Molecular Biology of the Gene (International Ed.)

755 Pages·2003·69.74 MB·English

by  James D. Watson

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F I F HT E D IT I O N
JamesD .W atson
ColSdp riHnagr bor
Laboratory
TaniAa. B aker
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ofT echnology
StepheP.n B ell
MascshausIentsttsi tute
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UniveorfCs ailtiyf ornia,
Berkeley
RicharLdo sick
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PEARSON
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Benjamin
Cummings
NOT FOR SALE
BenjamCiunm mings ColSdp riHnagr boLra boratPorreys s
PubliJsihSmem ri:t h PubliasnhSdep ro onrsiEndgi tJoorhI:nn glis
AssocPiraotjEeed coitrtA sl:e xaFnedlrlao wes, EditoDriiraelc tor:G aAnlne xander
JeanZnael esky Editorial DMeavnealgJoeaAprnrm: gne etnitn e
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PubliSsehricvneigIs n,c . Cryssttarlu citmuargee s: LeeTmoro r Joshua-
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rigrhetsse rMvaendu.f acitntu hrUeen di ted oSfAt maetreisc a. This
publiciapstr iootdne cbCtyoe pyraingdh tp ersmhiosusbliedo n
obtaifrnoemd p tuhbel ipsrhiteoorr parnoyh irbeiptreodd uction,
storianagr ee trsiyesvtaoelrtm r,a nsmiinas nsfyio ornom rb ya ny
meanesl,eo cntimrce,c hanical, photoocrlo ipkyeiwnigs,e .r ecording,
Too btpaeirnsm siiotnou( ssme)a teriatlh wifosrr poklm,e sausbem it
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190E0.L akAeve nuGel,e nvIiL6e 00w2,F5 o.ir n foartmiroeng arding
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Manoyf dtehsei gnuasteidom nabsny u facatnusdre elrltseod r iss tin­
guitshhep irro duacrctels a iamste rda demarks. Where those
designaaptepiaiornnt shb ioso akn,td h peu bslhiewra asw aorfea
tradecmlaartikhm de,e signhaatvibeoe nepsnr initne di nciatpisa l
ora lcla ps.
Iyfo puu rchtahsbieosdo wki thtihne UniteCda nSatydaoatu e s or
shoublead w atrheia htta s been wirmoprnotgewfdiu tlhltoyhua etp ­
provoaftl hP eu bliosrth heAeru thor.
PEARSON
-----­ 1 2 3 4 05- VHP6 -70 708 60 590 401 3
Benjamin
Cummings www.abwc-.com
wwwc.shlpress.com
NOT FOR SALE
Preface
s tfihfete hd itoifMo onl ceulBairo glyoo ft hGee ngeo etso
perscso,m pleotfti hohenu magne nosmeeq ueinsnc ole o nger
A
newsT.h iwsa sn ost omethtihnacgto usladf ehlayvb ee en
anticiwphaettneh fider setd itaipopaner eidn1 96e5v;ew nh en the
fouretdhi tciaomnoe u itn 1 978,fe iwa fn fyo rehsoawwq uicwkel y
woulmdo vien tawo o rwlhde rweh olgee no,mn eosjtu st individual
genecso,ub ledv isuaalnicdzo emdp arTehde.hr aebs e eanc ompara­
bllee aipnt he eluocfip draottsietoirnnu catswu erleTslh .u, is nt he
lasfetw y ea,rt she strouftc htheuu rgemeso lecmualcahri tnheast
dritvhebe a spirco cedsissecsu stshebidos o ikn- DtNrAac nrsiption,
replicpartoitsoeynihn,net sainssd,o f or-thahvlea rgbeelesyno lved
att haet omliecva enld, mdaentyao iftl hsei inrn er wroervkeianlgesd .
The needwi toifMo onel culBairoo glyo ft hGee nree fletchtess e
advaensac,n dm anoyt hebress idewsh.e wnBe us ta t dtoopw lna n
thliast veessrti ownew, e raelo lfa m intdh amtu cohf t hoer ganiza­
tioanns dc oopfte h oer igbionoasklh oublerd e taiTnheiwdsa. sn oat
mattoefcr o nven-iiennecveitianlb ilgoyhf,tt h der amatic changes
thahta dt akpelna ce tshliean escdtei ttihovena ,s t obftu hlteke xt
hadt ob ec omplerteewlryia tntyewnaa yn,da ltlh e raerntd ered
afresNho.t, h e reasosniimnptglh yamw toa,rst e h aenv eirnt hgies­
nomiecrt ah,e sreee maen de efdo arb ook etxhpalta wihnaegtde nes
araen d htohwew yo rakn,td h wiasse xacwthlaMyto elculBairol ogy
oft hGee nhea odr igibneaedlnel syi gtnode od.
Thu,ws eh avree sitshtteee dm pttatobi eocnoe mnec ycloorpt eod ic
delivneta ol ldiiesdc ipsluicanhsce esbl,il o glyAo.l swoe, wanted the
newe dittioro ent aaf ionc usp roinn cainpcdlo ensc ,ea pntostfheear­
tureof i tpsr edeceAsnsdso owr esi l.lu storuadrti es cusspsraiionng ly
witehx perimwehnitcashp, p ear imnab ionxleTysh .e sceo nesriad­
tioennss utrhebedo odki d not unbwecidoeyAml.se s tabtyei dta su ­
thoirnt hper eftaotc hefie r setd it"iOfotneI: n presenatn dba,e ­fact,
causoefl acokf s pacIe c,a nnooutt ltihneee x perimtehnatts
demonstirtast etv yaG.li ivdetinh ec hoibceet wedeenl eting an
imporptrainntco irgp ilveia nneg x perimdeentlta,aai mlIi ncltion ed
statthepe r iinpclTeh.ec" u rrienncta rnoafMt olieocnu Bliaorl oofgy
thGee naed heurneasp ologteott hipichsai lllhoyys .o p
Ano utloiftn heni esw e ditwiioltnlh u bsef amitloai nayro wnheo
hauss etdhb eo obke forbee.g (iWinePn a r1tw) i tahs eroifce hsa rpst e
(modiifinet hdce u rreednitt tihoanpt)l atchefie e lodfm olecular
biolioncg oyn tTehxetcs.he a ptseurmsm artihhzeie os rtoyf g enetics
anmdo lecbuilo_laaorng adyl spor estehnteti meclheesmsi pcrailn ci­
pletsh adte tertmhisent er ucatnudfr uen ctoifmo anc romol.e cules
Thtee txhte reiasofr tgearn tiofz oeldla fo awim liflaorw toofp iTchse.
natuorfte h e gemnaetteiricito asrl g,ia znatainodin tm sa intenance,
ardei scuisnsP aerd2t ;i na ddittioco hna ptoenDr NsA s truc,t ure
replicraetcioomnbo,in an,na drte ipanierwt, ot hpiasr otft hbeo oiks
a chapotnec rh romoscohmreosm,a atnidn ,nt uhcel eoTshoimse .
additrieoflnec cutrsr aepnptr ecoifah toiwto hnce o ntienxw th ich
ag ivgeenni efs o uinndfl ueintcfseu sn catnido nr egulation.
NOT FOR SALE
Thep assaogfe oirnamftifoornmg enteop reoitns-o-caglelneed
expre-sissic oonveirnePd a r3t; anPda r4it wn e d escrtihbee
reguloaftt ihoapntr ocAessw se.la lsc haptoentr hsbe a smiecc ha­
nismosfg enree guolnaP,ta ir4th acsh aptoentr hsre e guloaft ion
geneex presisnai noinm daelv elopamnedin ntt h eev oluotfi on
animdailv etryTs.h iecseh aptaegracsio non rfmt oat radietsitoanb ­
lishbeyed a rleideirt iaolnwsat:yh se hraebs e eanc hapotret rw o
linkbiansgmi ecc hanoifsm mosl ecbuiloalrto opg rye sbsiinogl ogi­
caqlu estiIontn hsec. u rreednitt tihoencs,he a ptienrvse stigate
perhtahpmeso sstt rirkeivnegl taotc ioomnfeor mc ompartihneg
complgeetneo me seoqfvu aernicaoenusis m :ad lifsfreenatn imals­
incdliunhgu mnas-contlaairng tehlseya mgee nes soad niedfr f­
encbeest wetehno asnei mmaulssrt e sullatr gfeorlmyc hangiens
how those egxepnreess saerde.
New toc utrhreeed nitt iisto hne fipnaarPlta -r5t- cmoprising
chaptoenre sx perimmeenttha-oltdh set echnioqfum eosl ecular
biolgoegnyo,m aincdbs i,o oirnamftci-sando nt hmeo del organisms
whosset udhya sr eveamlaendyo ft heu nderlying opfr inciples
molecbuiloaylr.o g
Wea llutdoet hdee x ploisnti honenu mbeorfas t omsitcr uctures
solvientd h lea st feTwh eysieena crlsnu.od oten lmya noyf t he
enzymtehsam te ditahtbeea spirco ceosfms oelse cbuiloaylr,ao ngd
manoyf t hper otethianrtse gulthaotpsere o cebsustteh se, nucleo­
somaesw elWlh.i ilrtee matirnutseh amta noyf btahseci ocn ceipnt s
molecbuiloalrco agbnye u nderswtiotohdro eulti oanns cter uctural
detla-iindeietid so noe f tshtree nogftt hhfiese ltdh atth iisst he
ca-sneeveerltehmsasn,my e chaniinissgthictcos m e foornmls ye eing
thesdee taAiclcso.rg ldyiw,nh erset ructsuhreledis g ohnth ow
thmeo leciunql euse stwioorwnke ,p restehnetam n;wd e d os oi na
consissttytelhnert o ugthhboeou otk .
Eacpha rotpn eeri ncluads ehsot retx t, owuhtalwtii nlbilen g
coveirnet dh e cocmhianpgta enrdasf ,e wp hotogrTahpehspsei. c ­
turfreosm,t hCeo ld SHparribLnoagrb oraAtrocrhyiw evreae,lt la ken
att hLea boraotnLo ornIygs latnhgder, e maatj oartit htSeyy mpiousm
hosttehdea rlem oesvtes ruym mseirn 1c9e3 C3a.p tiioednnsytw ihfo
isi ne acphi ctaunrwdeh einwt a st akWeent .h aCnlka Bruen caen d
thCeS HALr chfiovhree lwpi tthh ese.
Parotfts h ceu rreedninttg iroeowu otfa ni ntrodcuocutrosrey on mo­
lecublilaoorgt ya ugbhyot n oef (uRsLa )tH arvUanridv erasntidht iys,
authiosgr r atteoSf tuelHv aer riasnoJdni W ma ng wchoon tritbou ted
thciosu risnpe a syte aarnsd wihnoflsue einsrc eefl ecitne dC h6a pter
ande lsewWheeh raev.se h owsne ctoifot nhmsea nusctrovi aprti ous
colglueeaasn d tchoemimreh natvbsee emno svta luagbrleea,t ly improv­
intgh aec curaanacdcy c essoiftb hitele iaxtntfiyd g uersS.p ecifiwcea lly
thanJka:mC iaet Rei,c hEabrrhdit Mg.i kEei sCehnr,Fi rso mmIerH,aa ll,
AdriKarna iKnaerro,Ll uignBe riM.lc lG inMnaitMsti, c hLaieMlliy,r els,
NipaPma tCerla,Pi egt erMsaornPk,t ashUntet,aR ma jBhanadnadr y,
BrucSet illImnaa dnd.i tion, Craitgh seHe ucntotineot rnh ed rafted
worfmo Crh ap2t1eW.re alsot htowhshaeonp kr oviudswe idtfi hgu res,
ort hweh eerwitthoca rlelta hteeim n,c luSdeianCnga o:rl rSle,tD ha rst,
EdwaErgde lmGaeno,Hr agl dSetru,Ka irmtB ,i ll McSGtienPvnaeid s­,
docPkh,o eRbiecM ea,tS tc oPtett,Se orr gAenrd,rS zteajs TioamSk t,e itz,
DanVo ytaasn,Sd t eWvees t.
We armeo sgtr autlte foL eemJoors hTuoarw- hor endearleld
the strfugicutrueorsfe,t epnr oducmiunlgt ivpelres ions and
patiehnetllpyui sns gew eh icbhe s�t()'f e��'gA(_� needWeed .
arael sgor autle tfthoo wsheop rovitdheesdio trfw a1rP:ee Krr ualis,
RobeErstn oEutfh,aM ne rraintdBt ,ar rHyo niCgo.o rdiwneartee s
obtaifnoremd t heP rotDeaitnBa a n(kw wrwc.sb.bo/r;)ga /npdd
citattioto hnosws heo s oldv eeacsht rucatruiern ec luidnte hde
figluerge ends.
Oura rptr ogrwaams devaenldor epnedde brye tdaa lenatnedd
enthustieaasfmtr ioctm h Der agoMnefdliyGa r o,ul pebdy M ike
DemaraanyCd r aDiugr aRnetn.aH teel lmhieslspt eodd e vesloompe
ofo uirn istkieatlc ahnedps r oviedaerrdle yn deroifan n gusm boefr
figurTehsec. o viemra wgaes r endebryTe orndo N arashfoirmma an
authcoorn csekpettb cyE hr iBceaa (dMeB GCr pahics).
We thtahnokas teC olSdp riHnagr bLoarb oraPtroerwsyhs o
handdleevde lopomfte hnbitos o Jka.An r geinntdee,s phiatveit nog
enfotrhcdeee adlwianset sh,r ouglheoscusat j otlhiansngh wea s
tireleensgsaligyneh de lpuissn ogl tvhepe r obltehmesps ree sented.
MarlyiD zi ckekrespootrn g anizmeadso sfmt ahtee wreig aeln erated
anNdo rRai chee lpceodo rdianutahtmoeer e tianngds oatshpeerc ts
oft hper ojDeecnti.Ws eei sasn dE dA tkesporno dutchecedo ver
desiagnndJ; o hInn glwihsoi, n ititahticesod l labowraasot ni on,
hanwdi tahd viactce r itpiocianlit nts h per oceMsosso.tf a ll,
KaarJeann soseuenrd, i tkoerpe,tv erytahfilnwogia ttah n e nergy,
enthiuassamn,d a ctifvairt y beyondw eac noyutlrhdei ansgo n­
ablhya vaes kfeodtr h;i nsgism pwloyu lndoh ta vgeo ttot hipso int
withhoeurt.
Wea lswoi sthoa cknowltehwdeog reok ft hoastBe e njaCmuimn­
mings who cporoordduicnotaftit hoebendo oFkr.a nRkug girello
oversapwr otchees s cbayrJ riimSe mdi tKohau,yUt e noC.or inne
BensAolne,x aFnedlrlao JweeasnZ,na el ye,s kDaonndn Kaa lIanlg.r id
MounattE lmS trPeuetb lisSheirnvgic coepsec dh euelrlwfyi tthh e
manryo unodfs chtanoag raetns td e xt evleanit netv heperr yo cess.
MichSeolredw hii,pl aer otft hBee njaCmuimnm intgesah me,l ped
briunsag l tlo geitnh efitrrh spetl ace.
Andfi nalwleyg r ateafcuklnloyw loeudrfgma ei liaensdfi rends
whot,h rougthhoipuset r ipordo,v isduecsdht rosnugp podrets,p ite
havitnopg u utp w itohufr re quaebnste nacneddsi stractions.
JameDs.W atson
TaniAa.B aker
StepheP.n B ell
AlexanGdaenrn
MichaLeelv ine
RichaLrods ick
1 PerK raulgirsa ntepde rmissitooun s eM olScri(pKtr aulPi.}s 1 .9 91M.O LSCRIPTA:
Journoafl
progratmo p roducbeo thd etailaendd s chematpilco tosf p rotesitnr uctures.
ApplCireyadsl tlpoghrya
24:9 46-950R)o.b erEts nougfa vep ermissitooun s eB obScript
JournaMlo loefc Gurlpaahri cs
(EsnouRf. M.1 997. 15:1 32-134I)n.a dditioEnt,h an
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Merrigtatv eu su seo fR aster(3MDe rriEt.tA .a ndB aconD .J1.9 97R.a str3D:P hotoreal­
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grantepde rmissitoonu seG RASP (NicolAl.s,S harpK .A.a,n dH onigB .1 991P.r otein
foldianngd associaItnisoing:hf trso mt hei nterfaacnidat lh ermodynampirco pertoife s
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NOT FOR SALE
Aboutth eA uthors
JAMESD .W ATSON wasD ireocftC oorlS dp riHnagr bLoarb oratory
from1 96t8o1 99a3n idns o wi tPsr esiHdees npteh.ni tus n dergraduate
yeaarts Utnhiev eorfsC ihtiyca angdroe cehiivPeshd . iDn1. 9 5fr0o m
IndiUanniav eBrseittwy1e.e9 n5a 0n1 d9 5h3ed, i pdo stdoctoral research
inC openhaangCdea nm br,iE dnggeland.a tC Wahmiblr,eih debg eegan
thceo llabothraartte isouinl ntt ehdee l ucidoaftt hideoo nu bllei-chael
strucotfDu NrAie n1 95(3F.to hrd iiss coWvatesroyFn,r, ca niCsr iacnkd,
MaucreWi ilkinsa wwaerdrteehNd eo bel iPn1r 9i.6zL2)ea tien1r 9 53,
hew enttot hCea olrinfIinas toifTtec uhtneoo glHy.em ovetdoH arvianr d
195w5h,e rheet augahntd rdeisde oarnRc NhAs ynthaensdpi rso tein
synthuenst1ii9sl7 H6ew. a s fitrhsDeti reocftt hoNera tiCoennatlfe orr
GenoRmees eaorftc hNhea tiIonnsatli otfHu etaelfrsto hm1 9891 9t9o2 .
DrW.a tswoanss o aluet hooftr hfi er sste,c oanndtd,h ierddi toifMoo n­s
lecuBliaorlo ofgyt hGee en,a ndc oa- auotfth hoferuo r tehd itTihoens.e
werpeu bliisn1h 9e6d1 59,7 0, an1d19 9786r7,e specWtaitvseholanys .
albseoe inn volivnte wdoo t hteerx tb:ho ewo akossn oef otrhieg aiun­al
thoorfsM olceulBairo loogyft heC e,la lndi sa lsaon a uthoofr
ReconmabniDtN Aa:S hoCroutr se.
TANIAA .B AKER its hWeh itehPeraodf eosfBs ioogrly ao tt hMea ss­
achusIentsttsio tfTu ecthen olaongday nI nvestoiftg haHeto owra rd
HughMeesd iIcnaslt iSthurete ec.e aiB v.eiSdn.b i ochefrmoimst threy
UniveorfsW iitsyc onMsaidni,sa onnda,P h.iDnb. i ochemfriosmt ry
StanfUonridv eyi rns1 i89t8H.e r grraedsueawataresc h caoruritin e d
thlea boroaftP orroyf eAsrstohKruo rr nbaenrdfgc o usoendm echa­
nisomfsi nitioafDt NiAor ne plicSahtdeii opdno .s tdorcetsoeraarlc h
int hlea boroafDt ro.r y KiMyiozsuhuaictt h hiNe a tioInnaslt iotfu tes
Healstthu,d ytihmneeg c hanainsdm regoufl aDttNirAoa nn sposition.
Hecru rrreenste eaxrpclhmo ercehsa nainsrdme sg uloafgt einoenrt ei­c
combination, epnrzoytmueenoi-flncd aitanangld,y P zA-eTdde pendent
protdeeignr adationB.a kPerrreo cfeeitsvhsee2od 0r 0E1l Lii lly
ReseaArwacrhfrd o mt hAem eriScoactniy oe fM icroobgiayon ldt he
200M0I TS choooflS cienTecaec hiPnrgi fzoeUr n dergraduate
EducatSihoien sc. o -au(twhioAtrrh t hKuorr nboeftr hgbe)o oDkN A
RepclaitiSoenc,oE nddi tion.
STEPHENP. BELLi sa PreosfsoofrB iloogaytt hMea ssachusetts
InstiotfTu ecthen olaongday n A ssistant oIfnt vhHeeos wtairgda tor
HughMeesd iIcnaslt .i trHueetc eeBi.vAedd.e g rfreoemts h Dee partment
ofB oichemiMsotlreycB,u ilogalyrao, n Cde lBlio oglayn tdh Ien tegrated
Sciences atPN roorgtrhawmeU sntievrenar nsdiP thya. iDnb. i ochem­
istartty h Uen iveorfsC iatolynri ifBae,r keiln1e 99y1 H.i gsr aduraet­e
searwcahsc arroiueitdnt hlea boroafDt roR.ro yb eTrjtia annfd co used
one ukoatrtiyrca inpstcriHoedn i.pd o stdoctoriantl h lera ebsoeraar­ch
toroyfD rB.r uSctei laltCm oalnSd p riHnagr bLoarb orawtoorrkyoi,nn g
thien itioafet uikooantr iyDcN A replHiiccsau trirorenen.st e afro­ch
cusoens mtehec hanciosnmtsr tohldelu ipnlgi coafet uikooantr ciyhcr o­
mosomPerseo.sf sBoerl l retch2eei0 v0Ae1Sd B MBc-hSerPilnogu gh
ScienAtcihfiice veAmwaernadtn dth Eev erMeototrB ea kMeerm roial
Awarfdo Erx celilneU nndceer grTaeadcuhaiatnMtegI Ti n1 998.
NOT FORS ALE
ALEXANDEGRA NN isE ditoDriiraelco tfCo orl Sdp riHnargb or
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NOT FOSR ALE
BriCeofn tents
P A R 1T CHEMISTRYA ND GENETICS l
1 TheM endeliaVine wo ft heW orld 5
2 NucleiAcc idCso nveGye netiIcn formati1o9n
3 TheI mportanocfWe e akC hemicaIln teracti4o1n s
4 TheI mportanocfHe i gh-EnerBgoyn ds 55
s Weaka ndS tronBgo ndsD etermine
Macromolecular Str6u9c ture
PART 2 MAINTENANCEO FT HE GENOME 93
6 TheS tructuorfeD sN A andR NA 97
7 ChromosomeCsh,r omatiann,d t heN ucleosom1e2 9
o TheR eplicatoifoD nN A 181
9 TheM utabilaintdyR epaiorfD NA 235
to HomologouRse combinataitot nh eM oleculLaerv el2 59
11 Sitep-eScifiRce combinatainodnT ranspositoifoD nN A 293
PART 3 EXPRESSIOONF THE GENOME 343
12 MechanismosfT ranscripti3o47n
13 RNA Splicin3g7 9
14 Translatio41n1
15 TheG enetiCco de 461
ix
NOT FOR SALE