 655

709
Index
Pre-mRNA 112 R
Prenatal genetic diagnosis (PGD) 385 R group 57
Preserve 691 RBC 181
Primary bronchi 249 Reactant 61
Primary consumers 647 Reanneal 621
Primary oocyte 364 Reception 481
Primary response 520 Receptor potential 481
Primary structure 58 Receptors 481
Primers 627 Recessive 198
Probabilities 550 Recessive traits 546
Probe 624 Reciprocal cross 545
Probes 624 Recombinant plasmid 627
Producers 647 Recombination 571
Product rule 558 Rectum 134
Progesterone 283 Red bone marrow 196
Prokaryotes 73 Reduction 158
Prolactin (PRL) 283 Reductional division 328
Prometaphase 321 Reflexes 444
Propagated 459 Refractory period 458
Prophase 321 Relative fitness 614
Prophase I 329 Releasing hormones 282
Prostate cancer 352 Renal arteries 308
Prostate gland 351 Renal corpuscle 305
Protein-derived hormones 277 Renal pelvis 304
Proteins 56 Renal pyramids 304
Protons 7 Renal tubule 305
Proto-oncogenes 338 Renal veins 308
Proximal convoluted tubule (PCT) 306 Repolarization 455
Pulmonary arteries 209 Reproduction 15
Pulmonary circuit 209 Reproductive cloning 631
Pulmonary semilunar valve 211 Resilience 647
Pulmonary veins 211 Resistance 647
Pulse 232 Respiratory bronchioles 252
Punnett square 550 Responsiveness 13
Pupil 499 Resting membrane potential 454
Purines 63 Resting state 454
Pyloric sphincter 131 Restriction enzymes 626
Pyrimidines 63 Results 23
Pyruvate 160 Retina 499
Rh group 199
Q Rh negative 200
QRS complex 222 Rh positive 200
Quaternary structure 59 RhoGAM 205
Rib cage 390

710
Index
Ribonucleic acid (RNA) 63 Sensory nerve ending 481
Ribose 66 Sensory receptor 481
Ribosomal RNA (rRNA) 111 Sensory transduction 481
Ribosomes 93 Sertoli cells 354
Ribs 390 Set point 15
Right atrium 210 Sex chromosomes 317
Right ventricle 210 Sexual reproduction 326
RNA polymerase 110 Sickle cell anemia 190
Rods 501 Signal summation 464
Rough ER 96 Silent mutations 118
Simple diffusion 81
S Single covalent bond 41
S phase 319 Sinoatrial (SA) node 220
Sacrum 392 Sister chromatids 106
Salivary amylase 138 Skeletal muscle 126
Salivary glands 128 Skeletal muscle fiber 421
Saltatory conduction 461 Skin 485
Saltwater 654 Skull 389
Sarcolemma 421 Sleep apnea 269
Sarcomere 422 Sliding filament model 427
Sarcoplasmic reticulum (SR) 424 Small intestine 132
SARS-CoV-2 coronavirus 526 Smooth ER 95
Saturated fatty acids 53 Smooth muscle 126
Schizophrenia 475 Sodium ions (Na+1) 450
Schwann cells 447 Sodium-potassium pump 450
Scientific method 21 Solar energy 147
Scientific name 603 Solutes 84
Scrotum 350 Somatic cell 315
Second filial generation (F2) 543 Somatic nervous system (SNS) 444
Second law of thermodynamics 149 Somatosensation 484
Second messenger 277 Somatosensory receptors 485
Secondary consumers 647 Southern blotting 623
Secondary oocyte 364 Speciation 608
Secondary response 521 Species 608
Secondary structures 59 Species diversity 677
Secretin 143 Sperm 289
Selectively permeable 80 Spermatogenesis 354
Semen 351 Spermicides 375
Semi-conservative replication 106 Sphincter 130
Seminal vesicles 351 Sphygmomanometer 229
Seminiferous tubules 354 Spinal column 392
Sensation 443 Spinal cord 441
Sensor 16 Spliceosome 112
Sensory division 443 Splicing 112

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Index
Spongy bone 400 Target cells 274
Spontaneous mutations 109 Tastants 491
SRY gene 370 Taste buds 491
S-shaped curve 668 Taxonomy 602
Stabilizing selection 616 Tectorial membrane 495
Stapes 494 Telophase 321
Starch 51 Telophase I 333
Start codon 115 Telophase II 333
Stereocilia 495 Template strand 110
Sternum 392 Tendons 125
Steroid 55 Terminal electron acceptor 164
Stethoscope 219 Termination 108
Sticky ends 627 Terrestrial ecosystems 646
Stomach 131 Tertiary consumers 647
Stop codon 115 Tertiary structure 59
Stroke 246 Test cross 554
Structural genes 109 Testable 22
Subatomic particles 7 Testes 289
Substrate-level phosphorylation 157 Testis 350
Substrates 61 Testosterone 356
Sucrase 140 Tetrads 329
Sucrose 50 Thalamus 483
Sugar-phosphate backbone 63 Thalassemia 192
Sum rule 559 The Virginia Sterilization Act 381
Superior vena cava 209 Thick filaments 422
Suppressor T cells 518 Thin filaments 422
Surface area 72 Threshold 458
Surface area to volume ratio 72 Thrombocytes 181
Survival of the fittest 584 Thrombosis 195
Sympathetic pathway 444 Thymine 63
Sympatric speciation 609 Thymus 531
Synapse 462 Thyroid gland 283
Synapsis 329 Thyroid-stimulating hormone (TSH) 283
Synaptic cleft 424 Thyroxine (T4) 283
Synaptic end bulb 448 Tinnitus 497
Synaptic vesicles 462 Triplet code 112
Systemic circuit 209 Tissue 7
Systole 217 Tonicity 84
Systolic blood pressure 229 Trabeculae 401
Trachea 249
T Trans fats 54
T cells 517 Transcription 110
T wave 222 Transduction 481
Taq DNA polymerase 629 Transfer RNA (tRNA) 111

712
Index
Transgenic 637 Vas deferens 351
Transition reaction 161 Vasectomy 351
Translation 112 Vasoconstriction 237
Transverse (T) tubules 426 Vasodilation 237
Tricuspid valve 210 Vector 625
Triglyceride 52 Veins 209
Triiodothyronine (T3) 283 Ventricles 209
Triple-X 347 Venules 234
Trisomy 344 Vesicle 87
Trisomy 21 345 Vestigial structures 595
tRNA anticodon 115 Virus 636
Trophic levels 647 Visible light spectrum 498
Tropomyosin 429 Vitamin deficiency anemias 191
Troponin 429 Vitamin K 195
True breeding 542 Voltage-gated channel 451
Trypsin 140 Voltage-gated sodium channel 424
Tubal ligation 361 Volume 72
Tubular reabsorption 309 Voluntary motor response 444
Tubular secretion 309 Voluntary muscle 419
Tumor suppressor genes 339
Turner syndrome 347 W
Tympanum 494 Water cycle 654
Type 1 (juvenile) diabetes 299 Wavelength 493
Type 2 (adult onset) diabetes 299 WBC 181
Western blotting 624
U White matter 448
Ulcers 139
Unified cell theory 70 X
Universal donor 203 X inactivation 345
Universal recipient 203 X-linked 565
Uracil 63 X-linked dominant 577
Urea 174 X-linked recessive 577
Ureter 300
Urethra 301 Y
Urinalysis 297 Yellow marrow 401
Urine 296
Uterus 360 Z
Z lines 422
V Zygote 314
Vaccines 526
Vagina 362
Valence shell 36
Variable 23
Varicose veins 239

713