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Biology the dynamic science Third Edition, International
Edition / Hertz Digital Instant Download
Author(s): Hertz, Paul E.;Russell, Peter J.;McMillan, Beverly
ISBN(s): 9781133592051, 1133592058
Edition: Third edition, International edition /
File Details: PDF, 255.32 MB
Year: 2014
Language: english
BIOLOGY
the dynamic science third edition
Russell Hertz McMillan

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 Biology: The Dynamic Science, Third Edition, © 2014, 2011
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1 2 3 4 5 6 7 16 15 14 13 12
 Brief Contents

1 Introduction to Biological Concepts and 29 Seed Plants   630

Research   1 30 Fungi   649
31 Animal Phylogeny, Acoelomates, and
Unit One Molecules and Cells Protostomes   670
2 Life, Chemistry, and Water   22 32 Deuterostomes: Vertebrates and Their Closest
3 Biological Molecules: The Carbon Compounds Relatives   706
of Life   42
4 Energy, Enzymes, and Biological Reactions   71 Unit Five Plant Structure and Function
5 The Cell: An Overview   90 33 The Plant Body   748
6 Membranes and Transport   119 34 Transport in Plants   774
7 Cell Communication   141 35 Plant Nutrition   793
8 Harvesting Chemical Energy: Cellular 36 Reproduction and Development in Flowering
Respiration   161 Plants   811
9 Photosynthesis   182 37 Plant Signals and Responses to the Environment   838
10 Cell Division and Mitosis   205
Unit Six Animal Structure and Function
Unit Two Genetics 38 Introduction to Animal Organization and
11 Meiosis: The Cellular Basis of Sexual Physiology   867
Reproduction   225 39 Information Flow and the Neuron   883
12 Mendel, Genes, and Inheritance   239 40 Nervous Systems   903
13 Genes, Chromosomes, and Human Genetics   261 41 Sensory Systems   921
14 DNA Structure, Replication, and Organization   286 42 The Endocrine System   943
15 From DNA to Protein   310 43 Muscles, Bones, and Body Movements   966
16 Regulation of Gene Expression   339 44 The Circulatory System   981
17 Bacterial and Viral Genetics   369 45 Defenses against Disease   1002
18 DNA Technologies: Analyzing and Modifying 46 Gas Exchange: The Respiratory System   1024
Genes   391 47 Animal Nutrition   1043
19 Genomes and Proteomes   416 48 Regulating the Internal Environment   1069
49 Animal Reproduction   1096
Unit Three Evolutionary Biology 50 Animal Development   1118
20 Development of Evolutionary Thought   439
21 Microevolution: Genetic Changes within Unit Seven Ecology and Behavior
Populations   456
51 Ecology and the Biosphere   1145
22 Speciation   480
52 Population Ecology   1172
23 Paleobiology and Macroevolution   499
53 Population Interactions and Community Ecology   1199
24 Systematics and Phylogenetics: Revealing
54 Ecosystems   1229
the Tree of Life   528
55 Biodiversity and Conservation Biology   1254
Unit Four Biodiversity 56 Animal Behavior   1278

25 The Origin of Life   553

Appendix A: Answers   A-1
26 Prokaryotes: Bacteria and Archaea   567
Appendix B: Classification System   A-36
27 Protists   587
Glossary   G-1
28 Seedless Plants   611
Index   I-1

iii
 Peter J. Russell received a B.Sc. in Biology from the University of Sussex, England, in 1968
and a Ph.D. in Genetics from Cornell University in 1972. He has been a member of the Biology
faculty of Reed College since 1972 and is currently a professor of biology, emeritus. Peter taught
a section of the introductory biology course, a genetics course, and a research literature course
on molecular virology. In 1987 he received the Burlington Northern Faculty Achievement Award
from Reed College in recognition of his excellence in teaching. Since 1986, he has been the au-
thor of a successful genetics textbook; current editions are iGenetics: A Molecular Approach,
iGenetics: A Mendelian Approach, and Essential iGenetics. Peter’s research was in the area of
molecular genetics, with a specific interest in characterizing the role of host genes in the rep-
lication of the RNA genome of a pathogenic plant virus, and the expression of the genes of
the virus; yeast was used as the model host. His research has been funded by agencies includ-
ing the National Institutes of Health, the National Science Foundation, the American Cancer
Society, the Department of Defense, the Medical Research Foundation of Oregon, and the
Murdoch Foundation. He has published his research results in a variety of journals, including
Genetics, Journal of Bacteriology, Molecular and General Genetics, Nucleic Acids Research, Plasmid,
and Molecular and Cellular Biology. Peter has a long history of encouraging faculty research involv-
ing undergraduates, including cofounding the biology division of the Council on Undergraduate
Research in 1985. He was Principal Investigator/Program Director of a National Science Foundation
Award for the Integration of Research and Education (NSF–AIRE) to Reed College, 1998 to 2002.

Paul E. Hertz was born and raised in New York City. He received a B.S. in Biology from Stanford
University in 1972, an A.M. in Biology from Harvard University in 1973, and a Ph.D. in Biology from
Harvard University in 1977. While completing field research for the doctorate, he served on the Biol-
ogy faculty of the University of Puerto Rico at Rio Piedras. After spending two years as an Isaac Wal-
ton Killam Postdoctoral Fellow at Dalhousie University, Paul accepted a teaching position at Barnard
College, where he has taught since 1979. He was named Ann Whitney Olin Professor of Biology in
2000, and he received The Barnard Award for Excellence in Teaching in 2007. In addition to serving
on numerous college committees, Paul chaired Barnard’s Biology Department for eight years and
served as Acting Provost and Dean of the Faculty from 2011 to 2012. He is the founding Program Di-
rector of the Hughes Science Pipeline Project at Barnard, an undergraduate curriculum and research
program that has been funded continuously by the Howard Hughes Medical Institute since 1992. The
Pipeline Project includes the Intercollegiate Partnership, a program for local community college stu-
Aaron Kinard

dents that facilitates their transfer to four-year colleges and universities. He teaches one semester of
the introductory sequence for Biology majors and pre-professional students, lecture and laboratory
courses in vertebrate zoology and ecology, and a year-long seminar that introduces first-year students
to scientific research. Paul is an animal physiological ecologist with a specific research interest in the
thermal biology of lizards. He has conducted fieldwork in the West Indies since the mid-1970s, most
recently focusing on the lizards of Cuba. His work has been funded by the NSF, and he has published
his research in such prestigious journals as The American Naturalist, Ecology, Nature, Oecologia, and
Proceedings of the Royal Society. In 2010, he and his colleagues at three other universities received
funding from NSF for a project designed to detect the effects of global climate warming on the biol-
ogy of Anolis lizards in Puerto Rico.

Beverly McMillan has been a science writer for more than 25 years. She holds undergraduate
and graduate degrees from the University of California, Berkeley, and is coauthor of a college text
in human biology, now in its tenth edition. She has also written or coauthored numerous trade
books on scientific subjects and has worked extensively in educational and commercial publishing,
including eight years in editorial management positions in the college divisions of Random House
and McGraw-Hill.
Courtesy of Beverly McMillan

iv
 Preface

Welcome to the third edition of Biology: The Dynamic Science. thoughtful organization, a logical and seamless flow of topics
The book’s title reflects the speed with which our knowledge of within chapters, and carefully designed illustrations are key to
biology is growing. Although biologists have made enormous our approach.
progress in solving the riddles posed by the living world, every
discovery raises new questions and provides new opportunities Focusing on research to help students engage
for further research. As in the prior two editions, we have en-
the living world as scientists . . .
capsulated the dynamic nature of biology in the third edition by
explaining biological concepts—and the data from which they A primary goal of this book is to sharpen and sustain students’
are derived—in the historical context of each discovery and by curiosity about biology, rather than dulling it with a mountain
describing what we know now and what new discoveries will be of disconnected facts. We can help students develop the mental
likely to advance the field in the future. habits of scientists and a fascination with the living world by
conveying our passion for biological research. We want to excite
students not only with what biologists know about the living
Building on a strong foundation . . .
world but also with how they know it and what they still need to
The first two editions of this book provided students with the learn. In doing so, we can encourage some students to accept the
tools they need to learn fundamental biological concepts, pro- challenge and become biologists themselves, posing and an-
cesses, and facts. More important, they enabled students to swering important new questions through their own innovative
think like scientists. Our approach encourages students to think research. For students who pursue other careers, we hope that
about biological questions and hypotheses through clear exam- they will leave their introductory—and perhaps only—biology
ples of hypothesis development, observational and experimental course armed with intellectual skills that will enable them to
tests of hypotheses, and the conclusions that scientists draw evaluate future discoveries with a critical eye.
from their data. The many instructors and students who have In this book, we introduce students to a biologist’s “ways of
used the book have generously provided valuable feedback knowing.” Research biologists constantly integrate new obser-
about the elements that enhanced student learning. We have vations, hypotheses, questions, experiments, and insights with
also received comments from expert reviewers. As a result of existing knowledge and ideas. To help students engage the
these inputs, every chapter has been revised and updated, and world as biologists do, we must not simply introduce them to
some units have been reorganized. In addition, the third edition the current state of knowledge. We must also foster an appre-
includes new or modified illustrations and photos as well as ciation of the historical context within which those ideas devel-
some new features. oped, and identify the future directions that biological research
is likely to take.
To achieve these goals, our explanations are rooted in the
Emphasizing the big picture . . .
research that established the basic facts and principles of biol-
In this textbook, we have applied our collective experience as ogy. Thus, a substantial proportion of each chapter focuses on
teachers, researchers, and writers to create a readable and un- studies that define the state of biological knowledge today.
derstandable introduction that provides a foundation for stu- When describing research, we first identify the hypothesis or
dents who choose to enroll in more advanced biology courses in question that inspired the work and then relate it to the
the future. We provide straightforward explanations of funda- broader topic under discussion. Our research-oriented theme
mental concepts presented, where appropriate, from the evolu- teaches students, through example, how to ask scientific ques-
tionary perspective that binds the biological sciences together. tions and pose hypotheses, two key elements of the scientific
Recognizing that students in an introductory biology course process.
face a potentially daunting amount of material, we strive to pro- Because advances in science occur against a background
vide an appropriate balance between facts and concepts, taking of research, we also give students a feeling for how biologists of
great care to provide clear explanations while maintaining the the past formulated basic knowledge in the field. By fostering
narrative flow. In this way students not only see the big picture, an appreciation of such discoveries, given the information and
but they understand how we achieved our present knowledge. theories available to scientists in their own time, we can help
Having watched our students struggle to navigate the many ar- students understand the successes and limitations of what we
cane details of college-level introductory biology, we constantly consider cutting edge today. This historical perspective also
remind ourselves and each other to “include fewer facts, provide encourages students to view biology as a dynamic intellectual
better explanations, and maintain the narrative flow,” thereby enterprise, not just a collection of facts and generalities to be
enabling students to see the big picture. Clarity of presentation, memorized.

v
 We have endeavored to make the science of biology come as Escherichia coli, Drosophila, Arabidopsis, Caenorhabditis,
alive by describing how biologists formulate hypotheses and the mouse, and Anolis—and explain why they are used as
evaluate them using hard-won data; how data sometimes tell subjects for in-depth analysis.
only part of a story; and how the results of studies often end up
Three types of specially designed research figures provide
posing more questions than they answer. Although students
more detailed information about how biologists formulate and
might prefer simply to learn the “right” answer to a question,
test specific hypotheses by gathering and interpreting data. The
they must be encouraged to embrace “the unknown,” those
research figures are listed on the endpapers at the back of the
gaps in knowledge that create opportunities for further re-
book.
search. An appreciation of what biologists do not yet know will
draw more students into the field. And by defining why scien- • Experimental Research figures describe specific studies in
tists do not understand interesting phenomena, we encourage which researchers used both experimental and control treat-
students to think critically about possible solutions and to fol- ments—either in the laboratory or in the field—to test hy-
low paths dictated by their own curiosity. We hope that this potheses or answer research questions by manipulating the
approach will encourage students to make biology a part of system they studied.
their daily lives by having informal discussions and debates • Observational Research figures describe specific studies in
about new scientific discoveries. which biologists have tested hypotheses by comparing sys-
tems under varying natural circumstances.
• Research Method figures provide examples of important
Presenting the story line of the research techniques, such as the scientific method, cloning a gene,
process . . . DNA microarray analysis, plant cell culture, producing
monoclonal antibodies, radiometric dating, and cladistic
In preparing this book, we developed several special features to
analysis. Each Research Method figure leads a student
help students broaden their understanding of the material pre-
through the purpose of the technique and protocol and de-
sented and of the research process itself. A Visual Tour of these
scribes how scientists interpret the data it generates.
features and more begins on page xiii.
• The chapter openers, entitled Why It Matters . . . , are engag- Integrating effective, high-quality visuals
ing, short vignettes designed to capture students’ imagina-
into the narrative . . .
tions and whet their appetites for the topic that the chapter
addresses. In many cases, this feature tells the story of how a Today’s students are accustomed to receiving ideas and infor-
researcher or researchers arrived at a key insight or how bio- mation visually, making the illustrations and photographs in a
logical research solved a major societal problem, explained a textbook important. Our illustration program provides an ex-
fundamental process, or elucidated a phenomenon. The ceptionally clear supplement to the narrative in a style that is
Why It Matters . . . also provides a brief summary of the con- consistent throughout the book. Graphs and anatomical draw-
tents of the chapter. ings are annotated with interpretative explanations that lead
• To complement this historical or practical perspective, each students, step by step, through the major points they convey.
chapter closes with a brief essay entitled Unanswered For the second edition, we undertook a rigorous review of
Questions, prepared by an expert or experts in the field. all the art in the text. The publishing team identified the key
These essays identify important unresolved issues relating to elements of effective illustrations. In focus groups and surveys,
the chapter topic and describe cutting-edge research that instructors helped us identify the “Key Visual Learning Fig-
will advance our knowledge in the future. ures” covering concepts or processes that demand premier vi-
• Each chapter includes a short, boxed essay entitled Insights sual learning support. Each of these figures was critiqued by
from the Molecular Revolution, which describes how molec- our Art Advisory Board to ensure its usability and accuracy.
ular tools allow scientists to answer questions that they could For the third edition, we again evaluated each illustration and
not have posed even 30 years ago. Most Insights focus on a photograph carefully and made appropriate changes to im-
single study and include sufficient detail for its content to prove their use as teaching tools. New illustrations for the edi-
stand alone. tion were created in the same style as existing ones.
• Many chapters are further supplemented with one or more For the third edition, important figures were developed as
short, boxed essays involving three different aspects of re- Closer Look figures; a Summary and a concluding Think Like a
search. Focus on Basic Research essays describe how research Scientist question are designed to enhance student learning.
has provided understanding of basic biological principles. Many Closer Look figures involve key biological processes, such
Focus on Applied Research essays describe research designed as meiosis, transcription, muscle contraction, the cohesion-
to solve practical problems in the world, such as those relat- tension mechanism of water transport in plants, ecological in-
ing to health or the environment. Focus on Model Research teractions between predators and prey, and the haplodiploidy
Organisms essays introduce model research organisms—such genetic system in social insects.

vi Prefac e
Organizing chapters around important developed in the chapter to novel situations. Many end-of-
chapter questions also serve as good starting points for class
concepts . . .
discussions or out-of-class assignments.
As authors and college teachers, we understand how easily stu-
• Review Key Concepts provides a summary of important
dents can get lost within a chapter. When students request ad-
ideas developed in the chapter, referencing specific figures
vice about how to read a chapter and learn the material in it, we
and tables in the chapter. These Reviews are no substitute
usually suggest that, after reading each section, they pause and
for reading the chapter, but students may use them as a
quiz themselves on the material they have just encountered.
valuable outline of the material, filling in the details on
After completing all of the sections in a chapter, they should
their own.
quiz themselves again, even more rigorously, on the individual
• Understand & Apply includes five types of end-of-chapter
sections and, most important, on how the concepts developed
questions and problems that focus on the chapter’s factual
in the different sections fit together. Accordingly, we have ad-
content while encouraging students to apply what they have
opted a structure for each chapter to help students review con-
learned: (1) Test Your Knowledge is a set of 10 questions (with
cepts as they learn them.
answers in an appendix) that focus on factual material;
• The organization within chapters presents material in digest- (2) Discuss the Concepts involves open-ended questions that
ible sections, building on students’ knowledge and under- emphasize key ideas, the interpretation of data, and practical
standing as they acquire it. Each major section covers one applications of the material; (3) Design an Experiment ques-
broad topic. Each subsection, titled with a declarative sen- tions help students hone their critical thinking skills by ask-
tence that summarizes the main idea of its content, explores ing them to test hypotheses that relate to the chapter’s main
a narrower range of material. topic; (4) Interpret the Data questions help students develop
• Whenever possible, we include the derivation of unfamiliar analytical and quantitative skills by asking them to interpret
terms so that students will see connections between words graphical or tabular results of experimental or observational
that share etymological roots. Mastery of the technical lan- research experiments for which the hypotheses and methods
guage of biology will allow students to discuss ideas and pro- of analysis are presented; and (5) Apply Evolutionary Thinking
cesses precisely. At the same time, we have minimized the asks students to answer a question in relation to the princi-
use of unnecessary jargon. ples of evolutionary biology.
• Study Break questions follow every major section. These
questions encourage students to pause at the end of a section Helping students master key concepts
and review what they have learned before going on to the
throughout the course . . .
next topic within the chapter. Short answers to these ques-
tions appear in an appendix. Teachers know that student effort is an important determinant
of student success. Unfortunately, most teachers lack the time to
Encouraging active learning, critical thinking, develop novel learning tools for every concept—or even every
chapter—in an introductory biology textbook. To help address
and self-assessment of learning outcomes . . .
this problem, we are pleased to offer Aplia for Biology, an auto-
The third edition of Biology: The Dynamic Science includes a matically graded homework management system tailored to this
new active learning feature, Think Like a Scientist, which is de- edition. For students, Aplia provides a structure within which
signed to help students think analytically and critically about they can expand their efforts, master key concepts throughout
research presented in the chapter. Think Like a Scientist ques- the course, and increase their success. For faculty, Aplia can help
tions appear at the ends of Experimental Research figures, transform teaching and raise productivity by requiring more—
Observational Research figures, Closer Look figures, Insights from and more consistent—effort from students without increasing
the Molecular Revolution boxes, and Unanswered Questions. faculty workloads substantially. By providing students with con-
The new edition also includes Think Outside the Book, an tinuous exposure to key concepts and their applications
active learning feature introduced in the second edition. Think throughout the course, Aplia allows faculty to do what they do
Outside the Book activities have been designed to encourage best—respond to questions, lead discussions, and challenge the
students to explore biology directly or through electronic re- students.
sources. Students may engage in these activities either individ- We hope you agree that we have developed a clear, fresh,
ually or in small groups. and well-integrated introduction to biology as it is understood
Supplementary materials at the end of each chapter help by researchers today. Just as important, we hope that our efforts
students review the material they have learned, assess their un- will excite students about the research process and the biologi-
derstanding, and think analytically as they apply the principles cal discoveries it generates.

Preface vii
New to This Edition

The enhancements we have made in the third edition of Biology: structures in bacteria led scientists to propose and develop the
The Dynamic Science reflect our commitment to provide a text endosymbiotic theory. In Chapter 6 (Membranes and Trans-
that introduces students to new developments in biology while port), we point out that the close similarity of bilayer mem-
fostering active learning and critical thinking. As a part of this branes in all cells—prokaryotic and eukaryotic—is evidence
effort, we have added Closer Look figures that integrate a major that the basic structure of membranes evolved during the earli-
concept into a highlighted visual presentation. The key concept est stages of life on Earth, and has been conserved ever since. In
is stated briefly at the top, shown in detail through one or more Chapter 9 (Photosynthesis), a new section, Evolution of
illustrations, and summarized at the bottom. A Think Like a Photosynthesis and Cellular Respiration, summarizes the evolu-
Scientist question invites students to apply the figure concept(s) tionary development of these processes.
to a related problem or issue. We have also incorporated Think
Like a Scientist questions into Insights from the Molecular
Unit Two: Genetics
Revolution and Unanswered Questions, as well as into Experimental
Research and Observational Research figures. Chapter 11 (Meiosis: The Cellular Basis of Sexual Reproduc-
We have also made important changes in coverage to fol- tion) includes fuller descriptions of homologous chromosomes
low recent scientific advances. A new Chapter 19, Genomes and and sex chromosomes. Extensive revision and expansion of
Proteomes, introduces methods of genomics and proteomics Chapter 16 (Regulation of Gene Expression) provides more
along with examples of new discoveries and insights. In addi- thorough coverage of gene regulation and the operon model.
tion, we now devote two chapters to plant diversity, discussing We now introduce the role of DNA-binding proteins in pro-
seedless plants in Chapter 28 and seed plants in Chapter 29. karyotic as well as eukaryotic gene regulation and include more
Finally, we’ve consolidated our treatment of animal behavior detail on the activation of regulatory molecules. We have also
into a single Chapter 56 (Animal Behavior), which integrates added detail on combinatorial gene regulation, along with a fig-
various approaches to this subfield of biology. Beyond these ure showing a specific example, and have added new informa-
major organizational changes, we have made numerous im- tion and an illustration of how growth factors and growth-
provements to update and clarify scientific information and to inhibiting factors affect cell division. Chapter 17 (Bacterial and
engage students as interested readers and active learners, as Viral Genetics) includes new information on how horizontal
well as responsive scientific thinkers. The following sections gene transfer contributes to genome evolution in prokaryotes,
highlight some of the new content and organizational changes and evidence of its possible contribution to eukaryotic genome
in this edition. evolution.
A new Chapter 19 (Genomes and Proteomes) focuses on
the methods of genomics and the information it generates.
Unit One: Molecules and Cells
This chapter describes how genome sequences are determined
To make molecular and cellular processes easier to grasp, this and annotated, how genes in genomes are identified and char-
unit incorporates explanatory material into many more illustra- acterized, and how studies have generated new information
tions. For example, in Chapter 3 (Biological Molecules: The Car- on the evolution of genes and of genomes. It also includes ex-
bon Compounds of Life), Table 3.1 now presents more informa- amples of how genomics has become a source of new discov-
tion on the roles of functional groups of organic molecules and eries in many fields, including human physiology and evolu-
a new Figure 3.3 clarifies the concept of stereoisomers. In Chap- tionary biology.
ter 5 (The Cell: An Overview), we have combined the diagrams
of animal and plant cells in Figure 5.9 and labeled functions of
Unit Three: Evolutionary Biology
the organelles. In Chapter 8 (Harvesting Chemical Energy: Cel-
lular Respiration), a new overview diagram of glycolysis (Fig- In Chapter 21 (Microevolution: Genetic Changes within Popu-
ure 8.7) helps students understand basic concepts. Chapter 10 lations), Observational Research Figure 21.11 now shows more
(Cell Division and Mitosis) features a new discussion and illus- clearly how opposing forces of directional selection produce sta-
tration of the tight pairing of chromatids (sister chromatid co- bilizing selection. Plant speciation by alloploidy and polyploidy
hesion) during mitosis. is shown in parallel illustrations in one figure (22.16), allowing
New references to molecular aspects of evolution have been easy comparison. In Chapter 23 (Paleobiology and Macroevolu-
integrated into Unit One chapters to emphasize evolution as tion), Figure 23.15 clarifies our understanding of the rise and fall
the theme unifying the subfields of the biological sciences. For of plant lineages through evolutionary time. Chapter 24 (Sys-
example, in Chapter 5 (The Cell: An Overview), the discussion tematics and Phylogenetics: Revealing the Tree of Life) includes
of the mitochondrial matrix now highlights how equivalent a new example of how systematists construct phylogenetic trees

viii
with genetic distance data and includes a clarified discussion of refined diagrams of floral whorls and self-incompatibility, an
statistical methods used to construct phylogenetic trees. Re- updated Insights from the Molecular Revolution on trichome de-
worked phylogenetic trees throughout Chapter 24 are now fully velopment, and an updated Experimental Research figure on
consistent in presentation. studies of floral organ identity genes. Chapter 37 (Plant Signals
and Responses to the Environment) begins with a new Why It
Matters essay presenting the diverse adaptations of creosote
Unit Four: Biodiversity
bush (Larrea tridentata) to environmental challenges such as ex-
In Chapter 26 (Prokaryotes: Bacteria and Archaea), a new tended drought. The chapter also has been reorganized, with the
Insights from the Molecular Revolution describes how changes in discussion of signal transduction pathways and second messen-
gene expression in the bacterium that causes gingivitis help gov- ger systems now included in the introduction to plant hor-
ern its transition from a free-living state to a biofilm. A revised mones. New art illustrates current thinking on different signal
and expanded section discusses the five subgroups of proteo- transduction mechanisms in plant cells.
bacteria. In Chapter 27 (Protists), we’ve added the nucleariids to
the Opisthokont group, along with evidence that they may be
Unit Six: Animal Structure and Function
more closely related to fungi than to animals.
Plant diversity is now covered in two chapters. Chapter 28 In Chapter 43 (Muscles, Bones, and Body Movements), a new
(Seedless Plants) describes trends in land plant evolution and Insights from the Molecular Revolution presents experiments on
the characteristics of bryophytes and seedless vascular plants, exercise training in racehorses. We have updated and clarified
and Chapter 29 (Seed Plants) focuses on adaptations and dis- the discussion of immunity in Chapter 45 (Defenses against
tinguishing features of gymnosperms and flowering plants. Disease) and added new material on how microbial pathogens
Chapter 28 also includes a new Unanswered Questions essay, are detected and how pathogens may sometimes escape recog-
and Chapter 29 presents a new Insights from the Molecular nition by the immune system. In Chapter 47 (Animal Nutri-
Revolution feature on plant genome evolution. Chapter 30 tion), we have added detail on absorption in the small intestine.
(Fungi) presents an updated discussion of the evolution of mul- Chapter 48 (Regulating the Internal Environment) has ex-
ticellular animals and fungi from different opisthokont ances- panded coverage of mammalian kidney function and the role of
tors. Changes to Chapter 31 (Animal Phylogeny, Acoelomates, countercurrent heat exchanges in maintaining body tempera-
and Protostomes) include color-coding of anatomical illustra- ture. In Chapter 50 (Animal Development), we have revised and
tions of invertebrates to distinguish structures arising from en- expanded Section 50.5, The Cellular Basis of Development, in-
doderm, mesoderm, and ectoderm, and a new Table 31.1 pro- cluding new information on apoptosis during development and
viding a phylogenetic overview of the phyla presented in the on molecular mechanisms of induction.
chapter. In Chapter 32 (Deuterostomes: Vertebrates and Their
Closest Relatives), a new Insights from the Molecular Revolution
Unit Seven: Ecology and Behavior
feature describes a study of the evolutionary gains and losses of
genes that code for olfactory receptor proteins in various clades In Chapter 51 (Ecology and the Biosphere), improved illustra-
of mammals. Figure 32.38, which shows timelines for the spe- tions clarify the effects of latitudinal and seasonal variations in
cies of hominins, has been updated with recently discovered incoming solar radiation. In Chapter 52 (Population Ecology),
fossils. The discussion of human evolution includes recent ge- we have updated Figures 52.22 and 52.23 on human population
nomic studies of the relationship between Neanderthals and growth. In Chapter 53 (Population Interactions and Commu-
modern humans. nity Ecology), we have improved Figure 53.22 showing the food
web. We have also added informative labels to Figure 53.25,
which shows the effects of storms on corals. Chapter 55 (Biodi-
Unit Five: Plant Structure and Function
versity and Conservation Biology) features a new Figure 55.16
Chapter 33 (The Plant Body) features clearer illustrations of illustrating the species–area relationship. A unified Chapter 56
plant growth. Chapter 34 (Transport in Plants) has more fo- (Animal Behavior) concludes the text, integrating the discus-
cused discussions and illustrations of water movements in roots sions of genetic and experiential bases of animal behavior, the
and the physiology of stomatal function. A new Unanswered neurophysiological and endocrinological control of specific be-
Questions explores research in plant metabolomics. Chapter 36 haviors, and the ecology and evolution of several broad catego-
(Reproduction and Development in Flowering Plants) includes ries of animal behavior.

New to This Editio n ix

This page intentionally left blank
 THINK AND ENGAGE LIKE A SCIENTIST!
Develop a deep understanding of the core concepts in biology
and build a strong foundation for future courses.

Welcome to Biology: The Dynamic Science, Third

Edition, by Peter J. Russell, Paul E. Hertz, and
Beverly McMillan. The authors convey their passion
for biology as they guide you to an understanding
of what scientists know about the living world, how
they know it, and what they still need to learn. The
pages that follow highlight a few of the many ways
that they have made this book a great learning tool
for you. You’ll also find information about dynamic
online resources, as well as print materials that will
help you master key concepts and succeed in the
course.

Biology
the dynamic science third edition
Russell Hertz McMillan

Untitled-2 1 9/4/12 12:34 PM

Aplia for Biology, an interactive

online tool that complements
the text and helps you learn
and understand key concepts
through focused assignments,
an engaging variety of problem
types, exceptional text/art
integration, and immediate
feedback.

xi
 A BIG PICTURE FOCUS
Straightforward explanations of fundamental concepts bind the
biological sciences together and enable you to see the big picture.
Easy-to-use learning tools point out the topics covered
But atinthe
each
southern end of the Central Valley, adjacent
chapter, show why they are important, and help you learn rarely interbreed. Apparently, they have
subspecies
the material. differentiated to such an extent that they can no lon-
ger exchange genetic material directly.
Are the southernmost populations of this sala-
mander subspecies or different species? A biologist
who saw only the southern populations, which coex-

22 ist without interbreeding, might define them as

separate species. However, they still have the po-
tential to exchange genetic material through the
intervening populations that form the ring. Painted salamander
Hence, biologists recognize these populations as (E. e. picta)
belonging to the same species. Most likely, the
southern subspecies are in an intermediate stage

TOM VEZO/MINDEN PICTURES/

National Geographic Stock
of species formation.

CLINAL VARIATION When a species is distributed Yellow-eyed salamander

study outline Study Outline provides an overview of
Two closely related bird species, purple martins (Progne subis) and tree swallows (Tachycineta bicolor) perching together
(E. e. xanthoptica)
over a large, environmentally diverse area, some
on a branch in Crane Creek, Ohio.

22.1 What Is a Species?

22.2 Maintaining traits may exhibit a cline, a pattern mainofchapter topics and key concepts.
smooth varia-
Speciation
Reproductive Isolation
22.3 The Geography of tion along a geographical gradient. Each section breaks the material into a
For example, Tw
in
Speciation
22.4 Genetic Mechanisms
many birds and mammals in the northern
manageable amount of information, so p
of Speciation
hemisphere show clinal variation in body size (Fig-
you can develop understanding as you di
Why it matters . . . In 1927, nearly 100 years after Darwin in
boarded the Beagle, a young German naturalist namedure Ernst22.5)
Mayr em- and the relative length of theirknowledge.
acquire appendages. Monterey salamander
barked on his own journey, to the highlands of New Guinea. He was search-
In known
ing for rare “birds of paradise” (Figure 22.1). These birds were general,
in Eu- populations living in colder environ- (E. e. eschscholtzii)
rope only through their ornate and colorful feathers, which were used to
ments
decorate ladies’ hats. On his trek through the remote Arfak Mountains, have larger bodies and shorter appendages, a
pattern that is usually interpreted as a mechanism Figure 22.4 ring species. Six of the
Mayr identified 137 bird species (including many birds of paradise) based on
differences in their size, plumage, color, and other external characteristics.
to conserve body heat (see Chapter
To Mayr’s surprise, the native Papuans—who were untrained in the Why It48).Matters sections at the beginning
are distributed in a ring around California
ways of Western science, but who hunted these birds for food and feath-
of each chapter
Clinal variation usually results from gene
ers—had their own names for 136 of the 137 species he had identified. The capture the
geographicalexcitement
distributions overlap. Howe
close match between the two lists confirmed Mayr’s belief that the species south (marked with an arrow), the Monte
flowhasbetween
is a fundamental level of organization in nature. Each species a unique of biology
adjacent populations thatandarehelp
each you understand why
rarely interbreed.
combination of genes underlying its distinctive appearance and habits.
Thus, people who observe them closely—whether indigenous adaptinghunters orto slightly different
the conditions.
topic is How- ©and
important Cengagehow
Learning the
2014
Western scientists—can often distinguish one species from another.
Mayr also discovered some remarkable patterns inever, if populations at opposite
the geographical ends of a cline are
material that follows fits into the big
Bruce Beehler

distributions of the bird species in New Guinea. For example, each moun-
tain range he explored was home to some species that lived separated
nowhere else. by great distances, they may exchange very
picture.
Figure 22.1 Birds of paradise. A male Count Raggi’s bird of
paradise (Paradisaea raggiana) tries to attract the attention of a of unsuitable habitat. In 1942, Mayr little
published genetic
Closely related species often lived on different mountaintops, separated by
deep valleys the book material through reproduction. Thus, when a cline
female (not pictured) with his showy plumage and conspicuous
display. There are 43 known bird of paradise species, 35 of them
extends
Systematics and the Origin of Species, in which he described the role of ge-
ography in the evolution of new species; the book quickly became a corner-
over a large geographical gradient, distant populations
found only on the island of New Guinea.
Black rat snake
stone of the modern synthesis (which was outlined in Sectionmay be genetically and morphologically distinct.
20.3).

480 (E. o. obsoleta) Despite the geographical variation that many species exhibit, Chapt

even closely related species are genetically and morphologically

different from each other. In the next section, we consider the
mechanisms that maintain the genetic distinctness of closely re-
lated species by preventing their gene pools from mixing.

Study Break sections encourage you to Study Break 22.1

pause 1. How do the morphological, biological, and phylogenetic species
Yellow ratand
snakethink about the material you
concepts differ?
have just encountered before moving
(E. o. quadrivittata)
2. What is clinal variation?
to the next section.

>
think outside the book
Search the biological literature and the Internet for a definition
and application of the “ecological species concept.” How does
the ecological species concept differ from the three species
Figure 22.5 Clin
xii concepts described above? under what circumstances or for
clinal variation in ove
at snake Everglades rat snake what purpose would the ecological species concept be useful? of 16 skeletal features
spiloides) (E. o. rossalleni) © Cengage Learning 2014
 p from the white-flowered parent produces an
F1 generation of all Pp plants, which have
purple flowers because the P allele is
Gamete from dominant to the p allele. Because they have
parent with P
two different alleles of a gene, the plants are

THINK LIKE A SCIENTIST

purple flowers said to be heterozygous for that gene. The F1
Pp
heterozygote is called a monohybrid.

4. F1  F1 self
Your study of biology focuses not only on what scientists know
Mendel nowabout
performed a the
monohybrid cross by allowing
living world but also how they know it. Use these unique features
 toto self
F purple Pp plants learn
produce the F generation.
and 1
2
how scientists ask scientific questions,
Pp
pose hypotheses,
Pp
and test them.
5. F2 generation
 Gametes from
Pp F1 plant
Experimental Research

NEW!
Figure 12.5

P p
The Principle of Segregation: Inheritance of Flower Color in Garden Peas
Question: How is flower color in garden peas inherited?

Experiment: Mendel crossed a true-breeding purple-flowered plant with a true-breeding white-flowered plant and analyzed the progeny through
the F1 and F2 generations. We explain this cross here in modern terms.

“Think Like a Scientist” questions ask you to apply what  Gametes from P
1. P generation
Purple White
P is the dominant allele for p is the recessive allele for
purple; the true-breeding white; the true-breeding
purple-flowered parent has white-flowered parent has
the PP combination of alleles.  the pp combination of
PP Pp
you have learned beyond the material presented in the text. Pp F plant
The plant is homozygous for alleles. The plant is
PP pp
The P and p gametes fused to 1
the P allele. homozygous for the p allele.

produce the F generation. 2. Haploid gametes

2

These questions are incorporated into Experimental Research,

The two alleles separate during The two alleles separate during
gamete formation: only gamete formation: only
P p
p
gametes with the P allele are gametes with the p allele are
produced in a PP plant. produced in a pp plant.

Observational Research, and Closer Look figures, as well as

3. F1 generation
Pp pp Gamete from parent
with white flowers Fusion of the P gamete from the
purple-flowered parent with the p gamete
p from the white-flowered parent produces an
F1 generation of all Pp plants, which have

into Insights from the Molecular Revolution and Unanswered

purple flowers because the P allele is

Results: Mendel’s selfing of the F purple-flowered plants produced an F generation consisting of 3/4 purple-flowered and 1/4 white-flowered
Gamete from dominant to the p allele. Because they have
P
1 2 parent with
purple flowers
two different alleles of a gene, the plants are
Pp said to be heterozygous for that gene. The F1

plants. White flowers were inherited as a recessive trait, disappearing in the F and reappearing in the F . 1
heterozygote is called a monohybrid.
2

Questions boxes.

© Cengage Learning 2014

4. F1  F1 self

Conclusion: The results supported Mendel’s Principle of Segregation hypothesis that the pairs of alleles that control a character segregate as 
Mendel now performed a
monohybrid cross by allowing
F1 purple Pp plants to self and

gametes are formed, with half of the gametes carrying one allele, and the other half carrying the other allele. Pp Pp
produce the F2 generation.

5. F2 generation
 Gametes from
think like a scientist Suppose you pick at random one of the F2 purple-flowered plants and allow it to self. P
Pp F1 plant

What is the chance that the progeny will include both purple-flowered and white-flowered plants?
P
 Gametes from PP Pp
Pp F1 plant The P and p gametes fused to
produce the F2 generation.

Pp pp

InsIghts From the Molecular Revolution Results: Mendel’s selfing of the F1 purple-flowered plants produced an F2 generation consisting of 3/4 purple-flowered and 1/4 white-flowered
plants. White flowers were inherited as a recessive trait, disappearing in the F 1 and reappearing in the F2.

© Cengage Learning 2014

Hot Potatoes: Do plants use treatment resulted in an increase in UCP relatively recent evolutionary times with the Conclusion: The results supported Mendel’s Principle of Segregation hypothesis that the pairs of alleles that control a character segregate as

Insights from the Molecular Revolution essays highlight

uncoupling proteins to mRNA level in the leaves. appearance of birds and mammals, may be a gametes are formed, with half of the gametes carrying one allele, and the other half carrying the other allele.
much more ancient development.
generate heat? Conclusion think like a scientist Suppose you pick at random one of the F2 purple-flowered plants and allow it to self.
The research indicates that potato plants What is the chance that the progeny will include both purple-flowered and white-flowered plants?
Mammals use several biochemical and mo- think like a scientist The box shows
probably use the mitochondrial uncoupling
that potato plants produce a UCP that is

how molecular technologies allow researchers to answer

lecular processes to maintain body heat. One
process to warm tissues when they are similar to a mammalian UCP gene. What
process is shivering; the muscular activity of
stressed by low temperatures. Thus, mecha- are the evolutionary implications of that
shivering releases heat that helps keep body
nisms for warming body tissues once thought observation?
temperature at normal levels. Another mech-
to be the province only of animals appear to
anism operates through uncoupling proteins
be much more widespread. In particular,

questions that they could not even pose 20 or 30 years ago.

Source: M. Laloi et al. 1997. A plant cold-induced uncou-
(UCPs), which eliminate the mitochondrial pling protein. Nature 389:135–136.
UCPs, which were believed to have evolved in
H1 gradient by making the inner mitochon-
drial membrane leaky to protons. Electron
transfer and the oxidative reactions then run
at high rates in mitochondria without trap- Flowers—very high
ping energy in ATP. The energy is released as UCP mRNA level
Molecular Revolution acceptor molecule, while in anaerobic respiration the electrons raw materials to produce organic molecules in a process that
heat that helps maintain body temperature.
are transferred to an electron transfer system. However, in con- pushes the electrons back to high energy levels by absorbing the
Research Question trast to aerobic respiration, in anaerobic respiration the elec- energy of sunlight.
Do plants use UCPs to generate heat? trons are transferred through the electron transfer system to a
molecule other than oxygen, such as sulfate.
Experiments ºC This chapter traced the flow of high-energy electrons from Study Break 8.5
Research by Maryse Laloi and her colleagues 40
Stem—moderate fuel molecules to ATP. As part of the process, the fuels are bro- What is fermentation, and when does it occur? What are the two
UCP mRNA level ken into molecules of carbon dioxide and water. The next chap- types of fermentation?
at the Max Planck Institute for Molecular 30
20 ter shows how photosynthetic organisms use these inorganic
Plant Physiology in Germany shows that

Unanswered Questions
10
some tissues in plants may use the same pro-
0
cess involving UCPs to generate heat. Their unansweRed questions
evidence is as follows:
Leaf—low UCP
1. Potato plants (Solanum tuberosum) have a Glycolysis and energy metabolism are crucial for the normal functioning with altered genes so that they develop AD. (The generation of transgenic
mRNA level
gene with a DNA sequence similar to that of an animal. Research of many kinds is being conducted in this area, organisms is described in Chapter 18.)

explore important unresolved

of a mammalian UCP gene. The potato such as characterizing the molecular components of the reactions in The results of our experiments have demonstrated the levels that
detail and determining how they are regulated. The goal is to generate many mitochondrial proteins are altered in the brains of transgenic AD
gene protein is clearly related to the mam-
comprehensive models of cellular respiration and its regulation. Follow- mice. Interestingly, both down-regulated and up-regulated mitochondrial
malian protein, and also has the same Tuber—low UCP
ing is a specific example of ongoing research related to human disease proteins were identified in AD brains. These dysregulated mitochondrial
mRNA level
overall three-dimensional structure. caused by defects in cellular respiration. proteins participate in many different metabolic functions, including the
2. The researchers used the DNA of the po- citric acid cycle, oxidative phosphorylation, pyruvate metabolism, fatty

issues identified by experts

How do mitochondrial proteins change in patients with
Roots—very high acid oxidation, ketone body metabolism, metabolite transport, oxidative
tato UCP gene to probe for the presence alzheimer disease?
UCP mRNA level stress, mitochondrial protein synthesis, mitochondrial protein import,
of messenger RNA (mRNA), the gene Alzheimer disease (AD) is an age-dependent, irreversible, neurodegen-
and cell growth and apoptosis (a type of programmed cell death; see
erative disorder in humans. Symptoms include a progressive deteriora-
product that specifies the amino acid se- Figure 1 UCP mRNA levels of activity. Chapter 45). We have also determined that these changes in the mito-
tion of cognitive functions and, in particular, a significant loss of memory.
quence of proteins in the cytoplasm, with Neuropathologically, AD is characterized by the presence of extracellular
chondrial proteome occurs early in AD before the development of signifi-

in the field and describe

the results shown in Figure 1. These re- cant plaque and tangle pathologies. Future experiments will be directed
amyloid plaques, intracellular neurofibrillary tangles, and synaptic and
toward examining changes in the mitochondrial proteome in the brains
sults indicate that the potato UCP gene is neuronal loss. Reduced brain metabolism occurs early in the onset of AD.
of human AD patients. Ultimately, the results of our experiments may
active at different levels in various plant One of the mechanisms for this physiological change appears to be dam-
lead to the development of treatments that can slow or halt the progres-
20ºC After 1–3 days at 4ºC age to or reduction of key mitochondrial components, including enzymes
tissues, suggesting that certain tissues sion of AD in humans.
of the citric acid cycle and the oxidative phosphorylation system. How-
naturally need warming for optimal

cutting-edge research that

Low UCP Very high UCP ever, the complete scope of mitochondrial protein changes has not been
function. established, nor have detailed comparisons been made of mitochondrial
think like a scientist What significance do you attribute to the
mRNA level mRNA level statement in this essay that changes in levels of many mitochondrial
3. Laloi and her coworkers then tested protein changes among AD patients.
proteins occurs early in AD before the development of significant
To begin to address these questions, research is being carried out in
whether exposing potato plants to cold Figure 2 Results plaque and tangle pathologies? Develop a scientific hypothesis based
my laboratory at The University of Texas at Dallas to analyze quantitatively
temperatures could induce greater synthe- on this statement.
of exposure to cold the complete set of mitochondrial proteins in healthy versus AD brains.

will advance our knowledge

sis of the UCP mRNA (Figure 2). The cold temperature. The results will show the changes that occur in the mitochondrial pro-
teome in the two tissues. This approach is called quantitative compara- Gail a. Breen is an associate professor in the Department of
Molecular and Cell Biology at The University of Texas at Dal-
Courtesy of Gail A. Breen

© Cengage Learning 2014

tive proteomic profiling. (Proteomics is the complete characterization of las. Her current research focuses on mitochondrial biogene-
the proteins present in a cell, cell compartment, tissue, organ, or organ- sis and the role of mitochondria in neurodegenerative dis-
ism; see Chapter 19. The set of proteins identified in such a study is called eases, such as Alzheimer disease. To learn more about Dr.
Breen’s research go to http://www.utdallas.edu/biology/
the proteome.) We are using a transgenic mouse model of AD in the first

in the future.
faculty/breen.html.
stage of this research; that is, the mice have been genetically engineered
Some prokaryotic and eukaryotic cells are strict aerobes— Fermentation differs from anaerobic respiration, the form
that is, they have an absolute requirement for oxygen to survive of cellular respiration used by some prokaryotes for ATP pro-
and are unable to live solely by fermentations. Vertebrate brain duction (see Why It Matters for this chapter). In fermentation
cells are key examples of strict aerobes. the electrons carried by NADH are transferred to an organic review key concepts

178 U n i t On e MO l e cU l e s a n d c e l l s To access the course materials and companion resources for this
c h text,
a p t e r 8 ecule
h a rsuch
v e sas
t ioxygen;
n g c htheeMenergy
i c a lthat
e nisereleased
r g y: cdrives
e l lUATP
l a rsyn-
r e s p i r at i On
please visit www.cengagebrain.com. thesis (Figure 8.1).
• Oxidation–reduction reactions, called redox reactions, partially or
8.1 Overview of Cellular energy Metabolism completely transfer electrons from donor to acceptor atoms; the
• Plants and almost all other organisms obtain energy for cellular donor is oxidized as it releases electrons, and the acceptor is
activities through cellular respiration, the process of transferring reduced (Figure 8.2).
electrons from donor organic molecules to a final acceptor mol- • Cellular respiration occurs in three stages: (1) in glycolysis, glucose is
converted to two molecules of pyruvate through a series of enzyme-

Unit One M Olec Ules and cells chapter 8 har vesting cheM ical energy: cell Ular respirati O n 179

Malaria and the Plasmodium

Focus on Applied Research
tially in liver cells and then in red blood cells. ventative. However, many Plasmodium strains
Focus on Research boxes present research topics in more
depth.
Life Cycle Their growth causes red blood cells to rup- in Africa, India, and Southeast Asia have devel-
ture in regular cycles every 48 or 72 hours. oped resistance to the drugs. Vaccines have
Malaria is uncommon in the United States, The ruptured red blood cells clog vessels and proved difficult to develop; because vaccines
but it is a major epidemic in many other parts release the parasite’s metabolic wastes, caus- work by inducing the production of antibodies
of the world. From 300 million to 500 million ing cycles of chills and fever. that recognize surface groups on the parasites,
people get malaria each year in tropical re- The victim’s immune system is ineffective they are defeated by the same mechanisms the
gions, including Africa, India, Southeast Asia, because, during most of the infective cycle, the parasite uses inside the body to keep one step
the Middle East, Oceania, and Central and parasite is inside body cells and thus “hidden” ahead of the immune reaction.
South America. Of these, about 2 million die from antibodies. Further, Plasmodium regularly While in a malarial region, travelers
each year, which is twice as many deaths as changes its surface molecules, continually pro- should avoid exposure to mosquitoes by re-
from AIDS, worldwide. It is particularly deadly ducing new forms that are not recognized by maining indoors from dusk until dawn and
for children younger than six years old. antibodies developed against a previous form. sleeping inside mosquito nets treated with in-

Focus on Applied Research describes how scientific

Four different species of Plasmodium In this way, the parasite keeps one step ahead sect repellent. When out of doors, travelers
cause malaria. In the life cycle of the parasites of the immune system, often making malarial should wear clothes that expose as little skin
(Figure), sporozoites develop in a female infections essentially permanent. as possible and are thick enough to prevent
Anopheles mosquito, which transmits them by Travelers in countries with high rates of ma- mosquitoes from biting through the cloth. An

research has solved everyday problems.

its bite to human or bird hosts. The infecting laria are advised to use antimalarial drugs such insect repellent containing DEET should be
parasites divide repeatedly in their hosts, ini- as chloroquine, quinine, or quinidine as a pre- spread on any skin that is exposed.

Sporozoites
1 Plasmodium zygotes
undergo meiosis, Sporozoite
producing haploid
sporozoites in the gut
wall of a female
Anopheles mosquito.
The sporozoites migrate
to the mosquito’s
salivary glands. 2 When the infected
mosquito bites a human,
3 The sporozoites

Focus on Basic Research describes seminal research that

it injects sporozoites into
the blood, which carries reproduce asexually in liver
them to liver cells. cells, each producing
many merozoites.
Sinclair Stammers/Photo Researchers, Inc.

5 Some merozoites in
red blood cells develop
into immature male and
provided insight into an important problem.
female gamete cells,
Micrograph Steven L’Hernault

which are released into 4 The merozoites enter

the bloodstream. the bloodstream, invade
red blood cells, and
6 A female bites and reproduce asexually.
sucks blood from an Periodic breakdown of
infected human. Gamete red blood cells and

Focus on Model Research Organisms explains why

cells in the blood reach release of merozoites
her gut, mature, and fuse cause bouts of severe
Male gametocyte
by twos to form zygotes. chills and fever.
in red blood cell
Figure Life cycle of a Plasmodium species that causes malaria.

© Cengage Learning 2014

researchers use certain organisms as research subjects.

disease that is usually mild in adults but can cause severe brain plain. The flagella occur only on reproductive cells such as eggs
damage or even death to a fetus. and sperm, except in the golden algae, in which cells are flagel­
lated in all stages. The Stramenopila includes the Oomycota
STRAMENOPILA Stramenopila—stramenopiles—have two dif­ (water molds, white rusts, and mildews—formerly classified as
ferent flagella: one with hollow tripartite projections that give fungi), Bacillariophyta (diatoms), Chrysophyta (golden algae),
the flagellum a “hairy” appearance and a second one that is and Phaeophyta (brown algae).

Chapte r 27 pro ti sts 595

xiii
 ENGAGE LIKE A SCIENTIST
Be Active. Get involved in the process of learning and doing biology.

Research Figures provide information about how

biologists formulate and test specific hypotheses by Figure 10.7 Research Method

gathering and interpreting data.

Purpose: A karyotype is a display of chromosomes of an organism arranged in pairs.
Preparing a Human Karyotype
A normal karyotype has a characteristic appearance for each species. Examination of
the karyotype of the chromosomes from a particular individual indicates whether the
individual has a normal set of chromosomes or whether there are abnormalities in
number or appearance of individual chromosomes. A normal karyotype can be used
to indicate the species.

Protocol:

1. Add sample to culture medium that has stimulator 2. Stain the cells so that the chromosomes
for growth and division of cells (white blood cells in are distinguished. Some stains
the case of blood). Incubate at 37°C. Add colchicine, produce chromosome-specific

Research Method which blocks the formation of microtubules. As a

result, the spindle does not form and this causes
mitosis to arrest at metaphase.
banding patterns, as shown
in the photograph below.

Pair of sister chromatids closely

3. View the stained cells under a microscope equipped with a digital Pair of homologous aligned side-by-side by sister
imaging system and take a digital photograph. A computer processes chromosomes chromatid cohesion
the photograph to arrange the chromosomes in pairs and numbers
3. If the genotype frequencies in a population are 0.60 AA, 0.20 Aa, 9. The neutral variation hypothesis proposes that: them according to size and shape.

and 0.20 aa, and if the requirements of the Hardy–Weinberg a. complex structures in most organisms have not been fos-
Observational Research
principle apply, the genotype frequencies in the offspring genera-
Figure 22.18 Observational Research
tered by natural selection. 1 2 3 4 5

tion will be: b.Chromosomal

most mutations Similarities and have a strongly
Differences among harmful
the Great effect.
Apes
a. 0.60 AA, 0.20 Aa, 0.20 aa. c.Question:
some mutations
Does chromosome structure are not humans
differ between affected and theirby natural
closest selection.
relatives among the apes? 6 7 8 9 10 11 12

b. 0.36 AA, 0.60 Aa, 0.04 aa. d.evolutionary

natural selection cannot counteract the action of gene flow.
Hypothesis: Large scale chromosome rearrangements contributed to the development of reproductive isolation between species within the

© Medical-on-Line/Alamy
lineage that includes humans and apes. 13 14 15 16 17 18
c. 0.49 AA, 0.42 Aa, 0.09 aa. e.Prediction:
large populations are subject to stronger natural selection Interpreting the Results: The karyotype is evaluated with respect to the
Chromosome structure differs markedly between humans and their close relatives among the great apes: chimpanzees, gorillas, scientific question being asked. For example, it may identify a particular
d. 0.70 AA, 0.00 Aa, 0.30 aa. than small populations.
and orangutans. species, or it may indicate whether or not the chromosome set of a human
19 20 21 22 XY
e. 0.64 AA, 0.32 Aa, 0.04 aa. Method: Jorge J. Yunis and Om Prakash of the University of Minnesota Medical School used Giemsa stain to visualize the banding patterns on (fetus, child, or adult) is normal or aberrant.
10. Phenotypic
metaphase chromosome characteristics
preparations from humans, that increase
chimpanzees, the
gorillas, and fitness
orangutans. of individuals
They identified about 1,000 bands that are present in

4. The reason spontaneous mutations do not have an immediate ef- are called:
humans and in the three ape species. By matching the banding patterns on the chromosomes, the researchers verified that they were comparing
the same segments of the genomes in the four species. They then searched for similarities and differences in the structure of the chromosomes.
© Cengage Learning 2014

fect on allele frequencies in a large population is that: a.Results:mutations. Figure 14.9

Analysis of human chromosome 2 reveals that it was produced by the fusion of two smaller chromosomes that are still present in the
Experimental Research
be identified from this characteristic alone. Figure 10.7 shows how ceeds, the nucleolus reappears, RNA transcription resumes, and
a. mutations are random events, and mutations may be either b.other founder
chromosomes, effects.
three species. Although the position of the centromere in human chromosome 2 matches that of the centromere in one of the chimpanzee
in gorillas and orangutans it falls within an inverted segment of the chromosome.
The Meselson and Stahl Experiment
human chromosomes are prepared for analysis as a karyotype. Question: Does DNA replicate semiconservatively?
a new nuclear envelope forms around the chromosomes at each
Demonstrating the Semiconservative
beneficial or harmful. c. heterozygote advantages. Model for DNA Replication to Be Correct
ANAPHASE The proper alignment of chromosomes in metaphase
Experiment: Matthew Meselson and Franklin Stahl proved that pole
the
semiconservative model of DNA replication is correct and that the
producing the two
division is complete.
daughter nuclei. At this point, nuclear

b. mutations usually occur in males and have little effect on d. adaptive traits.
Human
triggers anaphase. An enzyme, separase, is activated and cleaves
1. Bacteria grown in 15N (heavy) 2. Bacteria transferred to 14N (light) medium
conservative and dispersive models are incorrect.
Mitosis produces two daughter nuclei, each with identical
eggs. e. polymorphisms. Centromere
entromere position is similar
in humans and chimpanzees.
the cohesin rings around the pairs of sister chromatids. This can-
medium. The heavy isotope is and allowed to grow and divide for several
cels the force opposing the pull of sister chromatids to opposite
incorporated into the bases of DNA, generations. All new DNA is light.
sets of chromosomes compared to the parental cell. Cytokine-
sis, the division of the cytoplasm, typically follows the nuclear
c. many mutations exert their effects after an organism has Chimpanzee resulting in all the DNA being heavy,
poles. Thus, 15during anaphase, the spindle separates sister chroma-
that is, labeled with N. division stage of mitosis, and produces two daughter cells each
stopped reproducing. Discuss the Concepts tids and pulls them to opposite spindle poles. The first signs of
chromosome movement can be seen at the centromeres, where
with one of the two daughter nuclei. Cytokinesis proceeds by
different pathways in the various kingdoms of eukaryotic or-
d. mutations are so rare that mutated alleles are greatly out-
Matching bands
tension developed by the spindle pulls14the kinetochores toward the ganisms. In animals, protists, and many fungi, a groove, the
numbered by nonmutated alleles. 1. MostGorillalarge commercial farms routinely administer antibiotics to poles. The movement
15N
medium continues untilmedium
N
the separated chromatids, furrow, girdles the cell and gradually deepens until it cuts the
igure 24.12 e.themost
principle of parsimony. farm animals to prevent the rapid spread of diseases through a now called daughter chromosomes, tion 15.1), mutations in the third codon position do not often
have reached the two poles. At cytoplasm into two parts (Figure 10.8). In plants, a new cell
mutations do not change the amino acid sequence of a
1st replication 2nd replication
this point, chromosome segregation has been completed. wall, called the cell plate, forms between the daughter nuclei
influence the amino acid composition of the protein for which
Compared
ompared to the chromosomes of humans and chimpanzees,

protein. flock or herd. Explain why you think that this practice is either the region that includes the centromere is inverted (its position
is reversed) in both gorillas and orangutans.
and grows laterally until it divides the cytoplasm in two (Fig-
a gene codes. As a result, third codon mutations are often se-
Orangutan 3. DNA extracted from bacteria cultured in 15N medium and after each generation in 14N medium. Extracted DNA
wise or unwise. wasTELOPHASE During
centrifuged in a special solutiontelophase,
to separate DNA the spindle
of different disassembles and
densities. ure 10.9). In both cases, the plane of cytoplasmic division is

5. The phenomenon in which chance events cause unpredictable the chromosomes at each spindle pole decondense, returning to
Results: Meselson and Stahl obtained the following results:
determined by the layer of microtubules that persist at the for-
lectively neutral (see Section 21.4), and they accumulate more
changes in allele frequencies is called: 2. Many human diseases are caused by recessive alleles that are not the extended state typical of interphase. As decondensation pro- mer spindle midpoint.

expressed in heterozygotes. Some people think that eugenics—

Conclusion: Differences in chromosome structure between humans and both gorillas and orangutans are more pronounced than they are between rapidly than do mutations in the first or second positions. Fi- 14N–14N
(light) DNA
a. gene flow. humans and chimpanzees. Structural differences in the chromosomes of these four species may contribute to their reproductive isolation. C h a p t e r 1 0 C e l l D i v i s i o n a nD M i t o s i s
nally, certain nucleotide substitutions are more common than
15N–14N
211

b. genetic drift. the selective breeding of humans

think like a scientist Do you think that the differences summaryto eliminate
in chromosome Based
undesirable
on this genetic
setandoftheir
five characters,
relatives amongthethe phylogenetic
15N–15N
hybrid DNA
15N–14N
hybrid DNA

others: transitions (the substitution of a purine for another pu-

structure between humans closest
traits—provides a way for us to ridwith ourfive populations
hypothesizedofevolutionary such
(heavy) DNA

c. inbreeding.
apes represent a prezygotic or postzygotic reproductive tree
isolating mechanism? changes (B) is more parsimo- DNA from DNA after one DNA after two

harmful alleles. Explain why eugenics nious thancannot the treeeliminate

that hypothesizes such ge- 10 evolutionary changes (C). In
15N
medium
rine or a pyrimidine for another pyrimidine) occur more fre-
replication in 14N replications in 14N

d. balanced polymorphism.
Source: J. J. Yunis and O. Prakash. 1982. The origin of man: A chromosomal pictorial legacy. Science 215:1525–1530.
The predicted DNA banding patterns for the three DNA replication models shown in Figure 14.8 were:
Conclusion:

netic traits from human populations. the absence of additional data, a systematist would accept the more quently than transversions (substitutions between purines and
e. stabilizing selection.
Experimental Research
© Cengage Learning 2014 N medium15 One replication in N Two replications in N 14 14

3. Using
different two
species.types of banding
In all species, beanspatterns
parsimonious tree as the best working hypothesis.
to represent
vary from one two have alleles at the same
a diploid complement gene whereas chim-
of 46 chromosomes,
(The characters
Semiconservative
pyrimidines).Matches
6. An Eastern European immigrant carrying the allele for Tay- chromosome segment to another. When researchers andfindclades
iden- of vascular
panzees, gorillas,plants are discussed
and orangutans in detailThein Chapter 28.)
have 48 chromosomes.
√ results
To avoid this problem, systematists develop statistical
Sachs disease settled in a small village on the St. Lawrence River. locus, design
tical banding patternsan exercisesegments
in chromosome to illustrate
from two orhow population
difference can be traced size
to the affects
fusion (that is, the joining to-
genetic
more related drift.
species, they know that they are examining com- gether) of two ancestral chromosomes into chromosome 2 of
Conservative
models of evolutionary change that take into account varia-
Many generations later, the frequency of the allele in that village
Does not
think
bandinglike a scientist Do the five characters in the table X match results
parable portions of the species’ genomes. Thus, the humans; the ancestral chromosomes are separate in the other
tions in the evolutionary rates at different nucleotide posi-
is statistically higher than it is in the immigrant’s homeland. The 4. Inpatterns
what ways
allow are
scientists to the
identifyeffects of sexual
specific chromosome
provide selection,
three species.
seg-sufficient disruptive
information selec-
to develop an hypothesis about the
ments and compare their positions in the chromosomes of dif- Moreover, banding patterns suggest that the position Dispersive
of tions or in different genes or species as well as changes in evo-
high frequency of the allele in the village probably provides an tion, and nonrandom mating different?
ferent species. evolutionarythe How are inthey
relationship
centromere similar?
of cycads
human chromosome and2 ginkgophytes?
closely matches that
Does not
X match results
lutionary rates over time. In one statistical approach, the
example of: The banding patterns of humans and their closest relatives of a centromere in one of the chimpanzee chromosomes, re-
among the apes—chimpanzees, gorillas, and orangutans—re- flecting their close evolutionary relationship. But this centro-
maximum likelihood method, systematists compare alterna-
The results support the semiconservative model.

a. natural selection. Design veal that an Experiment

whole sections of chromosomes have been©rearranged
Cengage Learning 2014falls within an inverted region of the chromosome in
mere
tivein trees with specific models about the rates of evolutionary
think like a scientist For the semiconservative replication model, what proportion of N– N, N– N, and 15 15 15 14
over evolutionary time (Figure 22.18). For example, humans gorillas and orangutans, reflecting their evolutionary diver-
b. the concept of relative fitness. 14
N– N 14
molecules would you expect after four and five replications N? 14

c. the Hardy–Weinberg genetic equilibrium. Design an experiment to test the hypothesis that the differences in change. The tree that is most likely to have produced the ob-
Source: M. Meselson and F. W. Stahl. 1958. The replication of DNA in Escherichia coli. Proceedings of the National Academy of Sciences USA 44:671–682.

C h a p t e r 2 2 S p e C i at i o n 495

d. phenotypic variation. size among adult guppies are determined by the amount of food they © Cengage Learning 2014
served distribution of molecular character states is identified
e. the founder effect. eat rather than by genetic factors. plausible explanation of any phenomenon is the best. If we as- as the best hypothesis.
7. If a storm kills many small sparrows in a population, but only a sume that any complex evolutionary change is an unlikely event, To illustrate how phylogenetic Apply Evolutionary trees are constructed from
Thinking

Interpret the Data Interpret the Data Bill depth

few medium-sized and large ones, which type of selection is then it is extremely unlikely that the same complex change DNA sequence data, we Captive tocite
a limitedansupplyexample using
of animals for a breeding the genetic
breeding programs for endangered species often have access
stock. As a result, their

probably operating? Peter and Rosemary Grant of Princeton evolved twice in one
University lineage.
have studied Thus, thewhen the principle is applied distance method, which logical calculates the overall organizationsproportion
offspring are at risk of being highly inbred. Why and how might zoo-
gardens and conservation avoid or minimize of
exercises, drawn from
inbreeding?
a. directional selection ecology and evolution of finches on the Galápagos Islands since the to phylogenetic analyses, it suggests that the “best” phylogenetic bases
1976 (before drought)
All 751 finches on the island that differ between two species. The genetic distance be-
tree is the onelarge that bills
hypothesizes
(as mea-the smallest number of evolu- tween closely related species is smaller than the genetic dis-
90
b. stabilizing selection early 1970s. They have shown that finches with
published biological
c. disruptive selection sured by bill depth; see Figure) cantionary eat both changes
small needed seeds and to account
large for the distribution of char- 60 tance between distantly related species, because the gene pools
Percentage of finches in each bill depth class

d. intersexual selection seeds, but finches with small bills can acter only states eatwithin
small aseeds. clade;In in 1977,
effect, athis approach minimizes30the of closely related species have accumulated distinctive muta-
research, help you build
e. intrasexual selection
8. Which of the following phenomena explains why the allele for
severe drought on the island of Daphne
tion by plants. After the birds consumed
number
similarwhatever
Major reduced seed
of homoplasies
traits) in small seeds
the tree
produc-
(that
(Figure
is, the independent evolution of
they24.12). To apply the principle, 0
tions for a shorter period of time. Systematists can construct a
phylogenetic tree from these data by making multiple compari-
found, only large seeds were still available. The resulting food shortage
your skills in analyzing
sickle-cell hemoglobin is common in some tropical and subtropi- computer programs evaluate the number of evolutionary sons of genetic distance between pairs of species and then be-
1978 (after drought)
The 90 surviving finches on the island

cal areas where the malaria parasite is prevalent? killed a majority of the medium ground changes fincheshypothesized(Geospiza byfortis) on
each phylogenetic tree they generate, 12
tween groups of species (Figure 24.13). Although not as powerful
a. balanced polymorphism Daphne Major; their population plummeted and the researcher from 751identifiesin 1976 the to just one with the fewest hypothe- as the maximum likelihood method, the genetic distance
figures and reading
b. heterozygote advantage 90 in 1978. The Grants’ research also
distributions of bill depths in the birds The
documented
sized changes as the most plausible.
fromprinciple
a
1976 to 1978,
change in the
as illus- also allows researchers 4 to
8

method does not depend on assumptions about the evolution-

c. sexual dimorphism of parsimony ary likelihood of different types of mutations. It also requires
graphs or tables.
d. neutral selection trated in the graphs to the right. Inidentify
the relationship between bill size and food size for these birds, inter-
light of what homologous you now know about
characters and infer their ancestral and 0
6 7
much8 9
less10 computing
11 12 13
power,
14
which is useful when comparing
e. stabilizing selection derived states. Once the most parsimonious phylogenetic tree billions of(mm)
Bill depth bases of homologous DNA sequences.
pret the change illustrated in the graph. What type of natural selection Grant, Peter R., Ecology and Evolution of Darwin’s Finches. © Princeton University Press. Reprinted by permission of

is identified, a researcher can visualize the distribution of de- Princeton University Press.

does this example illustrate? Source: P. R. Grant. 1986. Ecology and Evolution of Darwin’s Finches.
rived character states and pinpoint when each derivedPrinceton stateUniversity Press. Study Break 24.5
evolved. 1. How does outgroup comparison facilitate the identification of
ancestral and derived character states?
STATISTICAL APPROACHES When comparing two genome se-
2. What characteristics are used to group organisms in a cladistic
quences, each base in a strand of DNA can be treated as a analysis?
character with four possible states (A, G, T, and C). One could 3. How is the principle of parsimony applied in phylogenetic
a. Protein classes perform a parsimony analysis on molecular sequence > data, like analyses?
Think Outside the Book activities help Transfer/carrier protein (2%)
Structural protein (2%) Chaperone (1%)
thinkfor
the one illustrated
identify theEarlier
outside
phylogenetic tree we
in the chapter
the book
morphological
that
data in Figure 24.12, to
assumesthe
mentioned thefact
fewest muta-ani-
that cloned
tions. But the
malsapplication of the parsimony approach to mo- >
you think analytically and critically as you
Membrane traffic protein (3%)
Other (3%)
Protein of the may have many genes whose expression
immune system (1%) lecular data is complicated by several factors. First, given that
is abnormal
compared to gene expression in a noncloned animal. Individ- think outside the book
there are only four possible character
outlinestates at each
you position
would takeinex-
Unitexplore the biological world, either on ually or collaboratively, the steps access the web page for the tree of Life project at http://
478 Cytoskeletal
three evolU tiona ry B i o l o g y
protein (3%)
a nucleic acid, identicaltochanges
perimentally determine,in on
nucleotides oftenscale,
a genome-wide ariseif in-
genes www.tolweb.org/tree/. Select a group of animals or plants Chapter 21 MiCro
Enzyme (28%)
dependently.areSecond,
abnormallysegments
expressedof inDNA thatmammal.
a cloned do not code for
Your answer that is of interest to you, and study the structure of its phylo-
your own or as part of a team. Enzyme
modulator (7%) Unit three e v o lU t i o n a ry B i o l o g y
proteins areshould
less likely
includethan
how coding regionsreveals
the experiment to beboth
affected by
qualitative genetic tree. CHow many major clades does it include? On the479
h a p t e r 21 M iC ro e v o lUt i o n : g e n e t iC C h a n g e s w i t h i n p o pU l at i o n s

Signaling
natural selection. As a result,
and quantitative mutations
differences accumulate
in gene expression.faster in basis of what shared derived characters are those clades
molecule (7%) non-coding regions, causing them to evolve rapidly. Third, be- defined?
cause of the degeneracy of the genetic code (described in Sec-
Receptor (8%) Transcription

y B iology
factor (16%) 19.4 Genome Evolution
Chapter 24 Sy StematiCS and phylogen e t iC S: re v e a l i n g t h e t r e e o f li f e 543
Transporter (8%) DNA genomes with protein-coding genes are thought to have
Nucleic acid binding (11%)
evolved over 3.5 billion years ago, by the time of the earliest fos-
b. Biological classes sil microorganisms that have been discovered (see Chapter 25).
xiv Cell adhesion (3%) Those early cells probably had at most a few hundred protein-
Immune system Cell communication (11%) coding genes. New genes evolved as life evolved and became
process (7%) more complex, so that most present-day organisms have thou-
Cell cycle (5%) sands or tens of thousands of protein-coding genes. In this sec-
tion you will learn how genes and genomes have evolved, and
 VISUAL LEARNING
Spectacular illustrations—developed with great care—help you visualize
biological processes, relationships, and structures.

Closer look Figure 16.3 Regulation of the inducible lac operon by the Lac repressor in the
absence (A) and presence (B) of lactose.

a. Lactose absent from medium: structural genes not transcribed

lac operon

lacI Promoter Operator lacZ lacY lacA

DNA
Transcription
blocked 1 Active Lac repressor expressed
NEW!
1 3 from lacI gene binds to operator.

mRNA 2 RNA polymerase blocked

2
RNA polymerase
cannot bind
from binding to operator.

3 Transcription of structural genes does

not occur. (Repressor occasionally falls
“Closer Look” figures help you gain a
to promoter.
Lac repressor
(active)
off operator, allowing a very low rate of
transcription, resulting in a few molecules
of each protein being made.)
better understanding of a major concept
b. Lactose present in medium: structural genes transcribed
through a visual presentation, usually a
detailed, multistep diagram. The figures
lac operon

lacI Promoter Operator lacZ lacY lacA

end with a Summary and a Think Like a

DNA
Transcription 5
occurs.
RNA polymerase

Scientist question.
mRNA binds to promoter.
4 mRNA

Lac repressor 6 Translation 1 Permease molecules already present

(active) Inactive transport lactose into the cell.
3 repressor
Binding site 2 β–Galactosidase molecules already
for inducer Lactose present in the cell convert some of the
Allolactose catabolism lactose to the inducer allolactose.
(inducer) enzymes
3 Allolactose binds to the Lac repressor,
inactivating it by altering its shape so that
2 it cannot bind to the operator.

4 RNA polymerase binds

1 to the promoter.
Lactose 5 Transcription of the lac operon
structural genes occurs.
summary In the absence of lactose in the medium, there is no 6 Ribosomes recognize the ribosome
need for the lac operon genes to be active. In this case, the active Lac binding site upstream of each of the three
repressor binds to the operator and blocks RNA polymerase from coding sequences on the mRNA and
binding at the promoter. Therefore, transcription of the structural translation produces the three proteins.
genes does not occur. In the presence of lactose, the lac operon genes
are expressed so that catabolism of lactose can occur. In this case, the
think like a scientist If there was a mutation in the lacI gene Elongation is highly similar in prokaryotes and eukary- Researchers were surprised to discover that peptidyl transfer-
inducer allolactose binds to the Lac repressor inactivating it so that it
otes, with no significant conceptual differences. The elongation ase, the enzyme that forms peptide bonds in the elongation cycle,
does not bind to the operator. As a result, RNA polymerase binds to that results in a Lac repressor that could not bind to the operator,
cycle turns at the rate of about one to three times per second in is not a protein but a part of an rRNA of the large ribosomal sub-
the promoter and transcribes the structural genes. what effect would that have on the regulation of the lac operon? eukaryotes and 15 to 20 times per second in bacteria. Once it is unit. An RNA molecule that catalyzes a reaction like a protein
long enough, the growing polypeptide chain extends from the enzyme does is called a catalytic RNA or a ribozyme (ribonucleic
© Cengage Learning 2014 ribosome through the exit tunnel (see Figure 15.13) as elonga- acid enzyme). (Insights from the Molecular Revolution describes
tion continues. the experimental evidence showing that peptidyl transferase is a
ribozyme.)
duced to inactivate the repressor. The now-active repressor This quick turnover permits the cytoplasm to be cleared DNA

binds to the operator, blocking transcription of the struc- quickly of the mRNAs transcribed from an operon. The en- Figure 15.15 Translation elongation. A protein elongation factor (EF) complexes with
Transcription
tural genes. The controls are aided by the fact that bacterial coded proteins also have short lifetimes and are degraded Pre-mRNA the aminoacyl–tRNA to bring it to the ribosome, and another EF is needed for ribosome
translocation. For simplicity, the EFs are not shown in the figure.
mRNAs are very short-lived, about 3 minutes on the average. quickly. RNA processing © Cengage Learning 2014
mRNA

342 U ni t tW O Genetics Translation chapter 16 reGU lati On Of Gene e xpressi On

Illustrations of complex biological

Polypeptide
Ribosome with initiator Met
Met–tRNA bound to
Ribosome Peptidyl
the P site, and the
A site empty transferase

processes are annotated with numbered

Initiator
Empty tRNA tRNA
from E site
5' cap
4 When translocation is

step-by-step explanations that lead you

3' 1 An aminoacyl–tRNA
complete, the empty tRNA
E P A TP binds to the codon in the
in the E site is released. mRNA A site; TP is hydrolyzed
With the A site vacant and 1 2 3 4 5 DP
Peptidyl in this step.
the peptidyl–tRNA in the P +
Codons

through all the major points. Orientation

site, the ribosome repeats
P
the elongation cycle. In i
each cycle, the growing Peptidyl AA 2
polypeptide chain is Met transferase
transferred from the P site

diagrams are inset on figures and

tRNA to the amino acid on AA2 Met AA2
the A site tRNA.

Peptidyl–

help you identify the specific biological

tRNA Aminoacyl–
tRNA

3' 3'

process being depicted and where the

tion between their positive charges and the negatively charged DNA enters and leaves the core particle) and to the linker DNA. E P A E P A
phosphate groups of the DNA. This binding causes the nucleosomes to package into a coiled
Five types of histones exist in most eukaryotic cells: H1, structure 30 nm in diameter, called the 30-nm chromatin fiber. Exit 1 2 3 4 5 1 2 3 4 5

H2A, H2B, H3, and H4. The amino acid sequences of these One possible model for the 30-nm fiber is the solenoid model, Aminoacyl
3 The ribosome
proteins are highly similar among eukaryotes, suggesting that with the nucleosomes spiraling helically with about six nucleo-

process takes place.

translocates (moves) along
they perform the same functions in all eukaryotic organisms. somes per turn (see Figure 14.21).
the mRNA to the next
One function of histones is to pack DNA molecules into The arrangement of DNA in nucleosomes and the 30-nm
codon, using energy from Met 2 Peptidyl transferase, an enzyme
the narrow confines of the cell nucleus. For example, each fiber compacts the DNA and probably also protects it from TP hydrolysis. During GDP
human cell nucleus contains 2 meters of DNA. Combination chemical and mechanical damage. In the test tube, DNA translocation, the two + in the large ribosomal subunit,
AA2 cleaves the amino acid (here the
with the histones compacts this length so much that it fits into wound into nucleosomes and chromatin fibers is much moretRNAs remain bound to P
nuclei that are only about 10 µm in diameter. Another function resistant to attack by deoxyribonuclease (a DNA-digestingtheiren- respective codons, so i initiator methionine) from the tRNA
is the regulation of DNA activity. zyme) than when it is not bound to histone proteins. It is also this step positions the in the P site and forms a peptide
TP
less accessible to the proteins and enzymes required for gene bond between it and the amino acid
peptidyl–tRNA (the tRNA on the tRNA in the A site. When the
HISTONES AND DNA PACKING The histones pack DNA at sev- expression. Therefore, the association of the DNA with his- with the growing
eral levels of chromatin structure. In the most fundamental tones must loosen in order for a gene to become active (see reaction is complete, the
polypeptide) in the P site, polypeptide chain is attached to the
structure, called a nucleosome, two molecules each of H2A, Section 16.2). and generates a new vacant A site tRNA, and an “empty” tRNA
H2B, H3, and H4 combine to form a beadlike, eight-protein A site. The empty tRNA that
3' (a tRNA with no amino acid
nucleosome core particle around which DNA winds for almost PACKING OF EUKARYOTIC CHROMOSOMES AT STILL HIGHER was in the P site is now in
E P A attached) is in the P site.
two turns (Figure 14.21). A short segment of DNA, the linker, LEVELS: EUCHROMATIN AND HETEROCHROMATIN In inter- the E site.
extends between one nucleosome and the next. Under the elec- phase nuclei, chromatin fibers are loosely packed in some re- 1 2 3 4 5
tron microscope, this structure looks like beads on a string. The gions and densely packed in others. The loosely packed regions
diameter of the beads (the nucleosomes) gives this structure its are known as euchromatin (emu 5 true, regular, or typical), and
name—the 10-nm chromatin fiber (see Figure 14.21). the densely packed regions are called heterochromatin (hetero
Each nucleosome and linker includes about 200 base pairs 5 different). Chromatin fibers also fold and pack into the thick,
326 Unit t WO Genetics
of DNA. Nucleosomes compact DNA by a factor of about 7; rodlike chromosomes visible during mitosis and meiosis. Ex-
that is, a length of DNA becomes about 7 times shorter when it periments indicate that links formed between H1 histone mol-
is wrapped into nucleosomes. ecules contribute to the packing of chromatin fibers, both into
heterochromatin and into the chromosomes visible during nu-
Figure 9.3 the membranes Closer
tubular conducting cells. The organic productslook
HISTONES AND CHROMATIN FIBERS The fifth histone, H1,
of photosynthe-Figure 8.13 Oxidative phosphorylation: the mitochondrial electron transfer
clear division (see discussion in Section 10.2). However, the
and compartments of sis are distributed to all parts of the plant by other conductingsystem and chemiosmosis. brings about the next level of chromatin packing. One H1 mol- exact mechanism for the more complex folding and packing is
chloroplasts. cells (see Chapter 36). ecule binds both to the nucleosome (at the point where the not known.
javarman/Shutterstock.com

Macro-to-Micro views help

© Cengage Learning 2014
Oxidative phosphorylation involves the electron transfer system (steps 1–6), and chemiosmosis by ATP synthase
(steps 7–9). Blue arrows indicate electron flow; red arrows indicate H1 movement.
Study Break 9.1
1. What are the two stages of photosynthesis?
2. In which organelle does photosynthesis take place in 1 Complex I picks up high-energy 4 Complex IV accepts electrons 6 The resulting H+ gradient Histone tail
electrons from NADH and conducts from cytochrome c and delivers supplies the energy that drives Histone H1 binds to
plants? Where in that organelle
Glycolysis are theATP
two stages of them via two electron carriers, FMN them via electron carriers cyto- ATP synthesis by ATP synthase.

you visualize the levels of

Cutaway of a small One of the photosynthetic Inner nucleosomes and linker
photosynthesis carried out? mitochondrial (flavin mononucleotide) and an Fe/S chromes a and a3 to oxygen. Four Histone
section from the leaf cells, with green chloroplasts (iron–sulfur) protein, to ubiquinone. protons are added to a molecule 7 Because of the gradient, H+ DNA, causing nucleosomes
Pyruvate membrane flows across the inner membrane Solenoid
Leaf’s upper surface Photosynthetic cells oxidation of O2 as it accepts four electrons, to form coiled structure.
> 2 Complex II oxidizes FADH2 to FAD;
the two electrons released are
forming 2 H2O.
and into the matrix through a
channel in the ATP synthase.
think outside the
Citric book transferred to ubiquinone, and the two
Large central acid ATP 5 As electrons move through 8 The flow of H+ activates ATP
Scientists have been working cycle to develop an artifi- protons released go into the matrix. the electron transfer system, they
vacuole

organization of biological
Electrons that pass to ubiquinone by the synthase, making the headpiece
cial version of photosynthesis that can be used to release free energy. Part of the and stalk rotate.
complex II reaction bypass complex I of released energy is lost as heat, DNA Linker
produce liquid fuels from CO2 and H2O. Collabora-
Oxidative the electron transfer system. but some is used by the mito-
phosphorylation ATP 9 As a result of changes in
tively or individually, find an example of research 3 Complex III accepts electrons from chondrion to transport H+ across shape and position as it turns, the
on artificial photosynthesis and prepare an outline ubiquinone and transfers them through the inner mitochondrial headpiece catalyzes the synthesis Nucleosome: DNA
the electron carriers in the complex— membrane from the matrix to the of ATP from ADP and Pi. wound around a
of how the system works or is anticipated to work. cytochrome b, an Fe/S protein, and inter membrane compartment at
nucleosome core

structures and how systems

CO2 complexes I, III, and IV.
cytochrome c1—to cytochrome c, which particle consisting of
is free in the intermembrane space. 2 nm
Cytosol 2 molecules each of Chromosome
H2A, H2B, H3, H4 in metaphase
9.2 The Light-Dependent
Outer 10-nm chromatin fiber 30-nm chromatin fiber
mitochondrial
O2 Reactions of
membrane

function as a whole.
Stomata (through Nucleus
which O2 and CO2 Photosynthesis
Steve McKnight

H+
O.L. Miller, Jr.,

are exchanged with H+

In this section we discuss the light-dependent reac-
B. Hamkalo

the atmosphere)
tions (also referred to more simply as the light reac- H+ H+ H+
Figure 14.21 Levels of organization in
H+ H+
tions), in which light energy is convertedHto+
chemical H+ H+
eukaryotic chromatin and chromosomes.
© Cengage Learning 2014 Nucleosomes Linkers Chromatin fiber
energy. The light-dependent reactions involve two H+
Cutaway view of a chloroplast
main processes: (1) light absorption; and (2) synthesis
Intermembrane
compartment
Unit tWO Genetics of NADPH and ATP. We will describe each of these chapter 14 Dna strUct Ure, replicatiOn, an D OrGanizatiOn 305
Outer membrane
processes in turn. To keep the bigger picture in per- cyt b
cyt c
Inner membrane e– 4
spective, you may find it useful to refer periodically to e– 3 3 4
the summary of photosynthesis shown in Figure 9.2. e– 3 e–
cyt a e– cyt a3
Inner FMN e– 1 1 Ubiquinone Fe/S
3 8
mitochondrial Fe/S e– e– e–
(CoQ) Complex 4
Electrons in Pigment
membrane Molecules
e– Absorb Complex cyt c IV Stalk
Complex e– III
1
Light Energy in Photosynthesis I 5 H2O
1
Complex 2 H+
5 H +
The first process in photosynthesis is light absorption. II
5 H+
What is light? Visible light
NADHis a+ form
H+ of radiant energy. 2 H+ + 1/2 O2
H+ Head-
Thylakoids Stroma (space It makes up a small part of the electromagnetic
• light absorption by chlorophylls around thylakoids) piece
and carotenoids • light-independent
spectrum (Figure 9.4), which rangesNAD from
+
radio waves FADH2 FAD + 2 H+
• electron transfer reactions to gamma rays. The various forms of electromagnetic Low H+
Mitochondrial
• ATP synthesis by ATP synthase wavelength—the horizontal dis-
radiation differ in matrix 6
tance between the crests of successive waves. Radio 9
waves have wavelengths in the range of 10 meters to ADP + P ATP
i
hundreds of kilometers, and gamma rays have wave-
Electron transfer system Chemiosmosis
lengths in the range of one hundredth to one millionthElectrons flow through a series of proton (H+) ATP synthase catalyzes ATP
Granum
of a nanometer. The average wavelength of radiowaves pumps; the energy released builds an H+ gradient synthesis using energy from the H+
for an FM radio station, for example, is 3 m. across the inner mitochondrial membrane. gradient across the membrane.

The radiation humans detect as visible light has Oxidative phosphorylation

wavelengths between about 700 nm, seen as red light, Chapter 8 ha r v e s t i n g C h e m i C a l e n e r g y: C e l l u l a r r e s p i r at i o n

xv
Stromal Thylakoid Thylakoid and 400 nm, seen as blue light. We see the entire
lamella lumen membrane spectrum of wavelengths from 700 to 400 nm com-

ch a p t e r 9 phO t Os y n t h e s i s 185
Other documents randomly have
different content
 Agriculture - Complete Guide
First 2022 - School

Prepared by: Lecturer Williams

Date: July 28, 2025

Quiz 1: Literature review and discussion

Learning Objective 1: Critical analysis and evaluation
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Learning Objective 2: Historical development and evolution
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
[Figure 2: Diagram/Chart/Graph]
Learning Objective 3: Literature review and discussion
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Learning Objective 4: Literature review and discussion
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Learning Objective 5: Assessment criteria and rubrics
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 5: Diagram/Chart/Graph]
Important: Key terms and definitions
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 6: Theoretical framework and methodology
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 7: Diagram/Chart/Graph]
Practice Problem 7: Current trends and future directions
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 8: Diagram/Chart/Graph]
Example 8: Theoretical framework and methodology
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Practical applications and examples
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Appendix 2: Key terms and definitions
Practice Problem 10: Key terms and definitions
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Critical analysis and evaluation
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Remember: Comparative analysis and synthesis
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
[Figure 13: Diagram/Chart/Graph]
Definition: Key terms and definitions
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 14: Diagram/Chart/Graph]
Remember: Study tips and learning strategies
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 15: Diagram/Chart/Graph]
Important: Current trends and future directions
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 16: Diagram/Chart/Graph]
Important: Ethical considerations and implications
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Definition: Fundamental concepts and principles
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 18: Historical development and evolution
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Note: Assessment criteria and rubrics
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Lesson 3: Experimental procedures and results
Note: Problem-solving strategies and techniques
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Important: Current trends and future directions
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Ethical considerations and implications
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Key terms and definitions
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Critical analysis and evaluation
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 25: Key terms and definitions
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 26: Practical applications and examples
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Remember: Current trends and future directions
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Definition: Statistical analysis and interpretation
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Practice Problem 29: Key terms and definitions
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Part 4: Ethical considerations and implications
Example 30: Historical development and evolution
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Case studies and real-world applications
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Important: Current trends and future directions
• Learning outcomes and objectives
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Statistical analysis and interpretation
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Critical analysis and evaluation
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Example 35: Comparative analysis and synthesis
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Case studies and real-world applications
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Key terms and definitions
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 38: Study tips and learning strategies
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Learning outcomes and objectives
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Methodology 5: Ethical considerations and implications
Remember: Fundamental concepts and principles
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Definition: Current trends and future directions
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 42: Diagram/Chart/Graph]
Practice Problem 42: Literature review and discussion
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 43: Experimental procedures and results
• Experimental procedures and results
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Remember: Problem-solving strategies and techniques
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Example 45: Study tips and learning strategies
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Note: Problem-solving strategies and techniques
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Best practices and recommendations
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Note: Practical applications and examples
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 49: Literature review and discussion
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 50: Diagram/Chart/Graph]
Practice 6: Ethical considerations and implications
Note: Theoretical framework and methodology
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Key Concept: Historical development and evolution
• Learning outcomes and objectives
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Learning outcomes and objectives
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 53: Diagram/Chart/Graph]
Key Concept: Statistical analysis and interpretation
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Historical development and evolution
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 55: Diagram/Chart/Graph]
Note: Experimental procedures and results
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 56: Critical analysis and evaluation
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Note: Literature review and discussion
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Comparative analysis and synthesis
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 59: Ethical considerations and implications
• Assessment criteria and rubrics
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Background 7: Problem-solving strategies and techniques
Note: Learning outcomes and objectives
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Assessment criteria and rubrics
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Note: Ethical considerations and implications
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Definition: Practical applications and examples
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Historical development and evolution
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Practice Problem 65: Key terms and definitions
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Study tips and learning strategies
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Interdisciplinary approaches
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Current trends and future directions
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Practice Problem 69: Fundamental concepts and principles
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Section 8: Key terms and definitions
Note: Practical applications and examples
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Practice Problem 71: Fundamental concepts and principles
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Important: Experimental procedures and results
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Remember: Problem-solving strategies and techniques
• Historical development and evolution
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 74: Diagram/Chart/Graph]
Practice Problem 74: Interdisciplinary approaches
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Current trends and future directions
• Ethical considerations and implications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Note: Case studies and real-world applications
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Practice Problem 77: Assessment criteria and rubrics
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 78: Historical development and evolution
• Literature review and discussion
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 79: Diagram/Chart/Graph]
Important: Practical applications and examples
• Critical analysis and evaluation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Summary 9: Interdisciplinary approaches
Note: Critical analysis and evaluation
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Interdisciplinary approaches
• Learning outcomes and objectives
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Remember: Assessment criteria and rubrics
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Example 83: Problem-solving strategies and techniques
• Practical applications and examples
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Note: Research findings and conclusions
• Theoretical framework and methodology
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Comparative analysis and synthesis
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Key terms and definitions
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Note: Practical applications and examples
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Key Concept: Research findings and conclusions
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Statistical analysis and interpretation
• Best practices and recommendations
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Results 10: Ethical considerations and implications
Note: Literature review and discussion
• Case studies and real-world applications
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Literature review and discussion
• Key terms and definitions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Important: Study tips and learning strategies
• Problem-solving strategies and techniques
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Key Concept: Problem-solving strategies and techniques
• Current trends and future directions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Remember: Interdisciplinary approaches
• Statistical analysis and interpretation
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Definition: Study tips and learning strategies
• Comparative analysis and synthesis
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
[Figure 96: Diagram/Chart/Graph]
Important: Problem-solving strategies and techniques
• Study tips and learning strategies
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Formula: [Mathematical expression or equation]
Definition: Ethical considerations and implications
• Interdisciplinary approaches
- Sub-point: Additional details and explanations
- Example: Practical application scenario
Important: Learning outcomes and objectives
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
Remember: Case studies and real-world applications
• Research findings and conclusions
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Quiz 11: Key terms and definitions
Example 100: Statistical analysis and interpretation
• Fundamental concepts and principles
- Sub-point: Additional details and explanations
- Example: Practical application scenario
- Note: Important consideration
Formula: [Mathematical expression or equation]
[Figure 101: Diagram/Chart/Graph]
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