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Molecular Cell Biology (Lodish, Sixth Edition) PDF

1265 Pages·2007·181.93 MB·English
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BroJuapt )gpalsaer.)lL Oua oj ]l lannopp E;aepr ur.6asapr)qars ru ruunua slslar uqaqenrol C 4r1oa6 qsinptes lq a^Puons ]Ppt)s }qPl nrtnsa sau-rdlo {1o o^epa o nrluLlunu laesdousas eup }q naaei,s on riuJun uso{rsle Lpurslru6nrsqas lueup lrorlL ]oa C 6l1oa6q ieelqas ruuunuolo6{ 1 onupetbtepnel eslnpauls e 1Hpurer plurnarsrli leup nl1 PusLIsHgn: SaraT enney ExecurrvE Eonon: Katherine Ahr DpvEr-opneNrRI- Eorrons: Matthew Tontonoz, Erica PantagesF rost, Elizabeth Rice AssocIRrp Pnoyrcr MexecBR: Hannah Thonet AssrsreNr EorroR: Nick Tymoczko AsssocrerED nacton op MeRrrrrNc: Debbie Clare SeNIoRP no;ecr Eonon: Mary Louise Byrd Tsxr DpsrcNBR: Marsha Cohen Pacp Mnreup: Aptara, Inc. CovpR Drsrcrs:B lakeL ogan IlLuslReltoN CooRorNetoR: SusanT immrns ILrustR,q.rroNs: Nerwork Graphics, Erica Beade,H . Adam Steinberg Puoro Eorron: CeciliaV aras PHoro RpseencHrn: Christina Micek PRooucnoN CooRoneton: Susan\ 7ein Mron RNo Suppr-rvENTSE DrroR: Hannah Thonet Mron DEvgLoplns: Biostudio, Inc., Sumanas,I nc. CotvtposrrloN: Aptara, Inc. MeNupRcruRrNG: RR Donnelley & SonsC ompany About the cover: Mitotic PtK2 cellsi n late anaphases tainedb lue for DNA and greenf or tubulin. Courtesyo f Torsten\ gittman. Library of CongressC ataloging-in-PublicationD ata Molecular cell biology I Harvey Lodish . . . [et al.]. -6th ed. p. cm. Includesb ibliographicalr eferencesa nd index. 1. Cytology. 2. Molecular biology. I. Lodish, Harvey F. QH581.2.M6552 007 57L5-dc22 2007006188 ISBN-13 : 978 -0-71 67- 76 01-7 ISBN-10:0 -71 67- 7 601-4 @ 1986,1.990,1 995,2000,2004,2008 by tJf.H . Freemana nd Company All rights reserved. Printed in the United Stateso f America Secondp rinting W. H. Freemana nd Company 41 Madison Avenue,N ew York, NY 10010 Houndmills, Basingstoke RG21 6XS, England www.whfreeman.com To our studentsa nd to our teachers, from whom we continuet o learn, and to our familiesf,o r their support, encouragemenat,n d love PREFACE n writing the sixth edition of Molecular Cell Biology what we know. A number of experimental organisms,f rom I we have incorporated many of the spectaculara dvances yeastst o worms to mice, are used throughout so the student I I made over the past four years in biomedical science,d riven can seeh ow discoveriesm ade with a "lower organism" can in part by new experimental technologiest hat have revolu- Iead directly to insights even about human biology and dis- tionized many fields. High-velocity techniquesf or sequenc- ease.T his experimental approach, evident in the text itself, ing DNA, for example,h aveg eneratedt he completes equence has also been thoroughly integrated into the pedagogical of dozens of eukaryotic genomes;t hese in turn have led to framework. For example: important discoveriesa bout the organization of the human genome and regulation of gene expression,a s well as novel r Experimental Figures lead students through important insights into the evolution of life-forms and the functions of experimental results. individual members of multiprotein families. New imaging techniquesh ave generatedp rofound revelations about cell r Classic Experiments essaysf ocus on historically impor- organization and movement, and new molecular structures tant and Nobel Prize-winninge xperiments. have greatly increasedo ur understanding of life processes r New and revised Analyzet he Data problems at the end such as cell-cell signaling, photosynthesis,g ene transcrip- of each chapter require the student to synthesizer eal ex- tion, and chromatin structure. perimental data to answer a serieso f questions. New Author Team r Updated Perspectives for the Future essays explore potential applications of future discoveriesa nd unanswered Two new authors have been instrumental in refocusing this questionst hat lie ahead in research. book toward these exciting new developments. Anthony Bretscher of Cornell University is known for identifying and characterizing new components of the actin cytoskeleton and elucidating their biological functions in relation to cell polarity and membrane traffic. Hidde Ploegh, of the Massachusetts Institute of Technologg has made major contributions to our understanding of immune system behavior, particularly in regard to the various tactics that viruses employ to evade our immune responsesa nd the ways our immune systemr esponds. Both authors are widely recognized for their researcha s well as their classroomt eachinga bilities. 'We are grateful to Paul Matsudaira, Jim Darnell, Larry ZipurskS and David Baltimore for their exceptional contri- butions to the previous editions of Molecular Cell Biology. Much of their vision and insight is apparent at many places in this book. ExperimentaEl mphasis Fluorescenmceic roscopsyh owst he locationo f DNAa nd multiple The hallmark of Molecular Cell Biology has always beent he proteinws ithint he samec ell.l FromB N G Giepmaents a l, 2006S, cience use of experimentst o teach students how we have learned 3'12:21|7 vtl New DiscoveriesN,e w Methodologies Coiled-coisl talk Methodological advances continue to expand and enrich our knowledge of molecular cell biology and lead to new understanding. Following are just a selection of the new Motor head experimental methodologiesa nd cutting-edges ciencei ntro- Neckl inkers Microtubule (+) duced in this edition: r Expanded coverage of proteomics, including organelle proteome profiling and advances in mass spectroscopy (Chapter3 ) I Forwardm otorb indsB -tubulin, ^ releasinAgD P r Expandedc overageo f RNAi, including the useo f shRNAs to inhibit any gene of interest in a cultured cell or organism (Chapters5 , 8) r Updated discussionso f chromatin, including structure and condensation( Chapter6 ), control of genee xpressionb y chro- matin remodeling (Chapter 71, and chromatin-remodeling p Forwardh eadb indsA TP proteins and tumor development( Chapter 25) ? r Evolution of chromosomes and the mitochondrion (Chapter 6) r New molecular models, including pre-initiation complex and mediator complex (Chapter 7); annular phospholipids (Chapter 1,0);C az* ATPase( Chapter 11); rhodopsin, trans- p Conformationacl hangei n neckl inkerc ausesr ear ducin, and protein kinase A (Chapter 15); and myosin head to swing forward ATPase (Chapter 17) r Latest advancesi n light and electron microscopy,i nclud- ing cryoelectron tomography (Chapter 9) r Reactive oxygen species( ROS) (Chapter 12) [ rue* forwardh eadr eleaseAs DB ^ trailingh eadh ydrolyzesATP r Role of supercomplexesi n electron transport (Chapter 12) V andr eleasePs . r Human epidermal growth factor receptors (HERs) and treatment of cancer (Chapter 16) r Myosin ATPasec ycle (Chapter 17) r Kinesin-1M Pase cycle (Chapter 18) r Use of retrovirus infection for tracing cell lineage (Chapter2 1) r Axon guidance molecules (Chapter 23) r Somaticg ener earrangementi n immune cells (Chapter 24) r Cancers tem cells (Chapter2 5) r Use of DNA microarray analysis in tumor typing (Chapter2 5) Figure1 8-22K inesin-u1s esA TPt o "walk"d owna microtubule vill PREFACE lu)taeso 1p1er.r1 [eu Vp))assrqr;r1f 1 umr.r1ru3 lqe srxlq oprlrou rlo {rE^a lelau e sled qEJI eup es)ep ^q e.rr,\re, Es exderraura plaeJ{.rers ot qol qnupar.Srep- nelo eup 8repnela slnpeuls'reu po lo urp{o urelarreI tqel ,fEu ge penulluS ;or slnpouls uroJa N/o lrp^o eddroecqeqla' nlep lo qeleura onJ Jolare8a o;lqe lelesl spaunJrl pa^al- odruauls \^rlq slnpeuls. uaop Jor Jleer exdleuelrous oJ lqe ;nuperuaulels' eu p\yr.e lrel arulousrJrop onJ rourutllureul lo urelnuS qorq rqe couradls eu paxdarru.rauls utora eJlassrqlo' 1u derlrcnler, tlo qEAa rrudrorrap lqa cqudla roaeJrlreras' rur4ruS snJa lqet lqed 8rr,e slnpauts rqe qr8 drrlnra eup dro- ^rpe e sollp JonupElrou toJ {^qpl rs 10 N/3 qE^e elso ,fourO' srudq;rap rqa lex] eu pqavpru8s'reruor;ruS ler8ou \lgerelaJ dossrqla 'JIOEJ JouJedl-ouautap qeepruEs uom drorrrpe eu aesd-lo-roep roepured oJ eelq rqedler lenn gl6eurzellou laytltzap qnueu a6 6ra1 1sqonnlu r6.ue1a eu p]aJuele drounrlat' lfonJ]asI o L]-qa lauup UNtlsou vMprp goe]p I Iu ,fousnllelrou .^rt{r utore r{eu 196 droyassors ot leII qroyoSd ;rour €Jonup lga \^o qp^a la^rsap lqa oonulrd' lBqle oJ Jouleuls 10 nurla rolplop uelarrE IEUP urrror lqe Nan/ )qedlars or8eurzBlrou oJ urosl r{lq elso slrapulluep ?uP ,fonJses' lo,fnsap colere8a o; rueuI lodrcs g rt.,aor.SeurzruS selereI lqa srxrq epruou oJ wpacrln )a lgi toToSK rurlnpes tqree cqedters. eueglruS ns lo rrrel>e roou Jor tqa ppprrrou ot uarvr rqedlars: ua\^ urplarrpl' gel{r rqedter {rps Eu nd-to-pelo lrsr o JJeJar- eu,fes oJ leupruer{ slnpres €uJporudr.aqeusrL,e Jelre\^ ?ur- r "J{ra Wolernler JaII groloSd o ;qat,aloduraul., )J{rBdre; rles lqel prraD slnpauls eup ruslJnflors to urnfl. rPPprlrouEI 77 (dr.osaurs l{ro Jnuppuroulels oJ per,alodureul {oru Jorlrl- rzelrou lqron8q deuar.u \,^rl{ rru,freesap JoJns ou ruJoJur€lrou' ;or.utplrou( urEurul?IrEu per,elodruaut' r ,,Nar^e ]ells.. )3qedlar 79 (drorrrpes nurJre pcoriareSa o1 rqa ]e[ qroloSd o; lqesa sdonprza pca11s r'ucynpruS rga let- os plp^Eu,fes ru exoupI d?l{r ;rupruS eup sausord ra,fadlrou' ,r,1rurunuoyo3d,. )alredln 7y( eddlra sdr.rucrdlos o; rrtoler- nleJ raII qroloSd ro fqa ^arlagrplo rururnue sdslouI' wapt)el ualeneu)a 141eud upleuces ru q?srJ leIIFEr Eup u'lolatnler qroloS ql erza lep luoo.^\/, lJeeluauls 'oJ JpufaJ €up orqoJ srSurlrceul qnueu prsassos. lqase ureprlel exeudles er.e \lo^ou lqron8qonl tqe rqedlers mqa;a eddroduele lo 8rrre slnpeuts pu eddr.acrelrou Jor lqo pruloel eddlrcenous o; lqe qesr,f srreure lqod ere oJ 1oeruru8 .gryeud lqaso eddqcenous qru8e ou e porerlop nuparsleupruS oJ ruprrdr.ololu corudlaxas roua1ys-corudlaxas lqel cetpdze ca1 roSnlela CNV ruolelueuls' lreuscrtdlrou' JoorpruEla uolEqolrsu. ?up ,fouuerl lalls lo olqer rolls Bup Sxdeuueule l116nle 6-€ 0Cl]]aloulreltou ru lnllnra o ]drrueuf lo drolerus pup ,fffqoqdprEles ru l{rarr exlrErtollnlEr ou^rrou- l.uonsa selallt] a)uL^oqlesl( )alls tulo t'uns)la lolls lSonyas Io1 3 uaul. v Jourdlela IIsr oJ llra uaprJE IexEu.rdles ru lqo srxq ]ruarsou eup [ )qau 'gos]ou gtouap) uealsoet-)q lusltln]eI oprnou JEU q0 JonuP o^erlaPJ' dut lv)l IX

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