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Organization and New Content
Campbell BIOLOGY IN FOCUS, Second Edition, is organized UNIT 1 Chemistry and Cells
into an introductory chapter and seven units that cover core
concepts of biology at a thoughtful pace. When we adapted A succinct, two-chapter treatment of
Campbell BIOLOGY to write the first edition of this text, we basic chemistry (Chapters 2 and 3)
made informed choices about how to design each chapter of provides the foundation for this unit
Campbell BIOLOGY IN FOCUS to meet the needs of focused on cell structure and function.
instructors and students. In some chapters, we retained most The related topics of cell membranes
of the material; in other chapters, we pruned material; and in and cell signaling are consolidated into
still others, we completely reconfigured the material. In creating one c hapter (Chapter 5). Due to the im-
the Second Edition, we solicited feedback from reviewers and portance of the fundamental concepts
used their thoughtful critiques to further fine-tune the content in Units 1 and 2, much of the material
and pedagogy. We have also updated the content wherever in the rest of these two units has been retained from
appropriate, and in a few cases reintroduced material. Here, Campbell BIOLOGY.
we present synopses of the seven units and highlight the major For the Second Edition, a new table has been added to
revisions made to the Second Edition of Campbell BIOLOGY Chapter 2 detailing the elements in the human body, with an
IN FOCUS. associated Interpret the Data q uestion. Chapter 3 includes
a new section on isomers, with an accompanying figure
CHAPTER 1 Introduction: Evolution and the Foundations (Figure 3.5), and ends with a new Concept 3.7 that includes
of Biology cutting-edge coverage of DNA sequencing and introduces
genomics and proteomics, as well as bioinformatics. A new
Chapter 1 introduces the five biological Make Connections Figure (Figure 3.30) entitled “Contributions
themes woven throughout the text: of Genomics and Proteomics to Biology” provides an over-
the core theme of Evolution, together view of areas in which genomics and proteomics have had
with Organization, Information, significant impacts—including evolution, conservation biology,
Energy and Matter, and Interactions. paleontology, medical science, and species interactions—with
Chapter 1 also explores the process of the aim of inspiring and motivating students. A striking photo
scientific inquiry through a case study of thermophilic cyanobacteria has been added to Figure 6.16
describing experiments on the evolu- on environmental factors affecting enzyme activity. In
tion of coat color in the beach mouse. Chapter 7, a computer model of ATP synthase has been added
The chapter concludes with a discussion of the importance of to Figure 7.13. The icon for this enzyme in Chapters 7 and 8 has
diversity within the scientific community. been re-drawn to more closely represent its structure. A new
In the Second Edition, a new figure (Figure 1.8) on Make Connections Figure (Figure 8.20, “The Working Cell”)
gene expression uses lens cells in the eye as an example of integrates all the cellular activities covered in Chapters 3–8 in
DNA → RNA → protein and introduces the terms tran- the context of a single working plant cell.
scription and translation. This new figure and text equip
students from the outset with an understanding of how gene
sequences determine an organism’s characteristics. New UNIT 2 Genetics
text and a new photo (Figure 1.11) inform students about
the effects of climate change in general, and global warming Topics in this unit include meiosis and
in particular, on species survival and diversity. Concept 1.3 classical genetics as well as the chromo-
has been thoroughly revised to more realistically reflect somal and molecular basis for genetics
the process of science. A new section has been added on and gene expression (Chapters 10–14).
the Flexibility of the Scientific Process, accompanied by a We also include a chapter on the regu-
new Figure 1.19 that depicts the more realistic and complex lation of gene expression (Chapter 15)
process of science. The text now discusses searching the sci- and one on the role of gene regulation
entific literature, and a new question in the Chapter Review in development, stem cells, and cancer
asks students to use PubMed. (Chapter 16). Methods in biotechnology
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are integrated into appropriate chapters. The stand-alone UNIT 3 Evolution
chapter on viruses (Chapter 17) can be taught at any point in
the course. The final chapter in the unit, on genome evolution This unit provides in-depth coverage
(Chapter 18), provides both a capstone for the study of genet- of essential evolutionary topics, such
ics and a bridge to the evolution unit. as mechanisms of natural selection,
Chapter 10 of the Second Edition includes a new section population genetics, and speciation.
on “Crossing Over and Synapsis During Prophase I” that Early in the unit, Chapter 20 introduces
explains the events of prophase I in more detail, supported “tree thinking” to support students in
by new Figure 10.9, which clearly shows and describes these interpreting phylogenetic trees and
events. In Chapter 11, to incorporate more molecular biol- thinking about the big picture of evolu-
ogy into the discussion of Mendelian genetics, Figure 11.4 tion. Chapter 23 focuses on mechanisms that have influenced
on alleles has been enhanced and a new Figure 11.16 on long-term patterns of evolutionary change. Throughout the
sickle-cell disease has been added. Chapter 13 includes new unit, new discoveries in fields ranging from paleontology to
text and two new figures (Figures 13.29 and 13.30) cover- phylogenomics highlight the interdisciplinary nature of mod-
ing advances in sequencing technology. Also in this chapter, ern biology.
a new section, including new Figure 13.31, describes gene Revisions in the Second Edition aim to strengthen connec-
editing using the CRISPR-Cas9 system. In Chapter 15, the tions among fundamental evolutionary concepts. For example,
section on noncoding RNAs has been updated, and Concept 20.5 includes new text on horizontal gene transfer
Figure 15.14 on in situ hybridization has been expanded among eukaryotes, reinforcing the overall discussion of how
and enhanced to help students understand this important horizontal gene transfer has played an important role in the
technique. Chapter 16 includes a new Inquiry Figure evolutionary history of life. Also in Concept 20.5, a new
(Figure 16.16) on induced pluripotent stem cells (iPS cells). Scientific Skills Exercise walks students through the process
Material on embryonic stem cells and induced pluripotent of comparing and interpreting amino acid sequences to deter-
stem cells has been significantly updated. A new Make mine whether horizontal gene transfer may have occurred in
Connections Figure (Figure 16.21), “Genomics, Cell Signal- certain organisms. Chapter 20 also includes more discussion of
ing, and Cancer,” illustrates recent research on subtypes of tree thinking, as well as a new figure (Figure 20.11) that distin-
breast cancer, connecting content that students have learned guishes between paraphyletic and polyphyletic taxa. New ma-
in Chapters 5, 9, and 16. It also addresses treatment for terial in Chapter 21 clarifies the interplay between mutation,
one subtype of breast cancer as an example. In Chapter 17, genetic variation, and natural selection. A new Make Connec-
the discussion of the importance of cell-surface proteins in tions Figure (Figure 21.15, “The Sickle-Cell Allele”) integrates
determining host range has been enhanced. A new figure material from chapters across the book in exploring the sickle-
(Figure 17.9) presents the example of the receptor and co- cell allele and its impact from the molecular and cellular levels
receptor proteins for HIV. Coverage of the CRISPR system, to the allele’s global distribution in the human population.
as a bacterial “immune” system, has been added, supported Other changes in the unit include new examples and figures
by new Figure 17.6. Coverage of recent epidemics has been that reinforce evolutionary concepts. For example, a new
inserted (Ebola) or updated (H5N1). Chapter 18 has been introduction to Chapter 23 tells the story of the discovery of
significantly updated to reflect recent sequencing advances, whale fossils from the Sahara Desert, striking evidence of how
including a discussion of the results of the ENCODE organisms in the past differed from organisms living today.
project, information on the bonobo genome, and use In Chapter 22, a new figure (Figure 22.11) has been added to
of high-throughput techniques to address the problem support the expanded text discussion of allopolyploid specia-
of cancer. Regarding protein structure, the discussion tion in Tragopogon in the Pacific Northwest. Dates have also
of BLAST searches has been enhanced, and computer been revised in the text, Table 23.1 (The Geologic Record),
models of lysozyme and α-lactalbumin have been added to and figures in Chapter 23 and throughout the Second Edition
support the discussion of the evolution of genes with novel to reflect the International Commission on Stratigraphy 2013
functions. revision of the Geologic Time Scale.
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UNIT 4 The Evolutionary History of Life UNIT 5 Plant Form and Function
This unit employs a novel approach to The form and function of higher plants
studying the evolutionary history of are often treated as separate topics,
biodiversity. Each chapter focuses on thereby making it difficult for students
one or more major steps in the history to make connections between the two.
of life, such as the origin of cells or the In Unit 5, plant anatomy (Chapter 28)
colonization of land. Likewise, the cov- and the acquisition and transport of
erage of natural history and biological resources (Chapter 29) are bridged by
diversity emphasizes the evolutionary a discussion of how plant architec-
process—how factors such as the origin ture influences resource acquisition.
of key adaptations have influenced the rise and fall of different Chapter 30 provides an introduction to plant reproduction
groups of organisms over time. and examines controversies surrounding the genetic engineer-
In the Second Edition, we have expanded our coverage of ing of crop plants. The final chapter (Chapter 31) explores how
genomic and other molecular studies. Examples include a new plants respond to environmental challenges and opportunities
figure (Figure 24.25) and text on the potential use and signifi- and how the integration of this diverse information by plant
cance of CRISPR-Cas systems, a new Scientific Skills Exercise hormones influences plant growth and reproduction.
in Chapter 26 on genomic analyses of mycorrhizal and nonmy- In the Second Edition, a new micrograph of parenchyma
corrhizal fungi, and a new figure (Figure 27.36) and text related cells and new information relating to root hair density,
to evidence of gene flow between Neanderthals and modern length, and function have been added to Chapter 28. In
humans. In addition, many phylogenies have been revised to Chapter 29, a new Make Connections Figure (Figure 29.10,
reflect recent miRNA and genomic data. The unit also includes “Mutualism Across Kingdoms and Domains”) enables stu-
more connections to other chapters. For instance, a new Make dents to integrate what they have learned about plant mutu-
Connections Question in Figure 24.4 asks students to apply alisms with other examples across the natural realm. A new
material from Chapter 3 to explain how a membrane-like bi- Inquiry Figure (Figure 29.11) examines the metagenomics of
layer can self-assemble and form a vesicle, and a new Make soil bacteria. A discussion on mycorrhizae and plant evolu-
Connections Figure (Figure 26.14) explores the diverse struc- tion has also been added in Chapter 29. In Chapter 30, the
tural solutions for maximizing surface area that have evolved angiosperm life cycle figure and related text are more closely
in cells, organ systems, and whole organisms. Other changes integrated, with all the numbered steps now identified in the
enhance the evolutionary storyline of the unit. For example, text. Also, a discussion of coevolution of flowers and pollina-
in Chapter 26, the chapter title, Figure 26.2, Key Concept 26.2, tors has been added. The in-depth discussion of the devel-
and text in Concepts 26.1 and 26.2 have all been revised to em- opment from seed to flowering plant has been expanded to
phasize and explain that fungi are not closely related to plants, include the transition from vegetative growth to reproduc-
although they likely played a role in facilitating the colonization tive growth, making a connection to what students learned
of land by plants, and that fungi possess their own novel adap- about development in Chapter 28. In addition, the depictions
tations for terrestrial life. Likewise, in Chapter 27, the discus- of the structure of maize root systems and raspberry fruit
sion of the evolutionary impact of animals has been expanded, development have been improved. The information in Con-
and new text and four new figures (Figures 27.12, 27.13, 27.30, cept 31.4 concerning plant defenses against disease has been
and 27.31) on molluscs, birds, and mammals have been added. thoroughly revised and updated to reflect rapid advances in
The chapter also includes expanded coverage of human evo- our understanding of plant immunity. Updated information
lution, including three new figures (Figures 27.34, 27.35, and relates to the two types of plant immunity: PAMP-triggered
27.36). Supporting the extensive revision of Chapter 27, the immunity and effector-triggered immunity. New Figure 31.23
number of Key Concepts in this chapter has increased from highlights examples of physical, chemical, and behavioral de-
five to seven. fenses against herbivory.
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UNIT 6 Animal Form and Function UNIT 7 Ecology
In this unit, a focused exploration of This unit applies the key themes of
animal physiology and anatomy ap- the text, including evolution, interac-
plies a comparative approach to a tions, and energy and matter, to help
limited set of examples to bring out students learn ecological principles.
fundamental principles and conserved Chapter 40 integrates material on
mechanisms. Students are first intro- population growth and Earth’s environ-
duced to the closely related topics of ment, highlighting the importance of
endocrine signaling and homeostasis both biological and physical processes
in an integrative introductory chapter in determining where species are found.
(Chapter 32). Additional melding of interconnected material Chapter 43 ends the book with a focus on global ecology and
is reflected in chapters that combine treatment of circulation conservation biology. This chapter illustrates the threats to
and gas exchange, reproduction and development, neurons all species from increased human population growth and re-
and nervous systems, and motor mechanisms and behavior. source use. It begins with local factors that threaten individual
In the Second Edition, we re-envisioned the introductory species and ends with global factors that alter ecosystems,
chapter of this unit (Chapter 32), as conveyed by its new title, landscapes, and biomes.
“The Internal Environment of Animals: Organization and The increased emphasis throughout the Second Edition on
Regulation.” Endocrine signaling and the integration of ner- global climate change is capped by new discussions and figures
vous and endocrine system function now precede the intro- in Unit 7. Chapter 43, for example, includes a new figure on
duction of homeostasis and the consideration of the two major the greenhouse effect (Figure 43.26) as well as new text exam-
examples: thermoregulation and osmoregulation. Figures on ining aspects of climate change other than global warming.
simple hormone and neurohormone pathways (Figures 32.6 The chapter explores documented examples of the impacts to
and 32.7) and hormone cascades (Figure 32.8) have been sub- organisms in a new section on “Biological Effects of Climate
stantially revised to provide clear and consistent presentation Change” and a new Make Connections Figure (Figure 43.28,
of hormone function and of the regulation of hormone secre- “Climate Change Has Effects at All Levels of Biological Orga-
tion. The presentation of the mechanism for filtrate process- nization”). Throughout the unit, the presentation of several
ing in the kidney has been substantially revised, with a single other key topics has been revised. For example, in Chapter 40,
figure (Figure 32.22) in place of two and with the accompany- the discussion of each of the following concepts or models was
ing numbered text walking students through a carefully paced revised to standardize and clarify their meaning: life tables, per
tour of the nephron. In this chapter and throughout the unit, capita population growth, the per capita rate of increase (r),
figures illustrating homeostatic regulation have been revised to exponential population growth, and logistic population growth.
highlight the common principles and features of homeostatic The discussion of species interactions in Chapter 41 was
mechanisms. The unit includes two new Make Connections modified to group species interactions according to whether
Figures: Figure 32.3 illustrates shared and divergent solutions they have positive (+) or negative (–) effects on survival and
to fundamental challenges common to plants and animals, reproduction; as a result, there is a new section on “Exploita-
and Figure 37.8, on ion movements and gradients, explores the tion” (which includes predation, herbivory, and parasitism)
fundamental role of concentration gradients in life processes and another new section on “Positive Interactions” (which
ranging from osmoregulation and gas exchange to locomo- includes mutualism and commensalism). Material throughout
tion. Also in Chapter 37, the treatments of synaptic signaling, Chapter 42 was revised to reinforce the fact that energy flows
summation, modulating signaling, and neurotransmitters through ecosystems, whereas chemical elements cycle within
have been revised to highlight key ideas, ensuring appropriate ecosystems. New Figure Legend Questions give students
pacing and helping students focus on fundamental principles practice in actively interpreting results; see, for example, the
rather than memorization. Updates in Unit 6 informed by cur- new questions with Figure 43.22 (biological magnification of
rent research include new Figure 33.15 and text highlighting PCBs) and Figure 43.31 (a new figure on per capita ecological
the explosion of interest in and understanding of the microbi- footprints). The unit also includes a new Make Connections
ome. Chapter 38 opens with a new photograph and introduc- Figure (Figure 42.18, “The Working Ecosystem”) that ties
tory text that showcase the “brainbow” technique for labeling together population, community, and ecosystem processes in
individual brain neurons. the arctic tundra.
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About the Authors
The author team’s contributions reflect their biological expertise as researchers and their teaching sensibilities
gained from years of experience as instructors at diverse institutions. They are also experienced textbook
authors, having written Campbell BIOLOGY in addition to Campbell BIOLOGY IN FOCUS.
Lisa A. Urry
Lisa Urry (Chapter 1 and Units 1 and 2) is Professor of Biology and Chair of the Biology
Department at Mills College in Oakland, California, and a Visiting Scholar at the University of
California, Berkeley. After graduating from Tufts University with a double major in biology and
French, Lisa completed her Ph.D. in molecular and developmental biology at Massachusetts
Institute of Technology (MIT) in the MIT/Woods Hole Oceanographic Institution Joint P rogram.
She has published a number of research papers, most of them focused on gene expression dur-
ing embryonic and larval development in sea urchins. Lisa has taught a variety of courses, from
introductory biology to developmental biology and senior seminar. As a part of her mission to
increase understanding of evolution, Lisa also teaches a nonmajors course called Evolution for
Future Presidents and is on the Teacher Advisory Board for the Understanding Evolution website
developed by the University of California Museum of Paleontology. Lisa is also deeply committed
to promoting opportunities for women and underrepresented minorities in science.
Michael L. Cain
Michael Cain (Chapter 1 and Units 3, 4, and 7) is an ecologist and evolutionary biologist who is
now writing full-time. Michael earned a joint degree in biology and math at Bowdoin College,
an M.Sc. from Brown University, and a Ph.D. in ecology and evolutionary biology from Cornell
University. As a faculty member at New Mexico State University and Rose-Hulman Institute of
Technology, he taught a wide range of courses, including introductory biology, ecology, evolu-
tion, botany, and conservation biology. Michael is the author of dozens of scientific papers on
topics that include foraging behavior in insects and plants, long-distance seed dispersal, and
speciation in crickets. In addition to his work on Campbell BIOLOGY IN FOCUS, Michael is
also the lead author of an ecology textbook.
Steven A. Wasserman
Steve Wasserman (Chapter 1 and Unit 6) is Professor of Biology at the University of California,
San Diego (UCSD). He earned his A.B. in biology from Harvard University and his Ph.D. in bio-
logical sciences from MIT. Through his research on regulatory pathway mechanisms in the fruit
fly Drosophila, Steve has contributed to the fields of developmental biology, reproduction, and
immunity. As a faculty member at the University of Texas Southwestern Medical Center and
UCSD, he has taught genetics, development, and physiology to undergraduate, graduate, and
medical students. He currently focuses on teaching introductory biology. He has also served as
the research mentor for more than a dozen doctoral students and more than 50 aspiring scientists
at the undergraduate and high school levels. Steve has been the recipient of distinguished scholar
awards from both the Markey Charitable Trust and the David and Lucille Packard Foundation. In
2007, he received UCSD’s Distinguished Teaching Award for undergraduate teaching.
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Peter V. Minorsky
Peter Minorsky (Chapter 1 and Unit 5) is Professor of Biology at Mercy College in New York,
where he teaches introductory biology, evolution, ecology, and botany. He received his A.B.
in biology from Vassar College and his Ph.D. in plant physiology from Cornell University.
He is also the science writer for the journal Plant Physiology. After a postdoctoral fellowship
at the University of Wisconsin at Madison, Peter taught at Kenyon College, Union College,
Western Connecticut State University, and Vassar College. His research interests concern how
plants sense environmental change. Peter received the 2008 Award for Teaching Excellence at
Mercy College.
Jane B. Reece
The head of the author team for recent editions of Campbell BIOLOGY, Jane Reece was Neil
Campbell’s longtime collaborator. Earlier, Jane taught biology at Middlesex County College and
Queensborough Community College. She holds an A.B. in biology from Harvard University, an
M.S. in microbiology from Rutgers University, and a Ph.D. in bacteriology from the University
of California, Berkeley. Jane’s research as a doctoral student and postdoctoral f ellow focused
on genetic recombination in bacteria. Besides her work on the Campbell textbooks for biology
majors, she has been an author of Campbell Biology: Concepts & Connections, Campbell
Essential Biology, and The World of the Cell.
Neil A. Campbell
Neil Campbell (1946–2004) combined the investigative nature of a research scientist with the
soul of an experienced and caring teacher. He earned his M.A. in zoology from the University
of California, Los Angeles, and his Ph.D. in plant biology from the University of California,
Riverside, where he received the Distinguished Alumnus Award in 2001. Neil published numer-
ous research articles on desert and coastal plants and how the sensitive plant (Mimosa) and
other legumes move their leaves. His 30 years of teaching in diverse environments included
introductory biology courses at Cornell University, Pomona College, and San Bernardino V alley
College, where he received the college’s first Outstanding Professor Award in 1986. He was a
visiting scholar in the Department of Botany and Plant Sciences at the University of California,
Riverside. Neil was the lead author of Campbell Biology: Concepts & Connections, Campbell
Essential Biology, and Campbell BIOLOGY, upon which this book is based.
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Make Connections Visually
NEW! Ten Make
Connections Figures
integrate content from ▼ Figure 32.3
different chapters MAKE CONNECTIONS

and provide a visual
Life Challenges and Solutions
representation of “big
in Plants and Animals
picture” relationships. Multicellular organisms face a common set
of challenges. Comparing the solutions that
have evolved in plants and animals reveals
both unity (shared elements) and diversity
Make Connections (distinct features) across these two lineages.
Figures include:
Figure 3.30 Contributions of

Genomics and Proteomics
to Biology, p. 68
Figure 8.20 The Working Cell,

pp. 178–179
Figure 16.21 Genomics, Nutritional Mode

Cell Signaling, and Cancer, All living things must obtain energy and carbon from the
pp. 338–339 environment to grow, survive, and reproduce. Plants are
autotrophs, obtaining their energy through photosynthesis
Figure 21.15 The Sickle-Cell and their carbon from inorganic sources, whereas animals are
heterotrophs, obtaining their energy and carbon from food.
Allele, pp. 428–429 Evolutionary adaptations in plants and animals support these
different nutritional modes. The broad surface of many leaves
Figure 26.14 Maximizing (left) enhances light capture for photosynthesis. When hunting,
Surface Area, p. 526 a bobcat relies on stealth, speed, and sharp claws (right). (See
Figure 29.2 and Figure 33.14.)
Figure 29.10 Mutualism Across
Kingdoms and Domains, p. 603 Growth and Regulation
The growth and physiology of both plants and animals are
Figure 32.3 Life Challenges regulated by hormones. In plants, hormones may act in a local
and Solutions in Plants and area or be transported in the body. They control growth patterns,
flowering, fruit development, and more (left). In animals,
Animals, shown at right and hormones circulate throughout the body and act in specific
on pp. 666–667 target tissues, controlling
Environmental Response homeostatic processes and
Figure 37.8 Ion Movement and developmental events such
All forms of life must detect and respond
Gradients, p. 777 as molting (below).
appropriately to conditions in their
(See Table 31.1 and Figure 33.19.)
environment. Specialized organs sense
Figure 42.18 The Working environmental signals. For example,
Ecosystem, pp. 902–903 the floral head of a sunflower (left) and
an insect’s eyes (right) both contain
photoreceptors that detect light.
Figure 43.28 Climate Change
Environmental signals activate specific
Has Effects at All Levels of receptor proteins, triggering signal
Biological Organization, transduction pathways that initiate
cellular responses coordinated by
pp. 924–925 chemical and electrical communication.
(See Figure 31.12 and Figure 38.26.)
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Reproduction
In sexual reproduction,
specialized tissues and
structures produce and
exchange gametes. Offspring
are generally supplied with
nutritional stores that facilitate
Transport rapid growth and development.
All but the simplest For example, seeds (left) have
multicellular organisms stored food reserves that supply energy to the young seedling,
must transport nutrients and while milk provides sustenance for juvenile mammals (right).
waste products between locations in the body. A system of (See Figure 30.8 and Figure 32.7.)
tubelike vessels is the common evolutionary solution, while the
mechanism of circulation varies. Plants harness solar energy
to transport water, minerals, and sugars through specialized
tubes (left). In animals, a pump (heart) moves circulatory fluid
through vessels (right). (See Figure 28.9 and Figure 34.3.)
Gas Exchange
The exchange of certain
gases with the environment
is essential for life.
Respiration by plants and
animals requires taking up
oxygen (O2) and releasing carbon dioxide (CO2). In photosynthesis,
net exchange occurs in the opposite direction: CO2 uptake and O2
release. In both plants and animals, highly convoluted surfaces that
increase the area available for gas exchange have evolved, such as
the spongy mesophyll of leaves (left) and the alveoli of lungs (right).
(See Figure 28.17 and Figure 34.20.)
Absorption make connections Compare the adaptations that enable plants

Make Connections Questions
and animals to respond to the challenges of living in hot and cold
Organisms need to absorb nutrients. The root hairs ask students to relate
environments. See Concepts 31.3 and 32.3.
of plants (left) and the villi (projections) that line
the intestines of vertebrates (right) increase the
content in the chapter to
Visit the Study Area in MasteringBiology for the BioFlix®
surface area available for absorption. (See Figure 28.4
animation
3-d Animations on Water transport in Plants (chapter 29), material presented earlier in
and Figure 33.10.) Homeostasis: regulating Blood Sugar (chapter 33), and the course.
Gas Exchange (chapter 34).
ChApTEr 32 tHE intErnAl EnVironmEnt oF AnimAlS: orGAnizAtion And rEGulAtion 667
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M a k e C onnections V isually xiii

might be found. For example, scientists have constructed evolu- Species, most island species are closely related to species from
tionary trees for horses based on anatomical data. These trees and the nearest mainland or a neighboring island. He explained
Practice Scientific Skills

the ages of fossils of horse ancestors suggest that the genus that this observation by suggesting that islands are colonized by
includes present-day horses (Equus) originated 5 million years species from the nearest mainland. These colonists eventually
ago in North America. Geologic evidence indicates that at that give rise to new species as they adapt to their new environ-
time, North and South America were not yet connected, mak- ments. Such a process also explains why two islands with
ing it difficult for horses to travel between them. Thus, we would similar environments in distant parts of the world tend to be
predict that the oldest Equus fossils should be found only on the populated not by species that are closely related to each other,
continent on which the group originated—North America. This but rather by species related to those of the nearest mainland,
Scientific Skills Exercises in every chapter use real data to build
prediction and others like it for different groups of organisms have where the environment is often quite different.
been upheld, providing more evidence for evolution.
key skills needed for biology, including data analysis,
What Isgraphing,
Theoretical About Darwin’s
We can also use our understanding of evolution to explain
View of Life?
biogeographic data. For example, islands generally have many
experimental design, and math skills.
plant and animal species that are endemic (found nowhere
Some people dismiss Darwin’s ideas as “just a theory.” However,
else in the world). Yet, as Darwin described in The Origin of as we have seen, the pattern of evolution—the observation that
Scientific Skills Exercise
Making and Testing Predictions

Can Predation Result in Natural Selection for Color Patterns Data from the Experiment After 22 months (15 generations),
in Guppies? What we know about evolution changes constantly as researchers compared the color pattern data for guppies from the
new observations lead to new hypotheses—and hence to new ways source and transplanted populations.
to test our understanding of evolutionary theory. Consider the wild
Each Scientific Skills guppies (Poecilia reticulata) that live in pools connected by streams on 12 12
the Caribbean island of Trinidad. Male guppies have highly varied color
Exercise is based on an 10 10
Area of colored
colored spots
patterns that are controlled by genes that are only expressed in adult
spots (mm2)
Number of
8 8
males. Female guppies choose males with bright color patterns as mates
experiment related to more often than they choose males with drab coloring. But the bright
6 6
the chapter content. colors that attract females also can make the males more conspicuous to 4
2
4
2
predators. Researchers observed that in pools with few predator species,
the benefits of bright colors appear to “win out,” and males are more 0 0
Source Transplanted Source Transplanted
brightly colored than in pools where predation is more intense. population population population population
One guppy predator, the killifish, preys on juvenile guppies that
have not yet displayed their adult coloration. Researchers predicted Data from J. A. Endler, Natural selection on color patterns in Poecilia reticulata,
Evolution 34:76–91 (1980).
Most Scientific Skills
that if adult guppies with drab colors were transferred to a pool
with only killifish, eventually the descendants of these guppies Exercises use data
I N T E R P R E T T HE D ATA
would be more brightly colored (because of the female preference
for brightly colored males). 1. Identify the following elements of hypothesis-based science from published
How the Experiment Was Done Researchers transplanted
in this example: (a) question, (b) hypothesis, (c) prediction, research, cited in the
(d) control group, and (e) experimental group. (For additional
200 guppies from pools containing pike-cichlid fish, intense preda- information about hypothesis-based science, see Chapter 1 and exercise.
tors of adult guppies, to pools containing killifish, less active preda- the Scientific Skills Review in Appendix F and the Study Area of
tors that prey mainly on juvenile guppies. They tracked the number MasteringBiology.)
of bright-colored spots and the total area of those spots on male 2. Explain how the types of data the researchers chose to collect
guppies in each generation. enabled them to test their prediction.
3. What conclusion do you draw from the data presented above?
Guppies
transplanted
4. Predict what would happen if, after 22 months, guppies from
the transplanted population were returned to the source pool.
Questions build in
Describe an experiment to test your prediction. difficulty, walking
A related version of this Scientific Skills Exercise can be assigned students through new
in MasteringBiology.
skills step by step and
providing opportunities
for higher-level critical
Pools with thinking.
pike-cichlids Pools with killifish,
and guppies but no guppies
prior to transplant
392 UNIT THREE EVOLUTION
Every chapter has a Scientific Skills Exercise:

DESIGN SERVICES OF
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1. Interpreting a Pair of Bar Graphs, p. 18 13. Working with Data in a Table, p. 257
2. Interpreting a Scatter Plot with a Regression Line, p. 40 14. Interpreting a Sequence Logo, p. 294
3. Analyzing Polypeptide Sequence Data, p. 69 15. Analyzing DNA Deletion Experiments, p. 313
4. Using a Scale Bar to Calculate Volume and Surface Area 16. Analyzing Quantitative and Spatial Gene Expression Data, p. 325
of a Cell, p. 80 17. Analyzing a Sequence-Based Phylogenetic Tree to Understand
5. Interpreting a Scatter Plot with Two Sets of Data, p. 109 Viral Evolution, p. 353
6. Making a Line Graph and Calculating a Slope, p. 134 18. Reading an Amino Acid Sequence Identity Table, p. 370
7. Making a Bar Graph and Evaluating a Hypothesis, p. 155 19. Making and Testing Predictions, shown above and on p. 392
8. Making Scatter Plots with Regression Lines, p. 176 20. NEW! Using Protein Sequence Data to Test an Evolutionary
Hypothesis, p. 410
9. Interpreting Histograms, p. 196
21. Using the Hardy-Weinberg Equation to Interpret Data and Make
10. Making a Line Graph and Converting Between Units of
Predictions, p. 420
Data, p. 210
22. Identifying Independent and Dependent Variables, Making a
11. Making a Histogram and Analyzing a Distribution
Scatter Plot, and Interpreting Data, p. 441
Pattern, p. 227
23. Estimating Quantitative Data from a Graph and Developing
12. Using the Chi-Square (χ2) Test, p. 246
Hypotheses, p. 459
xiv P ractice S cienti f ic S k ills

Visit https://testbankfan.com
now to explore a rich
collection of testbank or
solution manual and enjoy
exciting offers!
Each Scientific Skills Exercise from the text also has an assignable,
interactive tutorial version in MasteringBiology that is automatically
graded and includes coaching feedback.
To learn more, visit www.masteringbiology.com
24. Making a Bar Graph and Interpreting Data, p. 493 34. Interpreting Data in Histograms, shown above and on p. 721
25. Interpreting Comparisons of Genetic Sequences, p. 501 35. Comparing Two Variables on a Common x-Axis, p. 748
26. NEW! Interpreting Genomic Data and Generating 36. Making Inferences and Designing an Experiment, p. 761
Hypotheses, p. 529 37. Interpreting Data Values Expressed in Scientific Notation, p. 787
27. Understanding Experimental Design and Interpreting 38. Designing an Experiment Using Genetic Mutants, p. 797
Data, p. 570
39. Interpreting a Graph with Log Scales, p. 825
28. Using Bar Graphs to Interpret Data, p. 582
40. Using the Logistic Equation to Model Population Growth, p. 860
29. Calculating and Interpreting Temperature
Coefficients, p. 597 41. Using Bar Graphs and Scatter Plots to Present and Interpret
Data, p. 870
30. Using Positive and Negative Correlations to Interpret
Data, p. 632 42. Interpreting Quantitative Data in a Table, p. 893
31. Interpreting Experimental Results from a Bar 43. Graphing Cyclic Data, p. 922
Graph, p. 656
32. Describing and Interpreting Quantitative
Data, p. 679
33. Interpreting Data from an Experiment with
Genetic Mutants, p. 704
P ractice S cienti f ic S k ills xv

Interpret Data
Campbell BIOLOGY IN FOCUS, Second Edition, and
MasteringBiology offer a wide variety of ways for students to
move beyond memorization and to think like a scientist.
NEW! Interpret the Data Questions

such genes change only slowly. But if the exact sequence of
90 throughout
amino the text
acids is less critical, fewerask
of thestudents to will
new mutations
number of mutations
beanalyze
harmful andamore
graph,will befigure, or table.
neutral. Such genes change
60 more quickly.
Potential Problems with Molecular Clocks

30
In fact, molecular clocks do not run as smoothly as would be
expected if the underlying mutations were selectively neutral.
0 Many irregularities are likely to be the result of natural selec-
0 30 60 90 120 tion in which certain DNA changes are favored over others.
Divergence time (millions of years) Indeed, evidence suggests that almost half the amino acid dif-
ferences in proteins of two Drosophila species, D. simulans
▲ Figure 20.19 A molecular clock for mammals. The number
and D. yakuba, are not neutral but have resulted from natural
of accumulated mutations in seven proteins has increased over time
in a consistent manner for most mammal species. The three green selection. But because the direction of natural selection may
data points represent primate species, whose proteins appear to have change repeatedly over long periods of time (and hence may
evolved more slowly than those of other mammals. The divergence average out), some genes experiencing selection can neverthe-
time for each data point was based on fossil evidence.
less serve as approximate markers of elapsed time.
I NT E R P R E T TH E D ATA Use the graph to estimate the divergence
time for a mammal with a total of 30 mutations in the seven proteins.
Another question arises when researchers attempt to ex-
tend molecular clocks beyond the time span documented by
the fossil record. Although some fossils are more than 3 billion
genetic differences—for example, nucleotide, codon, or amino years old, these are very rare. An abundant fossil record ex-
acid differences—against the dates of evolutionary branch tends back only about 550 million years, but molecular clocks
points that are known from the fossil record (Figure 20.19). have been used to date evolutionary divergences that occurred
The average rates of genetic change inferred from such graphs a billion or more years ago. These estimates assume that the
can then be used to estimate the dates of events that cannot clocks haveNEW! Every
been constant forInterpret the estimates
all that time. Such Data are
be discernedLearn
from more at record, such as the origin of the
the fossil highly uncertain.
www.masteringbiology.com
question from the text is assignable
silverswords discussed earlier. In some cases, problems may be avoided by calibrating
Of course, no gene marks time with complete precision. In in MasteringBiology.
molecular clocks with data on the rates at which genes have
fact, some portions of the genome appear to have evolved in evolved in different taxa. In other cases, problems may be
irregular bursts that are not at all clocklike. And even those avoided by using many genes rather just using one or a few
genes that seem to act as reliable molecular clocks are accurate genes. By using many genes, fluctuations in evolutionary rate
only in the statistical sense of showing a fairly smooth average due to natural selection or other factors that vary over time
NEW! Solve It Tutorials engage students
rate of change. Over time, there may still be deviations from may average out. For example, one group of researchers con-
that average rate. Furthermore, the same gene may evolve at in a multi-step
structed molecular clocks of vertebrate evolution frominvestigation
pub- of a
different rates in different groups of organisms. Finally, when “mystery”
lished sequence data for 658 nuclear or the
genes. Despite openbroadquestion in which
comparing genes that are clocklike, the rate of the clock may period of time covered (nearly 600 million
they years)analyze
must and the factreal data. These are
vary greatly from one gene to another; some genes evolve a that natural selection probably affected some of these genes,
million times faster than others. their estimates of divergence timesassignable
agreed closelyin MasteringBiology.
with fossil- Topics
based estimates. As this exampleinclude:
suggests, if used with care,
Differences in Clock Speed molecular clocks can aid our understanding of evolutionary
What causes such differences in the speed at which clocklike relationships. • Which Biofuel Has the Most Potential to Reduce
genes evolve? The answer stems from the fact that some mu- Our Dependence on Fossil Fuels?
tations are selectively neutral—neither beneficial nor detri-
Applying a Molecular Clock:
mental. Of course, many new mutations are harmful and are
Dating the Origin of HIV• Is It Possible to Treat Bacterial Infections
removed quickly by selection. But if most of the rest are neu- Researchers have used a molecularWithout
clock to date the origin Antibiotics?
Traditional of
tral and have little or no effect on survival and reproduction, HIV infection in humans. Phylogenetic analysis shows that
then the rate of evolution of those neutral mutations should HIV, the virus that causes AIDS,•is Which
descendedInsulin
from Mutations
viruses May Result in Disease?
indeed be regular, like a clock. Differences in the clock rate that infect chimpanzees and other primates. (Most of these
• Are You Getting the Fish You Paid For?
for different genes are a function of how important a gene viruses do not cause AIDS-like diseases in their native hosts.)
is. If the exact sequence of amino acids that a gene speci- When did HIV jump to humans?•There WhyisAre Honeyanswer,
no simple Bees Vanishing?
fies is essential to survival, most of the mutational changes because the virus has spread to humans more than once. The
• What Is Causing Episodes of Muscle Weakness in
will be harmful and only a few will be neutral. As a result, multiple origins of HIV are reflected in the variety of strains
a Patient?
• How Can the Severity of Forest Fires Be

Reduced?
C H A P T E R 2 0 Phylogeny 407
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Keep Current with New Scientific Advances
NEW! The Second Edition incorporates up-to-date content
on genomics, gene editing, human evolution, microbiomes,
climate change, and more.
▼ Figure 3.30
MAKE CONNECTIONS Paleontology

NEW! The Second Edition shows students how New DNA sequencing
techniques have allowed
our ability to sequence DNA and proteins rapidly Contributions of Genomics decoding of minute
and Proteomics to Biology quantities of DNA found
and inexpensively is transforming every subfield Nucleotide sequencing and

in ancient tissues from
our extinct relatives, the
the analysis of large sets Neanderthals (Homo
of biology, from cell biology to physiology to of genes and neanderthalensis).
Sequencing the Neander-
proteins can be done
ecology. For instance, the examples in this figure rapidly and inexpen-
thal genome has informed
our understanding of their
sively due to advances physical appearance as well
from Chapter 3 are explored in greater depth in technology and
information processing.
as their relationship with
modern humans. (See Figure 27.36.)
later in the text. Taken together, genomics
and proteomics have advanced
Medical Science
our understanding of biology
across many different fields. Identifying the genetic basis for human diseases like cancer helps
researchers focus their search for potential future treatments.
Currently, sequencing the sets of genes expressed in an individual’s
Evolution tumor can allow a more
A major aim of evolutionary biology is to understand targeted approach to
the relationships among species, both living and treating the cancer, a
highly successful way for researchers to “knock
extinct. out” (disable)
For example, a
genome sequence comparisons type of “personalized
Cas9 protein Guide RNA engineered to given gene to study what that gene does haveinidentified
an organism. the hippopotamus as the land mammal medicine.” (See
“guide” the Cas9 protein sharing the most recent common ancestor with Concept 9.3 and
to a target gene But what about using this system towhales.help (See
treatFiguregenetic dis-
19.20.) Figure 16.21.)
eases? Researchers have modified the technique so that the
CRISPR-Cas9 system can be used to repair a gene that has a
5′
3′ mutation. They introduce a segment from the normal (func-
Complementary tional) gene along with the CRISPR-Cas9 system. After Cas9
sequence that can cuts the target DNA, repair enzymes can use the normal DNA
Active sites that bind to a target gene
can cut DNA
as a template to repair the target DNA at the break point. In
Species Interactions
this way, the CRISPR-Cas9 system edits the defective gene so
Cas9–guide RNA complex Hippopotamus Most plant species exist in a
that it is corrected (see the bottom right of Figure 13.31, part b). Short-finned pilot whale mutually beneficial partnership
In 2014, a group of researchers reported success in correct- with fungi (right) and bacteria
1 Cas9 protein associated with the plants’ roots;
and guide RNA ing a genetic defect in mice using CRISPR technology. The lab Conservation Biology
these interactions improve plant
are allowed to mice had been genetically engineered to have a mutation in aThe tools of molecular genetics and growth. Genome sequencing
and analysis of gene expression
bind to each other, gene encoding a liver enzyme that metabolizes the amino acidgenomics are increasingly used
by forensic ecologists to identify have allowed characterization of
forming a complex
that is then introduced tyrosine, mimicking a fatal genetic disorder in humans called which species of animals and plant-associated communities.
plants are killed illegally. Such studies will help advance our
into a cell. tyrosinemia. A guide RNA molecule complementary to the In one case, genomic understanding of such interactions
mutated region of the gene was introduced into the mouse sequences of DNA from and may improve agricultural
illegal shipments of practices. (See the Chapter 26
along with the Cas9 protein and a segment of DNA from the elephant tusks were Scientific Skills Exercise and
CYTOPLASM same region of the normal gene for use as a template. Subse- used to track Figure 29.11.)
down poachers
quent analysis indicated that the faulty gene had been corrected and pinpoint
in enough of the liver cells that the amount of functional en- the territory
Cas9 active sites NUCLEUS where they were MAKE CONNECTIONS Considering the examples provided
zyme made was sufficient to alleviate the disease symptoms. operating. (See here, describe how the approaches of genomics and proteomics
Guide RNA There are still many hurdles to overcome before this approach Figure 43.8.) help us to address a variety of biological questions.
complementary
sequence can be tried in humans, but the CRISPR technology is sparking
68 U N I T O N E CHEMISTRY AND CELLS
widespread excitement among researchers and physicians alike.
2 In the nucleus, the 5′ 5′
3′ In this section, you have learned how understanding the
complementary 3′ 5′ elegant structure of DNA has led to powerful techniques for
sequence of the DESIGN SERVICES OF
guide RNA binds to part

genetic engineering. Earlier in the chapter, you also#saw 153397 Cust: Pearson / BC Au: Urry Pg. No. 68
how
URRY2751_02_C03_PR4.indd 68 Title: Campbell Biology in Focus, 2e
C/M/Y/K
Short / Normal / Long
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of the target gene. The DNA molecules are arranged in chromosomes and how DNA
active sites of the Cas9
protein cut the DNA
Part of the
target gene NEW! Chapter 13 describes gene
replication provides the copies of genes that parents pass to
on both strands. offspring. However, it is not enough that genes be copied and
editing using the CRISPR-Cas9
transmitted; the information they carry must be used by the
Resulting cut cell. In other words, genes must also be “expressed.” In the next
in target gene system, and Chapter 17 describes
few chapters, we’ll examine how the cell expresses the genetic
the basic biology of this system
information encoded in DNA. We’ll also return to the subject
of genetic engineering by exploring a few techniques for ana-
in bacteria.
lyzing gene expression.
3 The broken strands
of DNA are “repaired” Normal CONCEPT CHECK 13.4
by the cell in one (functional) 1. MAKE CONNECTIONS The restriction site for an enzyme
of two ways: gene for use
called PvuI is the following sequence:
as a template
5′-c G a T c G-3′
3′-G c T a G c-5′
(a) Scientists can disable
(“knock out”) the target gene
(b) If the target gene has a
mutation, it can be repaired.
NEW! Chapter 27 includes new
staggered cuts are made between the T and c on each strand.
What type of bonds are being cleaved? (see concept 3.6.)
to study its normal function. A normal copy of the gene is
material on human origins,
No template is provided, and provided, and repair 2. DRAW IT one strand of a Dna molecule has the following ▲ Figure 27.36 Fossil evidence
repair enzymes insert and/or
delete random nucleotides,
enzymes use it as a template,
restoring the normal including how sequencing
sequence: 5′-ccTTGacGaTcGTTaccG-3′. Draw the other DNA of human-Neanderthal
making the gene nonfunctional. gene sequence. strand. Will PvuI (see question 1) cut this molecule? if so,
extracted
draw the products. from this fossil jawbone interbreeding.
3. Describe the role of complementary base pairing during
Random nucleotides Normal nucleotides
recently
cloning, Pcr, Dna sequencing,revealed
and gene editingevidence
using the of
human-Neanderthal
crisPr-cas9 system.
For suggested answers, see Appendix A.
interbreeding.
▲ Figure 13.31 Gene editing using the CRISPR-Cas9 system.
CHAPTER 13 The Molecular Basis of inheriTance 275
Keep C urrent with N ew S cienti f ic A dvances xvii
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Focus on the Key Concepts
Each chapter is organized around a framework of 3 to 6 Key Concepts that
focus on the big picture and provide a context for the supporting details.
C H A P T E R
14 Gene Expression: From Gene to Protein
The list of Key Concepts KEY CONCEPTS

14.1 Genes specify proteins via
introduces the big ideas transcription and translation
14.2 Transcription is the DNA-directed
covered in the chapter. synthesis of RNA: a closer look
14.3 Eukaryotic cells modify RNA after
transcription
14.4 Translation is the RNA-directed
synthesis of a polypeptide:
a closer look
14.5 Mutations of one or a few
nucleotides can affect protein
structure and function
Every chapter opens

with a visually dynamic ▲ Figure 14.1 How does a single faulty gene result in the
dramatic appearance of an albino donkey?
photo accompanied by
an intriguing question The Flow of Genetic Information translated by cells into a specific trait, such as brown hair,
that invites students type A blood, or, in the case of an albino donkey, a total lack
T
he island of Asinara lies off the coast of Sardinia, an of pigment? The albino donkey has a faulty version of a key
into the chapter. Italian island. The name Asinara probably originated protein, an enzyme required for pigment synthesis, and this
from the Latin word sinuaria, which means “sinus- protein is faulty because the gene that codes for it contains
shaped.” A second meaning of Asinara is “donkey-inhabited,” incorrect information.
which is particularly appropriate because Asinara is home to a This example illustrates the main point of this chapter:
wild population of albino donkeys (Figure 14.1). The donkeys The DNA inherited by an organism leads to specific traits
were brought to Asinara in the early 1800s and abandoned by dictating the synthesis of proteins and of RNA molecules
there in 1885 when the 500 residents were forced to leave the involved in protein synthesis. In other words, proteins are the
island so it could be used as a penal colony. What is responsi- link between genotype and phenotype. Gene expression is
ble for the phenotype of the albino donkey, strikingly different the process by which DNA directs the synthesis of proteins
from its pigmented relative? (or, in some cases, just RNAs). The expression of genes that
Inherited traits are determined by genes, and the trait code for proteins includes two stages: transcription and
of albinism is caused by a recessive allele of a pigmenta- translation. This chapter describes the flow of information
tion gene (see Concept 11.4). The information content of from gene to protein and explains how genetic mutations
genes is in the form of specific sequences of nucleotides affect organisms through their proteins. Understanding the
along strands of DNA, the genetic material. But how does processes of gene expression, which are similar in all three
this information determine an organism’s traits? Put another domains of life, will allow us to revisit the concept of the gene
way, what does a gene actually say? And how is its message in more detail at the end of the chapter.
278
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After reading a Key Concept section,

disastrous effect on the resulting protein more often than sub- CONCEPT CHECK 14.5
students can check their understanding
stitutions do. Insertion or deletion of nucleotides may alter the 1. What happens when one nucleotide pair is lost from the
reading frameusing
of the the Concept
genetic message,Check questions:
the triplet grouping of middle of the coding sequence of a gene?
nucleotides on the mRNA that is questions
read duringasktranslation. 2. MAKE CONNECTIONS Individuals heterozygous for the sickle-
Make Connections students Such
cell allele show effects of the allele under some circumstances
a mutation, called a frameshift
to relate content inmutation, occurs
the chapter whenever
to material
(see Concept 11.4). Explain in terms of gene expression.
the number of nucleotides
presented inserted
earlier in theor course.
deleted is not a multiple
3. WHAT IF? DRAW IT The template strand of a gene includes
of three. All nucleotides downstream
What if? questions of the deletion
ask students or inser-
to apply what this sequence: 3′-TACTTGTCCGATATC-5′. It is mutated to
tion will be improperly grouped
they’ve learned. into codons; the result will be 3′-TACTTGTCCAATATC-5′. For both versions, draw the DNA,
extensive missense mutations, usually ending sooner or later the mRNA, and the encoded amino acid sequence. What is
Draw It Exercises ask students to put pencil
in a nonsensetomutation
paper andanddraw
premature termination.
a structure, Unless
annotate a the effect on the amino acid sequence?
the frameshift is veryornear
figure, the experimental
graph end of the gene, the protein is
data. For suggested answers, see Appendix A.
almost certain to be nonfunctional.
New Mutations and Mutagens

Focuscan
xviiiMutations onarise
the in
Ke a
y number of ways. Errors during DNA
C oncepts What Is a Gene? Revisiting the Question
replication or recombination can lead to nucleotide-pair Our definition of a gene has evolved over the past few chap-
substitutions, insertions, or deletions, as well as to mutations ters, as it has through the history of genetics. We began with
affecting longer stretches of DNA. If an incorrect nucleotide the Mendelian concept of a gene as a discrete unit of inheri-
is added to a growing chain during replication, for example, tance that affects a phenotypic character (Chapter 11). We saw
the base on that nucleotide will then be mismatched with the that Morgan and his colleagues assigned such genes to specific
nucleotide base on the other strand. In many cases, the error loci on chromosomes (Chapter 12). We went on to view a gene
The Summary of Key Concepts refocuses
14
Go to for Assignments, the eText, and the Study Area
What will be the results of with
chemically modifying
Activities,one nucleotide 9. DRAW IT Fill in the fo
chapter. ?
Animations, Vocab Self-Quiz, and practice Tests.
students on the main points
Chapterof theReview base of a gene? What role is played by DNA repair systems in
the cell? Type of RNA
VOCAB NEW! QR Codes and URLs PRACTICE Messenger RNA (mRNA)
SUMMARY OF KEY CONCEPTSTEST YOUR UNDERSTANDING
SELF-QUIZ • Eukaryotic
at the pre-mRNAs
end of every chapter
TEST RNA processing, which
undergo
14
Go to for Assignments, the eText, and the Study Area
Chapter Review
with Animations, Activities, Vocab Self-Quiz, and practice Tests.
includesquick
give students RNAaccess
splicing, the addition of a modified nucleotide
Transfer RNA (tRNA)
three times the number of aminoSUMMARY
acids in OF theKEY protein
CONCEPTS product. VOCAB
SELF-QUIZ • Eukaryotic pre-mRNAs undergo RNA processing, which Second mRNA base
Level 1: Knowledge/Comprehension
5′ cap toSelf-Quizzes
to Vocabulary the 5′ end, and the addition of a poly-A tail to the
CONCEPT 14.1 includes RNA splicing, the addition of a modified nucleotide
and Practice 3′ end.Tests The processed their mRNA includes an untranslated region
For example, it takes 300 nucleotides along an mRNA strand
CONCEPT 14.1
U
1. 5′ cap to the 5′ end, and the addition of a poly-A tail to the G
In eukaryotic C A
cells, transcription cannoton begin
(5′ UTRtablets, or 3′ UTR) and at each end of the coding segment.
3′ end. The processed mRNA includes an untranslated region
to code for the amino acids in a polypeptide
Genes specify proteins that Genes
via is 100 amino
transcription
specify proteins
UUU via UCU
transcription
UAU goo.gl/gbai8v
(5′ UTR or 3′ UTR) at each end of the coding segment.
until Cys
UGU
• Most eukaryotic genes are split into segments: They have introns U smartphones,
introns
and translation (pp. 279–284)
acids long. and translation (pp.UUC
phe
279–284)
tyr interspersed among the exons (regions included in the mRNA).
goo.gl/gbai8v computers. • Most eukaryotic genes aregoo.gl/CRZjvS split into segments:Primary They have transcript
(A) the two DNA strands have completely
• Beadle and Tatum’s studies of mutant strains of Neurospora led to
the one gene–one polypeptide hypothesis. During gene expression, UCC UAC UGC
In RNA splicing, introns are removed and exons joined. RNA C
U
interspersed among the exons (regions included in the mRNA).
splicing is typically carried out by spliceosomes, but in some
the information encoded in genes is used to make specific polypep- ser
cases, RNA alone catalyzes its own splicing. The catalytic
UAA stop UGAseparated
• Transcription is the synthesis•of RNA Beadle and Tatum’s studies UUA
of mutant UCA
strains of Neurospora stop A
led toand exposed the promoter.
tide chains (enzymes and other proteins) or RNA molecules.
Cracking the Code complementary to a Leu
ability of some RNA molecules, called ribozymes, derives
from the properties of RNA. The presence of introns allows for
In RNA splicing, introns are removed and exons Small RNAs in
joined. RNAthe spliceosom
UAG stop (B) UGGseveral trp Gtranscription factors have bound to
template strand of DNA. Translation is the synthesis of a alternative RNA splicing.
Molecular biologists cracked the geneticnucleotide sequence the
of life in the
in mRNA. one
polypeptide whose amino acid sequence is specified by the
code gene–one ?
UUG
polypeptide UCG
hypothesis. During gene expression,
What function do the 5′ cap and the poly-A tail serve on
splicing is typically carried out by spliceosomes, but in some
Third mRNA base (3′ end of codon)

CGUthe promoter.
First mRNA base (5′ end of codon)
a eukaryotic mRNA?
the
A codoninformation encoded CUU in genes is used to make specific polypep-
• Genetic information is encoded as a sequence of nonoverlap-
early 1960s when a series of elegant ping experiments disclosed alone catalyzes its own splicing. Level
nucleotide triplets, or codons. in messenger
RNA (mRNA) either is translated into an amino acid (61 of the
CCU
CONCEPT 14.4
CAU
His (C) the ?
5′
U
caps are removed from the mRNA.
cases, RNA 9. Fill in the following table:
What will be the results of chemically modifying one nucleotide
base of a gene? What role is played by DNA repair systems in The catalytic3: Synthesis/Ev
the amino acid translations of each of
64 codons)the RNA
or serves codons.
as a stop
tide chains (enzymes
The
signal (3 codons).
be read in the correct reading frame.
Codons must
Translationand CUC other proteins)
is the RNA-directed CCC
synthesis or
CAC RNA molecules.CGC C ability
the cell?
of some RNA molecules, called ribozymes, derives Type of RNA Functions
C (D) the DNA introns are removed from the template.
from the properties of RNA. The presence of10.
Leu
• Transcription
of a polypeptide: a closer look (pp. 288–298) pro
first codon was deciphered in 1961? by Marshall Nirenberg, of •is the synthesis ofprotein
RNA complementary Arg
to aAUNDERSTANDING
PRACTICE Messenger RnA (mRnA)
SCIENTIFIC INQUIRY
CGATEST YOUR
Describe the process of gene expression, by which a gene affects
introns allows for
TEST
the phenotype of an organism. A cell CUA
translates an mRNA messageCCA into CAA
using transfer
transfer RnA (tRnA)
the National Institutes of Health, along with his colleagues. template strandaminoacyl-tRNA
RNAs (tRNAs). After being bound to a specific amino acid by an
of DNA. Translation isup oftheGln synthesis Level 1: Knowledge/Comprehension
of athe Knowing that the gene
CONCEPT 14.2 CUG codon
at the complementary
synthetase, a tRNA lines up via its anticodon
CCG
on mRNA. A ribosome, madeCAG 2. Which
CGG1. In eukaryotic following
Gcells, transcription cannot beginis not true of a codon?
alternative RNA splicing.
Plays catalytic (ribozyme) roles and
structural roles in ribosomes
Nirenberg synthesized an artificial mRNA by linking identical uses molecular biologi
polypeptide whose
Transcription is the DNA-directed synthesis of RNA:
a closer look (pp. 284–286)
ribosomal
amino
with binding
RNAs (rRNAs)
sites for mRNAacidand
sequence is specified
proteins,
and tRNA.
facilitates this coupling
(A) by
until
It(A)maythe
the two DNA code forcompletely
strands have the same amino acid as another codon. goo.gl/CRZjvS Primary transcript
RNA nucleotides containing uracil as their base. No matter • Ribosomes AUU ACU AAU AGU (B) separated Uand exposed the promoter. What function do the 5′ cap and the poly-A tail serve on
mRNA. gene (shown in Figure
coordinate the three stages of translation: initiation,
nucleotide
to a DNA template sequence
elongation, in
small RnAs in the spliceosome
(B) It never
the promoter.codes for more than ? a eukaryotic
one amino acid.
• RNA synthesis is catalyzed by RNA polymerase, which links and termination. The formation of peptide bonds several transcription factors have bound to
where this message started or stopped, Asn ser
strand. Thisit could followscontain onlyrules as DNA
together RNA nucleotides complementary
process
• Genetic
the same base-pairing
information
between amino
move through AUC
is
acids is
encoded
catalyzed
the A andIle
by
P sites and
ribosomal
ACC
as a
RNAs as
sequence
exit through the E site.
tRNAs
AAC
of AGC (C) the 5′ caps
nonoverlap- C are removed from the mRNA. mRNA? cells will express it and
replication, except that in RNA,
one codon in repetition: UUU. Nirenberg added this “poly-U”
uracil substitutes for thymine. • AfterA translation, modifications to proteins can affectthr their shape. (C) It (D)extends
the DNA introns are from one
removed from end
the template. of a tRNA
Level 3:
molecule.
Synthesis/Evaluation
codons. Instead, the protein pro

Free ribosomes in the cytosol initiate synthesis of all proteins, but 10.
AUA of theA
ping nucleotide triplets, orpeptide ACA A codon AAA in messenger AGA2. Which
Transcription unit SCIENTIFIC INQUIRY
following is not true of a codon?
Lys (D) It(A) isIt maythe basic unit acid asof the codon.genetic Knowing code.
proteins with a signal
codon to a test-tube mixture that contained all 20 amino Promoter are synthesized on the ER. Arg code for the same amino
(B) It never codes for more than one amino acid.
another
uses
that the genetic code is almost universal, a scientist
molecular biological methods to insert the human β-globin
contain many fewer am
Polypeptide
metbyor
acids, ribosomes, and the other components required RNA for (mRNA) either
• A gene can AUG
ispolymerases
translated into an amino acid (61
be transcribed
start ACG AAG AGG of the
(C) It extends G from one end of a tRNA molecule.
Amino
CONCEPT
gene (shown in Figure 14.12) into bacterial cells, hoping the
14.4
3. The 3.anticodon of a particular tRNA Instead, molecule is is nonfunctional and is found to
5′ 3′ multiple RNA 3′ acid
(D) It is the basic unit of the genetic code. cells will express it and synthesize functional β-globin protein.
3′ 5′ tRNA
protein synthesis. His artificial system translated the64 5′

poly-Ucodons) or serves
simultaneously. Also, a single
as a
Template strand
mRNA molecule can stop
be trans-
GUU by a num- GCU signal (3 codons).
GAU Codons must
The anticodon of a particular tRNA molecule is
GGU (A) complementary U to the corresponding mRNA codon.
the protein produced
contain many fewer amino acids than does β-globin made by a eukaryotic cell. Explain
(A) complementary totriplet
theinTranslation
corresponding is mRNA the RNA-directed codon. synthesis
RNA polymerase of DNA
lated simultaneously eukaryotic cell. Explain why.
arebe read in the correct reading frame. GAC Asp GGC (B)(C)
RNA transcript complementary to the corresponding rRNA.
into a polypeptide containing many unitsstagesof the amino acid ber of ribosomes, forming a
GUC In bacteria, these GCC
E
C
A Anti-
the part of tRNA that bonds with a specific amino acid. 11. FOCUS ON EVOLUTION
11. FOCUS ON EVOLUTIO
(B) complementary toprocessing?
theofcorresponding triplet ain rRNA.
• The three of transcription initiation, elongation, polyribosome.
a polypeptide: closer can you look (pp. 288–298)
codon Most amino acids are coded for by a set of similar codons (see
phenylalanine (Phe), strung together
and termination. A promoter, often including a TATA box
as a establishes
polyphenylalanine G are coupled, but
processes
Val in
Ala
(D) catalytic, making the tRNA a ribozyme.
Glyof the following is not true of RNA Figure 14.6). What evolutionary explanation give for
in eukaryotes, where RNA synthesis is initiated.
Transcription factors help eukaryotic RNADescribe
eukaryotes they are separated
polymerase recog- the process
in time andGUAspaceofby thegene
nuclear expression, by which a
4. Which
gene affects
Codon
Most amino acids are c
Glu (C) the part of tRNA that bonds with a specific amino acid.
GCA GAA GGA A mRNA this pattern?
?andthe
(A) Exons are cut out before mRNA leaves the nucleus.
chain. Thus, Nirenberg determined that the mRNA forming a codon
using transfer
12.
function in RNA splicing. • A cell translates anthe unity
mRNA message into protein Figure
nize promoter sequences, transcription initiation FOCUS ON INFORMATION
14.6). What evo

membrane. Ribosome
(B) Nucleotides may be added at both ends of the RNA.
complex. Termination differs in bacteria
UUU specifies the amino acid phenylalanine. Soon, the amino phenotype
eukaryotes.
? of anGUG organism. GCG GAG (D)
GGGcatalytic,
(C) Ribozymes G maycanmaking
What function do tRNAs serve in the process of translation?
the tRNA a
Evolution accounts for
ribozyme.
and diversity of life, and the
continuity of life is based on heritable information in the form
RNAs (tRNAs).
(D) RNA splicing be catalyzed by spliceosomes.
Afteris inheritedbeing bound to a specific amino acid by(Hint:
an Th
What are the similarities and differences in the initiation
?
acids specified by the codons AAA, GGG, and CCC were
of gene transcription in bacteria and eukaryotes? 5. Which component is not directly involved in translation?
of DNA. In a short essay (100–150 words), discuss how the
fidelity with which DNA is related to the processes this pattern?
CONCEPT 14.3 ▲Mutations
Figure
CONCEPT 14.5
14.6 The codon table for mRNA. 4. Which
the three ofnucleotide
(A) GTP
(B) DNA the following (C) tRNA
is not
(D) ribosomes true ofofrepair
aminoacyl-tRNA RNA processing?
13.2.) synthetase, a tRNA lines up via its anticodon
evolution. (Review the discussion of proofreading and DNA
in Concept
try, and some less obvi
determined in the same way. Eukaryotic cells modify RNA after transcription of one or a few nucleotides can affect
designated here as(A)
CONCEPT 14.2 bases proteinof an mrnA
structure and functioncodon (pp.are298–300) Exons
the first,
Level second,
2: areandcut out beforeatmRNA
Application/Analysis 13.
leaves the nucleus.
the complementary codon on mRNA. A ribosome,
SYNTHESIZE YOUR KNOWLEDGE
made up of
Although more elaborate techniques(pp. 286–288)
were required to bases, reading in the 5′ → 3′ direction along the 6.mrnA.
• Small-scale mutations include point mutations, changes in
third
(B) Nucleotides
(practice
14.6, identify a 5′ → 3′ sequence of nucleotides in
the DNA template strand formay
Using Figure
an mRNA be coding added
for the polypep-at both ends of the RNA.
12. FOCUS ON INFORMAT
ribosomal RNAs (rRNAs) and proteins, facilitates this coupling
5′ Cap Poly-A tail
one DNA nucleotide pair, which may lead to production of non-
decode mixed triplets such as AUA and CGA, allTranscription 64 codons is
using the DNA-directed
this table by finding the codons synthesis
in Figure 14.5.)of the tideRNA:
codon AUG
5′ Exon Intron Exon Intron Exon 3′
Pre-mRNA functional proteins. Nucleotide-pair substitutions can cause
(C)but
sequence Phe-Pro-Lys.
Ribozymes Evolution accounts for
(B) 5′-GAACCCCTT-3′ may function in RNA splicing.
missense or nonsense mutations. Nucleotide-pair insertions
with binding sites for mRNA and tRNA.
(A) 5′-UUUCCCAAA-3′
notor only stands forframeshift
the amino acid methionine (met) also functions
were deciphered by the mid-1960s. As Figure 14.6
RNA splicing
a shows,
deletions may produce mutations.
closer look (pp.
mRNA
284–286)
as• Spontaneous
a “start” mutations can occur during DNA replication, re-
signal for ribosomes to begin translating
(D) RNAthe (C)mrnA at
splicing
5′-CTTCGGGAA-3′
can be catalyzed by spliceosomes. continuity of life is bas
61 of the 64 triplets code for amino acids. The three codons 5′ UTR Coding
segment that
combination, or repair. Chemical and physical mutagens cause
3′ UTR
DNA point.
damage thatthree
can alterofgenes.
the 64 codons function as “stop” 7.signals,
(D) 5′-AAACCCUUU-3′
marking • Ribosomes coordinate the three stages of translation: of DNA. initiation,
In a short ess
that do not designate amino acids are “stop” signals, • RNA or ter- synthesiswhere is catalyzed RNA polymerase,
by translation. which links
Which of the following mutations would be most likely to have a
elongation, andareresulttermination.
in proteins that function wellThe at one formation of peptide bonds
ribosomes end see Figure 3.18 for
5. a list of the onfull
Which (A) acomponent
deletion of three nucleotides is not directly involved
temperature butin translation?
harmful effect an organism? 301
fidelity with which DN
CHAPTER 14 GENE ExpRESSIoN: FRoM GENE To pRoTEIN
Some mutations
near the middle
mination codons, marking the end of translation. Notice together that RNA names nucleotides of all the amino complementaryacids. to a DNA template of a gene
CONCEPT between 14.2 amino
nonfunctional
in a gene acids
at a different (usually higher)
is catalyzed
that makes dark pig- by ribosomal RNAs as tRNAs
(A) GTP (C) tRNA
temperature. Siamese cats have such a “temperature-sensitive”
of evolution. (Review t
(B) a single nucleotide deletion in the middle of an intron
mutation encoding an enzyme
the codon AUG has a dual function: It codes for the strand. amino This process follows the same base-pairing rulessequence as DNA
(C) a single nucleotide deletion near the end of the coding
move through
point markings the Abodylearned
andcolor (seeP sites Usingand exit through the E site.
ment in the fur. The mutation results in the breed’s distinctive
(B) DNA Transcription (D) ribosomes
is the what youDNA-directed
DESIGN SERVICES OF
and lighter the photo).
repair in Concept 13.2
S4CARLISLE (D) a single nucleotide insertion downstream of, and close to,
acid methionine (Met) and • Salso
ummaryfunctions ofasKey a “start” replication,
Concepts signal, questions exceptcheck
Title: Campbell Biology in Focus, 2e
that in RNA, uracil substitutes for8.thymine.
students’
Short / Normal / Long this information and in the chapter, explain
URRY2751_02_C14_PR3.indd 301 Publishing Services 10/07/15 5:36 pm
• 14.23After translation, modifications to proteins can affect their shape.

the start of the coding sequence
the pattern of the cat’s fur pigmentation.
or initiation codon. Genetic messages usually begin with the

understanding of a key idea from each concept.Transcription unit probably be gibberish: for example, “her edd oga tet
foundheb ug.
in a eukaryotic” cell? Explain why orsynthesis
Would the coupling of the processes shown in Figure
why not.
be
of RNA: a
Free ribosomes in the cytosol initiate synthesis
closer look 13.of SYNTH
all proteins,
For selected answers, see Appendix A.
ESI ZEbut Y OUR K
mRNA codon AUG, which signals the protein-synthesizing The reading frame is also important inLevel 2: Application/Analysis
the molecular language
• Summary figures
machinery to begin translating the mRNA at that location. recap key information visually.
of cells. The short stretch of polypeptide shown in Figure 14.5, Now that proteins
we have with
considered a signalthe peptide
linguistic logic and evolution-
(Because AUG also stands for methionine, polypeptide chains Promoter for instance, will be made correctly only6.if the UsingmRNA Figure nucleo- 14.6, identify aare
ary significance 5′ → 3′
synthesized
of thesequencegenetic oncode, of
thenucleotides
weER. are ready toinreexamine Polypeptide
NEW!
the DNASynthesize template Your Knowledge
strand for an questions
mRNA coding ask students
for the toinapply
polypep- their
begin with methionine when they are synthesized. However, tides are read from left to right (5′ → 3′) in the groups of three •
transcription, A gene the can
first be
stage transcribed
of gene expression, by more detail.
Amino
understanding
tide sequence of the chapter content to explain an intriguing photo.
an enzyme may subsequently remove this starter amino 5′ acid shown in the figure: UGG UUU 3′ GGC UCA. Although a ge-3′Phe-Pro-Lys. multiple RNA polymerases acid
3′ 5′ Molecular
302
Components
UNIT TWO GenetiCs
ofaTranscription tRNA
from the chain.) netic message is written with no spaces between (A) 5′-UUUCCCAAA-3′
the codons, simultaneously. Also, single
Notice in Figure 14.6 that Evolution,
there is redundancy the in the geneticfundamental 5′
the cell’stheme protein-synthesizing of biology, machinery Template (B) the
reads 5′-GAACCCCTT-3′
strand
message as Messenger mRNA RNA, the moleculecarrier of can information be trans- from DNA to the
of DNA CONCEPT 14.2
DESIGN SERVICES OF
code, but no ambiguity. For example, although codons GAA a series of nonoverlapping RNA polymerase three-letter words. (C)The message is
5′-CTTCGGGAA-3′ cell’s protein-synthesizing
lated machinery,
# 153397 Cust: Pearson / BC Au: Urry
simultaneously is a
S4CARLISLE
by transcribed
Pg. No. 302
num- from the tem-

C/M/Y/K
is emphasized throughout. Foras aexample:

URRY2751_02_C14_PR5.indd 302 Title: Campbell Biology in Focus, 2e Short / Normal / Long Publishing Services 31/07/15 5:38 PM
and GAG both specify glutamic acid (redundancy), neither of RNA transcript
not read series of overlapping words—UGGUUU, and so
(D) 5′-AAACCCUUU-3′ ber of ribosomes, forming a plate strand of a gene. An enzyme called an RNA polymerase Transcription is t
them ever specifies any other amino acid (no ambiguity). The on—which would convey a very different message. pries the two strands of DNA apart and joins together RNA E A Anti-
• The three stages of transcription are initiation, elongation, polyribosome. In bacteria, these synthesis of RNA
(b) Pig7. Whicha of the following mutations would betomost likely to have a thus
redundancy in the code is not• altogether
Every Chapter random.Review In many (a) Tobacco plant expressing a nucleotides complementary the DNA template strand, codon
expressing jellyfish
includes a “Focus
andonacid
termination. A
Evolution promoter,
firefly gene. of the theyellow often
Genetic including
glow Codegene. Researchers a TATA
harmfulinjected box effecttheon an organism? processes are coupled,
elongating the RNA polynucleotide (Figure 14.8). Like the DNA but in Now that we have considere
cases, codons that are synonyms for a particular amino
in eukaryotes, establishes
is produced by wherea chemical RNA synthesis gene for isainitiated.
fluorescent protein eukaryotes they are separated Some Codon mutations of theresult
differ only in the third nucleotide Evolution”
base of the triplet. question We’ll con- EVOLUTION
reaction catalyzed The genetic by the code is nearly (A)
universal,
into fertilized apigdeletion
shared
eggs. One by of threepolymerases nucleotides that near
function the middle
in DNA replication, RNA polymerases ary significance geneti
sider the significance of this redundancy
(shown later above in the Transcription
chapter.
right). factors
organisms
protein product help
from the eukaryotic
of thesimplest
firefly bacteriaoftoRNA polymerase
thethe most
eggs ofcomplex
developed a gene recog-
into can assemble in time and spaceonly
a polynucleotide by in the its nuclear
5′ → 3′ direction. Unlike temperature
transcription, mRNA but
the firstare noo
stage
Our ability to extract the intended message from nize a writ-promoter plants sequences,
gene. and animals. forming The mRNA a transcriptioncodon thisCCG, initiation
fluorescent
for pig.
instance, is
(B) a single nucleotide deletion in the middle of an intronDNA membrane.
polymerases, however, RNA polymerases are able toRibosome
start a temperature. Siamese c
• Every chapter has a Molecular in Componen
ten language depends on reading the symbols in the complex. correct Termination
translated
▲ Figure 14.7 differs
as the amino
Expression in bacteria
acidofprolinegenes and in eukaryotes.
all
from organisms
different species.
(C) code,a single whosenucleotidechain deletionfrom scratch; near they the don’t
endneed ofdothe a primer.
coding
mutation a gene enc
section explicitly relating What function tRNAs serve in the process ofment translation?
groupings—that is, in the correct reading frame. Consider
What
Because diverse forms of life share
genetic code has been examined. In laboratory experiments,
are the
cansimilarities
be programmedand differences
a common genetic
in the initiation sequence
one species Specific ? sequences of nucleotides along the DNA mark where in the
Messenger RNA,fur.
theThe
carriermo
this statement: “The red dog atethe the chapter content to to produce proteins characteristic of a second
evolution
bug.” Group
(shown
?
the letters
atof
genes can
gene transcription
right). species by be transcribed
in bacteria and eukaryotes?
introducing DnA and the
from translated
second after being
species into trans-
(D) a single nucleotidewhere
the first.
transcription of a gene begins and ends. The DNA sequence
insertion downstream of, and close to,
point markings and lig
cell’s protein-synthesizing
plateinformation
strand of a gene.and
ma
An enz
incorrectly by starting at the wrong point, and the result will planted from one species to another, sometimes with quite RNA polymerase attaches and initiates transcription is this w
striking results, as shown in Figure 14.7. Bacteriathe canstart be of the coding knownsequence as the promoter; in bacteria, the sequence that signals
CONCEPT 14.5 the
pries the two strands of DNA
pattern of the cat’s
(a) Tobacco plant expressing the (b) nucleotides complementary to
CONCEPT 14.3 programmed by the insertion of human8.genes Wouldto synthesizethe coupling ofthe the end of transcription
processes
firefly gene. the shown is calledinaFigure Pig expressing
terminator.
14.23 be(Thea termi-
jellyfish
certain human proteins for medical use, such as insulin.

found in a Such
eukaryotic nation
cell? Mutations
mechanism
Explain
is producedwhy
yellow glow
of
byisa different
or
chemical one
why in not.or agene.
eukaryotes; few
gene for
researchers
anucleotides
we’ll
injected the
describe protein
fluorescent it can
For
elongating
affect
selected
the RNA polynucle
answers, see App
Eukaryotic cells modify RNA after transcription
C H A P T E R 1 4 Gene expression: From Gene to protein 283 polymerases that function in D
later.)protein structure
of the firefly and of function (pp.into298–300)
applications have produced many exciting developments in the reaction catalyzed
Molecular by the
biologists into fertilized
refer to the direction pig eggs. One
of transcription
protein product the eggs developed can assemble a polynucleotide
(pp. 286–288) area of genetic engineering (see Concept 13.4). as “downstream”
•
gene.
Small-scale
and the other directionthis
mutations
asfluorescent
include
“upstream.
point
” These
pig.
mutations, changes
DNA in however,
polymerases,
Despite a small number of exceptions in which a few co- terms are also used to describe the positions of nucleotide se-
5′ Cap Poly-A tail ▲ Figure 14.7 Expression of genes from different species.
one DNA nucleotide pair, which may lead to chain fromofscratch;
production non-they don’t
dons
5′ Exon differExon
Intron from the standard ones, the evolutionary signifi-
Intron quences within
Because theforms
diverse DNA of or
life RNA.
share aThus,
common thegenetic
promotercode,sequence
one species Specific sequences of nucl
DESIGN SERVICES OF Exon 3′
# 153397 Cust: Pearson / BC Au: Urry Pg. No. 283
Pre-mRNA cance of C/M/Y/K
the code’s near S4CARLISLE
universality is clear. A language shared in DNA
can functional
is
be said to
programmedbe proteins.
upstream
to produce fromNucleotide-pair
the
proteins terminator.
characteristic ofThe
a substitutions
stretch
second can cause
transcription of a gene begin
of DNAmissense nonsense mutations. insertions
31/07/15 5:38 PM species by introducing DnA from the second species into the first.
downstreamor Nucleotide-pair
RY2751_02_C14_PR5.indd 283 Publishing Services
by all living things must have been operating very early in the from the promoter that is transcribed into where RNA polymerase attac
RNA splicing
history of life—early enough to be present in the common an- an RNA or deletions
molecule
striking results,isascalled
shownmay Figure 14.7frameshift
a transcription
in produce Bacteria can bemutations.known as the promoter; in b
. unit.
cestormRNA
of all present-day organisms. A shared genetic vocabu- Bacteria have abysingle
• Spontaneous type of RNA
mutations polymerase
cangenesoccur that synthe-
during DNA
programmed the insertion
Focus of onhuman
the Ke y Ctooncepts
synthesize xixreplication, re-
the end of transcription is cal
lary is a reminder of the kinship that bonds all life on Earth. sizescertain
notcombination,
only mRNA
human butor
proteins also other types
forrepair.
medical use, of
Chemical RNA
such that func-
and
as insulin. physical
Such mutagens
nation mechanism
cause is differen
5′ UTR Coding 3′ UTR tionapplications
in protein synthesis, such many
segment DNA damage have produced thatascan ribosomal
exciting RNA.
alter genes. In contrast,
developments in the later.) Molecular biologists re
CONCEPT CHECK 14.1 eukaryotes have atengineering
area of genetic least three types of RNA13.4).
(see Concept polymerase in as “downstream” and the oth
1. MAKE CONNECTIONS in a research article about alkaptonuria their nuclei;
Despite thea small
one used number for pre-mRNA
of exceptionssynthesis
in whichisa called
few co- terms are also used to describ
published in 1902, Garrod suggested that humans inherit two dons differ from C H A P T E R 1 4 GENE ExpRESSIoN: FRoM GENE To pRoTEIN
the standard ones, the evolutionary signifi- quences within the301DNA or R
302 U N I T T W O GENETICS
RNA polymerase II. In the discussion that follows, we start
“characters” (alleles) for a particular enzyme and that both withcance of the code’s
the features of mRNA near universality is clear. Atolanguage
synthesis common shared
both bacteria in DNA is said to be upstream
parents must contribute a faulty version for the offspring to and by all living things
eukaryotes and then must have been
describe someoperating very early in the
key differences. of DNA downstream from th
URRY2751_02_FM_FINAL.indd 19 have the disorder. today, would this disorder be called domi- 03/09/15 5:58 PM
history of life—early enough to be present in the common an- an RNA molecule is called a
Exploring the Variety of Random
Documents with Different Content
12. EXTENSION OF KRONECKER'S
THEOREM ON ABELIAN FIELDS TO
ANY ALGEBRAIC REALM OF
RATIONALITY.
The theorem that every abelian number field arises from the
realm of rational numbers by the composition of fields of roots of
unity is due to Kronecker. This fundamental theorem in the theory of
integral equations contains two statements, namely:
First. It answers the question as to the number and existence of
those equations which have a given degree, a given abelian group
and a given discriminant with respect to the realm of rational
numbers.
Second. It states that the roots of such equations form a realm
of algebraic numbers which coincides with the realm obtained by
assigning to the argument in the exponential function all
rational numerical values in succession.
The first statement is concerned with the question of the
determination of certain algebraic numbers by their groups and their
branching. This question corresponds, therefore, to the known
problem of the determination of algebraic functions corresponding to
given Riemann surfaces. The second statement furnishes the
required numbers by transcendental means, namely, by the
exponential function .
Since the realm of the imaginary quadratic number fields is the
simplest after the realm of rational numbers, the problem arises, to
extend Kronecker's theorem to this case. Kronecker himself has
made the assertion that the abelian equations in the realm of a
quadratic field are given by the equations of transformation of elliptic
functions with singular moduli, so that the elliptic function assumes
here the same rôle as the exponential function in the former case.
The proof of Kronecker's conjecture has not yet been furnished; but I
believe that it must be obtainable without very great difficulty on the
basis of the theory of complex multiplication developed by H.
Weber[26] with the help of the purely arithmetical theorems on class
fields which I have established.
Finally, the extension of Kronecker's theorem to the case that, in
place of the realm of rational numbers or of the imaginary quadratic
field, any algebraic field whatever is laid down as realm of rationality,
seems to me of the greatest importance. I regard this problem as
one of the most profound and far-reaching in the theory of numbers
and of functions.
The problem is found to be accessible from many standpoints. I
regard as the most important key to the arithmetical part of this
problem the general law of reciprocity for residues of th powers
within any given number field.
As to the function-theoretical part of the problem, the
investigator in this attractive region will be guided by the remarkable
analogies which are noticeable between the theory of algebraic
functions of one variable and the theory of algebraic numbers.
Hensel[27] has proposed and investigated the analogue in the theory
of algebraic numbers to the development in power series of an
algebraic function; and Landsberg[28] has treated the analogue of
the Riemann-Roch theorem. The analogy between the deficiency of
a Riemann surface and that of the class number of a field of
numbers is also evident. Consider a Riemann surface of deficiency
(to touch on the simplest case only) and on the other hand a
number field of class . To the proof of the existence of an
integral everywhere finite on the Riemann surface, corresponds the
proof of the existence of an integer in the number field such that
the number represents a quadratic field, relatively unbranched
with respect to the fundamental field. In the theory of algebraic
functions, the method of boundary values (Randwerthaufgabe)
serves, as is well known, for the proof of Riemann's existence
theorem. In the theory of number fields also, the proof of the
existence of just this number offers the greatest difficulty. This
proof succeeds with indispensable assistance from the theorem that
in the number field there are always prime ideals corresponding to
given residual properties. This latter fact is therefore the analogue in
number theory to the problem of boundary values.
The equation of Abel's theorem in the theory of algebraic
functions expresses, as is well known, the necessary and sufficient
condition that the points in question on the Riemann surface are the
zero points of an algebraic function belonging to the surface. The
exact analogue of Abel's theorem, in the theory of the number field
of class , is the equation of the law of quadratic reciprocity[29]
which declares that the ideal is then and only then a principal ideal
of the number field when the quadratic residue of the number with
respect to the ideal is positive.
It will be seen that in the problem just sketched the three
fundamental branches of mathematics, number theory, algebra and
function theory, come into closest touch with one another, and I am
certain that the theory of analytical functions of several variables in
particular would be notably enriched if one should succeed in finding
and discussing those functions which play the part for any algebraic
number field corresponding to that of the exponential function in the
field of rational numbers and of the elliptic modular functions in the
imaginary quadratic number field.
Passing to algebra, I shall mention a problem from the theory of
equations and one to which the theory of algebraic invariants has led
me.
[26] Elliptische Funktionen und algebraische Zahlen.
Braunschweig, 1891.
[27] Jahresber. d. Deutschen Math-Vereinigung, vol. 6, and an
article soon to appear in the Math. Annalen [Vol. 55, p. 301]:
"Ueber die Entwickelung der algebraischen Zahlen in
Potenzreihen."
[28] Math. Annalen vol. 50 (1898).
[29] Cf. Hilbert, "Ueber die Theorie der relativ-Abelschen
Zahlkörper," Gött. Nachrichten, 1898.
13. IMPOSSIBILITY OF THE
SOLUTION OF THE GENERAL
EQUATION OF THE 7TH DEGREE BY
MEANS OF FUNCTIONS OF ONLY
TWO ARGUMENTS.
Nomography[30] deals with the problem: to solve equations by
means of drawings of families of curves depending on an arbitrary
parameter. It is seen at once that every root of an equation whose
coefficients depend upon only two parameters, that is, every function
of two independent variables, can be represented in manifold ways
according to the principle lying at the foundation of nomography.
Further, a large class of functions of three or more variables can
evidently be represented by this principle alone without the use of
variable elements, namely all those which can be generated by
forming first a function of two arguments, then equating each of
these arguments to a function of two arguments, next replacing each
of those arguments in their turn by a function of two arguments, and
so on, regarding as admissible any finite number of insertions of
functions of two arguments. So, for example, every rational function
of any number of arguments belongs to this class of functions
constructed by nomographic tables; for it can be generated by the
processes of addition, subtraction, multiplication and division and
each of these processes produces a function of only two arguments.
One sees easily that the roots of all equations which are solvable by
radicals in the natural realm of rationality belong to this class of
functions; for here the extraction of roots is adjoined to the four
arithmetical operations and this, indeed, presents a function of one
argument only. Likewise the general equations of the th and th
degrees are solvable by suitable nomographic tables; for, by means
of Tschirnhausen transformations, which require only extraction of
roots, they can be reduced to a form where the coefficients depend
upon two parameters only.
Now it is probable that the root of the equation of the seventh
degree is a function of its coefficients which does not belong to this
class of functions capable of nomographic construction, i. e., that it
cannot be constructed by a finite number of insertions of functions of
two arguments. In order to prove this, the proof would be necessary
that the equation of the seventh degree
is not solvable with the help of any
continuous functions of only two arguments. I may be allowed to add
that I have satisfied myself by a rigorous process that there exist
analytical functions of three arguments which cannot be
obtained by a finite chain of functions of only two arguments.
By employing auxiliary movable elements, nomography
succeeds in constructing functions of more than two arguments, as
d'Ocagne has recently proved in the case of the equation of the th
degree.[31]
[30] d'Ocagne, Traité de Nomographie, Paris, 1899.
[31] "Sur la resolution nomographiqne de l'équation du septième
degré." Comptes rendus, Paris, 1900.
14. PROOF OF THE FINITENESS OF
CERTAIN COMPLETE SYSTEMS OF
FUNCTIONS.
In the theory of algebraic invariants, questions as to the
finiteness of complete systems of forms deserve, as it seems to me,
particular interest. L. Maurer[32] has lately succeeded in extending
the theorems on finiteness in invariant theory proved by P. Gordan
and myself, to the case where, instead of the general projective
group, any subgroup is chosen as the basis for the definition of
invariants.
An important step in this direction had been taken already by A.
Hurwitz,[33] who, by an ingenious process, succeeded in effecting
the proof, in its entire generality, of the finiteness of the system of
orthogonal invariants of an arbitrary ground form.
The study of the question as to the finiteness of invariants has
led me to a simple problem which includes that question as a
particular case and whose solution probably requires a decidedly
more minutely detailed study of the theory of elimination and of
Kronecker's algebraic modular systems than has yet been made.
Let a number of integral rational functions
of the variables be given,
Every rational integral combination of must evidently
always become, after substitution of the above expressions, a
rational integral function of . Nevertheless, there may
well be rational fractional functions of which, by the
operation of the substitution , become integral functions in
. Every such rational function of , which
becomes integral in after the application of the
substitution , I propose to call a relatively integral function of
. Every integral function of is evidently
also relatively integral; further the sum, difference and product of
relative integral functions are themselves relatively integral.
The resulting problem is now to decide whether it is always
possible to find a finite system of relatively integral function
by which every other relatively integral function of
may be expressed rationally and integrally.
We can formulate the problem still more simply if we introduce
the idea of a finite field of integrality. By a finite field of integrality I
mean a system of functions from which a finite number of functions
can be chosen, in terms of which all other functions of the system
are rationally and integrally expressible. Our problem amounts, then,
to this: to show that all relatively integral functions of any given
domain of rationality always constitute a finite field of integrality.
It naturally occurs to us also to refine the problem by restrictions
drawn from number theory, by assuming the coefficients of the given
functions to be integers and including among the
relatively integral functions of only such rational
functions of these arguments as become, by the application of the
substitutions , rational integral functions of with
rational integral coefficients.
The following is a simple particular case of this refined problem:
Let integral rational functions of one variable with
integral rational coefficients, and a prime number be given.
Consider the system of those integral rational functions of which
can be expressed in the form
where is a rational integral function of the arguments
and is any power of the prime number . Earlier
investigations of mine[34] show immediately that all such expressions
for a fixed exponent form a finite domain of integrality. But the
question here is whether the same is true for all exponents , i. e.,
whether a finite number of such expressions can be chosen by
means of which for every exponent every other expression of that
form is integrally and rationally expressible.
From the boundary region between algebra and geometry, I will

mention two problems. The one concerns enumerative geometry and
the other the topology of algebraic curves and surfaces.
[32] Cf. Sitzungsber. d. K. Acad. d. Wiss. zu München, 1890, and
an article about to appear in the Math. Annalen.
[33] "Ueber die Erzeugung der Invarianten durch Integration,"
Nachrichten d. K. Geseltschaft d. Wiss. zu Göttingen, 1897.
[34] Math. Annalen, vol. 36 (1890), p. 485.
15. RIGOROUS FOUNDATION OF
SCHUBERT'S ENUMERATIVE
CALCULUS.
The problem consists in this: To establish rigorously and with an
exact determination of the limits of their validity those geometrical
numbers which Schubert[35] especially has determined on the basis
of the so-called principle of special position, or conservation of
number, by means of the enumerative calculus developed by him.
Although the algebra of to-day guarantees, in principle, the
possibility of carrying out the processes of elimination, yet for the
proof of the theorems of enumerative geometry decidedly more is
requisite, namely, the actual carrying out of the process of
elimination in the case of equations of special form in such a way
that the degree of the final equations and the multiplicity of their
solutions may be foreseen.
[35] Kalkül der abzählenden Geometrie, Leipzig, 1879.
16. PROBLEM OF THE TOPOLOGY OF
ALGEBRAIC CURVES AND SURFACES.
The maximum number of closed and separate branches which a
plane algebraic curve of the th order can have has been determined by
Harnack.[36] There arises the further question as to the relative position of
the branches in the plane. As to curves of the th order, I have satisfied
myself—by a complicated process, it is true—that of the eleven blanches
which they can have according to Harnack, by no means all can lie
external to one another, but that one branch must exist in whose interior
one branch and in whose exterior nine branches lie, or inversely. A
thorough investigation of the relative position of the separate branches
when their number is the maximum seems to me to be of very great
interest, and not less so the corresponding investigation as to the number,
form, and position of the sheets of an algebraic surface in space. Till now,
indeed, it is not even known what is the maximum number of sheets which
a surface of the th order in three dimensional space can really have.[37]
In connection with this purely algebraic problem, I wish to bring
forward a question which, it seems to me, may be attacked by the same
method of continuous variation of coefficients, and whose answer is of
corresponding value for the topology of families of curves defined by
differential equations. This is the question as to the maximum number and
position of Poincaré's boundary cycles (cycles limites) for a differential
equation of the first order and degree of the form
where and are rational integral functions of the th degree in and

. Written homogeneously, this is
where , and are rational integral homogeneous functions of the th
degree in , and the latter are to be determined as functions of the
parameter .
[36] Math. Annalen, vol. 10.
[37] Cf. Rohn. "Flächen vierter Ordnung," Preisschriften der Fürstlich
Jablonowskischen Gesellschaft, Leipzig, 1886.
17. EXPRESSION OF DEFINITE
FORMS BY SQUARES.
A rational integral function or form in any number of variables
with real coefficients such that it becomes negative for no real values
of these variables, is said to be definite. The system of all definite
forms is invariant with respect to the operations of addition and
multiplication, but the quotient of two definite forms—in case it
should be an integral function of the variables—is also a definite
form. The square of any form is evidently always a definite form. But
since, as I have shown,[38] not every definite form can be
compounded by addition from squares of forms, the question arises
—which I have answered affirmatively for ternary forms[39]—whether
every definite form may not be expressed as a quotient of sums of
squares of forms. At the same time it is desirable, for certain
questions as to the possibility of certain geometrical constructions, to
know whether the coefficients of the forms to be used in the
expression may always be taken from the realm of rationality given
by the coefficients of the form represented.[40]
I mention one more geometrical problem:
[39] Acta Mathematica, vol. 17.
[40] Cf. Hilbert: Grunglagen der Geometrie, Leipzig, 1899, Chap.
7 and in particular § 38.
18. BUILDING UP OF SPACE FROM
CONGRUENT POLYHEDRA.
If we enquire for those groups of motions in the plane for which
a fundamental region exists, we obtain various answers, according
as the plane considered is Riemann's (elliptic), Euclid's, or
Lobachevsky's (hyperbolic). In the case of the elliptic plane there is a
finite number of essentially different kinds of fundamental regions,
and a finite number of congruent regions suffices for a complete
covering of the whole plane; the group consists indeed of a finite
number of motions only. In the case of the hyperbolic plane there is
an infinite number of essentially different kinds of fundamental
regions, namely, the well-known Poincaré polygons. For the
complete covering of the plane an infinite number of congruent
regions is necessary. The case of Euclid's plane stands between
these; for in this case there is only a finite number of essentially
different kinds of groups of motions with fundamental regions, but for
a complete covering of the whole plane an infinite number of
congruent regions is necessary.
Exactly the corresponding facts are found in space of three
dimensions. The fact of the finiteness of the groups of motions in
elliptic space is an immediate consequence of a fundamental
theorem of C. Jordan,[41] whereby the number of essentially different
kinds of finite groups of linear substitutions in variables does not
surpass a certain finite limit dependent upon . The groups of
motions with fundamental regions in hyperbolic space have been
investigated by Fricke and Klein in the lectures on the theory of
automorphic functions,[42] and finally Fedorov,[43] Schoenflies[44]
and lately Rohn[45] have given the proof that there are, in euclidean
space, only a finite number of essentially different kinds of groups of
motions with a fundamental region. Now, while the results and
methods of proof applicable to elliptic and hyperbolic space hold
directly for -dimensional space also, the generalization of the
theorem for euclidean space seems to offer decided difficulties. The
investigation of the following question is therefore desirable: Is there
in -dimensional euclidean space also only a finite number of
essentially different kinds of groups of motions with a fundamental
region?
A fundamental region of each group of motions, together with
the congruent regions arising from the group, evidently fills up space
completely. The question arises: Whether polyhedra also exist which
do not appear as fundamental regions of groups of motions, by
means of which nevertheless by a suitable juxtaposition of congruent
copies a complete filling up of all space is possible. I point out the
following question, related to the preceding one, and important to
number theory and perhaps sometimes useful to physics and
chemistry: How can one arrange most densely in space an infinite
number of equal solids of given form, e. g., spheres with given radii
or regular tetrahedra with given edges (or in prescribed position),
that is, how can one so fit them together that the ratio of the filled to
the unfilled space may be as great as possible?
If we look over the development of the theory of functions in the

last century, we notice above all the fundamental importance of that
class of functions which we now designate as analytic functions—a
class of functions which will probably stand permanently in the
center of mathematical interest.
There are many different standpoints from which we might
choose, out of the totality of all conceivable functions, extensive
classes worthy of a particularly thorough investigation. Consider, for
example, the class of functions characterized by ordinary or partial
algebraic differential equations. It should be observed that this class
does not contain the functions that arise in number theory and
whose investigation is of the greatest importance. For example, the
before-mentioned function satisfies no algebraic differential
equation, as is easily seen with the help of the well-known relation
between and , if one refers to the theorem proved by
Holder,[46] that the function satisfies no algebraic differential
equation. Again, the function of the two variables and defined by
the infinite series
which stands in close relation with the function , probably

satisfies no algebraic partial differential equation. In the investigation
of this question the functional equation
will have to be used.

If, on the other hand, we are lead by arithmetical or geometrical
reasons to consider the class of all those functions which are
continuous and indefinitely differentiable, we should be obliged in its
investigation to dispense with that pliant instrument, the power
series, and with the circumstance that the function is fully determined
by the assignment of values in any region, however small. While,
therefore, the former limitation of the field of functions was too
narrow, the latter seems to me too wide.
The idea of the analytic function on the other hand includes the
whole wealth of functions most important to science, whether they
have their origin in number theory, in the theory of differential
equations or of algebraic functional equations, whether they arise in
geometry or in mathematical physics; and, therefore, in the entire
realm of functions, the analytic function justly holds undisputed
supremacy.
[41] Crelle's Journal, vol. 84 (1878), and Atti d. Reale Acad. di
Napoli, 1880.
[42] Leipzig, 1897. Cf. especially Abschnitt I, Chaplets 2 and 3.
[43] Symmetrie der regelmässigen Systeme von Figuren, 1890.
[44] Krystallsysteme und Krystallstruktur, Leipzig, 1891.
19. ARE THE SOLUTIONS OF
REGULAR PROBLEMS IN THE
CALCULUS OF VARIATIONS ALWAYS
NECESSARILY ANALYTIC?
One of the most remarkable facts in the elements of the theory
of analytic functions appears to me to be this: That there exist partial
differential equations whose integrals are all of necessity analytic
functions of the independent variables, that is, in short, equations
susceptible of none but analytic solutions. The best known partial
differential equations of this kind are the potential equation
and certain linear differential equations investigated by Picard;[47]

also the equation
the partial differential equation of minimal surfaces, and others. Most

of these partial differential equations have the common characteristic
of being the lagrangian differential equations of certain problems of
variation, viz., of such problems of variation
as satisfy, for all values of the arguments which fall within the range
of discussion, the inequality
itself being an analytic function. We shall call this sort of problem

a regular variation problem. It is chiefly the regular variation
problems that play a rôle in geometry, in mechanics, and in
mathematical physics; and the question naturally arises, whether all
solutions of regular variation problems must necessarily be analytic
functions. In other words, does every lagrangian partial differential
equation of a regular variation problem have the property of
admitting analytic integrals exclusively? And is this the case even
when the function is constrained to assume, as, e. g., in Dirichlet's
problem on the potential function, boundary values which are
continuous, but not analytic?
I may add that there exist surfaces of constant negative
gaussian curvature which are representable by functions that are
continuous and possess indeed all the derivatives, and yet are not
analytic; while on the other hand it is probable that every surface
whose gaussian curvature is constant and positive is necessarily an
analytic surface. And we know that the surfaces of positive constant
curvature are most closely related to this regular variation problem:
To pass through a closed curve in space a surface of minimal area
which shall inclose, in connection with a fixed surface through the
same closed curve, a volume of given magnitude.
[47] Jour. de l'Ecole Polytech., 1890.
20. THE GENERAL PROBLEM OF
BOUNDARY VALVES.
An important problem closely connected with the foregoing is
the question concerning the existence of solutions of partial
differential equations when the values on the boundary of the region
are prescribed. This problem is solved in the main by the keen
methods of H. A. Schwarz, C. Neumann, and Poincaré for the
differential equation of the potential. These methods, however, seem
to be generally not capable of direct extension to the case where
along the boundary there are prescribed either the differential
coefficients or any relations between these and the values of the
function. Nor can they be extended immediately to the case where
the inquiry is not for potential surfaces but, say, for surfaces of least
area, or surfaces of constant positive gaussian curvature, which are
to pass through a prescribed twisted curve or to stretch over a given
ring surface. It is my conviction that it will be possible to prove these
existence theorems by means of a general principle whose nature is
indicated by Dirichlet's principle. This general principle will then
perhaps enable us to approach the question: Has not every regular
variation problem a solution, provided certain assumptions regarding
the given boundary conditions are satisfied (say that the functions
concerned in these boundary conditions are continuous and have in
sections one or more derivatives), and provided also if need be that
the notion of a solution shall be suitably extended?[48]
[48] Cf. my lecture on Dirichlet's principle in the Jahresber. d.
Deutschen Math.-Vereinigung, vol. 8 (1900), p. 184.

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vi O R G A N I Z A T I O N AND NEW CONTENT

viii O R G A N I Z A T I O N AND NEW CONTENT

x A bout the A uthors

A bout the A uthors xi

different chapters MAKE CONNECTIONS

Figure 3.30 Contributions of

Figure 8.20 The Working Cell,

Figure 16.21 Genomics, Nutritional Mode

666 UNIT SIx AnimAl Form And Function

xii M a k e C onnections V isually

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ChApTEr 32 tHE intErnAl EnVironmEnt oF AnimAlS: orGAnizAtion And rEGulAtion 667

M a k e C onnections V isually xiii

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Practice Scientific Skills

Scientific Skills Exercise

Making and Testing Predictions

392 UNIT THREE EVOLUTION

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xiv P ractice S cienti f ic S k ills

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14 Gene Expression: From Gene to Protein

The list of Key Concepts KEY CONCEPTS

Every chapter opens

After reading a Key Concept section,

New Mutations and Mutagens

Third mRNA base (3′ end of codon)

codons. Instead, the protein pro

protein synthesis. His artificial system translated the64 5′

14.6). What evo

• 14.23After translation, modifications to proteins can affect their shape.

or initiation codon. Genetic messages usually begin with the

num- from the tem-

is emphasized throughout. Foras aexample: