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eighth edition neil a campbell jane b reece berkeley california lisa a urry mills college oakland california michael l cain bowdoin college brunswick maine steven a wasserman university of california san diego peter v minorsky mercy college dobbs ferry new york robert b jackson duke university durham north carolina -pearson bei iamin cummings san francisco boston new york cape town hong kong london madrid mexico city montreal munich paris singapore sydney tokyo toronto
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editor-in-chief beth wilbur executive director of development deborah gale supervising editors pat burner and beth n lvinickojj senior editorial manager ginnie simione /lllson managing editor michael early marketing managers lauren harp and josh frost developmental editors:john burner matt lee alice e fugate and library ofcongre.ss cataloging-in-publication data campbell neil a biology f neil a campbell jane b reece 8th i d p,cm [ndudes index [sbn-13 978-0-8053-6844-4 alk paj r isbn-io 0-8053-6844-2 alk paper 1 biology i reece jane b 11 title qh308.2.c34 2009 570 dc22 suzanne olivier developmental artists hi/air chism carla simmons kelly murph 2007042662 andrew recher and connie balek art editors laura murray and kelly murphy senior photo editor donna kalal photo researcher maureen spuhler editorial assistants benjamin pearson logan triglia and julia student package isbn 978 321-54325-7/0-321-54325-4 student text component isbn 978-0-8053-6844-4/0-8053-68 2 p-copy package [sbn 978 321-53172-8/0-321-53172-8 p-copy text component isbn 978-0-321 -54727-9/0-321-54727-6 copyright © 2008 pearson education [nc publishing as pearson benjamin cummings 1301 sansomi st san francisco ca 94111 all rights il served manufactured in the united states of america this publication is protected by copyright and j rmission should be obtained from the publisher prior to any prohibited reproduction storage in a retrieval system or transmission in any form or by any means electronic mechanical photocopying recording or likewise to obtain permissions to use material from this work please submit a written request to pearson education inc permissions department 1900 e lake ave glenview 1l6oo25 for information regarding permissions call 847 486-2635 many of the designations used by manufacturers and sellers to distinguish their products are claimed as trademarks where those designations appear in this book and the publisher was aware of a trademark claim the designations have been printed in initial caps or all caps pearson benjamin cummings is a trademark in the us andlor other countries of pearson education [nco or its affiliates 234567891o-crk-i1100908 benjamin cummings 1301 sansomi street san francisco ca 94111 khait production management composition and prepress s4cariisle publishing services illustrations precision graphics copy editor janet greenblatt art and design director mark ong design manager marilyn perry text design mark ong jennifer dunn jana anderson cover design yvo riezebos text printer courier kendallville cover printer moore langen/courier senior supplements project editor susan berge supplements production supervisor:jane brundage senior media producer jonathan ballard media project editors nora lally-graves and brienn buchanan associate web developer linda young on the cover magnolia flower chris foxicorbis credits continue following the appendices s pearson bel iamin cummings www.pearsonhighered.com
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1 introduction themes in the study of life 1 unit five unit one the chemistry of life the evolutionary history of biological diversity 2 3 4 5 the chemical context of life 30 water and the fitness of the environment 46 carbon and the molecular diversity of life 58 the structure and function of large biological molecules 68 unit two the cell 6 7 8 9 10 11 12 atour of the cell 94 membrane structure and function 125 an introduction to metabolism 142 cellular respiration harvesting chemical energy 162 photosynthesis 185 cell communication 206 the cell cycle 228 26 phylogeny and the tree of life 536 27 bacteria and archaea 556 28 protists 575 29 plant diversity i how plants colonized land 600 30 plant diversity ii the evolution of seed plants 618 31 fungi 636 32 an introduction to animal diversity 654 33 invertebrates 666 34 vertebrates 698 unit six plant form and function 35 36 unit three genetics plant structure growth and development 738 resource acquisition and transport in vascular plants 764 37 soil and plant nutrition 785 38 angiosperm reproduction and biotechnology 801 39 plant responses to internal and external signals 821 unit seven 13 14 15 16 17 18 19 20 21 meiosis and sexual life cycles 248 mendel and the gene idea 262 the chromosomal basis of inheritance 286 the molecular basis of inheritance 305 from gene to protein 325 regulation of gene expression 351 viruses 381 biotechnology 396 genomes and their evolution 426 animal form and function 852 unit four mechanisms of evolution 22 descent with modification a darwinian view of life 452 23 the evolution of populations 468 24 the origin of species 487 25 the history of life on earth 507 40 41 42 43 44 45 46 47 48 49 50 51 basic principles of animal form and function animal nutrition 875 circulation and gas exchange 898 the immune system 930 osmoregulation and excretion 954 hormones and the endocrine system 975 animal reproduction 997 animal development 1021 neurons synapses and signaling 1047 nervous systems 1064 sensory and motor mechanisms 1087 animal behavior 1120 unit eight ecology 52 an introduction to ecology and the biosphere 1148 53 population ecology 1174 54 community ecology 1198 55 ecosystems 1222 56 conservation biology and restoration ecology 1245
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neil a campbell neil campbell combined the investigative nature of a research scientist with the soul of an experienced and caring teacher he earned his m.a in zoology from ucla and his ph.d in plant biology from the university of california riverside where he received the distinguished alumnus award in 2001 neil published numerous research articles on desert and coastal plants and how the sensitive plant mimosa and other legumes move their leaves his 30 years of teaching in diverse environments included general biology courses at cornell university pomona college and san bernardino valley college where he received the college s first outstanding professor award in 1986 neil was a visiting scholar in the department of botany and plant sciences at the university of california riverside in addition to his authorship of this book he coauthored biology col1cepts col1l1ectiol1s and essential biology with jane reece neil died shortly after the initial planning of this revision but his legacy continues in biology eighth edition for the eighth edition jane reece is joined by five coauthors whose contributions reflect their biological expertise as scientific researchers and their teaching sensibilities gained from years of experience as instructors lisa a urry units 1-3 chapters 2-21 and chapter 47 lisa urry is a professor at mills college in oakland california and was a major contributor to the seventh edition of biology after graduating from tufts university with a double major in biology and french lisa completed her ph.d in molecular and developmental biology at mit following postdoctoral appointments at harvard medical school tufts university and uc berkeley she began teaching at mills college where she currently holds the letts-villard professorship and serves as chair of the biology department she has published research articles on various topics involving gene expression during embryonic development her current research interest is in sea urchin development lisa is also deeply committed to promoting opportunities for women in science education and research jane b reece lead author jane reece neil campbell s longtime collaborator has participated in every edition of biology-first as an editor and contributor then as an author her education includes an a.b in biology from harvard university an m.s in microbiology from rutgers university and a ph.d in bacteriology from the university of california berkeley before mi· grating to california from the northeast she taught biology at middlesex county college and queensborough commu· nity college at uc berkeley and later as a postdoctoral fellow in genetics at stanford university her research focused on genetic recombination in bacteria besides her work on biology she has been a coauthor on biology concepts conl1ections essential biology and the world ofthe cell.
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michael l cain units 4 and 5 chapters 22-34 michael cain is an ecologist and evolutionary biologist currently at bowdoin college michael earned a joint degree in biology and math from bowdoin college an m.sc from brown university and a ph.d in ecology and evolutionary biology from cornell university after postdoctoral work in plant ecology at the university of connecticut and molecular genetics at washington university in st louis michael went on to teach general biology ecology and evolution in a diverse range of settings including carleton college new mexico state university and rose-hulman instihlte of technology in indiana michael is the author of dozens of scientific papers on topics that include foraging behavior in insects and plants longdistance seed dispersal and speciation in crickets peter v minorsky unit 6 chapters 35-39 peter minorsky revised unit 6 for the sixth and seventh editions of biology and is a professor at mercy college in new york where he teaches evolution ecology botany and introductory biology he is also the science writer for the journal plant physiology he received his b.a in biology from vassar college and his ph.d in plant physiology from cornell university after a postdoctoral fellowship at the university of visconsin at madison peter taught at kenyon college union college western connecticut state university and vassar college he is an electrophysiologist who studies plant responses to stress and is currently exploring the possible effects of geomagnetism on plant growth steven a wasserman unit 7 chapters 40-46 48-51 steve wasserman is a professor at the university of california san diego he earned his a.b in biology from harvard university and his ph.d in biological sciences from mit since a postdoctoral sojourn at uc berkeley where he investigated topological transformations of dna he has focused on regulatory pathway mechanisms working with the fruit fly drosophila he has contributed to the fields ofembryogenesis reproduction and immunity as a faculty member at the university of texas southwestern medical center and uc san diego he has taught genetics development and physiology to undergraduate graduate and medical students he has also served as the research mentor for more than a dozen doctoral students and nearly 40 aspiring scientists at the undergraduate and high school level steve has been the recipient of distinguished scholar awards from both the markey charitable trust and the david and lucille packard foundation he recently received the 2007 academic senate distinguished teaching award for undergraduate teaching at uc san diego robert b jackson unit 8 chapters 52-56 rob jackson is a professor of biology and nicholas chair of environmental sciences at duke university he directed duke s program in ecology for many years and is currently the vice president of science for the ecological society of america rob holds a b.s in chemical engineering from rice university as well as m.s degrees in ecology and statistics and a ph.d in ecology from utah state university he was a postdoctoral scientist in stanford university s biology department and an assistant professor at the university of texas austin rob has received numerous awards including being honored at the white house with a presidential early career award in science and engineering from the national science foundation he has published a trade book about the environment the earth remains forever and a children s book of poetry about biology and animals called animal mischief his second children s book not again will be published in 2008.
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uch has changed in the world since the completion of the previous edition of biology in the realm of the biological sciences the sequencing of the genomes of many more species has had deep ramifications in diverse areas of research providing new insights for example into the evolutionary histories of numerous species m there has been an explosion of discovery about small rna molecules and their roles in gene regulation and at the other end of the size spectrum our knowledge of earth s biodiversity has expanded to encompass hundreds of new species in· eluding parrots monkeys and orchids and during the same period biology has become more prominent than ever in our daily lives the news is filled with stories about the promise of personalized medicine novel cancer treatments the possibility of producing biofuels with the help ofgenetic engineering and the use of genetic profiling in solving crimes other news stories report climate change and ecological disasters new drug-resistant strains of the pathogens that cause tuberculosis and parasitic infections and famine-crises in the world around us that are posing new challenges for biologists and their allies in the other sciences on a personal level many colleagues and i have missed our inspiring friend the late neil campbell even as our commitment to leadership in biological education has grown our changing world needs biologists and a scientifically literate citizenry as never be· fore and we are committed to working toward that goal tific expertise ranges from molecules to ecosystems and the schools where they teach range from small liberal arts colleges to large universities in addition both lisa and peter as major contributors to earlier editions had prior experience working on the book the six of us have collaborated unusually closely starting with book-wide planning meetings and continuing with frequent exchanges ofquestions and advice as we worked on our chapters for each chapter the revising author editors and itogether for· mulated a detailed plan subsequently my own role involved commenting on early drafts and polishing the final version together we have strived to extend the book s effectiveness for today s students and instructors while maintaining its core values our core values what are the core values ofthis book they start with getting the science right but then focus on helping students make sense of the science below i highlight our longtime values and describe how they ve been put into practice in the eighth edition you can see examples of many of the book s features in to the student how to use this book pp xiv-xix accuracy and currency getting the science right goes beyond making sure that the facts are accurate and up·to-date equally important is ensuring that our chapters reflect how scientists in the various subdisciplines ofbiology from cell biology to ecology currently view their area changes in the basic paradigms in various biological fields may call for us to reorganize some chapters and even create new ones in a new edition for example a new chapter 21 discusses genomes and their evolution and neurobiology is now covered in two chapters chapters 48 and 49 one focused on the cellular level and one at the organ system level on pages ix-x you can read more about new content and organizational improvements in the eighth edition the new coauthors the seventh edition of biology has been used by more stu· dents and instructors than any previous edition remaining the most widely used college textbook in the sciences with the privilege ofsharing biology with so many students comes the responsibility of improving the book to serve the biology community even better for that reason neil would have been delighted to see that this eighth edition fulfills our decade-long goal of expanding the author team as biological discoveries proliferated neil and i realized that it was becoming harder than ever to make judicious decisions about which biological concepts are most im portant to develop in depth in an introductory textbook we needed an author team with first-hand expertise across the bio-logical spectrum and we wanted coauthors who had honed their teaching values in the classroom our new coauthors-lisa urry michael cain steve wasserman peter minorsky and rob jack· son-represent the highest standards of scientific scholarship across a broad range of disciplines and a deep commitment to undergraduate teaching as described on pages iv-v their scien a framework of key concepts the explosion ofdiscoveries that makes biology so exciting today also threatens to suffocate students under an avalanche of infor mation our primary pedagogical goal is to help students build a framework for learning biology by organizing each chapter around a small number of key concepts typically three to six each chapter begins with a list ofits key concepts a photograph that raises an intriguing question and an overview section that addresses the question and introduces the chapter in the body of the chapter each key concept serves as a nwnbered heading for
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a major section in which the prose and pictures tell a more detailed story at the end of each concept section concept check questions enable students to assess their understanding of that concept before going on to the next concept students encounter the key concepts one last time when they reach the qlapter review at the end ofthe chapter the summary of key concepts restates them and offers succinct explanatory support in both words and summary diagrams-new to this edition active learning increasingly instructors tell us that they want their students to take a more active role in learning biology and to think about biological questions at a higher level in the eighth edition we provide several new ways for students to engage in active learning first the concept check questions in this edition build in difficulty and each set now ends with a new what if question that challenges students to integrate what they have learned and to think analytically there are also questions accompanying selected figures within the text each of these questions encourages students to delve into the figure and assess their understanding of its underlying ideas and new draw it exercises in every chapter ask students to put pencil to paper and draw a structure annotate a figure or graph experimental data in addition to appearing regularly in the chapter review a draw it question may show up in a concept check or figure legend finally the website that accompanies the book features two especially exciting new student tools both of which focus on biology s toughest topics masteringbiology tutorials and bioflix 3-d animations and tutorials these are described on page xx improved figures with the increased use of a more threedimensional art style where it can enhance understanding of biological structure at the same time we avoid excess detail which can obscure the main point of the figure we have also improved our popular exploring figures and have added more see the list on p xii each of these large figures is a learning unit that brings together a set of related illustrations and the text that describes them the exploring figures enable students to access dozens of complex topics very efficiently they are core chapter content not to be confused with some textbooks boxes which have content peripheral to the flow of a chapter modern biology is challenging enough without diverting students attention from a chapter s conceptual storyline telling the story al the righi level whether in pictures or prose we are committed to explaining biology at just the right level and we ve continued to use neil s quantum theory ofteaching biology as a touchstone according to this idea there are discrete levels at which a concept can be successfully explained and a successful explanation must avoid getting stuck bety ·een levels of course most seasoned instructors have independently recognized this issue also known as the too much-too little problem the author team has drawn upon both scientific expertise and teaching experience to tell the story of biology at an appropriate level the importance of scientific inquiry another of our core values is our belief in the importance of introducing students to the scientific way of thinking in both lecture hall and laboratory the authors and many of our colleagues are experimenting with diverse approaches for involving students in scientific inquiry the process by which questions about nature are posed and explored special features in the textbook and in inquiry-based supplements make this edition of bfology more effective than ever in helping instructors convey the process of science in their courses evolution and other unifying themes together with biologys emphasis on key concepts a thematic approach has always distinguished our book from an encyclopedia of biology in the eighth edition as previously the central theme is evolution evolution unifies all of biology by accounting for both the unity and diversity of life and for the remarkable adaptations of organisms to their environments the evolutionary theme is woven into every chapter of biology and unit four mechanisms of evolution has undergone a major revision in chapter i the other unifying themes have been streamlined from ten to six and throughout the book these themes are now referenced more explicitly in key concepts and subheadings the former themes of scientific inquir and science technology and society continue to be highlighted throughout the book not as biological themes but as aspects of how science is done and the role of science in our lives modeling inquiry by example every edition of biology has traced the history of many research questions and scientific debates to help students appreciate not just what we know but how we know and what we do not yet know in bfology seventh edition we strengthened this aspect of the book by introducing inquir figures which showcase examples of experiments and field studies in a format that is consistent throughout the book each of these inquiry cases begins with a research question followed by sections describing the experiment results and conclusion complementing the inquiry figures are research method figures which walk students through the techniques and tools of modern biology in the eighth edition we have added many more inquiry figures there is now at least one in every chapter and often more see the list of inquiry figures on pp xii-xiii each preface vii integration oftext and illustrations we regard text and illustrations as equal in importance and starting with the first edition have always developed them simultaneously the eighth edition has a number of new and
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inquiry figure now ends with a what ift question that requires students to demonstrate their understanding of the experiment described we have also expanded the usefulness of the inquiry figures in another important way in response to feedback from many instructors we now cite the journal article that is the source of the research providing a gateway to the primary literature and the full papers for nine of the inquiry figures are reprinted in inquiry in action interpreting scientific papers by ruth buskirk and christopher gillen this new supplement which can be ordered with the book for no additional charge provides background information on how to read scientific papers plus specific questions that guide students through the nine featured articles learning inquiry by practice biology eighth edition encourages students to practice thinking as scientists by tackling the what if questions in the concept checks and inquiry figures and occasional figure legends as well as the scientific inquiry questions in the chapter review many of those in the chapter reviews ask students to analyze data or to design an experiment the supplements for the eighth edition build on the textbook to provide diverse opportunities for students to practice scientific inquiry in more depth in addition to inquiry in action interpreting scientific papers these include new editions ofseveral other supplements that can be made available without cost one is biologicallnf[uiry a workbook ofinvestigative cases second edition by margaret waterman and ethel stanley another is practicing biology a student workbook third edition by jean heitz and cynthia giffen you can find out more about these and other student supplements both print and electronic on pages xx-xxiii courses could or should attempt to cover but given the great diversity of course syllabi we have opted for a survey broad enough and deep enough to support each instructor s particular emphases students also seem to appreciate biologys breadth and depth in this era when students sell many of their textbooks back to the bookstore more than 75 of students who have used biology have kept it after their introductory course in fact we are delighted to receive mail from upper division students and graduate students including medical students expressing their appreciation for the long-term value of biology as ageneral resource for their continuing education just as we recognize that few courses will cover all 56 chapters of the textbook we also understand that there is no single correct sequence of topics for a general biology course though a biology textbook s table of contents must be linear biology itself is more like a web of related concepts without a fixed starting point or a prescribed path diverse courses can navigate this network of concepts starting with molecules and cells or with evolution and the diversity of organisms or with the big-picture ideas of ecol· ogy we have built biology to be versatile enough to support these different syllabi the eight units ofthe book are largely selfcontained and for most ofthe units the chapters can be assigned in a different sequence ithout substantial loss ofcoherence for example instructors who integrate plant and animal physiology can merge chapters from unit six plant form and flulction and unit seven animal form and function to fit their courses aj another option instructors who begin their course with ecology and continue with this top-down approach can assign unit eight ecology right after chapter 1 which introduces the unifying themes that provide students with a panoramic view of biology no matter what the topic order ofthe course syllabus our partnership with instructors a core value underlying all our work as authors is our belief in the importance of our partnership with instructors our primary way of serving instructors of course is providing a textbook that serves their students well in addition benjamin cummings makes available a wealth of instructor resources in both print and electronic form see pp xx-xxiii however our rela tionship with instructors is nota one-way street in our continu ing efforts to improve the book and its supplements we benefit tremendously from instructor feedback not only in formal re views from hundreds ofscientists but also via informal communication in person and byphone and e-mai1 neil campbell built a vast network ofcolleagues throughout the world and my new coauthors and i are fully committed to continuing that tradition the real test of any textbook is how well it helps instructors teach and students learn we welcome comments from the students and professors who use biology please address your suggestions to me jane reece pearson benjamin cummings 1301 sansome street san francisco ca 94111 e-mail address janereece@cal.berkeley.edu the biology interviews a continuing tradition scientific inquiry is a social process catalyzed by communi cation among people who share a curiosity about nature one of the many joys of authoring biologyis the privilege of interviewing some of the world s most influential biologists eight new interviews one opening each unit of the textbook introduce students to eight of the fascinating individuals who are driving progress in biology and connecting science to society and in this edition each unit of the text includes an inquiry figure based upon the research of the unit s interviewee for example see inquiry figure 2.2 on page 31 the interviewees for this edition are listed on page xi aversatile book our book is intended to serve students as a textbook in their general biology course and also later as a useful tool for review and reference biology s breadth depth and versatile organization enable the book to meet these dual goals even by limiting our scope to a few key concepts per chapter biology spans more biological territory than most introductory viii preface
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this section provides just a few highlights of new content and organizational improvements in biology eighth edition unit one the chemistry of life new examples make basic chemistry more engaging for students including an explanation of why steam can burn your skin in chapter 3 the structures of the enantiomeric medications ibuprofen and albuterol in chapter 4 and information on trans fats in chapter 5 a new inquiry figure in chapter 3 relates acidity to the emerging global problem of ocean acidification and its effects on coral reefs the new inquiry figure in chapter 5 shows roger kornberg s 3-d model of the rna polymerase-dna-rna complex work for which he won the 2006 nobel prize in chemistry unit two lation goes awry material on bacterial genetics in seventh edition chapter 18 has been moved to other chapters within the genetics unit and to chapter 27 on prokaryotes chapter 19 now covers only viruses from seventh edition chapter 18 giving this chapter the flexibility to be assigned at any point in the course chapter 20 continues to cover biotechnology but genome sequencing and analysis have been moved to chapter 21 cloning and stem cell production are now in chapter 20 newly explained techniques include the screening of an arrayed library bac clones northern blotting rt-pcr and in situ hybridization the explosion of discoveries about genomes and their evolution led us to develop a chapter devoted to this subject the new chapter 21 this chapter consolidates new material with topics from chapters 19-21 of the seventh edition unit four the cell the judicious addition of recent research includes updated coverage of the sensory roles ofprimary cilia in chapter 6 new developments regarding the membrane model in chapter 7 and paul nurse s nobel prize-winning work on the cell cycle in chapter 12 chapter 11 now ends with a section on apoptosis formerly in chapter 21 new inquiry figures in this unit describe research on the role of microtubules in orienting cellulose in cell walls chapter 6 allosteric regulators of enzymes chapter 8 atp synthase chapter 9 yeast cell signaling chapter ll and a cell cycle regulator chapter 12 unit three mechanisms of evolution genetics chapter 14 now includes tips for genetics problems in chapter 15 sex linkage is discussed directly after the discussion of the white-eye trait in morgan s fruit flies chapter 16 covers replication of the bacterial chromosome and the structure of the eukaryotic chromosome including a new exploring figure formerly in chapters 18 and 19 respectively we have reorganized chapters 18-21 with the dual aims of telling a more coherent story and facilitating instructors coverage of molecular genetics regulation of gene expression for both bacteria and eukaryotes is now consolidated in chapter 18 which also includes a concept section on the crucial role of small rnas in eukaryotes we have streamlined material on the genetic basis of development formerly in chapter 21 and included it in chapter 18 where it provides the ultimate example of gene regulation chapter 18 ends with a section on the molecular basis of cancer previously in chapter 19 to demonstrate what happens when gene regu our revision emphasizes the centrality of evolution to biology and the breadth and depth of evidence for evolution new examples and inquiry figures present data from field and laboratory studies and reveal how scientists study evolution chapter 22 discusses how evolution can be viewed as both a pattern and a process and introduces three key observations about life that are explained by evolution the match between organisms and their environments adaptation the shared characteristics unity oflife and the diversity of life this discussion serves as a conceptual anchor throughout units four and five chapters 24 and 25 have been significantly reorganized chapter 24 is now more tightly focused on speciation enabling better pacing of this highly conceptual material a new concept section explores hybrid zones as naturallaboratories for studying speciation chapter 25 focuses on macroevolution incorporating topics formerly in chapters 24 and 26 such as the correlations between earth s geologic and biological history but the primary storyline concerns what we can learn from the fossil record about the evolutionary history of life new text and figures explore how the rise and fall of dominant groups of organisms are linked to large-scale processes such as continental drift mass extinctions and adaptive radiations coverage of evo-devo has been expanded phylogenetic trees are introduced earlier in a new section on tree-thinking in chapter 22 this material supports students in interpreting diagrams before studying phylogenetics more fully in chapter 26.
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unit five the evolutionary history of biological diversity unit seven animal form and function a new chapter 26 phylogeny and the tree of life introduces the unit extending material formerly in chapter 25 it describes how evolutionary trees are constructed and underscores their role as tools for understanding relationships rather than facts to be memorized new sections address common misconceptions in interpreting trees and help motivate students with practical applications chapter 27 has a new concept section on prokaryotic re production mutation and recombination formerly in chapter 18 this unifies the coverage of prokaryote biology and supports students in developing a fuller understanding of these microorganisms throughout unit five along with updating the phyloge· nies of various groups of organisms-introducing for exam· pie the usupergroup hypothesis of eukaryotic phylogeny in chapter 28 -we have found new opportunities to use the study of phylogeny as an opportunity to illustrate the iterative nature of the scientific process we aim to help students stay focused on the big picture of why biologists study evolutionary relationships each chapter also now includes an inquiry figure that models how researchers study organisms and their relationships at the same time in each chapter we highlight the key roles that various organisms play in the biosphere as well as their applied importance for humans unit six plant form and function an evolutionary perspective more strongly pervades this unit underscoring how environment and physical laws shape adaptations across animal groups each chapter now includes at least one inquiry figure together these figures highlight the wide range of methodologies used to study animal physiology including several experiments using molecular biology techniques students studied earlier in the book chapter 40 has been revised and reorganized to highlight functional relationships at all levels of organization in animal bodies thermoregulation serves as an extended example throughout the chapter chapter 43 the immune system has been extensively revised for instance we now contrast recognition of pathogen class in innate immunity with antigenspecific recognition in adaptive immunity helping overcome the common misconception that recognition is absent in innate immunity we have divided the former nervous system chapter into m o enabling us to better pace difficult material and high· light dynamic current research by focusing first on cellular processes in chapter 48 and then on nervous system or· ganization and function in chapter 49 chapter 50 rounds out the discussion of nervous system function by examining sensory and motor mechanisms this sequence leads naturally into chapter 51 on animal behavior formerly in unit eight which ties together aspects of genetics natural selection and physiology and provides a bridge to the ecology unit revisions to this unit draw more attention to the experimental basis of our understanding of plant biology new examples include recent progress toward identifying the flowering hormone chapter 39 featured in new inquiry figures are experiments demonstrating for example that trichomes affect insect feeding chapter 35 and that informational molecules transported through the symplast affect plant de· velopment chapter 36 in chapter 36 now titled resource acquisition and transport in vascular plants a new first concept section explores how architectural features of plants facilitate resource acquisition helping students relate the transport of water and nutrients to what they learned in chapter 35 about plant structure and growth another new concept section on symplastic transport discusses recent insights into changes in plasmodesmata shape and number and the transmission of electrical and molecular signals throughout the plant this unit now has more examples of practical applica· tions of plant biotechnology for instance chapter 37 dis· cusses how genetic modification has increased the resist· ance of some plants to aluminum toxicity and has improved the flood tolerance of rice crops chapter 38 elaborates on the principles of plant breeding and incorporates a new section on genetic engineering of biofuels x new to the eighth edition unit eight ecology this unit which now includes chapters 52·56 incorporates many new examples that demonstrate a range of methods and scales ofstudy for example a new figure in chapter 52 describes a large·scale field experiment in which researchers manipulated precipitation levels in forest plots while new research method figures describe determining population size using the mark-recapture method chapter 53 using molecular tools to measure diversity of soil microorganisms chapter 54 and determining primary production with satellite data chapter 55 by building on earlier units we hope to demonstrate how ecology represents a fitting capstone to the book we provide more microbial examples and more aquatic ones from diverse locations around the globe for instance chapter 52 now discusses the importance of salinity in determining the distribution ofaquatic organisms and chapter 54 s coverage of the intermediate disturbance hypothesis includes a new figure on a quantitative test of the hypothesis in new zealand streams the unit highlights the great relevance of ecology to society and to students lives a new concept section in chapter 54 for example discusses how community ecology helps us understand pathogen life cycles and control disease.
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unit one unit five the chemistry of life deborah m gordon stanford university the evolutionary history of biological diversity sean b carroll university of visconsin-madison interview 534 inquiry figure 684 interview 28 inquiry figure 31 unit two unit six the cell plant form and function paul nurse rockefeller university patricia zambryski university of california berkeley interview 92 inquiry figure 240 interview 736 inquiry figure 782 f unit three unit seven genetics animal form and function 1 1 · terry l orr-weaver of technology massachusetts institute masashi vanagisawa university of texas southwestern medical center interview 850 inquiry figure 908 interview 246 inquiry figure 322 unit four unit eight mechanisms of evolution ecology scott v edwards harvard university diana h wall colorado state university interview 450 inquiry figure 489 interview 1146 inquiry figure 1210
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exploring figures 4.10 5.21 6.9 6.32 7.20 1.4 i ewls of biological orpnization 4 some biologically important chemical groups 64 levels of protein structure 82 animalandpl.antcells 100 intercellular junctions in animal tissues 121 endocytosis in animal cells 139 11.7 membrane receptors 211 12.6 the mitotic division ofan animal cell 232 13.8 the meiotic division of an animal cell 254 16.21 chromatin packing in a eukaryotic chromosome 320 24.4 25.6 27.18 28.3 29.5 29.9 29.15 30.5 ]0.13 31.11 33.3 33.37 34.35 35.10 37.14 38.4 38.1 40.5 41.6 42.5 46.12 50.8 50.29 52.2 52.10 52.18 52.21 55.14 56.23 reproductive barriers 490 the origin ofmammals sf3 major groups ofbactcria 568 protistan diversity 578 derived traits of land plants 602 bryophyte diversity 608 seedlessvasruiar p ant diversity 6/4 gymnosperm ~ty 612 angiosperm diversity 630 fungal dinnity 642 invertebrate diversity 667 insect diversity 690 mammalian diversity 724 examples of differentiated plant cells 744 unusual nutritional adaptations in plants 798 flower pollination 804fruit and seed dispersal 8/1 structure and function in animal tissues 856 four main feeding mechanisms of animals 881 double circulation in vertebrates 902 humangametogenesis /008 the structure ofthe human ear 1093 the regulation ofskelet.d muscle contrdction /109 the scope of ecological research 1149 global oimate patterns jj aquatic biomes 1162 terrestrial biomes 1168 nutrientc des 1232 restoration ecology forldwide 1262 inquiry figures 1.27 does the presence of poisonous coral snakes affect predation rates on their mimics kingsnakes 22 ~2.2 what creates devil s gardens in the rain forest 31 3.11 what is the effect of carbonate ion concentration on coral reef calcification 55 4.2 can organk molecules form under conditions believed to simulate those on the early earth 59 5.25 hat can the 3-d shape of the enzyme rna polymerase ii tell us about its function 86 the original research paper and study questions are pro ided in inquiry in action erprcling xientific papers 6.29 what role do microtubules pby in orienting deposition of cellulose in ceii alis 119 7.6 do membrane proteins move 128 8.21 are there allosteric inhibitors ofcaspase enzymes 158 9.15 is the rotation of the internal rod in atp synthase responsible for atp synthesis 174 10.9 which wavelengths of light are most effective in driving photosynthesis 191 11.16 howdo signals induce directional cell growth in yeast 220 12.8 at which end do kinetochore mkrotubujes shorten during anaphase 235 12.13 do molecular signals in the cytoplasm regulate the cell cycle 238 12.16 how does the activity of a protein kinase essential for mitosis vary during the cell cycle 240 13.10 hat prevents the separation of sister chromatids at anaphase i of meiosis 257 14.3 hen fl hybrid pea plants are allowed to self-pollinate which traits appear in the f generation 264 14.8 do the alleles for one character assort into gametes dependently or independently of the alleles for a different character 268 15.4 in a cross between a wild-t pe female fruit fly and a mutant whit ed male what color e es wiu the f1 and f offspring have 289 15.9 how does linkage between t o genes affect inheritance of characters 293 16.2 can a genetic trait be transferred between different bacterial straim 306 16.4 is protein or dna the genetic material of phage t2 307 16.11 does dna replication follow the conservative semiconservative or dispersive model 312 16.22 what role does histone phosphorylation play in chromosomal behavior during meiosis 322 17.2 do individual genes specify the enzymes that function in a biochemical pathwdy 327 18.19 is bicoid a morphogen that determines the anterior end of a fruit fly 372 19.2 hatcausestobaccomosakdisease 382 20.16 can a differentiated plant cell devriop into a whole plant 412 20.17 can the nucleus from a differentiated animal cell direct deveiopmentofanorganism 413 21.16 what is the function ofa gene foxp2 that is rapidly evolving in the human lineage 444 22.13 can predation ll suit in natural selection for roior patterns in guppies 460 ~23.16 do females sdect mates based on traits indicative of good genes 482 24.3 does gene now occur between widel separated populations 489 24.9 can divergence ofallopatric populations lead to reproductive isolation 495 24.12 does sexual selection in cichlids result in reproductive isolation 497
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25.23 what causes the loss of spines in lake stickleback fish 528 26.6 what is the species identity offood being sold as whale meat 539 27.10 can prokaryotes evolve rapidly in response to environmental change 560 28.23 xrhat is the root of the eukaryotic tree 593 29.10 can bryophytes reduce the mte at which key nutrients are lost from soils 609 ·30.14 can flower shape influence speciation mte 632 31.21 do endophytes benefit a woody plant 648 32.6 did catenin play an ancient role in the molecular control ofgastrulation 658 33.28 did the arthropod body plan result from new hox genes 684 34.43 did neanderthals give rise to european humans 732 35.9 do soybean pod trichomcs deter herbivores 743 36.21 does phloem sap contain more sugar near sources than sinks 781 36.22 do alterations in symplastic communication affect plant development 782 ·37.13 does the invasive weed garlic mustard disrupt mutualistic associations between native tree seedlings and arbuscular mycorrhizal fungi 797 38.6 do gaba gradients playa role in directing pollen tubes to the eggs in arahidopsis 806 39.5 xrhat part ofa gmss coleoptile senses light and how is the signal tmnsmitted 825 39.6 does asymmctrical distribution of a growth-promoting chemical cause a coleoptile to grow toward the light 826 39.7 xrhat causes polar movement of auxin from shoot tip to base 828 39.17 how does the order of red and far-red illumination affect seed germination 836 40.14 how does a burmese python generate heat while incubating eggs 867 ·41.5 can diet influence the frequency of birth defects 880 41.24 xrhat are the roles of the ob and db genes in appetite regulation 895 42.12 how do endothelial cells control vasoconstriction 908 42.31 what is the basis for the pronghorn s unusually high me of 0 1 consumption 926 43.5 can a single antimicrobial peptide protect fruit flies against infection 932 44.7 how do seabirds eliminate excess salt from their bodies 958 44.20 can aquaporin mutations cause diabetes insipidus 971 45.4 xrhere in the cell is the receptor for melanocyte-stimulating hormone 978 45.22 xrhat role do hormones play in making a mammal male or female 993 46.9 xrhy is sperm usage biased when female fruit flies mate twice 1003 47.4 is the distribution of ca1 in an egg correlated with formation of the fertilization envelope 1024 47.19 is cadhcrin required for development of the blastula 1037 47.20 is an organized fibronectin matrix required for convergent extension 1037 47.23 how does distribution of the gray crescent affect the developmental potential of the first two daughter cells 1040 47.24 can the dorsal lip of the blastopore induce cells in another part of the amphibian embryo to change their developmental fate /041 47.26 xrhat role does the wne of polarizing activity zpa play in limb pattern formation in vertebmtes 1043 48.17 does the brain have a specific protein receptor for opiates 1061 49.12 xrhich cells control the circadian rhythm in mammals 1073 50.14 how do mammals detect different tastes /098 50.37 what are the energy costs of locomotion 16 51.11 does a digger wasp use landmarks to find her nest 127 51.14 are the songs of green lacewing species under the control of multiple genes 30 51.15 are differences in migmtory orientation within a species genetically determined 1131 52.8 does feeding by sea urchins limit seaweed distribution 1153 53.8 how does caring for offspring affect parental survival in kestrels 1180 54.3 can a species niche be influcnced by interspecific competition 1200 54.15 is pisaster ochraceus a keystone predator 1208 54.19 are soil nematode communities in antarctica controlled by bottom-up or top-down factors 1210 54.28 how does species richness relate to area /217 55.7 fhich nutrient limits phytoplankton production along the coast of long island /227 55.15 how does temperature affect litter decomposition in an ecosystem /234 ·56.11 xrhat caused the drastic decline of the illinois greater pmirie chicken population 1252 research method figures 2.6 6.3 6.4 6.5 7.4 10.8 13.3 14.2 14.7 15.11 20.4 20.7 20.8 20.9 20.11 20.12 20.13 20.15 20.18 20.25 26.15 radioactive tracers 34 light microscopy 96 electron microscopy 96 cell fmctionation 97 freeze-fracture 127 determining an absorption spectrum 191 preparing a karyotype 250 crossing pea plants 263 the testcross 267 constructing a linkage map 296 goning genes in bacterial plasmids 399 detecting a specific dna sequence by hybridization with a nucleic acid probe 4fj2 the polymemse chain reaction pcr 404 gel electrophoresis 405 southern blotting of dna fragments 4fj7 dideoxychain termination method for sequencing dna 4fj8 rt-pcr analysis of the expression of single genes 4fj9 dna microarray assay of gene expression levels 410 reproductive cloning of a mammal by nuclear transplantation 414 using the li plasmid to produce transgenic plants 421 applying parsimony to a problem in molecular systematics 546 using dendrochronology to study climate 753 hydroponic culture 790 intracellular recording 1052 determining population size using the mark-recapture method 1/75 determining microbial diversity using molecular tools 1205 determining primary production with satellites /225 35.21 37.6 48.8 53.2 54.10 55.5 featured figures xiii
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to the student how to use this book focus on the key concepts each chapter is organized around a framework of 3 to 6 key concepts that will help you stay focused on the big picture and give you a context for the the supporting details ofl m i ii 25.1 conditions on early earth made the origin of figure 25.1 who doe fo il eviden
ere this assumption might not be valid and explai how the estimate opopulation size would be affl ted for su9ge5ted an,wer see appendix a each set of concept check questions builds in difficulty ending with a what iff question that asks you to apply what you ve learned work through these questions on your own or in a study group-they re good practice for the kinds of questions you might be asked on an exam xiv
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