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Detailed Contents

1 Biology: The Study of Life 1

1.1 What Does It Mean to Say that Something Is

Alive? 2
1.2 Life Is Cellular and Replicates through Cell
Division 2
All Organisms Are Made of Cells 2
Where Do Cells Come From? 3
Life Replicates through Cell Division 4
1.3 Life Processes Information and Requires
Energy 4
The Central Dogma 5
Life Requires Energy 5 B.6 Separating and Visualizing Molecules 30
Using Electrophoresis to Separate Molecules 30
1.4 Life Evolves 6 Using Thin Layer Chromatography to Separate Molecules 32
What Is Evolution? 6 Visualizing Molecules 32
What Is Natural Selection? 6
B.7 Separating Cell Components by Centrifugation 34
1.5 The “Tree of Life” Depicts Evolutionary History 7
Using Genetic Sequences to Understand the Tree of Life 7 B.8 Using Spectrophotometry 35
How Should We Name Branches on the Tree of Life? 9 B.9 Using Microscopy 36
1.6 Doing Biology 10 Light and Fluorescence Microscopy 36
The Nature of Science 10 Electron Microscopy 36
An Introduction to Hypothesis Testing: Why Do Giraffes Have Studying Live Cells and Real-Time Processes 38
Long Necks? 10 B.10 Using Molecular Biology Tools and Techniques 39
An Introduction to Experimental Design: How Do Ants Making and Using cDNA Libraries 39
Navigate? 12 Amplifying DNA Using the Polymerase Chain Reaction
CHAPTER 1 REVIEW 15 (PCR) 40
Dideoxy Sequencing 41
END-OF-UNIT
CASE STUDY Mystery of the Newt 17 Shotgun Sequencing 42
DNA Microarrays 43
Doing Biology 18 B.11 Using Cell Culture and Model Organisms as Tools 44
Cell and Tissue Culture Methods 44
Model Organisms 45

BioSkills 20 B.12 Reading and Making Visual Models 48

Tips for Interpreting Models 48
B.1 Using the Metric System and Significant Figures 21 Tips for Making Your Own Models 49
Metric System Units and Conversions 21 Concept Maps 49
Significant Figures 22
B.13 Reading and Making Phylogenetic Trees 50
B.2 Reading and Making Graphs 23 Anatomy of a Phylogenetic Tree 50
Getting Started 24 How to Read a Phylogenetic Tree 51
Types of Graphs 25 How to Draw a Phylogenetic Tree 51
Getting Practice 26
B.14 Reading Chemical Structures 52
B.3 Interpreting Standard Error Bars and Using
B.15 Translating Greek and Latin Roots in Biology 53
Statistical Tests 26
Standard Error Bars 26 B.16 Reading and Citing the Primary Literature 53
Using Statistical Tests 27 What Is the Primary Literature? 53
Interpreting Differences: P Values and Statistical Getting Started 53
Significance 28 Citing Sources 55
Evaluating Causation versus Correlation 28 Getting Practice 55
B.4 Working with Probabilities 29 B.17 Recognizing and Correcting Misconceptions 55
The “Both-And” Rule 29
B.18 Using Bloom’s Taxonomy for Study Success 56
The “Either-Or” Rule 29
Categories of Human Cognition 56
B.5 Using Logarithms 29 Six Study Steps to Success 56

v

 How Can Biologists Use Phylogenetic Trees to Study the History Investigating the Human Microbiome 540
of Life? 514 Evaluating Molecular Phylogenies 541
Case Study: Where Do Whales Belong on the Tree of Life? 515
26.3 What Themes Occur in the Diversification of
25.2 Tools for Studying Life’s History: The Fossil Bacteria and Archaea? 541
Record 517 Genetic Variation through Gene Transfer 541
Biologists Study Many Types of Fossils 517 Morphological Diversity 543
What Are the Opportunities and Limitations of the Fossil Metabolic Diversity 544
Record? 518 Ecological Diversity and Global Impacts 547
How Are Fossils Used to Estimate Life’s Time Line? 519
26.4 Key Lineages of Bacteria and Archaea 550
25.3 Large-Scale Pattern in Life’s History: Adaptive Bacteria 550
Radiation 521 Archaea 550
Why Do Adaptive Radiations Occur? 522
CHAPTER 26 REVIEW 553
The Cambrian Explosion 522
25.4 Large-Scale Pattern in Life’s History: Mass
Extinction 525
27 Diversification of
How Do Mass Extinctions Differ from Background Eukaryotes 555
Extinctions? 525
27.1 Why Do Biologists Study Protists? 556
The End-Permian Extinction 525
Impacts on Human Health and Welfare 556
The End-Cretaceous Extinction 526
Ecological Importance of Protists 558
The Sixth Mass Extinction 527
27.2 How Do Biologists Study Protists? 559
CHAPTER 25 REVIEW 527
Microscopy: Studying Cell Structure 560
END-OF-UNIT Are Garter Snakes and Newts Engaged in an Evaluating Molecular Phylogenies 560
CASE STUDY
Arms Race? 530 Discovering New Lineages via Direct Sequencing 561
Evolution 532 27.3 What Themes Occur in the Diversification of
Protists? 562
What Morphological Innovations Evolved in Protists? 562
How Do Protists Obtain Food? 566
UNIT
THE DIVERSIFICATION OF LIFE 534 How Do Protists Move? 567
5 How Do Protists Reproduce? 568

26 Bacteria and Archaea 534

27.4 Key Lineages of Eukaryotes 571
Amoebozoa 571
Opisthokonta 571
26.1 Why Do Biologists Study Bacteria and Archaea? 535
Excavata 571
Biological Impact 535
Plantae 572
Some Prokaryotes Thrive in Extreme Environments 536
Rhizaria 573
Medical Importance 536
Alveolata 573
Role in Bioremediation 538
Stramenopila (Heterokonta) 574
26.2 How Do Biologists Study Bacteria and Archaea? 539
CHAPTER 27 REVIEW 575
Using Enrichment Cultures 539

28 Green Algae and Land

Using Metagenomics 540

Plants 577

28.1 Why Do Biologists Study Green Algae and Land

Plants? 578
Plants Provide Ecosystem Services 578
Plants Provide Humans with Food, Fuel, Fiber, Building
Materials, and Medicines 579
28.2 How Do Biologists Study Green Algae and Land
Plants? 580
Analyzing Morphological Traits 580
Using the Fossil Record 582
Evaluating Molecular Phylogenies 583
28.3 What Themes Occur in the Diversification of Land
Plants? 584
The Transition to Land I: How Did Plants Adapt to Dry
Conditions with Intense Sunlight? 584
Mapping Evolutionary Changes on the Phylogenetic Tree 587
The Transition to Land II: How Do Plants Reproduce in Dry
Conditions? 587
The Angiosperm Radiation 596

xii DETAILED CONTENTS

28.4 Key Lineages of Green Algae and Land Plants 597
Green Algae 598
Nonvascular Plants 598
Seedless Vascular Plants 598
Seed Plants: Gymnosperms and Angiosperms 598
CHAPTER 28 REVIEW 604

29 Fungi 606

29.1 Why Do Biologists Study Fungi? 607

Fungi Have Important Economic and Ecological
Impacts 607
Mycorrhizal Fungi Provide Nutrients for Land Plants 608
Saprophytic Fungi Accelerate the Carbon Cycle
on Land 609
29.2 How Do Biologists Study Fungi? 609
Analyzing Morphological Traits 610
Evaluating Molecular Phylogenies 612
29.3 What Themes Occur in the Diversification of
Fungi? 614
Fungi Often Participate in Symbioses 615
What Adaptations Make Fungi Such Effective
31 Protostome Animals 650

Decomposers? 618 31.1 What Is a Protostome? 651

Variation in Reproduction 619 The Water-to-Land Transition 652
Four Major Types of Life Cycles 621 Compartmentalized and Flexible Body Plans 653

29.4 Key Lineages of Fungi 624 31.2 What Is a Lophotrochozoan? 654

Microsporidia 624 What Is a Flatworm? 655
Chytrids 625 What Is a Segmented Worm? 657
Zygomycetes 625 What Is a Mollusk? 658
Glomeromycota 626 31.3 What Is an Ecdysozoan? 661
Basidiomycota 626 What Is a Roundworm? 662
Ascomycota 626 What Are Water Bears and Velvet Worms? 663
CHAPTER 29 REVIEW 626 What Is an Arthropod? 663
Arthropod Diversity 665

30 An Introduction to
Arthropod Metamorphosis 668
Take-Home Messages 669

Animals 629 CHAPTER 31 REVIEW 670

30.1 What Is an Animal? 630

30.2 What Key Innovations Occurred during the Origin of 32 Deuterostome Animals 672
Animal Phyla? 631
Origin of Multicellularity 632
32.1 What Is a Deuterostome? 673
Origin of Embryonic Tissue Layers and Muscle 634 32.2 What Is an Echinoderm? 674
Origin of Bilateral Symmetry, Cephalization, and the Nervous The Echinoderm Body Plan 674
System 635 Echinoderms Are Important Consumers 674
Origin of the Gut and Coelom 637
32.3 What Is a Chordate? 676
Origin of Protostomes and Deuterostomes 638
The Cephalochordates 676
Origin of Segmentation 639
The Urochordates 677
30.3 What Themes Occur in the Diversification within The Vertebrates 678
Animal Phyla? 639
32.4 What Is a Vertebrate? 678
Sensory Organs 640
Feeding 641 32.5 What Key Innovations Occurred during the
Movement 642 Evolution of Vertebrates? 679
Reproduction 644 Urochordates: Sister Group to Vertebrates 679
Life Cycles 645 First Vertebrates: Origin of the Cranium and Vertebrae 679
Gnathostomes: Origin of the Vertebrate Jaw 681
30.4 Key Lineages of Animals: Non-Bilaterian
Origin of the Bony Endoskeleton 683
Groups 646
Origin of the Lungs 683
Porifera (Sponges) 646
Tetrapods: Origin of the Limb 684
Ctenophora (Comb Jellies) 647
Amniotes: Origin of the Amniotic Egg 685
Cnidaria (Jellyfish, Corals, Anemones, Hydroids) 647
Mammals: Origin of Lactation and Fur 686
CHAPTER 30 REVIEW 648 Reptiles: Origin of Scales and Feathers Made of Keratin 687

DETAILED CONTENTS xiii

 Parental Care 689 The Root System 726
Take-Home Messages 690 The Shoot System 728
The Leaf 730
32.6 The Primates and Hominins 690
The Primates 690 34.2 Plant Cells and Tissue Systems 733
Fossil Humans 692 The Dermal Tissue System 734
The Out-of-Africa Hypothesis 695 The Ground Tissue System 734
Have Humans Stopped Evolving? 696 The Vascular Tissue System 736
CHAPTER 32 REVIEW 697 34.3 Primary Growth Extends the Plant Body 738
How Do Apical Meristems Produce the Primary Plant

33 Viruses 699
Body? 738
How Is the Primary Root System Organized? 740
33.1 Why Do Biologists Study Viruses? 700 How Is the Primary Shoot System Organized? 741
Viruses Shape the Evolution of Organisms 700 34.4 Secondary Growth Widens Shoots and Roots 741
Viruses Cause Disease 700 What Is a Cambium? 742
Current Viral Pandemics in Humans: AIDS 702 How Does a Cambium Initiate Secondary Growth? 743
33.2 How Do Biologists Study Viruses? 703 What Do Vascular Cambia Produce? 743
Analyzing Morphological Traits 703 What Do Cork Cambia Produce? 744
Analyzing the Genetic Material 704 The Structure of Tree Trunks 744
Analyzing the Phases of Replicative Growth 705 CHAPTER 34 REVIEW 745
Analyzing How Viruses Can Coexist with Host Cells 711
33.3 What Themes Occur in the Diversification of
Viruses? 712 35 Water and Sugar Transport in
Where Did Viruses Come From? 712
Emerging Viruses, Emerging Diseases 712
Plants 747

33.4 Key Lineages of Viruses 714 35.1 Water Potential and Water Movement 748
What Is Water Potential? 748
CHAPTER 33 REVIEW 718 What Factors Affect Water Potential? 748
END-OF-UNIT
Working with Water Potentials 749
CASE STUDY Are Newts Adapted to Kill Humans? 720 Water Potentials in Soils, Plants, and the Atmosphere 750

Diversity of Life 722 35.2 How Does Water Move from Roots to Shoots? 752
Movement of Water and Solutes into the Root 752
Water Movement via Root Pressure 754
UNIT Water Movement via Capillary Action 754
HOW PLANTS WORK 724
6 The Cohesion-Tension Theory 755
Plant Features That Reduce Water Loss through
Transpiration 757
34 Plant Form and Function 724 35.3 Translocation of Sugars 758
Tracing Connections between Sources and Sinks 759
34.1 Plant Form: Themes with Many Variations 725
The Anatomy of Phloem 760
The Importance of Surface Area to Volume
The Pressure-Flow Hypothesis 760
Relationships 726
Phloem Loading 762
Phloem Unloading 764
CHAPTER 35 REVIEW 765

36 Plant Nutrition 767

36.1 Nutritional Requirements of Plants 768

Which Nutrients Are Essential? 768
What Happens When Key Nutrients Are in Short Supply? 770
36.2 Soil: A Dynamic Mixture of Living and Nonliving
Components 771
The Importance of Soil Conservation 772
What Factors Affect Nutrient Availability? 773
36.3 Nutrient Uptake 774
Mechanisms of Nutrient Uptake 774
Mechanisms of Ion Exclusion 776
36.4 Nitrogen Fixation 778
The Role of Symbiotic Bacteria 779
What Is the Relationship between Plants and Nitrogen-Fixing
Bacteria? 779

xiv DETAILED CONTENTS

36.5 Nutritional Adaptations of Plants 781 38.5 Embryogenesis and Vegetative Development 827
Parasitic Plants 781 Embryogenesis 828
Epiphytic Plants 781 Meristem Formation 829
Carnivorous Plants 781 Which Genes Determine Body Axes in the Plant Embryo? 829
Which Genes Determine Leaf Structure and Shape? 830
CHAPTER 36 REVIEW 783
38.6 Reproductive Development 831
37 Plant Sensory Systems, The Floral Meristem and the Flower 831
The Genetic Control of Flower Structures 832
Signals, and Responses 785 CHAPTER 38 REVIEW 833
37.1 Information Processing in Plants 786 END-OF-UNIT Can Plant Compounds Perform a Role Similar to
How Do Cells Receive and Process an External Signal? 786 CASE STUDY
Tetrodotoxin? 836
How Do Cells Respond to Cell–Cell Signals? 786
Plant and Animal Form and Function 838
37.2 Blue Light: The Phototropic Response 788
Phototropins as Blue-Light Receptors 788
Auxin as the Phototropic Hormone 789
37.3 Red and Far-Red Light: Germination, Stem UNIT
HOW ANIMALS WORK 840
Elongation, and Flowering 792 7
The Red/Far-Red “Switch” 792
Phytochrome Is a Red/Far-Red Receptor 792
Signals That Promote Flowering 793 39 Animal Form and Function 840
37.4 Gravity: The Gravitropic Response 795 39.1 Form, Function, and Adaptation 841
The Statolith Hypothesis 795
The Role of Fitness Trade-Offs 841
Auxin as the Gravitropic Signal 796
Adaptation and Acclimatization 843
37.5 How Do Plants Respond to Wind and Touch? 797 39.2 Tissues, Organs, and Organ Systems: How Does
Changes in Growth Patterns 797
Structure Correlate with Function? 843
Movement Responses 797
Structure–Function Relationships at the Molecular and Cellular
37.6 Youth, Maturity, and Aging: The Growth Levels 844
Responses 798 Tissues Are Groups of Cells That Function as a Unit 844
Auxin and Apical Dominance 798 Organs and Organ Systems 847
Cytokinins and Cell Division 799
39.3 How Does Body Size Affect Animal Physiology? 848
Gibberellins and ABA: Growth and Dormancy 799
Surface Area to Volume Relationships: Theory 848
Brassinosteroids and Body Size 802
Surface Area to Volume Relationships: Data 849
Ethylene and Senescence 803
Adaptations That Increase Surface Area 850
An Overview of Plant Growth Regulators 804
39.4 Homeostasis 851
37.7 Pathogens and Herbivores: The Defense Homeostasis: General Principles 851
Responses 806 The Role of Regulation and Feedback 851
How Do Plants Sense and Respond to Pathogens? 806
How Do Plants Sense and Respond to Herbivore Attack? 808 39.5 Thermoregulation: A Closer Look 853
Mechanisms of Heat Exchange 853
CHAPTER 37 REVIEW 810 Thermoregulatory Strategies 853
Comparing Endothermy and Ectothermy 854
38 Flowering Plant Reproduction Countercurrent Heat Exchangers 855

and Development 813 CHAPTER 39 REVIEW 856

38.1 An Introduction to Flowering Plant

Reproduction 814 40 Water and Electrolyte
Asexual Reproduction 814 Balance in Animals 858
Sexual Reproduction and the Flowering Plant Life Cycle 815
40.1 Osmoregulation and Excretion 859
38.2 Reproductive Structures 816 What Is Osmotic Stress? 859
The General Structure of the Flower 816
Osmotic Stress in Seawater, in Fresh Water, and on Land 859
How Are Female Gametophytes Produced? 818
How Do Electrolytes and Water Move across Cell
How Are Male Gametophytes Produced? 819
Membranes? 860
38.3 Pollination and Fertilization 820 How Do Different Forms of Nitrogenous Waste Impact Water
Pollination 820 Balance? 861
Fertilization 822
40.2 Water and Electrolyte Balance in Marine and
38.4 Seeds and Fruits 823 Freshwater Fishes 862
Seed Maturation 823 Osmoconformation versus Osmoregulation in Marine
Fruit Development and Seed Dispersal 824 Fishes 862
Seed Dormancy 826 How Do Sharks Excrete Salt? 862
Seed Germination 826 How Do Freshwater Fishes Osmoregulate? 864

DETAILED CONTENTS xv
40.3 Water and Electrolyte Balance in Terrestrial
Insects 864 42 Gas Exchange and
How Do Insects Minimize Water Loss from the Body
Surface? 865
Circulation 896
How Do Insects Regulate the Amount of Water and Electrolytes 42.1 The Respiratory and Circulatory Systems 897
They Excrete? 865
42.2 Air and Water as Respiratory Media 897
40.4 Water and Electrolyte Balance in Terrestrial How Do Oxygen and Carbon Dioxide Behave
Vertebrates 866 in Air? 898
Structure of the Mammalian Kidney 866 How Do Oxygen and Carbon Dioxide Behave in
Function of the Mammalian Kidney: An Overview 867 Water? 898
Filtration: The Renal Corpuscle 868
42.3 Organs of Gas Exchange 899
Reabsorption: The Proximal Tubule 868
Physical Parameters: The Law of Diffusion 899
Creating an Osmotic Gradient: The Loop of Henle 870
How Do Gills Work? 900
Regulating Water and Electrolyte Balance: The Distal Tubule
How Do Insect Tracheae Work? 901
and Collecting Duct 872
How Do Vertebrate Lungs Work? 903
Urine Formation in Nonmammalian Vertebrates 873
Homeostatic Control of Ventilation 904
CHAPTER 40 REVIEW 875
42.4 How Are Oxygen and Carbon Dioxide Transported

41
in Blood? 905
Animal Nutrition 877 Structure and Function of Hemoglobin 905
CO2 Transport and the Buffering of Blood pH 908
41.1 Nutritional Requirements 878
42.5 Circulation 908
41.2 Capturing Food: The Structure and Function of What Is an Open Circulatory System? 909
Mouthparts 880 What Is a Closed Circulatory System? 909
Mouthparts as Adaptations 880 How Does the Heart Work? 912
A Case Study: The Cichlid Throat Jaw 880 Patterns in Blood Pressure and Blood Flow 916
41.3 The Structure and Function of Animal Digestive CHAPTER 42 REVIEW 918
Tracts 881
An Introduction to the Digestive Tract 881
An Overview of Digestive Processes 882 43 Animal Nervous Systems 921
The Mouth and Esophagus: Digestion and Ingestion 883
43.1 Principles of Electrical Signaling 922
The Stomach: Digestion 884
Types of Neurons 922
The Small Intestine: Digestion and Absorption 886
The Anatomy of a Neuron 923
The Large Intestine: Absorption and Elimination 891
An Introduction to Membrane Potentials 923
41.4 Nutritional Homeostasis—Glucose as a How Is the Resting Potential Maintained? 924
Case Study 891 Using Electrodes to Measure Membrane
Insulin’s Role in Glucose Homeostasis 891 Potentials 925
Diabetes Mellitus Has Two Forms 892 What Is an Action Potential? 925
The Type 2 Diabetes Mellitus Epidemic 892
43.2 Dissecting the Action Potential 926
CHAPTER 41 REVIEW 893 Distinct Ion Currents Are Responsible for Depolarization
and Repolarization 926
How Do Voltage-Gated Channels Work? 926
How Is the Action Potential Propagated? 928
43.3 The Synapse 930
Synapse Structure and Neurotransmitter
Release 930
What Do Neurotransmitters Do? 931
Postsynaptic Potentials 932
43.4 The Vertebrate Nervous System 934
What Does the Peripheral Nervous System Do? 934
Functional Anatomy of the CNS 934
How Do Learning and Memory Work? 938
CHAPTER 43 REVIEW 941

44 Animal Sensory Systems 944

44.1 How Do Sensory Organs Convey Information

to the Brain? 945
Sensory Transduction 945
Transmitting Information to the Brain 946

xvi DETAILED CONTENTS

44.2 Mechanoreception: Sensing Pressure Changes 946
How Do Sensory Cells Respond to Sound Waves and Other
Forms of Pressure? 947
The Mammalian Ear: Hearing 948
The Mammalian Ear: Equilibrium 950
Sensing Pressure Changes in Water 950
44.3 Photoreception: Sensing Light 952
The Insect Eye 952
The Vertebrate Eye 952
44.4 Chemoreception: Sensing Chemicals 956
Taste: Detecting Molecules in the Mouth 956
Olfaction: Detecting Molecules in the Air 957
44.5 Other Sensory Systems 959
Thermoreception: Sensing Temperature 959
Electroreception: Sensing Electric Fields 960
Magnetoreception: Sensing Magnetic Fields 960
CHAPTER 44 REVIEW 961

45 Animal Movement 964

45.1 How Do Muscles Contract? 965

Early Muscle Experiments 965 46.4 How Is the Production of Hormones Regulated? 997
The Sliding-Filament Model 965 The Hypothalamus and Pituitary Gland 998
How Do Actin and Myosin Interact? 966 Control of Epinephrine by Sympathetic Nerves 1000
How Do Neurons Initiate Contraction? 968
CHAPTER 46 REVIEW 1000
45.2 Classes of Muscle Tissue 969
Smooth Muscle 969
Cardiac Muscle 969 47 Animal Reproduction and
Skeletal Muscle 970
Development 1003
45.3 Skeletal Systems 972
Hydrostatic Skeletons (Hydrostats) 973 47.1 Asexual and Sexual Reproduction 1004
Endoskeletons 974 How Does Asexual Reproduction Occur? 1004
Exoskeletons 975 Switching Reproductive Modes in Daphnia: A Case
History 1004
45.4 Locomotion 976 Mechanisms of Sexual Reproduction: Gametogenesis 1005
How Do Biologists Study Locomotion? 976
Size Matters 979 47.2 Reproductive Structures and Their Functions 1008
The Male Reproductive System 1009
CHAPTER 45 REVIEW 981 The Female Reproductive System 1009

46 Chemical Signals in 47.3 Fertilization and Egg Development 1011

External Fertilization 1011
Animals 983 Internal Fertilization 1012
The Cell Biology of Fertilization 1013
46.1 Cell-to-Cell Signaling: An Overview 984 Why Do Some Females Lay Eggs, while Others Give Birth
Major Categories of Chemical Signals 984 to Live Offspring? 1014
Hormone Signaling Pathways 985
47.4 Embryonic Development 1016
What Structures Make Up the Endocrine System? 987
Cleavage 1016
How Do Researchers Identify a Hormone? 987
Gastrulation 1017
A Breakthrough in Measuring Hormone Levels 988
Organogenesis 1018
46.2 How Do Hormones Act on Target Cells? 988
47.5 The Role of Sex Hormones in Mammalian
Hormone Concentrations Are Small, but Their Effects Are
Reproduction 1021
Large 988
Which Hormones Control Puberty? 1021
The Three Chemical Classes of Hormones 988
Which Hormones Control the Menstrual Cycle in
Steroid Hormones Bind to Intracellular Receptors 989
Humans? 1022
Polypeptide Hormones Bind to Receptors on the Plasma
Membrane 990 47.6 Pregnancy and Birth in Mammals 1025
Why Do Different Target Cells Respond in Different Ways? 992 Gestation and Development in Marsupials 1025
Major Events during Human Pregnancy 1025
46.3 What Do Hormones Do? 992
How Does the Mother Exchange Materials with the Fetus? 1026
How Do Hormones Direct Developmental Processes? 992
Birth 1027
How Do Hormones Coordinate Responses to Stressors? 995
How Are Hormones Involved in Homeostasis? 996 CHAPTER 47 REVIEW 1028

DETAILED CONTENTS xvii

48 The Immune System in Population Ecology 1056
Community Ecology 1056

Animals 1030
Ecosystem Ecology 1056
Global Ecology 1056
48.1 Innate Immunity: First Response 1031 Conservation Biology Applies All Levels of Ecological
Barriers to Entry 1031 Study 1056
The Innate Immune Response 1032 49.2 What Determines the Distribution and Abundance
48.2 Adaptive Immunity: Recognition 1035 of Organisms? 1057
An Introduction to Lymphocytes 1035 Present Abiotic Factors 1057
Lymphocytes Recognize a Diverse Array of Antigens 1036 Present Biotic Factors 1058
How Does the Immune System Distinguish Self from Past Abiotic Factors 1059
Nonself ? 1039 Past Biotic Factors 1059
Looking to the Future 1060
48.3 Adaptive Immunity: Activation 1040
The Clonal Selection Theory 1040 49.3 Climate Patterns 1061
T-Cell Activation 1040 Why Are the Tropics Warm and the Poles Cold? 1061
B-Cell Activation and Antibody Secretion 1042 Why Are the Tropics Wet? 1061
What Causes Seasonality in Weather? 1062
48.4 Adaptive Immunity: Response and Memory 1044 What Regional Effects Do Mountains and Oceans Have
How Are Extracellular Pathogens Eliminated? on Climate? 1063
The Humoral Response 1044 Do Biotic Factors Affect Climate? 1064
How Are Intracellular Pathogens Eliminated?
The Cell-Mediated Response 1044 49.4 Types of Terrestrial Biomes 1064
Why Does the Immune System Reject Foreign Tissues and What Are the Major Natural Terrestrial Biomes? 1064
Organs? 1046 Human Land Use Is Displacing Natural Biomes 1067
Responding to Future Infections: Immunological Memory 1046 How Is Global Climate Change Affecting Terrestrial Biomes? 1067
49.5 Types of Aquatic Biomes 1069
48.5 What Happens When the Immune System Doesn’t
Salinity 1069
Work Correctly? 1048
Water Depth and Sunlight Availability 1069
Allergies 1048
Water Flow 1070
Autoimmune Diseases 1048
Nutrient Availability 1071
Immunodeficiency Diseases 1049
How Are Aquatic Biomes Affected by Humans? 1072
CHAPTER 48 REVIEW 1049
CHAPTER 49 REVIEW 1074
END-OF-UNIT Do Garter Snakes Resistant to TTX Experience
CASE STUDY
Trade-Offs? 1052

UNIT
ECOLOGY 1054
50 Behavioral Ecology 1076
8 50.1 An Introduction to Behavioral Ecology 1077
Proximate and Ultimate Causation 1077
49 An Introduction to Types of Behavior: An Overview 1078
Choices Involve Trade-Offs 1079
Ecology 1054
50.2 Choosing What, How, and When to Eat 1079
49.1 Levels of Ecological Study 1055 Proximate Causes: Foraging Alleles in Drosophila
Organismal Ecology 1055 melanogaster 1079
Ultimate Causes: Optimal Foraging 1080
50.3 Choosing a Mate 1082
Proximate Causes: How is Sexual Activity Triggered in Anolis
Lizards? 1083
Ultimate Causes: Sexual Selection 1083
50.4 Choosing Where to Go 1084
Proximate Causes: How Do Animals Navigate? 1084
Ultimate Causes: Why Do Animals Migrate? 1086
50.5 Communicating with Others 1086
Proximate Causes: How Do Honeybees Communicate? 1087
Ultimate Causes: Why Do Honeybees Communicate the Way
They Do? 1088
When is Communication Honest or Deceitful? 1088
50.6 Cooperating with Others 1089
Kin Selection 1089
Quantitative Methods 50.1 Calculating the Coefficient of
Relatedness 1089
Manipulation 1091
Reciprocal Altruism 1091

xviii DETAILED CONTENTS

 Cooperation and Mutualism 1092
Individuals Do Not Act for the Good of the Species 1092 53 Ecosystems and Global
Take-Home Messages 1092
Ecology 1141
CHAPTER 50 REVIEW 1093
53.1 How Does Energy Flow through Ecosystems? 1142

51 Population Ecology 1095

1142
How Efficient Are Autotrophs at Capturing Solar Energy?
What Happens to the Biomass of Autotrophs? 1143
Energy Transfer between Trophic Levels 1145
51.1 Distribution and Abundance 1096
Global Patterns in Productivity 1147
Geographic Distribution 1096
Sampling Methods 1097 53.2 How Do Nutrients Cycle through Ecosystems? 1149
Quantitative Methods 51.1 Mark–Recapture Studies 1098 Nutrient Cycling within Ecosystems 1149
Global Biogeochemical Cycles 1151
51.2 Demography and Life History 1098
Life Tables 1099 53.3 Global Climate Change 1155
The Role of Life History 1101 What Is the Cause of Global Climate Change? 1155
Quantitative Methods 51.2 Using Life Tables to Calculate How Much Is the Climate Changing? 1157
Population Growth Rates 1101 Biological Effects of Climate Change 1159
Consequences to Net Primary Productivity 1161
51.3 Population Growth 1103
Exponential Growth 1103 CHAPTER 53 REVIEW 1163
Quantitative Methods 51.3 Using Growth Models to Predict
Population Growth 1105
Logistic Growth 1106
54 Biodiversity and Conservation
What Factors Limit Population Size? 1107 Biology 1165
51.4 Population Dynamics 1108 54.1 What Is Biodiversity? 1166
Why Do Some Populations Crash? 1108 Biodiversity Can Be Measured and Analyzed at Several Levels 1166
Why Do Some Populations Cycle? 1108 How Many Species Are Living Today? 1168
How Do Metapopulations Change through Time? 1110 Where Is Biodiversity Highest? 1168
51.5 Case Study: Human Population Growth 1112 54.2 Threats to Biodiversity and Ecosystem Function 1170
Age Structure in Human Populations 1112
Multiple Interacting Threats 1171
Analyzing Change in the Growth Rate of Human Populations 1113
How Will These Threats Affect Future Extinction Rates? 1175
Take-Home Messages 1114
Quantitative Methods 54.1 Species–Area Plots 1176
CHAPTER 51 REVIEW 1114
54.3 Why Are Biodiversity and Ecosystem Function

52 Community Ecology 1117

Important? 1177
Biological Benefits of Biodiversity and Ecosystem Function 1177
Ecosystem Services: Economic and Social Benefits of
52.1 Species Interactions 1118
Biodiversity and Ecosystems 1179
Species Interaction: Commensalism 1118
An Ethical Dimension 1181
Species Interaction: Competition 1118
Species Interaction: Consumption 1122 54.4 Preserving Biodiversity and Ecosystem Function 1181
Species Interactions: Mutualism 1125 Addressing the Ultimate Causes of Loss 1182
Take-Home Messages 1127 Conservation Strategies to Preserve Genetic Diversity, Species,
and Ecosystem Function 1182
52.2 Community Structure 1128
Take-Home Message 1185
How Many Species Occur in Communities? 1128
How Do Species Interactions Form Networks? 1128 CHAPTER 54 REVIEW 1186
Quantitative Methods 52.1 Measuring Species Diversity 1129 END-OF-UNIT What Is the Larger Ecological Context of Toxic
Why Are Some Species More Important than Others in CASE STUDY
Newts? 1188
Structuring Communities? 1130
Ecology 1190
How Predictable Are Communities? 1131
52.3 Community Dynamics 1133
Disturbance and Change in Ecological Communities 1133
Succession: The Development of Communities after APPENDIX A Answers A-1
Disturbance 1134
APPENDIX B Periodic Table of Elements B-1
52.4 Geographic Patterns in Species Richness 1136
Predicting Species Richness: The Theory of Island Glossary G-1
Biogeography 1137
Global Patterns in Species Richness 1138 Credits Cr-1

CHAPTER 52 REVIEW 1139 Index I-1

DETAILED CONTENTS xix

About the Authors
A Letter from Scott Lizabeth A. Allison is Chancellor
Professor of Biology at the College
I started working on Biological Science in 1997 with a
of William & Mary. She received
simple goal: To help change the way biology is taught. After
her PhD in Zoology from the Uni-
just shy of 20,000 hours of work on four editions of this text,
versity of Washington, specializing
that goal still gets me out of bed in the morning. But instead
in molecular and cellular biology.
of focusing my energies on textbook writing, I’ve decided to
Before coming to William & Mary,
devote myself full-time to research on student learning and she spent eight years as a faculty
developing new courses for undergraduate and graduate member at the University of Can-
students at the University of Washington. terbury in New Zealand. Liz teaches
I have passed the torch to an all-star cast of leading introductory biology for majors and
scientists and educators who have enthusiastically taught upper division molecular biology
from, and contributed to, previous editions of Biological courses. She has mentored graduate students and more than 120
Science. The new team brings their passion, talent, and undergraduate research students, many of them coauthoring pa-
creativity to the book, with expertise that spans the breadth pers with her on intracellular trafficking of the thyroid hormone
of the life sciences. Just as important, they work beautifully receptor in normal and cancer cells. The recipient of numerous
together because they think alike. They are driven by a awards, including a State Council for Higher Education in Vir-
commitment to the craft of writing and a background in ginia (SCHEV) Outstanding Faculty Award in 2009, Liz received
evidence-based teaching. one of the three inaugural Arts and Sciences Faculty Awards for
These pages offer a brief introduction to Liz Allison, Teaching Excellence in 2011, and a Plumeri Award for Faculty
Michael Black, Greg Podgorski, Kim Quillin, Jeff Carmichael, Excellence in 2012. In addition to her work on this text, she is au-
and Emily Taylor. As a group, they’ve built on the book’s thor of ­Fundamental Molecular Biology, now in its second edition,
existing strengths and infused this edition with fresh energy, with a third edition under way.
perspective, and ideas. I’m full of admiration for what they Lead Author; Chapters 1, 33, 48 and BioSkills
have accomplished, and I’m excited about the impact this [email protected]
edition will have on biology students from all over the world.
—Scott Freeman
Michael Black received his PhD
in Microbiology and Immunology
Scott Freeman received a PhD from Stanford University School of
in Zoology from the University of Medicine as a Howard Hughes Pre­
Washington and was subsequently doctoral Fellow. After graduation,
awarded an Alfred P. Sloan Post- he studied cell biology as a Bur-
doctoral Fellowship in Molecular roughs Wellcome Postdoctoral Fel-
Evolution at Princeton University. low at the MRC Laboratory of Molec-
He has done research in evolution- ular Biology in Cambridge, England.
ary biology on topics ranging from His current research focuses on the
nest parasitism to the molecular use of molecules to identify and
systematics of the blackbird family track the transmission of microbes
and is coauthor, with Jon Herron, of in the environment. Michael is a professor of Cell and Molecular
the standard-setting undergraduate Biology at California Polytechnic State University in San Luis
text Evolutionary Analysis. Scott is the recipient of a Distinguished Obispo, where he teaches introductory and advanced classes for
Teaching Award from the University of Washington and is cur- majors in cell biology and microbiology. In addition to his teach-
rently a Principal Lecturer in the UW Department of Biology, ing and research activities, Michael serves as the director of the
where he teaches introductory biology for majors, a writing- Undergraduate Biotechnology Lab, where he works alongside
intensive course for majors called The Tree of Life, and a gradu- undergraduate technicians to integrate research projects and
ate seminar in college science teaching. Scott’s current research inquiry-based activities into undergraduate classes.
focuses on how active learning affects student learning and aca- Chapters 2–12
demic performance. [email protected]

xx
 Greg Podgorski received his PhD Jeff Carmichael received his BS in
in Molecular and Cellular Biology Biology from Slippery Rock Univer-
from Penn State University and has sity in Pennsylvania and his PhD in
been a postdoctoral fellow at the Plant Biology from the University of
Max Plank Institute for Biochemis- Georgia. As an undergraduate stu-
try and Columbia University. His re- dent, he spent some time studying
search interests are in biology edu- enzyme kinetics through a fellow-
cation, developmental genetics, and ship at Oak Ridge National Labo-
computational biology. Greg’s most ratory in Tennessee. His graduate
recent work has been in mathemati- work focused on sexual reproduc-
cal modeling of how patterns of dif- tion in an intriguing group of seed
ferent cell types emerge during de- plants. He has been teaching and
velopment and how tumors recruit new blood vessels in cancer. coordinating Introductory Biology at the University of North
Greg has been teaching at Utah State University for more than 20 Dakota (UND) for more than 20 years. He also works with the
years in courses that include introductory biology for majors and campus-wide Teaching Transformation and Development
­
for nonmajors, genetics, cell biology, developmental biology, and ­Academy, where he helps other faculty members incorporate
microbiology, and he has offered courses in nonmajors biology evidence-based best teaching practices in their courses. He has
in Beijing and Hong Kong. He has won teaching awards at Utah received excellence in teaching awards at UND and as a graduate
State University and has been recognized by the National Acad- student in Georgia.
emies as a Teaching Fellow and a Teaching Mentor. Chapters 26–29 and 34–38
Chapters 13–21 [email protected]
[email protected]

Kim Quillin received her BA in Bi- Emily Taylor earned a BA in ­English

ology at Oberlin College summa at the University of California,
cum laude and her PhD in Integra- Berkeley. She then earned a PhD
tive Biology from the University of in Biological Sciences from Ari-
California, Berkeley, as a National zona State University, where she
Science Foundation Graduate Fel- conducted research in the field of
low. Kim has worked in the trenches environmental physiology as a Na-
with Scott Freeman on every edition tional Science Foundation Graduate
of Biological Science, starting with Research Fellow. She is currently
the ground-up development of the a professor of Biological Sciences
illustrations in the first edition in at the California Polytechnic State
1999 and expanding her role in each University in San Luis Obispo. Her
edition. Kim currently serves as the Curriculum Coordinator for student-centered research program focuses on the endocrine
Introductory Biology for Salisbury University (SU), a member and reproductive physiology of free-ranging reptiles, especially
of the University System of Maryland, where she is actively in- rattlesnakes. She teaches numerous undergraduate and graduate
volved in the ongoing student-centered reform of the course for courses, including introductory biology, anatomy and physiol­
biology majors. She also serves as the Curriculum Facilitator for ogy, endocrinology, and herpetology, and received the California
the biology department, focusing on programmatic review and Faculty Association’s Distinguished Educator Award in 2010 and
alignment of the SU biology curriculum to the Vision and Change Cal Poly’s Distinguished Teaching Award in 2012.
core concepts and competencies. Chapters 39–47
Chapters 22–25, 30–32, 49–54 [email protected]
[email protected]

ABOUT THE AUTHORS xxi

Preface to Instructors

S ince its inception, Biological Science’s unique emphasis

on the process of scientific discovery and guiding stu-
dents to think like biologists has placed this book at the
forefront of change in the way we teach biology. The Seventh
Edition embraces this legacy and continues to exemplify the
• Integrative End-of-Unit Case Study Introduced in Chapter 1
and then revisited throughout the book, the end-of-unit case
study “Mystery of the Newt” features a cliff-hanger tale of poi-
sonous newts and resistant garter snakes. The unfolding story
illustrates how biology concepts and the various subdisciplines
principles outlined in the Vision and Change in Undergraduate of biology (genetics, evolution, physiology, ecology, etc.) are
­Biology Education report. As in previous editions, the cutting- ­connected across multiple levels from molecules and cells to
edge biology in the Seventh Edition is pitched at exactly the right ecology and evolution. Each unit concludes with a two-page
level for introductory students and is as accurate and exciting as spread that focuses on contemporary biological elements and
ever for instructors and students alike. New findings from edu- poses questions relevant to the unit and the case study. By high-
cation research about ways to increase student engagement in lighting the practice of science, we hope to encourage students to
the learning process continue to inform and inspire the coauthor envision themselves as scientists. The end-of-unit case study is
team. The Seventh Edition introduces novel approaches that will supported by Ready-to-Go teaching modules with instructional
allow instructors to attain an appropriate blend of both content resources that include in-class clicker questions and activities
and skill development in the classroom. Innovative features new and related MasteringTM Biology questions for pre- and post-
to this edition offer students even more opportunities to actively class assignments to promote critical thinking and student
apply concepts in new situations; evaluate experimental design, engagement in the classroom.
hypotheses, and data; synthesize results; and make and inter- • Interactive Figure Walk-through Videos Twenty-five
pret models. For instructors, additional resources are provided figure walk-through videos have been added throughout the
to help align course activities and learning goals with their text to help students with unpacking complex figures and to
assessment strategies. cultivate better engagement with the text and other scientific
sources. The videos will help students to better understand
Core Values quantitative and modeling skills, as well as key concepts,
In the Seventh Edition, the coauthor team has strived to extend by breaking down the information in figures step-by-step.
the vision and maintain the core values of Biological Science—to Interactive figures are embedded in Pearson’s new and engag-
provide a book and online resources for dedicated instructors ing eText for practice at the initial point of learning and
who embrace the challenge of helping students achieve higher included in assignable Mastering Biology activities to help
levels of learning, and to provide a book that helps students each students practice working with visuals.
step of the way in learning to think like scientists, regardless of • Three New Making Models Boxes Reports like Vision and
their starting point in the process. The Seventh Edition provides Change cite the importance of developing model-based rea-
tools to help students build their cognitive mastery in both trans- soning skills. To help attain this goal, Making Models boxes
ferrable skills and biology content—to learn at the level called for were introduced in the Sixth Edition to explicitly teach stu-
by the National Academy of Sciences, the Howard Hughes Medi- dents how to use visual models in learning and doing biol-
cal Institute, the American Association of Medical Academies, ogy. In the Seventh Edition, three new Making Models boxes
and the National Science Foundation. Reports such as Biology have been added: Tips on Drawing Carbohydrates (Ch. 5); Tips
2010, Scientific Foundations for Future Physicians, and Vision and on Drawing Arrows (Ch. 40); and Tips on Drawing Immune
Change all place a premium on fundamental skills and concepts System Processes (Ch. 48). Each Making Models box has three
as well as connecting core ideas across all levels of biology. components: instruction in interpreting or creating a spe-
cific type of model, an example of that type of model, and an
What’s New in This Edition application question so that students can immediately prac-
Many of the new or expanded features in the Seventh Edition are tice their skills. Additionally, whiteboard-style videos allow
designed to seamlessly intertwine delivery of cutting-edge biol- students to watch and interact with a dynamic presentation
ogy content with development of students’ skills in data analy- of modeling. These videos are embedded in the eText, acces-
sis and quantitative model-based reasoning, in an engaging and sible in the Study Area of Mastering Biology, and integrated
relevant way. Students will receive initial instruction in biology in assignable Mastering Biology activities that test students'
content and transferrable skills, followed by opportunities for understanding of models. Lastly, there are test bank questions
lots of practice in applying knowledge and skills to new contexts. aligned with the Making Models activities.
The ultimate goal is for students to learn to construct their own • Enhanced Interactive eText New and expanded features
knowledge and think like biologists. in the Seventh Edition have been designed not only to enhance

xxii
 the print text but also to make full use of the new, embedded students synthesize information about challenging topics that
Pearson eText platform. The eText presents over 250 videos span multiple chapters or units. Related activities are avail-
and animations that have been carefully selected to support able in Mastering Biology to help students work on higher-
content in the text. Resources include whiteboard Making order problems while synthesizing key ideas.
Models videos, interactive Figure Walk-through videos, HHMI • BioSkills Instructors recognize that biology students need
Biointeractive videos, and BioFlix® Animations and Tutorials to develop foundational science skills in addition to content
that engage students, help them learn, guide them in complet- knowledge. Since the Third Edition, Biological Science has pro-
ing assignments, and bring biology to life. vided a unique, robust set of materials and activities to guide
• In-Text Learning Objectives To help students navigate students who need extra help with the skills emphasized in the
content, a Learning Objective now appears at the beginning book. The BioSkills materials reside between Chapters 1 and
of each section of a chapter. Each Learning Objective makes 2 to emphasize their importance as a resource for success in
it clear up front what fundamental content students should doing biology, and to make it easier for students to access them
expect to learn in a particular section and what they should be throughout the course. The BioSkills are presented in five
able to do with what they’ve learned. broad categories: Quantifying Biology, Using Common Lab
• Key New Content Introduced in the Sixth Edition, end-of- Tools, Visualizing Biology, Reading Biology, and Monitoring
chapter Put It All Together case studies engage students by Your Own Learning. BioSkills include practice questions, are
asking them to connect what they’ve learned in the chapter cross-referenced throughout the text, and can be assigned
to an example of contemporary, relevant research. New case online in Mastering Biology.
studies have been added that are updated for more recent • Opportunities for Practice “Blue Thread” questions, inte-
developments, relevancy, and alignment with chapter con- grated throughout the text, are designed to help students
tent: Ch 30: “Why aren’t comb jellies the most diverse animals identify what they do and do not understand. Students are
on Earth?; Ch 31: “How could you count all the bees, beetles, encouraged to think like biologists and practice engaging with
and butterflies in one forest?; and Ch. 33: “How does Zika virus content at a higher level. The idea is that if students really
cause birth defects?” Also new to the Seventh Edition is a heav- understand a piece of information or a concept, they should be
ily revised Chapter 20, which includes expanded coverage of able to do something with it. As in the Sixth Edition, all ques-
CRISPR-Cas gene editing and synthetic biology. In addition, tions in the text are assigned a Bloom’s taxonomy level to help
coverage of climate change and other human impacts, as an both students and instructors understand whether a question
important context for biological systems, is woven throughout requires higher-order or lower-order cognitive skills. A sub-
relevant chapters, especially in the ecology unit. There is also stantial proportion of questions are higher order. In addition,
an increased emphasis on systems biology and big data; for skill-based question tags help students and instructors iden-
example, in the ecology unit students explore social networks tify opportunities to practice key skills. Questions are tagged
and community networks. to indicate the following: Process of Science questions explore
• Expanded Assessment Matrix Introduced in the Sixth Edi- the application of the scientific process; Model questions ask
tion, an assessment matrix for each chapter identifies for in- students to interpret or construct visual models; Society ques-
structors how each question is related to learning objectives, tions explore the relationship between science and society;
Bloom’s level, common misconceptions, and Vision and Change Quantitative questions help students perform quantitative
core concepts and competencies. New to the Seventh Edition, analysis and use mathematical reasoning; and Think Carefully
all in-text and test bank questions are integrated and tightly questions address topics that students often have common
aligned to two levels of learning objectives: broader “big pic- misconceptions about, and the answers to these questions
ture” objectives and more granular supporting objectives. We include information that addresses the misconception.
hope this tool will assist instructors in selecting the most appro- • In-text “You Should Be Able To” questions These
priate assessment items to align with the goals of their course. questions focus on topics and concepts that professors and
students have identified as most key or difficult in each
Hallmark Features of the Text chapter.
While we are excited to introduce the new features of the Seventh • Caption questions and exercises Students are chal-
Edition, we are committed to strengthening the hallmark fea- lenged to examine the information in a figure or table
tures that make this book unique. critically—not just absorb it.
• Road Maps Each chapter opens with a concept map that • Check Your Understanding boxes One to three tasks
visually groups and organizes information to help students are presented that students should be able to complete in
anticipate key ideas as well as recognize meaningful relation- order to demonstrate a mastery of learning objectives.
ships and connections among ideas. • End-of-chapter questions Questions are organized in
• Big Picture Spreads While the Road Maps help students three levels: Test Your Knowledge, Test Your Understand-
look forward as they engage with a chapter, Big Picture con- ing, and Test Your Problem-Solving Skills—so students
cept maps, typically found at the end of a unit, help students can build from lower- to higher-order cognitive levels of
review content. Words and visuals are integrated to help ­assessment.

PREFACE TO INSTRUCTORS xxiii

• Put It All Together Case Studies Case studies added in the • Case Study Questions Put It All Together case study ques-
Sixth Edition briefly introduce contemporary biology research tions from the end of each chapter are assignable in Mastering
in action, followed by questions that ask students to apply the Biology.
chapter’s content and skills to the research topic. Instructor • Solve It Tutorials These activities allow students to act like
resources include clicker questions to give instructors the scientists in simulated investigations. Each tutorial presents
opportunity to use the case studies as discussion prompts in an interesting, real-world question that students will answer
the classroom. An enduring feature of this text is its emphasis by analyzing and interpreting data.
on experimental evidence—on teaching how we know what we
• Experimental Inquiry Tutorials The call to teach stu­
know. The case studies expand this emphasis, requiring stu-
dents about the process of science has never been louder. To
dents to evaluate real data and to see how ongoing scientific
­support such teaching, there are 10 interactive tutorials on
research is related to core biological ideas.
classic scientific experiments—ranging from Meselson–Stahl
• Focus on Real Data Students now have expanded oppor- on DNA replication to the Grants’ work on Galápagos finches
tunities to develop skills in working with real data from the and Connell’s work on competition. Students who use these
primary literature. Sources of the data presented in research tutorials should be better prepared to think critically about
boxes, graphs, end-of-chapter case studies, and the end-of- experimental design and evaluate the wider implications of
unit case studies are cited to model good practice for students the data—preparing them to do the work of real scientists in
and to provide a resource for students and instructors who the future.
wish to evaluate the original data more deeply.
• BioFlix Animations and Tutorials BioFlix are movie-­
• Illustrated Summary Tables The art program is enhanced quality, 3-D animations that focus on the most difficult core
by illustrated summary tables that deliver content in a stream- topics and are accompanied by in-depth, online tutorials that
lined way and facilitate comparison and analysis by students. provide hints and feedback to guide student learning. Eighteen
Photographic summary tables in Unit 5, for instance, illus- BioFlix animations and tutorials tackle topics such as meiosis,
trate the diversity of life. These tables make subject areas mitosis, DNA replication, photosynthesis, homeostasis, and
more accessible to visual learners and reinforce a chapter’s the carbon cycle.
key concepts.
• HHMI BioInteractive Short Films Activities Documen­
tary-quality movies from HHMI are available in Mastering
Integration of Media Biology with assignable questions to make sure students
The textbook continues to be supported by Mastering Biology, understand key ideas.
the most powerful online homework, tutorial, and assessment • Galápagos Evolution Video Activities These incredible
system available. Tutorials follow the Socratic method, coach- videos, filmed on the Galápagos Islands by Peter and Rosemary
ing students to the correct answer by offering feedback specific Grant, bring to life the dynamic evolutionary processes that
to a student’s errors or misconceptions as well as supplying hints have an impact on Darwin’s finches on Daphne Major Island.
that students can access if they get stuck. Instructors can associ- Six videos explore important concepts and data from the
ate content with publisher-provided learning outcomes or create Grants’ field research, and assignable activities keep students
their own. Highlights include the following: focused on the important take-home points.
• eText The new eText provides an engaging learning experi- • GraphIt! Activities These activities use real data to help
ence for students. We have integrated over 250 videos and ani- students explore important topics while developing skills to
mations, including Making Models videos, Interactive Figure read, interpret, and create graphs.
Walk-through videos, BioFlix animations, and data visualiza-
• End-of-Chapter Questions A broad range of end-of-­
tions. The eText is accessible on computers, tablets, and smart
chapter questions are available to assign in Mastering Biology.
phones.
• Blue Thread Questions Over 500 questions based on
• Interactive Figure Walk-through Videos Twenty-five
the Blue Thread questions in the textbook are assignable in
figures that students find challenging to interpret are now
Mastering Biology.
enhanced with brief videos that break down the art and pro-
vide explanatory narration to help students better understand • Big Picture Concept Map Tutorials An engaging concept
key concepts and to develop critical thinking skills. The vid- mapping tool is the basis for highly interactive, challenging
eos are in the eText and included in assignable activities in concept map activities based on the Big Picture figures in the
Mastering Biology that allow students to apply what they’ve textbook. Students build their own concept maps, which are
learned. automatically graded, and then answer questions to make sure
they understand key ideas and make important connections.
• Making Models Activities Forty-six whiteboard videos
bring the Making Models feature from the book to life to help • BioSkills Activities Activities based on the BioSkills con-
students develop their visual modeling skills. The videos are tent in the textbook are assignable in Mastering Biology,
in the Study Area and are also included in assignable activi- including activities to support the new BioSkills.
ties that allow students to practice modeling and apply their • Reading Quiz Questions Every chapter includes reading
understanding to new situations. quiz questions that can be assigned to ensure students read

xxiv PREFACE TO INSTRUCTORS

 the textbook and understand the basics. These quizzes are • Vision and Change Filters All content in Mastering
perfect as a pre-lecture assignment to get students into the Biology is tagged to the Vision and Change core concepts and core
content before class, allowing instructors to use class time competencies, so instructors can identify assessment items in
more effectively. every chapter that are related to Vision and Change. Instructors
can also sort items by the learning outcomes for this book.
In addition, a few tools are of particular help to busy instructors.
• Ready-to-Go Teaching Modules Based on key topics that Serving a Community of Teachers
students struggle with, these modules provide instructors All members of the coauthor team are motivated by a deep
with assignments to use before class, in class, and after class. commitment to students and to supporting the efforts of
­
New modules will support the integrative end-of-unit case ­dedicated teachers. Our passion in life is doing and teaching
study about newts to help instructors engage students with a ­biology. At various points along our diverse paths, we have been
range of activities for in class and outside of class. inspired by our own teachers when we were students, and now
• Early Alerts A powerful algorithm looks at student perfor- we are inspired by our colleagues as we strive to become even bet-
mance and behavior to help identify struggling students as ter teacher-­scholars. In the tradition of all previous editions of
early as possible in the course. Instructors are notified and can Biological Science, we have tried to infuse this textbook with the
easily e-mail students from the Early Alerts dashboard to offer spirit and practice of evidence-based teaching. We welcome your
additional help to get students back on track. comments, suggestions, and questions.

PREFACE TO INSTRUCTORS xxv

Content Highlights of the Seventh Edition
Chapter 1 Biology—The Study of Life The new title and reor- the diversity of organelle structures. References to the central
ganized sections reflect a greater emphasis on the theme of five dogma, introduced in Chapter 1, are included when describing the
characteristics of life, within a framework of three unifying functions of the nucleoid, ribosome, and nuclear envelope. A new
theories: the cell theory, the theory of evolution, and the chro- figure shows how cytosolic proteins are imported into the nucleus
mosome theory of inheritance. Coverage of “life requires energy” after being experimentally modified to include a nuclear localiza-
is expanded to include more examples of energy acquisition and tion signal. The research box is replaced with a new one that dem-
use and emphasize its importance in the diversification of life. onstrates how the secretory pathway was identified using a pulse–
Chapter 2 Water and Carbon—The Chemical Basis of Life chase assay. The description of mitochondrial structure is expanded
Macromolecules are now introduced in this chapter to provide to include the existence of dynamic networks that undergo fusion
flexibility in how instructors organize their presentation of and fission between these organelles. Discussion of lysosomes and
content in Unit 1. The description of thermodynamic systems recycling is clarified and includes a new section on autophagy.
is expanded, including a new figure that illustrates the types of Chapter 8 Energy and Enzymes—An Introduction to
systems based on the exchange of energy and matter with the Metabolism A new figure illustrates the concept of entropy.
surrounding environment. To sharpen the focus on foundational Discussion of how temperature and concentration affect chemi-
concepts that support other chapters in the unit, coverage of the cal reactions is expanded, and the research box includes new data
models for chemical evolution is condensed. and a description of how the “iodine clock” experiment was per-
Chapter 3 Protein Structure and Function The presentation of formed. The discussion of how enzymes affect chemical reactions
how electron sharing gives peptide bonds characteristics similar to is reorganized to emphasize the relationship between catalyzed
double bonds is improved. Discussion of molecular chaperones is and uncatalyzed reactions.
expanded, and a new figure illustrates their role in the protein folding Chapter 9 Cellular Respiration and Fermentation The dis-
process. New assessment questions reinforce student understanding cussion of pyruvate oxidation and the citric acid cycle is revised
of peptide bonds, how proteins fold into different tertiary structures, to improve clarity. Figures illustrating electron transport are
and the relationship between protein structure and function. updated to clearly show how electrons are transferred to ubiqui-
Chapter 4 Nucleic Acids and the RNA World The description none via both complexes I and II, and illustrations in the research
of DNA secondary structure and the accompanying figures are box are improved to clearly show the coupling of electron trans-
updated to include the concept of base stacking and more accu- port and oxidative phosphorylation. A revised figure compar-
rately represent the geometry of nitrogenous bases relative to the ing cellular respiration and fermentation emphasizes how these
sugar–phosphate backbone. The varied functional roles of RNA pathways are related in terms of regenerating electron carriers.
are now discussed with reference to the Central Dogma, which Discussion of uncoupling proteins, and how they affect electron
is introduced in Chapter 1. A new figure illustrates how tertiary transport and oxidative phosphorylation efficiency, is expanded.
structures are formed from secondary structures in RNA. Chapter 10 Photosynthesis The revised introduction to the
Chapter 5 An Introduction to Carbohydrates The introduc- light-capturing reactions and Calvin cycle places greater empha-
tion of monosaccharides is revised to emphasize how variations sis on the interdependence of these two pathways. The role of
in the ring structure of the same monosaccharide will have pro- proteins in organizing and tuning pigments in light-harvesting
found functional consequences. A new Making Models figure is complexes is expanded. Material on carbon fixation and the
included to help students identify the effect of the ring struc- reduction of sugars is reorganized to emphasize their distinct
ture on glycosidic linkages between monosaccharides. Several roles and clarify the relationship between C3 and C4 pathways.
new questions in the text address how carbohydrate structure is Chapter 11 Cell–Cell Interactions The chapter opening image
related to function of the polymers. is replaced with a vibrant fluorescent micrograph that represents
Chapter 6 Lipids, Membranes, and the First Cells A new the chapter’s key concepts regarding intercellular connections.
figure illustrates how the chemical bonds within fatty acids are The introduction of plant and animal fiber composites is reor-
responsible for the large amount of potential energy stored in ganized to streamline discussion of animal extracellular matrix
fats. The discussion of selective permeability in membranes is (ECM) and intercellular connections. The figure on animal cell
streamlined to emphasize the impact of solute polarity, size, and ECM is supplemented with a new illustration on the assembly of
charge. The section on facilitated diffusion is reorganized, and collagen proteins into fibrils. Discussion of second messengers is
differences between how channel proteins and carrier proteins expanded to emphasize their roles in diversification and amplifi-
participate in this process are clarified. cation of intracellular signals.
Chapter 7 Inside the Cell Figures of generalized animal and plant Chapter 12 The Cell Cycle Centrosome replication and par-
cells are replaced with new and vibrant illustrations to represent ticipation in forming the mitotic spindle is now included in the

xxvi
discussion of mitosis. The summary table is revised to include of alternative splicing possibilities for a single gene; and how RISC
microtubules and microtubule motor proteins as structures binds and separates double-stranded RNA. A new summary table
involved in mitosis. A new section on the discovery of proteins highlights the components of transcriptional regulation.
responsible for M-phase-promoting factor (MPF) activity is Chapter 20 The Molecular Revolution—Biotechnology, Genom­
added, including a new research box on the expression of cyclins ics, and New Frontiers This chapter is extensively revised to
in the cell cycle that illustrates how scientists distinguish between reflect the rapid advances in biotechnology. Highlights include
correlation and causation. The introduction to the G1 checkpoint extensive coverage of CRISPR-Cas genome editing, including appli-
is revised to clarify how Rb and E2F are regulated. cations in agriculture and gene drives; many new examples of
Chapter 13 Meiosis Figures are updated to more accurately the applications of biotechnology across all areas of the chapter;
show how chiasma are maintained throughout metaphase I of expanded coverage of Next-Gen sequencing, GMOs, RNA-seq, and
meiosis. Discussion of human aneuploid conditions is expanded, gene therapy; a new section on synthetic biology; and a reorganiza-
and sections of the chapter are streamlined for a sharper focus tion of the chapter to present techniques first, followed by insights
on essential content: for example, by reducing the discussion of gained from these new methodologies and finally by coverage of
details of synapsis and crossing over during meiosis I. emerging areas in biology.
Chapter 14 Mendel and the Gene Coverage of epistasis is Chapter 21 Genes, Development, and Evolution Chapter sec-
included, and material on gene linkage is expanded. Revisions tions are reorganized and updated to improve comprehension
to improve clarity and understanding were made to the research and flow, and they now move up an organizational ladder from
box; to discussions of quantitative inheritance and multiple single cell properties to organizing multiple cells to creating the
alleles; and to the figures showing the genotypes and phenotypes body plan and finally to the link between development and evo-
of the ABO blood group and quantitative inheritance. lution. New figures illustrate the stem cell concept, the creation
Chapter 15 DNA and the Gene—Synthesis and Repair The of induced pluripotent stem cells, and the genetic regulatory cas-
description of the polarity of DNA strands and how the antiparal- cades in Drosophila development.
lel nature of DNA leads to a lagging strand and challenges in DNA Chapter 22 Evolution by Natural Selection More structure
replication is now more student friendly. The content and figures and practice are provided for students in the section on evidence
are updated throughout: for example, by clarifying the sliding of evolution. The mockingbird phylogeny is replaced with a finch
clamp’s relationship to DNA polymerase, discussing the role of phylogeny to align with the finch case study. Content is updated,
DNA helicase in nucleotide excision repair, and showing an exo- for example, by including reference to CRISPR technology, new
nuclease site in DNA polymerases that is distinct from the active data on pesticide and herbicide resistance, and organisms expe-
site for polymerization. riencing climate change. The summary table on common miscon-
Chapter 16 How Genes Work New assessment questions are ceptions is enlarged to help students identify and modify their
provided to enhance student understanding of the central dogma conceptions of evolution.
of molecular biology. The description of how the genetic code was Chapter 23 Evolutionary Processes The introduction to the
cracked is expanded. Updated content includes a new figure on chro- Hardy–Weinberg principle is more student friendly, including a
mosome structural alterations and their functional consequences. sea turtle example comparing alleles, genotypes, and phenotypes;
Chapter 17 Transcription, RNA Processing, and Translation an aligned new figure of the gene pool concept; and an aligned
Content is updated, including new findings on the prevalence of new figure helping students bridge the use of Punnett squares in
coupled transcription and translation in bacteria; modifications Mendelian genetics and their use in understanding the calcula-
to the figure on splicing to better illustrate how the spliceosome tions of the Hardy–Weinberg principle. A few examples have
assembles; enhanced discussion on the need for accuracy of ami- been either updated or replaced to align with examples used in
noacyl tRNA synthetases; and new information on translation other chapters (e.g., finches and whitefish). Discussion of sexual
elongation factors, the role of GTP in elongation, and termination selection is revised to be more culturally sensitive to stereotypes
of transcription in bacteria. in human gender identity and sexual orientation.
Chapter 18 Control of Gene Expression in Bacteria A broader Chapter 24 Speciation A new figure offers a friendly introduction
view of gene expression in bacteria is provided, including use of to the topic of speciation. Mechanisms of reproductive isolation are
the CAP protein in lac operon regulation as an example of positive reformatted in an illustrated summary table, followed by applica-
control, new coverage of the trp operon, and increased coverage tion questions to give students more practice. The seaside sparrow
of global gene regulation. Figures are updated for clarity, and a case study is replaced by an expanded case study on the number of
new figure on the SOS regulon is added. elephant species and includes a research box. The apple maggot fly
Chapter 19 Control of Gene Expression in Eukaryotes case study is replaced with a case study on sympatric speciation in
Sections were reordered to improve understanding and logical killer whales. New whitefish data on fusion is added to align with
flow. Coverage of epigenetic inheritance and RNA interference is the new disruptive selection example used in Chapter 23.
expanded. Extensive updates to the figures and accompanying Chapter 25 Phylogenies and the History of Life Chapter
text better illustrate key concepts, such as how promoter-­proximal structure is clarified, explaining how phylogenetics and fossil evi-
elements and enhancers are composed of multiple regulatory dence can be used in combination or separately to reveal insights
sequences; how base-pair projections in major and minor grooves about the history of life. The phylogenetics section is streamlined
of DNA helix can be recognized by transcription factors; the number and updated. Dates and events in the history of life are updated.

CONTENT HIGHLIGHTS OF THE SEVENTH EDITION xxvii

Discussion of the Cretaceous extinction is updated to include evi- Chapter 34 Plant Form and Function Several additional key
dence of volcanism in the Deccan Traps, including a new figure terms are added to expand coverage of fundamental concepts
that supplies geological context for the continental configuration and help students navigate this updated content. Figures are
at that time. Discussion of the sixth extinction is expanded, includ- enhanced to help clarify common misconceptions about plant
ing new data showing current extinction rates for vertebrates. form and function. Discussion of experimental studies on phe-
Chapter 26 Bacteria and Archaea The list of key terms is notypic plasticity in plants is expanded to give students greater
expanded to help students navigate content. Discussion of micro- exposure to the process of science.
bial communities is enhanced, including a new figure on human Chapter 35 Water and Sugar Transport in Plants Figures and
microbiomes. New figures illustrating DNA transfer by transfor- accompanying text are modified and improved to help clarify
mation and transduction are included, along with accompanying key concepts in water and sugar transport. Previous sections on
text to clarify these processes. forces that move water from roots to shoots and features that
Chapter 27 Diversification of Eukaryotes Although protists reduce water loss are streamlined and merged.
are still the primary focus, the title is changed to reflect the evo- Chapter 36 Plant Nutrition The text is updated and stream-
lutionary emphasis of the chapter. To support this evolutionary lined for clarity throughout the chapter to emphasize key
emphasis, figures are modified to better illustrate morphologi- concepts in plant nutrition. Students will get more practice
cal diversity and key life-cycle events of protists. A new figure is at interpreting data with the help of a new research box that
added to illustrate bioluminescence and its important role in pro- addresses the question of whether legumes regulate root nodule
tist defense mechanisms. development based on nitrogen availability.
Chapter 28 Green Algae and Land Plants Several figures and Chapter 37 Plant Sensory Systems Figures and accompanying
the accompanying text are modified to emphasize morphological text are enhanced to help clarify key concepts in plant sensory
diversity and key life-cycle events of green algae and land plants. systems. To help engage students in the process of science, discus-
A new figure is added that illustrates antheridia and archegonia sion of experiments on the relationship between photoperiodism
in seedless vascular plants. The fern life-cycle figure is modified to and flowering is expanded. Key terms are added to help students
better show how fern gametophytes facilitate cross-fertilization. navigate updated content.
Chapter 29 Fungi The text and some figures are updated and Chapter 38 Plant Reproduction and Development Key figures
improved to help clarify key concepts related to morphological and corresponding text are enhanced to help clarify important con-
diversity and key life-cycle events of fungi. The sections on repro- cepts in plant reproduction and development. Students will benefit
ductive structures in fungi are streamlined to minimize repetition. from greater exposure to the process of science and practice with
Chapter 30 An Introduction to Animals Revisions include interpreting data thanks to a newly added discussion of experiments
clarifications and updates, such as the removal of Xenoturbellida on the relationship between flower color and pollinator preference.
from the deuterostomes; a clearer distinction between germ lay- Chapter 39 Animal Form and Function The study on repro-
ers and tissues derived from germ layers; and an addition of the duction and immune function in crickets has been simplified
terms “deep homology” and “biradial symmetry.” A new case to better illustrate trade-offs. Content is updated, including a
study gives students practice in distinguishing traits that were description of the genetic basis of adaptation to high elevation in
important to the origin of animal phyla versus traits that were Tibet and enhanced descriptions of the composition of connec-
important in the diversification within phyla. tive tissues, bone, and cartilage. The treatment of epithelia has
Chapter 31 Protostome Animals More references to ecology are been expanded with an updated figure that shows simple and
added for context, such as the ecological implications of the water-to- stratified epithelia side by side. The mathematical explanation of
land transition and the increasing threat of extinction to protostomes surface-area-to-volume ratio is simplified to clarify this concept.
during the Anthropocene. In addition, a new case study emphasizes A new figure shows homeostatic thermoregulation in dogs.
sampling methods and measures of diversity in tropical forests. Chapter 40 Animal Water and Electrolyte Balance A new
Chapter 32 Deuterostome Animals Xenoturbellida is removed Making Models box shows tips for modeling osmoregulatory chal-
from the deuterostome phylogeny (they are no longer consid- lenges and solutions faced by animals. A new figure compares
ered deuterostomes). The walk-through of vertebrate evolution osmoregulation in marine bony fishes and osmoconformation in
includes a new figure showing the homology of placoderm head marine cartilaginous fishes. The sections on marine and freshwa-
shields to human skull and jaw, as well as an expanded expla- ter fishes are combined to better compare osmoregulation in the
nation of the origin of lungs. The human evolution section is two groups. In addition to filtration and reabsorption in the kidney,
updated, including Homo naledi and increased evidence of inter- secretion of substances into the nephron is now briefly discussed.
breeding among species of Homo. Chapter 41 Animal Nutrition Further detail is added on the
Chapter 33 Viruses New content is included on the decimation of actions of digestive enzymes in humans, including clarification
native populations in the Americas during the sixteenth to eighteenth of the function of lingual lipase; introduction of gastric lipase;
centuries by viral epidemics. Content on different modes of viral rep- clarification of carbohydrate digestion by salivary amylase and
lication is expanded. Coverage of the reemergence of Zika virus as a pancreatic amylase; introduction of brush border enzymes for
current significant international health problem is included, along digestion of disaccharides and dipeptides; and standardization of
with a new case study on how Zika virus causes birth defects. the name “enteropeptidase.” Information on how fat-soluble and

xxviii CONTENT HIGHLIGHTS OF THE SEVENTH EDITION

water-soluble vitamins are absorbed is included. A new section in the present abiotic, present biotic, past abiotic, and past biotic
covers the human gut microbiome. environments; and a new research box focusing on the effect of
Chapter 42 Animal Gas Exchange and Circulation The sec- climate change on the distribution of açaí palms. The chapter
tion on insect tracheae is revised for clarity. A brief discussion of now includes altitude as an abiotic factor, the effect of organisms
oxygen-carrying pigments other than hemoglobin is added. New on climate, and a definition of landscape ecology. Other changes
material and a new research box are added discussing the special improve clarity and timeliness of information.
anatomy of the alligator heart. Discussion of the mechanism by Chapter 50 Behavioral Ecology Revisions increase the focus
which blood pressure drops as blood moves away from the heart on human behavior, including a new case study on consolation
is revised to clarify key concepts. behavior in humans, elephants, and prairie voles. Behaviors
Chapter 43 Animal Nervous Systems Discussion of how the rest- are categorized in a summary table representing a continuum
ing membrane potential is established is clarified. Coverage of the between innate and learned behaviors. A new figure shows how
peripheral nervous system is expanded, specifically on the auto- behaviors are learned via social networks in bees. The section on
nomic (sympathetic and parasympathetic) nervous system. Content optimal foraging includes a new research box that looks at cuttle-
on the vertebrate brain is augmented by couching the main com- fish counting behavior and an introduction to game theory. The
ponents (cerebrum, cerebellum, diencephalon, and brainstem) in section on mate choice includes a new summary table on mating
terms of the three region (forebrain, midbrain, hindbrain) schema strategies, including language that helps students to interpret
from Chapter 32. Discussion on learning and memory is revised to the complexity of human mate choice and sexual identity in this
highlight how little scientists actually know about these processes. simplistic framework. There is increased emphasis on the impor-
tance of variability in behavior in the context of climate change.
Chapter 44 Animal Sensory Systems A brief discussion of
touch is now included in the mechanosensation section. A detailed Chapter 51 Population Ecology The chapter includes a new
discussion and new figure on the inner ear’s sensation of equilib- summary table of dispersion patterns with an emphasis on proxi-
rium are added. The section about the “dark current” associated mate and ultimate causation. Increased emphasis on life-history
with stimulation of rods in the eye is revised to describe the pro- patterns includes a new figure comparing the life-history patterns
cess at an appropriate level. of two lizard populations. The section on population dynamics
includes a new case study on the crash of the reindeer population
Chapter 45 Animal Movement Language is added throughout on St. Paul island to clarify the common misconception that the
to make the chapter more interactive (e.g., put your hand on your human population will gently peak at its carrying capacity, and
biceps and flex it). The figure on muscle cross-bridge cycling is the hare/lynx research box provides a more accessible data set.
updated to show it as a circular process. The description of car- The section on human population is updated. Application to con-
diac and smooth muscle is expanded, including discussions of servation biology is now more integrated throughout the chapter.
autorhythmicity and the reason smooth muscle appears smooth.
Chapter 52 Community Ecology Content now includes a new
Chapter 46 Animal Chemical Communication Coverage of summary table comparing consumption interactions, a new section
positive feedback is added to complement the discussion of nega- referencing the hare/lynx data in Chapter 51, a new figure illustrat-
tive feedback. The section on Berthold’s discovery of testosterone is ing how mutualists can be generalists or specialists, a new figure
updated to affirm that he discovered that chemical signal(s) affect of interaction networks to show examples other than food webs,
rooster development, although the terms “hormone” and “testoster- a Making Models box on species richness (brought forward from
one” were not coined until many years later. Discussion of hormone Chapter 54), a resurrection of Paine’s keystone species data (moved
receptors is updated and streamlined by removing detail on how the from Chapter 32), new data on latitudinal diversity, and increased
estradiol receptor was discovered and focusing on key content. emphasis throughout on human impacts, including climate change.
Chapter 47 Animal Reproduction and Development The Chapter 53 Ecosystems and Global Ecology Updated content
discussion of asexual reproduction in animals is updated to show includes a new opening image of global CO2 data, a new figure
that it does not always result in clones. For clarity, the process showing solar-induced fluorescence measured via satellite, a new
of cell migration during gastrulation is now described as “move- figure helping students to analyze the productivity pyramid while
ment” rather than “migration.” The concept of the evolution of also understanding the impacts of their food choices, increased
viviparity in reptiles and mammals is expanded. emphasis that the greenhouse effect and ozone layer are different
Chapter 48 The Immune System in Animals Content is phenomena, new reference to the Paris climate agreement, and
expanded to help students visualize how innate and adaptive new reference to ocean deoxygenation.
immunity work together, including a new Making Models box, Chapter 54 Biodiversity and Conservation Content is
“Tips on Drawing Immune Cells.” Updates to coverage of antigen updated throughout, including a new summary table compar-
presentation and the activation of B and T cells take a more com- ing species, phylogenetic and functional diversity; a revised
parative approach. map showing where global vertebrate biodiversity is highest; an
Chapter 49 An Introduction to Ecology The introductory sec- updated map of biodiversity hotspots; an updated data set of the
tion on levels of ecology increases the focus on variables that ecol- distribution of threats to terrestrial, freshwater, and saltwater
ogists measure. Section 49.2 is restructured as a case study on ecosystems; the inclusion of disease in the topic of invasive spe-
açaí palms, including a new niche model comparing açaí palms cies; and a new map illustrating the principle of “Nature Needs
and coconut palms; a new summary table comparing variables Half” as a conservation goal for a sustainable future.

CONTENT HIGHLIGHTS OF THE SEVENTH EDITION xxix

Another random document with
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denying herself food, clothes, and needed rest, to take care of the
one who had befriended her; but with all her care and kindness the
old woman faded day by day, and early in September died, invoking
with her last breath blessings on Lally’s name.
The few sticks of furniture were sold to give the old woman a decent
burial. Lally was out of money—out of everything. The
superintendent of the boys’ school refused to allow her to continue
the duties she had performed in the old woman’s name, alleging that
she was too young. And as a last blow, she was turned out of her
lodgings because of her inability to pay the rent.
At this crisis of her history, when as it seemed only death presented
an open door to her, she resolved to go down to Wyndham and look
once more on her husband’s face.
To think, with our desperate Lally, was to act. She set out to walk to
Wyndham, working in the hop-fields for sustenance as she went.
Thus she did three full days of work before she arrived near her
destination, and she had crept into the way-side thicket to rest before
continuing her journey to Wyndham, when she chanced to overhear
the conversation between Neva Wynde and Rufus Black.
Her despair, as she listened to the words of her young husband in
declaring his love for Neva, may be imagined. She did not dream
how bitterly he had mourned for his lost young wife; she did not
dream that she was dearer to him still than Neva could ever be. How
could she tell, when listening to his passionate vows of love to Miss
Wynde, that the young wife who had slept in his bosom was in his
thoughts by day and by night, and was regarded by him as a holy,
precious memory?
“It’s all over!” she sobbed, pressing her face down upon the dewy
turf. “I am forgotten—but why should I not be? I never was his wife.
He said so himself in his letter to me that I carry still next my heart.
Not his wife—but she will be! How beautiful she is! How lovely her
face was, how clear her voice. She would pity me if she knew, but
she is an heiress, I dare say, while I am only the poor outcast Rufus
has made me! Oh, Rufus, Rufus!”
She wailed aloud, but she had learned to bear her griefs in silence,
and presently she struggled to her feet and walked in the direction in
which the heiress and her lover had gone—the same way by which
Lally had recently come.
There was no need for her to go to Wyndham now. Her presence
there, or her appearance to Rufus, might embarrass his relations to
his newer love, and possibly interfere with his marriage. He thought
her dead, and had not even come forward to claim the body he
supposed to be hers. Ah, yes, she had never been his wife, and she
was forgotten. She would never cross his path again.
She staggered wearily along the road, in and out of the beaten foot-
path, with the twilight deepening around her, and with a deeper
twilight settling down upon her heart and brain. She passed the
Hawkhurst park, the picturesque stone lodge guarding the great
bronze gates, and here she paused.
The lodge was closed, and a faint light streamed out through the
dotted white curtains. Lally crept close to the great gates formed of
bronze spears tipped with gilt, like the gates of the Tuileries gardens
at Paris, and pressing her face against the cool rods, looked up the
avenue.
At the distance of half a mile or more, the great gray stone mansion
sat throned upon a broad ridge of land, and lights flared from the
wide uncurtained windows far upon the terrace, and the glass dome
of flowers was all alight, and the stately old house looked to the
homeless wanderer down by the gates like Paradise.
Her eager eyes searched the terrace, and then, inch by inch, the
great tree-arched avenue.
Midway up the avenue, walking slowly, as lovers walk, she saw her
young husband and Neva Wynde. With great jealous eyes she
watched their progress through the shadows, and, when they
paused in the stream of light upon the terrace, and Rufus Black bent
low toward the heiress, a great flame leaped into poor Lally’s sombre
eyes, and she caught her breath sharply.
The heiress and her suitor stood for some moments upon the
terrace, unconscious of the eyes upon them. Rufus declined to go
into the house that evening, alleging his agitation as an excuse.
Neva took her small parcel which he had carried, and he seized her
hand, uttering passionate words of love, and begging her to look
favorably upon his suit. Then not waiting for an answer, he pressed
her hand to his lips, and dashed down the avenue toward the gates,
while Neva entered the house.
And all this the jealous, disowned wife saw, with her face growing
death-like, and the flame burning yet more brightly in her sombre
eyes.
“She has accepted him,” she muttered. “She will not take the week to
consider his suit. They are betrothed. I was sure she lived here.
Perhaps she owns the place, and he will be its master. They will both
be rich and happy and beloved, while I—Ah, how swiftly he comes!
He walked like that the night I accepted him. But I am not his wife; I
never was, even when I thought myself so. He must not see me. No
shadow from the past must darken his happy life—his and hers. It is
all over—all over—and I shall never see his face again!”
With one last, long lingering look, and a sob that came from her very
soul, she turned and sped down the road like a mad creature—away
from Wyndham, and Rufus, and all her hopes—going, ah, where?
And Rufus, with his new love-dream glowing in his soul, came out of
the Hawkhurst grounds, and hurried toward his inn, never dreaming
how near he had been to his lost wife, nor how surely he had lost
her.
 CHAPTER XVIII.
ONE OF NEVA’S LOVERS DISPOSED OF.

Upon his return to the Wyndham inn, Rufus Black found his father
awaiting him in their private parlor. The elder Black arched his brows
inquiringly as his son came in, and Rufus bowed to him gayly, as he
said:
“Well, father, you ought to be pleased with me now. I have offered
myself to Miss Wynde.”
Craven Black started.
“She has accepted you?” he demanded.
“Not yet. She wants to think the matter over, and I have consented to
let the thing rest where it is for a week. I take it as a good sign that
she did not refuse me at once. Her hesitation implies a regard for me
—”
“Or a sense of duty toward some one else,” muttered Craven Black.
“Curse that letter. If I had seen the girl, I would never have written it.”
“What is it you say, father? I did not catch your words.”
“They were not meant for your ears. So, Miss Wynde demands a
week in which to consider your offer? It would be proper for you to
refrain from going to Hawkhurst to-morrow. I’ll explain to her that you
remained away from motives of delicacy.”
“Which I shall not do,” said Rufus doggedly. “I shall go to Hawkhurst
to-morrow evening. I will not leave the field clear to Lord Towyn. He’s
an earl, rich, handsome, and intellectual, the very man to capture a
girl’s heart, and if I know myself, I am not going to give him a clear
field. Why, he loves her better than I do even, and I can only come
out ahead of him by dint of sheer persistency. It’s a mystery to me
how she refrained from saying No to me, when she can have Lord
Towyn if she chooses. There is something behind her hesitation—
some hidden cause—”
“Which you will do well to let alone,” interposed his father. “‘Take the
goods the gods provide’ without questioning.”
Rufus was not satisfied, but concluded to act upon this advice.
The next morning Craven Black attired himself with unusual care,
and mounted his piebald horse, a new purchase, and set out alone,
at a slow canter, for Hawkhurst. He knew that the heiress usually
took a morning ride, attended only by her groom, and he knew in
what direction these rides usually lay. It was impossible for him to
demand a private interview with her at her home without exciting the
suspicions and jealousy of Lady Wynde, and he was determined to
see the heiress alone, and discover in what estimation she held him.
He was also determined not to accept quietly the four thousand a
year of the baronet’s widow until he knew, beyond all peradventure,
that he could not obtain the seventy thousand per annum of the
baronet’s daughter.
He rode up to Hawkhurst lodge, slackening his speed, but not
pausing. As it happened, a little boy, a son of the lodge keeper, was
playing in the road, and Craven Black tossed him a sixpence, and
demanded if Miss Wynde were out riding, and which way she had
gone.
“Dingle Farm way,” said the urchin, scrambling in the dust for the
shining coin. “She’s been gone a long time.”
“Who is with her?” asked Craven Black.
“Jim, the groom—that be all.”
Black put spurs to his horse and dashed on. He knew where the
Dingle Farm was, it having been pointed out to him by Lady Wynde,
as a portion of the Hawkhurst property. The ride was a favorite one
with Neva, being unusually diversified. The road led through the
Dingle wood, across a common, and skirted a chalk-pit of unusual
size and depth.
Craven Black turned off from the main road into a narrower one that
led across the country, and pursued this course until he entered into
the cool shadows of the Dingle wood. Still riding briskly, he came out
a little later upon the Dingle common, a square mile of unfenced
heath, covered with furze bushes. At the further edge of the common
was the chalk-pit, now disused. The road ran dangerously near to
the precipitous side of the pit, and there was no railing or fence to
serve as a safeguard. Beyond the chalk-pit lay the Dingle Farm, a
cozy, red brick farm-house, embowered with trees.
The morning was clear and bright, and the sun was shining. As
Craven Black emerged from the shadow of the wood he swept a
keen glance over the level common, and beheld a mile or more
away, beyond the chalk-pit, but approaching it, the figure of Miss
Wynde.
She was superbly mounted upon a thoroughbred horse, and was
followed at a little distance by her groom.
Even at that distance, Craven Black noticed how well Neva sat her
horse; how erectly she carried her lithe, light figure; how proudly the
little head was poised upon her shoulders. She was coming on
toward him at a sweeping gait, her long green robe fluttering in the
swift breeze she made.
“She will be a wife to be proud of,” thought Craven Black, with a
strange stirring at his heart. “How fearless she is. One would think
she would pass the chalk-pit at a walk, but it is evident she does not
intend to.”
He dashed on to meet her. Neva saw him coming, recognized him,
and the close grasp upon her bridle rein relaxed, and the fierce
gallop subsided into a quiet canter.
She was past the chalk-pit when he came up to her, and she bowed
to him coldly, but courteously.
“Good-morning, Miss Wynde,” said Mr. Black. “You were having a
mad ride here. I fairly shuddered when I saw you coming. A single
sheer on the part of your horse would have sent you over the
precipice.”
“Oh, Badjour and I understand each other,” said Neva lightly, patting
the horse’s proudly arched neck. “I never ride a horse, Mr. Black, if I
have not confidence in my ability to control him.”
“But the road is so narrow and dangerous at this point,” said Craven
Black, wheeling and riding slowly at her side.
“You are right, Mr. Black. The road must be fenced in. I will speak to
Lord Towyn about it.”
“And why not to Sir John Freise or Mr. Atkins, who are equally your
guardians?” asked Craven Black, with an attempt at playfulness.
“Because I presume I shall see Lord Towyn first,” replied Neva,
gravely. “What do you say to a race, Mr. Black? I see that you are
returning with me.”
Craven Black looked over his shoulder. The discreet groom had
fallen behind out of earshot. Now was the time to make his
declaration of love. Such an opportunity might not again occur.
“The truth is, Miss Wynde,” he exclaimed, “I came out to meet you. I
want to have a quiet talk with you, if you will hear me.”
Neva bowed her head gravely, and her reins fell loosely in her
gauntleted hand. They were out upon the wide common now, the
Dingle farm behind them. The Dingle wood ahead.
“You may guess the nature of the communication I have to make to
you, Miss Wynde,” said her elderly lover, with an appearance of
agitation, a portion of which was genuine. “That which I have to say
would be more fittingly said in some other position perhaps. I should
prefer to say it on my knees to you, as the knights made love in
olden times.”
“Oh!” said Neva. “Hadn’t we better move on faster, Mr. Black?”
“Coquettish like all of your sex!” said Craven Black, drawing nearer
to her. “You understand my meaning, Neva? You know that I love
you—I who never loved before—”
“Surely,” cried Neva, with an arch sparkle in her red-brown eyes,
“you did not perjure yourself when you married the mother of your
son?”
Craven Black bit his lips fiercely, but said smilingly:
“That marriage was one of convenience. No love entered into it, on
my side, at least. I never loved till I met you, fair Neva. You have
younger suitors, but not one among them all who will be to you what
I would be—your slave, your minister, your subject.”
“And I should want my husband to be my king,” murmured Neva
softly. “And I would be his queen.”
“That arrangement would suit me perfectly,” declared Craven Black,
feeling a little awkward at his love-making, not altogether sure Neva
was not secretly laughing at him, yet eagerly catching at the
assistance her words afforded him. “I would be your king, Miss Neva
—”
He paused in anger, as the girl’s light laugh made music in his ears
that he by no means appreciated. His anger deepened, as Neva
looked at him with a bright sauciness, a piquant witchery of eyes and
mouth.
“You are very kind,” the girl laughed, “but I do not think—pardon me,
Mr. Black—that you are of the stuff of which kings of the kind I meant
are made!”
Craven Black’s fair face flushed. He tugged at his light beard with
nervous fingers. An angry light glowered in his light eyes.
“I may not know the full meaning of your words, Miss Neva,” he said,
forcing himself to speak calmly. “A romantic young girl like you is
sure to have many fancies which time will prune. A young girl’s fancy
is like the overflowing of some graceful rose-tree. When time shall
have picked off a bud here, a leaf there, or a half-blown rose
elsewhere, the remainder of the blossoming will be more perfect. I
am no knight of romance, but I am not aware that there is anything
ridiculous in my face or figure. Ladies of the world have smiled
graciously upon me, and more than one peeress would have taken
my name had I but asked her. My heart is fresh and young, full of
romantic visions like yours. My love is honest, and a king could offer
no better. Miss Wynde, I ask you to be my wife!”
Neva’s face was grave now, but the sparkle was still in her eyes, as
she said:
“I am sure I beg your pardon, Mr. Black, but I thought you were a
suitor of Mrs. Artress. I never had an idea that your visits were
directed to me. I am deeply grateful for the honor you have done me
—I suppose that is the proper remark to make under the
circumstances; the ladies in novels always say it—but I must decline
it.”
“And why, if I may be allowed to ask?” demanded Craven Black, his
face deepening in hue nearly to purple. “Why this insulting refusal of
an honest offer of marriage, Miss Wynde?”
Neva regarded her angry suitor with cool gravity.
“I beg your pardon if the manner of my refusal seemed insulting,”
she said gently, “but the idea seems so singular—so preposterous!
At the risk of offending you again, Mr. Black, I must suggest that a
union with Mrs. Artress would be more suitable. I am only a girl, and
young still, as you know, and it is proper that youth should mate with
youth.”
“You prefer my son then?”
“To you? I do.”
“And you will marry him?”
The lovely face shadowed, but Neva answered quietly:
“Mr. Rufus has asked me that question, sir, and I prefer to have him
receive his answer from my lips. Whatever my feelings toward him, I
have no indecision in regard to you.”
“And you actually and decidedly refuse me?”
“Actually and decidedly, Mr. Black!”
“Is there no hope that you may change your mind Miss Wynde? Will
no devotion upon my part affect your resolution?”
“None whatever. I cannot even give your proposal serious
consideration, Mr. Black. I am willing to regard you as a friend. As a
lover, pardon me, you would be intolerable to me.”
Neva spoke with an honest frankness that increased Craven Black’s
anger. He saw that he had no chance of winning her love or her
fortune, and it behooved him not to lose the lesser fortune and lesser
charms of her step-mother. He tried to take his failure
philosophically, but in refusing his love, Neva had made him her
bitter and unscrupulous enemy.
“I accept my defeat, Miss Wynde,” he said bitterly, “and resign all my
pretensions to your hand. Pardon my folly, and forget it. I hope my
son will meet with better success in his suit. And may I ask as a
favor that you will keep my proposal secret, not even telling it to your
step-mother?”
“I am not in the habit of boasting of such things, even to Lady
Wynde,” said Neva, coldly. “Your proposal, Mr. Black, is already
forgotten.”
They were in Dingle wood now, and the heiress struck her horse
sharply and dashed away at a canter. Craven Black kept pace with
her, and at a discreet distance behind followed the liveried groom.
Neither spoke again until they were out of the wood, and had
traversed the cross-road and gained the highway. When the gray
towers of Hawkhurst loomed up in full view, their speed slackened,
and Craven Black said hastily:
“One word, Miss Wynde. I have your solemn promise, have I not,
that you will never betray the fact that I have proposed marriage to
you?”
Neva bowed haughtily.
“Since you have not confidence in my delicacy,” she said, “I will give
the promise.”
Craven Black’s face flushed with something of triumph. He was still
smarting with his anger and disappointment, still secretly foaming
with a bitter rage, but he desired to show Neva that he was not at all
crushed or humiliated.
“Thank you,” he said. “I shall rely upon that promise. The truth is,
Miss Neva, a betrayal of my secret would cause me serious trouble.
Ladies never pardon even a slight and temporary disaffection like
mine. I am engaged to be married, and my promised bride is the
most exacting of women. She would rage if she knew that I had
looked with love upon one so many years her junior.”
“Indeed! You will marry Artress then?”
“Artress?” ejaculated Black, in well-counterfeited amazement. “What,
marry the companion when I can have the mistress? No, indeed,
Miss Neva. I am engaged to Lady Wynde!”
“To Lady Wynde—to my father’s widow?”
Black bowed assent.
Neva was astounded. She had been too busy with her friends since
her return to Hawkhurst to detect the real object of Craven Black’s
visits, and both Lady Wynde and Black had conspired to hoodwink
her. She had never contemplated the possibility of Lady Wynde
marrying for the third time. The idea almost seemed sacrilegious.
Her father had seemed to her so grand and noble, so above other
men, that she had not deemed it possible for a woman who had
once been honored with his love to marry another.
“It is like Marie Louise, who married her chamberlain after having
been the wife of Napoleon,” she thought. “It is incredible. I refuse to
believe it!”
Her incredulity betrayed itself in her face.
“You don’t believe it?” said Black, with a mocking smile. “It is true, I
assure you. Lady Wynde and I became engaged before your return
from school. We are to be married next month. Her trousseau is
secretly preparing in London.”
His manner convinced Neva that he spoke the truth.
“And so,” she said, her lip curling, “when your wedding-day is so
near, and the woman you have won is making ready for your
marriage, you amuse yourself in talking love to me! And that is your
idea of honor, Mr. Black? You are well named. Craven by name, and
Craven by nature!”
She inclined her head haughtily and dashed on. Black, choking with
rage, hurried in close pursuit. The lodge gates swung open at their
approach, and they galloped up the avenue. Lady Wynde came out
upon the terrace to meet them. Neva dismounted at the carriage
porch, the terrace being only upon one side of the mansion, and with
a haughty little bow to Lady Wynde passed into the house.
Black dismounted and gave his horse in charge of the stable lad who
had taken in hand the horse of Neva, and then walked toward the
open drawing-room window with his betrothed wife.
“What is the matter between you and Neva, Craven?” asked Lady
Wynde jealously. “You look as black as a thundercloud, and she
looked like an insulted queen. What have you been saying to her?”
“I thought it time to divulge our secret to her, my darling,” said Black
hypocritically. “Our wedding-day is so near that I deemed it best to
inform her. I met her out riding, and seized upon the occasion to
declare the truth.”
“And what did she say?”
“She fairly withered me with her scorn; recommended me to marry
Matilda Artress; and seemed to regard my marriage with her father’s
widow as a species of sacrilege. I hate her!” he hissed between his
clenched teeth.
Lady Wynde smiled, well-pleased.
“And so do I,” she acknowledged frankly. “But it is for our interest to
counterfeit friendship for her. Be patient, Craven. Some day you and
I may bring down her haughty pride to the dust.”
“Suppose she refuses Rufus?”
“You and I will soon be married, Craven, and in our union is strength.
Tell Rufus to write to Neva, delaying her answer to his suit for a
month. By that time we shall be married. If she refuses then to
accept your son as her husband, we can contrive some way to
compel her obedience. I am her step-mother and guardian, and have
authority which I shall use if I am pushed to the wall. I promise you,
Craven, that we shall secure our ten thousand a year out of Neva’s
fortune, and that we shall compel the girl to marry your son. Leave it
all to me. Only wait and see!”
 CHAPTER XIX.
NEVA’S CHOICE FORESHADOWED.

In accordance with the advice of his scheming father, Rufus Black

wrote a letter to Neva Wynde entreating her to take a month or six
weeks, instead of the single week for which she had stipulated, for
the consideration of his suit. And Neva, struggling between
conflicting feelings, whose nature the reader already knows, and
glad to be relieved of the necessity for an immediate decision,
gratefully accepted the offered reprieve.
The engagement of Craven Black and Lady Wynde, now that it had
been declared to Neva, was no longer kept a secret from the world.
Mr. Black, in a moment of good-natured condescension, informed his
host at the Wyndham inn, and the amazed landlord bruited the story
through the village. The engagement was publicly announced in the
court papers, Craven Black himself writing the paragraph and
procuring its insertion, and this announcement was copied into the
Kentish journals.
As may be imagined, the news of Lady Wynde’s intended marriage
produced quite a sensation in the neighborhood of Hawkhurst. Sir
Harold Wynde’s former friends were scandalized that he should have
been so soon forgotten by the wife he had idolized, and that a man
so palpably inferior to the baronet in character and attributes should
have been chosen to take his place. Others, the three guardians of
Neva’s property among the number, were ill-pleased that Craven
Black should take his place during Neva’s minority as nominal
master of Hawkhurst, and accordingly one morning, a fortnight after
the publication of the engagement, Sir John Freise, Mr. Atkins, and
Lord Towyn, rode over to Hawkhurst, and demanded an interview
with Lady Wynde and Neva.
Miss Wynde appeared first in the drawing-room, simply dressed in
white, and fresh from a ramble in the park. She looked a little worn
and troubled, as if her nights were spent more in anxious thoughts
than in slumbers, but the radiance of her wonderful red-brown eyes
was undimmed, and her face had lost nothing of the piquant witchery
which was its chiefest charm.
Before time had been granted Neva to more than exchange
greetings with her guardians, Lady Wynde entered the room with an
indolent languor of motion, and welcomed her visitors with effusion.
“This is an unexpected pleasure, gentlemen,” said her ladyship, her
black eyes glancing from one to another. “You have come to
congratulate me upon the change in my prospects, I dare say. I have
been overwhelmed with calls during the past week, and begin to find
my connection with an old county family decidedly onerous,” and she
laughed softly. “All of Sir Harold’s friends have been to see me, and
really I believe that some of them have felt it their duty to condole
with Neva upon the misfortune of so soon possessing a step-papa.”
The three gentlemen had called for the purpose of discussing with
Lady Wynde and Neva the expected change in the prospects of her
ladyship, but the quiet audacity of the handsome widow’s speech
and manner half-confounded them.
Sir John Freise, being the eldest of the party, took upon himself the
office of spokesman.
“I was an old friend of Sir Harold, Lady Wynde,” he said, a little stiffly.
“I was a man when Sir Harold was a boy, but I knew him well, and I
loved him. I know how deeply he was attached to you, and it is for
his sake that I have now intruded upon you. You are still young, and
with your attractions and your fortune you are peculiarly liable to be
beset by fortune-hunters. As your late husband’s most intimate
friend, I desire to ask you if you have well considered this step you
are about to take?”
Lady Wynde bowed a cold assent.
“Your knowledge of the character of Mr. Black can be but slight,”
persisted Sir John Freise, leaning his chin upon the gold knob of his
walking-stick, and regarding the handsome widow with troubled
eyes. “He has been at Wyndham but a few months. I grant that he is
of attractive exterior, Lady Wynde, but what do you know of his
character? I have not come here to make any charges against Mr.
Black but those I am prepared to substantiate. These gentlemen who
have accompanied me will bear me out in the statement that I have
no personal prejudices in the matter, and that I am actuated only by
a desire for your ladyship’s happiness and that of Miss Wynde. I
have written to London since hearing the report of your engagement,
and yesterday received a reply of so much moment that I summoned
Lord Towyn from his marine villa and Mr. Atkins from Canterbury to
accompany me into your presence, and assist me to impart to you
the unpleasant news. Lady Wynde, this Craven Black, your accepted
lover, is a scoundrel, a gamester, a man unworthy your consideration
for a moment.”
“Indeed!” said Lady Wynde, with a slight sneer. “Mr. Black, to my
knowledge, goes in the first society. He visited at the Duke of
Cheltenham’s last year, and the duke is a perfect Puritan, as every
one knows.”
“The Duke of Cheltenham is a distant connection of Mr. Black, and
invited him to his house with the hope of winning him into better
courses,” said Sir John gravely. “But it is not Mr. Black’s high
connections, but the man himself, with whom your destiny is to be
linked, Lady Wynde. I implore you to consider your decision. Better
to remain for ever the honored widow of Sir Harold Wynde than to
become the wife of Mr. Craven Black.”
“I do not think so,” said her ladyship, her sneer deepening. “I believe
I am competent to choose for myself, Sir John, and it is my
happiness, you will be pleased to remember, which is at stake. I
resent your interference, as uncalled for and intrusive. I shall marry
Mr. Craven Black in two weeks from to-day, and if you do not
approve the marriage I presume you will be able to testify your
disapproval by remaining away from the wedding.”
Sir John looked deeply pained; Mr. Atkins looked disgusted. Lord
Towyn’s warm blue eyes were directed toward Neva rather than
toward Lady Wynde, but he lost nothing of the conversation.
“I have performed only my duty in warning you, Lady Wynde,” said
Sir John, after a pause. “You are bent upon this marriage with a man
who was a stranger to you three months since, and so soon after the
tragic death of Sir Harold Wynde in India?”
“I have waited a year and three months before marrying again,”
declared Lady Wynde, impatiently. “Why should I wait longer? Surely
a year of mourning is all that custom requires. And as to not knowing
Mr. Black, permit me to say that I know him well. I knew him before I
ever met Sir Harold. Frequenting the same circles in town, and
meeting more than once at the same houses in the country, it is
impossible that I should not have known him. And here I beg you will
drop the subject. I am in no mood to hear your aspersions of an
honorable man, and your jealousy for the memory of Sir Harold
Wynde need not blind you to the fact that virtue and honor did not
die with him.”
Sir John looked shocked and amazed. Neva’s face paled, and a
sudden indignation flamed in her eyes, but she remained silent.
“I think, with all deference to your opinion, Sir John,” said Mr. Atkins,
“that, as Lady Wynde suggests, we would better drop the subject of
Mr. Black. It is difficult to convey unpleasant information in a case
like this without giving offence. We have done our duty, and that
must content us. Let us now come to the actual business in hand.
Allow me to ask you, Lady Wynde, if you intend to continue your
residence at Hawkhurst after becoming Mrs. Craven Black?”
A flash of defiance shot from her ladyship’s black eyes.
“Certainly, I intend to reside here with my husband during the
minority of my step-daughter,” she declared boldly. “I am Neva’s
guardian, and my residence as such was assigned at Hawkhurst.”
“Sir Harold never contemplated a state of affairs such as you
propose Madam,” said Mr. Atkins doggedly. “To make this Mr.
Craven Black nominal master of the home of the Wyndes is
something utterly unlooked for.”
“Where I am mistress, my husband will be master!” asserted Lady
Wynde, with temper.
“It should be so,” declared Mr. Atkins, “but you see how inappropriate
it would be to make Mr. Black master of Hawkhurst. Good taste—
pardon my plainness—would dictate your ladyship’s retirement from
Hawkhurst upon the occasion of your third marriage, and we have
come to propose that Hawkhurst be closed, Miss Neva transferred to
the guardianship of Sir John Freise and Lady Freise, and that you
and your new husband take up your abode at Wynde Heights, your
dower house, or at any other place you may prefer.”
Lady Wynde frowned her anger and defiance.
“I shall remain at Hawkhurst,” she exclaimed haughtily. “If you desire
to remove me, you must do so by process of law. If you think her
father’s wife an unfit personal guardian for Miss Wynde, you can
have Sir Harold’s will set aside, or take legal proceedings to obtain
for her another guardian. I shall not relinquish my post, or the charge
my dead husband reposed in me, until I am compelled to do so.”
The young Lord Towyn’s face flushed, and he addressed Neva, in
his clear ringing voice:
“Miss Wynde, this matter concerns you above all others, and it is for
you to have a voice in it. The proposed marriage of Lady Wynde
completely vitiates your present relations to her. In becoming Mrs.
Craven Black, I consider that Lady Wynde throws off all allegiance to
Sir Harold Wynde, and ceases to be your step-mother. It is for you to
decide if you will choose a new personal guardian in her stead.”
All eyes turned upon the fair young girl. The young earl awaited her
reply with a breathless anxiety. Sir John Freise and Mr. Atkins fixed
their eager gaze upon her, and Lady Wynde regarded her sharply
and with some uneasiness.
“Before Neva comes to a decision,” said her ladyship hastily, “I have
a word to say to her. Have I not treated you with all kindness and
tenderness, Neva, since you came under this roof? Have I been
guilty of one act of neglect, of step-motherly cruelty, or want of
consideration? Have not your wishes been considered in all things?”
Neva could not answer these questions in the negative.
“There is no stipulation in Sir Harold’s will that I should not again
marry,” continued Lady Wynde. “Sir Harold, without mention of the
contingency of another marriage on my part, constituted me his
daughter’s personal guardian, with the request that I make
Hawkhurst my home until Neva marries or attains her majority. Not
one word is said about or against my marriage, you will observe; and
certainly Sir Harold Wynde was too sensible to expect me to remain
a widow long—at my age too. My marriage, therefore, does not
interfere with my relations toward Neva as her step-mother and
personal guardian. Any court of law will confirm this decision. If you
choose, Neva, to apply for a change of guardians, and to make a
scandal, and to make your name common on every lip, I can only
regret your ill-taste, and that you have yielded to such ill-guidance.”
Mr. Atkins felt a sentiment of admiration mingle with his dislike for
Lady Wynde.
“She ought to have been a lawyer,” he thought. “She’s a mighty
sharp woman, and we are sure to get the worst of it in a battle with
her. Pity we made the attack, if it is only to put her on her guard.”
Neva was still considering the matter intently. She had a thorough
contempt for Craven Black, and disliked the prospect of being under
the same roof with him, but she dreaded still more the publicity that
would be given to her application for change of guardians. She
remembered her father’s many injunctions to cling to Lady Wynde
until her own marriage, or the attainment of her majority. Lady
Wynde had not been unkind to her, nor illy fulfilled her duties as
chaperon. Neva had actually nothing of which to complain, save
Lady Wynde’s proposed marriage. She was a conscientious girl, and
she could not decide to throw off the yoke her father had placed
upon her shoulders, simply because Lady Wynde had chosen to
enter into new relations which were not likely to affect the old. She
felt that she was placed in a cruel position, but her duty, she thought,
was plain to her.
“Well, what is your decision, my child?” asked Sir John Freise
paternally.
“You are very kind to me, Sir John, and you also, Lord Towyn and Mr.
Atkins,” said the young girl tremulously, “and I cannot properly
express my gratitude to you for your concern for me. I appreciate all
you have said, all that you mean. I own that Lady Wynde’s intended
marriage is repugnant to me, and that I cannot understand how her
ladyship can take Mr. Craven Black into papa’s place, but I have
tried to reconcile myself to the change. And I think,” added Neva, her
tones gathering firmness, and a brave look shining in her eyes of red
gloom, “that I have not sufficient excuse for appealing to the law to
give me a change of guardians. I shall have little to do or say to Mr.
Craven Black, and Hawkhurst is large enough for us both. It was
papa’s wish that I should remain for a certain period under the care
of Lady Wynde, and I cannot forget that she was papa’s wife, and
that he loved her. And more,” concluded Neva very gently, “if Lady
Wynde is about to contract an imprudent marriage, and if she is
likely to know sorrow because of her false step, she will need my
friendship when the truth comes home to her. I thank you again, Sir
John, Lord Towyn, Mr. Atkins, but I do not think I should be justified
in taking the decided step you advise.”
“I don’t know but you are right, Neva,” said Sir John. “At any rate,
give your ideas of duty a fair trial, and if you change your mind let us
know. It is not as if you were going away from us. Mr. Black, finding
himself in a quiet, decorous neighborhood, may choose to settle
down, and become a better man. We shall see you frequently, and
my house will always be open to you, my dear, and my wife and girls
will always be glad to receive you as an inmate of our family.”
“I shall not forget your kindness, Sir John,” said Neva gratefully.
“Miss Neva has always a way of escape from an unpleasant
situation,” said the practical Mr. Atkins. “Her marriage will free her
from Lady Wynde’s guardianship without publicity of an unpleasant
description.”
Neva reddened vividly.
The frankness with which the conversation had been distinguished
had considerably surprised the young earl. No one seemed to
require the use of diplomacy in making plain an unpleasant meaning,
and even Lady Wynde did not seem offended at the utterance of
home truths from the lips of Mr. Atkins. It was an hour for plain-
dealing, which was freely indulged in.