Principal Campbell Biology: Concepts & Connections
Campbell Biology: Concepts & ConnectionsMartha R. Taylor, Eric J. Simon, Jean L. Dickey, Kelly A. Hogan, Jane B. Reece
Campbell Biology: Concepts & Connections
continues to introduce pedagogical innovations, which motivate students not only to learn, but also engage with biology. This bestselling textbook is designed to help students stay focused with its hallmark modular organization around central concepts and engages students in connections between concepts and the world outside of the classroom with Scientific Thinking, Evolution Connection and Connection essays in every chapter.
The 9th Edition offers students a framework organized around fundamental biological themes and encourages them to analyze visual representations of data with new Visualizing the Data figures. A reorganized Chapter One emphasizes the process of science and scientific reasoning, and robust instructor resources and multimedia allow students to engage with biological concepts in a memorable way. Unparalleled resources let instructors develop active and high interest lectures with ease.
continues to introduce pedagogical innovations, which motivate students not only to learn, but also engage with biology. This bestselling textbook is designed to help students stay focused with its hallmark modular organization around central concepts and engages students in connections between concepts and the world outside of the classroom with Scientific Thinking, Evolution Connection and Connection essays in every chapter.
The 9th Edition offers students a framework organized around fundamental biological themes and encourages them to analyze visual representations of data with new Visualizing the Data figures. A reorganized Chapter One emphasizes the process of science and scientific reasoning, and robust instructor resources and multimedia allow students to engage with biological concepts in a memorable way. Unparalleled resources let instructors develop active and high interest lectures with ease.
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Brief Contents 1 Biology: Exploring Life 1 Unit I The Life of the Cell 2 The Chemical Basis of Life 20 3 The Molecules of Cells 36 4 A Tour of the Cell 54 5 The Working Cell 76 6 How Cells Harvest Chemical Energy 92 7 Photosynthesis: Using Light to Make Food 110 U n i t II Cellular Reproduction and Genetics Unit V Animals: Form and Function 20 Unifying Concepts of Animal Structure and Function 416 21 Nutrition and Digestion 432 22 Gas Exchange 456 23 Circulation 470 24 The Immune System 488 25 Control of Body Temperature and Water Balance 508 26 Hormones and the Endocrine System 520 27 Reproduction and Embryonic Development 536 28 Nervous Systems 566 8 The Cellular Basis of Reproduction and Inheritance 128 29 The Senses 590 9 Patterns of Inheritance 156 30 How Animals Move 606 10 Molecular Biology of the Gene 184 11 How Genes Are Controlled 212 12 DNA Technology and Genomics 234 U n i t III Concepts of Evolution 13 How Populations Evolve 258 14 The Origin of Species 280 15 Tracing Evolutionary History 296 Unit IV Unit VI Plants: Form and Function 31 Plant Structure, Growth, and Reproduction 624 32 Plant Nutrition and Transport 646 33 Control Systems in Plants 664 U n i t V II Ecology The Evolution of Biological Diversity 34 The Biosphere: An Introduction to Earth’s Diverse Environments 682 16 Microbial Life: Prokaryotes and Protists 322 35 Behavioral Adaptations to the Environment 702 17 The Evolution of Plant and Fungal Diversity 344 36 Population Ecology 726 18 The Evolution of Invertebrate Diversity 368 37 Communities and Ecosystems 742 19 The Evolution of Vertebrate Diversity 392 38 Conservation Biology 764 CAMPBELL BIOLOGY C ONC E P T S & C ON N E C T ION S | MARTHA R. TAYLOR NINTH EDITION JEAN L. DICKEY Ithaca, New York Clemson University ERIC J. SIMON KELLY HOGAN New England College University of North Carolina, Chapel Hill JANE B. REECE Berkeley, California 330 Hudson Street, New York, NY 10013 Courseware Portfolio Management Director: Beth Wilbur Courseware Director of Content Development: Ginnie Simione Jutson Courseware Portfolio Management Specialist: Alison Rodal Development Editors: Julia Osborne, Susan Teahan, Mary Catherine Hager, Debbie Hardin, Evelyn Dahlgren Editorial Coordinator: Alison Cagle Director of Content Production and Digital Studio: Erin Gregg Managing Producer, Science: Michael Early Content Producers: Mae Lum, Courtney Towson Production Management and Composition: Alverne Ball, Integra Software Services, Inc. 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This publication is protected by copyright, and permission should be obtained from the publisher prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise. For information regarding permissions, request forms and the appropriate contacts within the Pearson Education Global Rights & Permissions department, please visit www.pearsoned.com/ permissions/. Acknowledgements of third party content appear on page A-26, which constitutes an extension of this copyright page or on the appropriate page within the text. PEARSON, ALWAYS LEARNING, MasteringBiology and BioFlix are exclusive trademarks in the U.S. and/or other countries owned by Pearson Education, Inc. or its affiliates. Unless otherwise indicated herein, any third-party trademarks that may appear in this work are the property of their respective owners and any references to third-party trademarks, logos or other trade dress are for demonstrative or descriptive purposes only. Such references are not intended to imply any sponsorship, endorsement, authorization, or promotion of Pearson’s products by the owners of such marks, or any relationship between the owner and Pearson Education, Inc. or its affiliates, authors, licensees or distributors. Library of Congress Cataloging-in-Publication Data Names: Reece, Jane B. | Taylor, Martha R. | Simon, Eric J. (Eric Jeffrey), 1967- | Dickey, Jean | Hogan, Kelly A. Title: Campbell biology : concepts & connections / Jane B. Reece, Berkeley, California, Martha R. Taylor, Ithaca, New York, Eric J. Simon, New England College, Jean L. Dickey, Clemson University, Kelly Hogan, University of North Carolina, Chapel Hill. Other titles: Biology Description: Ninth edition. | Boston : Pearson Education,  Identifiers: LCCN 2016019061 Subjects: LCSH: Biology. Classification: LCC QH308.2 .B56448 2017 | DDC 570—dc23 LC record available at https://lccn.loc.gov/2016019061 1 16 ISBN 10: 0-134-29601-X; ISBN 13: 978-0-134-29601-2 (Student Edition) ISBN 10: 0-134-44277-6; ISBN 13: 978-0-134-44277-8 (Books a la Carte Edition) ISBN 10: 0-134-65340-8; ISBN 13: 978-0-134-65340-2 (NASTA Edition) www.pearsonhighered.com About the Authors Martha R. Taylor has been teaching biology for more than 35 years. She earned her B.A. in biology from Gettysburg College and her M.S. and Ph.D. in science education from Cornell University. At Cornell, Dr. Taylor has served as assistant director of the Office of Instructional Support and has taught introductory biology for both majors and nonmajors. Most recently, she was a lecturer in the Learning Strategies Center, teaching supplemental biology courses. Her experience working with students in classrooms, in laboratories, and with tutorials has increased her commitment to helping students create their own knowledge of and appreciation for biology. She was the author of the Student Study Guide for ten editions of Campbell Biology. Eric J. Simon is a professor in the Department of Biology and Health Science at New England College in Henniker, New Hampshire. He teaches introductory biology to science majors and nonscience majors, as well as upper-level courses in tropical marine biology and careers in science. Dr. Simon received a B.A. in biology and computer science and an M.A. in biology from Wesleyan University, and a Ph.D. in biochemistry from Harvard University. His research focuses on innovative ways to use technology to improve teaching and learning in the science classroom. Dr. Simon also leads numerous international student field research trips and is a Scientific Advisor to the Elephant Conservation Center in Sayaboury, Laos. Dr. Simon is the lead author of the introductory nonmajors biology textbooks Campbell Essential Biology, Sixth Edition, and Campbell Essential Biology with Physiology, Fifth Edition, and the author of the introductory biology textbook Biology: The Core, Second Edition. Jean L. Dickey is Professor Emerita of Biological Sciences at Clemson University (Clemson, South Carolina). After receiving her B.S. in biology from Kent State University, she went on to earn a Ph.D. in ecology and evolution from Purdue University. In 1984, Dr. Dickey joined the faculty at Clemson, where she devoted her career to teaching biology to nonscience majors in a variety of courses. In addition to creating content-based instructional materials, she developed many activities to engage lecture and laboratory students in discussion, critical thinking, and writing, and implemented an investigative laboratory curriculum in general biology. Dr. Dickey is author of Laboratory Investigations for Biology, Second Edition, and coauthor of Campbell Essential Biology, Sixth Edition, and Campbell Essential Biology with Physiology, Fifth Edition. Kelly Hogan is a faculty member in the Department of Biology at the University of North Carolina at Chapel Hill, teaching introductory biology and genetics. Dr. Hogan teaches hundreds of students at a time, using active-learning methods that incorporate educational technologies both inside and outside of the classroom. She received her B.S. in biology at the College of New Jersey and her Ph.D. in pathology at the University of North Carolina, Chapel Hill. Her research interests focus on how large classes can be more inclusive through evidence-based teaching methods and technology. As the Director of Instructional Innovation at UNC, she encourages experienced faculty to take advantage of new professional development opportunities and inspires the next generation of innovative faculty. Dr. Hogan is the author of Stem Cells and Cloning, Second Edition, and co-author on Campbell Essential Biology with Physiology, Fifth Edition. Jane B. Reece has worked in biology publishing since 1978, when she joined the editorial staff of Benjamin Cummings. Her education includes an A.B. in biology from Harvard University, an M.S. in microbiology from Rutgers University, and a Ph.D. in bacteriology from the University of California, Berkeley. At UC Berkeley, and later as a postdoctoral fellow in genetics at Stanford University, her research focused on genetic recombination in bacteria. Dr. Reece taught biology at Middlesex County College (New Jersey) and Queensborough Community College (New York). During her 12 years as an editor at Benjamin Cummings, she played a major role in a number of successful textbooks. She is coauthor of Campbell Biology, Eleventh Edition, Campbell Biology in Focus, Second Edition, Campbell Essential Biology, Sixth Edition, and Campbell Essential Biology with Physiology, Fifth Edition. Neil A. Campbell (1946–2004) combined the inquiring nature of a research scientist with the soul of a caring teacher. Over his 30 years of teaching introductory biology to both science majors and nonscience majors, many thousands of students had the opportunity to learn from him and be stimulated by his enthusiasm for the study of life. While he is greatly missed by his many friends in the biology community, his coauthors remain inspired by his visionary dedication to education and are committed to searching for ever better ways to engage students in the wonders of biology. About the Authors iii See Connections New Features of the ninth edition of Campbell Biology: Concepts & Connections provide students with a framework for understanding biological concepts and encourage students to see connections between concepts and the world outside of the classroom. NEW! Unit Openers highlight the relevancy of the course to careers in a variety of fields. Connection and Evolution Connection Modules present engaging examples and relate chapter content to evolution. NEW! A reframed focus on Major Themes in Biology provide students with a framework for understanding and organizing biological concepts. Icons throughout the text call students attention to examples of specific themes within each chapter. iv Build Science Literacy Skills 1996 Cases documented in every U.S. state except Alaska 2002 Major outbreak in Hong Kong; no cases since 2004 TH Severe acute respiratory syndrome UALIZI E TA West Nile virus NG AIDS 1981 To date, 71 million infected with HIV; 34 million dead 1918 Deadliest outbreak ever; 20–50 million dead in 18 months VIS H1N1 flu DA NEW! Visualizing the Data Figures are eye-catching infographics designed to provide students with a fresh approach to understanding concepts illustrated by quantitative information. H1N1 flu 2009 A combination of bird, swine, and human viruses Avian flu Zika fever 2015 Transmitted by mosquitoes; spread via sexual contact Ebola 1976 Biggest outbreak from 2014 to 2016 in West Africa 1997 Rarely occurs in North America Scientific Thinking modules explore how scientists use the process of science and discovery. End-ofmodule questions prompt students to think critically. A new presentation of the process of science in chapter 1 demonstrates to students the iterative nature of scientific research. v Visualize Tough Topics EXPANDED! Visualizing the Concept Modules bring dynamic visuals and text together to walk students through tough concepts. The ninth edition features 28 of these immersive modules. Select modules are assignable in MasteringBiology as animated videos. Embedded text coaches students through key points and help address common misunderstandings. vi and Develop Understanding Streamlined text and illustrations step students through the concept. NEW! Topics in the ninth edition include: 1.3: Hierarchy of Life 6.9: Oxidative Phosphorylation 8.17: Crossing Over 13.14: Natural Selection 25.4: Osmoregulation vii Encourage Focus on Main headings allow students to see the big picture. A Central Concept at the start of each module helps students to focus on one concept at a time. Try This activities in every chapter encourage students to actively engage with the figures and develop positive study habits. viii Key Concepts and Active Learning Checkpoint questions at the end of every module let students check their understanding right away. Figures describing a process take students through a series of numbered steps keyed to explanations in the text. Chapter summaries include figures and text to help students review and check their understanding of the chapter concepts. ix Continuous Learning Before, During, and After Class Before Class Interactive assignments introduce students to key concepts NEW! Key Topic Overview videos introduce students to key concepts and vocabulary and are created by authors Eric Simon, Jean Dickey and Kelly Hogan. All 12 videos are delivered as a whiteboard style mini-lesson and are accompanied by assessment so that students can check their understanding. Dynamic Study Modules provide students with multiple sets of questions with extensive feedback so that they can test, learn, and retest until they achieve mastery of the textbook material. x with MasteringBiology Create pre-lecture assignments with 170 author created interactive coaching activities. EXPANDED! Give students extra practice with 18 assignable Visualizing the Concept videos, which pair with the select modules in the text. xi Continuous Learning Before, During, and After Class During Class Encourage engagement with dynamic videos and resources for in class activities NEW! HHMI Short Films are documentary-quality movies from the Howard Hughes Medical Institute with explorations from the discovery of the double helix to evolution and include assignable questions. Chapter 4: A Tour of the Cell Guided Reading Activities, in the MasteringBiology study area, accompany all chapters and are designed to help students stay on track and develop active reading skills. Big idea: The nucleus and ribosomes Answer the following questions as you read modules 4.5–4.6: 1. DNA and its associated proteins are referred to as ____________. 2. Which of the following cells would be preparing to divide? Briefly explain your answer. A Resources to help instructors plan dynamic lectures: 3. • NEW! Ready-to-Go Teaching Modules help instructors efficiently make use of the available teaching tools for the toughest topics. • The Instructor Exchange provides active learning techniques from biology instructors around the nation. Co-author Kelly Hogan moderates the exchange. xii B Complete the following table that compares rRNA to mRNA. rRNA mRNA Role in/part of . . . Made in . . . Travels to . . . 4. Briefly describe the relationship between the nucleus and ribosomes. Your answer should include the following key terms: mRNA, rRNA, and protein synthesis. with MasteringBiology Learning Catalytics is a “bring your own device” (laptop, smartphone, or tablet) engagement, assessment, and classroom intelligence system that allows for active learning and discussion. NEW! Try This questions in Learning Catalytics are easy to assign in-class active learning questions, based on the text “Try This” feature. NEW! Everyday Biology Videos briefly explore interesting and relevant biology topics that relate to concepts that students are learning in class. These 20 videos, produced by the BBC, can be assigned in MasteringBiology. xiii Engage in Biology anytime, anywhere AFTER CLASS Dynamic activities let students put skills into practice Scientific Thinking Activities help students develop an understanding of how scientific research is conducted. Examples of topics include: • What Is the Role of Peer Review in the Process of Science? • How Does “Citizen Science” Affect Scientific Data Collection? • Do the Microorganisms in Our Digestive Tract Play a Role in Obesity? Current Events Activities cover a wide range of biological topics to demonstrate to students how science connects to everyday life. xiv with MasteringBiology NEW! Evaluating Science in the Media Activities teach students to recognize validity, bias, purpose, and authority in everyday sources of information. NEW! eText 2.0 is now available on smartphones, tablets and computers, featuring seamlessly integrated videos, and concept check questions. The eText 2.0 mobile app is available for most iOS and Android phones. Powerful interactive and customization functions include integrated videos and concept check questions, accessible (screen-reader ready), note-taking, highlighting, bookmarking, search, and links to glossary terms. xv Preface I Connections Students are more motivated to study biology nspired by the thousands of students in our own classes over the years and by enthusiastic feedback from the many instructors who have used or reviewed our book, we are delighted to present this new, Ninth Edition. We authors have worked together closely to ensure that both the book and the supplementary material online reflect the changing needs of today’s courses and students, as well as current progress in biology. Titled Campbell Biology: Concepts & Connections to honor Neil Campbell’s founding role and his many contributions to biology education, this book continues to have a dual purpose: to engage students from a wide variety of majors in the wonders of the living world and to show them how biology relates to their own existence and the world they inhabit. Most of these students will not become biologists themselves, but their lives will be touched by biology every day. Understanding the concepts of biology and their connections to our lives is more important than ever. Whether we’re concerned with our own health or the health of our planet, a familiarity with biology is essential. This basic knowledge and an appreciation for how science works have become elements of good citizenship in an era when informed evaluations of health issues, environmental problems, and applications of new technology are critical. when they can connect it to their own lives and interests—for example, when they are able to relate science to health issues, economic problems, environmental quality, ethical controversies, and social responsibility. In this edition, purple Connection icons mark the numerous application modules that go beyond the core biological concepts. For example, the new Connection Module 32.6 describes how humans tap into plant transport mechanisms for harvesting such materials as maple syrup and latex. In addition, our Evolution Connection modules, identified by green icons, connect the content of each chapter to the grand unifying theme of evolution, without which the study of life has no coherence. For example, a new Evolution Connection in Chapter 14 uses data from studies by Rosemary and Peter Grant and their students to demonstrate the continuing effects of natural selection on Darwin’s finches. Explicit connections are also made between the chapter introduction and either the Evolution Connection module or the Scientific Thinking module in each chapter; high-interest questions introduce each chapter, drawing students into the topic and encouraging a curiosity to explore the question further when it appears again later in the chapter. And, connections are made in every chapter between key concepts and the core concepts of biology. Concepts and Connections New to This Edition Concepts Biology is a vast subject that gets bigger every New Focus on Five Underlying Themes of Biology year, but an introductory biology course is still only one or two semesters long. This book was the first introductory biology textbook to use concept modules to help students recognize and focus on the main ideas of each chapter. The heading of each module is a carefully crafted statement of a key concept. For example, “Helper T cells stimulate the humoral and cell-mediated immune responses” announces a key concept about the role of helper T cells in adaptive immunity (Module 24.12). Such a concept heading serves as a focal point, and the module’s text and illustrations converge on that concept with explanation and, often, analogies. The module text walks the student through the illustrations, just as an instructor might do in class. And in teaching a sequential process, such as the one diagrammed in Figure 24.12A, we number the steps in the text to correspond to numbered steps in the figure. The synergy between a module’s narrative and graphic components transforms the concept heading into an idea with meaning to the student. The checkpoint question at the end of each module encourages students to test their understanding as they proceed through a chapter. Finally, in the Chapter Review, all the key concept statements are listed and briefly summarized under the overarching section titles, explicitly reminding students of what they’ve learned. xvi Preface A major goal of this Ninth Edition is to provide students with an explicit framework for understanding and organizing the broad expanse of biological information presented in Concepts and Connections. This framework is based on the five major themes outlined in Vision and Change in Undergraduate Biology Education: A Call to Action published by the American Academy for the Advancement of Science. These major themes extend across all areas of biology: evolution, the flow of information, the correlation of structure and function, the exchange of energy and matter, and the interactions and interconnections of biological systems. Chapter 1 introduces each of these themes in a separate module. Specific examples of the themes are then called out in each chapter by green icons: INFORMATION , STRUCTURE AND FUNCTION , ENERGY AND MATTER , INTERACTIONS , and EVOLUTION CONNECTION (always in module form). Expanded Coverage of the Process of Science Chapter 1 also includes an enhanced focus on the nature of science and the process of scientific inquiry, setting the stage for both the content of the text and the process by which our biological knowledge has been built and continues to grow. We continue this emphasis on the process of scientific inquiry through our Scientific Thinking modules in every chapter, which are called out with an orange icon. New concept check questions for these modules focus on aspects of the process of science: the forming and testing of hypotheses; experimental design; variables and controls; the analysis of data; and the evaluation and communication of scientific results. Additional Visualizing the Concept Modules These modules, which were new to the Eighth Edition, have raised our hallmark art–text integration to a new level. Visualizing the Concept modules take challenging concepts or processes and walk students through them in a highly visual manner, using engaging, attractive art; clear and concise labels; and instructor “hints” called out in light blue bubbles. These short hints emulate the one-on-one coaching an instructor might provide to a student during office hours and help students make key connections within the figure. Examples of the eight new Visualizing the Concept modules include Module 6.9, Most ATP production occurs by oxidative phosphorylation; Module 8.17, Crossing over further increases genetic variability; Module 13.14, Natural selection can alter variation in a population in three ways; Module 28.6, Neurons communicate at synapses, and Module 34.18, The global water cycle connects aquatic and terrestrial biomes. New Visualizing the Data Figures Also new to this VIS NG TH TA edition are figures that present data in an infographic form, marked by Visualizing the Data icons. UALIZI These 19 eye-catching figures provide students E DA with a fresh approach to understanding the concepts illustrated by graphs and numerical data. Figure 10.19 maps emergent virus outbreaks, showing that they originate throughout the world. Figure 12.17 summarizes a wealth of bioinformatics data on genome sizes versus the number of genes found in various species. Figure 13.16 illustrates the growing threat of antibiotic resistant bacteria. Figure 21.14 allows students to directly compare caloric intake (via food) with caloric expenditure (via exercise). Figure 30.5B shows changes in bone mass during the human life span. Figure 36.11 offers an illuminating visual comparison of the per capita and national ecological footprints of several countries with world average and “fair share” footprints. Figure 38.3 shows graphic evidence of global warming by tracking annual global temperatures since 1880. New Unit Openers That Feature Careers Related to the Content of the Unit Expanding our emphasis on the connections of biology to students’ lives, each unit opener page now includes photos of individuals whose professions relate to the content of the unit. For instance, Unit I features a brewery owner and a solar energy engineer. Unit IV portrays a hatchery manager and a paleoanthropologist. These examples are intended to help students see how their biology course relates to the world outside the classroom and to their own career paths. New Design and Improved Art The fresh new design used throughout the chapters and the extensive reconceptualization of many figures make the book even more appealing and accessible to visual learners. Much of the art in Chapter 6, How Cells Harvest Chemical Energy, for example, has been revised to help students work through the complex reactions of cellular respiration. Other examples of improved art are found in Figures 5.15B, 10.11A, and 37.22B. The Latest Science Biology is a dynamic field of study, and we take pride in our book’s currency and scientific accuracy. For this edition, as in previous editions, we have integrated the results of the latest scientific research throughout the book. We have done this carefully and thoughtfully, recognizing that research advances can lead to new ways of looking at biological topics; such changes in perspective can necessitate organizational changes in our textbook to better reflect the current state of a field. For example, Chapter 12 uses both text and art to present the innovative CRISPR-Cas9 system for gene editing. You will find a unit-by-unit account of new content and organizational improvements in the “New Content” section on pages xix–xx following this Preface. MasteringBiology® MasteringBiology, the most widely used online tutorial and assessment program for biology, continues to accompany Campbell Biology: Concepts & Connections. In addition to 170 author-created activities that help students learn vocabulary, extend the book’s emphasis on visual learning, demonstrate the connections among key concepts (helping students grasp the big ideas), and coach students on how to interpret data, the Ninth Edition features new assignable videos. These videos bring this text’s Visualizing the Concept modules to life, help students learn how to evaluate sources of scientific information for reliability, and include short news videos that engage students in the many ways course concepts connect to the world outside the classroom. MasteringBiology® for Campbell Biology: Concepts & Connections, Ninth Edition, will help students to see strong connections through their text, and the additional practice available online allows instructors to capture powerful data on student performance, thereby making the most of class time. This Book’s Flexibility Although a biology textbook’s table of contents is by design linear, biology itself is more like a web of related concepts without a single starting point or prescribed path. Courses can navigate this network by starting with molecules, with ecology, or somewhere in-between, and courses can omit topics. Campbell Biology: Concepts & Connections is uniquely suited to offer flexibility and thus serve a variety of courses. The seven units of the book are largely self-contained, and in a number of the units, chapters can be assigned in a different order without much loss of coherence. The use of numbered modules makes it easy to skip topics or reorder the presentation of material. ■ ■ ■ Preface xvii For many students, introductory biology is the only science course that they will take during their college years. Long after today’s students have forgotten most of the specific content of their biology course, they will be left with general impressions and attitudes about science and scientists. We hope that this new edition of Campbell Biology: Concepts & Connections helps make those impressions positive and supports instructors’ goals for sharing the fun of biology. In our continuing efforts to improve the book and its supporting materials, we benefit tremendously from instructor and student feedback, not only in formal reviews but also via informal communication. Please let us know how we are doing and how we can improve the next edition of the book. xviii Preface Martha Taylor (Chapter 1 and Unit I), email@example.com Eric Simon (Units II and VI and Chapters 21 and 27), SimonBiology@gmail.com Jean Dickey (Units III, IV, and VII and Chapters 22 and 30), firstname.lastname@example.org Kelly Hogan (Chapters 20, 23–26, 28, and 29), email@example.com Jane Reece firstname.lastname@example.org New Content B elow are some important highlights of new content and organizational improvements in Campbell Biology: Concepts & Connections, Ninth Edition. Chapter 1, Biology: Exploring Life This chapter has been extensively reorganized and revised. Our expanded coverage of the nature of science and scientific inquiry has now moved to the forefront of Chapter 1. The first of the five modules in this section provides a general description of data, hypothesis formation and testing, the centrality of verifiable evidence to science, and an explanation of scientific theories. The module describing how hypotheses can be tested using controlled experiments now includes a subsection on hypothesis testing in humans. A new Scientific Thinking module entitled Hypotheses can be tested using observational data, describes how multiple lines of evidence, including DNA comparisons, have helped resolve the classification of the red panda. Another new module—The process of science is repetitive, nonlinear, and collaborative—presents a more accurate model of the process of science that includes four interacting circles: Exploration and Discovery; Forming and Testing Hypotheses: Analysis and Feedback from the Scientific Community; and Societal Benefits and Outcomes. The chapter concludes with the introduction of five core themes that underlie all of biology: evolution; information; structure and function; energy and matter; and interactions. Unit I, The Life of the Cell This unit guides students from basic chemistry and the molecules of life through cellular structures to cellular respiration and photosynthesis. Throughout the Ninth Edition, the five themes introduced in Chapter 1 are highlighted with specific references. Examples from Unit 1 include “Illustrating our theme of ENERGY AND MATTER , we see that matter has been rearranged, with an input of energy provided by sunlight” (Module 2.9); “The flow of genetic instruction that leads to gene expression, summarized as DNA S RNA S protein, illustrates the important biological theme of INFORMATION ” (Module 3.15); “The interconnections among these pathways provide a clear example of the theme of INTERACTIONS in producing the emergent property of a balanced metabolism” (Module 6.15); and “The precise arrangements of these membranes and compartments are essential to the process of photosynthesis—a classic example of the theme of STRUCTURE AND FUNCTION ” (Module 7.2). The theme of evolution is featured, as it is in every chapter, in an Evolution Connection module, such as Module 4.15, Mitochondria and chloroplasts evolved by endosymbiosis. Two new Visualizing the Concept modules are Module 2.6, Covalent bonds join atoms into molecules through electron sharing, and Module 6.9, Most ATP production occurs by oxidative phosphorylation. Both use new and highly revised art to guide students through these challenging topics. The Connection Module 2.2, Trace elements are common additives to food and water, uses added information on water fluoridation to emphasize the process of science and societal interactions. Two new Connection modules are Module 3.6, Are we eating too much sugar? (which includes a Visualizing the Data figure on recommended and actual sugar consumption), and Module 7.14, Reducing both fossil fuel use and deforestation may moderate climate change (which includes information on the 2015 Paris climate accord). New orientation diagrams help students follow the various stages of cellular respiration and photosynthesis in Chapters 6 and 7. Unit II, Cellular Reproduction and Genetics The purpose of this unit is to help students understand the relationship between DNA, chromosomes, and organisms and to help students see that genetics is not purely hypothetical but connects in many important and interesting ways to their lives, human society, and other life on Earth. The content has been reinforced with updated discussions of relevant topics, such as DCIS (also called stage 0 breast cancer), increased use of genetically modified organisms (GMOs), recent examples of DNA profiling, information about the 2015 California measles outbreak, a new infographic that charts emergent virus outbreaks, and new data on the health prospects of clones. This edition includes discussion of many recent advances in the field, such as an updated definition of the gene, and a largely new presentation of DNA technologies and bioinformatics, including extensive discussion in both text and art of the CRISPR-Cas9 system, GenBank, and BLAST searches. In some cases, sections within chapters have been reorganized to present a more logical flow of materials. Examples of new organization include an improved presentation of the genetics underlying cancer, a new Visualizing the Concept module on crossing over, a new circular genetic code chart that should improve student understanding, and a new Visualizing the Data that summarizes relevant information about different types of cancer and their survival rates. Material throughout the unit has been updated to reflect recent data, such as the latest statistics on cancer, cystic fibrosis, and Down syndrome, an improved model of ribosomes, new information about prions, expanded coverage of noncoding small RNAs, new human gene therapy trials, and recent information about Y chromosome inheritance. Unit III, Concepts of Evolution This unit presents the basic principles of evolution and natural selection, the overwhelming evidence that supports these theories, and their relevance to all of biology—and to the lives of students. For example, a new Visualizing the Data figure (13.16) illustrates the growing threat of antibiotic resistance. Chapter 13 also includes a new Visualizing the Concept module (13.14) on the effects of natural selection that shows experimental data along with hypothetical examples. Chapter 14 contains a new New Content xix Evolution Connection module (14.9) featuring the work of Rosemary and Peter Grant on Darwin’s finches. Modules 15.14 to 15.19 were revised to improve the flow and clarity of the material on phylogenetics and include updates from genomic studies and new art (for example, Figures 15.17 and 15.19A). Unit IV, The Evolution of Biological Diversity The diversity unit surveys all life on Earth in less than a hundred pages! Consequently, descriptions and illustrations of the unifying characteristics of each major group of organisms, along with a small sample of its diversity, make up the bulk of the content. Two recurring elements are interwoven with these descriptions: evolutionary history and examples of relevance to our everyday lives and society at large. With the rapid accumulation of molecular evidence, taxonomic revisions are inevitable. These changes are reflected in Chapter 16, Microbial Life, with a new module and figure (16.13) on protist supergroups, and in Chapters 18 and 19, Evolution of Invertebrate Diversity and Evolution of Vertebrate Diversity, with three modules about animal phylogeny (18.10, 18.11, and 19.1). The importance of metagenomics to the study of microorganisms is highlighted in Modules 16.1 and 16.7 (prokaryotes) and 17.14 (fungi). New examples of relevance include valley fever, a fungal disease linked to climate change (Module 17.19), and a Visualizing the Data figure (19.16) on the evolution of human skin color. Unit V, Animals: Form and Function This unit combines a comparative animal approach with an exploration of human anatomy and physiology. The introduction to Chapter 20, Unifying Concepts of Animal Structure and Function, begins with the question “Does evolution lead to the perfect animal form?” and the question is answered in the Evolution Connection, Module 20.1, in discussion of the lengthy laryngeal nerve in giraffes. By illustrating that a structure in an ancestral organism can become adapted to function in a descendant organism without being “perfected,” this example helps to combat a common student misconception about evolution. The main portion of every chapter in this unit is devoted to detailed presentations of human body systems, frequently illuminated by discussion of the health consequences of disorders in those systems. The Chapter 22 opening essay and Scientific Thinking module (22.7) were revised to compare the conclusions from long term studies on the health hazards of cigarette smoking with the very recent research on the effects of e-cigarettes. In Chapter 23, Circulation, the Scientific Thinking module (23.6) discusses the consequences of treating coronary artery disease with medicine or both medicine and stents. Chapter 29, The Senses, incorporates new material on common eye conditions, glaucoma and cataracts. In many areas, content has been updated to reflect newer issues in biology. New modules include 24.9 on the importance of community vaccination, 28.18 on neuronal plasticity, and 29.12 about the contribution of genes in one’s perception of the taste of cilantro. New Visualizing the Concept modules on osmoregulation (25.4) and neuronal synapses (28.6) help students better envision big concepts. New Visualizing the Data figures detail data on hypertension in the United States (23.9B), worldwide HIV infection and treatments (24.14B), and changes in bone mass during the human life span (30.5B). Chapter 21, Nutrition and Digestion, xx New Content includes a new discussion of human microbiome and microbiota as well as presentation of the forthcoming changes to food nutritional labels. Module 22.9, Breathing is automatically controlled, was heavily revised. The equation showing the formation and dissociation of carbonic acid now accompanies the discussion of how the medulla regulates breathing, a process illustrated by new art. Improvements to this unit also include a significant revision to the presentation of the kidney as a water-conserving organ (25.7) and a clearer four-step process by which a sensory stimulus results in a perception (Module 29.1). Chapter 27, Reproduction and Embryonic Development, presents data on the decreased incidence of cervical cancer due to early detection, a new Visualizing the Data (Figure 27.8) that summarizes different methods of contraception, and new information on reproductive technologies. Unit VI, Plants: Form and Function To help students gain an appreciation of the importance of plants, this unit presents the anatomy and physiology of angiosperms with frequent connections to the importance of plants to society. New Connections modules in this edition include an improved discussion of agriculture via artificial selection on plant parts and via plant cloning in Chapter 31; updated discussions of organic farming, human harvesting of plant transport products (such as maple syrup and rubber), and GMOs in Chapter 32; and a new discussion of caffeine as an evolutionary adaptation that can prevent herbivory in Chapter 33. Throughout the unit, the text has been revised with the goal of making the material more engaging and accessible to students. For example, the discussion of plant nutrients has been entirely reorganized into a large Visualizing the Data in Module 32.7, and the presentation of the potentially confusing topic of the effect of auxin on plant cell elongation benefits from a new visual presentation (Figure 33.3B). All of these changes are meant to make the point that human society is inexorably connected to the health of plants. Unit VII, Ecology In this unit, students learn the fundamental principles of ecology and how these principles apply to environmental problems. The Ninth Edition features a new Visualizing the Concept module that explains the global water cycle (34.18) and Visualizing the Data figures that compare ecological footprints (36.11), track global temperatures since 1880 (38.3A), and illustrate the results of a study on optimal foraging theory (35.12). Module 35.16 has been updated with new examples of the effects of endocrine-disrupting chemicals on animal behavior and the EPA’s progress in evaluating endocrine disruptors in pesticides as potential hazards to human health. Other content updates in this unit include human population data (36.9 and 36.10), species at risk for extinction (38.1), and the new federal law banning the use of microbeads in health and beauty products (38.2). Module 37.13 has been heavily revised to include more examples of invasive species. The unit-wide emphasis on climate change and sustainability continues in this edition. For example, the module on ecological footprints (36.11) has been updated and revised, and Module 37.23 includes a new emphasis on the role of wetlands in mitigating the effects of climate change. Figures 38.3B and 38.4A were updated with the most recent data available, and Module 38.3 was heavily revised. Acknowledgments T his Ninth Edition of Campbell Biology: Concepts & Connections is a result of the combined efforts of many talented and hardworking people, and the authors wish to extend heartfelt thanks to all those who contributed to this and previous editions. Our work on this edition was shaped by input from the biologists acknowledged in the reviewer list on pages xxii–xxiv, who shared with us their experiences teaching introductory biology and provided specific suggestions for improving the book. Feedback from the authors of this edition’s supplements and the unsolicited comments and suggestions we received from many biologists and biology students were also extremely helpful. In addition, this book has benefited in countless ways from the stimulating contacts we have had with the coauthors of Campbell Biology, Eleventh Edition. We wish to offer special thanks to the students and faculty at our teaching institutions. Marty Taylor thanks her students at Cornell University for their valuable feedback on the book. Eric Simon thanks his colleagues and friends at New England College, especially within the collegium of Natural Sciences and Mathematics, for their continued support and assistance. Jean Dickey thanks her colleagues at Clemson University for their expertise and support. And Kelly Hogan thanks her students for their enthusiasm and colleagues at the University of North Carolina, Chapel Hill, for their continued support. We thank Paul Corey, managing director of Higher Education Learning Services. In addition, the superb publishing team for this edition was headed up by courseware portfolio management specialist Alison Rodal, with the invaluable support of courseware portfolio management director Beth Wilbur. We cannot thank them enough for their unstinting efforts on behalf of the book and for their commitment to excellence in biology education. We are fortunate to have had once again the contributions of courseware director of content development Ginnie Simione Jutson. We are similarly grateful to the members of the editorial development team—Debbie Hardin, Evelyn Dahlgren, Julia Osborne, Susan Teahan, and Mary Catherine Hager—for their steadfast commitment to quality. We thank them for their thoroughness, hard work, and good humor; the book is far better than it would have been without their efforts. Thanks also to supplements project editor Melissa O’Conner on her oversight of the supplements program and to editorial coordinator Alison Cagle for the efficient and enthusiastic support she provided. This book and all the other components of the teaching package are both attractive and pedagogically effective in large part because of the hard work and creativity of the production professionals on our team. We wish to thank managing producer Mike Early and content producers Mae Lum and Courtney Towson. We also acknowledge copy editor Jon Preimesberger, proofreaders Pete Shanks and Joanna Dinsmore, and indexer Tim Engman. We again thank photo researcher Kristin Piljay for her contributions, as well as text permissions manager K. Ganesh and photo permissions manager Eric Shrader. Integra was responsible for composition, headed by production project manager Alverne Ball, and the art house Lachina, headed by project manager Whitney Philipp, who was responsible for overseeing the rendering of new and revised illustrations. We also thank manufacturing overseer Stacey Weinberger. We thank Elise Lansdon for creating a beautiful and functional interior design and a stunning cover, and we are again indebted to design manager Mark Ong for his oversight and design leadership. The value of Campbell Biology: Concepts & Connections as a learning tool is greatly enhanced by the hard work and creativity of the authors of the supplements that accompany this book: Ed Zalisko (Instructor’s Guide and PowerPoint® Lecture Presentations); Jean DeSaix, Kristen Miller, Justin Shaffer, and Suann Yang (Test Bank); Dana Kurpius (Active Reading Guide); Bob Iwan and Sukanya Subramanian (Reading Quizzes); Cheri LaRue (media correlator), and Brenda Hunzinger (Clicker Questions and Quiz Shows). In addition to supplements project editor Melissa O’Conner, the editorial and production staff for the supplements program included supplements production project manager Alverne Ball (Integra), Marsha Hall (PPS), and Jennifer Hastings (PPS). And the superlative MasteringBiology® program for this book would not exist without Lauren Fogel, Stacy Treco, Katie Foley, Sarah Jensen, Chloé Veylit, Jim Hufford, Charles Hall, Caroline Power, and David Kokorowski and his team. And a special thanks to Arl Nadel and Sarah Young-Dualan for their thoughtful work on the Visualizing the Concepts interactive videos. For their important roles in marketing the book, we are very grateful to marketing manager Christa Pelaez and vice president of marketing Christy Lesko. The members of the Pearson Science sales team have continued to help us connect with biology instructors and their teaching needs, and we thank them. Finally, we are deeply grateful to our families and friends for their support, encouragement, and patience throughout this project. Our special thanks to Paul, Dan, Maria, Armelle, and Sean (J.B.R.); Josie, Jason, Marnie, Alice, Jack, David, Paul, Ava, and Daniel (M.R.T.); Amanda, Reed, Forest, and dear friends Jamey, Nick, Jim, and Bethany (E.J.S.); Jessie and Katherine (J.L.D.); and Tracey, Vivian, Carolyn, Brian, Jake, and Lexi (K.H.) Jane Reece, Martha Taylor, Eric Simon, Jean Dickey, and Kelly Hogan Acknowledgments xxi Reviewers Reviewers of the Ninth Edition Ellen Baker, Santa Monica College Deborah Cardenas, Collin College Marc DalPonte, Lake Land College Tammy Dennis, Bishop State Community College Jean DeSaix, University of North Carolina, Chapel Hill Cynthia Galloway, Texas A&M University Jan Goerrissen, Orange Coast College Christopher Haynes, Shelton State Andrew Hinton, San Diego City College Duane Hinton, Washburn University Brenda Hunzinger, Lake Land College Robert Iwan, Inver Hills Community College Cheri LaRue, University of Arkansas, Fayetteville Barbara Lax, Community College of Allegheny County Brenda Leady, University of Toledo Sheryl Love, Temple University David Luther, George Mason University Steven MacKie, Pima County Community College Thaddeus McRae, Broward Community College Kristen Miller, University of Georgia Debbie Misencik, Community College of Allegheny County Justin Shaffer, University of California, Irvine Erica Sharar, Santiago Canyon College Patricia Steinke, San Jacinto College Central Jennifer Stueckle, West Virginia University Sukanya Subramanian, Collin County Community College Brad Williamson, University of Kansas Suann Yang, Presbyterian College Edward Zalisko, Blackburn College Media Review Panel, Ninth Edition Bob Iwan, Inver Hills Community College Cheri LaRue, University of Arkansas Linda Logdberg Lindsay Rush, Quinnipiac University Sukanya Subramanian, Collin County Community Reviewers of Previous Editions Michael Abbott, Westminster College Tanveer Abidi, Kean University Daryl Adams, Mankato State University Dawn Adrian Adams, Baylor University Olushola Adeyeye, Duquesne University Shylaja Akkaraju, Bronx Community College Felix Akojie, Paducah Community College Dan Alex, Chabot College John Aliff, Georgia Perimeter College Sylvester Allred, Northern Arizona University Jane Aloi-Horlings, Saddleback College Loren Ammerman, University of Texas at Arlington Dennis Anderson, Oklahoma City Community College xxii Reviewers Marjay Anderson, Howard University Steven Armstrong, Tarrant County College Bert Atsma, Union County College Yael Avissar, Rhode Island College Gail Baker, LaGuardia Community College Caroline Ballard, Rock Valley College Andrei Barkovskii, Georgia College and State University Mark Barnby, Ohlone College Chris Barnhart, University of San Diego Stephen Barnhart, Santa Rosa Junior College William Barstow, University of Georgia Kirk A. 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Carabelli, Broward Community College Jocelyn Cash, Central Piedmont Community College Cathryn Cates, Tyler Junior College Russell Centanni, Boise State University David Chambers, Northeastern University Ruth Chesnut, Eastern Illinois University Vic Chow, San Francisco City College Van Christman, Ricks College Craig Clifford, Northeastern State University, Tahlequah Richard Cobb, South Maine Community College Glenn Cohen, Troy University Mary Colavito, Santa Monica College Jennifer Cooper, Itawamba Community College Bob Cowling, Ouachita Technical College Don Cox, Miami University Robert Creek, Western Kentucky University Hillary Cressey, George Mason University Norma Criley, Illinois Wesleyan University Jessica Crowe, South Georgia College Mitch Cruzan, Portland State University Judy Daniels, Monroe Community College Michael Davis, Central Connecticut State University Pat Davis, East Central Community College Lewis Deaton, University of Louisiana Lawrence DeFilippi, Lurleen B. Wallace College James Dekloe, Solano Community College Veronique Delesalle, Gettysburg College Loren Denney, Southwest Missouri State University Jean DeSaix, University of North Carolina at Chapel Hill Mary Dettman, Seminole Community College of Florida Kathleen Diamond, College of San Mateo Alfred Diboll, Macon College Jean Dickey, Clemson University Stephen Dina, St. Louis University Robert P. 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Yeh, San Francisco, California William Yurkiewicz, Millersville University of Pennsylvania Gregory Zagursky, Radford University Martin Zahn, Thomas Nelson Community College Edward J. Zalisko, Blackburn College David Zeigler, University of North Carolina at Pembroke Uko Zylstra, Calvin College Detailed Contents 1 Biology: Exploring Life 1 2.3 Biology: The Scientific Study of Life 2 2.4 1.1 1.2 What is life? 2 Biologists arrange the diversity of life into three domains 3 1.3 VISUALIZING THE CONCEPT In life’s hierarchy of organization, new properties emerge at each level 4 The Process of Science 6 1.4 1.5 1.6 1.7 1.8 What is science? 6 Hypotheses can be tested using controlled experiments 7 Scientific Thinking Hypotheses can be tested using observational data 8 The process of science is repetitive, nonlinear, and collaborative 8 Connection Biology, technology, and society are connected in important ways 9 Five Unifying Themes in Biology 10 1.9 Theme: Evolution is the core theme of biology 10 1.10 Evolution Connection Evolution is connected to our everyday lives 12 1.11 Theme: Life depends on the flow of information 12 1.12 Theme: Structure and function are related 14 1.13 Theme: Life depends on the transfer and transformation of energy and matter 15 1.14 Theme: Life depends on interactions within and between systems 16 Chapter Review 17 Atoms consist of protons, neutrons, and electrons 24 Connection Radioactive isotopes can help or harm us 25 Chemical Bonds 26 2.5 2.6 2.7 2.8 2.9 The distribution of electrons determines an atom’s chemical properties 26 VISUALIZING THE CONCEPT Covalent bonds join atoms into molecules through electron sharing 27 Ionic bonds are attractions between ions of opposite charge 28 Hydrogen bonds are weak bonds important in the chemistry of life 28 Chemical reactions make and break chemical bonds 29 Water’s Life-Supporting Properties 30 Hydrogen bonds make liquid water cohesive 30 Water’s hydrogen bonds moderate temperature 30 Ice floats because it is less dense than liquid water 31 Water is the solvent of life 31 The chemistry of life is sensitive to acidic and basic conditions 32 2.15 Scientific Thinking Scientists study the effects of rising atmospheric CO2 on coral reef ecosystems 32 2.16 Evolution Connection The search for extraterrestrial life centers on the search for water 33 Chapter Review 34 2.10 2.11 2.12 2.13 2.14 3 The Molecules of Cells 36 Introduction to Organic Compounds 38 3.1 U nit I The Life of the Cell 19 2 The Chemical Basis of Life 20 Elements, Atoms, and Compounds 22 2.1 2.2 Organisms are composed of elements, usually combined into compounds 22 Connection Trace elements are common additives to food and water 23 3.2 3.3 Life’s molecular diversity is based on the properties of carbon 38 A few chemical groups are key to the functioning of biological molecules 39 Cells make large molecules from a limited set of small molecules 40 Carbohydrates 41 3.4 3.5 3.6 3.7 Monosaccharides are the simplest carbohydrates 41 Two monosaccharides are linked to form a disaccharide 42 Connection Are we eating too much sugar? 42 Polysaccharides are long chains of sugar units 43 Detailed Contents xxv Lipids 44 Fats are lipids that are mostly energy-storage molecules 44 3.9 Scientific Thinking Scientific studies document the health risks of trans fats 45 3.10 Phospholipids and steroids are important lipids with a variety of functions 46 3.11 Connection Anabolic steroids pose health risks 46 3.8 Proteins 47 3.12 Proteins have a wide range of functions and structures 47 3.13 Proteins are made from amino acids linked by peptide bonds 48 3.14 VISUALIZING THE CONCEPT A protein’s functional shape results from four levels of structure 49 Nucleic Acids 50 3.15 The nucleic acids DNA and RNA are informationrich polymers of nucleotides 50 3.16 Evolution Connection Lactose tolerance is a recent event in human evolution 51 Chapter Review 52 4 A Tour of the Cell 54 Introduction to the Cell 56 4.1 4.2 4.3 4.4 Microscopes reveal the world of the cell 56 The small size of cells relates to the need to exchange materials across the plasma membrane 58 Prokaryotic cells are structurally simpler than eukaryotic cells 59 Eukaryotic cells are partitioned into functional compartments 60 The Nucleus and Ribosomes 62 4.5 4.6 The nucleus contains the cell’s genetic instructions 62 Ribosomes make proteins for use in the cell and for export 63 The Endomembrane System 63 Many organelles are connected in the endomembrane system 63 4.8 The endoplasmic reticulum is a biosynthetic workshop 64 4.9 The Golgi apparatus modifies, sorts, and ships cell products 65 4.10 Lysosomes are digestive compartments within a cell 66 4.11 Vacuoles function in the general maintenance of the cell 66 4.12 A review of the structures involved in manufacturing and breakdown 67 4.7 Energy-Converting Organelles 67 4.13 Mitochondria harvest chemical energy from food 67 xxvi Detailed Contents 4.14 Chloroplasts convert solar energy to chemical energy 68 4.15 Evolution Connection Mitochondria and chloroplasts evolved by endosymbiosis 68 The Cytoskeleton and Cell Surfaces 69 4.16 The cell’s internal skeleton helps organize its structure and activities 69 4.17 Scientific Thinking Scientists discovered the cytoskeleton using the tools of biochemistry and microscopy 70 4.18 Cilia and flagella move when microtubules bend 70 4.19 The extracellular matrix of animal cells functions in support and regulation 71 4.20 Three types of cell junctions are found in animal tissues 72 4.21 Cell walls enclose and support plant cells 72 4.22 Review: Eukaryotic cell structures can be grouped on the basis of four main functions 73 Chapter Review 74 5 The Working Cell 76 Membrane Structure and Function 78 5.1 5.2 5.3 5.4 5.5 5.6 5.7 5.8 5.9 VISUALIZING THE CONCEPT Membranes are fluid mosaics of lipids and proteins with many functions 78 Evolution Connection The spontaneous formation of membranes was a critical step in the origin of life 79 Passive transport is diffusion across a membrane with no energy investment 79 Osmosis is the diffusion of water across a membrane 80 Water balance between cells and their surroundings is crucial to organisms 80 Transport proteins can facilitate diffusion across membranes 81 Scientific Thinking Research on another membrane protein led to the discovery of aquaporins 82 Cells expend energy in the active transport of a solute 82 Exocytosis and endocytosis transport large molecules across membranes 83 Energy and the Cell 84 5.10 Cells transform energy and matter as they perform work 84 5.11 Chemical reactions either release or store energy 85 5.12 ATP drives cellular work by coupling exergonic and endergonic reactions 86 How Enzymes Function 87 5.13 Enzymes speed up the cell’s chemical reactions by lowering energy barriers 87 5.14 A specific enzyme catalyzes each cellular reaction 88 5.15 Enzyme inhibition can regulate enzyme activity in a cell 89 5.16 Connection Many drugs, pesticides, and poisons are enzyme inhibitors 89 Chapter Review 90 6 How Cells Harvest Chemical Energy 92 Cellular Respiration: Aerobic Harvesting of Energy 94 6.1 6.2 6.3 6.4 6.5 Photosynthesis and cellular respiration provide energy for life 94 Breathing supplies O2 for use in cellular respiration and removes CO2 94 Cellular respiration banks energy in ATP molecules 95 Connection The human body uses energy from ATP for all its activities 95 Cells capture energy from electrons “falling” from organic fuels to oxygen 96 Stages of Cellular Respiration 97 Overview: Cellular respiration occurs in three main stages 97 6.7 Glycolysis harvests chemical energy by oxidizing glucose to pyruvate 98 6.8 After pyruvate is oxidized, the citric acid cycle completes the energy-yielding oxidation of organic molecules 100 6.9 VISUALIZING THE CONCEPT Most ATP production occurs by oxidative phosphorylation 102 6.10 Scientific Thinking Scientists have discovered heat-producing, calorie-burning brown fat in adults 103 6.11 Review: Each molecule of glucose yields many molecules of ATP 104 6.6 Fermentation: Anaerobic Harvesting of Energy 104 6.12 Fermentation enables cells to produce ATP without oxygen 104 6.13 Evolution Connection Glycolysis evolved early in the history of life on Earth 106 Connections Between Metabolic Pathways 106 6.14 Cells use many kinds of organic molecules as fuel for cellular respiration 106 6.15 Organic molecules from food provide raw materials for biosynthesis 107 Chapter Review 108 7 Photosynthesis: Using Light to Make Food 110 An Introduction to Photosynthesis 112 7.1 7.2 7.3 7.4 7.5 Photosynthesis fuels the biosphere 112 Photosynthesis occurs in chloroplasts in plant cells 113 Scientists traced the process of photosynthesis using isotopes 114 Photosynthesis is a redox process, as is cellular respiration 114 Photosynthesis occurs in two stages, which are linked by ATP and NADPH 115 The Light Reactions: Converting Solar Energy to Chemical Energy 116 7.6 7.7 7.8 7.9 Visible radiation absorbed by pigments drives the light reactions 116 Photosystems capture solar energy 117 Two photosystems connected by an electron transport chain convert light energy to the chemical energy of ATP and NADPH 118 VISUALIZING THE CONCEPT The light reactions take place within the thylakoid membranes 119 The Calvin Cycle: Reducing CO2 to Sugar 120 7.10 ATP and NADPH power sugar synthesis in the Calvin cycle 120 7.11 Evolution Connection Other methods of carbon fixation have evolved in hot, dry climates 121 The Global Significance of Photosynthesis 122 7.12 Photosynthesis provides food and O2 for almost all living organisms 122 7.13 Scientific Thinking Rising atmospheric levels of carbon dioxide may affect plants in various ways 123 7.14 Connection Reducing both fossil fuel use and deforestation may moderate climate change 124 Chapter Review 125 U nit I I Cellular Reproduction and Genetics 127 8 The Cellular Basis of Reproduction and Inheritance 128 Cell Division and Reproduction 130 8.1 Cell division plays many important roles in the lives of organisms 130 Detailed Contents xxvii 8.2 Prokaryotes reproduce by binary fission 131 The Eukaryotic Cell Cycle and Mitosis 132 The large, complex chromosomes of eukaryotes duplicate with each cell division 132 8.4 The cell cycle includes growth and division phases 133 8.5 Cell division is a continuum of dynamic changes 134 8.6 Cytokinesis differs for plant and animal cells 136 8.7 The rate of cell division is affected by environmental factors 137 8.8 Growth factors signal the cell cycle control system 138 8.9 Connection Growing out of control, cancer cells produce malignant tumors 139 8.10 Scientific Thinking The best cancer treatment may vary by individual 140 8.3 Meiosis and Crossing Over 140 8.11 Chromosomes are matched in homologous pairs 140 8.12 Gametes have a single set of chromosomes 141 8.13 Meiosis reduces the chromosome number from diploid to haploid 142 8.14 VISUALIZING THE CONCEPT Mitosis and meiosis have important similarities and differences 144 8.15 Independent orientation of chromosomes in meiosis and random fertilization lead to varied offspring 145 8.16 Homologous chromosomes may carry different versions of genes 146 8.17 VISUALIZING THE CONCEPT Crossing over further increases genetic variability 147 Alterations of Chromosome Number and Structure 148 8.18 Accidents during meiosis can alter chromosome number 148 8.19 A karyotype is a photographic inventory of an individual’s chromosomes 149 8.20 Connection An extra copy of chromosome 21 causes Down syndrome 150 8.21 Connection Abnormal numbers of sex chromosomes do not usually affect survival 151 8.22 Evolution Connection New species can arise from errors in cell division 151 8.23 Connection Alterations of chromosome structure can cause birth defects and cancer 152 Chapter Review 153 9 Patterns of Inheritance 156 Mendel’s Laws 158 9.1 9.2 9.3 xxviii The study of genetics has ancient roots 158 The science of genetics began in an abbey garden 158 Mendel’s law of segregation describes the inheritance of a single character 160 Detailed Contents Homologous chromosomes bear the alleles for each character 161 9.5 The law of independent assortment is revealed by tracking two characters at once 162 9.6 Geneticists can use a testcross to determine unknown genotypes 163 9.7 Mendel’s laws reflect the rules of probability 164 9.8 VISUALIZING THE CONCEPT Genetic traits in humans can be tracked through family pedigrees 165 9.9 Connection Many inherited traits in humans are controlled by a single gene 166 9.10 Connection New technologies can provide insight into one’s genetic legacy 168 9.4 Variations on Mendel’s Laws 170 9.11 Incomplete dominance results in intermediate phenotypes 170 9.12 Many genes have more than two alleles that may be codominant 171 9.13 A single gene may affect many phenotypic characters 172 9.14 A single character may be influenced by many genes 173 9.15 The environment affects many characters 174 The Chromosomal Basis of Inheritance 174 9.16 Chromosome behavior accounts for Mendel’s laws 174 9.17 Scientific Thinking Genes on the same chromosome tend to be inherited together 176 9.18 Crossing over produces new combinations of alleles 176 9.19 Geneticists use crossover data to map genes 178 Sex Chromosomes and Sex-Linked Genes 178 9.20 Chromosomes determine sex in many species 178 9.21 Sex-linked genes exhibit a unique pattern of inheritance 180 9.22 Connection Human sex-linked disorders affect mostly males 181 9.23 Evolution Connection The Y chromosome provides clues about human male evolution 181 Chapter Review 182 10 Molecular Biology of the Gene 184 The Structure of the Genetic Material 186 10.1 10.2 10.3 Scientific Thinking Experiments showed that DNA is the genetic material 186 DNA and RNA are polymers of nucleotides 188 DNA is a double-stranded helix 190 DNA Replication 192 10.4 10.5 DNA replication depends on specific base pairing 192 DNA replication proceeds in two directions at many sites simultaneously 192 The Flow of Genetic Information from DNA to RNA to Protein 194 10.6 10.7 10.8 10.9 10.10 10.11 10.12 10.13 10.14 10.15 10.16 Genes control phenotypic traits through the expression of proteins 194 Genetic information written in codons is translated into amino acid sequences 195 The genetic code dictates how codons are translated into amino acids 196 VISUALIZING THE CONCEPT Transcription produces genetic messages in the form of RNA 197 Eukaryotic RNA is processed before leaving the nucleus as mRNA 198 Transfer RNA molecules serve as interpreters during translation 198 Ribosomes build polypeptides 200 An initiation codon marks the start of an mRNA message 200 Elongation adds amino acids to the polypeptide chain until a stop codon terminates translation 201 Review: The flow of genetic information in the cell is DNA → RNA → protein 202 Mutations can affect genes 203 The Genetics of Viruses and Bacteria 204 10.17 Viral DNA may become part of the host chromosome 204 10.18 Connection Many viruses cause disease in animals and plants 205 10.19 Evolution Connection Emerging viruses threaten human health 206 10.20 The AIDS virus makes DNA on an RNA template 207 10.21 Prions are infectious proteins 207 10.22 Bacteria can transfer DNA in three ways 208 10.23 Bacterial plasmids can serve as carriers for gene transfer 209 Chapter Review 210 Cell signaling and waves of gene expression direct animal development 222 11.9 Connection Researchers can monitor the expression of specific genes 223 11.10 Signal transduction pathways convert messages received at the cell surface to responses within the cell 224 11.11 Evolution Connection Cell-signaling systems appeared early in the evolution of life 224 11.8 Cloning of Plants and Animals 225 11.12 Plant cloning shows that differentiated cells may retain all of their genetic potential 225 11.13 Scientific Thinking Biologists can clone animals via nuclear transplantation 226 11.14 Connection Therapeutic cloning can produce stem cells with great medical potential 227 The Genetic Basis of Cancer 228 11.15 Cancer results from mutations in genes that control cell division 228 11.16 Multiple genetic changes underlie the development of cancer 229 11.17 Faulty proteins can interfere with normal signal transduction pathways 230 11.18 Connection Lifestyle choices can reduce the risk of cancer 231 Chapter Review 232 12 DNA Technology and Genomics 234 Gene Cloning and Editing 236 12.1 Genes can be cloned in recombinant plasmids 236 12.2 VISUALIZING THE CONCEPT Enzymes are 11 How Genes Are Controlled 212 Control of Gene Expression 214 11.1 11.2 11.3 11.4 11.5 11.6 11.7 Proteins interacting with DNA turn prokaryotic genes on or off in response to environmental changes 214 Chromosome structure and chemical modifications can affect gene expression 216 Complex assemblies of proteins control eukaryotic transcription 218 Eukaryotic RNA may be spliced in more than one way 218 Later stages of gene expression are also subject to regulation 219 Noncoding RNAs play multiple roles in controlling gene expression 220 VISUALIZING THE CONCEPT Multiple mechanisms regulate gene expression in eukaryotes 221 12.3 12.4 12.5 used to “cut and paste” DNA 238 Nucleic acid probes can label specific DNA segments 239 Reverse transcriptase can help make genes for cloning 239 New techniques allow a specific gene to be edited 240 Genetically Modified Organisms 241 12.6 12.7 12.8 12.9 Recombinant cells and organisms can mass-produce gene products 241 Connection DNA technology has changed the pharmaceutical industry and medicine 242 Connection Genetically modified organisms are transforming agriculture 243 Scientific Thinking The use of genetically modified organisms raises questions and concerns 244 Detailed Contents xxix 12.10 Connection Gene therapy may someday help treat a variety of diseases 245 DNA Profiling 246 12.11 The analysis of genetic markers can produce a DNA profile 246 12.12 The PCR method is used to amplify DNA sequences 246 12.13 Gel electrophoresis sorts DNA molecules by size 247 12.14 Short tandem repeat analysis is used for DNA profiling 248 12.15 Connection DNA profiling has provided evidence in many forensic investigations 249 Genomics and Bioinformatics 250 12.16 Small segments of DNA can be sequenced directly 250 12.17 Genomics is the scientific study of whole genomes 251 12.18 Connection The Human Genome Project revealed that most of the human genome does not consist of genes 252 12.19 The whole-genome shotgun method of sequencing a genome can provide a wealth of data quickly 253 12.20 The field of bioinformatics is expanding our understanding of genomes 253 12.21 Evolution Connection Genomes hold clues to human evolution 254 Chapter Review 254 The Evolution of Populations 268 Mutation and sexual reproduction produce the genetic variation that makes evolution possible 268 13.9 Evolution occurs within populations 269 13.10 The Hardy-Weinberg equation can test whether a population is evolving 270 13.11 Connection The Hardy-Weinberg equation is useful in public health science 271 13.8 Mechanisms of Microevolution 272 13.12 Natural selection, genetic drift, and gene flow can cause microevolution 272 13.13 Natural selection is the only mechanism that consistently leads to adaptive evolution 273 13.14 VISUALIZING THE CONCEPT Natural selection can alter variation in a population in three ways 274 13.15 Sexual selection may lead to phenotypic differences between males and females 275 13.16 Evolution Connection The evolution of drugresistant microorganisms is a serious public health concern 276 13.17 Diploidy and balancing selection preserve genetic variation 276 13.18 Natural selection cannot fashion perfect organisms 277 Chapter Review 278 14 The Origin of Species 280 U nit I I I Concepts of Evolution 257 Defining Species 282 14.1 13 How Populations Evolve 258 Darwin’s Theory of Evolution 260 13.1 13.2 13.3 13.4 13.5 13.6 13.7 xxx A sea voyage helped Darwin frame his theory of evolution 260 The study of fossils provides strong evidence for evolution 262 Scientific Thinking Fossils of transitional forms support Darwin’s theory of evolution 263 Homologies provide strong evidence for evolution 264 Homologies indicate patterns of descent that can be shown on an evolutionary tree 265 Darwin proposed natural selection as the mechanism of evolution 266 Scientists can observe natural selection in action 267 Detailed Contents 14.2 14.3 The origin of species is the source of biological diversity 282 There are several ways to define a species 282 VISUALIZING THE CONCEPT Reproductive barriers keep species separate 284 Mechanisms of Speciation 286 14.4 14.5 14.6 14.7 14.8 In allopatric speciation, geographic isolation leads to speciation 286 Reproductive barriers can evolve as populations diverge 287 Sympatric speciation takes place without geographic isolation 288 Scientific Thinking Sexual selection can lead to speciation 289 Isolated islands are often showcases of speciation 290 Evolution Connection Long-term field studies document evolution in Darwin’s finches 291 14.10 Hybrid zones provide opportunities to study reproductive isolation 292 14.11 Speciation can occur rapidly or slowly 293 Chapter Review 294 14.9 15 Tracing Evolutionary History 296 Early Earth and the Origin of Life 298 15.1 Conditions on early Earth made the origin of life possible 298 15.2 Scientific Thinking 15.3 Experiments show that the abiotic synthesis of organic molecules is possible 299 Stages in the origin of the first cells probably included the formation of polymers, protocells, and self-replicating RNA 300 Major Events in the History of Life 301 15.4 15.5 15.6 The origins of single-celled and multicellular organisms and the colonization of land were key events in life’s history 301 The actual ages of rocks and fossils mark geologic time 302 The fossil record documents the history of life 302 Mechanisms of Macroevolution 304 15.7 15.8 15.9 15.10 15.11 15.12 15.13 Continental drift has played a major role in macroevolution 304 Connection Plate tectonics may imperil human life 306 Five mass extinctions have altered the course of evolution 306 Adaptive radiations have increased the diversity of life 308 Genes that control development play a major role in evolution 308 Evolution Connection Novel traits may arise in several ways 310 Evolutionary trends do not mean that evolution is goal directed 311 Phylogeny and the Tree of Life 312 15.14 Taxonomy names and classifies the diversity of life 312 15.15 Phylogenies based on homologies reflect evolutionary history 313 15.16 Shared characters are used to construct phylogenetic trees 314 15.17 An organism’s evolutionary history is documented in its genome 316 15.18 Molecular clocks help track evolutionary time 317 15.19 Constructing the tree of life is a work in progress 318 Chapter Review 319 U nit I V The Evolution of Biological Diversity 321 16 Microbial Life: Prokaryotes and Protists 322 Prokaryotes 324 Prokaryotes are diverse and widespread 324 16.2 External features contribute to the success of prokaryotes 324 16.3 Populations of prokaryotes can adapt rapidly to changes in the environment 326 16.4 Prokaryotes have unparalleled nutritional diversity 327 16.5 Connection Biofilms are complex associations of microbes 328 16.6 Connection Prokaryotes help clean up the environment 328 16.7 Bacteria and archaea are the two main branches of prokaryotic evolution 329 16.8 Archaea thrive in extreme environments—and in other habitats 330 16.9 Bacteria include a diverse assemblage of prokaryotes 330 16.10 Connection Some bacteria cause disease 332 16.11 Scientific Thinking Stomach microbiota affect health and disease 332 16.1 Protists 334 16.12 Protists are an extremely diverse assortment of eukaryotes 334 16.13 Protist diversity is organized in supergroups 335 16.14 The SAR supergroup represents the range of protist diversity 336 16.15 Connection Can algae provide a renewable source of energy? 338 16.16 Some excavates have modified mitochondria 338 16.17 Unikonts include protists that are closely related to fungi and animals 339 16.18 Archaeplastids include red algae, green algae, and land plants 340 16.19 Evolution Connection Multicellularity evolved several times in eukaryotes 341 Chapter Review 342 Detailed Contents xxxi 17 The Evolution of Plant 18.3 and Fungal Diversity 344 Plant Evolution and Diversity 346 17.1 17.2 Plants have adaptations for life on land 346 Plant diversity reflects the evolutionary history of the plant kingdom 348 Alternation of Generations and Plant Life Cycles 350 17.3 VISUALIZING THE CONCEPT Haploid and diploid generations alternate in plant life cycles 350 Seedless vascular plants dominated vast “coal forests” 352 17.5 Pollen and seeds are key adaptations for life on land 352 17.6 The flower is the centerpiece of angiosperm reproduction 354 17.7 The angiosperm plant is a sporophyte with gametophytes in its flowers 354 17.8 The structure of a fruit reflects its function in seed dispersal 356 17.9 Connection Angiosperms sustain us—and add spice to our diets 356 17.10 Evolution Connection Pollination by animals has influenced angiosperm evolution 357 17.11 Connection Plant diversity is vital to the future of the world’s food supply 358 17.4 Diversity of Fungi 359 17.12 Fungi absorb food after digesting it outside their bodies 359 17.13 Fungi produce spores in both asexual and sexual life cycles 360 17.14 Fungi are classified into five groups 360 17.15 Connection Fungi have enormous ecological benefits 362 17.16 Connection Fungi have many practical uses 362 17.17 Lichens are symbiotic associations of fungi and photosynthetic organisms 363 17.18 Scientific Thinking Mycorrhizae may have helped plants colonize land 364 17.19 Connection Parasitic fungi harm plants and animals 365 Chapter Review 366 18 The Evolution of Invertebrate Diversity 368 Animal Evolution and Diversity 370 18.1 18.2 xxxii What is an animal? 370 Animal diversification began more than half a billion years ago 371 Detailed Contents 18.4 VISUALIZING THE CONCEPT Animals can be characterized by basic features of their “body plan” 372 Body plans and molecular comparisons of animals can be used to build phylogenetic trees 373 Invertebrate Diversity 374 Sponges have a relatively simple, porous body 374 Cnidarians are radial animals with tentacles and stinging cells 375 18.7 Flatworms are the simplest bilateral animals 376 18.8 Nematodes have a body cavity and a complete digestive tract 377 18.9 Diverse molluscs are variations on a common body plan 378 18.10 Annelids are segmented worms 380 18.11 Arthropods are segmented animals with jointed appendages and an exoskeleton 382 18.12 Evolution Connection Insects are the most successful group of animals 384 18.13 Scientific Thinking The genes that build animal bodies are ancient 386 18.14 Echinoderms have spiny skin, an endoskeleton, and a water vascular system for movement 387 18.15 Our own phylum, Chordata, is distinguished by four features 388 18.16 Connection Invertebrate diversity is a valuable but threatened resource 389 Chapter Review 390 18.5 18.6 19 The Evolution of Vertebrate Diversity 392 Vertebrate Evolution and Diversity 394 19.1 19.2 19.3 19.4 19.5 19.6 19.7 19.8 Shared derived characters define the major clades of chordates 394 Hagfishes and lampreys lack hinged jaws 395 Jawed vertebrates with gills and paired fins include sharks, ray-finned fishes, and lobe-finned fishes 396 Evolution Connection New fossil discoveries are filling in the gaps of tetrapod evolution 398 Amphibians are tetrapods—vertebrates with two pairs of limbs 400 Reptiles are amniotes—tetrapods with a terrestrially adapted egg 401 Birds are feathered reptiles with adaptations for flight 402 Mammals are amniotes that have hair and produce milk 403 Primate Diversity 404 19.9 VISUALIZING THE CONCEPT Many primate characters are adaptations to life in the trees 404 19.10 The human story begins with our primate heritage 406 Hominin Evolution 407 19.11 The hominin branch of the primate tree includes species that coexisted 407 19.12 Australopiths were bipedal and had small brains 408 19.13 Larger brains mark the evolution of Homo 409 19.14 From origins in Africa, Homo sapiens spread around the world 410 19.15 Scientific Thinking New discoveries raise new questions about the history of hominins 410 19.16 Evolution Connection Human skin color reflects adaptations to varying amounts of sunlight 411 19.17 Connection Our knowledge of animal diversity is far from complete 412 Chapter Review 413 U nit V Animals: Form and Function 415 20 Unifying Concepts of Animal Structure and Function 416 Structure and Function in Animal Tissues 418 20.1 20.2 20.3 20.4 20.5 20.6 20.7 Evolution Connection An animal’s form is not the perfect design 418 Structure fits function at all levels of organization in the animal body 419 Tissues are groups of cells with a common structure and function 420 Epithelial tissue covers the body and lines its organs and cavities 420 Connective tissue binds and supports other tissues 421 Muscle tissue functions in movement 422 Nervous tissue forms a communication network 422 Organs and Organ Systems 423 Organs are made up of tissues 423 Connection Bioengineers are learning to produce organs for transplants 423 20.10 Organ systems work together to perform life’s functions 424 20.11 The integumentary system protects the body 426 20.12 Scientific Thinking Well-designed experiments help answer scientific questions 427 20.8 20.9 External Exchange and Internal Regulation 428 20.13 Structural adaptations enhance exchange with the environment 428 20.14 Animals regulate their internal environment 429 20.15 Homeostasis depends on negative feedback 430 Chapter Review 430 21 Nutrition and Digestion 432 Obtaining and Processing Food 434 21.1 21.2 21.3 Animals obtain and ingest their food in a variety of ways 434 Overview: Food processing occurs in four stages 435 Digestion occurs in specialized compartments 436 The Human Digestive System 437 21.4 21.5 21.6 21.7 21.8 21.9 21.10 21.11 21.12 21.13 The human digestive system consists of an alimentary canal and accessory organs 437 Digestion begins in the oral cavity 438 After swallowing, peristalsis moves food through the esophagus to the stomach 438 Connection The Heimlich maneuver can save lives 439 The stomach stores food and breaks it down with acid and enzymes 440 Connection Digestive ailments include acid reflux and gastric ulcers 441 The small intestine is the major organ of chemical digestion and nutrient absorption 442 The liver processes and detoxifies blood from the intestines 444 The large intestine reclaims water and compacts the feces 444 Evolution Connection Evolutionary adaptations of vertebrate digestive systems relate to diet 445 Nutrition 446 21.14 An animal’s diet must provide sufficient energy 446 21.15 An animal’s diet must supply essential nutrients 447 21.16 A proper human diet must include sufficient vitamins and minerals 448 21.17 Connection Food labels provide nutritional information 450 21.18 Connection Dietary deficiencies can have a number of causes 450 21.19 Evolution Connection The human health problem of obesity may reflect our evolutionary past 451 21.20 Scientific Thinking Scientists use a variety of methods to test weight loss claims 452 Detailed Contents xxxiii 21.21 Connection Diet can influence risk of cardiovascular disease and cancer 453 Chapter Review 454 22 Gas Exchange 456 Mechanisms of Gas Exchange 458 22.1 22.2 22.3 22.4 22.5 Gas exchange in humans involves