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147 views112 pages

(Ebook) Biology: The Dynamic Science, Vol. 1, 2Nd Edition by Peter J. Russell, Paul E. Hertz, Beverly Mcmillan Isbn 0538493720 PDF Download

Educational material: (Ebook) Biology: The Dynamic Science, Vol. 1, 2nd Edition by Peter J. Russell, Paul E. Hertz, Beverly McMillan ISBN 0538493720 Available Instantly. Comprehensive study guide with detailed analysis, academic insights, and professional content for educational purposes.

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Biology: The Dynamic Science, Second Edition, © 2011, 2008 Brooks/Cole, Cengage Learning
Volume 1
ALL RIGHTS RESERVED. No part of this work covered by the copyright herein
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Brief Contents

1 Introduction to Biological Concepts and Research 1

Unit One Molecules and Cells


2 Life, Chemistry, and Water 22
3 Biological Molecules: The Carbon Compounds of Life 42
4 Energy, Enzymes, and Biological Reactions 70
5 The Cell: An Overview 88
6 Membranes and Transport 116
7 Cell Communication 137
8 Harvesting Chemical Energy: Cellular Respiration 155
9 Photosynthesis 176
10 Cell Division and Mitosis 199

Unit Two Genetics


11 Meiosis: The Cellular Basis of Sexual Reproduction 219
12 Mendel, Genes, and Inheritance 234
13 Genes, Chromosomes, and Human Genetics 256
14 DNA Structure, Replication, and Organization 281
15 From DNA to Protein 305
16 Regulation of Gene Expression 333
17 Bacterial and Viral Genetics 362
18 DNA Technologies and Genomics 383

Appendix A: Answers A-1


Appendix B: Classification System A-13
Glossary G-1
Index I-1

41248_00_FM_V1_i-xxx.indd v 11/15/10 10:16 AM


About the Authors

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; he is currently a professor of biology. Peter teaches a section of the intro-
ductory 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 recog-
nition of his excellence in teaching. Since 1986, he has been the author of a successful genetics text-
book; current editions are iGenetics: A Molecular Approach, iGenetics: A Mendelian Approach, and
Essential iGenetics. Peter’s research is in the area of molecular genetics, with a specific interest in
characterizing the role of host genes in the replication of the RNA genome of a pathogenic plant
virus, and the expression of the genes of the virus; yeast is used as the model host. His research has
been funded by agencies including the National Institutes of Health, the National Science Founda-
tion, and the American Cancer Society. 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
involving undergraduates, including cofounding the biology division of the Council on Undergradu-
ate Research in 1985. He was Principal Investigator/Program Director of a National Science Founda-
tion (NSF) Award for the Integration of Research and Education to Reed College, 1998–2002.

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 Biology
faculty of the University of Puerto Rico at Rio Piedras. After spending two years as an Isaac Walton
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. He is also
the Program Director 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 commu-
nity college students that facilitates their transfer to four-year colleges and universities. He teaches one
Aaron W. Kinard

semester of the introductory sequence for Biology majors and preprofessional 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 pub-
lished his research in such prestigious journals as The American Naturalist, Ecology, Nature, Oecologia,
and Proceedings of the Royal Society. In 2010, he received funding from NSF for a project designed to
detect the effects of global climate warming on the biology of Anolis lizards in Puerto Rico.

Beverly McMillan has been a science writer for more than 25 years and is coauthor of a college
text in human biology, now in its eighth edition. She has worked extensively in educational and
commercial publishing, including eight years in editorial management positions in the college divi-
sions of Random House and McGraw-Hill. In a multifaceted freelance career, Bev also has written
or coauthored 10 trade books, as well as story panels for exhibitions at the Science Museum of
Virginia and the San Francisco Exploratorium. She has worked as a radio producer and speech-
writer for the University of California system and as a science writer and media relations advisor
for the Virginia Institute of Marine Science of the College of William and Mary. She holds under-
graduate and graduate degrees from the University of California, Berkeley.

vi

41248_00_FM_V1_i-xxx.indd vi 11/15/10 10:16 AM


Preface

W elcome to the second edition of Biology: The Dynamic Sci-


ence. The title of our book reflects the explosive growth in
knowledge of the living world over the past few decades—indeed,
research. We want to amaze students not only with what biologists
know about the living world, but also with how they know it and
what they still need to learn. In doing so, we can encourage some
even over the few years since the first edition appeared. Although students to accept the challenge and become biologists them-
the rapid pace of discovery makes biology the most exciting of all selves, posing and answering important new questions through
the natural sciences, it also makes it the most difficult to teach. their own innovative research. For students who pursue other ca-
How can college instructors—not to mention their students— reers, we hope that they will leave their introductory—and per-
absorb the ever-growing body of ideas and information? The task haps only—biology course armed with intellectual skills that will
is daunting, especially in introductory courses that provide a enable them to evaluate future discoveries with a critical eye.
broad overview of the discipline. In this book, we introduce students to a biologist’s “ways of
knowing.” Scientists constantly integrate new observations, hy-
potheses, experiments, and insights with existing knowledge
Building on a strong foundation . . .
and ideas. To help students engage the world as scientists do, we
As scientists and authors, we viewed the first edition of this book must not simply introduce them to the current state of knowl-
as an experiment: we presented a focused view of the essential edge. We must also foster an appreciation of the historical con-
knowledge that we thought students would need to begin their text within which those ideas developed, and identify the future
careers as biologists. Like all good experiments, our first edition directions that biological research is likely to take.
generated lots of data: the many instructors and students who To achieve these goals, our explanations are grounded in the
used the book offered positive feedback about elements that en- research that established the basic facts and principles of biology.
hanced students’ learning as well as valuable suggestions for pos- Thus, a substantial proportion of each chapter focuses on studies
sible modifications. We also received input from a small army of that defi ne the state of biological knowledge today. When de-
expert reviewers as well as media and art advisory boards. As a scribing research, we fi rst identify the hypothesis or question
result of these efforts, every chapter has been revised and updated, that inspired the work and then relate it to the broader topic
and some units have been reorganized. In addition, the second under discussion. Our research-oriented theme teaches stu-
edition includes many new or modified illustrations and photos, dents, through example, how to ask scientific questions and pose
and we have taken great pains to make the text and the art even hypotheses, two key elements of the “scientific process.”
more tightly integrated than they were in the first edition. Because advances in science occur against a background of
past research, we also give students a feeling for how biologists of
Emphasizing the big picture . . . the past formulated basic knowledge in the field. By fostering an
appreciation of such discoveries, given the information and theo-
In this textbook, we have applied our collective experience as col- ries available to scientists in their own time, we can help students
lege teachers, researchers, and writers to create a readable and understand the successes and limitations of what we consider
understandable introduction to our field of study. We provide cutting edge today. This historical perspective also encourages
straightforward explanations of fundamental concepts, presented students to view biology as a dynamic intellectual enterprise, not
from the evolutionary perspective that binds the biological sci- just a list of facts and generalities to be memorized.
ences together. Having watched our students struggle to navigate We have endeavored to make the science of biology come
the many arcane details of college-level introductory biology, we alive by describing how biologists formulate hypotheses and
constantly remind ourselves and each other to “include fewer evaluate them using hard-won data; how data sometimes tell
facts, provide better explanations, and maintain the narrative only part of a story; and how studies often end up posing more
flow,” thereby enabling students to see the big picture. Clarity of questions than they answer. Although students might prefer
presentation, thoughtful organization, a seamless flow of topics simply to learn the “right” answer to a question, they must be
within chapters, and spectacularly useful illustrations are central encouraged to embrace “the unknown,” those gaps in knowledge
to our approach. that create opportunities for further research. An appreciation of
what biologists don’t know will draw more students into the field.
Focusing on research to help students engage And by defining why scientists don’t understand interesting phe-
nomena, we encourage students to think critically about possible
the living world as scientists . . . solutions and to follow paths dictated by their own curiosity. We
A primary goal of this book is to sustain students’ curiosity about hope that this approach will encourage students to make biology
the living world instead of burying it under a mountain of discon- a part of their daily lives—to have informal discussions about
nected facts. We can help students develop the mental habits and new scientific discoveries, just as they do about politics, sports,
fascination of scientists by conveying our passion for biological or entertainment.

vii

41248_00_FM_V1_i-xxx.indd vii 11/15/10 10:16 AM


Presenting the story line of the research provides an exceptionally clear supplement to the narrative in a
style that is consistent throughout the book. Graphs and anatomi-
process . . .
cal drawings are annotated with interpretative explanations that
In preparing this book, we developed several special features to lead students, step by step, through the major points they convey.
help students broaden their understanding of the material pre- In preparing this edition, we undertook a rigorous review of
sented and of the research process itself. A Visual Tour of these all the art in the text. The publishing team has made an extraor-
features and more begins on page xiii. dinary effort to identify the key elements of effective illustra-
• The chapter openers, entitled Why It Matters. . . , tell the story of tions. In focus groups and surveys, instructors helped us identify
how a researcher arrived at a key insight or how biological re- the “Key Visual Learning Figures” covering concepts or pro-
search solved a major societal problem, explained a fundamen- cesses that demand premier visual learning support. Each of
tal process, or elucidated a phenomenon. These engaging, short these figures has been critiqued by our Art Advisory Board, re-
vignettes are designed to capture students’ imaginations and vised, and revised again to insure its usability and accuracy.
whet their appetites for the topic that the chapter addresses.
• To complement this historical or practical perspective, each chap-
ter closes with a brief essay, entitled Unanswered Questions, pre- Organizing chapters around important
pared by an expert in the field. These essays identify important concepts . . .
unresolved issues relating to the chapter topic and describe cut-
As authors and college teachers, we know how easily students can
ting-edge research that will advance our knowledge in the future.
get lost within a chapter that spans 15 or more pages. When stu-
• Each chapter also includes a short, boxed essay entitled Insights
from the Molecular Revolution, which describes how molecular
dents request advice about how to approach such a large task, we
tools allow scientists to answer questions that they could not usually suggest that, after reading each section, they pause and
have even posed 20 or 30 years ago. Each Insight focuses on a quiz themselves on the material they have just encountered. After
single study and includes sufficient detail for its content to stand completing all of the sections in a chapter, they should quiz them-
alone. selves again, even more rigorously, on the individual sections and,
• Most chapters are further supplemented with one or more short, most important, on how the concepts developed in different sec-
boxed essays entitled Focus on Research. Some of these essays tions fit together. To assist these efforts, we have adopted a struc-
describe seminal studies that provided a new perspective on an ture for each chapter that will help students review concepts as
important question. Others describe how basic research has they learn them.
solved everyday problems relating to health or the environment. • The organization within chapters presents material in digestible
Another set introduces model research organisms—such as chunks, building on students’ knowledge and understanding as
Escherichia coli, Drosophila, Arabidopsis, Caenorhabditis, and they acquire it. Each major section covers one broad topic. Each
Anolis—and explains why they have been selected as subjects for subsection, titled with a declarative sentence that summarizes the
in-depth analysis. main idea of its content, explores a narrower range of material.
• Whenever possible, we include the derivation of unfamiliar
Three types of specially designed Research Figures provide
terms so that students will see connections between words that
more detailed information about how biologists formulate and share etymological roots. Mastery of the technical language of
test specific hypotheses by gathering and interpreting data. In biology will allow students to discuss ideas and processes pre-
the second edition we have included one or more Research Fig- cisely. At the same time, we have minimized the use of unneces-
ures in practically every chapter (see the list on the endpapers at sary jargon as much as possible.
the back of the book). • Sets of embedded Study Break questions follow every major sec-
• Research Method figures provide examples of important tech- tion. These questions encourage students to pause at the end of
niques, such as gel electrophoresis, the use of radioisotopes, and a section and review what they have learned before going on to
cladistic analysis. Each Research Method figure leads a student the next topic within the chapter. Short answers to these ques-
through the technique’s purpose and protocol and describes tions appear in an appendix.
how scientists interpret the data it generates.
• Observational Research figures describe specific studies in which
biologists have tested hypotheses by comparing systems under Encouraging active learning, critical thinking,
varying natural circumstances. and self-assessment of learning outcomes . . .
• Experimental Research figures describe specific studies in which
The second edition of Biology: The Dynamic Science includes a new
researchers used both experimental and control treatments—
active learning feature, Think Outside the Book, which will help
either in the laboratory or in the field—to test hypotheses by
manipulating the system they studied.
students think analytically and critically about the material as they
are learning it. Think Outside the Book activities have been de-
Integrating spectacular visuals into signed to encourage students to explore the biological world di-
rectly or through high-quality electronic resources. Students can
the narrative . . . engage in these activities either individually or in small groups.
Today’s students are accustomed to receiving ideas and informa- Supplementary materials at the end of each chapter help
tion visually, making the illustrations and photographs in a text- students review the material they have learned, assess their un-
book more important than ever before. Our illustration program derstanding, and think analytically as they apply the principles

viii P R E FA C E

41248_00_FM_V1_i-xxx.indd viii 11/15/10 10:16 AM


developed in the chapter to novel situations. Many of the end-of- Helping students master key concepts
chapter questions also serve as good starting points for class dis-
throughout the course. . .
cussions or out-of-class assignments.
• Review Key Concepts references figures and tables in the chapter, As teachers, we know that student effort is an important determi-
providing a summary of important ideas developed in the chap- nant of student success. Unfortunately, many of us simply cannot
ter. These Reviews are much too short to serve as a substitute for spare the time to develop novel learning tools for every concept—
reading the chapter. Instead, students may use them as an out- or even every chapter—in a large and complex introductory text-
line of the material, filling in the details on their own. book. To help address this problem, we are pleased to offer Aplia
• Each chapter also closes with Test Your Knowledge, a set of for Biology, an automatically graded homework management
10 questions that focus on factual material. system tailored to this edition. For students, Aplia provides a
• Several open-ended Discuss the Concepts questions emphasize structure within which they can expand their efforts, master
key ideas, the interpretation of data, and practical applications key concepts throughout the course, and increase their success.
of the material. For faculty, Aplia can help us transform our teaching and raise
• To help students hone their critical thinking ability, another
our productivity by allowing us to require more—and more
question asks students to Design an Experiment to test hypoth-
consistent—effort from students without adding to our work
eses that relate to the chapter’s main topic.
load. By providing students with continuous exposure to key con-
• To help students develop analytical and quantitative skills, each
cepts and their applications throughout the course, Aplia allows
chapter also includes an Interpret the Data feature. The hypoth-
esis and methods of an experimental or observational study are us to do what we do best—respond to questions, lead discussions,
summarized, and some of its results are presented in either and challenge our students.
graphical or tabular format. Students are asked to interpret these We hope you agree that we have developed a clear, fresh, and
results in the context of the experimental design. well-integrated introduction to biology as it is understood by re-
• Apply Evolutionary Thinking asks students to interpret a relevant searchers today. Just as important, we hope that our efforts will
topic in relation to the principles of evolutionary biology. excite students about the research process and the new discover-
• The Express Your Opinion exercise allows students to weigh both ies it generates.
sides of an issue by reading pro/con articles, and then making
their opinion known through an online voting process.

P R E FA C E ix

41248_00_FM_V1_i-xxx.indd ix 11/15/10 10:16 AM


New to This Edition

T his section highlights the changes we made to enhance the


effectiveness of the text. Every chapter has been updated to
ensure currency of information. We made organizational changes
important unresolved issues relating to the chapter topic. The es-
says now conclude with a Think Critically question, which en-
courages students to think about the unanswered question, pon-
to more closely link related topics and reflect preferred teaching der possible next steps in the research, or consider the benefits of
sequences. New features in the text have been developed to help answering the question.
students actively engage in their study of biology. Enriched media The Research Figures in the text provide more detailed infor-
offerings provide students a broad spectrum of learning mation about how biologists formulate and test specific hypoth-
opportunities. eses by gathering and interpreting data. We have developed
26 new Research Figures. In addition, references to the original
papers have been added to each figure so that the research can be
Organizational Changes explored in more depth, as needed.
The genetic and molecular regulation of development has been The second edition also includes new features that are de-
integrated into Chapter 16 (Regulation of Gene Expression), signed to encourage students to engage with the material and
which introduces students to the mechanisms that produce a develop the quantitative skills necessary for biological study and
complex multicellular organism from a single cell. The new sec- investigation. A feature called Think Outside the Book helps stu-
tion expands their understanding of the scope of gene regulation dents think critically about the material as they are learning it.
by showing how sequential regulatory events such as induction, The questions presented in these boxes are related to topics that
determination, and differentiation drive the formation of a com- the student has just studied and are designed to help explore the
plex organism, and it prepares them for the application of these biological world directly or through high-quality electronic re-
principles more specifically to plants and animals in Chapters 34 sources. In addition, new Interpret the Data exercises are in-
and 48, respectively. cluded in the study material at the end of each chapter. These
Coverage of viruses is now complete in Chapter 17 (Bacterial exercises, most of which are drawn from published biological re-
and Viral Genetics); brief coverage of viroids and prions is also search, help students build their skills in analyzing figures and in
included in that chapter. Placing the entire coverage of viruses in reading graphs and tables.
one chapter makes it easier for instructors to tailor the treatment
of viruses in their course.
The Ecology Unit has been reorganized to begin with a
Enhanced Art Program
large-scale overview of the science of ecology and the biosphere Helping today’s students understand biological processes requires
(combining part of the first edition’s Chapter 49 with the first effective visual learning support. In preparation for this edition
edition’s Chapter 52). This reorganized Chapter 49 lays out a con- we undertook a rigorous review of all the art in the text. Focus
ceptual framework for the scope of ecology and describes the groups and instructor surveys helped identify key figures that are
patterns of distribution of life on Earth. After this orientation, essential teaching and learning tools. The figures identified in this
the chapters continue in order with Chapter 50 focusing on pop- process were scrutinized by our Art Advisory Board and other
ulation ecology, Chapter 51 treating population interactions and content area experts to insure their effectiveness in the classroom.
community ecology, and Chapter 52 covering ecosystems. In some cases we developed new views of structures to facilitate a
three-dimensional visualization. We also included additional
“orientation” diagrams to help students visualize levels of organi-
New Features zation and how systems function as a whole.
We carefully considered all of the existing features and made sev- The effective integration of text and illustration has also
eral improvements to enhance their effectiveness as learning and been a top priority for the development of the art. We have in-
teaching tools. The Insights from the Molecular Revolution, which creased the number of illustrations supported by numbered
showcase exciting and interesting research based on molecular step-by-step annotations placed directly on the figure. These an-
techniques, have been rewritten to improve readability and ease notations help students interpret detailed illustrations and de-
of understanding. The topics of these features have been updated velop deeper understanding of complex processes.
to reflect recent discoveries, and references to the original papers
have been included. In addition, more than 40 illustrations have
been developed to add clarity and stimulate student interest in
Enriched Media
this feature. New to the second edition is Aplia for Biology, an automatically
The Unanswered Questions essays appearing at the end of graded homework management system tailored to this edition.
every chapter, each prepared by an expert in the field, identify Aplia courses are customized to fit with each instructor’s syllabus

41248_00_FM_V1_i-xxx.indd x 11/15/10 10:16 AM


and provide automatically graded homework with detailed, im- the animations ensure that students have the necessary founda-
mediate feedback on every question. Aplia’s interactive tools serve tion for understanding these important concepts.
to increase student engagement and understanding. Also new to the second edition are clips from the BBC Mo-
New, interactive 3-D animations have been developed that tion Gallery. Th is diverse and robust library of high-quality
help students visualize processes in a more dynamic way. These videos features clips from well-respected scientists and natural-
animations promote in-depth understanding of key biological ists, including Sir David Attenborough. The clips can be used in
topics, including cellular respiration, photosynthesis, DNA rep- conjunction with the text to spark discussion and help students
lication, and evolutionary processes. Embedded assessments in connect the material to their lives outside of the classroom.

We now invite you and your students to preview the


many exciting features that will help them think and
engage like scientists . . .

NEW TO THIS EDITION xi

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This page intentionally left blank
THE BIG PICTURE
Each chapter is carefully organized and presented in “digestible chunks” so you
can stay focused on the most important concepts. Easy-to-use learning tools
point out the topics covered in each chapter, show why they are important,
and help you learn the material.

8
study outline ❮ Study Outline The Study Outline
provides an overview of all the topics
8.1 Overview of Cellular
and key concepts in the chapter. Each

Dr. Donald Fawcett/Visuals Unlimited, Inc.


Energy Metabolism
8.2 Glycolysis: Splitting the
Sugar in Half
section breaks the material into a
8.3 Pyruvate Oxidation and manageable amount of information,
the Citric Acid Cycle
8.4 Oxidative
building on knowledge and under-
Mitochondrion (colorized TEM). Mitochondria are the sites of cellular respiration.
Phosphorylation:
The Electron
standing as you acquire it.
Transfer System and
Chemiosmosis
8.5 Fermentation

Harvesting Chemical Energy:


Cellular Respiration
Why It Matters. . . In the early 1960s, Swedish physician Rolf Luft mulled over
❮ Why It Matters Engaging
some odd symptoms of a patient. The young woman felt weak and too hot all the time (with a body
temperature of up to 38.4°C). Even on the coldest winter days she never stopped perspiring, and her introductory sections capture the
skin was always flushed. She was also underweight (40 kg), despite consuming about 3,500 calories
per day. excitement of biology and help you
Luft inferred that his patient’s symptoms pointed to a metabolic disorder. Her cells were
very active, but much of their activity was being dissipated as metabolic heat. He decided to
understand why the topic is important
order tests to measure her metabolic rate, the amount of energy her body was expending. The and how the material you are about to
results showed the patient’s oxygen consumption was the highest ever recorded—about twice
the normal rate! read fits into the Big Picture.
Luft also examined a tissue sample from the patient’s skeletal muscles. Using a microscope,
he found that her muscle cells contained many more mitochondria—the ATP-producing organ-
elles of the cell—than are normally present in muscle cells. In addition, her mitochondria were
abnormally shaped and their interior was packed to an abnormal degree with cristae, the infold-
ings of the inner mitochondrial membrane (see Section 5.3). Other studies showed that the mito-
chondria were engaged in cellular respiration—their prime function—but little ATP was being
generated.
The disorder, now called Luft syndrome, was the first disorder to be linked directly to a defective
mitochondrion. By analogy, someone with this disorder functions like a city with half of its power
plants shut down. Skeletal and heart muscles, the brain, and other hardworking body parts with
high energy demands are hurt the most by the inability of mitochondria to provide enough energy
for metabolic demands. More than 100 mitochondrial disorders are now known.

155

❯ Study Break Encourages you to pause and think STUDY BREAK 8.2 <
about the key concepts you have just encountered 1. What are the energy-requiring and energy-releasing steps of
glycolysis?
before moving to the next section. 2. What is the redox reaction in glycolysis?
3. How is ATP synthesized in glycolysis?
4. Why is phosphofructokinase a target for inhibition by ATP?

xiii

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THINK LIKE A SCIENTIST
Your study of biology focuses not only on what scientists now know about the
living world, but also how they know it. Use these unique features to learn through
example how scientists ask scientific questions, pose hypotheses, and test them.

❮ Insights from the Molecular Revolution Showcases


INSIGHTS FROM THE Molecular Revolution

Hot Potatoes: Do Plants Use 3. Laloi and her coworkers then tested stressed by low temperatures. Thus, mecha-
exciting and interesting research relying on molecular techniques;
Uncoupling Proteins to whether exposing potato plants to cold nisms for warming body tissues once thought
Generate Heat?
Mammals use several biochemical and
temperatures could induce greater synthe-
sis of the UCP mRNA (Figure 2). The cold
treatment resulted in an increase in UCP
to be the province only of animals appear to
be much more widespread. In particular,
UCPs, which were believed to have evolved in
these molecular tools allow researchers to answer questions that
molecular processes to maintain body heat.
One process is shivering; the muscular
activity of shivering releases heat that helps
mRNA level in the leaves.

Conclusion
The research indicates that potato plants
relatively recent evolutionary times with the
appearance of birds and mammals, may be a
much more ancient development.
they could not even pose 20 or 30 years ago.
keep body temperature at normal levels.
Another mechanism operates through un- probably use the mitochondrial uncoupling Source: M. Laloi et al. 1997. A plant cold-induced
coupling proteins (UCPs), which eliminate process to warm tissues when they are uncoupling protein. Nature 389:135–136.
the mitochondrial H⫹ gradient by making the
inner mitochondrial membrane leaky to
Flowers—very high
protons. Electron transfer and the oxidative
UCP mRNA level
reactions then run at high rates in mitochon-

❮ Unanswered Questions Top


dria without trapping energy in ATP. The
energy is released as heat that helps main-
tain body temperature. UNANSWERED QUESTIONS
ºC
Research Question
Do plants use UCPs to generate heat?
Experiments
40
30
20
Stem—moderate
UCP mRNA level
Do asexual organisms form species? researchers have identified one to three
As you learned in this chapter, the biological species concept applies only
Research by Maryse Laloi and her colleagues
at the Max Planck Institute for Molecular Plant
Physiology in Germany shows that some
10
0
to sexually reproducing organisms because only those organisms can important unresolved questions that
tissues in plants may use the same process Leaf—low UCP evolve barriers to gene flow (asexual organisms reproduce more or less
involving UCPs to generate heat. Their evi-
dence is as follows:
mRNA level
clonally). Nevertheless, research is starting to show that organisms whose relate to the chapter topic and talk about
1. Potato plants (Solanum tuberosum) have reproduction is almost entirely asexual, such as bacteria, seem to form
a gene with a DNA sequence similar to
that of a mammalian UCP gene. The
potato gene protein is clearly related to
Tuber—low UCP
mRNA level distinct and discrete clusters in nature. (These clusters could be consid-
ered “species.”) That is, bacteria and other asexual forms may be as dis-
what it would mean to the field if the
questions were answered.
the mammalian protein, and also has the Roots—very high
same overall three-dimensional structure. FIGURE 1
UCP mRNA level tinct as the species of birds described by Ernst Mayr in New Guinea.
2. The researchers used the DNA of the UCP mRNA levels of activity. Workers are now studying the many species of bacteria in nature (only a
potato UCP gene to probe for the presence
of messenger RNA (mRNA), the gene small number of which have been discovered) to see if they indeed fall
product that specifies the amino acid
20ºC After 1–3 days at 4ºC
into distinct groups. If they do, g then scientists will need a special theory,
sequence of proteins in the cytoplasm,
with the results shown in Figure 1. These
results indicate that the potato UCP gene
is active at different levels in various plant
Low UCP
mRNA level
Very high UCP
mRNA level
independent of reproductive
tists are now working
oductive isolation, to explain this distinctness. Scien-
ng onThink Critically
theories of whether the existence of discrete Questions at the end of each
ecological niches in nature Professor Coyne suggested
might explain the possiblethatdiscreteness
biologists will
of need to devise a special
tissues, suggesting that certain tissues
naturally need warming for optimal
function.
FIGURE 2
Results of exposure
to cold temperature.
asexual “species.” theory—or an additional type of species definition—to explain the dis-
tinctness of clusters of asexual organisms that are observed in nature. Do
essay help you think critically
How often does speciation
eciiatio
sympatrically or parapatrically?
rap
n occur
oyou
patrrlogical
think allopatrically versus
that this new explanation would be more similar to the morpho-
ically? species concept or the phylogenetic species concept as described
about the unanswered question,
Scientists do not know ow how in often speciation occurs between populations
this chapter?
solated geographically (allopatric speciation), com-
that are completely isolated ponder possible next steps in the
pared to how often itt occurs in populationsJerry thatCoyne conducts
exchange research
genes on speciation and teaches at the
(para-
peciation). The relative University
patric or sympatric speciation). frequencyofof
Chicago.
these To learn more
modes of about his research go to
http://pondside.uchicago.edu/ecol-evol/faculty/coyne_j.html.
research, or consider the benefits
speciation in nature is an active area of research. The ongoing work in-
cludes studies on smallmall isolated islands: if an invading species divides of answering the question.
Jerry Coyne

into two or more speciescies in this situation, it probably did so sympatrically


or parapatrically, sincece geographical isolation of populations in small is-
lands is unlikely. In addition,
ddition, biologists are reconstructing the evolutionary
41248_08_0155-0175.indd 171 7/9/10 1:39:48 PM
history of speciation using molecular tools and correlating this history
with the species’ geographical distributions. If this line of research were
to show, for example, that the most closely related pairs of species always
had geographically isolated distributions, it would imply that speciation
was usually allopatric. These lines of research should eventually answer C

❯ Focus on Research Special boxes in most chapters present research topics in more depth.

FOCUS ON Basic Research


Focus on Basic Research describes
Two-Dimensional Paper cells to the labeled carbon dioxide. Then, at vari- to the label, radioactivity showed up in G3P and
seminal research that provided insight
Chromatography and ous times, cells were removed and placed in hot in more complex substances including a variety
the Calvin Cycle
The first significant progress in unraveling the
alcohol, which instantly stopped all the photo-
synthetic reactions of the algae. Radioactive car-
bohydrates were then extracted from the cells
of six-carbon sugars, sucrose, and starch. The re-
searchers also examined the effect of reducing
the amount of CO2 available to the Chlorella cells
into an important problem.
light-independent reactions was made in the and two-dimensional paper chromatography so that photosynthesis worked slowly even in
1940s, when newly developed radioactive com- was done to separate and to identify them chem- bright light. Under these conditions, RuBP
pounds became available to biochemists. One ically, as shown in the figure. accumulated in the cells, suggesting that it is
substance, CO2 labeled with the radioactive car- By analyzing the labeled molecules revealed the first substance to react with CO2 in the light-
bon isotope 14C (discussed in Focus on Applied by the two-dimensional chromatography tech- independent reactions and that it accumulates if
Research in Chapter 2), was critical to this nique in the extracts prepared from Chlorella CO2 is in short supply. Most of the intermediate
research.
Beginning in 1945, Melvin Calvin, Andrew A.
Benson, and their colleagues at the University
cells under different conditions, Calvin and his
colleagues were able to reconstruct the reactions
of the Calvin cycle. For example, in carbohydrate
compounds between CO2 and six-carbon sugars
were identified in similar experiments.
Using this information, Calvin and his col-
FOCUS ON Applied Research
Focus on Applied Research describes
of California at Berkeley, combined 14C-labeled extracts made within a few seconds after the cells leagues pieced together the light-independent re-
CO2 with a widely used technique called two-
dimensional paper chromatography to trace the
pathways of the light-independent reactions in a
were exposed to the labeled CO2, most of the ra-
dioactivity was found in 3PGA, indicating that it
actions of photosynthesis and showed that they
formed a continuous cycle. Melvin Calvin was Using Radioisotopes to Trace tope 14C. Then they extracted various substances doses can kill cells (radiation generates highly re-
how scientific research has solved
is one of the earliest products of photosynthesis. awarded a Nobel Prize in 1961 for his work on Reactions and Save Lives from the cells at intervals, separated them on a active chemical groups that break and disrupt bi-
green alga of the genus Chlorella. The research-
ers exposed actively photosynthesizing Chlorella
In extracts made after longer periods of exposure the assimilation of carbon dioxide in plants.
In 1896, the French physicist Henri Becquerel
wrapped a rock containing uranium in paper
piece of paper based on their different solubili-
ties in particular solvents, and placed the paper
on a photographic film. The particular sub-
ological molecules). Dangerously overactive thy-
roid glands are treated by giving patients a dose
of radioactive iodine calculated to destroy just
everyday problems.
and tucked it into a desk drawer on top of a
stances that exposed spots on the film because enough thyroid cells to reduce activity of the
case containing an unexposed photographic
they were radioactive, as well as their order of gland to normal levels. In radiation therapy, can-
plate. When he opened the case containing the
appearance in the cells, allowed the researchers cer cells are killed by bombarding them with radi-
plate a few days later, he was surprised to find
to piece together the sequence of reactions in ation emitted by radium-226 or cobalt-60. As
an image of the rock on the plate—apparently
drop of
photosynthesis, as described and illustrated in much as is possible, the radiation is focused on
Turn paper 90° caused by energy emitted from the rock. One of
extract Focus on Basic Research in Chapter 9. the tumor to avoid destroying nearby healthy tis-
his coworkers, Marie Curie, named the phe-
Radioisotopes are widely used in medicine to sues. In some forms of chemotherapy for cancer,
nomenon “radioactivity.” Although radioactivity
diagnose and cure disease, to produce images of patients are given radioactive substances at lev-
can be dangerous to life (more than one re-
Solvent 1 Solvent 1 Solvent 2 diseased body organs, and, as in biological re- els that kill cancer cells without also killing the
searcher, including Marie Curie, has died from
search, to trace the locations and routes followed patient.
its effects), it has been harnessed and put to
by individual substances marked for identifica-
highly productive use for scientific and medical
tion by radioactivity. One example of their use is
purposes.
1 A drop of extract containing 2 The extracted molecules in the 3 The paper is dried, turned 90°, and touched to a 4 The paper is dried and in the diagnosis of thyroid gland disease. The
radioactive carbohydrates is spot dissolve and are carried second solvent (Calvin used a water solution of phenol covered with a sheet of The radiation released by unstable isotopes
thyroid is the only structure in the body that ab-
placed at one corner of a piece upward through the paper as the for this part of the experiment). As this solvent moves photographic film. Radioactive can be detected by placing a photographic film
sorbs iodine in quantity. The size and shape of
of chromatography paper. The solvent rises. The rates of through the paper, the molecules again migrate upward molecules expose the film in over samples containing the isotopes (as Bec-
paper is placed in a jar with its movement of the molecules vary from the spots produced by the first dimension, but at spots over their locations in the thyroid, which reflect its health, are measured
querel discovered) or by using an instrument
edge touching a solvent. (Calvin according to their molecule size rates different from their mobility in the first solvent. the paper. Developing the film by injecting a small amount of a radioactive io-
used a water solution of butyl and solubility. The resulting vertical This step, the second dimension of the two-dimensional reveals the locations of the known as a scintillation counter. These tech-
dine isotope into the patient’s bloodstream. After
alcohol and propionic acid.) line of spots is the first dimension technique, separates molecules that, although different, radioactive spots. The spots niques allow researchers to use isotopes as trac-
of the two-dimensional technique. had produced a single spot in the first solvent because on the film are compared with
the isotope is concentrated in the thyroid, the
ers in chemical reactions. Typically, organisms
they had migrated at the same rate. The individual the spots on the paper to gland is then scanned by an apparatus that uses
Photo by Gary Head

spots
p are identified byy comparing
p g their locations with identifyy the molecules that
are exposed to a reactant chemical that has been
the radioactivity to produce an imag image of the
the positions of spots made by known molecules when were radioactive. “labeled” with a radioactive isotope such as 14C
gland on a photo
photographic
graphic film Examples
lm. Exa m of what
the “knowns” are run through the same procedure. or 3H. After being exposed to the tracer, the
presented
the scans may show are presente nted in the figure.
chemical products in which the isotope appears,
Another application uses the factt tha that radioactive
and their sequence of appearance, can be de-
regions
thallium is not taken up by region ns oof the heart
FOCUS ON Model Research Organisms tected by their radioactivity and identifi fied.
identified.
muscle with poor circulation to dete detect coronary
For example, algae and plants use carbon di-
artery disease. Other isotopes aree us used to detect
Normal Enlarged Cancerous
The Yeast Saccharomyces oxide (CO2cul-
The cells growing in Saccharomyces ) as a raw material
mammals.in photosynthesis.
Genetic studies with Saccharomyces
bone injuries and defects, including injured, ar-
To trace Ifthe reactions ofwere
photosynthesis,
also the firstMelvin Scans of human thyroid glands after iodine-123
cerevisiae tures in the research lab are haploid. the to show the thritic,
genes orcarried in
abnormally growing segsegments
gme of bone.
was injected into the bloodstream. The radioactive
culture conditions are kept at Calvin
optimaland his coworkers
levels thegrew
DNAalgal cells in a
of mitochondria and their patternsof
Treatment ofdisease with radiois
radioisotopes takes
Saccharomyces cerevisiae, commonly known as iodine becomes concentrated in the thyroid gland.
(which requires only a source medium with CO2 that inheritance.
of a fermentable contained the radioiso-
The complete DNA sequenceofof
advantage radioactivity
the fact that radioac ctivi in large
baker’s yeast or brewer’s yeast, was probably
sugar such as glucose, a nitrogen source, and S. cerevisiae, which includes more than 12 mil-
the first microorganism to have been grown
minerals), the cells reproduce asexually by lion base pairs that encode about 6,000 genes,
and kept in cultures—a beer-brewing vessel is
budding. Saccharomyces has two mating was the first eukaryotic genome to be obtained.
basically a Saccharomyces culture. Favorite
types. If two yeast cells of different mating Analysis of the genome sequence revealed that
strains of baker’s and brewer’s yeast have been
types contact one another, they fuse—mate— yeast has many genes related to those in ani-
kept in continuous cultures for centuries. The
producing a diploid cell. Diploid cells can also mals, including mammals, making this rela-
yeast has also been widely used in scientific re-
reproduce asexually by budding. Under cer- tively simple microorganism an excellent sub-
search; its microscopic size and relatively
tain conditions, diploid yeast cells undergo ject for research that can be applied to the
short generation time make it easy and inex-
meiosis, producing haploid spores which are more complex animals of interest.
pensive to culture in large numbers in the
laboratory.
resistant to harsh conditions in the environ-
ment. Yeast spores germinate into haploid
cells, which then reproduce asexually.
The genes of yeast can also be manipu-
lated easily using genetic engineering tech-
niques. This has made it possible for re-
Focus on Model Research Organisms explains
Genetic studies with Saccharomyces led to
the discovery of some of the genes that control
searchers to alter essentially any of the yeast
genes experimentally to test their functions why researchers use certain organisms as
Adrian Warren/Ardea/London

the eukaryotic cell cycle. Many of these genes, and to introduce genes or DNA samples from
after their first discovery in yeast cells, were other organisms for testing or cloning. Sac-
found to have counterparts in animals and
plants. Defective versions of the genes often
charomyces has been so important to genetic
studies in eukaryotes that it is often called the
research subjects.
contribute to the development of cancer in eukaryotic E. coli.

xiv

41248_00_FM_V1_i-xxx.indd xiv 11/15/10 10:17 AM


ENGAGE LIKE A SCIENTIST
Be Active. Engage yourself in the process of learning and doing biology.

❯ Research Figures Specially designed figures provide information


FIGURE 1.14 Research Method
about how biologists formulate and test specific hypotheses by gathering The Scientific Method Purpose: The scientific method is a method of inquiry that allows researchers to crystallize
their thoughts about a topic and devise a formal way to test their ideas by making observa-
and interpreting data. tions and collecting measurable data.

Protocol:

1. Make detailed observations about a phenomenon of interest.

Observations

Research Method Inductive


reasoning
2. Use inductive reasoning to create a testable hypothesis that provides a working
explanation of the observations. Hypotheses may be expressed in words or in
mathematical equations. Many scientists also formulate alternative hypotheses
Hypothesis (that is, alternative explanations) at the same time.

Deductive
FIGURE 1.16 Observational Research reasoning
3. Use deductive reasoning to make predictions about what you would observe if
the hypothesis were applied to a novel situation.
A Field Study Using a Null Hypothesis Hypothesis: Anolis cristatellus, the crested anole, uses patches of sun and shade to regulate
its body temperature. Predictions

Null Hypothesis: Because this lizard does not regulate its body temperature, individuals se-
Observational Research lect perching sites at random with respect to environmental factors that influence body
temperature.

Anolis cristatellus Method: The researchers created a set of hollow, copper lizard models, each equipped with 4. Design and conduct a controlled experiment (or new observational study) to
a temperature-sensing wire. At study sites where the lizards live in Puerto Rico, the research- test the predictions of the hypothesis. The experiment must be clearly defined
ers hung 60 models at random positions in trees. They observed how often live lizards and so that it can be repeated in future studies. It must also lead to the collection of
the randomly positioned copper models were perched in patches of sun or shade, and they Experiments measurable data that other researchers can evaluate and reproduce if they
measured the temperatures of live lizards and the copper models. Data from the randomly choose to repeat the experiment themselves.
positioned copper models define the predictions of the null hypothesis.

Results: The researchers compared the frequency with which live lizards and the copper
models perched in sun or shade as well as the temperatures of live lizards andd thee copper
c
models. The data revealed that the behavior and temperatures of A. cristatelluss we
ere differ-
were IInterpreting
Interpret
t t ing
i the
th l t Compare the results of the experiment or new observations with those predicted by the hypothesis. Scientists
h Results:
R esults: Sc often use
origginal
ent from those of the randomly positioned models, therefore confirming the original formal statistical tests to determine whether the results match the predictions of the hypothesis.
FIGURE 8.15 Experimental Research
Alejandro Sanchez

hypothesis.

The Racker and Stoeckenius Experiment Question: Does an H⫹ gradient power ATP synthase-catalyzed ATP synthesis, thereby sup-

Demonstrating That an Hⴙ Gradient Powers porting Mitchell’s chemiosmotic hypothesis?

Copper Anolis model Percentage of models and Temperatures of ATP Synthesis by ATP Synthase Experiment: Efraim Racker of Cornell University and Walther Stoeckenius of University of
lizards perched in sun or shade models and lizards California, San Francisco, made membrane vesicles that had a proton pump and ATP syn-
Kevin de Queiroz, National Museum of Natural History, Smithsonian

Anolis cristatellus Anolis cristatellus thase to determine whether proton-motive force drives ATP synthesis.
100 30 1. The researchers constructed synthetic
Percentage in sun or shade

Percentage of observations

20 phospholipid membrane vesicles con-


Purple Light
75 In the habitat where Lizards Body tem
temperatures
mperatures taining a segment of purple surface H+
A. cristatellus lives, 10 of A. cristatellus
cristtatellus were membrane Bacteriorhodopsin
membrane from an archaean. The
nearly all models significantly
fican ntly higher segment
50 perched in shade, 30 than thos
thosese of the purple membrane contained only
but most lizards omlyy placed bacteriorhodopsin, a protein that
randomly
perched in sun.
20
models.
els. resembles rhodopsin, the visual H+
25 Models H+
10 H+ H+
pigment of animals (see Chapter Synthetic
KEY H+ H+ +
39). Bacteriorhodopsin is a light-acti- vesicle H
H+
Models Lizards 20 30 40
Perched in sun vated proton pump. The researchers
Institution

Perched in shade Temperature ( oC) illuminated the vesicles and then ana-
lyzed the concentration of H⫹ in them.

Conclusion: A. cristatellus uses patches of sun and shade to regulate its body temperature. Cutaway view of a synthetic vesicle

Source: P.E. Hertz. 1992. Temperature regulation in Puerto Rican Anolis lizards: A field test using null hypotheses. Ecology 73:1405–1417.
Result: H⫹ is pumped into the vesicles, creating an H⫹ gradient.

2. The researchers next made synthetic vesicles containing


Dark Light
ATP synthase from both bacteriorhodopsin and bovine H+
heart mitochondria. The ATP synthase molecule was ori- P ADP ADP
i
ented so that the ATP-synthesizing headpiece was on P
ADP i
P
the outside of the vesicles. They added ADP and Pi to i

the medium containing the vesicles and tested whether H+


H+ H+
ATP was produced in the dark and after a period of H+
+
illumination. H+ H H+

Experimental Research ADP


P
i
ADP
P
i

ADP
ATP ADP
synthase Headpiece end
ADP + P ATP
i

Result: In the dark, no ATP was synthesized. Result: In the light, ATP was synthesized.

Together, these results showed that light activated the bacteriorhodopsin to produce an
H⫹ gradient, with H⫹ moving from the outside to the inside of the vesicle (like the move-
ment from the mitochondrial matrix to the intermembrane compartment in Figure 8.13),
and that the energy from the H⫹ gradient drove ATP synthesis by ATP synthase.

Conclusion: An H⫹ gradient—and, therefore, proton-motive force—powers ATP synthesis


by ATP synthase. The results support Mitchell’s chemiosmotic hypothesis for ATP synthesis
in mitochondria.
Source: E. Racker and W. Stoeckenius. 1974. Reconstitution of purple membrane vesicles catalyzing light-driven
proton uptake and adenosine triphosphate formation. Journal of Biological Chemistry 249:662–663.

❯ Interpret the Data These exercises, most of Interpret the Data


80 KEY
Some cancer treatments target rapidly dividing cells while leaving
which are drawn from published biological research, non-proliferating cells undisturbed. The chemicals 5-fluorouracil Schedule A: 5-FU alone
Cell proliferation (%)

(5-FU) and cisplatin (CDDP) are drugs that work in this way. 5-FU 60 Schedule B: CDDP (8h)
help you build your skills in analyzing figures and in inhibits DNA replication, while CDDP binds to DNA causing changes
that cannot be corrected by DNA repair enzymes so that programmed
followed by 5-FU
Schedule C: 5-FU and CDDP
cell death (see Chapter 7) is triggered. Researchers suspected that 40
reading graphs and tables. these drugs might be useful in treating human gastric cancer and
(8h) followed by 5-FU
Schedule D: 5-FU (8h), 5-FU
tested their effectiveness in gastric cancer cells growing in culture. and CDDP (8h), followed
20
They added 5-FU alone, or 5-FU and CDDP in various timed combi- by 5-FU
nations (schedules) to cultured gastric cancer cells and measured the Schedule E: 5-FU (16h), 5-FU
inhibitory effects of the drugs on cell proliferation compared with and CDDP (8h), followed
untreated cells. The results in the figure to the right shows for each A B C D E
by 5-FU
schedule the % cell proliferation, meaning the proliferation of treated Schedule
cells/proliferation of control, untreated cells ⫻ 100%.
1. Which drug schedule was the most effective? Source: H. Cho, et al. 2002. “In-vitro effect of a combination of
2. How did the drug schedule in the most effective treatment differ 5-fluorouracil (5-FU) and cisplatin (CDDP) on human gastric cancer
from the schedules in all the other treatments? cell lines: timing of cisplatin treatment.” Gastric Cancer 5:43–46.

>
❯ Think Outside the Book These individual, THINK OUTSIDE THE BOOK
team, or Internet research activities encourage active Scientists have been working to develop an arti-
ficial version of photosynthesis that can be used
learning and peer discussion. They foster critical to produce liquid fuels from CO2 and H2O. Col-
laboratively or individually, find an example of
thinking skills by engaging you in the topic. research on artificial photosynthesis and pre-
pare an outline of how the system works or is
anticipated to work.

xv

41248_00_FM_V1_i-xxx.indd xv 11/15/10 10:17 AM


VISUAL LEARNING
Spectacular illustrations—developed with great care—help you visualize
biological processes, relationships, and structures.

❯ Illustrations of complex biological processes are annotated with


numbered step-by-step explanations that lead you through all Glycolysis ATP
1 Complex I picks up
high-energy electrons from
4 Complex IV accepts electrons
from cytochrome c and delivers
6 The resulting H+ gradient
supplies the energy that drives
NADH and conducts them via them via electron carriers cyto- ATP synthesis by ATP synthase.

the major points. Orientation diagrams are inset on figures and Pyruvate
Inner
mitochondrial
two electron carriers, FMN (flavin
mononucleotide) and an Fe/S
(iron–sulfur) protein, to
chromes a and a3 to oxygen. Four
protons are added to a molecule
of O2 as it accepts four electrons,
7 Because of the gradient, H+
oxidation
flows across the inner membrane
help you identify the specific biological process being depicted membrane ubiquinone. forming 2 H2O. and into the matrix through a
channel in the ATP synthase.
Citric
2 Complex II oxidizes FADH2 to 5 As electrons move through
and where the process takes place. acid
cycle
ATP FAD; the two electrons released
are transferred to ubiquinone, and
the electron transfer system, they
release free energy. Part of the
8 The flow of H+ activates ATP
synthase, making the headpiece
the two protons released go into released energy is lost as heat, and stalk rotate.
the matrix. Electrons that pass to but some is used by the mito-
ubiquinone by the complex II chondrion to transport H+ across
Oxidative 9 As a result of changes in
ATP reaction bypass complex I of the the inner mitochondrial
phosphorylation shape and position as it turns, the
electron transfer system. membrane from the matrix to the
headpiece catalyzes the synthesis
inter membrane compartment at
of ATP from ADP and Pi.
3 Complex III accepts electrons complexes I, III, and IV.
from ubiquinone and transfers
them through the electron carriers
in the complex—cytochrome b, an
Image copyright javarman, 2010. Used under

Fe/S protein, and cytochrome


FIGURE 9.3 c1—to cytochrome c, which is free
The membranes and compart- Cytosol in the intermembrane space.

ments of chloroplasts.
license from Shutterstock.com

Outer
mitochondrial
membrane

6
H+ H+ High H+
H+ H+
H+ H+ H+ H+ H+
H+ H+ H+
Cutaway of a small One of the photosynthetic H+ H+ H+ H+ H+
H+
section from the leaf cells, with green chloroplasts H+
ATP
synthase Stator
Leaf’s upper surface Photosynthetic cells Intermembrane
compartment
cyt c
Large central cyt b e– 4
4 Basal
Basal
vacuole e– 3 3
3 cyt a e– cyt a3 7 unit
unit
e– e–
Inner FMN e– 1 1 Ubiquinone Fe/S
3 8
mitochondrial Fe/S e– e– e–
(CoQ) Complex 4
membrane e– Complex
cyt c1 IV Stalk
Complex e– III
5 H2O
I
1 Complex 2 H+
II
5 H+
CO2 5
NADH + H+ H+ 2 H+ + 1/ O2
H+
2
Head-
piece
NAD+ FADH2 FAD + 2 H+

Mitochondrial Low H+
matrix 6
O2 9
Stomata (through Nucleus
ADP + P ATP
i
which O2 and CO2
are exchanged with Electron transfer system Chemiosmosis
the atmosphere) Electrons flow through a series of proton (H+) ATP synthase catalyzes ATP
pumps; the energy released builds an H+ gradient synthesis using energy from the H+
across the inner mitochondrial membrane. gradient across the membrane.

Oxidative phosphorylation
Cutaway view of a chloroplast
FIGURE 8.13
Oxidative phosphorylation: The mitochondrial electron transfer system and chemiosmosis. Oxidative phosphorylation involves the electron transfer
Outer membrane 1system
66 (stepsU N1–6),
I T Oand
N E chemiosmosis
M O L E C U L by
E SATP
A Nsynthase
D C E L L(steps
S 7–9). Blue arrows indicate electron flow; red arrows indicate H⫹ movement.
Inner membrane

Thylakoids Stroma (space


❮ From Macro to Micro: Multiple views help you
around thylakoids)
• light absorption by chlorophylls
and carotenoids • light-independent visualize the levels of organization of biological
• electron transfer reactions
• ATP synthesis by ATP synthase structures and how systems function as a whole.

Granum

Stromal Thylakoid Thylakoid


hylakoid
lamella lumen membrane
FIGURE 14.21
Levels of organization in eukaryotic chro-
matin and chromosomes. Histone tail
Histone H1 binds to
Histone nucleosomes and linker
DNA, causing nucleosomes
to form coiled structure Solenoid

❯ Electron micrographs are keyed to selected


illustrations to help clarify biological DNA Linker
structures.
Nucleosome: DNA
wound around core of
2 molecules each of
H2A, H2B, H3, H4
2 nm
Chromosome
in metaphase
10-nm chromatin fiber 30-nm chromatin fiber

Nucleosomes Linkers Chromatin fiber

xvi

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REVIEW
End-of-chapter material encourages you to review the content, assess your under-
standing, think analytically, and apply what you have learned to novel situations.

❯ Review Key Concepts This brief review review key concepts


references figures and tables in the chapter
and provides an outline summary of Go to CENGAGENOW at www.cengage.com/login to access quizzing, phorylation, which is comprised of the electron transfer system and
animations, exercises, articles, and personalized homework help. chemiosmosis, high-energy electrons produced from the first two
important ideas developed in the chapter. stages pass through the transfer system, with much of their energy
8.1 Overview of Cellular Energy Metabolism being used to establish an H⫹ gradient across the membrane that
drives the synthesis of ATP from ADP and Pi (Figure 8.3).
• Plants and almost all other organisms obtain energy for cellular
activities through cellular respiration, the process of transferring • In eukaryotes, most of the reactions of cellular respiration occur in
electrons from donor organic molecules to a final acceptor mol- mitochondria. In prokaryotes, glycolysis, pyruvate oxidation, and
ecule such as oxygen; the energy that is released drives ATP syn- the citric acid cycle occur in the cytosol, while the rest of cellular
thesis (Figure 8.1). respiration occurs on the plasma membrane (Figure 8.4).
• Oxidation–reduction reactions, called redox reactions, partially or Animation: The functional zones in mitochondria
completely transfer electrons from donor to acceptor atoms; the
donor is oxidized as it releases electrons, and the acceptor is 8.2 Glycolysis: Splitting the Sugar in Half
reduced (Figure 8.2).
• In glycolysis, which occurs in the cytosol, glucose (six carbons) is
• Cellular respiration occurs in three stages: (1) In glycolysis, glucose oxidized into two molecules of pyruvate (three carbons each).
is converted to two molecules of pyruvate through a series of Electrons removed in the oxidations are delivered to NAD⫹, pro-
enzyme-catalyzed reactions; (2) in pyruvate oxidation and the citric
c ducing NADH. The reaction sequence produces a net gain of
acid cycle, pyruvate is converted to an acetyl compound that is 2 ATP, 2 NADH, and 2 pyruvate molecules for each molecule of
understand and apply oxidized completely to carbon dioxide; and (3) in oxidative pho phos- glucose oxidized (Figures 8.5 and 8.7).

Test Your Knowledge 4. Which of the following statements is false? Imagine that you in-
1. What is the final acceptor for electrons in cellular respiration? gested three chocolate bars just before sitting down to study this
a. oxygen chapter. Most likely:
b. ATP a. your brain cells are using ATP.
c. carbon dioxide b. there is no deficit of the initial substrate to begin glycolysis.
d. hydrogen c. the respiratory processes in your brain cells are moving
e. water atoms from glycolysis through the citric acid cycle to the
electron transfer system.
2. In glycolysis: d. after a couple of hours, you change position and stretch to ❮ Understand and Apply End-of-chapter
a. free oxygen is required for the reactions to occur. rest certain muscle cells, which removes lactate from these
b. ATP is used when glucose and fructose-6-phosphate are muscles. questions focus on factual content in the
phosphorylated, and ATP is synthesized when 3-phospho- e. after 2 hours, your brain cells are oxygen-deficient.
glycerate and pyruvate are formed.
5. If ADP is produced in excess in cellular respiration, this excess chapter while encouraging you to apply
c. the enzymes that move phosphate groups on and off the
ADP will:
molecules are uncoupling proteins.
a. bind glucose to turn off glycolysis. what you have learned.
d. the product with the highest potential energy in the pathway
is pyruvate. b. bind glucose-6-phosphate to turn off glycolysis.
e. the end product of glycolysis moves to the electron transfer c. bind phosphofructokinase to turn on or keep glycolysis
system. turned on.
d. cause lactate to form.
3. Which of the following statements about phosphofructokinase is e. increase oxaloacetate binding to increase NAD+ production.
false?
a. It is located and has its main activity in the inner mitochon- 6. Which of the following statements is false? In cellular respiration:
drial membrane. a. one molecule of glucose can produce about 32 ATP.
b. It catalyzes a reaction to form a product with the highest b. oxygen combines directly with glucose to form carbon dioxide.
potential energy in the pathway. c. a series of energy-requiring reactions is coupled to a series of
c. It can be inactivated by ATP at an inhibitory site on its energy-releasing reactions.
surface. d. NADH and FADH2 allow H+ to be pumped across the inner
d. It can be activated by ADP at an excitatory site on its surface. mitochondrial membrane.
e. It can cause ADP to form. e. the electron transfer system occurs in the inner mitochon-
drial membrane.

Design an Experiment Design an Experiment challenges your understanding of


There are several ways to measure cellular respiration experimentally. the chapter and helps deepen your understanding of the
For example, CO2 and O2 gas sensors measure changes over time in scientific method as you consider how to develop and test
the concentration of carbon dioxide or oxygen, respectively. Design
two experiments to test the effects of changing two different variables hypotheses about a situation that relates to a main chapter
or conditions (one per experiment) on the respiration of a research
organism of your choice. topic.

Discuss the Concepts Discuss the Concepts enables you to


1. Why do you think nucleic acids are not oxidized extensively as a participate in discussions on key questions
cellular energy source?
2. A hospital patient was regularly found to be intoxicated. He de- to build your knowledge and learn from
nied that he was drinking alcoholic beverages. The doctors and
nurses made a special point to eliminate the possibility that the
others.
patient or his friends were smuggling alcohol into his room, but
he was still regularly intoxicated. Then, one of the doctors had an
idea that turned out to be correct and cured the patient of his in-
toxication. The idea involved the patient’s digestive system and
41248_08_0155-0175.indd 175 one of the oxidative reactions covered in this chapter. What was 7/9/10 1:56:48 PM
the doctor’s idea?

Express Your Opinion Express Your Opinion encourages you to weigh both sides
Developing new drugs is costly. There is little incentive for pharma- of an issue by reading pro/con articles, and then make your
ceutical companies to target ailments that affect relatively few indi-
viduals, such as Luft syndrome. Should governments allocate some opinion known through an online voting process.
funds to private companies that search for cures for rare disorders? Go
to CengageNOW to investigate both sides of the issue and then vote
online.

xvii

41248_00_FM_V1_i-xxx.indd xvii 11/15/10 10:17 AM


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41248_00_FM_V1_i-xxx.indd xviii 11/15/10 10:18 AM


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