89892_FM_i-xxxviii.indd 1 10/21/15 11:23 AM (cid:13)(cid:96)(cid:136)(cid:204)(cid:105)(cid:96)(cid:202)(cid:220)(cid:136)(cid:204)(cid:133)(cid:202)(cid:204)(cid:133)(cid:105)(cid:202)(cid:96)(cid:105)(cid:147)(cid:156)(cid:202)(cid:219)(cid:105)(cid:192)(cid:195)(cid:136)(cid:156)(cid:152)(cid:202)(cid:156)(cid:118)(cid:202) (cid:22)(cid:152)(cid:118)(cid:136)(cid:221)(cid:202)(cid:42)(cid:192)(cid:156)(cid:202)(cid:42)(cid:12)(cid:19)(cid:202)(cid:13)(cid:96)(cid:136)(cid:204)(cid:156)(cid:192)(cid:202) (cid:47)(cid:156)(cid:202)(cid:192)(cid:105)(cid:147)(cid:156)(cid:219)(cid:105)(cid:202)(cid:204)(cid:133)(cid:136)(cid:195)(cid:202)(cid:152)(cid:156)(cid:204)(cid:136)(cid:86)(cid:105)(cid:93)(cid:202)(cid:219)(cid:136)(cid:195)(cid:136)(cid:204)(cid:92)(cid:202) (cid:220)(cid:220)(cid:220)(cid:176)(cid:136)(cid:86)(cid:105)(cid:152)(cid:136)(cid:176)(cid:86)(cid:156)(cid:147)(cid:201)(cid:213)(cid:152)(cid:143)(cid:156)(cid:86)(cid:142)(cid:176)(cid:133)(cid:204)(cid:147) 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. Copyright 2017 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. 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Obviously, right? But the right way to go about it isn’t always so obvious. Go digital to get the grades. MindTap’s customizable study tools and eTextbook give you everything you need all in one place. Engage with your course content, enjoy the flexibility of studying anytime and anywhere, stay connected to assignment due dates and instructor notifications with the MindTap Mobile app... and most of all…EARN BETTER GRADES. TO GET STARTED VISIT WWW.CENGAGE.COM/STUDENTS/MINDTAP Copyright 2017 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s). Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it. 89892_ES3_SE.indd 1 10/22/15 3:45 PM (cid:13)(cid:96)(cid:136)(cid:204)(cid:105)(cid:96)(cid:202)(cid:220)(cid:136)(cid:204)(cid:133)(cid:202)(cid:204)(cid:133)(cid:105)(cid:202)(cid:96)(cid:105)(cid:147)(cid:156)(cid:202)(cid:219)(cid:105)(cid:192)(cid:195)(cid:136)(cid:156)(cid:152)(cid:202)(cid:156)(cid:118)(cid:202) (cid:22)(cid:152)(cid:118)(cid:136)(cid:221)(cid:202)(cid:42)(cid:192)(cid:156)(cid:202)(cid:42)(cid:12)(cid:19)(cid:202)(cid:13)(cid:96)(cid:136)(cid:204)(cid:156)(cid:192)(cid:202) (cid:47)(cid:156)(cid:202)(cid:192)(cid:105)(cid:147)(cid:156)(cid:219)(cid:105)(cid:202)(cid:204)(cid:133)(cid:136)(cid:195)(cid:202)(cid:152)(cid:156)(cid:204)(cid:136)(cid:86)(cid:105)(cid:93)(cid:202)(cid:219)(cid:136)(cid:195)(cid:136)(cid:204)(cid:92)(cid:202) (cid:220)(cid:220)(cid:220)(cid:176)(cid:136)(cid:86)(cid:105)(cid:152)(cid:136)(cid:176)(cid:86)(cid:156)(cid:147)(cid:201)(cid:213)(cid:152)(cid:143)(cid:156)(cid:86)(cid:142)(cid:176)(cid:133)(cid:204)(cid:147) Biology The Dynamic Science Fourth Edition Russell Hertz McMillan Australia • Brazil • Mexico • Singapore • United Kingdom • United States Copyright 2017 Cengage Learning. 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Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it. 89892_FM_i-xxxviii.indd 1 10/21/15 11:23 AM (cid:13)(cid:96)(cid:136)(cid:204)(cid:105)(cid:96)(cid:202)(cid:220)(cid:136)(cid:204)(cid:133)(cid:202)(cid:204)(cid:133)(cid:105)(cid:202)(cid:96)(cid:105)(cid:147)(cid:156)(cid:202)(cid:219)(cid:105)(cid:192)(cid:195)(cid:136)(cid:156)(cid:152)(cid:202)(cid:156)(cid:118)(cid:202) (cid:22)(cid:152)(cid:118)(cid:136)(cid:221)(cid:202)(cid:42)(cid:192)(cid:156)(cid:202)(cid:42)(cid:12)(cid:19)(cid:202)(cid:13)(cid:96)(cid:136)(cid:204)(cid:156)(cid:192)(cid:202) (cid:47)(cid:156)(cid:202)(cid:192)(cid:105)(cid:147)(cid:156)(cid:219)(cid:105)(cid:202)(cid:204)(cid:133)(cid:136)(cid:195)(cid:202)(cid:152)(cid:156)(cid:204)(cid:136)(cid:86)(cid:105)(cid:93)(cid:202)(cid:219)(cid:136)(cid:195)(cid:136)(cid:204)(cid:92)(cid:202) (cid:220)(cid:220)(cid:220)(cid:176)(cid:136)(cid:86)(cid:105)(cid:152)(cid:136)(cid:176)(cid:86)(cid:156)(cid:147)(cid:201)(cid:213)(cid:152)(cid:143)(cid:156)(cid:86)(cid:142)(cid:176)(cid:133)(cid:204)(cid:147) Biology: The Dynamic Science, Fourth Edition © 2017, 2014 Cengage Learning Peter J. 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All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s). Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it. 89892_FM_i-xxxviii.indd 3 10/21/15 11:23 AM (cid:13)(cid:96)(cid:136)(cid:204)(cid:105)(cid:96)(cid:202)(cid:220)(cid:136)(cid:204)(cid:133)(cid:202)(cid:204)(cid:133)(cid:105)(cid:202)(cid:96)(cid:105)(cid:147)(cid:156)(cid:202)(cid:219)(cid:105)(cid:192)(cid:195)(cid:136)(cid:156)(cid:152)(cid:202)(cid:156)(cid:118)(cid:202) (cid:22)(cid:152)(cid:118)(cid:136)(cid:221)(cid:202)(cid:42)(cid:192)(cid:156)(cid:202)(cid:42)(cid:12)(cid:19)(cid:202)(cid:13)(cid:96)(cid:136)(cid:204)(cid:156)(cid:192)(cid:202) (cid:47)(cid:156)(cid:202)(cid:192)(cid:105)(cid:147)(cid:156)(cid:219)(cid:105)(cid:202)(cid:204)(cid:133)(cid:136)(cid:195)(cid:202)(cid:152)(cid:156)(cid:204)(cid:136)(cid:86)(cid:105)(cid:93)(cid:202)(cid:219)(cid:136)(cid:195)(cid:136)(cid:204)(cid:92)(cid:202) (cid:220)(cid:220)(cid:220)(cid:176)(cid:136)(cid:86)(cid:105)(cid:152)(cid:136)(cid:176)(cid:86)(cid:156)(cid:147)(cid:201)(cid:213)(cid:152)(cid:143)(cid:156)(cid:86)(cid:142)(cid:176)(cid:133)(cid:204)(cid:147) 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 monuen sietym ceosltleerg oe fs tthued einnttrso tdhuatc tfoarcyil isteaqteuse tnhceei rf otrra Bnisofelor gtyo mfoaujro-ryse aanr dco pllreeg-epsr oafnedss uionnivael rsstiutdieesn. tHs,e l etecatuchree s 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. Millan Mc Courtesy of Beverly iv Chapter 8 harvesting ChemiCal energy: Cellular respiration Copyright 2017 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s). Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it. 89892_FM_i-xxxviii.indd 4 10/21/15 11:23 AM (cid:13)(cid:96)(cid:136)(cid:204)(cid:105)(cid:96)(cid:202)(cid:220)(cid:136)(cid:204)(cid:133)(cid:202)(cid:204)(cid:133)(cid:105)(cid:202)(cid:96)(cid:105)(cid:147)(cid:156)(cid:202)(cid:219)(cid:105)(cid:192)(cid:195)(cid:136)(cid:156)(cid:152)(cid:202)(cid:156)(cid:118)(cid:202) (cid:22)(cid:152)(cid:118)(cid:136)(cid:221)(cid:202)(cid:42)(cid:192)(cid:156)(cid:202)(cid:42)(cid:12)(cid:19)(cid:202)(cid:13)(cid:96)(cid:136)(cid:204)(cid:156)(cid:192)(cid:202) (cid:47)(cid:156)(cid:202)(cid:192)(cid:105)(cid:147)(cid:156)(cid:219)(cid:105)(cid:202)(cid:204)(cid:133)(cid:136)(cid:195)(cid:202)(cid:152)(cid:156)(cid:204)(cid:136)(cid:86)(cid:105)(cid:93)(cid:202)(cid:219)(cid:136)(cid:195)(cid:136)(cid:204)(cid:92)(cid:202) (cid:220)(cid:220)(cid:220)(cid:176)(cid:136)(cid:86)(cid:105)(cid:152)(cid:136)(cid:176)(cid:86)(cid:156)(cid:147)(cid:201)(cid:213)(cid:152)(cid:143)(cid:156)(cid:86)(cid:142)(cid:176)(cid:133)(cid:204)(cid:147) 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 the living world as scientists . . . learning. We have also received comments from expert review- 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 harvesting ChemiCal energy: Cellular respiration v Copyright 2017 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s). Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. Cengage Learning reserves the right to remove additional content at any time if subsequent rights restrictions require it. 89892_FM_i-xxxviii.indd 5 10/21/15 11:23 AM (cid:13)(cid:96)(cid:136)(cid:204)(cid:105)(cid:96)(cid:202)(cid:220)(cid:136)(cid:204)(cid:133)(cid:202)(cid:204)(cid:133)(cid:105)(cid:202)(cid:96)(cid:105)(cid:147)(cid:156)(cid:202)(cid:219)(cid:105)(cid:192)(cid:195)(cid:136)(cid:156)(cid:152)(cid:202)(cid:156)(cid:118)(cid:202) (cid:22)(cid:152)(cid:118)(cid:136)(cid:221)(cid:202)(cid:42)(cid:192)(cid:156)(cid:202)(cid:42)(cid:12)(cid:19)(cid:202)(cid:13)(cid:96)(cid:136)(cid:204)(cid:156)(cid:192)(cid:202) (cid:47)(cid:156)(cid:202)(cid:192)(cid:105)(cid:147)(cid:156)(cid:219)(cid:105)(cid:202)(cid:204)(cid:133)(cid:136)(cid:195)(cid:202)(cid:152)(cid:156)(cid:204)(cid:136)(cid:86)(cid:105)(cid:93)(cid:202)(cid:219)(cid:136)(cid:195)(cid:136)(cid:204)(cid:92)(cid:202) (cid:220)(cid:220)(cid:220)(cid:176)(cid:136)(cid:86)(cid:105)(cid:152)(cid:136)(cid:176)(cid:86)(cid:156)(cid:147)(cid:201)(cid:213)(cid:152)(cid:143)(cid:156)(cid:86)(cid:142)(cid:176)(cid:133)(cid:204)(cid:147) 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- into the narrative . . . tions and whet their appetites for the topic that the chapter addresses. In many cases, this feature tells the story of how 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 prefaCe prefaCe Copyright 2017 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s). Editorial review has deemed that any suppressed content does not materially affect the overall learning experience. 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