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About the cover
The red salamander (Pseudotriton ruber) is common in cold, rocky springs
and streams in wooded and open areas of the eastern United Sates. Adults
sometimes inhabit moist sphagnum or soil under rocks or logs, leaf litter,
and crevices adjacent to streams. Like all salamanders, P. ruber is predatory,
typically feeding on invertebrate prey and on smaller salamanders.
Adults grow to 95–190 mm in total length. Although males may court
and inseminate females in wet terrestrial habitats, red salamanders move to
streams in the autumn to lay 30–130 eggs under submerged rocks and logs.
Eggs hatch in late winter or spring, and the metamorphosis of larvae into
adults occurs 1.5 to 3.5 years later.
Although not listed as threatened or endangered, many red salamander
populations have suffered from habitat loss and degradation, largely attrib-
utable to acid drainage from coal mines, other forms of pollution, and the
accumulation of silt in streams.
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Biology
The Dynamic Science
Fourth Edition
Russell Hertz McMillan
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Peter J. Russell, Paul E. Hertz, WCN: 02-200-203
Beverly McMillan
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Printed in Canada
Print Number: 01 Print Year: 2015
Brief Contents
1 Introduction to Biological Concepts 30 Fungi 684
and Research 1 31 Animal Phylogeny, Acoelomates, and Protostomes 706
32 Deuterostomes: Vertebrates and Their Closest
Unit One Molecules and Cells Relatives 743
2 Life, Chemistry, and Water 24
3 Biological Molecules: The Carbon Compounds of Life 44 Unit Five Plant Structure and Function
4 Cells 73 33 The Plant Body 786
5 Membranes and Transport 104 34 Transport in Plants 813
6 Energy, Enzymes, and Biological Reactions 126 35 Plant Nutrition 833
7 Cellular Respiration: Harvesting Chemical Energy 146 36 Reproduction and Development in Flowering
8 Photosynthesis 169 Plants 852
9 Cell Communication 192 37 Plant Signals and Responses to the Environment 881
10 Cell Division and Mitosis 214
Unit Six Animal Structure and Function
Unit Two Genetics 38 Introduction to Animal Organization and
Physiology 912
11 Meiosis: The Cellular Basis of Sexual Reproduction 237
39 Information Flow and the Neuron 930
12 Mendel, Genes, and Inheritance 251
40 Nervous Systems 951
13 Genes, Chromosomes, and Human Genetics 274
41 Sensory Systems 972
14 DNA Structure and Replication 300
42 The Endocrine System 994
15 From DNA to Protein 323
43 Muscles, Bones, and Body Movements 1016
16 Regulation of Gene Expression 354
44 The Circulatory System 1031
17 Bacterial and Viral Genetics 385
45 Defenses against Disease 1052
18 DNA Technologies: Making and Using Genetically Altered
Organisms, and Other Applications 407 46 Gas Exchange: The Respiratory System 1074
19 Genomes and Proteomes 432 47 Animal Nutrition 1093
48 Regulating the Internal Environment 1118
Unit Three Evolutionary Biology 49 Animal Reproduction 1144
20 Development of Evolutionary Thinking 457 50 Animal Development 1166
21 Microevolution: Genetic Changes within
Populations 476 Unit Seven Ecology and Behavior
22 Speciation 501 51 Ecology and the Biosphere 1193
23 Paleobiology and Macroevolution 521 52 Population Ecology 1222
24 Systematics and Phylogenetics: Revealing the Tree of 53 Population Interactions and Community Ecology 1251
Life 550 54 Ecosystems and Global Change 1283
55 Biodiversity and Conservation Biology 1309
Unit Four Biodiversity 56 Animal Behavior 1334
25 The Origin of Life 577
26 Prokaryotes: Bacteria and Archaea 593 Appendix A: Answers A-1
27 Protists 614 Appendix B: Classification System A-36
28 Seedless Plants 639 Glossary G-1
29 Seed Plants 661 Index I-1
iii
Peter J. Russell received a B.Sc. in Biology from the University of Sussex, England, in 1968 and
a Ph.D. in Genetics from Cornell University in 1972. He has been a member of the Biology faculty
of Reed College since 1972 and is currently a Professor of Biology, Emeritus. Peter taught a section
of the introductory biology course, a genetics course, and a research literature course on molecular
virology. In 1987 he received the Burlington Northern Faculty Achievement Award from Reed College
in recognition of his excellence in teaching. Since 1986, he has been the author of a successful genetics
textbook; current editions are iGenetics: A Molecular Approach, iGenetics: A Mendelian Approach, and
Essential iGenetics. Peter’s research was in the area of molecular genetics, with a specific interest in
characterizing the role of host genes in the replication of the RNA genome of a pathogenic plant virus,
and the expression of the genes of the virus; yeast was used as the model host. His research has been
funded by agencies including the National Institutes of Health, the National Science Foundation, the
American Cancer Society, the Department of Defense, the Medical Research Foundation of Oregon,
and the Murdoch Foundation. He has published his research results in a variety of journals, including
Genetics, Journal of Bacteriology, Molecular and General Genetics, Nucleic Acids Research, Plasmid,
and Molecular and Cellular Biology. Peter has a long history of encouraging faculty research involving
undergraduates, including cofounding the biology division of the Council on Undergraduate Research
in 1985. He was Principal Investigator/Program Director of a National Science Foundation Award for
the Integration of Research and Education (NSF–AIRE) to Reed College, 1998 to 2002.
Paul E. Hertz was born and raised in New York City. He received a B.S. in Biology from Stanford
University in 1972, an A.M. in Biology from Harvard University in 1973, and a Ph.D. in Biology
from Harvard University in 1977. While completing field research for the doctorate, he served on
the Biology faculty of the University of Puerto Rico at Rio Piedras. After spending two years as an
Isaac Walton Killam Postdoctoral Fellow at Dalhousie University, Paul accepted a teaching position
at Barnard College, where he has taught since 1979. He was named Ann Whitney Olin Professor of
Biology in 2000, and he received The Barnard Award for Excellence in Teaching in 2007. In addition
to serving on numerous college committees, Paul chaired Barnard’s Biology Department for eight
years and served as Acting Provost and Dean of the Faculty from 2011 to 2012. He is the founding
Program Director of the Hughes Science Pipeline Project at Barnard, an undergraduate curriculum
and research program that has been funded continuously by the Howard Hughes Medical Institute
since 1992. The Pipeline Project includes the Intercollegiate Partnership, a program for local com-
Aaron Kinard
munity college students that facilitates their transfer to four-year colleges and universities. He teaches
one semester of the introductory sequence for Biology majors and pre-professional students, lecture
and laboratory courses in vertebrate zoology and ecology, and a year-long seminar that introduces
first-year students to scientific research. Paul is an animal physiological ecologist with a specific
research interest in the thermal biology of lizards. He has conducted fieldwork in the West Indies
since the mid-1970s, most recently focusing on the lizards of Cuba. His work has been funded by the
NSF, and he has published his research in The American Naturalist, Ecology, Nature, Oecologia, and
Proceedings of the Royal Society. In 2010, he and his colleagues at three other universities received
funding from NSF for a project designed to detect the effects of global climate warming on the
biology of Anolis lizards in Puerto Rico.
Beverly McMillan has been a science writer for more than 25 years. She holds undergraduate
and graduate degrees from the University of California, Berkeley, and is coauthor of a college text
in human biology, now in its eleventh edition. She has also written or coauthored numerous trade
books on scientific subjects and has worked extensively in educational and commercial publishing,
including eight years in editorial management positions in the college divisions of Random House
and McGraw-Hill.
Courtesy of Beverly McMillan
iv
Preface
Welcome to the fourth edition of Biology: The Dynamic Science. understandable foundation for students who choose to enroll
The book’s title reflects the speed with which our knowledge of in more advanced biology courses in the future. Where appro-
biology is growing. Although biologists have made enormous priate, we provide straightforward explanations of fundamen-
progress in solving the riddles posed by the living world, every tal concepts from the evolutionary perspective that bind
discovery raises new questions and provides new opportunities together all of the biological sciences. Recognizing that stu-
for further research. As in previous editions, we have encapsu- dents in an introductory biology course face a potentially
lated the dynamic nature of biology in the fourth edition by daunting quantity of ideas and information, we strive to pro-
explaining biological concepts—and the data from which they vide an appropriate balance between factual and conceptual
are derived—in the historical context of each discovery and by material, taking great care to provide clear explanations of how
describing what we know now and what new discoveries will be scientists draw conclusions from empirical data. Our approach
likely to advance the field in the future. helps students understand how we achieved our present knowl-
edge. Having watched our students struggle to navigate the
many arcane details of college-level introductory biology, we
Building on a strong foundation . . . constantly remind ourselves and each other to “include fewer
The first three editions of this book provided students with the facts, provide better explanations, and maintain the narrative
tools they need to learn fundamental biological concepts and flow,” thereby enabling students to see the big picture. Clarity
processes. More important, the previous editions encouraged of presentation, thoughtful organization, a logical and seamless
students to think like scientists by applying the process of sci- flow of topics within chapters, and carefully designed illustra-
ence. Our approach encourages students to think about bio- tions are key to our approach. With this edition, full integra-
logical questions and hypotheses through clear examples of tion with MindTap engages students with appealing and useful
hypothesis development, observational and experimental tests exercises that encourage them to learn biology by thinking like
of hypotheses, and the conclusions that scientists draw from scientists.
data. The many instructors and students who have used previ-
ous editions have generously provided valuable feedback that Focusing on research to help students engage
has allowed us to strengthen the elements that enhance student
learning. We have also received comments from expert review-
the living world as scientists . . .
ers. As a result of these inputs, every chapter has been revised A primary goal of this book is to sharpen and sustain students’
and updated with recent studies, including many based on curiosity about biology, rather than dulling it with a mountain
genomic and proteomic analyses. In addition, the chapters in of disconnected facts. We can help students develop the mental
Unit One (Molecules and Cells) have been reorganized. This habits of scientists and a fascination with the living world by
edition also includes new or modified illustrations and photos, conveying our passion for biological research. We want to
as well as some new features. excite students not only with what biologists know about the
The fourth edition of Biology: The Dynamic Science repre- living world but also with how they know it and what they still
sents a fully integrated package of print and media that will need to learn. In doing so, we can encourage some students to
appeal to today’s students. Although the traditional format of accept the challenge and become biologists themselves, posing
the printed text can stand alone for both instructors and stu- and answering important new questions through their own
dents, MindTap, the most engaging and easily personalized innovative research. For students who pursue other careers, we
online solution in biology, enables instructors to deliver what hope that they will leave their introductory— and perhaps
they know is best for their students. MindTap offers an online only—biology course armed with intellectual skills that will
version of the text, as well as before-class and in-class exercises, enable them to evaluate future knowledge with a critical eye.
assignable and gradable homework exercises drawn from the In this book, we introduce students to a biologist’s “ways of
book’s content, and other resources and features that allow stu- learning.” Research biologists constantly integrate new obser-
dents to assess their learning as they progress through their vations, hypotheses, questions, experiments, and insights with
study of biology. existing knowledge and ideas. To help students engage the
world as biologists do, we must not simply introduce them to
the current state of knowledge. We must also foster an appre-
Emphasizing the big picture . . . ciation of the historical context within which those ideas devel-
In this textbook, we have applied our collective experience as oped, and identify the future directions that biological research
teachers, researchers, and writers to create a readable and is likely to take.
Chapter 8 h a r v e s t i n g c h e m i c a l e n e r g y: c e l lu l a r r e s p i r at i o n v
To achieve these goals, our explanations are rooted in the • To complement this historical or practical perspective,
research that established the basic facts and principles of biol- each chapter closes with a brief essay titled Unanswered
ogy. Thus, a substantial proportion of each chapter focuses on Questions, prepared by an expert or experts in the field.
studies that define the state of biological knowledge today. These essays identify important unresolved issues relating
When describing research, we first identify the hypothesis or to the chapter topic and describe cutting-edge research
question that inspired the work and then relate it to the that will advance our knowledge in the future.
broader topic under discussion. Our research-oriented theme • Most chapters include a short, boxed essay titled Molecular
teaches students, through example, how to ask scientific ques- Insights (formerly called Insights from the Molecular
tions and pose hypotheses, two key elements of the scientific Revolution), which describes how molecular tools allow
process. scientists to answer questions that they could not have
Because advances in science occur against a background of posed even 30 years ago. Most Molecular Insights focus on
research, we also give students a feeling for how biologists of a single study and include sufficient detail for their content
the past formulated basic knowledge in the field. By fostering to stand alone.
an appreciation of such discoveries, given the information and • Many chapters are further supplemented with one or more
theories available to scientists in their own time, we can help short, boxed essays called Focus on Research. Each essay
students understand the successes and limitations of what we focuses on one of three different aspects of research. Focus
consider cutting edge today. This historical perspective also on Research: Basic Research essays describe how research
encourages students to view biology as a dynamic intellectual has provided understanding of basic biological principles.
enterprise, not just a collection of facts and generalities to be Focus on Research: Applied Research essays describe
memorized. research designed to solve practical problems in the world,
We have endeavored to make the science of biology come such as those relating to health or the environment. Focus
alive by describing how biologists formulate hypotheses and on Research: Model Organisms essays introduce model
evaluate them using hard-won data; how data sometimes tell research organisms—such as Escherichia coli, Drosophila,
only part of a story; and how the results of studies often end up Arabidopsis, Caenorhabditis, the mouse, and Anolis—and
posing more questions than they answer. Although students explain why they are used as subjects for in-depth analysis.
might prefer simply to learn the “right” answer to a question, • Three types of specially designed research figures provide
they must be encouraged to embrace “the unknown,” those more detailed information about how biologists formulate
gaps in knowledge that create opportunities for further specific hypotheses and test them by gathering and inter-
research. An appreciation of what biologists do not yet know preting data. The research figures are listed on the end
will draw more students into the field. And by defining why papers at the back of the book. Experimental Research
scientists do not understand interesting phenomena, we figures describe specific studies in which researchers used
encourage students to think critically about possible solutions both experimental and control treatments—either in the
and to follow paths dictated by their own curiosity. We hope laboratory or in the field—to test hypotheses or answer
that this approach will encourage students to make biology a research questions by manipulating the system they stud-
part of their daily lives by having informal discussions and ied. Observational Research figures describe specific stud-
debates about new scientific discoveries. ies in which biologists have tested hypotheses by comparing
systems under varying natural circumstances. Research
Method figures provide examples of important techniques,
Presenting the story line of the research such as light and electron microscopy, the polymerase
process . . . chain reaction, making a knockout mouse, DNA microar-
ray analysis, plant cell culture, producing monoclonal anti-
In preparing this book, we developed several special features,
bodies, radiometric dating, and cladistic analysis. Each
all of which are included in MindTap, to help students broaden
Research Method figure leads a student through the pur-
their understanding of the material presented and of the
pose of the technique and protocol and describes how sci-
research process itself. A Visual Tour of these features and
entists interpret the data it generates.
more begins on page xiv.
• The chapter openers, titled Why it matters . . ., are engag- Integrating effective, high-quality visuals
ing, short vignettes designed to capture students’ imagina-
tions and whet their appetites for the topic that the chapter
into the narrative . . .
addresses. In many cases, this feature tells the story of how Today’s students are accustomed to receiving ideas and infor-
a researcher or researchers arrived at a key insight or how mation visually, making the illustrations and photographs in
biological research solved a major societal problem, a textbook and the fully integrated online resources critically
explained a fundamental process, or elucidated a phenom- important. From the first edition, our illustration program
enon. The Why it matters . . . feature also provides a brief has provided an exceptionally clear supplement to the narra-
summary of the contents of the chapter. tive in a style that is consistent throughout the book. Graphs
vi P r e fa c e
and anatomical drawings are annotated with interpretive Encouraging active learning, critical thinking,
explanations that lead students, step by step, through the
major points they convey.
and self-assessment of learning outcomes . . .
Over subsequent editions, we have enhanced the illustra- In the third edition we introduced an active learning feature,
tion program, focusing on features that reviewers and users of Think Like a Scientist, which is designed to help students think
the book identified as the most useful pedagogical tools. For analytically and critically about research presented in the chap-
this most recent edition, we focused explicitly on helping stu- ter. Think Like a Scientist questions appear at the end of
dents to think like scientists. A revised Figure 1.14 illuminates Experimental Research figures, Observational Research figures,
the intellectual steps that collectively lead researchers to new Closer Look figures, Molecular Insights boxes, and Unanswered
scientific discoveries. These steps—observation, hypothesis, Questions. In this new edition, Experimental Research figures
prediction, experiment, and interpretation—represent the and Observational Research figures include a new icon that
fundamentals of our “think like a scientist” theme in this identifies the particular step in the process of science that the
book. Think Like a Scientist question addresses: Observe, Hypothe-
In revising the text, we reevaluated each illustration and size, Predict, Experiment, or Interpret. (Sample shown.)
photograph and made appropriate changes to improve their Observe
utility as teaching tools. New illustrations for the fourth edi- think like Hypothesize
Predict
tion were created in the same style as existing ones. In addi- a scientist Experiment
tion, some illustrations of key biological processes were Interpret
recast as Closer Look figures in which a Summary and a con- This edition also continues the popular Think Outside the
cluding Think Like a Scientist question enhance student Book active learning feature. Think Outside the Book activities
learning. have been designed to encourage students to explore biology
directly or through electronic resources, working either indi-
vidually or collaboratively.
Organizing chapters around important Supplementary materials at the end of each chapter—a ll of
concepts . . . which are fully integrated into MindTap—help students review
the material they have learned, assess their understanding, and
As authors and college teachers, we understand how easily stu-
think analytically as they apply the principles developed in the
dents can get lost within a chapter. When students request
chapter to novel situations. Many end-of-chapter questions also
advice about how to read a chapter and learn the material in it,
serve as good starting points for class discussions or out-of-
we usually suggest that, after reading each section, they pause
class assignments.
and quiz themselves on the material they have just encoun-
Review Key Concepts provides a summary of important
tered. After completing all of the sections in a chapter, they
ideas developed in the chapter, referencing specific figures and
should quiz themselves again, even more rigorously, on the
tables in the chapter. These Reviews are no substitute for read-
individual sections and, most important, on how the concepts
ing the chapter, but students may use them as a valuable outline
developed in the different sections fit together. Accordingly, we
of the material, filling in the details on their own.
have adopted a structure for each chapter to help students
Test Your Knowledge includes four types of end-of-chapter
review concepts as they learn them.
questions and problems that focus on the chapter’s factual con-
• The organization within chapters presents material in tent while encouraging students to apply what they have
digestible sections, building on students’ knowledge and learned: (1) Multiple-choice questions (with answers in an
understanding as they acquire it. Each major section cov- appendix) focus on factual material; (2) Discuss Concepts ques-
ers one broad topic. Each subsection, titled with a declara- tions involve open-ended issues that emphasize key ideas, the
tive sentence that summarizes the main idea of its content, interpretation of data, and practical applications of the mate-
explores a narrower range of material. rial; (3) Design an Experiment questions help students hone
• Whenever possible, we include the derivation of unfamil- their critical thinking skills by asking them to test hypotheses
iar terms so that students will see connections between that relate to the chapter’s main topic; and (4) Apply Evolutionary
words that share etymological roots. Mastery of the tech- Thinking questions ask students to answer a question in rela-
nical language of biology will allow students to discuss tion to the principles of evolutionary biology.
ideas and processes precisely. At the same time, we have In this edition, Test Your Knowledge questions are orga-
minimized the use of unnecessary jargon. nized according to Bloom’s taxonomy into three sections:
• Study Break questions follow every major section. These Remember/Understand, Apply/Analyze, and Evaluate/Create.
questions encourage students to pause at the end of a sec- This structure allows students to review the material in a
tion and review what they have learned before going on to sequence that moves from the basic knowledge of factual mate-
the next topic within the chapter. Short answers to these rial to more challenging and sophisticated applications of that
questions appear in an appendix. knowledge to novel situations.
P r e fa c e vii
Interpret the Data questions, highlighted in a distinctive
format, help students develop analytical and quantitative skills
by asking them to interpret graphical or tabular results of
experimental or observational research studies for which the
hypotheses and methods of analysis are presented.
Effectively introducing digital solutions into

your classroom—online or in class—is now
easier than ever . . .
altrendo images/Getty Images.

The fourth edition of Biology: The Dynamic Science represents a
fully integrated package of print and media, providing compre-
hensive learning tools and flexible delivery options. In prepar-
ing this edition we conducted extensive research to determine
how instructors prefer to present online learning opportuni-
ties. The result of this research is a new MindTap course orga-
nized around the instructor’s preferred workflow. Instructors • Many Experimental Research and Observational Research
can now select just the content they want to assign, chosen figures now include an additional activity to help students
from a comprehensive set of learning materials provided with engage with the scientific process.
the course for each chapter. Many types of learning activities • The end-of-chapter Design an Experiment feature is pre-
are assignable and offer students immediate feedback and auto- sented online as a guided learning activity that takes the
mated instructor assessment. student through the process of designing an experiment.
Research also indicates that online content is most effec-
The Instructor Resource Center provides everything you
tive when it enhances conceptual understanding through the
need for your course in one place. This collection of lecture
use of relevant applications. In this edition, we have devel-
and class tools is available online for instructors only via
oped three new assessable, online learning activities that
www.cengage.com/login. There you can access and download
align with important book features and provide students the
PowerPoint presentations, images, the Instructor’s Manual,
opportunity to explore and practice biology the way scientists
videos, and more.
practice biology:
We hope you agree that we have developed a clear, fresh,
• The Interpret the Data feature at the end of every chapter is and well-integrated introduction to biology as it is understood
enhanced by an additional online exercise to further by researchers today. Just as important, we hope that our efforts
develop student quantitative analysis and mathematical will excite students about the research process and the biologi-
reasoning skills. cal discoveries it generates.
Peter J. Russell Paul E. Hertz Beverly McMillan
viii P r e fa c e
New to This Edition
The enhancements we have made in the fourth edition of new subsection on molecule geometry and function in the cell,
Biology: The Dynamic Science reflect our commitment to pro- and a new Unanswered Questions essay on the effect of climate
vide a text that introduces students to new developments in change on marine ecosystems.
biology while fostering active learning and critical thinking. Chapter 3 (Biological Molecules: The Carbon Compounds
The new icon for Think Like a Scientist questions not only helps of Life) has molecular models added to Table 3.1 on functional
readers identify those questions easily, but it also helps students groups and to Table 3.2 on major protein functions.
reinforce their understanding of the process of science. In Chapter 4 (Cells), Section 4.2 on Prokaryotic Cells has
We have also made important changes in coverage to follow been reorganized into subsections on structure and organiza-
recent scientific advances. In the third edition, we added a new tion, and evolutionary divergence of bacteria and archaea, and
Chapter 19, Genomes and Proteomes, that discusses methods of Section 4.3 on Eukaryotic Cells has added proteomics content
genomics and proteomics along with examples of new discover- in the endomembrane system discussion, updated discussion of
ies and insights. To reflect the importance of these approaches, the Golgi complex and vesicle traffic, and enhanced discussion
genomics and proteomics coverage has been integrated through- of the movement of organelles and vesicles. A new Molecular
out the book. In addition, 35 Molecular Insights boxes have been Insights describes a genome-wide analysis study identifying
revised to include recently published research, and many of human proteins that regulate secretion.
them focus on genomics or proteomics. Beyond these changes, Chapter 5 (Membranes and Transport) adds the kiss-and-
we have made numerous improvements to update and clarify run model of exocytosis.
scientific information and to engage students as interested read- Chapter 7 (Cellular Respiration: Harvesting Chemical
ers and active learners, as well as responsive scientific thinkers. Energy) adds more examples of human disorders related to cel-
The following sections highlight some of the new content and lular respiration, discussion of evolution of mitochondria, and
organizational changes in this edition. discussion of anaerobic respiration. A new Section 7.6 on Inter-
relationships of Catabolic and Anabolic Pathways combines
material previously in earlier sections with new material to dis-
Chapter 1 cuss how food substances feed into glycolysis, how biosynthetic
In the introductory chapter, Section 1.1 (What is Life? Character- pathways link to glycolysis and the citric acid cycle (with a new
istics of Living Organisms) has been revised to provide a more figure), and the regulation of cellular respiration (with a new
detailed discussion of the function and transmission of genetic figure). A new Molecular Insights describes experiments study-
information in living systems; and the accompanying figure ing the mitochondrial proteome and its variation among organs
(Figure 1.4) provides a more accurate portrayal of nucleic acids and organisms.
and the flow of information from DNA to RNA to proteins. In In Chapter 8 (Photosynthesis), a new Experimental Research
addition, Section 1.4 (Biological Research) now has a more explicit figure describes the Engelmann experiment. Discussion of the
discussion of the process of science, including revisions to evolution of photosynthesis and cellular respiration has been
Figure 1.14 (now a Closer Look figure), which introduces the new enhanced. A new Molecular Insights describes proteomics experi-
icon for Think Like a Scientist questions in Experimental Research ments on the effect of water deficit on growth and photosynthesis
figures and Observational Research figures. Finally, the introduc- in a C3 plant.
tion of genomics and proteomics has been refined, and systems Chapter 9 (Cell Communication) has expanded discussion
biology is introduced (with a new Figure 1.17) as an inclusive of the different types of signaling along with an updated figure.
approach for exploring complex biological phenomena. Sections on receptor tyrosine kinases and G-protein–coupled
receptors are now combined with new material on ligand-gated
ion channels into a single section on signaling pathways trig-
Unit One: Molecules and Cells gered by surface receptors. Nitric oxide is added to the internal
Unit One has been reorganized. The former Chapter 4 (Energy, receptor section.
Enzymes, and Biological Reactions) is now Chapter 6, preced- Chapter 10 (Cell Division and Mitosis) has enhanced dis-
ing Chapter 7 (Cellular Respiration: Harvesting Chemical cussion of chromosomes now including presentation of chro-
Energy) and Chapter 8 (Photosynthesis) to provide a more logi- mosome structure and levels of chromosome organization
cal conceptual flow of metabolism. The former Chapter 7 (Cell (including a figure) moved from Chapter 14 (DNA Structure
Communication) is now Chapter 9. and Replication). A new Molecular Insights describes pro-
Chapter 2 (Life, Chemistry, and Water) has a new Why it teomics experiments on the regulation of proteins in the cell
matters . . . relating biology to underlying chemical reactions, a cycle of humans.
ix
Chapter 18 is retitled DNA Technologies: Making and
Unit Two: Genetics Using Genetically Altered Organisms, and Other Applications,
In Chapter 12 (Mendel, Genes, and Inheritance), the terms gene to reflect a reorganization of the chapter around the theme of
marker, genetic marker, and DNA marker are now introduced making and using genetically altered organisms for basic and
to relate to discussions later in the unit. applied research. The three main sections of the chapter are
Chapter 13 (Genes, Chromosomes, and Human Genetics) now Key DNA Technologies for Making Genetically Altered
adds discussions of pedigrees and pedigree analysis with four Organisms, Applications of Genetically Altered Organisms,
new figures, and of the usefulness of mitochondrial DNA anal- and Other Applications of DNA Technologies (not involving
ysis in genealogy and forensics. A new Molecular Insights genetically altered organisms). PCR is now emphasized more as
describes experiments on the involvement of noncoding RNAs a widely used technique, and Southern and northern blotting
in X-chromosome inactivation. are no longer discussed. A new figure is added on making
Chapter 14 (DNA Structure and Replication) adds discus- knockout mice by gene targeting, and a new Focus on Research
sion of the loading and unloading of the sliding clamp in box describes programmable RNA-guided genome editing
human DNA replication, a new figure and description of the based on CRISPR-Cas. The Why it matters . . . is new, discuss-
replisome complex for DNA replication, discussion of the role ing the relationship between historical genetics approaches to
of telomeres as chromosome caps, and a description of how modern-day genomics approaches. A new Molecular Insights
newly replicated DNA is assembled into nucleosomes. describes metabolomic experiments on the nutritional quality
Section 14.4 is retitled Repair of Errors in DNA to reflect the of genetically modified food.
addition of new material on excision repair mechanisms, and a In Chapter 19 (Genomes and Proteomes), lncRNAs are
new figure on thymine dimers and their repair. added to the discussions of important genome sequences, the
Chapter 15 (From DNA to Protein) now discusses Garrod’s ENCODE project is added to the discussion of the profile of the
classic experiment in Why it matters . . . adds discussion of the human genome, and tables are updated.
coupling of transcription, pre-mRNA processing and export of
mRNA from the nucleus, and adds new material on spontane-
Unit Three: Evolutionary Biology
ous and induced mutations and mutagens, transposable ele-
ment content of genomes, and the types of changes in gene Chapter 20 (The Development of Evolutionary Thinking) has
expression eukaryotic transposable elements can cause. A new been revised to contextualize the study of evolution within the
Molecular Insights describes genomics/proteomics experiments science process described in Chapter 1, and Figure 20.10 pre-
on the effect of sleep–wake timing on gene expression. senting Darwin’s observations, hypotheses, and predictions has
In Chapter 16 (Regulation of Gene Expression), Section 16.2 been restructured to reflect the process of science theme we
on Regulation of Transcription in Eukaryotes is reorganized to have developed. In addition, the discussion of the fossil record
present chromatin modification and methylation before the has been updated to reflect new discoveries about the relation-
molecular details of transcription initiation at promoters. Dis- ships of birds to non-avian dinosaurs, and Figure 20.13 now
cussion of DNA methylation and gene regulation is expanded, includes a phylogenetic tree to highlight when key adaptations
including analysis of DNA methylomes. New material is added arose. Chapter 20 also includes a new Molecular Insights on the
on the interference of transcription by long noncoding RNAs genetics of dog domestication.
(lncRNAs), on the cataloging of human promoters and enhanc- In Chapter 21 (Microevolution: Genetic Changes within
ers by genome analysis, and on the role of alternative splicing in Populations), the discussion of variation in DNA sequences
adjusting gene output to match physiological requirements. In now includes the importance of single-nucleotide polymor-
Section 16.4, Genetic and Molecular Regulation of Develop- phisms in biological research. The Focus on Research box about
ment, the discussion of Hox genes is rewritten and expanded, the Hardy–Weinberg genetic equilibrium has been subdivided
and lncRNAs are added to the discussion of noncoding RNAs to make the steps in the analysis more explicit. The figure on
and their roles in development. Section 16.5, Genetics of Cancer, the distribution of color and striping patterns in European gar-
is changed to The Genetics and Genomics of Cancer to reflect den snails (Figure 21.15) has been revised to emphasize the
added content. Discussion of genes involved in cancer is rewrit- associations between snail phenotypes and habitat types. The
ten to include new knowledge from genomic approaches on the chapter includes a new example of frequency-dependent selec-
types of genes involved in cancer, and the cellular processes tion, using flower color in a European orchid (Figure 21.16). In
affected by mutations associated with cancer. Also added is dis- Table 21.2, we have added a column about the fitness conse-
cussion of cancer therapy and the prospect of personalized quences of each of the agents of evolution. Finally, Chapter 21
medicine, and how cancer genomic analysis identifies cancer includes a new Molecular Insights about the genetic and physi-
subtypes. A new Molecular Insights describes experiments on ological mechanisms that produce exaggerated horns in male
the role of a key long noncoding RNA in cardiac development. rhinoceros beetles.
In Chapter 17 (Bacterial and Viral Genetics), a new In Chapter 22 (Speciation), a new introduction to the dis-
Molecular Insights describes genomics/proteomics experiments cussion of reproductive isolation in Section 22.2 provides a
that identified the largest known giant virus. broader context for the discussion of the mechanisms of
x N e w to T h i s E d i t i o n
reproductive isolation. The discussion of parapatric speciation understanding of land plant evolution. The chapter’s new
was deleted from Section 22.3 on the geography of speciation in Molecular Insights, Comparative Genomics Probes Plant
favor of a more extended discussion of the possible conse- Evolution, reinforces these ideas. The chapter now concludes
quences of secondary contact after a period of allopatry. The with a new section that surveys the ecological, economic, and
example of a hybrid zone between oriole (Icterus) species has research importance of seedless plants. New figures accompany
been enhanced with additional text and a revised figure (22.12). chapter discussions of early adaptation for water transport and
The discussion of allopolyploidy and speciation in wheat has the evolutionary shift from homospory to heterospory.
been updated with new genomic studies and a greatly revised Chapter 29 (Seed Plants) begins with a new Why it
figure (22.16). Chapter 22 includes a revised figure in the Focus matters . . . that introduces students to the core concept that the
on Research box about speciation in Hawaiian Drosophila, as evolution of the seed, together with pollen and pollination, was
well as a new Molecular Insights about the genetics of ecological crucial in the radiation of vascular plants into nearly every land
and behavioral isolation between two Drosophila species. environment. A new Section 29.1, The Rise of Seed Plants,
In Chapter 23 (Paleobiology and Macroevolution), the examines key innovations in the evolution of the seed; new line
table (Table 23.1) outlining geological time and the history of art illustrates key stages hypothesized for one of those innova-
life has been consolidated and now includes more references to tions, the evolution of the ovule. This new section provides the
major geological events. The chapter also includes a revised dis- conceptual foundation for subsequent sections on gymno-
cussion of biogeographical realms with a revised figure (23.10) sperms and angiosperms, both of which have been reorganized
based on a study published in 2013. The discussions of both and expanded. Section 29.3, on angiosperms, includes new sub-
adaptive radiations and mass extinctions have also been sections that underscore the adaptive roles of flowers, double
updated along with a new figure (23.14) showing the pattern of fertilization, fruits and seeds. Expanded coverage in Section 29.4
mass extinctions more explicitly. The discussion of feathers in (previously 29.3), Insights from Plant Genome Research,
non-avian dinosaurs and birds has been updated with new includes the hypothesized role of whole genome duplication in
information, and a new figure (23.24) illustrates age-related plant polyploidy and examples of how genome sequencing is
changes in dinosaur plumage. advancing research on the biology and evolutionary relation-
In Chapter 24 (Systematics and Phylogenetics: Revealing ships of various seed plant lineages. A new, concise Section 29.5,
the Tree of Life), the taxonomic hierarchy is outlined in a new Seed Plants and People, reminds students of the major roles of
table (Table 24.1) that cross-references information in seed plants in human affairs. A new Molecular Insights describes
Figure 1.10. The discussion about reading a phylogenetic tree insights that are emerging from the sequencing of the loblolly
has been consolidated, and the accompanying figures have pine genome.
been combined into a Closer Look figure (24.2). The discussion Revisions to Chapter 30 (Fungi) include a discussion of the
of homology and homoplasy has been clarified. Chapter 24 also proposed new fungal phylum Cryptomycota and a new
includes a new Molecular Insights (with accompanying phylo- Molecular Insights, Researching Relationships of “Hidden
genetic tree) about the evolution of electric organs in fishes. Fungi.” Much of the third edition’s discussion of mycorrhizae
now appears in Chapter 34 on plant nutrition. A new conclud-
ing section discusses the effects of fungi on ecosystems and
Unit Four: Biodiversity human endeavors.
In Chapter 25 (The Origin of Life), Section 25.1 on the forma- Changes to Chapter 31 (Animal Phylogeny, Acoelomates,
tion of molecules necessary for life was revised to add material and Protostomes) include an updated description of how the
about what is needed to understand the origin of life, about molecular phylogeny for animals was constructed. The chapter
questions asked in the scientific study of the origin of life, and now also includes descriptions of a recently discovered preda-
about the possible role of alkaline hydrothermal vents in the tory sponge, all-female species of bdelloid rotifers, and a revised
origin of life. Section 25.2 on the origin of cells was largely analysis of annelid systematics to reflect recent research. The
rewritten, now to discuss the evolution of molecular replica- discussion of insects includes a new description of the evo–
tors, the evolution of cellular membranes, and the evolution of devo origin of insect wings. Although Molecular Insights
biological energy sources, as well as adding to the discussion of addresses the same topic (relationships among arthropods) as
prokaryotic cells as the first living cells. in prior editions, it is now based on research published in 2010
Chapter 26 (Prokaryotes: Bacteria and Archaea) adds a dis- and includes a revised phylogenetic tree for arthropods.
cussion of microbiomes. In Chapter 32 (Deuterostomes: Vertebrates and Their Clos-
Chapter 27 (Protists) adds a discussion of the apicoplast est Relatives), revisions include an expanded discussion of con-
and its origin. A new Molecular Insights describes experiments odont elements and of placoderms as a paraphyletic group. The
on two different types of bacteria associated with a cellular chapter also includes a greatly expanded discussion of the ori-
slime mold. gin of limbs with a completely revised figure (32.21). The dis-
Chapter 28 (Seedless Plants) includes examples of compar- cussion of amniote relationships, including the phylogenetic
ative genomics that shed light on the evolution of vascular tis- tree (Figure 32.24), reflects new insights into the relationships
sue in land plants and of new fossil finds that expand our of turtles to archosaurs and recognizes the newly defined group
N e w to T h i s E d i t i o n xi
Archelosauromorpha. The phylogenetic tree for primates gametes now includes a new subsection on evolutionary
(Figure 32.34) now includes characters that distinguish trends in flower structure with accompanying photographs
clades, and the discussion of human evolution now includes that allow students to readily visualize and compare key dif-
a description of the Denisovans and an enhanced hominin ferences in flower structure. Two other new subsections
timeline (Figure 32.39). expand the chapter’s coverage of fruit diversity and seed dis-
persal mechanisms. Section 36.4, Asexual Reproduction of
Flowering Plants, provides expanded examples of vegetative
Unit Five: Plant Structure and Function reproduction. The discussion of tissue culture methods
This edition’s chapters on plant anatomy and physiology also emphasizes somatic embryogenesis, including the creation of
have been significantly revised, updated, and reorganized. In artificial seeds.
Chapter 33 (The Plant Body), a new Why it matters . . . intro- Chapter 37 (Plant Signals and Responses to the Environ-
duces the main chapter topic using the historical domestica- ment) also features important revisions and additions. The
tion of grasses (rice, wheat, corn) to underscore the relevance introduction to plant hormones in Section 37.1 is reorganized
of plant parts to human concerns. Sections on root and shoot to consider auxins, cytokinins, and gibberellins, in that order.
systems are reordered, with root systems considered first It also includes strigolactones (SLs) as a major plant hormone
(Section 33.3) followed by primary shoot systems (Section 33.4). family, with accompanying line art reinforcing the varied
A new Molecular Insights features current research on the functions of SLs in enhancing nutrient access and optimizing
complex genetic events governing the formation of secondary the growth of shoot and root parts. The subsection on ethyl-
cell walls. ene now includes the triple response in seedling growth. A
In Chapter 34 (Transport in Plants), a new Why it brief comparison of constitutive and inducible defenses is
matters . . . uses the example of the centuries-old Angel Oak added to Section 37.2, Plant Chemical Defenses. One new
on Johns Island in South Carolina to introduce transport in subsection, Defensive Chemicals Reflect the Coevolution of
plants. The discussion of water potential is revised for clarity, Plants, Pathogens, and Herbivores, discusses bioactive spe-
with new subsections providing a more straightforward pre- cialized compounds including phytoalexins, plant alkaloids,
sentation on solute and pressure potential, turgor pressure, terpenes, and phenolics. Another new subsection on inducible
and wilting. Section 34.2, formerly Transport in Roots, is responses to specific threats introduces pattern recognition
now titled Roots: Moving Water and Minerals into the Plant. receptors and their role in detecting pathogen-associated
A new Figure (34.7) provides an enhanced visual to support molecular patterns (PAMPs). A new Molecular Insights dis-
the text discussion of the Casparian strip. Section 34.3, cusses the recent work of Chunyang Wang and colleagues in
Transport of Water and Minerals in the Xylem, now includes exploring the evolution of hormone signaling in plants.
a subsection on effects of humidity, temperature, and wind
on transpiration. The functioning and regulation of stomata
are now considered in a separate section (Section 34.4). A
Unit Six: Animal Structure and Function
new Molecular Insights, Going with the Phloem, looks at Chapter 38 (Introduction to Animal Organization and Physi-
recent experiments exploring the role of the transcription ology) expands discussion of anchoring junctions, tight junc-
factors called NAC proteins in shaping the development of tions, and gap junctions, and of the basics of epithelial
sieve-tube elements. structure and structure, including new discussion and a fig-
Enhancements to Chapter 35 (Plant Nutrition) include ure on pseudostratified columnar epithelium. A new Molec-
new photographs (Figure 35.2) to illustrate a range of nutri- ular Insights describes miRNA regulation of epithelial cell
ent deficiency symptoms in plants. A new subsection dis- differentiation in the lung and its relationship to lung cancer.
cusses the use of fertilizers to remedy nutrient deficiencies in In the presentation of homeostasis, a new subsection dis-
soil. Section 35.2 on soil characteristics now includes the role cusses how set points can change due to biorhythms or
of weathering in soil formation. New subsections clearly dis- altered environmental conditions.
tinguish the effects of organic and inorganic components of Chapter 39 (Information Flow and the Neuron) adds
soils and discuss the role of proper soil management (includ- information about the proteomic analysis of chemical syn-
ing no-till farming) in sustainable agriculture. Section 35.3, apses, and the discussion of equilibrium potential now
Root Adaptations for Obtaining and Absorbing Nutrients, includes presentation of the Nernst equation.
now presents much of the discussion of mycorrhizae that Chapter 40 (Nervous Systems) adds discussion of the
previously was in Chapter 30 (Fungi). The subsection on evolutionary changes in brain regions, brain size, and rela-
nitrogen fixation now includes a concise description of tive size of brain regions in vertebrate brains, and expanded
denitrification. description of the knee-jerk reflex. A new Molecular Insights
In Chapter 36 (Reproduction and Development in Flow- describes experiments on sex differences in the neural con-
ering Plants), Section 36.2 on the formation of flowers and nections of the human brain.
xii N e w to T h i s E d i t i o n
Chapter 41 (Sensory Systems) adds a new subsection on the the take-home message about the effects of predation on
evolutionary history of olfactory receptor genes revealed by guppy life histories. The chapter also includes a new Molecular
bioinformatics analysis of genomes. A new Molecular Insights Insights on the construction of a life table for ant colonies.
describes experiments on taste neuron changes associated with Chapter 53 (Population Interactions and Community
the emergence of an adaptive behavior in cockroaches. Ecology) includes new examples of Batesian mimicry
Chapter 42 (The Endocrine System) adds discussion of (between birds and insects) and fundamental and realized
nongenomic action of some steroid hormones. A new niches (cane toads) with dramatic new figures. In addition,
Molecular Insights describes experiments on the fear- the figure about primary succession (53.28) has been con-
enhancing effects of some oxytocin receptors in mice. verted into the Closer Look format.
Chapter 43 (Muscles, Bones, and Body Movements) Chapter 54 (Ecosystems and Global Change) has under-
expands the discussion of synovial joints. A new Molecular gone substantial revision. The distinction between detrital
Insights describes aspects of the genetics of bone formation and grazing food webs has been eliminated, and the discus-
learned from analysis of the elephant shark genome. sion of biological magnification (previously a Focus on
Chapter 44 (The Circulatory System) adds a figure on the Research box) has been tightened and incorporated into the
control of red blood cell production. text. Ecological pyramids are now represented in a single fig-
Chapter 45 (Defenses against Disease) adds a new flow- ure (54.8) instead of three; and new figures on the effects of
chart figure of antibody-mediated and cell-mediated immune temperature and precipitation on primary productivity
responses, and adds discussion of microbiome composition (54.4), seasonal changes in primary productivity (54.5), the
in preventing pathogen attack. A new Molecular Insights greenhouse effect (54.15), rising carbon dioxide levels and
describes genomics experiments revealing the unique global temperature (54.16), and anthropogenic nitrogen fixa-
immune system of the Atlantic cod. tion (54.17) have been added. In addition, Section 54.4 (now
In Chapter 46 (Gas Exchange: The Respiratory System), a titled Human Activities and Anthropogenic Global Change)
new Molecular Insights describes genomics-based experi- includes expanded and updated discussions of disruptions to
ments on the evolution of altitude adaptation in Tibetans. the carbon and nitrogen cycles and a completely new discus-
Chapter 47 (Animal Nutrition) adds discussion of gut sion of the impact of global change on ecosystems, including
microbiomes and their roles in digestion and nutrition. A new ocean acidification, declining primary productivity, and
Molecular Insights describes experiments on the association dead zones in shallow marine environments. The Molecular
of intestinal bacterial populations with obesity in humans. Insights box has been condensed.
Chapter 48 (Regulating the Internal Environment) Chapter 55 (Biodiversity and Conservation Biology) includes
expands discussion of temperature regulation in endotherms. updated and expanded discussions of overfishing, invasive spe-
A new Molecular Insights describes experiments on the cies (updated Figure 55.10 on hemlock woolly adelgids), dam
involvement of miRNAs with the development of polycystic removals, amphibian declines, and vulture mortality in South
kidney disease. Asia. It also features a new example of habitat fragmentation with
In Chapter 49 (Animal Reproduction), discussion of hor- a new figure (55.6). Section 55.3 (now titled Ecosystem Services
monal regulation of male reproductive functions is enhanced That Biodiversity Provides) includes discussions of provisioning,
and includes a new replacement figure. regulating, and support services. Section 55.4 (Which Species and
In Chapter 50 (Animal Development), a new Molecular Ecosystems Are Most Threatened by Human Activities?) now
Insights describes experiments showing an essential role of includes a discussion of the IUCN Red List of Threatened Species
protein O-mannosylation in embryonic development. and an accompanying new figure (55.13). Section 55.5 includes a
new discussion on the cost of preserving biodiversity. The
Molecular Insights box on DNA barcoding has been updated and
Unit Seven: Ecology and Behavior includes a new figure, and the Focus on Research box about popu-
In Chapter 51 (Ecology and the Biosphere), the chapter lation viability analysis has been tightened and condensed.
opener is now more focused on natural climate cycles and Chapter 56 (Animal Behavior) has added emphasis on
less on disastrous weather events. The chapter also includes cost/benefit analyses in the chapter opener. The discussion of
expanded discussions of positive feedback loops between cli- honeybee communication has been updated to reflect new
mate warming and melting of permafrost in tundra, as well research on the waggle dance (with revisions to Figure 56.19).
as more detailed descriptions of deep-sea environments. A We have also substantially revised the discussion of altruism,
new Molecular Insights describes the genetic basis of adapta- inclusive fitness, and kin selection to include Hamilton’s
tion to extreme cold in polar octopuses. inequality as a means of predicting altruistic behavior; and
In Chapter 52 (Population Ecology), we have updated Fig- the discussion of haplodiploidy and eusociality has been
ures 52.22 and 52.23 on human population growth. We have expanded to include alternative hypotheses for the evolution
subdivided the Focus on Research box into sections to clarify of this complex behavior.
N e w to T h i s E d i t i o n xiii
Welcome to Biology: The Dynamic Science 4e
Russell/Hertz/McMillan, Biology:
The Dynamic Science 4e and
MindTap engage students so they
learn not only what scientists know,
but how they know it, and what they
still need to learn.
Customize your students’ learning experience

with unparalleled content options presented in
easily edited folders. ▼
▲ Elevate learning and conceptual understanding with assignable and gradeable exercises that build
on material presented in the text. Fully editable content delivery options are organized into categories to match
course workflow. The InClass folder is expanded to show the pre-loaded content in this category.
xiv
Science as a Process
Immerse students in the process of doing biology, while building skills
students need to succeed in more advanced courses.
Matej Kastelic/Shutterstock.com
wavebreakmedia/Shutterstock.com
Milles Studio/Shutterstock.com
Science as a Process is emphasized throughout the ▶

text. Research Figures provide information about how
biologists formulate and test specific hypotheses by
gathering and interpreting data.
◀ Build Quantitative Skills: Interpret

the Data exercises in every chapter develop
quantitative analysis and mathematical
reasoning skills. Exercises are assignable
and gradeable in MindTap!
▲ Apply the Process of Science: Think Like a Scientist

questions throughout the text ask students to apply what ▲ Virtual Biology Laboratory (VBL)
they have learned beyond the material presented in the enables students to “do” science by acquiring
book. New icon on selected figures relates the questions to data, performing experiments, and using that
steps in the scientific process. Selected Think Like a Scientist data to explain biological concepts or
exercises are assignable and gradeable in MindTap! phenomena.
xv
Engagement Taken to the Next Level
Comprehensive content and flexible course delivery options...because
you know what is best for your students.
Engage with your students before

class. We provide a collection of learning
exercises and activities including high quality
videos for you to choose from.
Or insert your own content such as

PowerPoints, videos, and animations! ▼
Hero Images/Corbis
Straightforward explanations and carefully developed illustrations are followed by extensive opportunity
▲ Spectacular illustrations such as this Closer Look ▲ Summary figures help students see the big
figure help students visualize complex processes. Numbered picture and understand important connections.
step-by-step explanations lead students through all the
major concepts.
xvi
Full student engagement. On these two pages, take a tour of how you can design a dynamic
learning path for students to help them learn important concepts. Our case example is from
Chapter 7: Cellular Respiration: Harvesting Chemical Energy.
iStockphoto.com/Steve Debenport
“The explanations when you receive the

wrong answer on a homework question
Engage with your students during class. We provide a were really good and helped me under-
collection of dynamic activities. Or (of course!) insert your stand many concepts that I didn’t originally
own content. Assess student understanding on a topic during understand.”
class with interactive figures and immediate feedback with —Timothy, California State University of Northridge
explanations. ▶
for reinforcement and practice. Sample content for learning about Cellular Respiration is shown below.
▲ Post Learning Assessments offer

students opportunities for more practice
before a full chapter test. Practice Tests are
assignable and gradeable in MindTap!
◀ Conceptual exercises in MindTap, repeatable in

alternate versions, help students learn the material.
Exercises are assignable and gradeable in MindTap!
xvii
Active Learning
Features that engage your students in the process of learning because an
engaged student is a successful student.
Unanswered Questions explore important unresolved
issues identified by experts in the field and describe cutting-
edge research that will advance knowledge in the future.
Think Like a Scientist questions encourage students to think
critically about the research projects described. ▼
▲ Molecular Insights boxes in each chapter describe

how molecular tools allow scientists to answer questions
that they could not have posed even 30 years ago. Most
Molecular Insights focus on a single study and include
sufficient detail for its content to stand alone. 25 new
Molecular Insights boxes are focused on genomics or
proteomics.
▲ Study Break questions at the end of every section in a

chapter encourage students to pause and think about the
material just encountered before moving to the next
section. Answers to Study Break questions are provided in
Appendix A.
▲ Think Outside the Book activities help students ▲ End-of-Chapter review questions focus on both
think analytically and critically as they explore the biological factual and conceptual questions. Now organized according
world, either on their own or as part of a team. to Bloom’s Taxonomy.
xviii
MindTap Course Development: Simple & Powerful
Auto-graded learning activities are easily assignable by instructors and offer students immediate
feedback and coaching. Request a demo today: www.cengage.com/learningsolutions.
▲ Easily assign content and assignments with due dates. ▲ Comprehensive course analytics to assess student
learning and understanding in real time.
Effectively introducing digital solutions into your classroom — online or on-ground —

is now easier than ever. We’re with you every step of the way.
◀ Our Digital Course

“The technical help provided by Cengage staff in
Support Team
When you adopt from using MindTap was superb. The Digital Support
Cengage Learning, you Team was proactive in training and prompt in
have a dedicated team of answering follow-up questions for their excellent
Digital Course Support online learning resources.”
professionals, who will
provide hands-on, start- —K. Sata Sathasivan, The University of Texas at Austin
to-finish support, making
digital course delivery a
success for you and your
students.
xix
Acknowledgments
Revising a text—both the printed version and its wholly inte- Peter Russell thanks Joel Benington of St. Bonaventure
grated digital applications—from one edition to the next is a University for his extensive reviewing and contributions to
complex, though exciting and rewarding, project. The helpful Chapter 19, Genomes and Proteomes, and Chapter 25, The Ori-
assistance of many people enabled us to accomplish the task in gin of Life; and Sharon Thoma of the University of Wisconsin–
a timely manner. Madison for her expert input and advice during the revision of
Kristen Sutton, Life Sciences Product Manager, provided the Unit Six chapters on Animal Structure and Function. Paul
the essential vision, leadership, and support needed to bring Hertz thanks Hilary Callahan, John Glendinning, Jennifer
our unified course solution to fruition. Kristen and Nicole Mansfield, and Brian Morton of Barnard College for their gen-
Hurst, Product Development Specialist, worked tirelessly gain- erous advice on many phases of this project; Eric Dinerstein of
ing instructor input to ensure that the MindTap Learning Path RESOLVE for his contributions to the discussion of Conserva-
reflects the current state of biological knowledge conveyed with tion Biology; and Joel Benington for his expertise on genomic
cutting edge and effective pedagogical approaches. issues in systematics. Paul especially thanks Jamie Rauchman
Our Content Developers, Suzannah Alexander, Elesha Hyde, for extraordinary patience and endless support as this book
and Jake Warde, served as pilots for the generation of core con- was written (and rewritten, and rewritten again), as well as
tent. They provided very helpful guidance as the manuscript his thousands of past students at Barnard College, who have
matured. They compiled, interpreted, and sometimes decon- taught him how to be an effective teacher and mentor. Beverly
structed reviewer comments; their analyses and insights have McMillan once again thanks John A. Musick, Acuff Professor
helped us tighten the narrative and maintain a steady course. Emeritus at the College of William and Mary—and an award-
We are grateful to Kellie Petruzelli for coordinating winning teacher and mentor to at least two generations of col-
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We offer many thanks to John Anderson, Lauren Oliveira, biological science.
and Colleen O’Rourke, who spearheaded the MindTap course The authors dedicate this edition of the text to the memory
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teaching. seen the light of day.
We appreciate the help of the production staff led by Con- We would also like to thank our advisors and
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Pasternak at Cengage and Dan Fitzgerald and Megan Knight at
Graphic World. Supplements Authors
We also wish to acknowledge Tom Ziolkowski, our Mar- David Asch, Youngstown State University
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to Steve McEntee and to Dragonfly Media Group (DMG), led by Brent Ewers, University of Wyoming
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were never coming down. We’ve had breakfast ages ago; and Uncle Bexley
and the others, all except Mr. Karne, are already out on the moors.”
“How is it Mr. Karne has not gone?” Lady Marjorie asked wonderingly;
for Herbert was an enthusiastic sportsman.
“I don’t know. He is having a smoke in the lounge. P’raps he’s waiting to
give you your present. I mustn’t tell you what it is—it’s a surprise, you
know,—but I’m sure you will like it awf’lly. Uncle says it’s a very striking
likeness of me.”
“Tut-tut, Master Bobbie,” put in Janet, warningly. “You are letting the
cat out of the bag;” and the boy promptly clapped his hand to his lips.
Lady Marjorie found Karne deep in thought, watching with half-closed
eyes the smoke as it curled upwards from his cigar.
He rose at her approach, and having wished her many happy returns of
the day, presented her with a beautifully painted pastel of her boy.
Her face lit up with pleasure as she thanked him, for the gift had
evidently occasioned him much thought.
“I shall hang it up in my boudoir at Durlston,” she said, when she had
expressed her admiration of the portrait, “next to the one you painted of
Bobbie as a baby. Heigho, how time flies! I feel dreadfully old to-day—
because it is my birthday, I suppose.”
“One is never old whilst the heart is young,” he answered, with a swift
glance from his deep eyes. He was just thinking how delightfully fresh and
young she looked.
Lady Marjorie met his eyes and blushed. Then she sat down at a small
table and, unfolding a daily paper, glanced through the morning’s news.
“Are you tired of the shooting?” she inquired presently. “I was quite
surprised when Bobbie informed me that you were still indoors.”
“I am afraid there will be no more shooting for me this year,” he replied
regretfully, taking up a time-table which had recently occupied his attention.
“I have just packed my traps previous to taking my departure. This
morning’s post brought me two letters containing news which makes it
necessary for me to go to Brighton immediately. I am more sorry than I can
say to have to bring this enjoyable visit to such an abrupt termination.”
Lady Marjorie’s face fell perceptibly. “Then you are going away!” she
exclaimed in dismay. “You have not received bad news, I hope?”
“Well, that depends on how one looks at it,” he answered, noting her
crestfallen expression with a vague pang of self-reproach. “Celia’s visit to
Woodruffe has cost her dear; it has probably been the means of making her
lose her entire fortune.”
Lady Marjorie gave vent to an ejaculation of amazement.
“How could that possibly be?” she asked, her eyes distended in surprise.
The announcement almost took her breath away.
“She has decided to become a Christian,” he replied, as if apprising her
of some calamity. “And by doing so, according to the terms of her father’s
will, forfeits all claim to his wealth, which will go to build a Jewish hospital
in South Africa.”
Lady Marjorie stared at him blankly. “The little goose!” she exclaimed.
Then she corrected herself. “No, I didn’t mean that. Of course she must act
according to her belief. But I wonder what made her father insert such a
nonsensical stipulation in his will. I suppose she is aware of it?”
“No; judging by her letter, I do not think she is,” the artist answered,
with troubled brow. “I blame myself very much that I did not inform her of
it when I received the copy of the will, but I never dreamt of such a thing as
this happening. Her fiancé knows, however—Bernie Franks must have told
him himself,—and he is in a dreadful way about it. He is staying at Mrs.
Rosen’s house in Brighton, and begs me to join him there without delay.
Celia’s baptism is fixed for next Sunday; and, of course, if that is allowed to
take place, nothing can be done. Salmon writes that we must prevent that at
all costs, but I don’t see how we can if the girl has thoroughly made up her
mind to it.”
“No, I suppose not, as she is of age. But you may be able to persuade her
to postpone her baptism for a few months or so. It is possible that her
opinions may yet undergo another change. Does she seem very enthusiastic
over the matter?”
For answer Herbert handed her Celia’s letter to read. It consisted of eight
closely written pages; and judging by the frequent erasures, had evidently
been a difficult one to indite.
Lady Marjorie perused it carefully, reading several passages two or three
times in order to fully comprehend their meaning. At length she replaced it
in the envelope, and returned it without comment.
“Well?” interrogated Karne, briefly. “What do you think about it?”
“I hardly know. You see, I’m a Christian myself—though not a good
one, I’m afraid,—and I can understand how Celia feels about it. Religion is
a strange and fascinating subject; and it has evidently taken strong hold of
her. I do not think you will be able to deter her from carrying out her
intention. She seems to take it for granted that you will not blame her for
what she is doing. But I should not think she is aware of the loss of fortune
her conversion entails.”
“Oh, I do not blame her,” he said quickly. “If she imagines she can be
happier as a Christian, let her be one by all means. I do not suppose there
will be anything gained by attempting to argue the question with her. She
will probably prefer to be guided by the instinct she calls faith than to
consider any reasoning of mine.”
A clock in the adjoining hall struck eleven. Herbert glanced at his watch.
“I suppose you will go by the 12.50?” Lady Marjorie said, with a sigh.
“We must have an early luncheon; and then I will drive down to the station
to see you off. I shall miss you when you are gone,” she added regretfully.
“We’ve had a nice time up here together, haven’t we? Do you know, of all
Bexley’s guests, you are the only one whose society I have really enjoyed.
If it hadn’t been for you, I don’t think I should have stayed in Scotland all
this time. I am terribly outspoken, am I not? But one cannot always bottle
up one’s feelings.”
Again a touch of self-reproach smote Karne’s breast. He glanced into
Lady Marjorie’s eyes—such blue eyes, as clear and innocent as a child’s;
then feeling that he was expected to say something, expressed the pleasure
his visit had given him, and thanked her for her own and Bexley’s kindness.
He did not respond, however, in the way she had hoped he would; and
his words struck coldly upon her ears. Why did he always repel her
whenever she tried to make their friendship a little closer, she wondered,
with a vague feeling of disappointment at her heart.
It was the same at the railway station, where she lingered until the train
moved off. She gave him plenty of opportunity for pretty farewell speeches,
but he didn’t make them; and as she drove home again with Bobbie, tears of
mortification welled up into her eyes. It was quite ridiculous of her to care
so much, she told herself, as she choked them down.
Bobbie noticing her emotion, endeavoured to console her.
“Don’t cry, mother dear,” he said sympathetically. “We shall see Mr.
Karne again in Durlston next month. If you cry on your birthday, you’ll cry
all the year round, you know.”
Lady Marjorie thought she detected amusement in the expression of the
footman’s broad back.
“Nonsense!” she exclaimed, with a feeble smile. “Crying, indeed! It’s a
speck of dust in my eye.”
And another white lie was added to the list on her conscience.
CHAPTER VIII
THE RING RETURNED
“Well, what do you think of this d——d nonsense about Celia?” was
David Salmon’s polite greeting when he met Herbert Karne in the King’s
Road, Brighton, the next day.
He was so full of his grievance that he did not trouble to exchange the
customary civilities with the artist. Instead, he broke into a torrent of abuse
against the Wiltons, Lady Marjorie Stonor, and even Karne himself, for
having combined to lead his fiancée astray. He had been up to Woodruffe
that morning, he said, in order to give the Wiltons a piece of his mind, and
to implore Celia not to persist in her tomfoolery; but the girl was as
obstinate as a mule.
“Did you tell her what the consequence of her act will be, so far as
money is concerned?” asked Karne, who was not favourably impressed
with Salmon’s blustering manner.
“Yes, of course; but that didn’t seem to make the slightest difference.
She just went a bit white, and looked at me in a queer sort of way; then said
some stuff about ‘renunciation,’ and that was all. It’s my opinion that those
Wiltons must have worked upon her until her mind has become diseased;
and the sooner she gets away from them, the better. I have never heard of
such an idiotic affair in my life.”
Celia did not look, however, as if she possessed a morbid or diseased
mind. Her brother went over to Woodruffe in the afternoon, and found her
playing tennis. The exercise had lent a healthy glow to her cheeks; and she
looked much better and brighter than when he had last seen her in London.
The Wiltons received him kindly, although they were not sure whether
his visit were hostile as Mr. Salmon’s had been, or whether he was disposed
to be friendly; but their doubts were set at rest when he cordially invited
Enid to accompany Celia back to the Towers for the fortnight before her
rehearsals for the Haviland play began, and the invitation was accepted with
alacrity.
After tea they tactfully left the brother and sister alone, thinking, with
kindly consideration, that the two would have much to say to each other.
They were not mistaken. Herbert immediately began to ply Celia with a
volley of questions; and was some little time in eliciting all the information
he desired. Then he bade her consider well the gravity of her intended
action—an action that would cut her adrift from her own people, and make
her, for ever, an outcast in Israel.
“Do you know what your father would do, if he were alive?” he said
seriously. “He would sit shiva,[15] and mourn for you as one dead.”
But he did not blame her, nor did he cavil at her faith. He was kind, even
sympathetic; and all he asked her to do, for the present, was to wait awhile.
Celia, however, would not hear of procrastination in this matter; for the
Rev. Ralph Wilton was about to return to his parish, and she particularly
desired him to assist at the baptismal ceremony before he left. Besides,
there was nothing to be gained by waiting, she declared; her mind was fully
made up, her determination taken.
Herbert then advanced the monetary consideration, urging her not to
yield to a rash impulse she would probably live to regret; but, as he had
expected, this plea influenced her not at all.
“If the early Christians had allowed themselves to be guided by social
expediency, there would probably be little Christianity in the world to-day,”
she returned convincingly. “I must do what I feel to be my duty. But you
need not fear for me, Herbert. I am young and strong; and I have my voice.”
“And what of David Salmon? Have you considered him at all? You
know, it comes rather hardly upon him, after having been led to expect that
you would bring him a fortune.”
Celia’s eyes fell. “If he really loved me, he would be just as willing to
marry me poor as rich,” she rejoined.
“True; but I am afraid that he is not so unworldly as yourself. Tell me, sis
dear, would it hurt you very much if he were to give you up?”
Her heart beat fast; she had never thought of such a possibility.
“Do you think he would do that?” she asked, evading his question; and
her brother did not omit to notice the eager light in her eyes.
“Well, I had a lengthy conversation with him this morning,” he answered
slowly. “And it appears to me that this affair has brought out a new side to
his character; not a very commendable one, either, I am afraid. Of course
he, in common with the Friedbergs and Rosens, is shocked and disgusted;
not so much because of your change of faith—although the idea of his
marrying a converted Jewess is repugnant to them all—but because, by so
doing, you are deliberately throwing away a fortune. He informed me that,
on his marriage, Mr. Rosen intended taking him into partnership; but were
he to marry you without your money, the scheme would, of necessity, fall
through. Then he asked me what dowry I would give you, in the event of
your losing your inheritance. Now, you may be sure, dear sis, that I shall
always do my best to make ample provision for you; and you shall never
want, I trust, whilst I am alive; but I thought I would just meet Salmon on
his own ground. So I told him that I lived up to my income, pretty well—
which is quite true,—and that, having never foreseen this contingency, I
found myself utterly unable to provide you with a marriage portion. I don’t
think he quite believed that; anyway, he suggested my raising a mortgage
on the Towers, or something of that sort. Then, when he saw that I was
obdurate, he said that, much as he likes you, he could not afford to marry a
girl without money; so that, if you persist in what he calls your madness,
the engagement will have to be broken off. Finally, he asked me to persuade
you to reconsider your decision; and sincerely hoped that I would bring him
back good news.”
Celia was filled with indignation; but, because she had never really
loved him, the avariciousness of her fiancé occasioned her no grief. Rather,
she was relieved that his true nature was thus manifested before it was too
late.
“It is a wonder he did not suggest my singing or acting as a means of
support,” she said.
“He did; but I told him that I did not believe in a woman working to
keep her husband, unless he happened to be incapacitated by illness, or
there were some other urgent necessity. So it remains with you to decide
whether you will marry him or not. From what Marjie—Lady Marjorie, I
mean—has told me, I do not think your affections were deeply involved, so
that I can guess pretty well what your answer will be—eh, Celia?”
The girl slowly drew off her engagement ring. “Yes,” she replied
seriously, “I do not think I could marry him now, even were I to retain my
inheritance. My respect for him seems to have been suddenly obliterated.
Will you take him back this ring, please? And tell him that the man I marry
must love me for myself alone. Say, also, that, as I mean to carry out my
intention of joining the Christian Church, I am sure that there would often
be contention between us on that account; therefore the best thing—the only
thing—that I can do, is to dissolve our engagement.”
“And your decision is final?”
“Absolutely.”
Herbert made a wry face. “I cannot say I relish being the bearer of such a
message,” he said, placing the ring in his pocket-book. “Still, as you have
given it to me, I suppose I had better deliver it. I dare say Salmon will
round on me for having incensed you against him; and perhaps he will
prefer to receive your refusal from your own lips. I am afraid there will be a
mauvais quart d’heure for me when I get back to Brunswick Terrace.”
There was. David Salmon received the news with an oath, and broke into
a fit of passionate rage. After having cursed women in general, and Celia
Franks in particular, he declared that he would take to drink. When he had
calmed down, however, he thought better of it, and decided to console
himself with Dinah Friedberg. Dinah, so he said, besides being madly in
love with him, possessed no silly notions about religion, and her father,
although he did not make a pretence of being well off—as did Karne—
would at least endeavour to provide his daughter with a suitable marriage
dowry.
The next morning he presented himself at Woodruffe as though nothing
had happened. Celia would have preferred not to see him, but could not
very well refuse him the interview.
It was a painful one for both of them; and Celia, at least, felt relieved
when it was over. David implored, beseeched, and entreated her to
reconsider her decision, and refused at first to take back the few presents he
had given her, although he accepted them in the end. Finding that all his
pleading was of no avail, he revenged himself by indulging in cheap sneers
at her new-found faith, taunting her in the way best calculated to wound her
feelings. Finally, he encountered Ralph Wilton just as he was going out, and
told the clergyman what he thought of him in no measured terms.
Wilton himself was calm and unresentful, and his demeanour had the
effect of making Salmon a little bit ashamed of himself. He had the grace to
attempt an apology, at any rate, and even went so far as to shake hands
when he left.
Mr. Wilton accompanied him as far as the gate; then returned to the
drawing-room, to find Celia in tears.
The sight filled him with dismay. “Miss Franks!” he exclaimed, hardly
knowing how to express himself. “I—I am so sorry. I wish I could help you.
All this has been too much for you, I am afraid.”
Celia dried her eyes and smiled at him through her tears, reminding the
young clergyman of a burst of sunshine after a shower of rain.
“It—was—dreadfully weak of me!” she murmured in a small voice.
“But I couldn’t help it. Mr. Salmon did say such cruel things; and although I
know it’s foolish, they—they rankle. He made me feel as if I were about to
commit a crime.”
Ralph Wilton looked at her with deep sympathy in his eyes.
“The crown of thorns does indeed press hard upon your brow,” he said
compassionately. “You are being deprived of your fortune and your lover at
one blow. But do not lose heart, Miss Franks; I feel sure there is much
sunshine in store for you yet. Who can tell? Your self-sacrifice may lead to
happiness you know not of. Only trust and believe, and all will yet be well.”
“Oh, I am not at all unhappy,” she responded hastily, not wishing him to
be falsely impressed. “There is really no self-sacrifice in what I am doing.”
She did not add that the breaking of her engagement came as an unexpected
and not unwelcome release. Nevertheless, she felt it to be such, although it
was some little time before she could altogether realize that she was indeed
free.
The news of her conversion and its pecuniary consequence spread with
astonishing rapidity, even leaking into the Jewish and society papers.
Jewish people criticized her action as disgraceful, non-Jews as quixotic; and
both unanimously agreed that by foregoing a public confession of faith—
meaning the ceremony of baptism—she might have retained her fortune.
But public opinion caused Celia no concern, for she knew that no other
course than the one she had taken would have been possible to her for any
length of time. If she had acted foolishly according to the world’s standard,
she had at least done what she had felt to be her duty in the sight of God.
If she left Woodruffe the poorer in one way for her visit there, she was
richer in another; and never, during the whole course of her life, did she
ever wish her action undone.
CHAPTER IX
AN OUTCAST IN ISRAEL
“An outcast in Israel!” The words recurred to Celia with persistent

frequency during the next few weeks; for she went back to Durlston to find
herself ostracized by the little Jewish colony in whom she had taken interest
for so long a time.
Almost the first day after her return she went among them, as was her
custom when at home, taking with her toys for the children, articles of
adornment for the women, tobacco pouches for the men—all little
evidences of her thought for them whilst away. Never dreaming that her
conversion would make the slightest difference to them, the reception they
gave her stung her to the quick. The kindly greetings with which she was
wont to accost them died on her lips as she detected the look of scorn on
their faces. Mothers drew their little ones away from her, as though her very
touch meant contamination. Her gifts they regarded as so many briberies to
retain their good will, and therefore refused them with disdain.
Almost dumbfounded, and grieved to the heart, the girl sought refuge in
Mrs. Strelitzki’s cottage. Surely Anna would not turn against her, she
thought confidently, remembering the many kindnesses she had performed
for her in bygone days.
But even Anna Strelitzki, although she did not slam the door in her face,
as some of the others had done, received her without the slightest display of
cordiality. With embarrassment plainly discernible in her manner, she
offered her a seat by the fire, and then bolted the cottage door—a
proceeding which struck Celia as decidedly strange. Then, without
speaking, she went on with her washing, occasionally glancing furtively at
the window, apparently apprehensive of some unpleasant interruption.
“What is the meaning of all this, Anna?” Celia asked passionately. “Why
do they shun me as if I were some evil creature? I have done them no
harm!”
Mrs. Strelitzki trifled nervously with the corner of her apron, refusing to
meet the steady gaze from the girl’s clear eyes.
“M’shumadas!”[16] she exclaimed laconically, evidently deeming the
word sufficient explanation in itself, for she relapsed into silence, and went
on with her washing. Her manner was certainly strange.
Celia did not quite catch the meaning of the epithet; and, with tightly
clenched hands and compressed lips, waited for more. But no sound broke
the stillness save the ticking of the clock, and the measured breathing of a
sleeping child.
Suddenly the shrill toot of the factory horn, announcing the acquittal of
the workers, broke upon their ears. The child woke up with a fretful cry;
and the mother, drying her hands, came forward to quiet him.
“Oh, miss, I wish you would go home, if you don’t mind,” she said,
turning towards her visitor with an air of apology. “It’s getting near Jacob’s
dinner-time; and I dunno what ’ud happen if he were to coom back and find
you here. He’d half kill me, I think. He told me to have nowt to do with
you.”
“But why? I have done no harm,” the girl repeated, almost piteously. “Is
it because I have become a Christian?”
The woman nodded. “M’shumadas—traitress to the Faith,” she said in
the tone of one who repeats a watchword. “The people here are all good
Jews. They despise m’shumadim. They don’t want you to come and convert
their children, or give them tracts out of a black bag.”
“But I have no black bag,” Celia put in, with a faint smile, although there
were tears in her eyes. “And I have brought toys—not tracts. It is very
unkind of you all to treat me like this. I should not have thought it of you,
especially, Anna.”
“Good Jews despise m’shumadim,” the woman reiterated half sullenly,
and unbolted the door.
Celia drew on her gloves, and took her leave. With flaming cheeks and
quivering lips she hurried past the factory and down the high road. The men
were pouring out of the workshops, most of them wending their way
homewards. A few months ago they would have lifted their caps with a
courteous “Good morning, miss.” Now, they passed her with a scowl. Some
of the recently-arrived workers were informed as to her identity, and Celia
caught the word m’shumadas as it passed from lip to lip.
Arrived at the Towers, she burst into the library, where her brother and
Enid Wilton were writing, and impetuously told them of the insult she had
received. It was so uncalled-for, so nonsensical, so absolutely absurd, she
declared tremulously. She had done nothing to merit such treatment.
Enid Wilton listened with sympathy. Herbert Karne flung down his pen
with annoyance.
“So they mean war, do they?—the blockheads!” he exclaimed, with an
angry laugh. “I ought to have prepared you for this, Celia: you must not go
near them any more.”
“But why?” the girl asked quickly, as she threw her hat down on the
couch, and lifted Souvie up to be petted. “Do they not know that by
insulting me, they offend you also?”
The artist shrugged his shoulders. “They don’t much care if they do. For
some unaccountable reason I have lost my popularity amongst them. You
cannot imagine how terribly those people have disappointed me,” he added,
turning towards Enid Wilton with a touch of bitterness. “After having spent
much thought, time, and money on their education and the improvement of
their surroundings, I find them, in spite of it all, still dominated by the
instincts of the untutored savage: unprincipled, ungrateful, uncouth,
irresponsible, ignorant and superstitious in the extreme. The first few
batches of men I had down here responded admirably, and appreciated to
the full my efforts for them, but these present ones are absolutely
incorrigible. It is disheartening, is it not?—for I was confident of success in
my undertaking.”
“But what has happened to turn them against you?” asked his sister with
surprise. “Have you offended them also?”
“It seems like it. For the last six months there seems to have been an evil
influence among them; sometimes I think the poison of anarchy lurks in
their veins. They have taken a violent and senseless dislike to all the
influential men in the neighbourhood; they grudge them their wealth and
position, I suppose. Latterly, I have myself been included under the ban.”
“How strange!” exclaimed Celia, deeply interested, but vexed withal.
“A little while ago,” Karne continued, “I was commissioned to paint two
pictures for the Duke of Downshire’s private chapel, one on the subject of
the Annunciation, the other on the Crucifixion. I do not go in for religious
paintings as a rule, you know, but for several reasons I undertook these.
Well, these people from Mendel’s factory happened to see the pictures
through the studio window when they visited my grounds on the Sunday
after their completion, and took it into their stupid heads to imagine that
because I painted pictures on those subjects, I must of necessity be trending
towards Christianity myself. The news of Celia’s conversion coming on top
of that must have strengthened that idea, hence our unpopularity.”
“How narrow-minded they must be,” said Enid Wilton, thoughtfully.
“But surely it is against their own interest to offend you and your equals, is
it not?”
“Decidedly,” Herbert assented. “That is where their madness comes in;
they spite themselves, not us. However, I intend to close the night-school,
the dispensary, and the club for a few weeks. I must do something to bring
them to their senses.”
“What a pity!” Celia said, regretfully. “Enid and I were going to get up
such a nice concert for them next week; and Lady Marjorie had promised to
allow Bobbie to dance the hornpipe. The little fellow will be so
disappointed.”
She was disappointed herself—more keenly than she cared to confess—
and brooded on the inimical attitude of Mendel’s people, until the thought
of it quite distressed her. Had it not been for Enid Wilton’s companionship
she would have felt inclined to give way to depression; but Enid was bright
and entertaining, and did her best to divert her friend’s mind into other
channels.
The two girls avoided the vicinity of the factory as much as possible, but
were obliged to pass it on their way to Durlston House, whither Lady
Marjorie had recently returned. Occasionally they met some of the
workpeople or their relatives; but Celia always passed them without a sign
of recognition, for she knew that to speak to them would be to invite an
insult.
One day they came across a small Jewish maiden who was sitting by the
road-side alone in a sorry plight. She was some distance from the factory,
and had evidently been sent on an errand, for clutched in her grasp was a
basket of provisions. A bottle of olive oil, too unwieldy for her to manage,
had accidentally fallen out. She was surrounded by broken pieces of glass,
and her thinly-clad feet had been painfully cut and scratched. Judging by
her appearance, one might have credited her with having taken an oil bath,
for, from her curly black ringlets down to her toes, she was literally covered
with the greasy fluid.
The girls’ kind hearts were touched by the sight. Celia, forgetting all
strife in her compassion for the little one, bent down and inquired her name.
After some amount of coaxing, she discovered that it was “Blume
Horwitz;” that her feet hurt her so much that she could not walk; that her
mother was waiting for the oil to fry the fish, and that she would be
welcomed with a beating when she did arrive home. Her tale of woe ended
in a fit of sobbing and gulping pitiful to behold.
The girls consulted as to what they should do. They could not leave her
there, on the chance of one of her people picking her up, nor could they
carry her home, saturated with oil as she was.
At length Celia decided to go home as quickly as she could for the pony-
chaise, leaving her friend to stay with the child. This she accordingly did,
and in less than twenty minutes was back again with the conveyance.
The coachman gingerly covered the little girl with an overall belonging
to the stable-boy, and lifted her into the chaise. Celia had brought some lint
back with her, and between them the two girls skilfully bound up her
wounds, which were not so severe as they had at first supposed. When they
arrived at the Towers, a messenger was immediately despatched to inform
Mrs. Horwitz of the accident, and to procure a change of clothing for
Blume. Meanwhile the child’s wants were attended to in Celia’s pretty
bedroom.
An hour later, the coachman, Roberts, drove her home; clean,
comfortable, and well-fed. He found the cottage shut up, for Mrs. Horwitz
was always out at that time of the day; but a man was waiting at the wicket
in anticipation of Blume’s arrival. Possessing small cunning eyes with an
unpleasant leer in them, an aquiline nose, heavy jaw, and cruel mouth, his
countenance was decidedly unattractive; and his burly form suggested an
ample reserve of brute force. He was Anna’s husband, Jacob Strelitzki, who
had recently returned after a year’s absence from the factory, spent no one
knew where. Roberts pulled up at the wicket, and alighting from the chaise,
eyed the man with disfavour.
“Hello!” he said bluntly. “Strelitzki, is it? Thought I’d seen that ugly
face before. So you’ve come back, have you? Been in quod, I suppose?
Lost your curly wig, anyhow. Where is this kid’s mother?”
If looks could kill, the coachman would have been exterminated on the
spot. Scowling savagely, Strelitzki bade him hold his tongue, for the child
had fallen asleep, and he did not wish her to be awakened. With more
gentleness than was his custom, he lifted her out of the chaise, and,
unlocking the cottage door, laid her carefully down on the couch.
Then he returned to the wicket, and informed the coachman that he
might consider himself dismissed. Roberts, however, was apparently not
quite satisfied.
“ ’Ere, where’s the kid’s mother?” he asked again. “My mistress said I
was to see that the little girl was all right. She has cut her foot, and has got
to lay up. You ain’t any relation, are you?”
“Yes; I am her uncle,” the man replied briefly. “Rachael Horwitz has told
me all about the accident. She ought not to have sent such a little thing so
far on an errand. She’s got slipper-work at the factory, so you’ll have to
leave the child with me.” And without further remark, Roberts drove away.
Strelitzki bolted the door after him, and quietly moved to where the child
lay. She was still fast asleep, but stirred uneasily as he watched her. Fearing
that the light might awaken her, Jacob carefully shut the lattice. His
movement suggested mystery; but all his caution was for the purpose of
performing an apparently trivial action.
Taking a small packet out of his coat-pocket, he cut the string and
unfolded the tissue-paper. Inside lay a tiny crucifix composed of black
wood and nickel silver—truly a strange emblem to be in his possession.
From another pocket he produced a piece of slightly-faded blue ribbon.
Then twisting the ribbon through the ring at the top of the crucifix, he tied it
securely round Blume’s neck, tucking it under her pinafore. This
accomplished, he gave a sigh—which was almost a chuckle—of relief.
The action disturbed the child, who awoke with a feeble cry of pain. For
the moment she could not quite take in her surroundings, and blinked at the
daylight in bewilderment. When she recollected what had happened, she
began to cry, fearing her mother’s anger on account of the broken bottle of
oil; but her uncle assured her that the accident had been explained, and that
her mother would be back directly, grieved to find her in pain.
Strelitzki lit his pipe and professed to read the newspaper; at the same
time watching the little girl out of the corner of his eye. Her feet still
smarted painfully, and she moved her position frequently in order to obtain
greater ease. In doing so, the crucifix slipped out, and hung suspended from
her neck above her pinafore.
“Hello!” exclaimed Strelitzki. “Where did you get that?”
Blume examined it with wide-open eyes. She had not the faintest idea of
the meaning of the symbol, or, indeed, that it was a symbol at all; but the
blue ribbon and silver figure pleased her, and in her childish mind she
considered it a fine ornament, to be put on a par with her mother’s lozenge-
shaped earrings, and only to be worn on Shabbos[17] and Yomtov.[18]
“I don’t know,” she replied truthfully, wishing it had escaped her uncle’s
observation.
“Nonsense!” said Jacob Strelitzki. “Of course, you must know. I expect
Miss Celia—the lady with the carroty-golden hair—gave it to you when she
changed your things, didn’t she?”
That seemed very likely, so Blume agreed to it. She did not remember
Miss Celia giving it to her, it is true; but she had given her a box of
chocolates and a “Cinderella” picture-book, so no doubt the ornament came
from her as well.
“Yes,” she assented, readily. “Miss Celia gave it to me.”
Strelitzki grunted satisfaction. “Well, tuck it under your frock,” he
advised. “Or some one may want to take it off you. If your mammy should
find it when she puts you to bed, say that Miss Celia said you were to keep
it and not give it away.”
The child acquiesced; and Strelitzki went on reading his paper. He
seemed to find it difficult to concentrate his thoughts, however, for he soon
tossed it aside, and stared into the fire with his shaggy brows contracted,
and an evil smile on his heavy face.
“So—so, Herbert Karne,” he muttered softly in his native jargon. “You
and I hate each other; and we have a long-standing account to settle.
Revenge grows keener with delay. It shall be settled soon!”
CHAPTER X
STRELITZKI PAVES THE WAY FOR HIS REVENGE
The yard at Mendel’s factory was filled to its utmost capacity. Men jostled
each other’s elbows, and trod on each other’s corns with good-natured
indiscrimination. A jargon of Polish, Yiddish, Roumanian, and English of
the Lancashire dialect smote the air with Babel-like confusion; and as each
man spoke to his neighbour at the precise moment that his neighbour spoke
to him, the amount of comprehension on either side was reduced to nil.
They had met for the discussion of a grievance. Herbert Karne, after
further provocation, had put his threat into execution: the night-school, the
dispensary, and the club were closed. A notice was pasted on the doors
stating that they would remain closed until he received, signed by each one
of the men, a full and satisfactory apology for the gratuitous insults levelled
at his sister and himself; together with a promise of better behaviour in
future.
The news produced a sensation, some of the men utterly refusing to
believe it until they saw the notice for themselves. The club had been
opened so long, and occupied such a prominent position in the recreative
part of their work-a-day lives, that they had lost sight of the fact that it was
kept up entirely at Herbert Karne’s expense. Nearly every evening they
repaired thither to while away an hour or two in the comfortable reading or
smoke rooms; which were always well heated in winter, well ventilated in
summer. Here they could chat, or schmooze,[19] as they called it, to their
heart’s content. They were also at liberty to play solo-whist, so long as they
played for nominal stakes only, gambling being strictly prohibited; and in
the winter evenings, Herbert Karne arranged numerous entertainments for
their benefit, to which their women folks, in their Sabbath clothes, came as
well.
The club closed, they would be obliged to have recourse to the bar-
parlours of the public-houses; for the gregarious instinct was strong within
them, and their home-life more or less unattractive. But they knew that,
being foreigners and abstemious, they would not receive a cordial welcome
there; nor, indeed, did they desire the society of public-house frequenters.
They had the greatest respect for the British workman when sober; but they
were aware that having waxed convivial by the aid of beer, he was apt to
indulge in uncomplimentary remarks concerning “them furriners;” and
being extremely sensitive, they did not care for jocularity at their own
expense.
It became evident, therefore, that they must endeavour to get the club re-
opened; and it was in order to effect this end, that the meeting was being
held.
In the centre of the yard a number of heavy boxes had been piled up to
serve as a rostrum; and from this a slender olive-skinned man addressed his
fellow-workers. He was Emil Blatz, the foreman of the factory and manager
of the club.
Their present attitude to their benefactor, he told them—when he could
command silence—was senseless to the last degree. They had been
indulging in foolish spleen, and incurring serious harm to themselves, as the
closing of the club and dispensary testified. They were simply running their
heads against a brick wall when they imagined they could go against a man
in Mr. Karne’s position. He advised them to sign an apology which he
himself would prepare; and voted that they should do all in their power to
renew their former friendly relations with Herbert Karne.
His address was received with expressions of mingled approval and
dissent. The majority of them were half inclined to think that it would be
wiser in the end to cease hostility, especially as the winter was approaching.
They remembered the numerous creature comforts which had been
provided every year at the artist’s expense.
Jacob Strelitzki, with a wild light in his eyes, elbowed his way through
the crowd and sprang on to the platform.
“Mates!” he shouted energetically, “do you want to be turned into bacon-
eating m’shumadim by Herbert Karne and his sister?”
A vigorous reply in the negative rolled towards him like the answer of
one man.
“Well, then, don’t apologize, don’t play into their hands! Herbert Karne
is no true friend of ours! He has taken an interest in our welfare simply that
he might convert us all in the end! Four years ago he did his best to make a
m’shumad of me, but I resisted before it was too late. We have our wives
and children to consider—suppose he converts them against our will? Let
us make a firm stand against it, and swear that that shall never be!”
Murmurs of indignation and applause came from every throat; but the
foreman Blatz held up his hand to still them.
“It is false!” he cried in a voice that could be heard at the furthermost
corner of the yard. “Mr. Karne is our true friend, and he is not a m’shumad.
He has told us over and over again that he wishes us to be good Jews and
upright men; he has never attempted to teach us any creed but our own.
What right, then, have we to say that he is not a good Jew?”
“Every right!” replied the dark-bearded man vehemently. “If Herbert
Karne were a good Jew, he would not have received his sister into his house
after she became a Christian. He should have treated her as Bernie Franks
would have done had he lived; he ought to have cast her adrift. Listen here,
friends, Strelitzki is right. If we allow ourselves to be ruled by the people at
the Towers, we shall find our wives and children being led astray. Only
yesterday my little girl Blume met with a slight accident whilst out on an
errand. Miss Celia Franks used it as an excuse to entice her to the Towers,
where she kept her for some time. What she said to the child I do not know,
but when my wife undressed Blume at night, she discovered this”—lifting a
crucifix high above their heads—“hung round her neck. Comrades, are we
to stand by without protest in the face of an insult such as this?”
“No, no!” responded the angry crowd, their ire aroused at the sight of the
offending emblem. “Stamp on it! Crush the trumpery thing! Down with
those who dare to tamper with our religion! Down with m’shumadim!”
A crucifix around a Jewish child’s neck! It was the worst indignity that
could have been offered to them, for nothing could have shocked them
more. Here was proof positive of Celia Franks’ intention to convert their
children by force; here was virtually their call to arms.
Even the foreman Blatz knew not what to think; like the rest of them he
was amazed and shocked. In vain now did he urge them to establish peace;
the incident of the crucifix decided what their course of action should be.
They accused Blatz himself of apostasy when he again pleaded in favour
of the artist. They would do without Mr. Karne’s gifts rather than be robbed
of the faith of their forefathers. They would ask one of the Rothschilds or
Montefiores to build them a club; they would accept nothing more from
Herbert Karne.
The meeting broke up in noisy confusion, a motion being carried to
arrange further proceedings the following night. The men dispersed in twos
and threes, each discoursing volubly with his neighbour in whatever his
native language happened to be.
Emil Blatz went on his way alone, with heavy heart and thoughtful brow.
Usually he himself, as foreman, took the lead in factory affairs, but to-night
he had been superseded. The men had been swayed by Strelitzki and
Horwitz, who by common consent had established themselves as leaders,
and their temper boded no good towards Herbert Karne.
Blatz possessed a strong admiration for the artist, who had done him
many a good turn. He could not forget a certain eventful night, when his
boy lay dying, and Karne had kept vigil with him for eight weary hours,
until, at dawn, the little soul had fled into the dim unknown. He felt he
owed him a debt of gratitude for that, which, if it were in his power, he must
repay.
Almost involuntarily his steps turned towards the Towers, although he
had only a vague idea as to what he intended to do. Without giving himself
time for thought he pressed the visitors’ bell. Noiselessly the gate swung
back, gaining him admittance to the grounds. The coachman’s wife peered
out at him as he drew near the lodge, but offered no resistance: and with
careful steps he passed along the gravelled path which bounded the lawn,
until the house with its ornamental turrets loomed clear against the
blackness of the night.
Presently the sound of music made him pause; the mellow tones of a
piano, and then a woman’s voice, full, rich, and clear. Blatz listened with
eager attention, for he was a musician born. Softly and sweetly the notes
floated towards him through the half-open windows. He recognized the
melody; it was an aria from Elijah.
Moving a few steps to the right he found himself in full view of the
drawing-room. The blinds had not been lowered; and through the
transparent curtains he could see the interior of the room.
The scene struck him strangely, being in such marked contrast to the one
he had just left. It was as if, in the midst of turbulent strife, he had suddenly
come upon a haven of rest. Here for one short moment he might breathe the
atmosphere of peace and refinement. Although but a humble factory
worker, Blatz possessed a passionate love of the beautiful; and this
luxurious apartment, with its dainty touches of femininity, awakened a keen
thrill of pleasure within his breast.
There were five occupants of the room, all of whom were known to the
foreman, except the dark-haired girl at the piano. Herbert Karne stood with
his back to the fireplace exhibiting a book of sketches to the white-haired
vicar of Durlston. Seated on a low chair in the roseate glow of the lamp was
the vicar’s daughter, her fingers busily plying a piece of fancy-work; and
facing her, by the side of the grand piano, stood Celia Franks, singing with
all her heart.
“Hear ye, Israel! hear what the Lord speaketh: Oh, hadst thou heeded,
heeded My commandments!”
Sweetly and half reproachfully she sang the words to their melodious
accompaniment. Her eyes were dimly fixed on the dark swaying trees in the
garden; her thoughts were far from the lighted room.
Then more solemnly she enunciated the question: “Who hath believed
our report? To whom is the arm of the Lord revealed?” Afterwards
recurring to the exhortation, “Hear ye!” and closing with the pathetic appeal
in the minor key, “Israel!... Israel!”
A wave of emotion swept over Emil Blatz as he listened; the mellifluous
beauty of the melody almost carried him away. He knew not whom he
envied the more: Mendelssohn for having composed such music, or the
young singer for her power to interpret it in that way.
The words, too, sounded in his ear with peculiar significance; they
seemed like a justification of the singer’s faith.
Suddenly the voice ceased its tender note of appeal; and after a few bars
of recitative, burst forth into a triumphant assurance of divine protection,
followed by the sublime meditation:—
“Say, who art thou, that art afraid of a man that shall die? And forgettest
the Lord thy Maker, Who hath stretched forth the heavens, and laid the
earth’s foundations? Be not afraid, for I, thy God, will strengthen thee!”
To Blatz there was a note of defiance in the girl’s rendering of the
dramatic music: the very poise of her head, as she sang the “Be not afraid,”
seemed like a challenge to those who were her enemies. In his simplicity he
forgot that she was quite unconscious of her uninvited listener, and that the
words were not her own.

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THE RING RETURNED

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STRELITZKI PAVES THE WAY FOR HIS REVENGE

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