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Molecular Cell Biology (Lodish, Sixth Edition) PDF

1265 Pages·2007·181.93 MB·English
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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 anaphase s 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 famil iesf,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. r Updated Perspectives for the Future essays explore New Author Team 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 v t l Coiled-coisl talk New DiscoveriesN,e w Methodologies 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 p Conformationacl hangei n (Chapter 1,0);C az* ATPase( Chapter 11); rhodopsin, trans- 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 ^ trail ingh 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 vi l l PREFACE XI l )v l tud 'JPalre^o P unoJ0 q UEJu onrpo Ie]nltlsulq )-teosaleu) pauotgu o]sog' uaq) [ pueu osraur] qxrs oql ur seldr.uExe I EJrpau arll Jo rsII aleldruoJ v .luau 3 1oI saynoSl sllol al)snu'to lut slla) (lselqo^Lua) ]tllalesa snou.l -uorr^uo rElnllotrErlxe rrar{l ur selErpdqoqfff, pup surelord ol fueurrd] o arnl lnl ur uotlerluola]] lC0 €-6 eln611l elueuuedxS puB sllor reqlo ol sllal lreuuof, pu? .usrloqElou alEurprooJ 'uorldtrcsuerl 'slueulelour VNC alelnSor 1ac ezdptec leql saxalduroc-sy1auor saxalduroc ulolo.rdrrpur Jo Surpuelsrapun polrerop e uo e8urq suonecqdde osaql dueyrg. 8urureo1 Jo ere doql eruerrs f,rseq eql ;o suonecrldde leolurp oql roJ uorlerca.rddeu p stuepnls errr8o l eleudordde a;aqm sreldeqr eqt lnoq8norql uo^ol\ e.res elduexe lelrperu esaql .sossasrp ueunq luecrlruSrsJ oqro pu€ JafupJ Jo' sluauleeJl ,/\^.ooul pel azreqlS olorq relntalol'u puE rEFIIel Jrs?qu r secuelpu due141 a)uenelau le)tpaw 'Iuolsds eunrururo lprgalra^ aqf or dSolorq IIar Jeln -relotrr ;o soldrcur.rds arldde (y7 nlderla) .,d3oyounurur1,r, 'uorldaf,ar drosuasp ue Surpur; r{l?d Ipuoxe ur sef,uE^ppls o -tel agr Surpnycur's 11acp azrpnods aseql ;o dSolorq [e] aqr 1o aSerairocp erJruns eprrrord( 97 raldeq3) ..slle] e^raN,, r 'tuaurdole,rep uErI?luruEru (uorlptu.ro; 'uorl€ruJoJur uo snJoJp aseerf,ur {lr^,\ u.rauedq 8norql uorlezr -lrlroJ uro{ luerudola,repJ o sleluoruppunJo r{l sruaso.rd( 77 IEuorlrpPPr .lfnru ot srolfnJlsur pue sluapnls Darrp leql selr ;erdBr{J) ,.luarudola,taq ; o dSolorg IIaJ relnreloW ar{J" r -ru? ^\erleJ e,Lrsueqa.rduropJu € serpnls {reurpuel Jo sef,ue -raJeJ J o rsrl olep-ot-dn uE spr{ retdeqr r{leg 'lprralpru ^\au :sraldeqr rvrau to uorrrppp aqt roJ uoor e>lerrro l sn Surlgeue .sretdeqc eerqt sepnlrur KSoTotg il a) nlrcapw Jo uourpe qrxrs aql Iereles SurzrueS.roa,.trg scrdol Iueur ;o a8ereloc pasnf,ol 'sesJnof, Pu? peulluparls osle ql{r lsoru Jo uorlBzrue8ro sraldeq) /naN eql rorrru PUEI Erraleu polplor alrun 01 slueluoJ Jo elqBl 'drlunoo aql pasr^al a^pq o^\ agl punoJ€ ruor; dSoyorq IIel to srossayord6 91 ue{r erotu r{tr^. uorlellnsuof, uI I p]eogp rpMvu osltNU puuala q-L]o Isa]Jnofl ' talrnuord e leuJap] ue a1eu6.ru lnnoqs1 1a6r 6a ueunq paztl tyal uollezrue6lg nnel reldeqr qlee Jo derupeor peor-ol-dsea ue eprrrordm ou sEurpeeqp atuauo-ldeJuoJ JEOIJ 'alqrssod Jaleregl\ uo8rel Sur;rourer's8urpvaqp ue ]xel aqr par;qdurs 'Oruof, osle e^Eq 3/N 01 sr lpq^{ Jot uorlEpunoJ pllos e epr^ -ord pue arnlrrd 8rq eqr stuapnls e,rr8d eql teql aJnsS ur4rur 'sarerlrJeao r alduqc aql parrordurr aAEq olt ,relncrlred u1 'olqrssalJe arotu sluar.urradxap ue sldaruoc eqr qroq Sunleru ol luerullturuor Jno porJrsuolura lerl e.ryp\ ue 'sleluaurepun; eql Jo suorleueldxe reelJ roJ poau .sluepnls qlr^\ sluaurdo -la^ap lrJnuaps lselel eql;o a8eraloJ Jno arueleq ol peln 'alqeqceordde o^prl o/N aJoru sluopnls ro; Sullunep eg uEf, leqt Ierraleru o{pru ol op uer'sluepnls alenper8 pue olen -perS.rapunq loq to srer.{Jeapl aruarredxes E ,er\,re. q^ pe)se pue IJEq dels e ualel a^Er{o lr uorlrpo qlxrs eql 3ur1r.rmu 1 f1r;rqrssa))pVu e[ 1r.re1p1o seat)ul

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