Campbell Biology: Concepts and Connections / Jane B

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

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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. Alison Rodal Kristin Piljay Executive Editorial Manager: Photo Permissions Ginnie Simione Jutson Management: Editorial Project Manager: Bill Smith Group Debbie Hardin Director of Editorial Content Development Editors: MasteringBiology®: Debbie Hardin, Susan Teahan Tania Mlawer Editorial Assistant: Development Editor, Libby Reiser MasteringBiology®: Juliana Tringali Senior Supplements Project Copyright © 2015, 2012, 2009 Pearson Education, Inc. All rights Editor: Senior Mastering® Media reserved. Manufactured in the United States of America. This Susan Berge Producer: publication is protected by Copyright, and permission should be Katie Foley obtained from the publisher prior to any prohibited reproduction, Supplements Production storage in a retrieval system, or transmission in any form or by any Project Manager: Associate Mastering® Media means, electronic, mechanical, photocopying, recording, or likewise. Jane Brundage Producer: Taylor Merck To obtain permission to use material from this work, please submit a Manager, Text Permissions: written request to Pearson Education, Inc., Permissions Department, Editorial Media Producer: Tim Nicholls One Lake Street, Upper Saddle River, NJ 07458, or you can fill out one Daniel Ross Project Manager, Text of our forms at http://www.pearsoned.com/permissions/. Permissions: Senior Manager Web Readers may view, browse, and/or download material for temporary Alison Bruckner Development: Steve Wright copying purposes only, provided these uses are for noncommercial Text Permissions Specialist: personal purposes. Except as provided by law, this material may not Web Development Lead: James Toftness, Creative be further reproduced, distributed, transmitted, modified, adapted, Dario Wong Compliance, LLC performed, displayed, published, or sold in whole or in part, without Director of Production: Vice President of Marketing: prior written permission from the publisher. Erin Gregg Christy Lesko Many of the designations used by manufacturers and sellers to Executive Marketing Manager: Managing Editor: distinguish their products are claimed as trademarks. Where those Lauren Harp Michael Early designations appear in this book, and the publisher was aware of a Production Project Manager: Senior Marketing Manager: trademark claim, the designations have been printed in initial caps or Lori Newman Amee Mosley all caps. 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 introductory 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 g Gene Clonin Big Ideas help you connect the 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 Modi e 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. to foster considera ives Transgenic c borne on a rapidl aluable export cr . A stion continues nologies aect our l e used in tary staple and a v t a few decades ago que our diet, DNA tech and animals ar P paya is both a die med doomed jus ion to GMOs in edical and in- . relate biology to your life and the world In Hawaii, pa apaya industry see t the islands In addit used to produce m e and medicine g today, Hawaii’s p ad spread throughou s: Gene cloning is sic sci- agricultur ConnECtion your life Although thrivin spot virus (PRV) h tists from in many other way ed the eld of foren led the papaya ring ulation. But scien roling has chang adly pathogen cal he papaya plant pop al products, DNA p iological research, outside the classroom. de pletely eradicate t w, genetically engi- dustri aluable data for b ed to com eating ne produce v is ▹ and appeared pois the industry by cr and e, new technologies cal questions. In th ly modi ed were able to rescue once again vibrant— enc investigate histori Are genetical iversity of Hawaii papaya industry is can even be used to lso consider the the Un papaya. Today, the and DNA pplications. We’ll a s safe? resistant strains of rganisms (GMOs). uss each of these a he social, organism neered PRV- etically modied o the Hawai- chapter, we’ll disc lied, and some of t ? ii’s papayas are gen ing the recovery of , how they are app ling t majority of Hawa stances surround e c techniques used technologies. DNA Pro the vas y about the circum onsumption in th speci aised by the new 2.16) t everyone is happ s are approved for c cal issues that are r (12.11–1 However, no y modied papaya safety con- legal, and ethi lthough geneticall e critics have raised papaya industry. A d vegetables), som ee-year kers can be used NEW! Each chapter ian her GMO fruits an ccasions over a thr Genetic mar es (as are many ot ment. On three o ely match a DNA 16.5 Bio lms are complex associations of microbes United Stat and for the environ to de nitiv ople who eat them . opens with a high- cerns—for the pe sample to an individual In many natural environments, prokaryotes attach bacteria can also form on implanted medical devices such as to surfaces in highly organized colonies called catheters, replacement joints, or pacemakers. e complex- CONNECTION bio lms. A biolm may consist of one or several ity of biolms makes these infections especially dicult to interest question to species of prokaryotes, and it may include protists defeat. Antibiotics may not be able to penetrate beyond the

▹ and fungi as well. Biolms can form on almost any outer layer of cells, leaving much of the community intact. spark your interest in support, including rocks, soil, organic material (including For example, some biolm bacteria produce an enzyme that living tissue), metal, and plastic. You have a biolm on your breaks down penicillin faster than it can diuse inward. Genomics teeth—dental plaque is a biolm that can cause tooth decay. Biolms that form in the environment can be dicult to the topic. Questions Biolms can even form without a solid foundation, for ex- eradicate, too. A variety of industries spend billions of dollars (12.17–12.21) ample, on the surface of stagnant water. every year trying to get rid of bio- e DNA are revisited later in The study of complet Biolm formation begins when prokaryotes secrete signal- lms that clog and corrode pipes, ing molecules that attract nearby cells into a cluster. Once the gum up lters and drains, and coat sets helps us learn about . cluster becomes suciently large, the cells produce a gooey the hulls of ships (Figure 16.5). olutionary history the chapter, in either ev coating that glues them to the support and to each other, Biolms in water distribution making the biolm extremely dicult to dislodge. For exam- pipes may survive chlorination, a Scientific Thinking or ple, if you don’t scrub your shower, you could nd a biolm the most common method of en- growing around the drain—running water alone is not strong suring that drinking water does enough to wash it away. As the biolm gets larger and more not contain any harmful micro- Evolution Connection complex, it becomes a “city” of microbes. Communicating by organisms. For example, biolms chemical signals, members of the community coordinate the of Vibrio cholera, the bacterium division of labor, defense against invaders, and other activi- that causes cholera, found in water ▲ Figure 16.5 A bio lm module. fouling the insides of a pipe ties. Channels in the biolm allow nutrients to reach cells in pipes were capable of withstanding the interior and allow wastes to leave, and a variety of envi- levels of chlorine 10 to 20 times higher than the concentrations ronments develop within it. routinely used to chlorinate drinking water. Biolms are common among bacteria that cause disease in

humans. For instance, ear infections and urinary tract infec- Why are bio lms dicult to eradicate?

erior of the bio lm. the of erior t n i the o t n i ting a r penet om r f es c

tions are oen the result of biolm-forming bacteria. Cystic ? substan

obial obial r timic n a t n e v e r p y a m ells c of er y a l er t ou the other; each o t

brosis patients are vulnerable to pneumonia caused by bac- stick bio lm e up the the up e k ma t a th ells c the and on, esides r it e c fa r su the o t sticks bio lm he T

teria that form biolms in their lungs. Biolms of harmful ●

231 16.6 Prokaryotes help clean up the environment e characteristics that have made prokaryotes so widespread and successful—their nutritional CONNECTION diversity, adaptability, and caEvolutionpacity for forming biolms—also make them useful for cleaning up S OF DESIGN SERVICE contaminants in the environment. 8/11/13 11:15 AM C / M / Y / K S4CARLISLE Bioremediation is the use of organisms to remove pol- eece Pg. No. 231 Publishing Services ConnECtion Au: R Short / Normal lutants from soil, air, or water. Prokaryotic decomposers are earson Education / NJ / CHET Server: 230 # 110463 Cust: P the mainstays of sewage treatment facilities. Raw sewage ◃ Evolution Connection S OF Title: Campbell Biology: Concepts & Connections 0e is rst passed through a series of screens and shredders, DESIGN SERVICE M12_REEC5325_08_SE_CH12.indd 231 8/11/13 11:15 AM C / M / Y / K S4CARLISLE and solid matter settles out from the liquid waste. is modules present eece Pg. No. 230 Publishing Services solid matter, called sludge, is then gradually added to a Au: R Short / Normal Rotating earson Education / NJ / CHET 10.19 Emerging viruses threaten human health # 110463 Cust: P Server: culture of anaerobic prokaryotes, including both bacte- spray arm ria and archaea. e microbes decompose the organic Title: Campbell Biology: Concepts & Connections 0e Emerging viruses are ones that seem to burst on to ▼ Figure 10.19 A Hong Kong concrete examplesRock bed coated M12_REEC5325_08_SE_CH12.indd 230 the scene, becoming apparent to the medical matter in the sludge into healthmate-rcaiarle t whoartker ca pnr epabe rpeslac toe culld in with aerobic EVOLUTION a chicken to help prevent the spread community quite suddenly. ere are a landll or used as fertilizer. prokaryotes CONNECTION Liquid wastes are treatofe dthe se apvianara utel yvirus fro (shm otwnhe insl udtheg e. of the evidenceand fungi for many familiar examples, such as the inset) In Figure 16.6A, yoTEM 180,000 u can see a trickling lter system,

2009 H1N1 in uenza virus (discussed ed in the chapter introduction). Another one type of mechai z nism for treating liquid wastes. e evolution within each olo r example is HIV (human immunodeciency long horizontal pipC es rotate slowly, spraying liquid wastes Liquid wastes Outow virus), the virus that causes AIDS (acquired through the air onto a thick bed of rocks, the lter. Biolms ▲ Figure 16.6Achapter, The trickling lter s yprovidingstem at a sewage treatment pla yount immunodeciency syndrome). HIV ap- of aerobic bacteria and fungi growing on the rocks remove peared in New York and California in the early 1980s, seemingly out of nowhere. Yet 324 CHAPTER 16 Microbial Life: Prokaryotes and Protists with a coherent theme ABC News Videos and another example is the deadly Ebola virus, ◃ recognized initially in 1976 in central Africa; for the study of life. it is one of several emerging viruses that cause Current Events articles from DESIGN SERVICES OF hemorrhagic fever, an oen fatal syndrome char- # 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. 324 C / M / Y / K S4CARLISLE acterized by fever, vomiting, massive bleeding, and M16_REEC5325_08_SE_CH16.inddTitle: 324 Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services 8/11/13 11:19 AM The New York Times connect circulatory system collapse. A number of other dangerous newly recognized viruses cause encephalitis, in amma- what you learn in biology tion of the brain. One example is the Why are viral West Nile virus, which appeared in North America in 1999 and has since diseases such a class to fascinating stories spread to all 48 contiguous U.S. states. constant threat? West Nile virus is spread primarily by mosquitoes, which carry the virus in in the news. blood sucked from one victim and can transfer it to another victim. West Nile virus cases surged in 2012, especially in Texas. Severe acute respiratory syndrome (SARS) rst appeared in China in 2002. Within eight months, about 8,000 people were infected, and 10% died. Researchers quickly identied the infectious agent as a previously unknown, iv single-stranded RNA coronavirus, so named for its crown- birds appeared to stop that outbreak (Figure 10.19). Beginning v like “corona” of spikes. in 2002, however, new cases of human infection by this bird How do such viruses emerge suddenly, giving rise to new strain began to spread to Europe and Africa. As of 2012, the diseases? And why are viral diseases so hard to eradicate? disease caused by this virus, now called “avian u,” has killed ree processes contribute to the emergence of viral diseases: more than 300 people, and more than 100 million birds either mutation, contact among species, and spread from isolated have died from the disease or have been killed to prevent the populations. spread of infection. e H1N1 in uenza strain is another e mutation of existing viruses is a major source of example of a virus spreading from animals to humans; aer DESIGN SERVICES OF new viral diseases. RNA viruses tend to have unusually high circulating among pigs for many years, a mutated form began # 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. iv rates of mutation because errors in replicating their RNA to infect humans, creating the 2009 pandemic. A01_REEC5325_08_SE_FM_NASTA.indd 4 C / M / Y / K 11/6/13 7:47 AM S4carlisle genomes are not subject to the kind of proofreading and e spread of a viral disease from a small, isolated human Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services repair mechanisms that help reduce errors in DNA replica- population can also lead to widespread epidemics. For in- tion. Some mutations change existing viruses into new strains stance, AIDS went unnamed and virtually unnoticed for (genetic varieties) that can cause disease in individuals who decades before it began to spread around the world. In this have developed immunity to ancestral strains. at is why case, technological and social factors—including aordable we need yearly u vaccines: Mutations create new in uenza international travel, blood transfusions, sexual practices, and virus strains to which previously vaccinated people have no the abuse of intravenous drugs—allowed a previously rare immunity. human disease to become a global scourge. If we ever manage New viral diseases oen arise from the spread of exist- to control HIV and other emerging viruses, that success will ing viruses from one host species to another. Scientists esti- likely develop out of our understanding of molecular biology. mate that about three-quarters of new human diseases have W hy doesn’t a u shot one year give us immunity to u in

originated in other animals. For example, in 1997, at least

? subsequent years?

. d e t a cin c a

18 people in Hong Kong were infected with a strain of u v

e been been e v ’ e w which o t ones the om r f t n e r e di be ely k li most will er t a l us t c e f

virus called H5N1, which was previously seen only in birds. in t t a th ains r st the , s thu tion; a mut t n eque r f y b apidly r e v ol v e viruses n uenza I

A mass culling of all of Hong Kong’s 1.5 million domestic ●

202 CHAPTER 10 Molecular Biology of the Gene

B IG IDEA S g Gene Clonin Big Ideas help you connect the 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 Modi e span, sumably a Geneti

humans. For instance, ear infections and urinary tract infec- Why are bio lms dicult to eradicate?

erior of the bio lm. the of erior t n i the o t n i ting a r penet om r f es c

tions are oen the result of biolm-forming bacteria. Cystic ? substan

obial obial r timic n a t n e v e r p y a m ells c of er y a l er t ou the other; each o t

brosis patients are vulnerable to pneumonia caused by bac- stick bio lm e up the the up e k ma t a th ells c the and on, esides r it e c fa r su the o t sticks bio lm he T

teria that form biolms in their lungs. Biolms of harmful ●

2009 H1N1 in uenza virus (discussed ed in the chapter introduction). Another one type of mechai z nism for treating liquid wastes. e evolution within each olo r example is HIV (human immunodeciency long horizontal pipC es rotate slowly, spraying liquid wastes Liquid wastes Outow virus), the virus that causes AIDS (acquired through the air onto a thick bed of rocks, the lter. Biolms ▲ Figure 16.6Achapter, The trickling lter s yprovidingstem at a sewage treatment pla yount immunodeciency syndrome). HIV ap- of aerobic bacteria and fungi growing on the rocks remove peared in New York and California in the early 1980s, seemingly out of nowhere. Yet 324 CHAPTER 16 Microbial Life: Prokaryotes and Protists with a coherent theme ABC News Videos and another example is the deadly Ebola virus, ◃ recognized initially in 1976 in central Africa; for the study of life. it is one of several emerging viruses that cause Current Events articles from DESIGN SERVICES OF hemorrhagic fever, an oen fatal syndrome char- # 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. 324 C / M / Y / K S4CARLISLE acterized by fever, vomiting, massive bleeding, and M16_REEC5325_08_SE_CH16.inddTitle: 324 Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services 8/11/13 11:19 AM The New York Times connect circulatory system collapse. A number of other dangerous newly recognized viruses cause encephalitis, in amma- what you learn in biology tion of the brain. One example is the Why are viral West Nile virus, which appeared in North America in 1999 and has since diseases such a class to fascinating stories spread to all 48 contiguous U.S. states. constant threat? West Nile virus is spread primarily by mosquitoes, which carry the virus in in the news. blood sucked from one victim and can transfer it to another victim. West Nile virus cases surged in 2012, especially in Texas. Severe acute respiratory syndrome (SARS) rst appeared in China in 2002. Within eight months, about 8,000 people were infected, and 10% died. Researchers quickly identied the infectious agent as a previously unknown, iv single-stranded RNA coronavirus, so named for its crown- birds appeared to stop that outbreak (Figure 10.19). Beginning v like “corona” of spikes. in 2002, however, new cases of human infection by this bird How do such viruses emerge suddenly, giving rise to new strain began to spread to Europe and Africa. As of 2012, the diseases? And why are viral diseases so hard to eradicate? disease caused by this virus, now called “avian u,” has killed ree processes contribute to the emergence of viral diseases: more than 300 people, and more than 100 million birds either mutation, contact among species, and spread from isolated have died from the disease or have been killed to prevent the populations. spread of infection. e H1N1 in uenza strain is another e mutation of existing viruses is a major source of example of a virus spreading from animals to humans; aer new viral diseases. RNA viruses tend to have unusually high circulating among pigs for many years, a mutated form began DESIGN SERVICES OF # 110463 Cust: Pearsonrates of m uEducationtation because er ro/ rNJs in r/ep liCHETcating their RNAu:A Reeceto infePg.ct hu mNo.ans, c rvea ting the 2009 pandemic.C / M / Y / K A01_REEC5325_08_SE_FM_NASTA.indd 5 11/6/13 7:47 AM Title: Campbell Biology:genom esConcepts are not subject &to t hConnectionse kind of proofreadin g0e and Server: e spread of a viral disease from a small, isolated human S4carlisle repair mechanisms that help reduce errors in DNA replica- population can also lead to widespread epShortidemics. F o/r inNormal- Publishing Services tion. Some mutations change existing viruses into new strains stance, AIDS went unnamed and virtually unnoticed for (genetic varieties) that can cause disease in individuals who decades before it began to spread around the world. In this have developed immunity to ancestral strains. at is why case, technological and social factors—including aordable we need yearly u vaccines: Mutations create new in uenza international travel, blood transfusions, sexual practices, and virus strains to which previously vaccinated people have no the abuse of intravenous drugs—allowed a previously rare immunity. human disease to become a global scourge. If we ever manage New viral diseases oen arise from the spread of exist- to control HIV and other emerging viruses, that success will ing viruses from one host species to another. Scientists esti- likely develop out of our understanding of molecular biology. mate that about three-quarters of new human diseases have W hy doesn’t a u shot one year give us immunity to u in

originated in other animals. For example, in 1997, at least

? subsequent years?

. d e t a cin c a

18 people in Hong Kong were infected with a strain of u v

e been been e v ’ e w which o t ones the om r f t n e r e di be ely k li most will er t a l us t c e f

virus called H5N1, which was previously seen only in birds. in t t a th ains r st the , s thu tion; a mut t n eque r f y b apidly r e v ol v e viruses n uenza I

A mass culling of all of Hong Kong’s 1.5 million domestic ●

202 CHAPTER 10 Molecular Biology of the Gene

DESIGN SERVICES OF # 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. 202 C / M / Y / K S4CARLISLE M10_REEC5325_08_SE_CH10.inddTitle: 202 Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services 8/11/13 11:22 AM In addition to making secretory proteins, rough ER is a membrane-making machine for the cell. It grows in place by Now let’s follow a transport vesicle carrying products of adding membrane proteins and phospholipids to its own mem the rough ER to the Golgi apparatus. brane. As polypeptides destined to be membrane proteins grow- from bound ribosomes, they are inserted into the ER mem - brane. Phospholipids are made by enzymes of the rough ER Explain why we say that the endoplasmic reticulum is a and also inserted into the membrane. Thus, the ER membrane ? biosynthetic workshop.

grows, and portions of it are transferred to other components

for membranes, other organelles, and secretion by the cell. the by secretion and organelles, other membranes, for To the Student: How to use this book and MasteringBiology® ribosomes) bound by (synthesized proteins and hormones, steroid membranes,

of the endomembrane system in the form of transport vesicles. 1 The ER produces a huge variety of molecules, including phospholipids for cell cell for phospholipids including molecules, of variety huge a produces ER The 2.9 ● Genetic ally modi ed organisms raise health concerns As soon as scientists realize d the power of DNA tech- their n nology, they bega ew genes to related species in nearb SCIENTIFIC n to worry about potential dangers. y wild areas, dis- 4.9 Earl turbing the composition of th The Golgi apparatus modifies, sorts, and ships cell products THINKING y concerns focused on the possibility e natural ecosystem. Critics of that r GMO crops can po ecombinant DNA technology might Are gene int to several studies that do cre tically indeed show unin After leaving the ER, many transport vesicles travel to the ate new pathogens. To guard against tended gene transfer from en- rogue micr modi ed gineered crops to n Golgi apparatus. Using a light microscope and a staining functions as a depot, dispatching its products in vesicles that obes, scientists developed a set of earby wild relatives. But GMO Stay focused on Learn how to to think guidelines in organisms safe? advocates count technique he developed, Italian scientist Camillo Golgi dis- bud off and travel to other sites. cluding strict laboratory safety and er that no lasting or detrimental containm e ects from s ent procedures, the genetic cripplin uch transfers have been demon- covered this membranous organelle in 1898. The electron How might ER products be processed during their transit g of transgenic strated, and th the key concepts like a scientist organisms to ensure that they cann at some GMOs (such as bacteria eng through the Golgi? Various Golgi enzymes modify the carbo ot survive outside the lab- bre ineered to microscope confirmed his discovery more than 50 years later, oratory, and a prohibitio ak down oil spills) can actively help t - n on certain dangerous experiments. he environment. hydrate portions of the glycoproteins made in the ER, removing Today, most pu revealing a stack of flattened sacs, looking much like a pile of blic concern centers on GMOs used f Labeling Althoug some sugars and substituting others. Molecular identification or food. h the majority of several staple crops Central concepts pita bread. A cell may contain many, even hundreds, of these New Scientific Thinking Human Safety grown in the United Stat ▹ Genetically modied organi es—including corn and soybeans— tags, such as phosphate groups, may be added that help the Golgi sms are used in are geneticall stacks. The number of Golgi stacks correlates with how active crop production because they are m y modied, products made from GM summarize the key modules explore ore nutritious or because req Os are not the cell is in secreting proteins—a multistep process that, as sort molecules into different batches for different destinations. they are cheaper to pro uired to be labeled in any way. Chan duce. But do these advantages come ces are you ate a food Finished secretory products, packaged in transport at a cost to the healt containing GMOs today, but the lack o you have just seen, is initiated in the rough ER. h of people consuming GMOs? When f labeling means you topic of each how scientists use the investigating com probably can’t say for certain. Lab The Golgi apparatus serves as a molecular warehouse and vesicles, move to the plasma membrane for export from the plex questions like this one, scientis eling of foods containing use m ts oen more than trace amoun cell. Alternatively, finished products may become part of the ultiple experimental methods. A 2012 a ts of GMOs is required in Europe, module, helping you processing station for products manufactured by the ER. You processes of science nimal study Japan, Austra involved 104 pigs that were divide lia, China, Russia, and other countr plasma membrane itself or part of another organelle, such as d into two groups: e in ies. Label- can follow these activities in Figure 4.9 rst was fed a diet con g advocates point out that the inform . Note that the flat- taining 39% GMO corn and the other ation would allow stay focused as you tened Golgi sacs are not connected, as are ER sacs. a lysosome, which we discuss next. for discovery. Each a closely related n consumers to decide for themselv on-GMO corn. e health of the pigs wa es whether they wish to ➊ One measur s be exposed to GMO foo side of a Golgi stack serves as a receiving dock for transport ed over the short term (31 days), the m ds. Some biotechnology advocates, study. module concludes with a edium term however, res What is the relationship of the Golgi apparatus to the ER in a (110 days), and the normal genera pond that similar demands were not m vesicles produced by the ER. ➋ ? tional life span. e re- “tra ade when A vesicle fuses with a Golgi protein-secreting cell? searchers reported no sig nsgenic” crop plants produced by tradi sac, adding its membrane and contents to the “receiving” nicant di erences between the two tional breeding

question that challenges groups and no traces o techniques were put on the market. F proteins are secreted. are proteins f foreign DNA in the slaught or example, triticale (a

side. ➌ Products of the ER are modified as a Golgi sac pro- A ered pigs. crop used primarily in a proteins and dispatches transport vesicles to the plasma membrane, where the the where membrane, plasma the to vesicles transport dispatches and proteins lthough pigs are a good model o nimal feed but also in some h

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The “shipping” side of the Golgi hat human data are required to draw bining the genomes of The Golgi receives transport vesicles budded from the ER that contain contain that ER the from budded vesicles transport receives Golgi The Checkpoint ● wheat and rye—t conclusions about the safety of dieta wo plants that do not interbreed in n ry GMOs for people. e Tritic ature. scientist. results of one human stud ale is now sold worldwide without a y, conducted jointly by Chinese and ny special labeling. questions at the end American scientists, w Scientists and the public need to w “Receiving” side of Golgi apparatus ere published in 2012. Sixty-eight C eigh the possible ben- nese sch hi- ets versus risks on a cas oolchildren (ages 6–8) were fed Golden R e-by-case basis. e best scenario ice, spinach would be to pr of each module help New Scientific Thinking (a natural source of beta-carotene), o oceed with caution, basing our decisio ▹ r a capsule containing soun ns on Transport vesicle pure beta-carotene. Over 21 d d scientic information rather than o ➊ topics include: ays, blood samples were drawn n either irrational you stay on track. from the ER to measure how m fear or blind optimism. uch vitamin A the body produced fr each foo om d source. e data show that the bet ▸ Module 2.15 — Scientists study the a-carotene in Why might crop plan both Golden Rice and the c ts engineered to be resistant to w ➋ apsules was converted to vitamin ? killer pose a danger t eed

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nica herbicide or f n genes he tly less v T itamin A (Figure 12.9). ● coral reef ecosystems e results led resear chers to conclude that GMO rice can thinking indeed be e ective in p art provides clear reventing vitamin A deciency. ▸ Despite its positive 75 Module 8.10 — Tailoring treatment ndings, this study caused an uproar. Chinese authorities ca visuals to help you ➌ lled the study an unethical “scandal,” to each patient may improvecomp cancerlaining th Colorized, TEM 145,000× at U.S. scientists had used Chin ese schoolchil- bed and Golgi apparatus dren as laboratory su 50 understand key therapy bjects. e project leaders countered that proper per o vitamin A mission and consent had been obta bo ined in th China and the U ed t topics. Selected nited States. e controversy highlig t ▸ hts tage absor Module 25.3 — Coordinatedone of wavesthe dicu er 25 ➍ lties in conducting r v esearch on human en

c nutrition: Animal studies a on c figures include of movement in huddles help re of limited value, but human er studies may be un P ethical. To date, no study has docum heal ented 0 numbered steps penguins thermoregulate th risks in humans from GMO foods, a Transport vesicle nd there is gen- Capsule of pur eral agreement among scientists t e Golden rice Spinach from the Golgi hat the GMO foods on the beta-carotene ▸ market are safe. Howeve that are keyed to Module 26.3 — A widely used weed r, it is not yet possible to measure the long-term e ec ▲ Figure 12.9 Vitamin A production af ts (if any) of GMOs on huma ter consumption of di erent n health. sources of beta-carotene explanations in killer demasculinizes maleEn frogsvironmental S afety Adv Data from G. Tang et al., B “Shipping” side of Golgi apparatus ocates of a cautious approach eta-carotene in Golden Rice is as good as beta- toward GM in oil at providing vitamin A t carotene O crops fear that transgenic p o children, American Journal of Clinic the text. ▸ Module 29.2 — The model for lants might pass 658–64 (2012). al Nutrition 96(3): ▲ Figure 4.9 The Golgi apparatus receiving, processing, and shipping products 240 CHAPTE R 12 magnetic sensory reception is DNA Technology and Genomics incomplete The Endomembrane System 61 # 110463 Cust: Pearson Education / NJ / CHET M12_REEC5325_08_SE_CH12.indd 240 Au: Reece Pg. No. 240 Title: Campbell Biology: Concepts & Connections 0e C / M / Y / K DESIGN SERVICES OF Server: Short / Normal S4CARLISLE Publishing Services 8/22/13 7:46 AM # 110463 Cust: Pearson Education / NJ / CHET Au: Reece M04_REEC5325_08_SE_CH04.inddTitle: 61 Campbell Biology: Concepts & Connections 0e Pg. No. 61 C / M / Y / K DESIGN SERVICES OF Server: S4carliSle Short / Normal Publishing Services 10/29/13 1:27 PM

◃ Connecting the Concepts ◃ NEW! Scientific Thinking activities link one biological activities teach you how to concept to another. practice important scientific skills like understanding variables and making predictions. Specific wrong-answer feedback coaches you to the correct response.

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. vi C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 6 S4carlisle 11/6/13 7:47 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services In addition to making secretory proteins, rough ER is a membrane-making machine for the cell. It grows in place by Now let’s follow a transport vesicle carrying products of adding membrane proteins and phospholipids to its own mem the rough ER to the Golgi apparatus. brane. As polypeptides destined to be membrane proteins grow- from bound ribosomes, they are inserted into the ER mem - brane. Phospholipids are made by enzymes of the rough ER Explain why we say that the endoplasmic reticulum is a and also inserted into the membrane. Thus, the ER membrane ? biosynthetic workshop.

grows, and portions of it are transferred to other components for membranes, other organelles, and secretion by the cell. the by secretion and organelles, other membranes, for To the Student: How to use this book and MasteringBiology® ribosomes) bound by (synthesized proteins and hormones, steroid membranes,

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Before microscopes were rst used in the 1600s, no one knew that living organisms were composed of the tiny units we call cells. e rst microscopes were light microscopes, like the ones you may use in a biology laboratory. In a light microscope (LM), visible light is passed through a specimen, such as a microorganism or a thin slice of animal or plant tissue, and then through glass lenses. e lenses bend the light in such a way that the image of the specimen is magni ed as it is projected into your eye or a camera. Magni cation is the increase in an object’s image size Figure 4.1A

compared with its actual size. shows a micrograph LM 230 × of a single-celled organism called Paramecium. e notation “LM 230* ” printed along the right edge tells you that this mi- ▲ Figure 4.1A Light micrograph of a unicellular organism, Paramecium crograph was taken through a light microscope and that the image is 230 times the actual size of the organism. e size of this Paramecium is about 0.33 millimeter in length. Table 4.1 shows the most common units of length that biologists use. To the Student: How to use this book and MasteringBiology® An important factor in microscopy is resolution, a measure of the clarity of an image. Resolution is the ability to distinguish two nearby objects as separate. For example, what you see as a single star in the sky may be resolved as twin stars with a telescope. Each optical instrument—be it an eye, a × telescope, or a microscope—has a limit to its resolution. e ee C Cyyclescles human eye can distinguish points as close together as 0.1 mil- Maximize your learning tionstions and and Plant Plant Li Liff ed SEM 580 tion of Genera i z ▹▹ AlAltternernaation of Genera ee C Cyyclecle limeter (mm), about the size of a very ne grain of sand. A AA F Fernern Li Liff olo r and success typical light micro scope cannot resolve detail ner than about C t liefe c cyyclescles ernernaattee in in pla plannt lif ▲ Figure 4.1B Scanning electron micrograph of Paramecium tionstions al altt No matter how many times the image of such a small cell is New Visualizing the Concept INGG 17.317.3 Haploid Haploid and and diploid diploid gener generaa ▹ VIVISSUUALIZALIZIN DiploidDiploid (2 (2n)n) Haploid (n) t (tn ()n)magni ed, the light microscope cannot resolve the details of TTHHEE C COONNCCEPEPTT YYCLECLE KeKyey Haploid (n) GameGametoptophyhtyet plae plan n modules walk you through ANTANT LIFE LIFE C C phyteyte its structure. Indeed, light microscopes can eectively magnify THETHE P PLL e aphaploliodi dgamet gametopho y ThThe h tse s fromfrom ours. ours. Humans Humans are are diploid diploid individuals— individuals— ds psorpore edi divivdeides sb yb uces haploidi dg agmeamete AA sin singlge-le-ceclelelled prpordoudces haplo tosis.is. objects only about 1,000 times. that is, each of us has two sets of chromosomes, lopsp s ndn deggs eggs) by) by mi mitos challenging concepts and that is, each of us has two sets of chromosomes, mimitotsiosis sa ndand deve develo (s(pespermrm a a ular oneone from from each each parent parent (Module (Module 8.12). 8.12). Gametes Gametes (sperm (sperm and and intointo a amul multiceticellulllar GGameametoptophyhtyete From the time that Hooke discovered cells in 1665 until MM Spemrm (n ()n) y MiMtoitsoissis mm swim swim to t theo the metoetophytphyte.e. platn (tn ()n) itoito Sper e edivides divides b yb SpeSper r eggs)eggs) are are the the only only haploid haploid stage stage in in the the human human life life cycle. cycle. gagam d d plan si si A singl-ec-elledcelled spo spor r emale complex processes. n n t t s s A single ot oa a the middle of the 1900s, bioloegggeggi sints in the theh femalead f o nly light microscopes a aenen elopselops in itn PlantsPlants have have an an alternation alternation of of generations generations: The: The diploid diploid is is EggEgg (n ()n) mimitosistosis and and de dev v iumium s s mm ytyet.e. gametangametang g o o p p opoph h The male t t o o ellularellular game gamet t for Tvheie malewing cells. With these microscopes and various stain- andand haploid haploid stages stages are are distinct, distinct, multicellular multicellular bodies. bodies. i i l l multimultic c e e liezses M rm fertirltiiz iumium eses (n ()n) M v v The life cycles A Aspe sperm fe gametangametang g The haploid generation of a plant produces gametes SpoSpor r e e The life cycles of awtaetr.er. The haploid generation of a plant produces gametes d m. of w d n egg,res resulutilngting poduroducesces spe sperm.r ofof all all plants plants follow follow the the ana egg, d d ing tperchniques to increase contrast between parts of cells, The brief narrative works andand is is called called the the gametophyte gametophyte. The. The diploid diploid generation generation loid dz yzgyogteote. . nan t t ini na adip diploi as nen patternpattern shown. shown. Be Be sure sure issi e UndersideUnderside Spemrm ▹ × s Sper producesproduces spores spores and and is is called called the the sporophyte sporophyte. In. In a a o o mpm these early biologists discovered microorganisms, animal and it it opo thatthat you you understand understand l el of ofgametophyte: gametophyte: plant’splant’s life life cycle, cycle, these these two two generations generations alternate alternate in in M Mve v tion e e together with the artwork thisthis diagram; diagram; then then FeFretilizrtilizationa d d actualactual size size 0.5 0.5 cm cm producing each other. In mosses, as in all nonvascular plant cells, and eTvheTenhe female femalesome structures within cells. By the mid- producing each other. In mosses, as in all nonvascular Meiosiseiosis SpoSporesres acrossacross M reviewreview it itafter after studying studying gametaniumgium plants,plants, the the gametophyte gametophyte is is the the larger, larger, more more obvious obvious stage stage ZyZgoygotet e(2 (2n)n) 1800s, this accumgametanulatigon of evidence led to the cell theory, eacheach life life cycle cycle to to see see prpoduroducesces TEM 9,375 to help you visualize and ofof the the life life cycle. cycle. Ferns, Ferns, like like most most plants, plants, have have a alife life cycle cycle howhow the the pattern pattern d d t t . . gg ed n n n n celledcelled which states that analan egl eglgigving things are coEmggE posed of cells and dominated by the sporophyte. Today, about 95% of all a a e e ThTeh esi sngle-ingle- dominated by the sporophyte. Today, about 95% of all applies.applies. opophyhtyet e i z is is m m es by TheThe spo spor r s s p p zgoygotet edi divividesd by plants,plants, including including all all seed seed plants, plants, have have a adominant dominant e totoo o zy es spoesres b yb y understand the topic. ThThe i i l el oposp s prpoduroducesc spor that all cells come from other cells. MM e nadn dd eveldevel olo r yhtyet e evev mmitoitsoisi as anangia.gia. AlthoughAlthough eggs eggs and and sperm sperm sporophytesporophyte in in their their life life cycle. cycle. The The life life cycles cycles of of all all plants plants spsorporopoph C opophyhtyete d d lctiellcellulualra r meiosismeiosis in in spo spor r roducduceses SpoSpor r initnot oa am muluti areare usually usually produced produced in inseparate separate followfollow a apattern pattern shown shown here. here. prop t (2n)n) Our knowledge of cell structure took a giant leap forward id s psorporese s plaplantn (2 sporsporophophyteyte. . locationslocations on on the the same same gametophyte, gametophyte, hahaplpolido FeFretilizrtilizationation byb ym meieoisisosis. . as bioalo varietya varietygis ofts ofmechanisms mechanismsbegan promote upromotesin g the electron microscope in the ▲ Figure 4.1C Transmission electron micrograph of Toxoplasma cross-fertilizationcross-fertilization between between (This parasite of cats can be transmitted to humans, causing the disease ee C Cyyclecle MMatuaturere 1950s. Instgametophytes.egametophytes.ad of using light, an electron microscope (EM) AA Moss Moss Li Liff MMeiosiseiosis sposporoprophyhtyete toxoplasmosis.) InIn plants, plants, gametes gametes are are focuses a beam of electrons through a specimen or onto its MMitosisitosis producedproduced by by mitosis. mitosis. Try This Describe a major diﬀerence between the Paramecium in Figure 4.1B and the protist ZyZygogotet e TheThe green, green, cushiony cushiony in this ﬁgure. (Hint: Compare the notations along the right sides of the micrographs.) mossmoss we we see see consists consists n TABLE 4.1 METRIC MEASUREMENT EQUIVALENTS Hints embedded within the e eg agmeametantangigiumum in i mm to t oth teh eegg egg in i n TheThe new new ofof gametophytes. gametophytes. ThTh SpeSpermrm swi swi yete lele g agmetametophophytytee mmetangietangiumum sposporoprophyht ndnd t t a ama ma thethe fem femaleale ga ga a a enen esces s pespermrm. . grgorwosw fsr omfrom the the is is m prpordoudcu module emulate the guidance osos pmp ytyet.e. it it lolo gamegame1top tmeoph hter (m) = 100 cm = 1,000 mm = 39.4 inches MM e e surface. Electron microscopes can distinguish biological evev d d d d - 2 n n t t 1 centimeter (cm) = 10 (1/100) a amn n = 0.4 inch that you might receive during is is e e structures as small as about 2 nanometers (nm), a 100-fold s s m m SpeSpermrm to top p s psporeore - 3 i i lo lo - 1 AA s isnigngle-le-celcellelde d M Me e New! Try This activities help you actively an and d 1 millimeter (mm) = 10 (1/1,000)evev m = 10 (1/10) cm improvement over the light microscope. is high resolution didividvideses b by ym mitosisitosis d d ▵ class or in a tutoring session. elellullaular r eveevelopslops in itonto multic multic - 6 - 3 dd ThThe eg agmeametantangigiumum ClustersClusters of ofsporangia sporangia = 10 m = 10 mm has enabled biologists to explore cell ultrastructure, the com- ggametopametophyhtyet.e. engage with the figures and develop inin a afe femamalele onon this this fern fern look look like like - 9 - 3 Figures 4.1B 4.1C These hints provide additional opophyhtyet ep lpalnatnst s(n ()n) hythytee 1 nanometer (nm) = 10 m = 10 ygoygotet dividese divides plex internal anatomy of a cell. and show GaGamemet t gagmeametotopp EgEggg brownbrown dots. dots. TheThe singl single-ecelled-celled z z ana negg egg. . elopselops in itnot ao a SpSporeores s(n ()n) prpordoudcuesces byb miy mitosistosis and and de dev v positiveimages prod studyuced by thabits.wo kinds of electron microscopes. opophyhtyet.e. information to deepen your giguimum multimulticellularcellular spo spor r SpSoranporan g rogroww SpoSporoprophyhtyetse s(2 (n2)n) g amgametoetophyphytetes.s. frformom 52 CHAPTER 4 A Tour of the Cell understanding of the topic. FeFretilizrtilizationation poroorophyphytete TheThe tiny tiny gametophyte gametophyte soon soon ThThe es posporoprophytehyte SSp pprorodduceuces ssp sporeores s disintegrates,disintegrates, and and the the sporophyte sporophyte growsgrows independently. independently. bby ym meieosiosisi sin in th thee TheThe sporophyte sporophyte cannot cannot spsporoarngangiuimum. . photosynthesize—itphotosynthesize—it is isdependent dependent s s onon the the gametophyte. gametophyte. 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Chapter Title 574 CHAPTE R 00 To the Teacher: Implement active learning in your classroom

Resources save you hours of time preparing for class

▹ NEW! Learning ◃ Instructor Exchange, moderated Catalytics™ is a “bring by co-author Kelly Hogan, offers a your own device” student library of active learning strategies engagement, assessment, contributed by educators from and classroom intelligence across the country. system. This technology has grown out of twenty years of cutting-edge research, innovation, and implementation of ▿ BioFlix activities offer students 3-D interactive teaching and animations to help them visualize peer instruction. and learn challenging topics.

Assign tutorials to help students prepare for class ▿ Video Tutor Sessions and MP3 Tutor Sessions, hosted by co-author Eric Simon, provide on- the-go tutorials focused on key concepts and vocabulary. Connect your class to current topics ◃ Campbell Current Topics PowerPoint slides help you prepare a high-impact discussion developed around current issues. Topics include cancer, global climate change, athletic cheating, nutrition, and more.

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Resources save you hours of time preparing for class

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MasteringBiology® is an online assessment and tutorial system designed Gain insight into student to help you teach more efficiently. It offers a variety of interactive activities to engage students and help them to succeed in the class. progress at a glance ▹ Get daily diagnostics. Access students’ results with easy-to-interpret student performance data Gradebook Diagnostics provide unique insight into class performance. With a ◃ Gradebook single click, see a summary of how your students are struggling or progressing. • Every assignment is automatically graded.

• At a glance, shades of red highlight vulnerable MasteringBiology® is easy for you students and challenging and your students to use assignments. ◃ The Mastering platform is the most effective and widely used online tutorial, homework, and assessment system for the sciences.

▹ Student performance data reveal how students are doing compared to a national average and which topics they’re struggling with. With MasteringBiology®, you can: • Assign publisher-created pre-built ▵ Efficacy studies Wrong answer summaries assignments to get started quickly. ▹ Go to the “Proven Results” tab at give unique insight into your • Easily edit any of our questions or answers to students’ misconceptions and match the precise language you use. www.masteringbiology.com to see efficacy studies. support just-in-time teaching. • Import your own questions and begin compiling meaningful data on student performance. • Easily export grades to Microsoft®Excel or other course-management systems.

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A01_REEC5325_08_SE_FM_NASTA.indd 13 11/6/13 7:48 AM Preface

nspired by the thousands of students in our own classes Connection icons mark the numerous application modules over the years and by enthusiastic feedback from the that go beyond the core biological concepts. For example, many educators who have used our book, we are de- the new Connection Module 26.12 describes the potential Ilighted to present this new, Eighth Edition. We authors have role oxytocin plays in human–dog bonding. In addition, our worked together closely to ensure that both the book and the Evolution Connection modules, identified by green icons, supplementary material online reflect the changing needs of connect the content of each chapter to the grand unifying today’s courses and students, as well as current progress in theme of evolution, without which the study of life has no biology. Titled Campbell Biology: Concepts & Connections to coherence. Explicit connections are also made between the honor Neil Campbell’s founding role and his many contri- chapter introduction and either the Evolution Connection butions to biology education, this book continues to have a module or the new Scientific Thinking module in each dual purpose: to engage students in the wonders of the living chapter; new high-interest questions introduce each chapter, world and to show them how biology relates to their own drawing students into the topic and encouraging a curiosity existence and the world they inhabit. Most of these students to explore the question further when it appears again later in will not become biologists themselves, but their lives will be the chapter. touched by biology every day. Understanding the concepts of biology and their connections to our lives is more important New to This Edition than ever. Whether we’re concerned with our own health or the health of our planet, a familiarity with biology is essential. New Scientific Thinking Modules In this edition we placed This basic knowledge and an appreciation for how science greater emphasis on the process of scientific inquiry through works have become elements of good citizenship in an era the addition to each chapter of a new type of module called when informed evaluations of health issues, environmental Scientific Thinking, which is called out with a purple icon. problems, and applications of new technology are critical. These modules cover recent scientific research as well as underscore the spirit of inquiry in historical discoveries. Concepts and Connections All Scientific Thinking modules strive to demonstrate to students what scientists do. Each of these modules identifies Concepts Biology is a vast subject that gets bigger every key attributes of scientific inquiry, from the forming and year. This book was the first introductory biology textbook testing of hypotheses to the analysis of data to the evaluation to use concept modules to help students recognize and focus and communication of scientific results among scientists and on the main ideas of each chapter. The heading of each with society as a whole. For example, the new Module 2.15 module is a carefully crafted statement of a key concept. describes how scientists use both controlled experiments and For example, “A nerve signal begins as a change in the observational field studies to document the effects of rising membrane potential” announces a key concept about the atmospheric CO2 on coral reef ecosystems. Module 13.3 generation of an action potential (Module 28.4). Such a describes the scientific search for the common ancestor of concept heading serves as a focal point, and the module’s whales, using different lines of inquiry from early fossil clues, text and illustrations converge on that concept with molecular comparisons, and a series of transitional fossils explanation and, often, analogies. The module text walks that link whales to cloven-hoofed mammals, animals that live the student through the illustrations, just as a teacher might on land. And to prepare students for the renewed focus in the do in class. And in teaching a sequential process, such as book on how biological concepts emerge from the process the one diagrammed in Figure 28.4, we number the steps of science, we have significantly revised the introduction in in the text to correspond to numbered steps in the figure. Chapter 1, Biology: Exploring Life. These changes will better The synergy between a module’s narrative and graphic equip students to think like scientists and emphasize the components transforms the concept heading into an idea connections between discovery and the concepts explored with meaning to the student. The checkpoint question at throughout the course. the end of each module encourages students to test their New Visualizing the Concept Modules Also new to this edition understanding as they proceed through a chapter. Finally, in are modules that raise our hallmark art–text integration to the Chapter Review, all the key concept statements are listed a new level. These Visualizing the Concept modules take and briefly summarized under the overarching section titles, challenging concepts or processes and walk students through explicitly reminding students of what they’ve learned. them in a highly visual manner, using engaging, attractive Connections Students are more motivated to study biology art; clear and concise labels; and teacher hints called out in when they can connect it to their own lives and interests— light blue bubbles. These short teacher hints emulate the one- for example, when they are able to relate science to health on-one coaching a teacher might provide to students during issues, economic problems, environmental quality, ethical school hours and help students make key connections within controversies, and social responsibility. In this edition, blue the figure. Examples of this new feature include Module 9.8,

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xiv C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 14 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services which demonstrates to students the process of reading and for a new representation of a molecule of tRNA binding to an analyzing a family pedigree; Module 17.3, which introduces enzyme molecule. the concept of plant life cycles through a combination of The Latest Science Biology is a dynamic field of study, and photographs and detailed life cycle art displayed across an we take pride in our book’s currency and scientific accuracy. impressive two-page layout; and Module 26.8, which walks For this edition, as in previous editions, we have integrated students through the concept of homeostatic controls in the results of the latest scientific research throughout blood glucose levels. the book. We have done this carefully and thoughtfully, New “Try This” Tips One theme of the revision for the Eighth recognizing that research advances can lead to new ways of Edition is to help all students learn positive study habits they looking at biological topics; such changes in perspective can can take with them throughout their college careers and, in necessitate organizational changes in our textbook to better particular, to encourage them to be active in their reading reflect the current state of a field. You will find a unit-by-unit and studying. To foster good study habits, several figures account of new content and organizational improvements in in each chapter feature a new “Try This” study tip. These the “New Content” section on pp. xvii–xviii following this action-oriented statements or questions direct students to Preface. study a figure more closely and explain, interpret, or extend New MasteringBiology® A specially developed version of what the figure presents. For example, in Figure 3.13B, MasteringBiology, the most widely used online tutorial and students are asked to “Point out the bonds and functional assessment program for biology, continues to accompany groups that make the R groups of these three amino acids Campbell Biology: Concepts & Connections. In addition either hydrophobic or hydrophilic.” Figure 6.10B is a new to 170 author-created activities that help students learn figure illustrating the molecular rotary motor ATP synthase, vocabulary, extend the book’s emphasis on visual learning, and the accompanying Try This tip asks students to “Identify demonstrate the connections among key concepts (helping the power source that runs this motor. Explain where this students grasp the big ideas), and coach students on how to ‘power’ comes from.” Figure 36.7, on the effect of predation interpret data, the Eighth Edition features two additional new on the life history traits of guppies, offers the following Try activity types. New Scientific Thinking activities encourage This tip: “Use the figure to explain how the hypothesis was students to practice the basic science skills explored in the tested.” in-text Scientific Thinking feature, allowing students to try Improvements to End-of-Chapter Section The Testing Your out thinking like a scientist and allowing teachers to assess Knowledge questions are now arranged to reflect Bloom’s this understanding; new Visualizing the Concept activities Taxonomy of cognitive domains. Questions and activities are take students on an animated and narrated tour of select grouped into Level 1: Knowledge/Comprehension, Level 2: Visualizing the Concept modules from the text, offering Application/Analysis, and Level 3: Synthesis/Evaluation. In students the chance to review key concepts in a digital addition, a new Scientific Thinking question has been added learning modality. MasteringBiology® for Campbell Biology: to each chapter that connects to and extends the topic of the Concepts & Connections, Eighth Edition, will help students Scientific Thinking module. Throughout the Chapter Review, to see strong connections through their print textbook, new questions have been added that will help students better and the additional practice available online allows teachers engage with the chapter topic and practice higher-level to capture powerful data on student performance, thereby problem solving. making the most of class time. See program components for New Design and Improved Art The fresh new design MasteringBiology access information on the next page. used throughout the chapters and the extensive reconceptualization of many figures make the book even This Book’s Flexibility more appealing and accessible to visual learners. The cellular Although a biology textbook’s table of contents is by design art in Chapter 4, A Tour of the Cell, for example, has been linear, biology itself is more like a web of related concepts completely reimagined for more depth perspective and without a single starting point or prescribed path. Classes richer color. The new big-picture diagrams of the animal can navigate this network by starting with molecules, and plant cells are vibrant and better demonstrate the with ecology, or somewhere in between, and teachers can spatial relationships among the cellular structures with omit topics. Campbell Biology: Concepts & Connections is an almost three-dimensional style. The illustrations of uniquely suited to offer flexibility and thus serve a variety cellular organelles elsewhere in Chapter 4 include electron of classes. The seven units of the book are largely self- micrographs overlaid on diagrams to emphasize the contained, and in a number of the units, chapters can be as- connection between the realistic micrograph depiction and signed in a different order without much loss of coherence. the artwork. Figure 4.9, for example, shows a micrograph The use of numbered modules makes it easy to skip topics of an actual Golgi apparatus paired with an illustration; or reorder the presentation of material.

an accompanying orientation diagram—a hallmark of ■■■ Concepts and Connections—continues to act as a roadmap that reminds students of how an organelle fits within the Long after today’s students have forgotten most of the spe- overall cell structure. Finally, throughout the book we have cific content of their biology class, they will be left with gen- introduced new molecular art; for example, see Figure 10.11B eral impressions and attitudes about science and scientists.

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xv C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 15 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services We hope that this new edition of Campbell Biology: Concepts Jane Reece, [email protected] & Connections helps make those impressions positive and Martha Taylor (Chapter 1 and Unit I), [email protected] supports teachers’ goals for sharing the fun of biology. In Eric Simon (Units II and VI and Chapters 21 and 27), [email protected] our continuing efforts to improve the book and its supporting materials, we benefit tremendously from teacher and Jean Dickey (Units III, IV, and VII and Chapters 22, 23, and 30), student feedback, not only in formal reviews but also via [email protected] informal communication. Please let us know how we are do- Kelly Hogan (Chapters 20, 24–26, 28, and 29), [email protected] ing and how we can improve the next edition of the book.

Program Components Instructor Resource Center (IRC). Upon adoption or to preview, please go to www.pearsonschool.com/access_ request and select Instructor Resource Center. You will be required to complete a brief one-time registration subject Upon textbook purchase, students and teachers are granted to verification of educator status. Upon verification, access access to MasteringBiology. High school teachers can obtain information and instructions will be sent to you via email. preview or adoption access for MasteringBiology in one of Once logged into the IRC, enter ISBN 978-0-13-348039-9 in the following ways: the “Search our Catalog” box to locate resources. Electronic Preview Access teacher supplements are also available within the Instruc- • Teachers can request preview access online by visiting tor’s tab of MasteringBiology®. PearsonSchool.com/Access_Request, select Initial Access Instructor Resource DVD then Option 2. Preview Access information will be sent to This DVD set combines all the instructor media for Camp- the teacher via email. bell Biology: Concepts & Connections, Eighth Edition, into Adoption Access one chapter-by-chapter resource including PowerPoint® • With the purchase of this program, a Pearson Adoption slides, animations, Discovery Channel video clips, lecture Access Card with Instructor Manual will be delivered with presentations, lecture questions to stimulate class discus- your textbook purchase. (ISBN: 0-13-034391-9) sions, quiz games, and digital transparencies. The Test • Ask your sales representative for an Pearson Adoption Ac- Bank provides a variety of test questions, many art- or cess Card with Instructor Manual. (ISBN: 0-13-034391-9) scenario-based, in both TestGen® and Microsoft® Word. Selected materials from the Instructor Resources DVD can OR be downloaded from the “Instructor Resources” area of • Visit PearsonSchool.com/Access_Request, select Initial MasteringBiology®. Access then Option 3. Adoption access information will Instructor Guide be sent to the teacher via email. This comprehensive guide provides chapter-by-chapter ref- Students, ask your teacher for access. erences to all the media resources available to students. The guide also includes objectives, key terms, word roots, lecture For the Teacher outlines, student misconceptions and concerns, and teaching tips. Some of the teacher supplements and resources for this text are available electronically to qualified adopters on the

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elow are some important highlights of new content Unit II, Cellular Reproduction and Genetics The purpose of this and organizational improvements in Campbell Biology: unit is to help students understand the relationship between Concepts & Connections, Eighth Edition. DNA, chromosomes, and organisms and to help them see B that genetics is not purely hypothetical but connects in many Chapter 1, Biology: Exploring Life The snowy owl, our cover important and interesting ways to their lives, human society, organism for the Eighth Edition, is featured in the chapter and other life on Earth. In preparing this edition, we worked introduction. The discussion of the evolutionary adaptations to clarify difficult concepts, enhancing text and illustrations of these owls to life on the arctic tundra links to a new and providing timely new applications of genetic principles. Scientific Thinking module on testing the hypothesis that The content is reinforced with updated discussions of relevant camouflage coloration protects some animals from predation. topics, such as personalized cancer therapy, the H1N1 and An expanded module on evolution as the core theme of H5N1 influenza viruses, umbilical cord blood banking, and biology now includes a phylogenetic tree of elephants to the science and controversy surrounding genetically modified enhance the discussion of the unity and diversity of life. foods. This edition includes discussion of many recent advances in the field. Some new topics concern our basic understanding Unit I, The Life of the Cell Throughout the Eighth Edition, of genetics and the cell cycle, such as how sister chromatids are the themes introduced in new chapter introductions are physically attached during meiosis, how chemical modifications expanded and further explored in either Scientific Thinking such as methylation and acetylation affect inheritance, and the or Evolution Connection modules. For instance, in this roles of activators and enhancers in controlling gene expression. unit, Chapter 5, The Working Cell, begins with the question Other topics include recent advances in our understanding “How can water flow through a membrane?” and an essay of genetics, such as the analysis of recent human evolution of that describes the role these water channels play in kidney high-altitude-dwelling Sherpas, expanded roles for microRNAs function; the essay is illustrated with a computer model of in the control of genetic information, and our improved aquaporins spanning a membrane. Module 5.7, a Scientific understanding of the cellular basis of health problems in Thinking module, then details the serendipitous discovery cloned animals. In some cases, sections within chapters have of aquaporins and presents data from a study that helped been reorganized to present a more logical flow of materials. identify their function. Chapter 7, Photosynthesis: Using Examples of new organization include the discussion of human Light to Make Food, begins with the question “Will global karyotypes and the diagnosis of chromosomal abnormalities climate change make you itch?” and uses the example of (Modules 8.18–8.20) and the processes of reproductive proliferation of poison ivy to introduce this chapter on and therapeutic cloning (Modules 11.12–11.14). Material photosynthesis. Then, Module 7.13, another Scientific throughout the unit has been updated to reflect recent data, Thinking module, explores various ways that scientists test such as the latest cancer statistics and results from whole- the effects of rising atmospheric CO2 levels on plant growth and presents results from a study on poison ivy growth. genome sequencing. The Scientific Thinking question at the end of the chapter Unit III, Concepts of Evolution This unit presents the continues this theme, with data from a study on pollen basic principles of evolution and natural selection, the production by ragweed under varying CO2 concentrations, overwhelming evidence that supports these theories, beginning with the question “Will global climate change and their relevance to all of biology—and to the lives of make you sneeze as well as itch?” This unit also has three students. A new chapter introduction in Chapter 13, How of the new Visualizing the Concept modules: Module 3.14: Populations Evolve, highlights the role that evolution plays A protein’s functional shape results from four levels of in thwarting human attempts to eradicate disease. The structure; Module 5.1: Membranes are fluid mosaics of lipids chapter has been reorganized so that the opening module on and proteins with many functions; and Module 7.9: The Darwin’s development of the theory of evolution is followed light reactions take place within the thylakoid membranes. immediately by evidence for evolution, including a Scientific These modules use both new and highly revised art to Thinking module on fossils of transitional forms. Another guide students through these challenging topics in a visual, new module (13.4) assembles evidence from homologies, highly intuitive manner. Chapter 6, How Cells Harvest including an example of “pseudogenes.” New material in this Chemical Energy, now includes a new figure and expanded unit also supports our goal of directly addressing student explanation of the amazing molecular motor, ATP synthase. misconceptions about evolution. For example, a new chapter The art program in Chapter 4, A Tour of the Cell, has been introduction and Scientific Thinking module in Chapter 14, completely reimagined and revised. The beautiful new The Origin of Species, tackle the question “Can we observe diagrams of animal and plant cells and their component parts speciation occurring?” and a new chapter introduction in are designed to help students appreciate the complexities of Chapter 15, Tracing Evolutionary History, poses the question cell structure and explore the relationship between structure (answered in Module 15.12) “How do brand-new structures and function. arise by evolution?”

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xvii C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 17 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services Unit IV, The Evolution of Biological Diversity The diversity newer issues in biology. The chapter introduction and unit surveys all life on Earth in less than a hundred pages! new Scientific Thinking module in Chapter 26, Hormones Consequently, descriptions and illustrations of the unifying and the Endocrine System, discuss the consequences of characteristics of each major group of organisms, along endocrine disruptors in the environment. The Scientific with a small sample of its diversity, make up the bulk of the Thinking module in Chapter 23 describes large clinical trials content. Two recurring elements are interwoven with these investigating the hypothesis that heart attacks are caused by descriptions: evolutionary history and examples of relevance the body’s inflammatory response. Chapter 27, Reproduction to our everyday lives and society at large. For the Eighth and Embryonic Development, has a new chapter Edition, we have improved and updated those two elements. introduction on viral STDs, improved figures presenting For example, Chapter 16, Microbial Life: Prokaryotes and embryonic development, as well as a Visualizing the Concept Protists, opens with a new introduction on human microbiota module on human pregnancy. Improvements to this unit also and the question “Are antibiotics making us fat?” The related include a significant revision to the presentation of nutrition Scientific Thinking module (16.11) updates the story of in Modules 21.14 to 21.21 and a reorganization of text and Marshall’s discovery of the role of Helicobacter pylori in ulcers art in Modules 25.6 and 25.7 to guide students through the with a new hypothesis about a possible connection between anatomy and physiology of the kidneys. H. pylori and obesity. A new chapter introduction and Scientific Thinking module in Chapter 17, The Evolution of Unit VI, Plants: Form and Function To help students gain an Plant and Fungal Diversity, highlight the interdependence of appreciation of the importance of plants, this unit presents plants and fungi. The alternation of generations and the life the anatomy and physiology of angiosperms with frequent cycle in mosses and ferns are presented in an attractive two- connections to the importance of plants to society. New page Visualizing the Concept module (17.3), while Connections in this edition include an increased discussion details of the pine life cycle have been replaced with a of the importance of agriculture to human civilization new Module 17.5 that emphasizes pollen and seeds as key (including presentation of genomic data investigating this adaptations for terrestrial life. The animal diversity chapters question) in Chapter 31, issues surrounding organic farming (18, The Evolution of Invertebrate Diversity; and 19, The (including presentation of data on the nutritional value of Evolution of Vertebrate Diversity) also have new opening organic versus conventionally grown produce) in Chapter 32, essays. A Visualizing the Concept module (18.3) beautifully an expanded discussion of phytoestrogens, as well as a illustrates features of the animal body plan. A new Module new discussion on the production of seedless vegetables in 18.16 calls attention to the value of invertebrate diversity. Chapter 33. Throughout the unit, the text has been revised Chapter 19 includes a Visualizing the Concept module (19.9) with the goal of making the material more engaging and on primate diversity and also updates the story of hominin accessible to students. For example, the difficult topic of evolution, including the recently described Australopithecus transpiration is now presented in an entirely new, visual style sediba. within a Visualizing the Concept module (Module 32.3), and streamlined and simplified discussions were written for such Unit V, Animals: Form and Function This unit combines topics as the auxin hormones and phytochromes. All of these a comparative approach with an exploration of human changes are meant to make the point that human society is anatomy and physiology. Many chapters begin with an inexorably connected to the health of plants. overview of a general problem that animals face and a comparative discussion of how different animals address Unit VII, Ecology In this unit, students learn the fundamental this problem, all framed within an evolutionary context. principles of ecology and how these principles apply to For example, the introduction to Chapter 20, Unifying environmental problems. Along with a new introduction in Concepts of Animal Structure and Function, begins with each chapter, the Eighth Edition features many new photos the question “Does evolution lead to the perfect animal and two Visualizing the Concept modules (35.7 and 37.9)— form?” Module 20.1 is a new Evolution Connection that one focuses on whether animal movement is a response to discusses the long, looped laryngeal nerve in vertebrates stimuli or requires spatial learning and the other explores (using the giraffe as an example) to illustrate that a structure the interconnection of food chains and food webs. Scientific in an ancestral organism can become adapted to function Thinking modules sample the variety of approaches to in a descendant organism without being “perfected,” studying ecology, including the classic field study that led thereby combating common student misconceptions about to the concept of keystone species (37.11); the “natural evolution. The main portion of every chapter is devoted to experiment” of returning gray wolves to the Yellowstone detailed presentations of human body systems, frequently ecosystem (38.11); and the combination of historical records, illuminated by discussion of the health consequences of long-term experimentation, and modern technology to disorders in those systems. For example, Chapter 28, Nervous investigate the snowshoe hare–lynx population cycle (36.6). Systems, includes new material describing a genetic risk for The pioneering work of Rachel Carson (34.2) and Jane developing Alzheimer’s disease, the long-term consequences Goodall (35.22) is also described in Scientific Thinking of traumatic brain injury, and how some antidepressants modules. Modules that present data on human population may not be as effective at combating depression as once (36.3, 36.9–36.11), declining biodiversity (38.1), and global thought. In many areas, content has been updated to reflect climate change (38.3, 38.4) have all been updated.

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xviii C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 18 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services Acknowledgments

his Eighth Edition of Campbell Biology: Concepts Piljay for their contributions, as well as project manager for text & Connections is a result of the combined efforts permissions Alison Bruckner. S4Carlisle Publishing Services of many talented and hardworking people, and the was responsible for composition, headed by senior project edi- Tauthors wish to extend heartfelt thanks to all those who con- tor Emily Bush, with help from paging specialist Donna Healy; tributed to this and previous editions. Our work on this edi- and Precision Graphics, headed by project manager Amanda tion was shaped by input from the biologists acknowledged Bickel, was responsible for rendering new and revised illustra- in the reviewer list on pages xx–xxii, who shared with us tions. We also thank manufacturing buyer Jeffrey Sargent. their experiences teaching introductory biology and provided We thank Gary Hespenheide for creating a beautiful and specific suggestions for improving the book. Feedback from functional interior design and a stunning cover, and we are the authors of this edition’s supplements and the unsolicited again indebted to design manager Marilyn Perry for her over- comments and suggestions we received from many biologists sight and design leadership. The new Visualizing the Concept and biology students were also extremely helpful. In addition, modules benefited from her vision, as well as from the early this book has benefited in countless ways from the stimulat- input of art editor Elisheva Marcus and the continuing con- ing contacts we have had with the coauthors of Campbell tributions of artist Andrew Recher of Precision Graphics. Biology, Tenth Edition. Art editor Kelly Murphy envisioned the beautiful new cell art We wish to offer special thanks to the students and faculty throughout the book. at our teaching institutions. Marty Taylor thanks her students The value of Campbell Biology: Concepts & Connections as at Cornell University for their valuable feedback on the book. a learning tool is greatly enhanced by the hard work and cre- Eric Simon thanks his colleagues and friends at New England ativity of the authors of the supplements that accompany this College, especially within the collegium of Natural Sciences and book: Ed Zalisko (Instructor’s Guide and PowerPoint® Lecture Mathematics, for their continued support and assistance. Jean Presentations); Jean DeSaix, Tanya Smutka, Kristen Miller, Dickey thanks her colleagues at Clemson University for their and Justin Shaffer (Test Bank); Dana Kurpius (Active Reading expertise and support. And Kelly Hogan thanks her students for Guide); Robert Iwan and Amaya Garcia (Reading Quizzes and their enthusiasm and thanks her colleagues at the University of media correlations); and Shannon Datwyler (Clicker Questions North Carolina, Chapel Hill, for their continued support. and Quiz Shows). In addition to senior supplements project We thank Paul Corey, president, Science, Business, and editor Susan Berge, the editorial and production staff for the Technology, Pearson Higher Education. In addition, the supplements program included supplements production proj- superb publishing team for this edition was headed up by ac- ect manager Jane Brundage, PowerPoint® Lecture Presentations quisitions editor Alison Rodal, with the invaluable support of editor Joanna Dinsmore, and project manager Sylvia Rebert of editor-in-chief Beth Wilbur. We cannot thank them enough Progressive Publishing Alternatives. And the superlative Mas- for their unstinting efforts on behalf of the book and for their teringBiology® program for this book would not exist without commitment to excellence in biology education. We are for- Lauren Fogel, Stacy Treco, Tania Mlawer, Katie Foley, Sarah tunate to have had once again the contributions of executive Jensen, Juliana Tringali, Daniel Ross, Dario Wong, Taylor director of development Deborah Gale and executive edito- Merck, Caroline Power, and David Kokorowski and his team. rial manager Ginnie Simione Jutson. We are similarly grateful And a special thanks to Sarah Young-Dualan for her thought- to the members of the editorial development team—Debbie ful work on the Visualizing the Concepts interactive videos. Hardin, who also served as the day-to-day editorial project For their important roles in marketing the book, we are manager, and Susan Teahan—for their steadfast commitment very grateful to senior marketing manager Amee Mosley, ex- to quality. We thank them for their thoroughness, hard work, ecutive marketing manager Lauren Harp, and vice president and good humor; the book is far better than it would have of marketing Christy Lesko. We also appreciate the work of been without their efforts. Thanks also to senior supplements the executive marketing manager for MasteringBiology®, project editor Susan Berge for her oversight of the supple- Scott Dustan. The members of the Pearson Science sales team ments program and to editorial assistants Rachel Brickner, have continued to help us connect with biology instructors Katherine Harrison-Adcock, and Libby Reiser for the effi- and their teaching needs, and we thank them. cient and enthusiastic support they provided. Finally, we are deeply grateful to our families and friends This book and all the other components of the teaching for their support, encouragement, and patience throughout package are both attractive and pedagogically effective in large this project. Our special thanks to Paul, Dan, Maria, Armelle, part because of the hard work and creativity of the production and Sean (J.B.R.); Josie, Jason, Marnie, Alice, Jack, David, professionals on our team. We wish to thank managing editor Paul, Ava, and Daniel (M.R.T.); Amanda, Reed, Forest, and Mike Early and production project manager Lori Newman. We dear friends Jamey, Nick, Jim, and Bethany (E.J.S.); Jessie and also acknowledge copyeditor Joanna Dinsmore, proofreader Katherine (J.L.D.); and Tracey, Vivian, Carolyn, Brian, Jake, Pete Shanks, and indexer Lynn Armstrong. We again thank and Lexi (K.H.) senior photo editor Donna Kalal and photo researcher Kristin Jane Reece, Martha Taylor, Eric Simon, Jean Dickey, and Kelly Hogan

Acknowledgments xix

# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xix C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 19 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services Reviewers

Visualizing the Concept Review Panel, Dawn Adrian Adams, Baylor University Warren Buss, University of Northern Colorado Eighth Edition Olushola Adeyeye, Duquesne University Linda Butler, University of Texas at Austin Shylaja Akkaraju, Bronx Community College Jerry Button, Portland Community College Erica Kipp, Pace University Felix Akojie, Paducah Community College Carolee Caffrey, University of California, David Loring, Johnson County Community College Dan Alex, Chabot College Los Angeles Sheryl Love, Temple University John Aliff, Georgia Perimeter College George Cain, University of Iowa Sukanya Subramanian, Collin County Community Sylvester Allred, Northern Arizona University Beth Campbell, Itawamba Community College College Jane Aloi-Horlings, Saddleback College John Campbell, Northern Oklahoma College Jennifer J. Yeh, San Francisco, California Loren Ammerman, University of Texas at John Capeheart, University of Houston, Downtown Arlington James Cappuccino, Rockland Community College Reviewers of the Eighth Edition Dennis Anderson, Oklahoma City Community M. Carabelli, Broward Community College Steven Armstrong, Tarrant County College College Jocelyn Cash, Central Piedmont Community Michael Battaglia, Greenville Technical College Marjay Anderson, Howard University College Lisa Bonneau, Metropolitan Community College Bert Atsma, Union County College Cathryn Cates, Tyler Junior College Stephen T. Brown, Los Angeles Mission College Yael Avissar, Rhode Island College Russell Centanni, Boise State University Nancy Buschhaus, University of Tennessee at Gail Baker, LaGuardia Community College David Chambers, Northeastern University Martin Caroline Ballard, Rock Valley College Ruth Chesnut, Eastern Illinois University Glenn Cohen, Troy University Andrei Barkovskii, Georgia College and State Vic Chow, San Francisco City College Nora Espinoza, Clemson University University Van Christman, Ricks College Karen E. Francl, Radford University Mark Barnby, Ohlone College Craig Clifford, Northeastern State University, Jennifer Greenwood, University of Tennessee at Chris Barnhart, University of San Diego Tahlequah Martin Stephen Barnhart, Santa Rosa Junior College Richard Cobb, South Maine Community College Joel Hagen, Radford University William Barstow, University of Georgia Mary Colavito, Santa Monica College Chris Haynes, Shelton State Community College Kirk A. Bartholomew, Central Connecticut State Jennifer Cooper, Itawamba Community College Duane A. Hinton, Washburn University University Bob Cowling, Ouachita Technical College Amy Hollingsworth, The University of Akron Michael Battaglia, Greenville Technical College Don Cox, Miami University Erica Kipp, Pace University Gail Baughman, Mira Costa College Robert Creek, Western Kentucky University Cindy Klevickis, James Madison University Jane Beiswenger, University of Wyoming Hillary Cressey, George Mason University Dubear Kroening, University of Wisconsin, Tania Beliz, College of San Mateo Norma Criley, Illinois Wesleyan University Fox Valley Lisa Bellows, North Central Texas College Jessica Crowe, South Georgia College Dana Kurpius, Elgin Community College Ernest Benfield, Virginia Polytechnic Institute Mitch Cruzan, Portland State University Dale Lambert, Tarrant County College Rudi Berkelhamer, University of California, Irvine Judy Daniels, Monroe Community College David Loring, Johnson County Community College Harry Bernheim, Tufts University Michael Davis, Central Connecticut State Mark Meade, Jacksonville State University Richard Bliss, Yuba College University John Mersfelder, Sinclair Community College Lawrence Blumer, Morehouse College Pat Davis, East Central Community College Andrew Miller, Thomas University Dennis Bogyo, Valdosta State University Lewis Deaton, University of Louisiana Zia Nisani, Antelope Valley College Lisa K. Bonneau, Metropolitan Community Lawrence DeFilippi, Lurleen B. Wallace College Camellia M. Okpodu, Norfolk State University College, Blue River James Dekloe, Solano Community College James Rayburn, Jacksonville State University Mehdi Borhan, Johnson County Community Veronique Delesalle, Gettysburg College Ashley Rhodes, Kansas State University College Loren Denney, Southwest Missouri State Lori B. Robinson, Georgia College & State Kathleen Bossy, Bryant College University University William Bowen, University of Arkansas Jean DeSaix, University of North Carolina at Ursula Roese, University of New England at Little Rock Chapel Hill Doreen J. Schroeder, University of St. Thomas Robert Boyd, Auburn University Mary Dettman, Seminole Community College of Justin Shaffer, North Carolina A&T State Bradford Boyer, State University of New York, Florida University Suffolk County Community College Kathleen Diamond, College of San Mateo Marilyn Shopper, Johnson County Community Paul Boyer, University of Wisconsin Alfred Diboll, Macon College College William Bradshaw, Brigham Young University Jean Dickey, Clemson University Ayesha Siddiqui, Schoolcraft College Agnello Braganza, Chabot College Stephen Dina, St. Louis University Ashley Spring, Brevard Community College James Bray, Blackburn College Robert P. Donaldson, George Washington Thaxton Springfield, St. Petersburg College Peggy Brickman, University of Georgia University Linda Brooke Stabler, University of Central Chris Brinegar, San Jose State University Gary Donnermeyer, Iowa Central Community Oklahoma Chad Brommer, Emory University College Patrick Stokley, East Central Community College Charles Brown, Santa Rosa Junior College Charles Duggins, University of South Carolina Lori Tolley-Jordan, Jacksonville State University Carole Browne, Wake Forest University Susan Dunford, University of Cincinnati Jimmy Triplett, Jacksonville State University Becky Brown-Watson, Santa Rosa Junior College Lee Edwards, Greenville Technical College Lisa Weasel, Portland State University Delia Brownson, University of Texas at Austin and Betty Eidemiller, Lamar University Martin Zahn, Thomas Nelson Community College Austin Community College Jamin Eisenbach, Eastern Michigan University Michael Bucher, College of San Mateo Norman Ellstrand, University of California, Virginia Buckner, Johnson County Community Riverside Reviewers of Previous Editions College Thomas Emmel, University of Florida Michael Abbott, Westminster College Joseph C. Bundy, Jr., University of North Carolina Cindy Erwin, City College of San Francisco Tanveer Abidi, Kean University at Greensboro Gerald Esch, Wake Forest University Daryl Adams, Mankato State University Ray Burton, Germanna Community College David Essar, Winona State University

xx Reviewers

# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xx C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 20 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services Cory Etchberger, Longview Community College Janet Haynes, Long Island University Margaret Maile Lam, Kapiolani Community Nancy Eyster-Smith, Bentley College Jean Helgeson, Collin County Community College College William Ezell, University of North Carolina at Ira Herskowitz, University of California, MaryLynne LaMantia, Golden West College Pembroke San Francisco Mary Rose Lamb, University of Puget Sound Laurie Faber, Grand Rapids Community College Paul Hertz, Barnard College Dale Lambert, Tarrant County College, Northeast Terence Farrell, Stetson University Margaret Hicks, David Lipscomb University Thomas Lammers, University of Wisconsin, Shannon Kuchel Fehlberg, Colorado Christian Jean Higgins-Fonda, Prince George’s Community Oshkosh University College Carmine Lanciani, University of Florida Jerry Feldman, University of California, Phyllis Hirsch, East Los Angeles College Vic Landrum, Washburn University Santa Cruz William Hixon, St. Ambrose University Deborah Langsam, University of North Carolina Eugene Fenster, Longview Community College Carl Hoagstrom, Ohio Northern University at Charlotte Dino Fiabane, Community College of Philadelphia Kim Hodgson, Longwood College Geneen Lannom, University of Central Oklahoma Kathleen Fisher, San Diego State University Jon Hoekstra, Gainesville State College Brenda Latham, Merced College Edward Fliss, St. Louis Community College, Kelly Hogan, University of North Carolina at Liz Lawrence, Miles Community College Florissant Valley Chapel Hill Steven Lebsack, Linn-Benton Community College Linda Flora, Montgomery County Community John Holt, Michigan State University Karen Lee, University of Pittsburgh at Johnstown College Laura Hoopes, Occidental College Tom Lehman, Morgan Community College Dennis Forsythe, The Citadel Military College of Lauren Howard, Norwich University William Lemon, Southwestern Oregon Community South Carolina Robert Howe, Suffolk University College Robert Frankis, College of Charleston Michael Hudecki, State University of New York, Laurie M. Len, El Camino College James French, Rutgers University Buffalo Peggy Lepley, Cincinnati State University Bernard Frye, University of Texas at Arlington George Hudock, Indiana University Richard Liebaert, Linn-Benton Community Anne Galbraith, University of Wisconsin Kris Hueftle, Pensacola Junior College College Robert Galbraith, Crafton Hills College Barbara Hunnicutt, Seminole Community College Kevin Lien, Portland Community College Rosa Gambier, State University of New York, Brenda Hunzinger, Lake Land College Harvey Liftin, Broward Community College Suffolk County Community College Catherine Hurlbut, Florida Community College Ivo Lindauer, University of Northern Colorado George Garcia, University of Texas at Austin Charles Ide, Tulane University William Lindsay, Monterey Peninsula College Linda Gardner, San Diego Mesa College Mark Ikeda, San Bernardino Valley College Kirsten Lindstrom, Santa Rosa Junior College Sandi Gardner, Triton College Georgia Ineichen, Hinds Community College Melanie Loo, California State University, Gail Gasparich, Towson University Robert Iwan, Inver Hills Community College Sacramento Janet Gaston, Troy University Mark E. Jackson, Central Connecticut State David Loring, Johnson County Community College Shelley Gaudia, Lane Community College University Eric Lovely, Arkansas Tech University Douglas Gayou, University of Missouri at Charles Jacobs, Henry Ford Community College Paul Lurquin, Washington State University Columbia Fred James, Presbyterian College James Mack, Monmouth University Robert Gendron, Indiana University of Ursula Jander, Washburn University David Magrane, Morehead State University Pennsylvania Alan Jaworski, University of Georgia Joan Maloof, Salisbury State University Bagie George, Georgia Gwinnett College R. Jensen, Saint Mary’s College Joseph Marshall, West Virginia University Rebecca German, University of Cincinnati Robert Johnson, Pierce College, Lakewood Campus Presley Martin, Drexel University Grant Gerrish, University of Hawaii Roishene Johnson, Bossier Parish Community William McComas, University of Iowa Julie Gibbs, College of DuPage College Steven McCullagh, Kennesaw State College Frank Gilliam, Marshall University Russell Johnson, Ricks College Mitchell McGinnis, North Seattle Community Patricia Glas, The Citadel Military College of John C. Jones, Calhoun Community College College South Carolina Florence Juillerat, Indiana University at James McGivern, Gannon University David Glenn-Lewin, Wichita State University Indianapolis Colleen McNamara, Albuquerque TVI Robert Grammer, Belmont University Tracy Kahn, University of California, Riverside Community College Laura Grayson-Roselli, Burlington County College Hinrich Kaiser, Victor Valley College Caroline McNutt, Schoolcraft College Peggy Green, Broward Community College Klaus Kalthoff, University of Texas at Austin Scott Meissner, Cornell University Miriam L. Greenberg, Wayne State University Tom Kantz, California State University, Joseph Mendelson, Utah State University Sylvia Greer, City University of New York Sacramento Timothy Metz, Campbell University Eileen Gregory, Rollins College Jennifer Katcher, Pima Community College Iain Miller, University of Cincinnati Dana Griffin, University of Florida Judy Kaufman, Monroe Community College Robert Miller, University of Dubuque Richard Groover, J. Sargeant Reynolds Community Marlene Kayne, The College of New Jersey V. Christine Minor, Clemson University College Mahlon Kelly, University of Virginia Brad Mogen, University of Wisconsin, River Falls Peggy Guthrie, University of Central Oklahoma Kenneth Kerrick, University of Pittsburgh at James Moné, Millersville University Maggie Haag, University of Alberta Johnstown Jamie Moon, University of North Florida Richard Haas, California State University, Fresno Joyce Kille-Marino, College of Charleston Juan Morata, Miami Dade College Martin Hahn, William Paterson College Joanne Kilpatrick, Auburn University, Richard Mortensen, Albion College Leah Haimo, University of California, Riverside Montgomery Henry Mulcahy, Suffolk University James Hampton, Salt Lake Community College Stephen Kilpatrick, University of Pittsburgh at Christopher Murphy, James Madison University Blanche Haning, North Carolina State University Johnstown Kathryn Nette, Cuyamaca College Richard Hanke, Rose State College Lee Kirkpatrick, Glendale Community College James Newcomb, New England College Laszlo Hanzely, Northern Illinois University Peter Kish, Southwestern Oklahoma State Zia Nisani, Antelope Valley College David Harbster, Paradise Valley Community University James Nivison, Mid Michigan Community College College Cindy Klevickis, James Madison University Peter Nordloh, Southeastern Community College Sig Harden, Troy University Montgomery Robert Koch, California State University, Fullerton Stephen Novak, Boise State University Reba Harrell, Hinds Community College Eliot Krause, Seton Hall University Bette Nybakken, Hartnell College Jim Harris, Utah Valley Community College Dubear Kroening, University of Wisconsin, Michael O’Donnell, Trinity College Mary Harris, Louisiana State University Fox Valley Steven Oliver, Worcester State College Chris Haynes, Shelton State Community College Kevin Krown, San Diego State University Karen Olmstead, University of South Dakota

Reviewers xxi

# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxi C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 21 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services Steven O’Neal, Southwestern Oklahoma State Steven Roof, Fairmont State College Kathy Thompson, Louisiana State University University Jim Rosowski, University of Nebraska Laura Thurlow, Jackson Community College Lowell Orr, Kent State University Stephen Rothstein, University of California, Anne Tokazewski, Burlington County College William Outlaw, Florida State University Santa Barbara John Tolli, Southwestern College Phillip Pack, Woodbury University Donald Roush, University of North Alabama Bruce Tomlinson, State University of New York, Kevin Padian, University of California, Berkeley Lynette Rushton, South Puget Sound Community Fredonia Kay Pauling, Foothill College College Nancy Tress, University of Pittsburgh at Titusville Mark Paulissen, Northeastern State University, Connie Rye, East Mississippi Community College Donald Trisel, Fairmont State College Tahlequah Linda Sabatino, State University of New York, Kimberly Turk, Mitchell Community College Debra Pearce, Northern Kentucky University Suffolk County Community College Virginia Turner, Harper College David Pearson, Bucknell University Douglas Schamel, University of Alaska, Fairbanks Mike Tveten, Pima College Patricia Pearson, Western Kentucky University Douglas Schelhaas, University of Mary Michael Twaddle, University of Toledo Kathleen Pelkki, Saginaw Valley State University Beverly Schieltz, Wright State University Rani Vajravelu, University of Central Florida Andrew Penniman, Georgia Perimeter College Fred Schindler, Indian Hills Community College Leslie VanderMolen, Humboldt State University John Peters, College of Charleston Robert Schoch, Boston University Cinnamon VanPutte, Southwestern Illinois Gary Peterson, South Dakota State University Brian Scholtens, College of Charleston College Margaret Peterson, Concordia Lutheran College John Richard Schrock, Emporia State University Sarah VanVickle-Chavez, Washington University Russell L. Peterson, Indiana University of Julie Schroer, Bismarck State College John Vaughan, Georgetown College Pennsylvania Fayla Schwartz, Everett Community College Martin Vaughan, Indiana University Paula Piehl, Potomac State College Judy Shea, Kutztown University of Pennsylvania Mark Venable, Appalachian State University Ben Pierce, Baylor University Daniela Shebitz, Kean University Ann Vernon, St. Charles County Community Jack Plaggemeyer, Little Big Horn College Thomas Shellberg, Henry Ford Community College Barbara Pleasants, Iowa State University College Rukmani Viswanath, Laredo Community College Kathryn Podwall, Nassau Community College Cara Shillington, Eastern Michigan University Frederick W. Vogt, Elgin Community College Judith Pottmeyer, Columbia Basin College Lisa Shimeld, Crafton Hills College Mary Beth Voltura, State University of New York, Donald Potts, University of California, Santa Cruz Brian Shmaefsky, Kingwood College Cortland Nirmala Prabhu, Edison Community College Mark Shotwell, Slippery Rock University Jerry Waldvogel, Clemson University Elena Pravosudova, University of Nevada, Reno Jane Shoup, Purdue University Robert Wallace, Ripon College James Pru, Belleville Area College Michele Shuster, New Mexico State University Dennis Walsh, MassBay Community College Rongsun Pu, Kean University Linda Simpson, University of North Carolina at Patricia Walsh, University of Delaware Charles Pumpuni, Northern Virginia Community Charlotte Lisa Weasel, Portland State University College Gary Smith, Tarrant County Junior College James Wee, Loyola University Kimberly Puvalowski, Old Bridge High School Marc Smith, Sinclair Community College Harrington Wells, University of Tulsa Rebecca Pyles, East Tennessee State University Michael Smith, Western Kentucky University Jennifer Wiatrowski, Pasco-Hernando Community Shanmugavel Rajendran, Baltimore City Phil Snider, University of Houston College Community College Sam C. Sochet, Thomas Edison Career and Larry Williams, University of Houston Bob Ratterman, Jamestown Community College Technical Education High School Ray S. Williams, Appalachian State University Jill Raymond, Rock Valley College Gary Sojka, Bucknell University Lura Williamson, University of New Orleans Michael Read, Germanna Community College Ralph Sorensen, Gettysburg College Sandra Winicur, Indiana University, South Bend Brian Reeder, Morehead State University Ruth Sporer, Rutgers University Robert R. Wise, University of Wisconsin Oshkosh Bruce Reid, Kean College Linda Brooke Stabler, University of Central Mary E. Wisgirda, San Jacinto College David Reid, Blackburn College Oklahoma Mary Jo Witz, Monroe Community College Stephen Reinbold, Longview Community College David Stanton, Saginaw Valley State University Neil Woffinden, University of Pittsburgh at Erin Rempala, San Diego Mesa College Amanda Starnes, Emory University Johnstown Michael Renfroe, James Madison University John Stolz, Duquesne University Michael Womack, Macon State University Tim Revell, Mt. San Antonio College Ross Strayer, Washtenaw Community College Patrick Woolley, East Central College Douglas Reynolds, Central Washington University Donald Streuble, Idaho State University Maury Wrightson, Germanna Community College Fred Rhoades, Western Washington University Megan Stringer, Jones County Junior College Tumen Wuliji, University of Nevada, Reno John Rinehart, Eastern Oregon University Mark Sugalski, New England College Mark Wygoda, McNeese State University Laura Ritt, Burlington County College Gerald Summers, University of Missouri Tony Yates, Seminole State College Lynn Rivers, Henry Ford Community College Marshall Sundberg, Louisiana State University William Yurkiewicz, Millersville University of Bruce Robart, University of Pittsburgh at Christopher Tabit, University of West Georgia Pennsylvania Johnstown David Tauck, Santa Clara University Gregory Zagursky, Radford University Jennifer Roberts, Lewis University Hilda Taylor, Acadia University Martin Zahn, Thomas Nelson Community College Laurel Roberts, University of Pittsburgh Franklin Te, Miami Dade College Edward J. Zalisko, Blackburn College Luis A. Rodriguez, San Antonio Colleges Gene Thomas, Solano Community College David Zeigler, University of North Carolina at Duane Rohlfing, University of South Carolina Kenneth Thomas, Northern Essex Community Pembroke Jeanette Rollinger, College of the Sequoias College Uko Zylstra, Calvin College

xxii Reviewers

# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxii C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 22 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services Detailed Contents

▹ Chemical Bonds 22 1 Biology: Exploring Life 1 2.5 The distribution of electrons determines an atom’s chemical properties 22 ▹ Themes in the Study of 2.6 Covalent bonds join atoms into molecules through Biology 2 electron sharing 23 1.1 All forms of life 2.7 Ionic bonds are attractions between ions of opposite share common charge 24 properties 2 2.8 Hydrogen bonds are weak bonds important in the 1.2 In life’s hierarchy of chemistry of life 24 organization, new 2.9 Chemical reactions make and break chemical properties emerge at bonds 25 each level 3 1.3 Cells are the structural ▹ Water’s Life-Supporting Properties 26 and functional units of life 4 2.10 Hydrogen bonds make liquid water cohesive 26 1.4 Organisms interact with their environment, exchanging 2.11 Water’s hydrogen bonds moderate temperature 26 matter and energy 5 2.12 Ice floats because it is less dense than liquid water 27 ▹ Evolution, the Core Theme of Biology 6 2.13 Water is the solvent of life 27 1.5 The unity of life is based on DNA and a common 2.14 The chemistry of life is sensitive to acidic and basic genetic code 6 conditions 28 1.6 The diversity of life can be arranged into three 2.15 SCIENTIFIC THINKING Scientists study the effects of rising domains 6 atmospheric CO2 on coral reef ecosystems 28 1.7 Evolution explains the unity and diversity of life 8 2.16 EVOLUTION CONNECTION The search for extraterrestrial life centers on the search for water 29 The Process of Science 10 ▹ Chapter Review 30 1.8 In studying nature, scientists make observations and form and test hypotheses 10 1.9 SCIENTIFIC THINKING Hypotheses can be tested using controlled field studies 11 3 The Molecules of Cells 32 Biology and Everyday Life 12 ▹ ▹ Introduction to Organic Compounds 34 1.10 EVOLUTION CONNECTION Evolution is connected to our 3.1 Life’s molecular diversity is based on everyday lives 12 the properties of carbon 34 1.11 CONNECTION Biology, technology, and society are 3.2 A few chemical groups are connected in important ways 12 key to the functioning of Chapter Review 13 biological molecules 35 3.3 Cells make large molecules UNIT I from a limited set of small molecules 36 The Life of the Cell 15 ▹ Carbohydrates 37 3.4 Monosaccharides are the simplest carbohydrates 37 3.5 Two monosaccharides are linked to form a 2 The Chemical Basis of Life 16 disaccharide 38 3.6 CONNECTION What is high-fructose corn syrup, and is it to ▹ Elements, Atoms, and Compounds 18 blame for obesity? 38 2.1 Organisms are composed of 3.7 Polysaccharides are long chains of sugar units 39 elements, in combinations called compounds 18 ▹ Lipids 40 2.2 CONNECTION Trace elements are 3.8 Fats are lipids that are mostly energy-storage common additives to food and molecules 40 water 19 3.9 SCIENTIFIC THINKING Scientific studies document the health 2.3 Atoms consist of protons, risks of trans fats 41 neutrons, and electrons 20 3.10 Phospholipids and steroids are important lipids with a 2.4 CONNECTION Radioactive isotopes can help or harm variety of functions 42 us 21 3.11 CONNECTION Anabolic steroids pose health risks 42

Detailed Contents xxiii

# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxiii C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 23 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services ▹ Proteins 43 4.17 SCIENTIFIC THINKING Scientists discovered the 3.12 Proteins have a wide range of functions and cytoskeleton using the tools of biochemistry structures 43 and microscopy 66 4.18 3.13 Proteins are made from amino acids linked by peptide Cilia and flagella move when microtubules bend 66 bonds 44 4.19 The extracellular matrix of animal cells functions in 3.14 VISUALIZING THE CONCEPT A protein’s functional shape support and regulation 67 results from four levels of structure 45 4.20 Three types of cell junctions are found in animal tissues 68 46 ▹ Nucleic Acids 4.21 Cell walls enclose and support plant cells 68 3.15 DNA and RNA are the two types of nucleic acids 46 4.22 Review: Eukaryotic cell structures can be grouped on 3.16 Nucleic acids are polymers of nucleotides 46 the basis of four main functions 69 3.17 EVOLUTION CONNECTION Lactose tolerance is a recent event Chapter Review 70 in human evolution 47 Chapter Review 48 5 The Working Cell 72

4 A Tour of the Cell 50 ▹ Membrane Structure and Function 74 5.1 VISUALIZING THE CONCEPT ▹ Introduction to the Cell 52 Membranes are fluid 4.1 Microscopes reveal the world of mosaics of lipids and the cell 52 proteins with many 4.2 The small size of cells relates functions 74 to the need to exchange 5.2 EVOLUTION CONNECTION The materials across the plasma spontaneous formation membrane 54 of membranes was a 4.3 Prokaryotic cells are structurally critical step in the origin simpler than eukaryotic of life 75 cells 55 5.3 Passive transport is 4.4 Eukaryotic cells are diffusion across a partitioned into functional membrane with no energy compartments 56 investment 75 5.4 Osmosis is the diffusion of water across a ▹ The Nucleus and Ribosomes 58 membrane 76 4.5 The nucleus contains the cell’s genetic instructions 58 5.5 Water balance between cells and their surroundings is 4.6 Ribosomes make proteins for use in the cell and for crucial to organisms 76 export 59 5.6 Transport proteins can facilitate diffusion across ▹ The Endomembrane System 59 membranes 77 4.7 Many organelles are connected in the endomembrane 5.7 SCIENTIFIC THINKING Research on another membrane system 59 protein led to the discovery of aquaporins 78 4.8 The endoplasmic reticulum is a biosynthetic 5.8 Cells expend energy in the active transport of a workshop 60 solute 78 4.9 The Golgi apparatus modifies, sorts, and ships cell 5.9 Exocytosis and endocytosis transport large molecules products 61 across membranes 79 4.10 Lysosomes are digestive compartments within a ▹ Energy and the Cell 80 cell 62 5.10 Cells transform energy as they perform work 80 4.11 Vacuoles function in the general maintenance of the 5.11 Chemical reactions either release or store energy 81 cell 62 5.12 ATP drives cellular work by coupling exergonic and 4.12 A review of the structures involved in manufacturing endergonic reactions 82 and breakdown 63 ▹ How Enzymes Function 83 ▹ Energy-Converting Organelles 63 5.13 Enzymes speed up the cell’s chemical reactions by 4.13 Mitochondria harvest chemical energy from food 63 lowering energy barriers 83 4.14 Chloroplasts convert solar energy to chemical 5.14 A specific enzyme catalyzes each cellular energy 64 reaction 84 4.15 EVOLUTION CONNECTION Mitochondria and chloroplasts 5.15 Enzyme inhibition can regulate enzyme activity in a evolved by endosymbiosis 64 cell 85 ▹ The Cytoskeleton and Cell Surfaces 65 5.16 CONNECTION Many drugs, pesticides, and poisons are 4.16 The cell’s internal skeleton helps organize its structure enzyme inhibitors 85 and activities 65 Chapter Review 86

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxiv C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 24 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services 7.3 Scientists traced the process of photosynthesis using 6 How Cells Harvest Chemical isotopes 110 7.4 Photosynthesis is a redox process, as is cellular Energy 88 respiration 110 7.5 The two stages of photosynthesis are linked by ATP and ▹ Cellular Respiration: Aerobic Harvesting NADPH 111 of Energy 90 6.1 Photosynthesis and cellular ▹ The Light Reactions: Converting Solar Energy to Chemical respiration provide energy for Energy 112 life 90 7.6 Visible radiation absorbed by pigments drives the light reactions 112 6.2 Breathing supplies O2 for use in cellular respiration and 7.7 Photosystems capture solar energy 113 removes CO2 90 7.8 Two photosystems connected by an electron transport 6.3 Cellular respiration banks energy chain generate ATP and NADPH 114 in ATP molecules 91 7.9 VISUALIZING THE CONCEPT The light reactions take place 6.4 CONNECTION The human body uses within the thylakoid membranes 115 energy from ATP for all its activities 91 ▹ The Calvin Cycle: Reducing CO2 to Sugar 116 6.5 Cells capture energy from electrons “falling” from organic fuels to oxygen 92 7.10 ATP and NADPH power sugar synthesis in the Calvin cycle 116 ▹ Stages of Cellular Respiration 93 7.11 EVOLUTION CONNECTION Other methods of carbon fixation 6.6 Overview: Cellular respiration occurs in three main have evolved in hot, dry climates 117 stages 93 ▹ The Global Significance of Photosynthesis 118 6.7 Glycolysis harvests chemical energy by oxidizing glucose to pyruvate 94 7.12 Photosynthesis makes sugar from CO2 and H2O, providing food and O2 for almost all living 6.8 Pyruvate is oxidized in preparation for the citric acid organisms 118 cycle 96 7.13 SCIENTIFIC THINKING Rising atmospheric levels of carbon 6.9 The citric acid cycle completes the oxidation of organic dioxide and global climate change will affect plants in molecules, generating many NADH and FADH2 various ways 119 molecules 96 7.14 Scientific research and international treaties have 6.10 Most ATP production occurs by oxidative helped slow the depletion of Earth’s ozone layer 120 phosphorylation 98 Chapter Review 121 6.11 SCIENTIFIC THINKING Scientists have discovered heat- producing, calorie-burning brown fat in adults 99 6.12 Review: Each molecule of glucose yields many molecules of ATP 100 UNIT II ▹ Fermentation: Anaerobic Harvesting of Energy 100 6.13 Fermentation enables cells to produce ATP without Cellular Reproduction oxygen 100 and Genetics 123 6.14 EVOLUTION CONNECTION Glycolysis evolved early in the history of life on Earth 102 ▹ Connections Between Metabolic Pathways 102 8 The Cellular Basis of Reproduction 6.15 Cells use many kinds of organic molecules as fuel for cellular respiration 102 and Inheritance 124 6.16 Organic molecules from food provide raw materials for biosynthesis 103 ▹ Cell Division and Reproduction 126 Chapter Review 104 8.1 Cell division plays many important roles in the lives of organisms 126 7 Photosynthesis: Using Light to Make 8.2 Prokaryotes reproduce by binary Food 106 fission 127 ▹ The Eukaryotic Cell Cycle and ▹ An Introduction to Mitosis 128 Photosynthesis 108 8.3 The large, complex chromosomes 7.1 Photosynthesis fuels the of eukaryotes duplicate with each cell biosphere 108 division 128 7.2 Photosynthesis occurs 8.4 The cell cycle includes growing and division in chloroplasts in plant phases 129 cells 109 8.5 Cell division is a continuum of dynamic changes 130

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxv C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 25 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services 8.6 Cytokinesis differs for plant and animal cells 132 9.10 CONNECTION New technologies can provide insight into 8.7 Anchorage, cell density, and chemical growth factors one’s genetic legacy 164 affect cell division 133 ▹ Variations on Mendel’s Laws 166 8.8 Growth factors signal the cell cycle control system 134 9.11 Incomplete dominance results in intermediate 8.9 CONNECTION Growing out of control, cancer cells produce phenotypes 166 malignant tumors 135 9.12 Many genes have more than two alleles in the 8.10 SCIENTIFIC THINKING Tailoring treatment to each patient population 167 may improve cancer therapy 136 9.13 A single gene may affect many phenotypic ▹ Meiosis and Crossing Over 136 characters 168 8.11 Chromosomes are matched in homologous pairs 136 9.14 A single character may be influenced by many 8.12 Gametes have a single set of chromosomes 137 genes 169 9.15 8.13 Meiosis reduces the chromosome number from diploid The environment affects many characters 170 to haploid 138 ▹ The Chromosomal Basis of Inheritance 170 8.14 VISUALIZING THE CONCEPT Mitosis and meiosis have 9.16 Chromosome behavior accounts for Mendel’s laws 170 important similarities and differences 140 9.17 SCIENTIFIC THINKING Genes on the same chromosome tend 8.15 Independent orientation of chromosomes in meiosis to be inherited together 172 and random fertilization lead to varied offspring 141 9.18 Crossing over produces new combinations of 8.16 Homologous chromosomes may carry different versions alleles 172 of genes 142 9.19 Geneticists use crossover data to map genes 174 8.17 Crossing over further increases genetic variability 142 ▹ Sex Chromosomes and Sex-Linked Genes 174 ▹ Alterations of Chromosome Number and Structure 144 9.20 Chromosomes determine sex in many species 174 8.18 Accidents during meiosis can alter chromosome 9.21 Sex-linked genes exhibit a unique pattern of number 144 inheritance 176 8.19 A karyotype is a photographic inventory of an 9.22 CONNECTION Human sex-linked disorders affect mostly individual’s chromosomes 145 males 177 8.20 CONNECTION An extra copy of chromosome 21 causes 9.23 EVOLUTION CONNECTION The Y chromosome provides clues Down syndrome 146 about human male evolution 177 8.21 CONNECTION Abnormal numbers of sex chromosomes do Chapter Review 178 not usually affect survival 147 8.22 EVOLUTION CONNECTION New species can arise from errors in cell division 147 10 Molecular Biology of the Gene 180 8.23 CONNECTION Alterations of chromosome structure can cause birth defects and cancer 148 ▹ The Structure of the Genetic Material 182 Chapter Review 149 10.1 SCIENTIFIC THINKING Experiments showed that DNA is the genetic Patterns of Inheritance 152 material 182 9 10.2 DNA and RNA are polymers of nucleotides 184 Mendel’s Laws 154 ▹ 10.3 DNA is a double-stranded 9.1 The study of genetics has ancient helix 186 roots 154 9.2 The science of genetics began in an ▹ DNA Replication 188 abbey garden 154 10.4 DNA replication depends on 9.3 Mendel’s law of segregation specific base pairing 188 describes the inheritance of a single 10.5 DNA replication proceeds character 156 in two directions at many sites 9.4 Homologous chromosomes bear the alleles for each simultaneously 188 character 157 ▹ The Flow of Genetic Information from DNA to RNA to 9.5 The law of independent assortment is revealed by Protein 190 tracking two characters at once 158 10.6 Genes control phenotypic traits through the expression 9.6 Geneticists can use a testcross to determine unknown of proteins 190 genotypes 159 10.7 Genetic information written in codons is translated into 9.7 Mendel’s laws reflect the rules of probability 160 amino acid sequences 191 9.8 VISUALIZING THE CONCEPT Genetic traits in humans can be 10.8 The genetic code dictates how codons are translated tracked through family pedigrees 161 into amino acids 192 9.9 CONNECTION Many inherited traits in humans are 10.9 VISUALIZING THE CONCEPT Transcription produces genetic controlled by a single gene 162 messages in the form of RNA 193

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxvi C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 26 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services 10.10 Eukaryotic RNA is processed before leaving the nucleus ▹ Cloning of Plants and Animals 221 as mRNA 194 11.12 Plant cloning shows that differentiated cells may retain 10.11 Transfer RNA molecules serve as interpreters during all of their genetic potential 221 translation 194 11.13 SCIENTIFIC THINKING Biologists can clone animals via 10.12 Ribosomes build polypeptides 196 nuclear transplantation 222 10.13 An initiation codon marks the start of an mRNA 11.14 CONNECTION Therapeutic cloning can produce stem cells message 196 with great medical potential 223 10.14 Elongation adds amino acids to the polypeptide chain until a stop codon terminates translation 197 ▹ The Genetic Basis of Cancer 224 11.15 10.15 Review: The flow of genetic information in the cell is Cancer results from mutations in genes that control cell division 224 DNA S RNA S protein 198 11.16 10.16 Mutations can affect genes 199 Multiple genetic changes underlie the development of cancer 225 ▹ The Genetics of Viruses and Bacteria 200 11.17 Faulty proteins can interfere with normal signal 10.17 Viral DNA may become part of the host transduction pathways 226 chromosome 200 11.18 CONNECTION Lifestyle choices can reduce the risk of 10.18 CONNECTION Many viruses cause disease in animals and cancer 227 plants 201 Chapter Review 228 10.19 EVOLUTION CONNECTION Emerging viruses threaten human health 202 DNA Technology and 10.20 The AIDS virus makes DNA on an RNA 12 template 203 Genomics 230 10.21 Viroids and prions are formidable pathogens in plants and animals 203 ▹ Gene Cloning 232 10.22 Bacteria can transfer DNA in three ways 204 12.1 Genes can be cloned in 10.23 Bacterial plasmids can serve as carriers for gene recombinant plasmids 232 transfer 205 12.2 VISUALIZING THE CONCEPT Enzymes Chapter Review 206 are used to “cut and paste” DNA 234 12.3 Cloned genes can be stored in 11 How Genes Are Controlled 208 genomic libraries 235 12.4 Reverse transcriptase can help ▹ Control of Gene Expression 210 make genes for cloning 235 11.1 Proteins interacting with DNA turn 12.5 Nucleic acid probes identify prokaryotic genes on or off in response clones carrying specific genes 236 to environmental changes 210 ▹ Genetically Modified Organisms 236 11.2 Chromosome structure and 12.6 Recombinant cells and organisms can mass-produce chemical modifications can affect gene products 236 gene expression 212 12.7 CONNECTION DNA technology has changed the 11.3 Complex assemblies of pharmaceutical industry and medicine 238 proteins control eukaryotic 12.8 CONNECTION Genetically modified organisms are transcription 214 transforming agriculture 239 11.4 Eukaryotic RNA may be spliced in 12.9 SCIENTIFIC THINKING Genetically modified organisms raise more than one way 214 health concerns 240 11.5 Small RNAs play multiple roles in controlling gene 12.10 CONNECTION Gene therapy may someday help treat a expression 215 variety of diseases 241 11.6 Later stages of gene expression are also subject to regulation 216 ▹ DNA Profiling 242 11.7 VISUALIZING THE CONCEPT Multiple mechanisms regulate 12.11 The analysis of genetic markers can produce a DNA gene expression in eukaryotes 217 profile 242 11.8 Cell signaling and waves of gene expression direct 12.12 The PCR method is used to amplify DNA animal development 218 sequences 242 11.9 CONNECTION Scientists use DNA microarrays to test for 12.13 Gel electrophoresis sorts DNA molecules by size 243 the transcription of many genes at once 219 12.14 Short tandem repeat analysis is commonly used for 11.10 Signal transduction pathways convert messages DNA profiling 244 received at the cell surface to responses within 12.15 CONNECTION DNA profiling has provided evidence in the cell 220 many forensic investigations 245 11.11 EVOLUTION CONNECTION Cell-signaling systems appeared 12.16 RFLPs can be used to detect differences in DNA early in the evolution of life 220 sequences 246

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxvii C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 27 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services ▹ Genomics 247 13.16 EVOLUTION CONNECTION The evolution of drug- 12.17 Genomics is the scientific study of whole genomes 247 resistant microorganisms is a serious public health concern 272 12.18 CONNECTION The Human Genome Project revealed 13.17 that most of the human genome does not consist of Diploidy and balancing selection preserve genetic genes 248 variation 272 13.18 12.19 The whole-genome shotgun method of sequencing a Natural selection cannot fashion perfect genome can provide a wealth of data quickly 249 organisms 273 Chapter Review 274 12.20 Proteomics is the scientific study of the full set of proteins encoded by a genome 249 12.21 EVOLUTION CONNECTION Genomes hold clues to human evolution 250 14 The Origin of Species 276 Chapter Review 250 ▹ Defining Species 278 14.1 The origin of species is the source of biological diversity 278 UNIT III 14.2 There are several ways to define a species 278 Concepts of Evolution 253 14.3 VISUALIZING THE CONCEPT Reproductive barriers keep species separate 280 ▹ Mechanisms of Speciation 282 13 How Populations Evolve 254 14.4 In allopatric speciation, geographic isolation leads to speciation 282 ▹ Darwin’s Theory of 14.5 Reproductive barriers can evolve as Evolution 256 populations diverge 283 13.1 A sea voyage helped 14.6 Sympatric speciation takes place without geographic Darwin frame his theory isolation 284 of evolution 256 14.7 EVOLUTION CONNECTION The origin of most plant species 13.2 The study of fossils can be traced to polyploid speciation 285 provides strong evidence 14.8 Isolated islands are often showcases of speciation 286 for evolution 258 14.9 SCIENTIFIC THINKING Lake Victoria is a living laboratory for 13.3 SCIENTIFIC THINKING Fossils studying speciation 287 of transitional forms support Darwin’s theory 14.10 Hybrid zones provide opportunities to study of evolution 259 reproductive isolation 288 13.4 Homologies provide strong evidence for evolution 260 14.11 Speciation can occur rapidly or slowly 289 13.5 Homologies indicate patterns of descent that can be Chapter Review 290 shown on an evolutionary tree 261 13.6 Darwin proposed natural selection as the mechanism of evolution 262 15 Tracing Evolutionary History 292 13.7 Scientists can observe natural selection in action 263 ▹ The Evolution of Populations 264 ▹ Early Earth and the Origin of Life 294 15.1 13.8 Mutation and sexual reproduction produce the genetic Conditions on early Earth made the variation that makes evolution possible 264 origin of life possible 294 15.2 SCIENTIFIC THINKING 13.9 Evolution occurs within populations 265 Experiments show that the abiotic synthesis 13.10 The Hardy-Weinberg equation can test whether a of organic molecules is population is evolving 266 possible 295 13.11 CONNECTION The Hardy-Weinberg equation is useful in 15.3 Stages in the origin of the first cells public health science 267 probably included the formation ▹ Mechanisms of Microevolution 268 of polymers, protocells, and self- replicating RNA 296 13.12 Natural selection, genetic drift, and gene flow can cause microevolution 268 ▹ Major Events in the History of Life 297 13.13 Natural selection is the only mechanism that 15.4 The origins of single-celled and multicellular organisms consistently leads to adaptive evolution 269 and the colonization of land were key events in life’s 13.14 Natural selection can alter variation in a population in history 297 three ways 270 15.5 The actual ages of rocks and fossils mark geologic 13.15 Sexual selection may lead to phenotypic differences time 298 between males and females 271 15.6 The fossil record documents the history of life 298

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxviii C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 28 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services ▹ Mechanisms of Macroevolution 300 16.8 Archaea thrive in extreme environments—and in other 15.7 Continental drift has played a major role in habitats 326 macroevolution 300 16.9 Bacteria include a diverse assemblage of 15.8 CONNECTION Plate tectonics may imperil human prokaryotes 326 life 302 16.10 CONNECTION Some bacteria cause disease 328 15.9 During mass extinctions, large numbers of species are 16.11 SCIENTIFIC THINKING Stomach microbiota affect health and lost 302 disease 328 15.10 Adaptive radiations have increased the diversity of ▹ Protists 330 life 304 16.12 Protists are an extremely diverse assortment of 15.11 Genes that control development play a major role in eukaryotes 330 evolution 304 16.13 EVOLUTION CONNECTION Endosymbiosis of unicellular algae 15.12 EVOLUTION CONNECTION Novel traits may arise in several is the key to much of protist diversity 331 ways 306 16.14 The “SAR” supergroup represents the range of protist 15.13 Evolutionary trends do not mean that evolution is goal diversity 332 directed 307 16.15 CONNECTION Can algae provide a renewable source of ▹ Phylogeny and the Tree of Life 308 energy? 334 15.14 Phylogenies based on homologies reflect evolutionary 16.16 Some excavates have modified mitochondria 334 history 308 16.17 Unikonts include protists that are closely related to 15.15 Systematics connects classification with evolutionary fungi and animals 335 history 308 16.18 Archaeplastids include red algae, green algae, and land 15.16 Shared characters are used to construct phylogenetic plants 336 trees 310 16.19 EVOLUTION CONNECTION Multicellularity evolved several 15.17 An organism’s evolutionary history is documented in its times in eukaryotes 337 genome 312 Chapter Review 338 15.18 Molecular clocks help track evolutionary time 313 15.19 Constructing the tree of life is a work in progress 314 Chapter Review 315 17 The Evolution of Plant and Fungal Diversity 340

UNIT IV ▹ Plant Evolution and Diversity 342 17.1 Plants have adaptations for life on The Evolution of Biological land 342 17.2 Plant diversity reflects the Diversity 317 evolutionary history of the plant kingdom 344 ▹ Alternation of Generations and 16 Microbial Life: Prokaryotes and Plant Life Cycles 346 Protists 318 17.3 VISUALIZING THE CONCEPT Haploid and diploid generations ▹ Prokaryotes 320 alternate in plant life cycles 346 16.1 Prokaryotes are diverse and widespread 320 17.4 Seedless vascular plants dominated vast “coal forests” 348 16.2 External features contribute to the success of 17.5 Pollen and seeds are key adaptations for life on prokaryotes 320 land 348 16.3 Populations of prokaryotes 17.6 The flower is the centerpiece of angiosperm can adapt rapidly to changes reproduction 350 in the environment 322 17.7 The angiosperm plant is a sporophyte with 16.4 Prokaryotes have gametophytes in its flowers 350 unparalleled nutritional 17.8 The structure of a fruit reflects its function in seed diversity 323 dispersal 352 16.5 CONNECTION Biofilms are complex associations of 17.9 CONNECTION Angiosperms sustain us—and add spice to microbes 324 our diets 352 16.6 CONNECTION Prokaryotes help clean up the 17.10 EVOLUTION CONNECTION Pollination by animals has environment 324 influenced angiosperm evolution 353 16.7 Bacteria and archaea are the two main branches of 17.11 CONNECTION Plant diversity is vital to the future of the prokaryotic evolution 325 world’s food supply 354

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxix C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 29 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services ▹ Diversity of Fungi 355 18.14 Echinoderms have spiny skin, an endoskeleton, and a 17.12 Fungi absorb food after digesting it outside their water vascular system for movement 383 bodies 355 18.15 Our own phylum, Chordata, is distinguished by four 17.13 Fungi produce spores in both asexual and sexual life features 384 cycles 356 18.16 CONNECTION Invertebrate diversity is a valuable but 17.14 Fungi are classified into five groups 356 threatened resource 385 Chapter Review 386 17.15 CONNECTION Fungi have enormous ecological benefits 358 17.16 CONNECTION Fungi have many practical uses 358 17.17 Lichens are symbiotic associations of fungi and 19 The Evolution of Vertebrate photosynthetic organisms 359 Diversity 388 17.18 SCIENTIFIC THINKING Mycorrhizae may have helped plants colonize land 360 Vertebrate Evolution and Diversity 390 17.19 CONNECTION Parasitic fungi harm plants and ▹ animals 361 19.1 Derived characters define the major clades of Chapter Review 362 chordates 390 19.2 Hagfishes and lampreys lack hinged jaws 391 19.3 Jawed vertebrates with 18 The Evolution of Invertebrate gills and paired fins include sharks, ray-finned Diversity 364 fishes, and lobe-finned fishes 392 ▹ Animal Evolution and Diversity 366 19.4 EVOLUTION CONNECTION New 18.1 What is an animal? 366 fossil discoveries are filling 18.2 Animal diversification in the gaps of tetrapod began more than half a evolution 394 billion years ago 367 19.5 Amphibians are tetrapods— 18.3 VISUALIZING THE CONCEPT vertebrates with two pairs Animals can be of limbs 396 characterized by basic 19.6 Reptiles are amniotes—tetrapods with features of their “body a terrestrially adapted egg 397 plan” 368 19.7 Birds are feathered reptiles with adaptations for 18.4 Body plans and molecular flight 398 comparisons of animals 19.8 Mammals are amniotes that have hair and produce can be used to build milk 399 phylogenetic trees 369 ▹ Primate Diversity 400 ▹ Invertebrate Diversity 370 19.9 VISUALIZING THE CONCEPT Primates include lemurs, 18.5 Sponges have a relatively monkeys, and apes 400 simple, porous 19.10 The human story begins with our primate body 370 heritage 402 18.6 Cnidarians are radial animals with tentacles and ▹ Hominin Evolution 403 stinging cells 371 19.11 The hominin branch of the primate tree includes 18.7 Flatworms are the simplest species that coexisted 403 bilateral animals 372 19.12 Australopiths were bipedal and had small 18.8 Nematodes have a body cavity and a complete digestive brains 404 tract 373 19.13 Larger brains mark the evolution of Homo 405 18.9 Diverse molluscs are variations on a common body 19.14 From origins in Africa, Homo sapiens spread around the plan 374 world 406 18.10 Annelids are segmented worms 376 19.15 SCIENTIFIC THINKING New discoveries raise new questions 18.11 Arthropods are segmented animals with jointed about the history of hominins 406 appendages and an exoskeleton 378 19.16 EVOLUTION CONNECTION Human skin color reflects 18.12 EVOLUTION CONNECTION Insects are the most successful adaptations to varying amounts of sunlight 407 group of animals 380 19.17 CONNECTION Our knowledge of animal diversity is far 18.13 SCIENTIFIC THINKING The genes that build animal bodies are from complete 408 ancient 382 Chapter Review 409

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxx C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 30 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services UNIT V ▹ The Human Digestive System 433 21.4 The human digestive system consists of an alimentary Animals: Form and canal and accessory glands 433 Function 411 21.5 Digestion begins in the oral cavity 434 21.6 After swallowing, peristalsis moves food through the esophagus to the stomach 434 21.7 CONNECTION The Heimlich maneuver can save 20 Unifying Concepts of Animal lives 435 Structure and Function 412 21.8 The stomach stores food and breaks it down with acid and enzymes 436 ▹ Structure and Function in 21.9 CONNECTION Digestive ailments include acid reflux and Animal Tissues 414 gastric ulcers 437 20.1 EVOLUTION CONNECTION An 21.10 The small intestine is the major organ of chemical animal’s form is not the digestion and nutrient absorption 438 perfect design 414 21.11 The liver processes and detoxifies blood from the 20.2 Structure fits function at all levels intestines 440 of organization in the animal 21.12 The large intestine reclaims water and compacts the body 415 feces 440 20.3 Tissues are groups of cells 21.13 EVOLUTION CONNECTION Evolutionary adaptations of with a common structure and vertebrate digestive systems relate to diet 441 function 416 20.4 Epithelial tissue covers the body and ▹ Nutrition 442 lines its organs and cavities 416 21.14 An animal’s diet must provide sufficient energy 442 20.5 Connective tissue binds and supports 21.15 An animal’s diet must supply essential nutrients 443 other tissues 417 21.16 A proper human diet must include sufficient vitamins 20.6 Muscle tissue functions in movement 418 and minerals 444 20.7 Nervous tissue forms a communication network 418 21.17 CONNECTION Food labels provide nutritional information 446 Organs and Organ Systems 419 ▹ 21.18 CONNECTION Dietary deficiencies can have a number of 20.8 Organs are made up of tissues 419 causes 446 20.9 CONNECTION Bioengineers are learning to produce organs 21.19 EVOLUTION CONNECTION The human health problem of for transplants 419 obesity may reflect our evolutionary past 447 20.10 Organ systems work together to perform life’s 21.20 SCIENTIFIC THINKING Scientists use a variety of methods to functions 420 test weight-loss claims 448 20.11 The integumentary system protects the body 422 21.21 CONNECTION Diet can influence risk of cardiovascular 20.12 SCIENTIFIC THINKING Well-designed studies help answer disease and cancer 449 scientific questions 423 Chapter Review 450 ▹ External Exchange and Internal Regulation 424 20.13 Structural adaptations enhance exchange with the environment 424 22 Gas Exchange 452 20.14 Animals regulate their internal environment 425 ▹ Mechanisms of Gas Exchange 454 20.15 Homeostasis depends on negative feedback 426 22.1 Gas exchange in humans Chapter Review 426 involves breathing, transport of gases, and exchange with body 21 Nutrition and Digestion 428 cells 454

22.2 Animals exchange O2 and ▹ Obtaining and Processing CO2 across moist body Food 430 surfaces 454 21.1 Animals obtain and 22.3 VISUALIZING THE CONCEPT ingest their food in a Gills are adapted for gas variety of ways 430 exchange in aquatic environments 456 21.2 Overview: Food 22.4 The tracheal system of insects provides direct exchange processing occurs in between the air and body cells 457 four stages 431 22.5 EVOLUTION CONNECTION The evolution of lungs facilitated 21.3 Digestion occurs in specialized compartments 432 the movement of tetrapods onto land 458

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxxi C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 31 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services ▹ The Human Respiratory System 458 22.6 In mammals, branching tubes convey air to lungs 24 The Immune System 484 located in the chest cavity 458 22.7 SCIENTIFIC THINKING Warning: Cigarette smoking is ▹ Innate Immunity 486 hazardous to your health 460 24.1 All animals have innate 22.8 Negative pressure breathing ventilates your immunity 486 lungs 460 24.2 Inflammation mobilizes the 22.9 Breathing is automatically controlled 461 innate immune response 487 ▹ Transport of Gases in the Human Body 462 ▹ Adaptive Immunity 488 22.10 Blood transports respiratory gases 462 24.3 The adaptive immune response counters specific invaders 488 22.11 Hemoglobin carries O2, helps transport CO2, and buffers the blood 462 24.4 The lymphatic system becomes a crucial battleground during infection 489 22.12 CONNECTION The human fetus exchanges gases with the mother’s blood 463 24.5 Lymphocytes mount a dual defense 490 Chapter Review 464 24.6 Antigen receptors and antibodies bind to specific regions on an antigen 491 24.7 VISUALIZING THE CONCEPT Clonal selection mobilizes defensive forces against specific antigens 492 23 Circulation 466 24.8 The primary and secondary responses differ in speed, strength, and duration 493 ▹ Circulatory Systems 468 24.9 The structure of an antibody matches its function 494 23.1 Circulatory systems 24.10 CONNECTION Antibodies are powerful tools in the lab and facilitate exchange with clinic 495 all body tissues 468 24.11 SCIENTIFIC THINKING Scientists measure antibody levels to 23.2 EVOLUTION CONNECTION look for waning immunity after HPV vaccination 496 Vertebrate cardiovascular 24.12 Helper T cells stimulate the humoral and cell-mediated systems reflect evolution 469 immune responses 497 ▹ The Human Cardiovascular System 24.13 Cytotoxic T cells destroy infected body cells 498 and Heart 470 24.14 CONNECTION HIV destroys helper T cells, compromising 23.3 VISUALIZING THE CONCEPT The human cardiovascular the body’s defenses 498 system illustrates the double circulation of 24.15 EVOLUTION CONNECTION The rapid evolution of HIV mammals 470 complicates AIDS treatment 499 23.4 The heart contracts and relaxes rhythmically 471 24.16 The immune system depends on our molecular 23.5 The SA node sets the tempo of the heartbeat 472 fingerprints 500 23.6 SCIENTIFIC THINKING What causes heart attacks? 473 ▹ Disorders of the Immune System 500 ▹ Structure and Function of Blood Vessels 474 24.17 CONNECTION Immune system disorders result from self- directed or underactive responses 500 23.7 The structure of blood vessels fits their functions 474 24.18 CONNECTION Allergies are overreactions to certain environmental antigens 501 23.8 Blood pressure and velocity reflect the structure and arrangement of blood vessels 475 Chapter Review 502 23.9 CONNECTION Measuring blood pressure can reveal cardiovascular problems 476 23.10 Smooth muscle controls the distribution of blood 477 25 Control of Body Temperature and 23.11 Capillaries allow the transfer of substances through Water Balance 504 their walls 478 ▹ Structure and Function of Blood 479 ▹ Thermoregulation 506 25.1 23.12 Blood consists of red and white blood cells suspended An animal’s regulation of body in plasma 479 temperature helps maintain homeostasis 506 23.13 CONNECTION Too few or too many red blood cells can be unhealthy 480 25.2 Thermoregulation involves adaptations 23.14 Blood clots plug leaks when blood vessels are that balance heat gain and injured 480 loss 506 23.15 CONNECTION Stem cells offer a potential cure for blood cell 25.3 SCIENTIFIC THINKING Coordinated waves diseases 481 of movement in huddles help penguins Chapter Review 482 thermoregulate 507

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxxii C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 32 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services ▹ Osmoregulation and Excretion 508 27.2 Sexual reproduction results in the generation of genetically unique offspring 534 25.4 Animals balance their levels of water and solutes through osmoregulation 508 ▹ Human Reproduction 536 25.5 EVOLUTION CONNECTION Several ways to dispose of 27.3 The human female reproductive system includes the nitrogenous wastes have evolved in animals 509 ovaries and structures that deliver gametes 536 25.6 The urinary system plays several major roles in 27.4 The human male reproductive system includes the homeostasis 510 testes and structures that deliver gametes 538 25.7 Reabsorption and secretion refine the filtrate 512 27.5 The formation of sperm and egg cells requires 25.8 Hormones regulate the urinary system 513 meiosis 540 25.9 CONNECTION Kidney dialysis can save lives 513 27.6 Hormones synchronize cyclic changes in the ovary and Chapter Review 514 uterus 542 27.7 SCIENTIFIC THINKING Sexual activity can transmit disease 544 Hormones and the Endocrine 27.8 CONNECTION Contraception can prevent unwanted 26 pregnancy 545 System 516 ▹ Principles of Embryonic Development 546 The Nature of Chemical 27.9 Fertilization results in a zygote and triggers embryonic ▹ development 546 Regulation 518 27.10 26.1 Cleavage produces a blastula from the Chemical and electrical zygote 548 signals coordinate body functions 518 27.11 Gastrulation produces a three-layered embryo 549 26.2 Hormones affect target cells using two main signaling 27.12 Organs start to form after gastrulation 550 mechanisms 519 27.13 Multiple processes give form to the developing 26.3 SCIENTIFIC THINKING A widely animal 552 used weed killer demasculinizes male frogs 520 27.14 EVOLUTION CONNECTION Pattern formation during embryonic development is controlled by ancient ▹ The Vertebrate Endocrine System 520 genes 552 26.4 The vertebrate endocrine system consists of more than a dozen major glands 520 ▹ Human Development 554 26.5 The hypothalamus, which is closely tied to the pituitary, 27.15 The embryo and placenta take shape during the first connects the nervous and endocrine systems 522 month of pregnancy 554 27.16 VISUALIZING THE CONCEPT Human pregnancy is divided into ▹ Hormones and Homeostasis 524 trimesters 556 26.6 The thyroid regulates development and 27.17 Childbirth is induced by hormones and other chemical metabolism 524 signals 558 26.7 The gonads secrete sex hormones 525 27.18 CONNECTION Reproductive technologies increase our 26.8 VISUALIZING THE CONCEPT Pancreatic hormones regulate reproductive options 559 blood glucose level 526 Chapter Review 560 26.9 CONNECTION Diabetes is a common endocrine disorder 527 26.10 The adrenal glands mobilize responses to stress 528 26.11 EVOLUTION CONNECTION A single hormone can perform a 28 Nervous Systems 562 variety of functions in different animals 529 26.12 CONNECTION Hormones can promote social ▹ Nervous System Structure and behaviors 530 Function 564 Chapter Review 530 28.1 Nervous systems receive sensory input, interpret it, and send out 27 Reproduction and Embryonic commands 564 Development 532 28.2 Neurons are the functional units of nervous systems 565 ▹ Asexual and Sexual Reproduction 534 ▹ Nerve Signals and Their 27.1 Asexual reproduction Transmission 566 results in the generation 28.3 Nerve function depends on of genetically identical charge differences across offspring 534 neuron membranes 566

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxxiii C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 33 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services 28.4 A nerve signal begins as a change in the membrane 29.5 The inner ear houses our organs of balance 594 potential 566 29.6 CONNECTION What causes motion sickness? 594 28.5 The action potential propagates itself along the axon 568 ▹ Vision 595 28.6 Neurons communicate at synapses 569 29.7 EVOLUTION CONNECTION Several types of eyes have evolved among animals 595 28.7 Chemical synapses enable complex information to be processed 570 29.8 Humans have single-lens eyes that focus by changing shape 596 28.8 A variety of small molecules function as neurotransmitters 570 29.9 CONNECTION Artificial lenses or surgery can correct focusing problems 597 28.9 CONNECTION Many drugs act at chemical synapses 571 29.10 The human retina contains two types of photoreceptors: rods and cones 598 28.10 SCIENTIFIC THINKING Published data are biased toward positive findings 572 ▹ Taste and Smell 599 An Overview of Animal Nervous Systems 573 29.11 Taste and odor receptors detect chemicals present in ▹ solution or air 599 28.11 EVOLUTION CONNECTION The evolution of animal nervous systems reflects changes in body symmetry 573 29.12 CONNECTION “Supertasters” have a heightened sense of taste 599 28.12 Vertebrate nervous systems are highly centralized 574 29.13 Review: The central nervous system couples stimulus with response 600 28.13 The peripheral nervous system of vertebrates can be divided into functional components 575 Chapter Review 600 28.14 The vertebrate brain develops from three anterior bulges of the neural tube 576 ▹ The Human Brain 576 How Animals Move 602 28.15 The structure of a living supercomputer: The human 30 brain 576 28.16 The cerebral cortex is a mosaic of specialized, ▹ Movement and interactive regions 578 Locomotion 604 28.17 CONNECTION Injuries and brain operations provide insight 30.1 Locomotion into brain function 579 requires energy to overcome 28.18 CONNECTION fMRI scans provide insight into brain friction and structure and function 580 gravity 604 28.19 The reticular formation is involved in arousal and 30.2 Skeletons function in sleep 580 support, movement, and 28.20 The limbic system is involved in emotions and protection 606 memory 581 The Vertebrate Skeleton 608 28.21 CONNECTION Changes in brain physiology can produce ▹ neurological disorders 582 30.3 EVOLUTION CONNECTION Vertebrate skeletons are Chapter Review 584 variations on an ancient theme 608 30.4 Bones are complex living organs 609 30.5 CONNECTION Healthy bones resist stress and heal from injuries 610 29 The Senses 586 30.6 Joints permit different types of movement 611 ▹ Sensory Reception 588 ▹ Muscle Contraction and Movement 611 29.1 Sensory receptors convert 30.7 The skeleton and muscles interact in stimulus energy to action movement 611 potentials 588 30.8 Each muscle cell has its own contractile 29.2 SCIENTIFIC THINKING The apparatus 612 model for magnetic 30.9 A muscle contracts when thin filaments slide along sensory reception is thick filaments 612 incomplete 589 30.10 Motor neurons stimulate muscle 29.3 Specialized sensory contraction 614 receptors detect five 30.11 CONNECTION Aerobic respiration supplies most of the categories of stimuli 590 energy for exercise 615 ▹ Hearing and Balance 592 30.12 SCIENTIFIC THINKING Characteristics of muscle fiber affect 29.4 The ear converts air pressure waves to action potentials athletic performance 616 that are perceived as sound 592 Chapter Review 617

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxxiv C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 34 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services UNIT VI 32.4 Guard cells control transpiration 647 32.5 Phloem transports sugars 648 Plants: Form and Function 619 ▹ Plant Nutrients and the Soil 650 32.6 Plant health depends on obtaining all of the essential inorganic nutrients 650 31 Plant Structure, Growth, 32.7 CONNECTION Fertilizers can help prevent nutrient and Reproduction 620 deficiencies 651 32.8 Fertile soil supports plant growth 652 ▹ Plant Structure and Function 622 32.9 CONNECTION Soil conservation is essential to human 31.1 SCIENTIFIC THINKING The life 653 domestication of crops 32.10 SCIENTIFIC THINKING Organic farmers follow principles changed the course of human meant to promote health 654 history 622 32.11 CONNECTION Agricultural research is improving the yields 31.2 The two major groups and nutritional values of crops 654 of angiosperms are the monocots and the ▹ Plant Nutrition and Symbiosis 655 eudicots 623 32.12 Most plants depend on bacteria to supply 31.3 A typical plant body contains nitrogen 655 three basic organs: roots, 32.13 EVOLUTION CONNECTION Plants have evolved mutually stems, and leaves 624 beneficial symbiotic relationships 656 31.4 Many plants have modified 32.14 The plant kingdom includes epiphytes, parasites, and roots, stems, and leaves 625 carnivores 657 31.5 Three tissue systems make up Chapter Review 658 the plant body 626 31.6 Plant cells are diverse in structure and function 628 33 Control Systems in Plants 660 Plant Growth 630 ▹ ▹ Plant Hormones 662 31.7 Primary growth lengthens roots and shoots 630 33.1 SCIENTIFIC THINKING A 31.8 Secondary growth increases the diameter of woody series of experiments by plants 632 several scientists led to ▹ Reproduction of Flowering Plants 634 the discovery of a plant hormone 662 31.9 The flower is the organ of sexual reproduction in angiosperms 634 33.2 Botanists have identified several major types of 31.10 The development of pollen and ovules culminates in hormones 664 fertilization 635 33.3 Auxin stimulates the 31.11 The ovule develops into a seed 636 elongation of cells in young 31.12 The ovary develops into a fruit 637 shoots 664 31.13 Seed germination continues the life cycle 638 33.4 Cytokinins stimulate cell division 666 31.14 Asexual reproduction produces plant clones 638 33.5 Gibberellins affect stem elongation and have numerous 31.15 EVOLUTION CONNECTION Evolutionary adaptations help some other effects 666 plants to live very long lives 640 33.6 Abscisic acid inhibits many plant processes 667 Chapter Review 640 33.7 Ethylene triggers fruit ripening and other aging processes 668 33.8 CONNECTION Plant hormones have many agricultural 32 Plant Nutrition and Transport 642 uses 669 ▹ Responses to Stimuli 670 ▹ The Uptake and Transport of 33.9 Tropisms orient plant growth toward or away from Plant Nutrients 644 environmental stimuli 670 32.1 Plants acquire nutrients from 33.10 Plants have internal clocks 671 air, water, and soil 644 33.11 Plants mark the seasons by measuring photoperiod 672 32.2 The plasma membranes of root cells control solute 33.12 Phytochromes are light detectors that help set the uptake 645 biological clock 673 32.3 VISUALIZING THE CONCEPT 33.13 EVOLUTION CONNECTION Defenses against herbivores and Transpiration pulls water up infectious microbes have evolved in plants 674 xylem vessels 646 Chapter Review 675

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxxv C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 35 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services UNIT VII 35.2 Fixed action patterns are innate behaviors 700 35.3 Behavior is the result of both genetic and environmental Ecology 677 factors 702 ▹ Learning 703 35.4 Habituation is a simple type of learning 703 34 The Biosphere: An Introduction to 35.5 Imprinting requires both innate behavior and Earth’s Diverse Environments 678 experience 704 35.6 CONNECTION Imprinting poses problems and ▹ The Biosphere 680 opportunities for conservation programs 705 34.1 Ecologists study how 35.7 VISUALIZING THE CONCEPT Animal movement may be a organisms interact with response to stimuli or require spatial learning 706 their environment at several 35.8 A variety of cues guide migratory movements 707 levels 680 35.9 Animals may learn to associate a stimulus or behavior 34.2 SCIENTIFIC THINKING The with a response 708 science of ecology provides 35.10 Social learning employs observation and imitation of insight into environmental others 708 problems 681 35.11 Problem-solving behavior relies on cognition 709 34.3 Physical and chemical factors influence life in the biosphere 682 ▹ Survival and Reproductive Success 710 34.4 EVOLUTION CONNECTION Organisms are adapted to abiotic 35.12 Behavioral ecologists use cost–benefit analysis to study and biotic factors by natural selection 683 foraging 710 34.5 Regional climate influences the distribution of 35.13 Communication is an essential element of interactions terrestrial communities 684 between animals 711 ▹ Aquatic Biomes 686 35.14 Mating behavior often includes elaborate courtship rituals 712 34.6 Sunlight and substrate are key factors in the distribution of marine organisms 686 35.15 Mating systems and parental care enhance reproductive success 713 34.7 Current, sunlight, and nutrients are important abiotic factors in freshwater biomes 688 35.16 CONNECTION Chemical pollutants can cause abnormal behavior 714 ▹ Terrestrial Biomes 689 Social Behavior and Sociobiology 715 34.8 Terrestrial biomes reflect regional variations in ▹ climate 689 35.17 Sociobiology places social behavior in an evolutionary context 715 34.9 Tropical forests cluster near the equator 690 35.18 Territorial behavior parcels out space and 34.10 Savannas are grasslands with scattered trees 690 resources 715 34.11 Deserts are defined by their dryness 691 35.19 Agonistic behavior often resolves confrontations 34.12 Spiny shrubs dominate the chaparral 692 between competitors 716 34.13 Temperate grasslands include the North American 35.20 Dominance hierarchies are maintained by agonistic prairie 692 behavior 716 34.14 Broadleaf trees dominate temperate forests 693 35.21 EVOLUTION CONNECTION Altruistic acts can often be 34.15 Coniferous forests are often dominated by a few species explained by the concept of inclusive fitness 717 of trees 693 35.22 SCIENTIFIC THINKING Jane Goodall revolutionized our 34.16 Long, bitter-cold winters characterize the tundra 694 understanding of chimpanzee behavior 718 34.17 Polar ice covers the land at high latitudes 694 35.23 Human behavior is the result of both genetic and 34.18 The global water cycle connects aquatic and terrestrial environmental factors 719 biomes 695 Chapter Review 720 Chapter Review 696 36 Population Ecology 722 Behavioral Adaptations to the 35 ▹ Population Structure and Dynamics 724 Environment 698 36.1 Population ecology is the study of how and why ▹ The Scientific Study of populations change 724 Behavior 700 36.2 Density and dispersion 35.1 Behavioral ecologists patterns are important ask both proximate population variables 724 and ultimate 36.3 Life tables track survivorship questions 700 in populations 725

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxxvi C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 36 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services 36.4 Idealized models predict patterns of population 37.19 The carbon cycle depends on photosynthesis and growth 726 respiration 753 36.5 Multiple factors may limit population growth 728 37.20 The phosphorus cycle depends on the weathering of 36.6 SCIENTIFIC THINKING Some populations have “boom-and- rock 754 bust” cycles 729 37.21 The nitrogen cycle depends on bacteria 754 36.7 EVOLUTION CONNECTION Evolution shapes life histories 730 37.22 CONNECTION A rapid inflow of nutrients degrades aquatic 36.8 CONNECTION Principles of population ecology have ecosystems 756 practical applications 731 37.23 CONNECTION Ecosystem services are essential to human ▹ The Human Population 732 well-being 757 Chapter Review 758 36.9 The human population continues to increase, but the growth rate is slowing 732 36.10 CONNECTION Age structures reveal social and economic 38 Conservation Biology 760 trends 734 36.11 CONNECTION An ecological footprint is a measure of The Loss of Biodiversity 762 resource consumption 735 ▹ 38.1 Loss of biodiversity includes the Chapter Review 736 loss of ecosystems, species, and genes 762 37 Communities and Ecosystems 738 38.2 CONNECTION Habitat loss, invasive species, overharvesting, pollution, and climate change are major ▹ Community Structure and Dynamics 740 threats to biodiversity 764 37.1 A community includes all 38.3 CONNECTION Rapid warming is the organisms inhabiting a changing the global climate 766 particular area 740 38.4 CONNECTION 37.2 Human activities are responsible for rising Interspecific interactions are concentrations of greenhouse gases 767 fundamental to community structure 740 38.5 Global climate change affects biomes, ecosystems, communities, and populations 768 37.3 Competition may occur when a shared resource is limited 741 38.6 EVOLUTION CONNECTION Climate change is an agent of natural selection 769 37.4 Mutualism benefits both partners 741 37.5 EVOLUTION CONNECTION Predation leads to diverse ▹ Conservation Biology and Restoration Ecology 770 adaptations in prey species 742 38.7 Protecting endangered populations is one goal of 37.6 EVOLUTION CONNECTION Herbivory leads to diverse conservation biology 770 adaptations in plants 742 38.8 Sustaining ecosystems and landscapes is a conservation 37.7 Parasites and pathogens can affect community priority 771 composition 743 38.9 Establishing protected areas slows the loss of 37.8 Trophic structure is a key factor in community biodiversity 772 dynamics 744 38.10 Zoned reserves are an attempt to reverse ecosystem 37.9 VISUALIZING THE CONCEPT Food chains interconnect, disruption 773 forming food webs 745 38.11 SCIENTIFIC THINKING The Yellowstone to Yukon 37.10 Species diversity includes relative abundance and Conservation Initiative seeks to preserve biodiversity by species richness 746 connecting protected areas 774 37.11 SCIENTIFIC THINKING Some species have a disproportionate 38.12 CONNECTION The study of how to restore degraded impact on diversity 747 habitats is a developing science 776 37.12 Disturbance is a prominent feature of most 38.13 Sustainable development is an ultimate goal 777 communities 748 Chapter Review 778 37.13 CONNECTION Invasive species can devastate communities 749 Appendix 1 Metric Conversion Table A-1 ▹ Ecosystem Structure and Dynamics 750 Appendix 2 The Periodic Table A-2 37.14 Ecosystem ecology emphasizes energy flow and chemical cycling 750 Appendix 3 The Amino Acids of Proteins A-3 37.15 Primary production sets the energy budget for ecosystems 750 Appendix 4 Chapter Review Answers A-4 37.16 Energy supply limits the length of food chains 751 Appendix 5 Credits A-26 37.17 CONNECTION A pyramid of production explains the ecological cost of meat 752 Glossary G-1 37.18 Chemicals are cycled between organic matter and abiotic reservoirs 752 Index I-1

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# 110463 Cust: Pearson Education / NJ / CHET Au: Reece Pg. No. xxxvii C / M / Y / K DESIGN SERVICES OF A01_REEC5325_08_SE_FM_NASTA.indd 37 S4carlisle 11/6/13 7:48 AM Title: Campbell Biology: Concepts & Connections 0e Server: Short / Normal Publishing Services