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Brief contents 1 Evolution, the Themes of Biology, and Scientific 28 Protists 591 Inquiry 2 29 Plant Diversity I: How Plants Colonized Land 616 30 Plant Diversity II: The Evolution of Seed Unit 1 thE chEmistry oF liFE 27 Plants 634 31 Fungi 652 2 The Chemical Context of Life 28 32 An Overview of Animal Diversity 671 3 Water and Life 44 33 An Introduction to Invertebrates 684 4 Carbon and the Molecular Diversity of Life 56 34 The Origin and Evolution of Vertebrates 716 5 The Structure and Function of Large Biological Molecules 66 Unit 6 Plant Form and FUnction 755 Unit 2 thE cEll 92 35 Vascular Plant Structure, Growth, and Development 756 6 A Tour of the Cell 93 36 Resource Acquisition and Transport in Vascular 7 Membrane Structure and Function 126 Plants 782 8 An Introduction to Metabolism 143 37 Soil and Plant Nutrition 803 9 Cellular Respiration and Fermentation 164 38 Angiosperm Reproduction and Biotechnology 820 10 Photosynthesis 187 39 Plant Responses to Internal and External 11 Cell Communication 212 Signals 840 12 The Cell Cycle 234 Unit 7 animal Form and FUnction 870 Unit 3 gEnEtics 253 40 Basic Principles of Animal Form and Function 871 13 Meiosis and Sexual Life Cycles 254 41 Animal Nutrition 896 14 Mendel and the Gene Idea 269 42 Circulation and Gas Exchange 919 15 The Chromosomal Basis of Inheritance 294 43 The Immune System 950 16 The Molecular Basis of Inheritance 314 44 Osmoregulation and Excretion 975 17 Gene Expression: From Gene to Protein 335 45 Hormones and the Endocrine System 997 18 Regulation of Gene Expression 363 46 Animal Reproduction 1017 19 Viruses 396 47 Animal Development 1041 20 DNA Tools and Biotechnology 413 48 Neurons, Synapses, and Signaling 1065 21 Genomes and Their Evolution 440 49 Nervous Systems 1083 50 Sensory and Motor Mechanisms 1105 Unit 4 mEchanisms oF EvolUtion 465 51 Animal Behavior 1137 22 Descent with Modification: A Darwinian View of Life 466 Unit 8 Ecology 1161 23 The Evolution of Populations 484 52 An Introduction to Ecology and the Biosphere 1162 24 The Origin of Species 504 53 Population Ecology 1188 25 The History of Life on Earth 523 54 Community Ecology 1212 55 Ecosystems and Restoration Ecology 1236 Unit 5 thE EvolUtionary history 56 Conservation Biology and Global Change 1258 oF Biological divErsity 550 26 Phylogeny and the Tree of Life 551 27 Bacteria and Archaea 571 C a m p b e l l B i o l o G Y ElEvEnth Edition Lisa A. Urry Michael L. Cain Steven A. Wasserman Mills College, oakland, Bowdoin College, BrunswiCk, university of California, California Maine san diego Peter V. Minorsky Jane B. Reece MerCy College, doBBs ferry, Berkeley, California new york 330 Hudson Street, New York, NY 10013 Courseware Portfolio Management Director: Beth Wilbur Art/Text/Photo Permissions Specialist: Courseware Portfolio Management Specialist: Josh Frost Integra Software Services, Inc. Courseware Director, Content Development: Senior Content Developer, MasteringBiology™: Ginnie Simione Jutson Sarah Jensen Supervising Editors: Beth N. 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Field Marketing Manager: Kelly Galli Illustrations: Lachina Manufacturing Buyer: Stacey Weinberger Design Manager: Marilyn Perry Text & Cover Printer: Courier/Kendallville Cover and Text Design: Elise Lansdon Cover Photo: Helianthus annuus (cultivated sunflower) Rights & Permissions Manager: Ben Ferrini Cover Photo Credit: © Radius Images/Getty Images Photo Researcher: Maureen Spuhler Copyright © 2017, 2014, 2011 Pearson Education, Inc. All Rights Reserved. Printed in the United States of America. 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/. Readers may view, browse, and/or download material for temporary copying purposes only, provided these uses are for noncommercial personal purposes. Except as provided by law, this material may not be further reproduced, distributed, transmitted, modified, adapted, performed, displayed, published, or sold in whole or in part, without prior written permission from the publisher. Acknowledgments of third-party content appear on page CR-1, which constitutes an extension of this copyright page. PEARSON, ALWAYS LEARNING, MasteringBiology™ and BioFlix® are exclusive trademarks owned by Pearson Education, Inc. or its affiliates in the U.S. and/or other countries. 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., author. | Urry, Lisa A., author. | Campbell, Neil A., 1946-2004, author. Title: Campbell biology / Lisa Urry, Michael Cain, Steven Wasserman, Peter Minorsky, Jane Reece. Description: Eleventh edition. | Hoboken : Pearson Higher Education, 2016. | Previous edition: Campbell biology / Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson. Tenth edition. 2014. | 1st edition: Biology by Neil A. Campbell, 1987. Identifiers: LCCN 2016025573 Subjects: LCSH: Biology. Classification: LCC QH308.2 .C34 2016 | DDC 570—dc23 LC record available at https://lccn.loc.gov/2016025573 1 16 ISBN 10: 0-134-09341-0; ISBN 13: 978-0-134-09341-3 (Student Edition) ISBN 10: 0-134-15412-6; ISBN 13: 978-0-134-15412-1 (Books a la Carte Edition) About the Authors Lisa A. Urry is Professor of Biology and Chair of the Biology Department at Mills College. After earning a B.A. at Tufts University, she completed her Ph.D. at the Massachusetts Institute of Technology (MIT). Lisa has conducted research on gene expression during embryonic and larval development in sea urchins. Deeply committed to promoting opportunities in science for women and underrepresented minorities, she has taught courses ranging from introductory and developmental biology to a nonmajors course called Evolution for Future Presidents. Lisa is a coauthor of Campbell Biology in Focus. Michael L. Cain is an ecologist and evolutionary biologist who is now writing full-time. Michael earned an A.B. from Bowdoin College, an M.Sc. from Brown University, and a Ph.D. from Cornell University. As a faculty member at New Mexico State University, he taught introductory biology, ecology, evolution, botany, and conservation biology. Michael is the author of dozens of scientific papers on topics that include foraging behavior in insects and plants, long-distance seed dispersal, and speciation in crickets. He is a coauthor of Campbell Biology in Focus and of an ecology textbook. Steven A. Wasserman is Professor of Biology at the University of California, San Diego (UCSD). He earned an A.B. from Harvard University and a Ph.D. from MIT. Working on the fruit fly Drosophila, Steve has done research on developmental biology, reproduction, and immunity. Having taught genetics, development, and physiology to undergraduate, graduate, and medical students, he now focuses on introductory biology, for which he has been honored with UCSD’s Distinguished Teaching Award. He is a coauthor of Campbell Biology in Focus. Peter V. Minorsky is Professor of Biology at Mercy College in New York, where he teaches introductory biology, ecology, and botany. He received his A.B. from Vassar College and his Ph.D. from Cornell University. Peter taught at Kenyon College, Union College, Western Connecticut State University, and Vassar College; he is also the science writer for the journal Plant Physiology. His research interests concern how plants sense environmental change. Peter received the 2008 Award for Teaching Excellence at Mercy College and is a coauthor of Campbell Biology in Focus. Jane B. Reece, the head of the author team for Editions 8–10 of Campbell BIOLOGY, was Neil Campbell’s longtime collaborator. Jane taught biology at Middlesex County College and Queensborough Community College. She holds an A.B. from Harvard University, an M.S. from Rutgers University, and a Ph.D. from the University of California, Berkeley. Jane’s research as a doctoral student at UC Berkeley and postdoctoral fellow at Stanford University focused on genetic recombination in bacteria. Besides her work on Campbell BIOLOGY, Jane has been a coauthor on all the Campbell texts. Neil A. Campbell (1946–2004) earned his M.A. from the University of California, Los Angeles, and his Ph.D. from the University of California, Riverside. His research focused on desert and coastal plants. Neil’s 30 years of teaching included introductory biology courses at Cornell University, Pomona College, and San Bernardino Valley College, where he received the college’s first Outstanding Professor Award in 1986. For many years he was also a visiting scholar at UC Riverside. Neil was the founding author of Campbell BIOLOGY. aBout tHe autHors iii Preface We are honored to present the more extensive “Solve It” investigations Eleventh Edition of Campbell to further explore a given topic. BIOLOGY. For the last three decades, Ready-to-Go Teaching Modules Campbell BIOLOGY has been the leading on key topics provide instructors with college text in the biological sciences. It assignments to use before and after has been translated into 19 languages and class, as well as in-class activities that has provided millions of students with a use clickers or Learning Catalytics™ solid foundation in college-level biology. for assessment. This success is a testament not only to Neil Integrated text and media: Media Campbell’s original vision but also to the references in the printed book direct dedication of hundreds of reviewers (listed students to the wealth of online on pages xxviii–xxxi), who, together with self-study resources available to editors, artists, and contributors, have them in the Study Area section of shaped and inspired this work. MasteringBiology. In eText 2.0 Our goals for the Eleventh Edition include: (which will be available for Fall 2017 classes), these resources will also be integrated directly into the eText. The new online increasing visual literacy through new figures, questions, learning tools include: and exercises that build students’ skills in understanding and Get Ready for This Chapter questions provide a creating visual representations of biological structures and quick check of student understanding of the background processes information needed to learn a new chapter’s content, with asking students to practice scientific skills by applying feedback to bolster their preparation. scientific skills to real-world problems Figure Walkthroughs guide students through key supporting instructors by providing teaching modules figures with narrated explanations, figure markups, and with tools and materials for introducing, teaching, and questions that reinforce important points. Additional assessing important and often challenging topics questions can be assigned in MasteringBiology. integrating text and media to engage, guide, and inform More than 450 animations and videos bring students in an active process of inquiry and learning biology to life. These include resources from HHMI Our starting point, as always, is our commitment to craft- BioInteractive that engage students in topics from ing text and visuals that are accurate, are current, and reflect the discovery of the double helix to evolution. our passion for teaching biology. QR codes and URLs within the Chapter Review provide easy access to Vocabulary Self-Quizzes and Practice New to This Edition Tests for each chapter that can be used on smartphones, tablets, and computers. Here we provide an overview of the new features that we have Interviews from the First Edition through the Eleventh developed for the Eleventh Edition; we invite you to explore Edition of Campbell BIOLOGY are referenced in the pages ix–xxvi for more information and examples. chapter where they are most relevant. The interviews show Visualizing Figures and Visual Skills Questions students the human side of science by featuring diverse give students practice in interpreting and creating visual scientists talking about how they became interested in representations in biology. The Visualizing Figures have what they study, how they began, and what inspires them. embedded questions that guide students in exploring The impact of climate change at all levels of the biological how diagrams, photographs, and models represent and hierarchy is explored throughout the text, starting with a reflect biological systems and processes. Assignable new figure (Figure 1.12) and discussion in Chapter 1 and questions are also available in MasteringBiology to concluding with a new Make Connections Figure (Figure give students practice with the visual skills addressed in 56.30) and expanded coverage on causes and effects of the figures. climate change in Chapter 56. Problem-Solving Exercises challenge students to apply As in each new edition of Campbell BIOLOGY, the Eleventh scientific skills and interpret data in solving real-world Edition incorporates new content and pedagogical problems. These exercises are designed to engage students improvements. These are summarized on pp. vi–viii, through compelling case studies and provide practice following this Preface. Content updates reflect rapid, ongoing with data analysis skills. Problem-Solving Exercises have changes in technology and knowledge in the fields of assignable versions in MasteringBiology. Some also have genomics, gene editing technology (CRISPR), evolutionary iv PrefaCe biology, microbiology, and more. In addition, significant and the unit-opening interviews, our standard-setting Inquiry revisions to Unit 8, Ecology, improve the conceptual Figures deepen the ability of students to understand how we framework for core ecological topics (such as population know what we know. Scientific Inquiry Questions give stu- growth, species interactions, and community dynamics) dents opportunities to practice scientific thinking, along with and more deeply integrate evolutionary principles. the Problem-Solving Exercises, Scientific Skills Exercises, and Interpret the Data Questions. Together, these activities pro- Our Hallmark Features vide students practice in both applying the process of science and using quantitative reasoning, addressing additional core Teachers of general biology face a daunting challenge: to help competencies outlined in Vision and Change. students acquire a conceptual framework for organizing an ever- expanding amount of information. The hallmark features of Campbell BIOLOGY provide such a framework, while promoting a deeper understanding of biology and the process of science. MasteringBiology, the most widely used online assessment As such, they are well aligned with the core competencies out- and tutorial program for biology, provides an extensive library of lined by the 2009 Vision and Change national conference. homework assignments that are graded automatically. In addi- Furthermore, the core concepts defined by Vision and Change tion to the new Get Ready for This Chapter Questions, have close parallels in the unifying themes that are introduced Figure Walkthroughs, Problem-Solving Exercises, and in Chapter 1 and integrated throughout the book. Chief among Visualizing Figures, MasteringBiology offers Dynamic Study the themes of both Vision and Change and Campbell BIOLOGY is Modules, Adaptive Follow-Up Assignments, Scientific Skills Exer- evolution. Each chapter of this text includes at least one Evolu- cises, Interpret the Data Questions, Solve It Tutorials, HHMI Bio- tion section that explicitly focuses on evolutionary aspects of Interactive Short Films, BioFlix® Tutorials with 3-D Animations, the chapter material, and each chapter ends with an Evolution Experimental Inquiry Tutorials, Interpreting Data Tutorials, Connection Question and a Write About a Theme Question. BLAST Tutorials, Make Connections Tutorials, Video Field Trips, To help students distinguish the “forest from the trees,” Video Tutor Sessions, Get Ready for Biology, Activities, Reading each chapter is organized around a framework of three to seven Quiz Questions, Student Misconception Questions, 4,500 Test carefully chosen Key Concepts. The text, Concept Check Bank Questions, and MasteringBiology Virtual Labs. Mastering- Questions, Summary of Key Concepts, and MasteringBiology Biology also includes the Campbell BIOLOGY eText, Study Area, resources all reinforce these main ideas and essential facts. Instructor Resources, and Ready-to-Go Teaching Modules. See Because text and illustrations are equally important for learn- pages ix–xxiii and www.masteringbiology.com for more details. ing biology, integration of text and figures has been a hallmark of this text since the First Edition. In addition to the Our Partnership with Instructors new Visualizing Figures, our popular Exploring Figures and Make and Students Connections Figures epitomize this approach. Each Exploring Figure is a learning unit of core content that brings together re- A core value underlying our work is our belief in the impor- lated illustrations and text. Make Connections Figures reinforce tance of a partnership with instructors and students. One fundamental conceptual connections throughout biology, help- primary way of serving instructors and students, of course, is ing students overcome tendencies to compartmentalize informa- providing a text that teaches biology well. In addition, Pear- tion. The Eleventh Edition features two new Make Connections son offers a rich variety of instructor and student resources, in Figures. There are also Guided Tour Figures that walk students both print and electronic form (see pp. ix–xxiii). In our con- through complex figures as an instructor would. tinuing efforts to improve the book and its supplements, we To encourage active reading of the text, Campbell BIOLOGY benefit tremendously from instructor and student feedback, includes numerous opportunities for students to stop and think not only in formal reviews from hundreds of scientists, but about what they are reading, often by putting pencil to paper to also via e-mail and other avenues of informal communication. draw a sketch, annotate a figure, or graph data. Active reading The real test of any textbook is how well it helps instructors questions include Visual Skills Questions, Draw It Questions, teach and students learn. We welcome comments from both Make Connections Questions, What If? Questions, Figure Leg- students and instructors. Please address your suggestions to: end Questions, Summary Questions, Synthesize Your Knowl- Lisa Urry (Chapter 1 and Units 1–3) edge Questions, and Interpret the Data Questions. Answering [email protected] these questions requires students to write or draw as well as Michael Cain (Units 4, 5, and 8) think and thus helps develop the core competency of communi- [email protected] cating science. Peter Minorsky (Unit 6) Finally, Campbell BIOLOGY has always featured scientific [email protected] inquiry, an essential component of any biology course. Com- Steven Wasserman (Unit 7) plementing stories of scientific discovery in the text narrative [email protected] PrefaCe v Highlights of New Content This section highlights selected new content and pedagogical where in the three life cycles haploid cells undergo mitosis, and changes in Campbell BIOLOGY, Eleventh Edition. what type of cells are formed. Chapter 14 includes new informa- tion from a 2014 genomic study on the number of genes and genetic variants contributing to height. Figure 14.15b now uses CHAPTER 1 Evolution, the Themes of Biology, “inability to taste PTC” rather than “attached earlobe.” Chapters and Scientific Inquiry 14 and 15 are more inclusive, clarifying the meaning of the term Chapter 1 opens with a new introduction to a case study “normal” in genetics and explaining that sex is no longer thought on the evolution of coloration in mice. New text and a new to be simply binary. Other updates in Chapter 15 include new photo (Figure 1.12) relate climate change to species survival. research in sex determination and a technique being developed to avoid passing on mitochondrial diseases. New Visualizing UNIT 1 THE CHEMISTRy OF LIFE Figure 16.7 shows students various ways that DNA is illustrated. Chapter 17 has a new opening photo and story about albino don- In Unit 1, new content engages students in learning this founda- keys to pique student interest in gene expression. To help students tional material. The opening of Chapter 3 and new Figure 3.7 understand the Beadle and Tatum experiment, new Figure 17.2 show organisms affected by loss of Arctic sea ice. Chapter 5 explains how they obtained nutritional mutants. A new Problem- has updates on lactose intolerance, trans fats, the effects of Solving Exercise asks students to identify mutations in the insulin diet on blood cho- gene and predict their effect on the protein. lesterol, protein Figure 3.7 Effects of climate change Chapters 18–21 are extensively updated, driven by exciting sequences and on the Arctic. new discoveries based on DNA sequencing and gene-editing tech- structures, and nology. Chapter 18 has updates on histone modifications, nuclear intrinsically disor- location and the persistence of transcription factories, chromatin dered proteins. remodeling by ncRNAs, long noncoding RNAs (lncRNAs), the New Visualizing role of master regulatory genes in modifying chromatin struc- Figure 5.16 helps ture, and the possible role of p53 in the low incidence of cancer students understand in elephants. Make Connections Figure 18.27, “Genomics, Cell various ways pro- Signaling, and Cancer,” has teins are depicted. been expanded to include Figure 20.14 Gene editing A new Problem- more information on cell using the CRISPR-Cas9 system. Solving Exercise signaling. Chapter 19 fea- engages students by tures a new section that having them com- covers bacterial defenses pare DNA sequences against bacteriophages in a case of possible and describes the CRISPR- fish fraud. Cas9 system (Figure 19.7); updates include the Ebola, UNIT 2 THE CELL Chikungunya, and Zika viruses (Figure 19.10) and Our main goal for this unit was to make the material more acces- discovery of the largest sible and inviting to students. New Visualizing Figure 6.32 shows virus known to date. A the profusion of molecules and structures in a cell, all drawn to discussion has been added scale. In Chapter 7, a new figure illustrates levels of LDL receptors in of mosquito transmission people with and without familial hypercholesterolemia. Chapter 8 of diseases and concerns includes a beautiful new photo of a geyser with thermophilic bac- about the effects of global teria in Figure 8.17, bringing to life the graphs of optimal tempera- climate change on disease tures for enzyme function. Chapter 10 discusses current research transmission. Chapter 20 trying to genetically modify rice (a C crop) so that it is capable has a new photo of next- 3 of carrying out C photosynthesis to increase yields. Chapter 11 generation DNA sequenc- 4 includes a new Problem-Solving Exercise that guides students ing machines (Figure 20.2) through assessing possible new treatments for bacterial infections and a new illustration of by blocking quorum sensing. In Chapter 12, the mechanism of the widely used technique chromosome movement in bacteria has been updated and more of RNA sequencing cell cycle control checkpoints have been added, including one pro- (Figure 20.13). A new sec- posed by researchers in 2014. tion titled Editing Genes and Genomes has been added describing the UNIT 3 GENETICS CRISPR-Cas9 system In Chapters 13–17, we have incorporated changes that help (Figure 20.14) that students grasp the more abstract concepts of genetics and their has been developed to chromosomal and molecular underpinnings. For example, a new edit genes in living cells. Visual Skills Question with Figure 13.6 asks students to identify Information has also been vi HigHligHts of new Content added later in the chapter on use of the CRISPR-Cas9 system, UNIT 5 THE EVOLUTIONARy HISTORy including a study in which a genetic mutation for the disease tyrosinemia was corrected in mice. Finally, the discussion of ethi- OF BIOLOGICAL DIVERSITy cal considerations has been updated to include a recent report of In keeping with our goal of improving how students interpret scientists using the CRISPR-Cas9 system to edit a gene in human and create visual representations in biology, we have added a new embryos, along with a discussion of the ethical questions raised figure (Figure 26.5, “Visualizing Phylogenetic Relationships”) by such experiments, such as its usage in the gene drive approach that introduces the visual conventions used in phylogenetic to combat carrying of diseases by mosquitoes. In Chapter 21, in trees and helps students understand what such trees do and addition to the usual updates of sequence-related data (speed of don’t convey. Students are also provided many opportunities to sequencing, number of species’ genomes sequenced, etc.), there practice their visual skills, with more than ten new Visual Skills are several research updates, including some early results from Questions on topics ranging from interpreting phylogenetic the new Roadmap Epigenomics Project and results from a 2015 trees to predicting which regions of a bacterial flagellum are study focusing on 414 important yeast genes. hydrophobic. The unit also contains new content on tree think- ing, emphasizing such key points as how sister groups provide a UNIT 4 MECHANISMS OF EVOLUTION clear way to describe evolutionary relationships and how trees do not show a “direction” in evolution. Other major content A major goal for this revision was to strengthen how we help changes include new text in Concepts 26.6, 27.4, and 28.1 on students understand and interpret visual representations of evolu- the 2015 discovery of the Lokiarchaeota, a group of archaea that tionary data and concepts. Toward this end, we have added a new may represent the sister group of the eukaryotes, new text and figure (Figure 25.8), “Visualizing the Scale of Geologic Time,” and a new figure (Figure 26.22) on horizontal gene transfer from a new figure (Figure 23.12) on gene flow. Several figures have prokaryotes to eukaryotes, new text in Concept 27.6 describ- been revised to improve the presentation of data, including Figure ing the CRISPR-Cas9 system and a new figure (Figure 27.21) that 24.6 (on reproductive isolation in mosquitofish), Figure 24.10 (on illustrates one example of how CRISPR-Cas 9 technology has allopolyploid speciation), and Figure 25.25 (on the origin of the opened new avenues of research on HIV, and new material in insect body plan). The Figure 23.12 Gene flow and local Concept 29.3 describing how early forests contributed to global unit also features new adaptation in the Lake Erie water climate change (in this case, global cooling). A new Problem- material that connects snake (Nerodia sipedon). Solving Exercise in Chapter 34 engages students in interpret- evolutionary concepts ing data from a study investigating whether frogs can acquire and societal problems. resistance to a fungal pathogen through controlled exposure to Examples include text it. Other updates include the revision of many phylogenies to in Chapter 22 on the reflect recent phylogenomic data, new chapter-opening stories in 2015 discovery of teix- Chapter 31 (on how mycorrhizae link trees of different species) obactin, an antibiotic and Chapter 33 (on the “blue dragon,” a mollusc that preys on the that is effective against highly toxic Portuguese man-of-war), new text and a new figure some hard-to-treat patho- (Figure 34.37) on the adaptations of the kangaroo rat to its arid envi- gens, a new discussion in ronment, and new material in Concept 34.7, including a new figure Chap ter 24 on the impact (Figure 34.52) describing fossil and DNA evidence indicating that of climate change on humans and Neander- hybrid zones, and a new thals interbred, produc- Figure 34.53 Fossils of hand and Problem-Solving Exercise ing viable offspring. The foot bones of Homo naledi. in Chapter 24 on how discussion of human hybridization may have evolution also includes led to the spread of insec- new text and a new figure ticide resistance genes in (Figure 34.53) on Homo mosquitoes that transmit naledi, the most recently malaria. The unit also discovered member of includes new chapter- the human evolutionary opening stories in lineage. Chapter 22 (on a moth whose features illustrate the concepts of unity, diversity, and adaptation) and Chapter 25 (on the discovery of whale bones UNIT 6 PLANT FORM AND FUNCTION in the Sahara Desert). Additional changes include new text in A major aim in revising Chapter 35 was to help students better Concept 22.3 emphasizing how populations can evolve over understand how primary and secondary growth are related. short periods of time, a new table (Table 23.1) highlighting the New Visualizing Figure 35.11 enables students to picture growth five conditions required for a population to be in Hardy-Weinberg at the cellular level. Also, the terms protoderm, procambium, equilibrium, and new material in Concept 25.1 describing how and ground meristem have been introduced to underscore the researchers recently succeeded for the first time in constructing transition of meristematic to mature tissues. A new flowchart a “protocell” in which replication of a template strand of RNA (Figure 35.24) summarizes growth in a woody shoot. New could occur. text and a figure (Figure 35.26) focus on genome analysis of Arabidopsis ecotypes, relating plant morphology to ecol- ogy and evolution. In Chapter 36, new Figure 36.8 illus- trates the fine branching of leaf veins, and information on HigHligHts of new Content vii phloem-xylem water transfer has been updated. New Make UNIT 8 ECOLOGy Connections Figure 37.10 highlights mutualism across king- doms and domains. Figure 37.13 and the related text include The Ecology Unit has been extensively revised for the Eleventh new findings on how some soil nitrogen derives from weather- Edition. We have reorganized and improved the conceptual ing of rocks. New Figure 38.3 clarifies how the terms carpel and framework with which students are introduced to the follow- pistil are related. The text on flower structure and the angio- ing core ecological topics: life tables, per capita population sperm life cycle figure identify carpels as megasporophylls growth, intrinsic rate of increase (“r ”), exponential popula- and stamens as microsporophylls, correlating with the plant tion growth, logistic population growth, density dependence, evolution discussion in Unit 5. In Concept 38.3, the current species interactions (in particular, parasitism, commensalism, problem of glyphosate-resistant crops is discussed in detail. A and mutualism), and MacArthur and Wilson’s island bioge- revised Figure 39.7 helps students visualize how cells elongate. ography model. The revision also includes a deeper integra- Figure 39.8 now addresses apical dominance in a Guided Tour tion of evolutionary principles, including a new Key Concept format. Information about the role of sugars in controlling api- (52.5) and two new figures (Figures 52.22 and 52.23) on the cal dominance has been added. In Concept 39.4, a new Prob- reciprocal effects of ecology and evolution, new material in lem-Solving Exercise highlights how global climate change Concept 52.4 on how the geographic distributions of spe- affects crop productivity. Figure 39.26 on defense responses cies are shaped by a combination of evolutionary history and against pathogens has been simplified and improved. ecological factors, and five new Make Connections Questions that ask students to examine how ecological and evolutionary UNIT 7 ANIMAL FORM AND FUNCTION mechanisms interact. In keeping with our goal of expanding and strengthening our coverage of climate change, we have A major goal of the Unit 7 revision was to transform how stu- added a new discussion and a new figure (Figure 52.20) on dents interact with and learn from representations of anatomy how climate change has affected the distribution of a key- and physiology. For example, gastrulation is now introduced stone species, a new Figure 55.8 Climate change, with a Visualizing Figure (Figure 47.8) that provides a clear and section of text in wildfires, and insect outbreaks. carefully paced introduction to three-dimensional processes Concept 55.2 on that may be difficult for students to grasp. In addition, a num- how climate change ber of the new and revised figures help students explore spatial affects NPP, a new relationships in anatomical contexts, such as the interplay of figure (Figure 55.8) lymphatic and cardiovascular circulation (Figure 42.15) and on how climate the relationship of the limbic system to overall brain structure change has caused (Figure 49.14). A new Problem-Solving Exercise in Chapter 45 an increase in wild- taps into student interest in medical mysteries through a case fires and insect study that explores the science behind laboratory testing outbreaks, a new and diagnosis. Content updates help students appreciate the Problem-Solving continued evolution of our understanding of even familiar Exercise in Chapter phenomena, such as the sensation of thirst (Concept 44.4) 55 that explores and the locomotion of kangaroos and jellies (Concept 50.6). how insect out- Furthermore, new text and figures introduce students to breaks induced by cutting-edge technology relating to such topics as RNA-based climate change can antiviral defense in invertebrates (Figure 43.4) and rapid, com- cause an ecosystem prehensive characterization of viral exposure (Figure 43.24), as to switch from a car- well as recent discoveries regarding brown fat in adult humans bon sink to a carbon (Figure 40.16), the microbiome (Figure 41.17), partheno- source, a new figure genesis (Concept 46.1), and magnetoreception (Concept 50.1). (Figure 56.29) on As always, there is fine-tuning of pedagogy, as in discussions of the greenhouse effect, new text in Concept 56.4 on biologi- the complementary roles of inactivation and voltage gating of cal effects of climate change, and a new Make Connections ion channels during action potential formation (Concept 48.3) Figure (Figure 56.30) on how climate change affects all levels and of the experimental characterization of genetic determi- of biological organization. Additional updates include a new nants in bird migration (Figure 51.24). figure (Figure 53.25) on per capita ecological footprints, a new chapter-opening story in Chapter 54 on a seemingly unlikely mutualism between a shrimp and a much larger predatory Figure 41.17 Variation in human gut microbiome at fish, new text in Concept 54.1 emphasizing that each partner different life stages. in a mutualism experiences both benefits and costs, new text in Concept 54.1 describing how the outcome of an eco- logical interaction can change over time, two new figures (Figures 54.29 and 54.30) on the island equilibrium model, a new figure (Figure 54.31) documenting two shrew species as unexpected hosts of the Lyme disease, new text in Concept 56.1 comparing extinction rates today with those typically seen in the fossil record, and a new discussion and figure (Figure 56.22) on the restoration of a degraded urban stream. viii HigHligHts of new Content

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