CAMPBELL BICON CEPTSOL & COOGNNECTIONYS E I G H T H E D I T I O N 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 Boston Columbus Indianapolis New York San Francisco Upper Saddle River Amsterdam Cape Town Dubai London Madrid Milan Munich Paris Montréal Toronto Delhi Mexico City São Paulo Sydney Hong Kong Seoul Singagore Taipei Tokyo A01_REEC5325_08_SE_FM_NASTA.indd 1 11/6/13 7:47 AM Editor-in-Chief: Development Artists: Cover Photo Credit: Beth Wilbur Kelly Murphy; Andrew Recher, Andy Rouse/Nature Picture Library Executive Director of Precision Graphics Credits and acknowledgments for materials borrowed from Development: Senior Photo Editor: other sources and reproduced, with permission, in this Deborah Gale Donna Kalal textbook appear on the appropriate page within the text or Acquisitions Editor: Photo Researcher: on p. A-26. 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Manufacturing Buyer: Production Management and MasteringBiology® and BioFlix® are registered trademarks, in the U.S. Jeffrey Sargent Composition: and/or other countries, of Pearson Education, Inc. or its affiliates. S4Carlisle Publishing Services Cover Printer: Library of Congress Cataloging-in-Publication Data Design Manager: Lehigh-Phoenix Marilyn Perry Text Printer: Reece, Jane B. Cover and Interior Designer: Courier/Kendallville Campbell biology: concepts and connections / Jane B. Reece [and four Hespenheide Design others].—Eighth edition. pages cm Illustrations: Previous edition: Campbell biology: concepts & connections, 2012. Precision Graphics ISBN 978-0-321-88532-6 1. Biology. I. Title. II. Title: Biology. QH308.2.B56448 2013 570—dc23 2013024409 ISBN 10: 0-13-348039-9 (High School Binding) ISBN 13: 978-0-13-348039-9 (High School Binding) 1 2 3 4 5 6 7 8 9 10—CRK—18 17 16 15 14 PearsonSchool.com/Advanced A01_REEC5325_08_SE_FM_NASTA.indd 2 11/6/13 7:47 AM About the Authors Jane B. Reece has worked in biology Jean L. Dickey is Professor Emerita of publishing since 1978, when she joined Biological Sciences at Clemson Univer- the editorial staff of Benjamin Cum- sity (Clemson, South Carolina). After mings. Her education includes an A.B. receiving her B.S. in biology from Kent in biology from Harvard University, State University, she went on to earn a an M.S. in microbiology from Rutgers Ph.D. in ecology and evolution from University, and a Ph.D. in bacteriol- Purdue University. In 1984, Dr. Dickey ogy from the University of California, joined the faculty at Clemson, where Berkeley. At UC Berkeley, and later as a she devoted her career to teaching biol- postdoctoral fellow in genetics at Stan- ogy to nonscience majors in a variety ford University, her research focused on genetic recombination of courses. In addition to creating content-based instructional in bacteria. Dr. Reece taught biology at Middlesex County materials, she developed many activities to engage lecture and College (New Jersey) and Queensborough Community Col- laboratory students in discussion, critical thinking, and writ- lege (New York). During her 12 years as an editor at Benjamin ing, and implemented an investigative laboratory curriculum Cummings, she played a major role in a number of successful in general biology. Dr. Dickey is author of Laboratory Investiga- textbooks. She is coauthor of Campbell Biology, Tenth Edition, tions for Biology, Second Edition, and coauthor of Campbell Campbell Biology in Focus, Campbell Essential Biology, and Essential Biology, Fifth Edition, and Campbell Essential Biology Campbell Essential Biology with Physiology, Fourth Edition. with Physiology, Fourth Edition. Martha R. Taylor has been teach- Kelly Hogan is a faculty member ing biology for more than 35 years. in the Department of Biology at the She earned her B.A. in biology from University of North Carolina at Chapel Gettysburg College and her M.S. and Hill, teaching introductory biology and Ph.D. in science education from Cornell introductory genetics to science majors. University. At Cornell, she has served Dr. Hogan teaches hundreds of students as assistant director of the Office of at a time, using active-learning methods Instructional Support and has taught that incorporate technology such as cell introductory biology for both majors phones as clickers, online homework, and nonmajors. Most recently, she was a and peer evaluation tools. Dr. Hogan lecturer in the Learning Strategies Center, teaching supplemen- received her B.S. in biology at the College of New Jersey and tal biology courses. Her experience working with students in her Ph.D. in pathology at the University of North Carolina, classrooms, in laboratories, and with tutorials has increased her Chapel Hill. Her research interests relate to how large classes commitment to helping students create their own knowledge of can be more inclusive through evidence-based teaching meth- and appreciation for biology. She has been the author of the Stu- ods and technology. She provides faculty development to other dent Study Guide for all ten editions of Campbell Biology. instructors through peer-coaching, workshops, and mentor- ing. Dr. Hogan is the author of Stem Cells and Cloning, Second Eric J. Simon is a professor in the De- Edition, and is lead moderator of the Instructor Exchange, a partment of Biology and Health Science site within MasteringBiology® for instructors to exchange class- at New England College (Henniker, room materials and ideas. New Hampshire). He teaches introduc- tory biology to science majors and non- Neil A. Campbell (1946–2004) science majors, as well as upper-level combined the inquiring nature of a re- courses in tropical marine biology and search scientist with the soul of a caring careers in science. Dr. Simon received teacher. Over his 30 years of teaching a B.A. in biology and computer science introductory biology to both science and an M.A. in biology from Wesleyan majors and nonscience majors, many University, and a Ph.D. in biochemistry from Harvard Univer- thousands of students had the oppor- sity. His research focuses on innovative ways to use technology tunity to learn from him and be stimu- to improve teaching and learning in the science classroom, par- lated by his enthusiasm for the study of ticularly for nonscience majors. Dr. Simon is the lead author of life. While he is greatly missed by his the introductory nonmajors biology textbooks Campbell Essen- many friends in the biology community, his coauthors remain tial Biology, Fifth Edition, and Campbell Essential Biology with inspired by his visionary dedication to education and are com- Physiology, Fourth Edition, and the author of the introductory mitted to searching for ever better ways to engage students in biology textbook Biology: The Core. the wonders of biology. About the Authors iii A01_REEC5325_08_SE_FM_NASTA.indd 3 11/6/13 7:47 AM To the Student: How to use this book and MasteringBiology® BIG IDEA S Cloning Big Ideas help you connect the Gene Make (12.1–12.5) ◃ tory A variety of labora overarching concepts that are o important techniques can be used t copy and combine DNA explored in the chapter. molecules. connections R E chnology between DNA Te ▹ ii were hacked big island of Hawa HAPT on the omics ds of papaya trees t against d en thousan s a protes cally Modie span, sumably a Geneti C d G pre biological an darkness, der the cover of nal behavior, isms C. ey are down un condone such crimi Organ oaded with vitamin though few would ps? is 12.10) elow, are sweet and l l climates. GMO crops. Al safety of GMO cro (12.6– 2 the photograph b ows only in tropica ncerned about the . Connection modules in every chapter 1 fruit, shown in apaya) that gr e in fact be co disagreement ts, concepts and apaya plant (Carica p should w ble debate and ells, plan ▿ y growing treelike op.