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Global Global
edition edition
edition
Global
For these Global Editions, the editorial team at Pearson has
Biology in Focus
Campbell
collaborated with educators across the world to address a wide range
of subjects and requirements, equipping students with the best possible
learning tools. This Global Edition preserves the cutting-edge approach
and pedagogy of the original, but also features alterations, customization,
and adaptation from the North American version.
Lisa A. Urry
Michael L. Cain
Steven A. Wasserman
Peter V. Minorsky
second
edition
Jane B. Reece
Minorsky • Reece
Urry • Cain • Wasserman
This is a special edition of an established title widely
used by colleges and universities throughout the world.
Campbell
Pearson published this exclusive edition for the benefit
of students outside the United States and Canada. If you
purchased this book within the United States or Canada,
you should be aware that it has been imported without
Biology in Focus
the approval of the Publisher or Author. second edition
Pearson Global Edition
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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
6 O R G A N I Z A T I O N AND NEW CONTENT
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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.
8 O R G A N I Z A T I O N AND NEW CONTENT
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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.
O R G A N I Z A T I O N A N D N E W C O N T E N T 9
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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.
10 A B O U T THE AUTHORS
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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.
A B O U T T H E A U T H O R S 11
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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. 112
Figure 8.20 The Working Cell,

pp. 222–223
Figure 16.21 Genomics, Nutritional Mode

Cell Signaling, and Cancer, All living things must obtain energy and carbon from the
pp. 382–383 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. 472–473 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. 570 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. 647 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. 710–711 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. 821 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. 946–947 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. 968–969 chemical and electrical communication.
(See Figure 31.12 and Figure 38.26.)
710 UNIT SIx AnimAl Form And Function
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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 711
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M A K E C O N N E C T I O N S V I S U A L LY 13

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. 62 13. Working with Data in a Table, p. 301
2. Interpreting a Scatter Plot with a Regression Line, p. 84 14. Interpreting a Sequence Logo, p. 338
3. Analyzing Polypeptide Sequence Data, p. 113 15. Analyzing DNA Deletion Experiments, p. 357
4. Using a Scale Bar to Calculate Volume and Surface Area 16. Analyzing Quantitative and Spatial Gene Expression Data, p. 369
of a Cell, p. 124 17. Analyzing a Sequence-Based Phylogenetic Tree to Understand
5. Interpreting a Scatter Plot with Two Sets of Data, p. 153 Viral Evolution, p. 397
6. Making a Line Graph and Calculating a Slope, p. 178 18. Reading an Amino Acid Sequence Identity Table, p. 414
7. Making a Bar Graph and Evaluating a Hypothesis, p. 199 19. Making and Testing Predictions, shown above and on p. 436
8. Making Scatter Plots with Regression Lines, p. 220 20. NEW! Using Protein Sequence Data to Test an Evolutionary
Hypothesis, p. 454
9. Interpreting Histograms, p. 240
21. Using the Hardy-Weinberg Equation to Interpret Data and Make
10. Making a Line Graph and Converting Between Units of
Predictions, p. 464
Data, p. 254
22. Identifying Independent and Dependent Variables, Making a
11. Making a Histogram and Analyzing a Distribution
Scatter Plot, and Interpreting Data, p. 485
Pattern, p. 271
23. Estimating Quantitative Data from a Graph and Developing
12. Using the Chi-Square (χ2) Test, p. 290
Hypotheses, p. 503
14 P R A C T I C E SCIENTIFIC SKILLS

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. 537 34. Interpreting Data in Histograms, shown above and on p. 765
25. Interpreting Comparisons of Genetic Sequences, p. 545 35. Comparing Two Variables on a Common x-Axis, p. 792
26. NEW! Interpreting Genomic Data and Generating 36. Making Inferences and Designing an Experiment, p. 805
Hypotheses, p. 573 37. Interpreting Data Values Expressed in Scientific Notation, p. 831
27. Understanding Experimental Design and Interpreting 38. Designing an Experiment Using Genetic Mutants, p. 841
Data, p. 614
39. Interpreting a Graph with Log Scales, p. 869
28. Using Bar Graphs to Interpret Data, p. 626
40. Using the Logistic Equation to Model Population Growth, p. 904
29. Calculating and Interpreting Temperature
Coefficients, p. 641 41. Using Bar Graphs and Scatter Plots to Present and Interpret
Data, p. 914
30. Using Positive and Negative Correlations to Interpret
Data, p. 676 42. Interpreting Quantitative Data in a Table, p. 937
31. Interpreting Experimental Results from a Bar 43. Graphing Cyclic Data, p. 966
Graph, p. 700
32. Describing and Interpreting Quantitative
Data, p. 723
33. Interpreting Data from an Experiment with
Genetic Mutants, p. 748
PR AC TICE SCIENTIFIC SKILLS 15

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
treatFigure genetic 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
112 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#112 153397 Cust: Pearson / BC Au: Urry Pg. No. 112
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Campbell Biology in Focus, 2e
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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 319
K E E P C U R R E N T W I T H N E W S C I E N T I F I C A D V A N C E S 17
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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.
322
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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

18
Mutations can
FOCUS O N Tarise
H E K Ein
Y a
COnumber
N C E P T S of ways. Errors during DNA 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. G
In eukaryotic
5′ cap to the 5′ end, and the addition of a poly-A tail to the 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. 323–328)
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. the spliceosom
RNA
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′ caps
U
are removed from the
cases, RNA Level 3: Synthesis/Evaluation
What will be the results of chemically modifying one nucleotide
mRNA.
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
be read in the correct reading frame. tide chains (enzymes
The
signal (3 codons). Codons must
translationand CUC other proteins)
is the rNa-directed CCC
synthesis or
CAC RNA molecules.
CGC C ability
the cell? 10.
of A some
researcher wants RNA to produce molecules,
a eukaryotic protein of 45 kDa called ribozymes, derives
SCIENTIFIC INQUIRY
C (D) the DNA introns are removed from the template.

first codon was deciphered in 1961? by Marshall Nirenberg, • Transcription of of a polypeptide:
•is the synthesis
Describe the process of gene expression, by which a gene affects Leu
a closer look (pp. 332–342)
ofprotein
RNA pro
complementary CGAtESt YoUR
Arg
to aAUnDERStAnDinG from thefrom
(around 350 amino acids). The researcher amplifies the gene
properties of RNA. The

PRACTICE
10. SCIENTIFIC INQUIRY
codon presence of introns allows for
TEST
the phenotype of an organism. A cell CUA
translates an mRNA messageCCA into CAA
using transfer genomic DNA taking care to eliminate the eukaryotic
promoter and starts the amplification at the start ATG
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 oftheGlnsynthesis Level 1: Knowledge/Comprehension
ofof theathe 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. Which Gfollowingfollowing
is not true about mRNAs? is not true ofand
alternative a
cloned codon?
RNA
into splicing.
stops it right at the termination codon. Finally, the gene is
a bacterial expression vector right after a bacterial
Nirenberg synthesized an artificial mRNA byDNa-directed
linking identical uses molecular biologi
polypeptide whose amino acid sequence is specified by the
ribosomal RNAs (rRNAs) and proteins, facilitates this coupling (A) Eukaryotic mRNA slack introns. promoter. When the researcher tries to produce the protein in
RNA nucleotides containing uracil

transcription is the
a closer look (pp. 328–330)
as their base. No matter
synthesis of rNa:
with binding sites for mRNA and tRNA. (A) It(C)may(B) Prokaryotic code for the same amino ever,
mRNAs do not have poly-A tails. acid as another codon.
bacteria no protein of the expected size can be detected. How- goo.gl/CRZjvS
nucleotide
to a DNA template sequence
• Ribosomes AUU
elongation, in mRNA.
coordinate the three stages ACU AAU
of translation: initiation, AGU (D) Prokaryotic U cells do not cap their mRNAs on the 5′. Whatthatfunction dopairs.the
Explain 5′
what iscap
the researcher is able to identify some RNA for the gene
and the poly-A tailgene serve on
(shown in Figure
(B) It never codes for more ? 11.
than one amino acid.
• RNA synthesis is catalyzed by RNA polymerase, which links and termination. The formation of peptide bonds Pre-mRNAs do not have 5′ cap. is around 10,000 base happening in
where this message started or stopped, Asn ser
strand. Thisit could followscontain onlyrules as DNA
together RNA nucleotides complementary this experiment.
C for the same amino acid as another codon. a eukaryotic mRNA?
between amino acids is catalyzed by ribosomal RNAs as tRNAs
AUC the A andIle ACC AAC AGC2. Which of the following is not true of a codon?
cells will express it and
• Genetic information is encoded as a sequence of nonoverlap-
process the same base-pairing move through P sites and exit through the E site.
(C) It (B)extends from than oneone end of a tRNA molecule.
(A) It may code FOCUS ON EVOLUTION
replication, except that in RNA, uracil substitutes for thymine. • AfterA
one codon in repetition: UUU. Nirenberg added this “poly-U” thr
translation, modifications to proteins can affect their shape. It never codes for more amino acid. The genetic code has a built-in redundancy as several codons
codons. Instead, the protein pro

Free ribosomes in the cytosol initiate synthesis of all proteins, but
AUA Afromunitoneofend can code for a single amino acid. This is the result of the avail-
ping nucleotide triplets, orpeptide ACA A codon AAA in messenger AGA (C) ItIt extends
transcription unit
of a tRNA molecule.
Lys (D) It(D)Arg is isthethe basicbasic unit code. of the genetic ability code.
proteins with a signal
codon to a test-tube mixture that contained all 20 amino promoter are synthesized on the ER.
the genetic of greater possible base combinations in codons (64) than
availability of amino acids. Take a look at table 14.6 and you will
contain many fewer am
polypeptide
metbyor
RNA for (mRNA) either ispolymerases
translated into an amino acid (61 of the
• A gene can AUG 3. The anticodon of a particular tRNA molecule is
be transcribed ACG AAG AGG G realize that the third base of a codon or several amino can be
acids, ribosomes, and the other components required start
CONCEPT 14.4
Amino
3. The anticodon
(A) complementary to the corresponding mRNA codon.
ofcorresponding
a particulartriplet in rRNA. tRNA implicationsmolecule isin mutation and evolution.
5′ 3′ multiple RNA 3′ acid changed without changing the translation. Discuss the possible
3′ 5′ trnA
(B) complementary to the
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
(C) the part of tRNA that bonds with a specific amino acid.
GGU (D) catalytic,Umaking the tRNA a ribozyme. 12.
this could have
eukaryotic cell. Explain
(A) complementary to theTranslationcorresponding Evolutionis mRNA
the codon.
RNA-directed synthesis
rnA polymerase of DnA FOCUS ON INFORMATION
lated simultaneously
reading frame. GAC Asp GGC4. Which
accounts for the unity and diversity of life, and the
arebe read in the correct
rnA transcript
into a polypeptide containing many unitsstagesof the amino acid ber of ribosomes, forming a
GUC In bacteria, these GCC
of the following is not true of RNA processing?
E A Anti-
(A) Exons areCcut out before mRNA leaves the nucleus.
continuity of life is based on heritable information in the form
11. FOCUS ON EVOLUTIO
(B) complementary
(B) Nucleotides may be added at bothto ends the corresponding triplet aincloserisrRNA.
• The three of transcription initiation, elongation, polyribosome.
Ribozymes may function in RNA splicing. of a polypeptide:

related to the look (pp. 288–298)
codon of DNA. In a short essay (100–150 words), discuss how the
phenylalanine (Phe), strung together
and termination. A promoter, often including a TATA box
as a establishes
polyphenylalanine G are coupled, but
processes in of the RNA.
Val Ala (C)Gly
fidelity with which DNA is inherited processes
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 gene(D) RNAaffects
Codon
Most amino acids are c
Glu (C) the part of tRNA that bonds with ain specific amino acid.
GCA GAA GGA A mrnA of evolution. (Review the discussion of proofreading and DNA
?andthe
splicing can be catalyzed by spliceosomes.
chain. Thus, Nirenberg determined that the mRNA forming a codon
repair Concept 13.2.)
nize promoter sequences, transcription initiation membrane.
•tRNA A cell translates an mRNA message into protein Figure using transfer
14.6). What evo
ribosome
5. Which of the following is not required for transcription?
complex. Termination differs in bacteria
UUU specifies the amino acid phenylalanine. Soon, the amino phenotype
eukaryotes.
? of anGUG organism. GCG GAG (D)
GGGcatalytic, G making
What function do tRNAs serve in the process of translation?
(A) RNA polymerase
the
(C) Deoxynucletides.
a ribozyme.
13. SYNTHESIZE YOU R K NOWLEDGE
RNAs (tRNAs). After being bound to a specific amino acid by(Hint:

an Th
What are the similarities and differences in the initiation
this pattern?
(B) Transcription factors (D) DNA promoter
? of gene transcription in bacteria and eukaryotes?
acids specified by the codons AAA, GGG, and CCC were
cONcept 14.3 ▲Mutations
Figure
cONcept 14.5
14.6 The codon table for mRNA. 4. Which
the three ofnucleotide
the
Level 2: Application/Analysis
following is not true of RNA processing?
aminoacyl-tRNA synthetase, a tRNA lines up via try, its
and anticodon
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)
6. Using Figure 14.6, identify a 5′ → 3′ sequence of nucleotides in
CONCEPT 14.2 bases proteinof an mrnA

structure and functioncodon (pp.are342–344) Exons
the first, tide sequenceare
the second,
DNA templateand
cut out beforeatmRNA
strand for an mRNA coding for the polypep-
leaves the nucleus.
the complementary codon on mRNA. A ribosome, made up of
12.
Phe-Pro-Lys.
Although more elaborate techniques(pp. 330–332)
were required to bases, reading in the 5′ → 3′ direction along the mrnA.
• Small-scale mutations include point mutations, changes in
third (practice FOCUS ON INFORMAT
(B) of Nucleotides
(A) 5′-UUUCCCAAA-3′
(B) 5′-GAACCCCTT-3′ may be added at both ends of the RNA.
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
thisproteins.
table by finding the codons synthesis
can causein Figure 14.5.) the (C)RNA:
codon AUG
5′ exon intron exon intron exon 3′
Pre-mRNA functional
missense or nonsense
Nucleotide-pair substitutions
mutations. Nucleotide-pair insertions
(C) Ribozymes
5′-CTTCGGGAA-3′
also functionsmay function in RNA splicing.
Some mutations result in proteins that function well at one Evolution accounts for
with binding sites for mRNA and tRNA.
(D) 5′-AAACCCUUU-3′
notor only stands forframeshift
the amino acid methionine (met) but
were deciphered by the mid-1960s. As Figure 14.6
rnA splicing
shows,
a closer look (pp.
deletions may produce
284–286)
mutations.
7. The following sequence of a DNA template strand: 5′-TAT
temperature but are nonfunctional at a different (usually higher)
continuity of life is bas
mRNA
as• Spontaneous
a “start” mutations can occur during DNA replication, re-
signal for ribosomes to begin translating the mrnA at temperature. Siamese cats have such a “temperature-sensitive”
61 of the 64 triplets code for amino acids. The three codons 5′ Utr Coding
combination, or repair. Chemical and physical mutagens cause
3′ Utr (D) RNA would havesplicing
GGT GCA-3′ codes for a
can be catalyzed
peptide. Determine the mutation that
• Ribosomes by spliceosomes.
ment in coordinate
mutation
the fur. The mutation resultsthe three
in a gene encoding an enzyme that makes dark pig-
stages of translation: initiation,
of DNA. In a short ess
the most severe effect.
segment thatDNA point.
damage thatthree
can alterofgenes.
the 64 codons function as “stop” signals, (A) 5′-TATmarking
in the breed’s distinctive
that do not designate amino acids are “stop” signals, or ter- • RNA synthesis is catalyzed by RNA polymerase, which links GGT ACA-3′ (C) 5′-GAT GGT GCA-3′
elongation,
point markings and lighter body color (see the photo). Using
andin termination. The explain formation of peptide bonds
where ribosomes end translation. see Figure 3.18 for a list of the full
5. Which component isshown
not directly involved cat’stranslation?
(B) 5′-TAT CGT GCA-3′ 345 (D) 5′-TAT GTT GCA-3′ this information and what you learned in the chapter,
fidelity with which DN
chapter 14 Gene expression: From Gene to protein
together that RNA names nucleotides complementary to a DNA template

8. Would the coupling of the processes in Figure 14.23 be the pattern of the fur pigmentation.
of all the amino acids.
mination codons, marking the end of translation. Notice
(A) GTP
found in a eukaryotic cell? Explain why or why CONCEPT
not.
between 14.2(C) amino tRNA acids is catalyzed by ribosomal RNAs as tRNAs
the codon AUG has a dual function: It codes for the strand. amino This process follows the same base-pairing rules as DNA
9. Fill in the following table: of evolution. (Review t
For selected answers, see Appendix A.
# 153397 Cust: Pearson / BC Au: Urry Pg. No. 345

S4carliSle C/M/Y/K
(B) DNA
DESIGN SERVICES OF
Transcription move through (D)

Type of RNA ribosomes
is the
the A DNA-directed
and P sites and exit throughrepair
Functions the Einsite. Concept 13.2
acid methionine (Met) and • Salso functions ofasKey a “start” replication,
signal, questions exceptcheck that in RNA, uracil substitutes for thymine.
M14_URRY9589_02_GE_CH14.indd Title: Campbell Biology in Focus, 2e Short / Normal / Long
ummary Concepts students’
345 Publishing Services 22/01/16 1:08 PM
• After translation, modifications to proteins can affect their shape.

Messenger RnA (mRnA)
or initiation codon. Genetic messages usually begin

understanding of a key with idea the from each probably be gibberish: for example, “her edd oga tet heb ug.”
concept. synthesis of RNA: a closer look
transfer RnA (tRnA)
Free ribosomes in the cytosol initiate synthesis 13.of SYNTHESI
all proteins, ZEbut YO UR K
mRNA codon AUG, which signals the protein-synthesizing
Transcription unit
The reading frame is also important in the molecular language Level 2: Application/Analysis Plays catalytic (ribozyme) roles and
structural roles in ribosomes
• 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 thatproteins
we have with
considered
Primary transcript a signal
the peptide
linguistic logic and evolution-
Promoter for instance, will be made correctly only6.if the Using Figure 14.6, identify aare
ary significance 5′ → 3′
synthesized
of the sequencegenetic oncode,of
thenucleotides
weER. are ready toinreexamine
small RnAs in the spliceosome
(Because AUG also stands for methionine, polypeptide chains mRNA nucleo- Polypeptide
NEW!the DNASynthesize template Your strandKnowledge 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
346
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 informationbe 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, is a
# 153397 Cust: Pearson / BC Au: Urry
simultaneously S4carliSle
by transcribed
num- from the tem-
Pg. No. 346 C/M/Y/K
is emphasized throughout. Foras aexample:

346 Campbell Biology in Focus, 2e Short / Normal / Long Publishing Services 22/01/16 3:45 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 is ainitiated.
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 in DNAmiddle 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 RNA
firefly bacteriaofto polymerase
the
the 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,
fluorescentinitiation
for pig.
instance, is
(B) a single nucleotide deletion in the middle of an intron DNA 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
acidofproline
genes and in eukaryotes.
all
from organisms
different species.
(C) code,a single whosenucleotidechain deletion from 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 mo
carrier
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 thetrans-
(D) a single nucleotidewhere
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 synthesize the coupling ofthe the end of transcription
processes
firefly gene. the shown is called inaFigure Pig expressing
terminator.
14.23 (Thea termi-
be 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 or agene.
in not.
eukaryotes; genefew researchers
for 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
promoter
code,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
Title: Campbell Biology in Focus, 2e Short / Normal / Long
of DNAmissense nonsense mutations. insertions
RY2751_02_C14_PR5.indd 283 31/07/15 5:38 PM species by introducing DnA from the second species into the first.
downstreamor Nucleotide-pair
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 that synthe-
programmed the insertion N Tcan
F O C U SofOhuman occur
H Egenes
K E Y to
CO during
synthesize DNA19replication,
N C E P T S 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
ChemicalsuchRNA 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
A01_URRY9589_02_GE_FM.indd 19 have the disorder. today, would this disorder be called domi- 23/02/16 2:46 PM
history of life—early enough to be present in the common an- an RNA molecule is called a
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The Project Gutenberg eBook of Pen and pencil sketches of Faröe and
Iceland
This ebook is for the use of anyone anywhere in the United
States and most other parts of the world at no cost and with
almost no restrictions whatsoever. You may copy it, give it away
or re-use it under the terms of the Project Gutenberg License
included with this ebook or online at www.gutenberg.org. If you
are not located in the United States, you will have to check the
laws of the country where you are located before using this
eBook.
Title: Pen and pencil sketches of Faröe and Iceland

With an appendix containing translations from the
Icelandic and 51 illustrations engraved on wood by W.
J. Linton
Author: Andrew James Symington
Illustrator: W. J. Linton
Translator: Ólafur Pálsson
Release date: October 23, 2023 [eBook #71936]
Language: English
Original publication: London: Longman, Green, Longman, and

Roberts, 1862
Credits: Charlene Taylor, Gísli Valgeirsson, Bryan Ness and the

Online Distributed Proofreading Team at
https://www.pgdp.net (This file was produced from
images generously made available by The Internet
Archive/American Libraries.)
*** START OF THE PROJECT GUTENBERG EBOOK PEN AND

PENCIL SKETCHES OF FARÖE AND ICELAND ***
PEN AND PENCIL SKETCHES
OF
FARÖE AND ICELAND.
“To the ocean now I fly,

And those northern climes that lie
Where Day never shuts his eye.”
Comus.
THE GREAT GEYSER IN ERUPTION.—See page 117.
OF
FARÖE AND ICELAND
WITH AN APPENDIX
CONTAINING TRANSLATIONS FROM THE ICELANDIC
AND 51 ILLUSTRATIONS ENGRAVED ON WOOD BY W. J. LINTON
BY
ANDREW JAMES SYMINGTON

Author of “Harebell Chimes,” “The Beautiful in Nature, Art, and Life,” &c.
LONDON
LONGMAN, GREEN, LONGMAN, AND ROBERTS
1862
TO
LAURENCE EDMONDSTON, Esq., M.D.,

OF SHETLAND,
CORRESPONDING MEMBER OF THE ROYAL PHYSICAL AND WERNERIAN SOCIETIES,

EDINBURGH;
HONORARY MEMBER OF THE YORKSHIRE PHILOSOPHICAL AND MANCHESTER
NATURAL HISTORY SOCIETIES, ETC.,
THIS VOLUME
IS AFFECTIONATELY INSCRIBED BY HIS SON-IN-LAW
A.J.S.
May 1862.
PREFACE.
The greater part of this volume consists of a diary jotted down in presence of
the scenes described, so as to preserve for the reader, as far as possible, the
freshness of first impressions, and invest the whole with an atmosphere of
human interest.
The route taken may be thus shortly indicated: Thorshavn; Portland Huk; the
Westmanna Islands; Reykjavik; the Geysers; then, by sea, round the south
coast of the island, with its magnificent Jökul-range of volcanoes; along the
east coast, with its picturesque Fiords, as far north as Seydisfiord; and thence
home again, by the Faröe Isles.
The aim, throughout, has been both to present pictures and condense
information on matters relating to Faröe and Iceland. In obtaining the latter I
have had the advantages of frequent intercourse with Icelanders, both personal
and by letter, since my visit to the North in the summer of 1859, and would
here mention, in particular, the Rev. Olaf Pálsson, Dean and Rector of
Reykjavik Cathedral; Mr. Jón Arnason, Secretary to the Bishop, and Librarian;
Mr. Gísli Brynjúlfsson, the Icelandic poet and M.P.; Mr. Sigurdur Sivertsen, a
retired merchant, and Mr. Jacobson.
And so too with the Faröese.
I acknowledge obligations to Dr. David Mackinlay of Glasgow, Dr. Lauder
Lindsay of Perth, and several other friends who have visited Iceland and
rendered me assistance of various kinds. Thanks are also due to Mr. P. L.
Henderson, for transmitting, by the Arcturus, letters, books and newspapers to
and from the north.
The Appendix comprises thirteen Icelandic stories and fairy tales translated by
the Rev. Olaf Pálsson; specimens of old Icelandic poetry; poems on northern
subjects in English and Icelandic; information for intending tourists; a
glossary; and lastly, a chapter on our Scandinavian ancestors—treating of race,
history, characteristics, language and tendencies. This paper, originally intended
for an introduction, may be perused either first or last, at the option of the
reader. There is also a copious Index to the volume.
The illustrations, engraved by Mr. W. J. Linton, are all from original drawings
by the writer, with the exception of half a dozen,[1] taken from plates in the
large French folio which contains the account of Gaimard’s Expedition.
Should these pages induce photographers and other artists to visit this strange
trahytic island resting on an ocean of fire in the lone North Sea, or students to
become familiar with its stirring history and grand old literature, I shall feel
solaced, under a feeling almost akin to regret, that this self-imposed task—
which, in spite of sundry vexatious delays and interruptions, has afforded me
much true enjoyment—should at length have come to an end.
A.J.S.
May 1862.
CONTENTS.
PAGE
PREFACE v
LEITH TO THORSHAVN 1
WESTMANNA ISLANDS—REYKJAVIK 35
RIDE TO THE GEYSERS 69
REYKJAVIK 143
JÖKUL-RANGES AND VOLCANOES ON THE SOUTH COAST 160
Kötlugjá’s Eruptions 163
Icelandic Statistics 180
Eruption of Skaptár Jökul 187
Volcanic History of Iceland 193
THE EAST COAST. BREIDAMERKR—SEYDISFIORD 197
Seydisfiord, by Faröe, to Leith 208
APPENDIX.
I. Icelandic Stories and Fairy Tales

Stories of Sæmundur Frodi called the learned.
I. The dark School 219
II. Sæmund gets the living of Oddi 221
III. The Goblin and the Cowherd 222
IV. Old Nick made himself as little as he was able 224
V. The Fly 224
VI. The Goblin’s Whistle 225
Fairy Tales
Biarni Sveinsson and his sister Salvör 226
Una the Fairy 235
Gilitrutt 240
Hildur the Fairy Queen 244
A Clergyman’s daughter married to a Fairy Man 253
The Clergyman’s daughter in Prestsbakki 256
The Changeling 257
II. Specimens of old Icelandic Poetry

From the “Völuspá” 260
From the “Sólar Ljód” or “Sun Song” 262
From the Poems relating to Sigurd & Brynhild 265
“The Hávamál” or “High Song of Odin” 265
III. Poems on Northern Subjects

The Lay of the Vikings, by M.S.E.S. 278
Do. Translated into Icelandic by the Rev. Olaf Pálsson 279
The Viking’s Raven, by M.S.E.S. 281
Death of the Old Norse King, by A.J.S. 286
Do. Translated into Icelandic by the Rev. Olaf Pálsson 287
IV. Information for intending Tourists 289
V. Glossary 292
VI. Chapter on Our Scandinavian Ancestors 293
INDEX 309
LIST OF ILLUSTRATIONS.
PAGE
1. The Great Geyser in Eruption Frontispiece.
2. Little Dimon—Faröe 1
3. Foola 10
4. Naalsöe 15
5. Thorshavn, the capital of Faröe 20
6. Fort, at Thorshavn 23
7. From Thorshavn—showing Faröese Boats 27
8. Hans Petersen; a Faröese boatman 28
9. Basalt Caves—South point of Stromoe 32
10. Portland Huk—looking south 35
11. Needle Rocks or Drongs—off Portland Huk 38
12. Bjarnarey 43
13. Westmanna Skerries 43
14. Cape Reykjanes—showing Karl’s Klip (cliff) 45
15. Coast near Reykjavik 45
16. Eldey 46
17. Icelandic shoes, snuff-box, distaff, head-dress, & fishermen’s two-thumbed 53
mits
18. Reykjavik, from behind the town 67
19. Icelandic Lady in full dress, from a Photograph 68
20. View on the Route to Thingvalla 69
21. Ravine 73
22. Descent into the Almannagjá 81
23. Almannagjá 82
24. Fording the Oxerá 83
25. Priest’s House at Thingvalla 84
26. Althing and Lögberg from behind the church 87
27. Lake of Thingvalla from the Lögberg 89
28. Waterfall of the Oxerá as seen from the Lögberg 90
29. Vent of Tintron 94
30. Cinder-range of Vari-coloured Hills 95
31. Crossing the Bruará 104
32. The Great Geyser 129
33. Skaptár Jökul 134
34. Mount Hekla 135
35. Lake of Thingvalla from the north-west 139
36. Icelandic Farm, two hours’ ride from Reykjavik 142
37. Music in an Icelandic home (playing the langspiel) 145
38. Common Gull (Larus canus) 159
39. Oræfa Jökul, the highest mountain in Iceland 160
40. Snæfell Jökul, from fifty miles at sea 161
41. Part of Myrdals Jökul and Kötlugjá range 184
42. Oræfa Jökul, from the sea 185
43. Entrance to Reydarfiord—east coast 197
44. Near the entrance to Hornafiord 198
45. Mr. Henderson’s Factory at the head of Seydisfiord 202
46. Farm House, Seydisfiord 206
47. Seydisfiord, looking east towards the sea 207
48. Brimnæs Fjall 208
49. Naalsöe—Faröe 212
50. Entrance to the Sound leading to Thorshavn 213
51. Stromoe—Faröe, looking north-east from below the Fort at Thorshavn 216
LITTLE DIMON—FARÖE.

OF
FARÖE AND ICELAND.

LEITH TO THORSHAVN.
Can Iceland—that distant island of the North Sea, that land of Eddas and
Sagas, of lava-wastes, snow-jökuls, volcanoes, and boiling geysers—be visited
during a summer’s holiday? This was the question which for years I had
vaguely proposed to myself. Now I wished definitely to ascertain particulars,
and, if at all practicable, to accomplish such a journey during the present
season.
Three ways presented themselves—the chance of getting north in a private
yacht—to charter a sloop from Lerwick—or to take the mail-steamer from
Copenhagen. The first way seemed very doubtful; I was dissuaded from the
second by the great uncertainty as to when one might get back, and the earnest
entreaties of friends, who, with long faces, insinuated that these wild northern
seas were not to be trifled with. However, the uncertainty as to time, and the
expense, which for one person would have been considerable, weighed more
with me than any idea of danger. Of the mail-steamer it was difficult to obtain
any information.
One morning, when in this dilemma, my eye fell on an advertisement in the
Times, headed “Steam to Iceland,” informing all whom it might concern that
the Danish mail-steamer “Arcturus,” would, about the 20th of July, touch at
Leith on its way north, affording passengers a week to visit the interior of the
island, and would return to Leith within a month. I subsequently ascertained
that it was to call at the Faröe and Westmanna Isles, and that it would also sail
from Reykjavik round to Seydisfiord, on the east of Iceland, so that one might
obtain a view of the magnificent range of jökuls and numerous glaciers along
the south coast.
The day of sailing was a fortnight earlier than I could have desired, but such an
opportunity was not to be missed. Providing myself with a long waterproof
overcoat, overboots of the same material—both absolutely essential for riding
with any degree of comfort in Iceland, to protect from lashing rains, and when
splashing through mud-puddles or deep river fordings—getting together a
supply of preserved meats, soups, &c. in tin cans, a mariner’s compass,
thermometer, one of De La Rue’s solid sketch-books, files of newspapers, a
few articles for presents, and other needful things, my traps were speedily put
up; and, on Wednesday the 20th of July, I found myself on board the
“Arcturus” in Leith dock.
It was a Clyde-built screw-steamer, of 400 tons burden. Captain Andriessen, a
Dane, received me kindly; the crew, with the exception of the engineer, a
Scotchman, were all foreigners. In the first cabin were eight fellow-passengers,
strangers to each other; but, as is usual at sea, acquaintanceships were soon
formed; by degrees we came to know each other, and all got along very
pleasantly together.
There was only one lady passenger, to whom I was introduced, Miss Löbner,
daughter of the late governor of Faröe, who had been south, visiting friends in
Edinburgh. Afraid of being ill, she speedily disappeared, and did not leave her
cabin till we reached Thorshavn. Of our number were Professor Chadbourne,
of William’s College, Massachusetts, and Bowdoin College, Maine, U.S.; Capt.
Forbes, R.N.; Mr. Haycock, a gentleman from Norfolk, who had recently
visited Norway in his yacht; Mr. Cleghorn, lately an officer in the Indian army;
Mr. Douglas Murray, an intelligent Scottish farmer, from the neighbourhood
of Haddington, taking his annual holiday; Dr. Livingston, an American M.D.;
and Capt. B——, a Danish artillery officer, en route from Copenhagen to
Reykjavik.
There were also several passengers in the second cabin, some of whom were
students returning home from their studies in the Danish universities.
There was a large boat to be got on board, for discharging the steamer’s cargo
at Iceland, which took several hours to get fastened aloft on the right side of
the hurricane deck—with the comfortable prospect of its top-heaviness acting
like a pendulum, and adding considerably to the roll of the ship, should the
weather prove rough.
Shortly after seven P.M. we got fairly clear of the dock. Strange to think, as the
last hawser was being cast off, that, till our return, we should hear no
postman’s ring, receive no letters with either good tidings or annoyances—for
we carry the mail,—and see no later newspapers than those we take with us!
Friends may be well or ill. The stirring events of the Continent, too, leave us to
speculate on changes that may suddenly occur in the aspect of European
affairs, with the chances of peace, or declarations of war.
However, allowing such thoughts to disturb me as little as possible, and
trusting that, under a kind Providence, all would be well with those dear to me,
hopefully, and not without a deep feeling of inward satisfaction that a long
cherished dream of boyhood was now about to be realised, I turned my face to
the North.
A dense mist having settled on the Frith of Forth, the captain deemed it
prudent to anchor in the roads. During the night it cleared off, and at five
o’clock on Thursday morning, 21st July, our star was in the ascendant, and the
“Arcturus” got fairly under way.
The morning, bright and clear, was truly splendid; the day sunny and warm;
many sails in sight, and numerous sea-birds kept following the ship.
Breakfast, dinner, and tea follow each other in regular succession, making, with
their pleasant reunions and friendly intercourse, a threefold division of the day.
On shipboard the steward’s bell becomes an important institution, a sort of
repeating gastronomical chronometer, and is not an unpleasant sound when
the fresh sea-air has sharpened one’s appetite into expectancy.
The commissariat supplies were liberal, and the department well attended to
by a worthy Dane, who spoke no English, and who was only observed to smile
once during the voyage. Captain Andriessen’s fluent English, and the obliging
Danish stewardess’ German, enabled us all to get along in a sort of way;
although the conversation at times assumed a polyglot aspect, the ludicrous
olla-podrida nature of which afforded us many a good hearty laugh.
The chief peculiarities in our bill of fare were lax or red-smoked salmon; the
sweet soups of Denmark, with raisins floating in them; black stale rye-bread;
and a substantial dish, generally produced thrice a day, which, in forgetfulness
of the technical nomenclature, we shall venture to call beef-steak fried with
onions or garlic—that bulb which Don Quixote denounced as pertaining to
scullions and low fellows, entreating Sancho to eschew it above all things when
he came to his Island. At sea, however, we found it not unpalatable. There
must ever be some drawbacks on shipboard. One of these was the water
produced at table, of which Captain Forbes funnily remarked, that it “tasted
badly of bung cloth—and dirty cloth, too!” But, such as it was, the Professor
and I preferred it to wine.
Thus much of culinary matters, for, with the exception of a few surprises,
which, according to all our previous ideas, confused the chronology of the
dishes—making a literal mess of it—and sundry minor variations in the cycle
of desserts proper, the service of one day resembled that of another.
There was only wind enough to fill the mainsail, and in it, on the lea side of
the boom, as if in a hammock, sheltered from the broiling sun, I lay resting for
hours. Off Peterhead, we saw innumerable fishing boats—counted 205 in one
fleet. Off Inverness, far out at sea, we counted as many, ere we gave in and
stopped. Their sails were mostly down, and we, passing quite near, could
observe the process of the fishermen shooting their nets; the sea to the north-
east all thickly dotted with boats, which appeared like black specks. A steamer
was sailing among them, probably to receive and convey the fish ashore.
Perilous is the calling of the fisherman! Calm to-day, squalls may overtake him
on the morrow—
“But men must work, and women must weep,
Though storms be sudden, and waters deep,
And the harbour bar be moaning.”
As the sun went down, from the forecastle we watched a dense bank of cloud
resting on the sea; its dark purple ranges here and there shewing openings,
with hopeful silver linings intensely bright—glimpses, as it were, into the land
of Beulah. Then the lights and shadows grandly massed themselves, gradually
assuming a sombre hue; while starry thoughts of dear ones at home rose,
welling up within us, as the daylight ebbed slowly away over the horizon’s rim.
Friday morning, July 22.—Rose at seven; weather dull; neither land, sky, nor sail,
visible; our position not very accurately known. At four in the morning the
engine had been stopped, the look-out having seen breakers a-head—no
observation to be had. Our course to the North Sea lay between the Orkney
and Shetland Islands. After breakfast it cleared, and on the starboard bow, we
saw Fair Isle, so that our course was right, although we had not known in what
part of it we were.
There was cause for thankfulness that the Orkneys had been passed in safety.
Where the navigation is intricate and requires care at best, our chances of
danger during the uncertainty of the night had doubtless been great. The south
of the Shetland Isles also appeared to rise from the sea, dim and blue, resting
on the horizon, like clouds ethereal and dreamlike.
At 11 o’clock A.M., sailing past Fair Isle, made several sketches of its varied
aspects, as seen from different points. Green and fair, this lonely island lies
about thirty miles south-west of the Shetland group, and in the very track of
vessels going north.
It has no light-house, and is dreaded by sailors; for many are the shipwrecks
which it occasions. Before now, we had heard captains, in their anxiety, wish it
were at the bottom of the sea. Could not a light be placed upon it by the
Admiralty, and a fearful loss of life thus be averted?
The island contains about a hundred inhabitants, who live chiefly by fishing
and knitting. They are both skilful and industrious. During the winter months,
the men, as well as the women, knit caps, gloves, and waistcoats; and for
dyeing the wool, procure a variety of colours from native herbs and lichens.
True happiness, springing as it ever does from above and from within, may
have its peaceful abode here among those lonely islanders quite apart from the
noise and bustle of what is called the great world, although the stranger sailing
past is apt to think such places “remote, unfriended, melancholy, slow.”[2]
Ere long we could distinguish the bold headland of Sumburgh, which is the
southern extremity of Shetland; and a little to the north-west of it, by the aid
of an opera-glass, Fitful Head,[3] rendered famous by Sir Walter Scott as the
dwelling place of Norna, in “The Pirate.”
Last summer I visited this the most northern group of British islands, famed
alike for skilful seamen, fearless fishermen, and fairy-fingered knitters; for its
hardy ponies, and for that soft, warm, fleecy wool which is peculiar to its
sheep.
Gazing on the blue outline of the islands, I now involuntarily recalled their
many voes, wild caves, and splintered skerries, alive with sea gulls and
kittiwakes. The magnificent land-locked sound of Bressay too, where her
Majesty’s fleet might ride in safety, and where Lerwick—the capital of the
islands, and the most northerly town in the British dominions—with its quaint,
foreign, gabled aspect, rises, crowning the heights, from the very water’s edge,
so that sillacks might be fished from the windows of those houses next the
sea. Boating excursions and pony scamperings are also recalled; the Noss
Head, with its mural precipice rising sheer from the sea to a height of 700 feet,
vividly reminding one of Edgar’s description of Dover Cliff, in “Lear,” or of
that which Horatio pictured to Hamlet—

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

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

10 A B O U T THE AUTHORS

different chapters MAKE CONNECTIONS

Figure 3.30 Contributions of

Figure 8.20 The Working Cell,

Figure 16.21 Genomics, Nutritional Mode

710 UNIT SIx AnimAl Form And Function

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ChApTEr 32 tHE intErnAl EnVironmEnt oF AnimAlS: orGAnizAtion And rEGulAtion 711

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Practice Scientific Skills

Scientific Skills Exercise

Making and Testing Predictions

392 UNIT THREE EVOLUTION

Every chapter has a Scientific Skills Exercise:

14 P R A C T I C E SCIENTIFIC SKILLS

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To learn more, visit www.masteringbiology.com

PR AC TICE SCIENTIFIC SKILLS 15

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NEW! Interpret the Data Questions

Potential Problems with Molecular Clocks

• How Can the Severity of Forest Fires Be

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MAKe ConneCtionS Paleontology

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guide RNA binds to part

chapter 13 The Molecular Basis of inheriTance 319

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14 Gene Expression: From Gene to Protein

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Every chapter opens

After reading a Key Concept section,

New Mutations and Mutagens

Third mRNA base (3′ end of codon)

C (D) the DNA introns are removed from the template.

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Instructors can easily incorporate active