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Ann Rev Cell BioI 98 : 89Copyright 98 by Annual eviews nc. rights reserved

CHR GRGATN

N AND

ndrew W urray and ak W zostak

Depame o Moleula Biology, assahuses Geeal ospial,Boso, Massauses 021 4

NENS

INTRODUCION

A MODEL FOR CHROMOSOM BHAVIOR DURNG TOSIS AND IOSIS

ThMitoic Cll Cyclnterphase

MetaphaseAnaphase

MeiosisXPRNA VINC .................

ttc C Cyc ......................Time f kinetchre duplication Centrmeric DNA replicatin DNA replication and catenation Prometaphase chromoome movement Micrmanipulation f metaphase chrmsomes Anaphase Artcial chromosmes

Mi:...Pairing and disjnctin Yeast meitic mutants l meiotic mutants Meiosi veu meioi

VOUON O MIOSIS

INTRODUCTION

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3

The aihul iheiae o eei omaio epes o he oelysegegaio o homosomes i miosis a meiosis Mioi segegaiopoues geeially ieial aughe ells, hile meioi segegaiopoues ells ha oai oly oe membe o eah homosome pai ha

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290 RRY & SOSK

as pese i he paea ce homosome segegaio is a exemelyaae pess: hmsme oss i misis i yeas us e i eey105 iisios 1 hile abeaios i meioic chomosome seegaiooccu oce i eey 104 eioses i boh yeas 2 a Dphil (eiee

i ).Rece eies hae iscusse miosis ih efeece o mechaisms of

chomosome moeme (6), he oe of membaes i he behao of hemioic spile 7), a he sucue a behaio of kieochoe (8). Thegeeic cool of meiosis has bee eiee fo yeas ) Dphil (), aohe ogaisms 10). Uasucual aspecs of meiosis hae aso beeeiee ( 1 1 1 2) e o o aemp o eie he oumious ieaue oal aspecs o miosis a meiosis hee, bu coceae o he fuameal

pobem of chomosome segegaio he aue of he mechaisms haesue ha ise chomais (i misis a he seco meioi iisio) ohmogus hmsmes (i meiosis segegae o pposie poes f hespile e emphasie suies o geeicaly acable ogaism, especi-ally bake's yeas (acchamyc cviia) e use he em kieochoeo escibe he sucua compoes ha ae assembe o he ceomeic A eseig he em ceomee fo he A ise e use heem spie poe boy (SPB) o efe o he geeal oep of spile poeogaiig aciiy, alhough he mophological B is fou oly i fugi

I aima cels he ceosome is he micoubue ogaiig ee.Miosis a meiosis hae bee suie i may ogaisms usig a ie

aiey of echiques e hae ie o iegae ou iscussio by co-sieig he expeimea eiece i he oex of a specic moe fo

chomosome segegaio The moel ess o a syhesis of ieas ha haeal bee pu foa a leas oce by ohe okes. ix ceal cocepsuelie he moe:

1. omal chomosome segegaio is epee he iegiy o

mcoubules (eee 4 a 52 Sepaae mechasms exis ha a geeae foe chmosomes

eihe oas o aay fom he spie pole o hich hey ae aache(eiee i 4

The sabe aachme of a chomosome pai o he spie eques hegeeaio of fces ha pull he o kieochoes of ha chomosmepai oas opposie poes f he spile, ceaig esio a eiblelage beee e o chmosoma uis o be segegae

4 he epiai of A moecues ca geeae iage f he augheA mecues i a chomosome by ieiig (caeaio) of heA upexes aeaio ca be esoye uig aaphase by heaciaio of ype II A opoisomeases, hich caale he passage ooe A upex hough ahe (eiee i 18

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CHROOSOE SEGREGON N OSS ND EOSS 29

In mot oanm noma aton o homoo at mo I dndnt on ntc comnaton 1.

6 Th dnc twn th hao o t chomatd n mo Iom tha n mto and mo II th t o dnc n th

oanzaton o th chomoom ath than that o th nd 6.

A MODL FOR CHROMOSOM BHAVIORDURING MITOSIS AND MIOSIS

Th oown mod hghy cuatv ut ha th advantag o o-dn a und nttaton o a wd vaty o data and o makntta dcton

he ioi el yleP Ou mod o th moc c cyc hown n gu th nnn o th c cyc ach chomoom ha a n kntochoW oow ndn & ahay n oon that th conton that an a o ntocho a t a ut o th nkatwn th oducd y th ntcatnaton o t chomatd 19.Intcatnaon a whn du DN catd n th anc otoooma actvty. Und th condton ach douhca tun o

th anta d contd nto on nttwnn o th dahtmoc F 2 undn & ahay ood that na a atth ont wh caton o mt a a ut o th tc cuon oDN toooma y th caton com 7.

ntcatnaton ta om to mto thn ach chomatd houd catnatd to t t at ach o th many ont wh caton okmt. taty mot ntcatnaton cod od dun G2. Inth ca th catnaton that w oo nk t chomatd togth atmtaha coud gnatd y th at caton o th cntomcDN 19 chum & Caon ood that th yat cntom acta a ock to th aa o caton o 20 Ra o th oc atoha woud aow caton o th cntomc DNA wth tntono t chomatd nkag a a oca catnaton o th DN at hcntom.

EPHSE Ou ooa o th mchanc o chomoom movmntdun mto ad on a cnt vw y Pcktta t a Mcotuu n th two ha o th nd ha oot oat anda tho wthn on ha ha th am oaty 2, 22 ca o thod oaty o th nd th a no ndamnta dct n

nvntng chm that w uy th ctoa chomoom movmntqud at anaha

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292 Y & SZOSK

Pcka a 4 uggd ha th knocho can ntact whmcouu n wo ontaon In th oa P ontaon th knocho ud owad th nd o om whch h mcotuuogna W ugg hat attachmn n h P onaon ac a a

mocua atcht: A ong a h P oc ood by a oc acting n thooie ieion men i be. e ineoore o wo ie

chomad a aachd n P onao to mcotuu om ooto o th nd hy w mo owad h o o whch hy a

a

Figure A model fo mitotic chromosome segegation. (a) DNA repication eads to twsiste chomatids attached to each othe by catenation (each sister chromatid is epesented b

a singe ine (b) Metaphase: stabe attachment to the spinde is achieved when thkinetochoes ae attached to opposite poes (c) Anaphase DNA topoisomease activit)decatenates the siste chromatids and aows tem to move to opposite poes of the spinde

Symbos tiangle kinetochoe; cicle spinde poe.

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CHROOSOE SEGREGON OSS ND OSS 293

Fgue 2 Gnton of tnton Whn two plton fok t th lt - tu ofh hlx nw h h pl

che un he lne beween hem reche o h he P orce onone neochore oppoe b h cn n he oppoe recon on

er. Achmen of neochore o mcrobe n he oppoepor el he npolr (AP) orenon. In h orenon heneochore enere force h move w from he pole from whchhe mcrobue orne. Becue l chmen mephe re n he orenon chmen n he A orenon m be nble

NPHSE We propoe h urn mephe he epron o erchrom prevene b her cenon We ue h feebcmechnm e h cve DA opoomere II onl fer ll he

neochore re che o mcro ble n he orenon Thecvon of opoomere mr he rnon from mephe onphe he erucon o he cenon beween er

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294 UY & SOSK

cromas a allows em o searae an move oewars uner enuence of e force.

eisis

We arue a e meoc cel cyce reresens a moc ce cyce moeby e aon of a secon ucaon of e SB wou an nervenn Sase aown wo rouns of cromosome sereaon o occur n a snece cycle. Seccaly, we roose a neocore ucaon ancenromere relcaon s eaye un e comeon of meoss . Durnmeoss II e cenromere DA s relcae, eavn e neocoresaace o eac oer by nercaenaon, exacy as ey are n mocmeaphase.

We roose a e roer sjuncon of omoos n meoss I asoreures nercaenaon beween serean cromosomes, an a snae arses as e resu of recombnaon If e sser cromas amae u a omoo are caenae a many ons as a resu of e cosonof recaon fors n meoc S ase en any recroca crossover (buno ene converson evens beween nonsser cromas wl eneraenae of e omoos (ure ) Aer recombnaon eac recombnanchromatid w be caenaed to its sster centromere roximal to the oint ofrecombnaon, an o s nonsser sa o e se of crossover. (We

ene e sse of a recombnan croma as ben e croma wwc sares a neocore n meoss .

If omoos are ne by nercaenaon, s lnae mus be esroyeby e acvaon of a ye II oosomerase a e meaaeanaaseranson of meoss I. Te oosomerase acvaon w aso esroy e

nercaenaon of sser cromas. We roose a e nae sreenerae a meoss II as e resu of recaon of e enromercDA aer e comleon of meoss .

Te eaye relcaon of e neocores srucural eemens an e

centromec D means that at meioss I the homologs each have only oneneocore an no searaon of sser cromas can occur (ure Ins sceme neocore ucaon comms e ce o a moc vson.

Two mocaons are reure o allow meoss o evove from moss( nroucon of a recocous SB ucaon a alows meoss I ooccur before e mosse meoss II an e suresson onecore ucaon un afer e comeon of meoss I. If ereuaon of SB ucaon an sne assemby n meoss I s eren

from a n meoss an moss we exec o n muaons a boce assemby of e meoss I sne, ean o meoc vsons n wcrecombnaon s normal bu meoss I fals o occur Suc muans are n

fac nown )

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CHOOSOE SEGEGON N OSS ND EOSS 295

MEIOSIS

ISlsnFgue 3 A mdl f hmsm sggan n mss n mss w hmlgus pasf ss hmads a shwn Eah ss hmad pa has a sngl knh and hnm DA has n bn plad (a) Rmbnan lads h anan f ssand nnss hmads, pvdng a xbl lnkag bwn h w pas f sshmads () aphas sabl aahmn h spnd us whn h knhsa aahd pps pls f h spnd () Anaphas DA psmas dsys h

anan allwng h ss hmad pas sga pps ps f h spnd(d) Rplan f h nm DA gnas a pa f anad ss hmads (nyn f h w pdus f mss s shwn) (e) aphas ; () Anaphas DApsmas danas h ss hmads allwng hm sgga pps plsf h spnd. Symbls: tange = knh; e = spnd pl

XPRIMNTAL VIDNC

he o ell yle OF KNTOCHO PLICION We sues a suura om-onens o e kneoore bu no e enomer DA ae ulaeurn G 1 aou e mn of kneooe uaon s no essena

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296 MY & SZSK

o the integity of the mode Thee is no die assay aailable fokinetohoe dupiaion. mmunouoesene studies on mammalian es

using anikineohoe sea fom paients suering fom auoimmunediseases show that the kinetohoes appear as single sruures in mostinephase ells 25, 26). n a small faion of G2 els he kineohoesappea as pairs of osey apposed dots 26) Thee ae appoximaey hesame number of paired dots as here ae homosomes in he el ine,suggesting that he staining reeas pairs of sister kinetohores.

n the fungus Spge kineohores are isible throughout th elye and ae aways assoiaed with mioubues 27) tron miosopy eeas that he morphologia dupliaion of the kinetohoe ous

immediatey before mitosis 27) Howeer it is possibe hat the sruturalomponens of he kineohoe aualy dupliate muh ealie in he ellye, but hat the morphologia dupliation is no isible until heenomei DNA epiaes during G2 Beause the kineohoes of Sceeve ae no isibe, een in eeon miogaphs we do no know hey emain assoiaed with miroubues houghou metaphase, ahoughnuea mioubues ae pesent i his peiod eiewed in mmunouoesene of yeast es wih anitubuln antisea oes not show

extensie miotubue arays in the interphase nuleus 2) suggesting thatin inephase he miroubues may exend ony a sho distane fom theSPB.

Thee is no ompeling reason to ague against the dupliaion of hesuual ompnents of the kietohoe in S o in ealy G2 although it isinteresing o noe ha during miosis of es in whih endoreupiaionhas ourred the siser hromaids ha aise fom the extra round of DNArepiaion fai to segegate fom eah other 29). This suggess that his paiof DNA dupexes shaes a single kinetohoe, and thus hat kinetohoedupliation does not ou during S phase. The yeast el ye muantcdc may also be efetie in kinetohoe dupiation At the nonpemissie emperatue the dupliaion of the spinde poe body whih nomalyous in G, is boked 0) Howee he SPB ineases in sie, and hepoesses of DNA repliaion and he iniia stages of spinle formationour as norma 0). The els ares in nuea diision and eleronmirosopy shows tha a normal spindle forms tha has a SPB ony a oneen 0). This raises obious quesions abou he roe of he SPB in spinde

fomation hen cdc els ae gown at a semipemissie tempeatuehei poidy ineases 1) This suggests that unde these onditions sisterhomatis fai to separae so that al the homosomes go o a singe pole.e suggest tha the CDC gene podut is equied fo kineohoe as wellas SPB dupliaon, onsisten wih a simultaneous dupliaion of he SPBand kineohoes in G1. One unesoled quesion is whethe i is he

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CHOMOSOME SEGEGTON MTOSS N OSS 297

mohe o daughe ell ha eeies he homosmes when a cdc saindiides a he semiemissie emeaue

Anhe yeas gene pdu ha may be equied f ineheduliaion is ha of he gene. In oldsensiie dc muans he

loidy of he ells ineases when hey ae gown a semiemissieemeaues. A he nonemissie emeaue ais of ells aumulaein whih one f he geny inheis all of he DNA as judged by saningwih he DNAsei dye DAP alhough he sindle aeas nomal byaniubulin immunouoesene he ell ha inheis he DNA an beeihe he mohe o he daughe ell J hmas & D Bosein esonalommuniain) he behaio of his muan is omaible wih hepssibiliy ha lak f he funinal NDC gene podu bloks kineo-hoe duliaion s ha sise homaids fail o sepaae.

Kinehoes may emain aahed o ne of he SPBs houghouinephase. his wuld pide an aaie eplanain f he segega-on of omads whse odes sand s o he same age whh has beenbseed in yeas () Neup () and sme mammalian ells (4). Ifineohoe duiain was nseaie suh ha he de inehoesemained aahed boh he oldes DNA sands and o one of he wSPBs hen all he ld inehes wuld segegae o he same spindepole aying wih hem he homaids ha onain he oldes DNA

sands In yeas he mohe and daughe ells wee equally likely o inheihe old se of hmaids (),whih suggess ha he SPB aahed o heold kinehoes is equally likely o be inheied by eihe ell; his isnsisen wih u expanain f he behai f he d muan

CENTOMIC N ELCTON he ime a whih dieen homosomalegions eliae has been inesigaed in lan and animal ells. In genealhe eno mees and elomees ae lae eliaing eiewed in 5Hwee sine hese egions ae also he sies of saellie DNA sequenes

(6) whih ae en lae eliaing (6) i is no ssible ell if hefunional enomees and elomees ae hemseles lae eliaing.

N ELCTON N CNTON Sundin & Vashasky 17) de-monsaed ha he eliaion of he animal ius SV40 oduedaenaed dimes hey suggesed ha sise homaids ould also beomeaenaed as a esu DNA eaion and ha his migh hel diemii hmsme segegaion 19). isomease II was shown beequied fo he sepaaion f baeial hmsmes in i and aenaed

homosmes oud be indued seaae in i by he ain ofpisomease (8). ould he same esul be bained wih euayoihomsomes isolaed fm meahase ells?

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298 MUY & SZSK

n h ya p muan whih arri a mpraurnii DNAopoiomra ll arr uring nuar iiion a h nonprmiimpraur 9. n adiion h 2 m ir bom highy inranad39 No inranaion of h irl i n a h prmii mpraur

39 whih ugg ha h halflif of ana form i hor anhrfor ha danaion our prior o anapha. f ir hromaiinranaion gnra uing S pha uri from h n of Shroughmapha h rmoal of anaion by opoiomra aiiymu our muh mor lowly for ong linar DNA moul han i ofor hor iruar on.

By prforming hifup an hifown xprimn on rain arrying ampraurnii top2muaion olm & D Boin ha hownha h aiiy of h TOP2 gn prou i rquir a mioi pronaommuniaion Th xmn do o ra whhr opoiomra i ao ai a ohr poin in h ll yl. n h abn 0opoiomra aiiy h ll arr a mioi an l dah rapilynu Th ah of h mioi ll an b prnd by ainnooazo a rug ha inhibi ubulin polymrizaion bfor hiinl o h nonprmii mpraur c Holm & Boin pronalommuniaion Thi ugg ha ah i u o hromoom braagor anuploiy ha our whn l amp o nr anapha wihanad ir hromaid.

hn diploi ya l ar rad for prolong prio wih hmiroubu dpolymriing rug mhyl bnzimaazol arbamaMB a high fraion of anuploi ll ar n in ll ru on rugfr mdium 0 On inrpraion of hi nding i ha anupoidy aribau h anaion bwn ir hromai lowly ay in haar arr in mioi allowing a fi anapha paraion pro our f hi ida i orr top2 mun houd how a muh owfrquny of anupoidy whn hy ar arr in mioi by MB a hnonprmii han a h prmii mpraur.

A mapha in iaom h ir inohor i o o hirpi po parad from ah ohr by abou 0 m Thuh proximiy of h inohor anno b wha alow hm o brogni a ir Th hypohi ha ir hromaid inranaionormaly hol aughr DNA mou oghr a mapha proixay h or of xibl long rang onnion hi obraion ruir

S CMS MVMN iHap an hi olaboraor ha mad xni udi of promapha hromoom mo-mn in iaom whih o h onp of an A for riw in rom xprimn uing ohiin an drug ha prn AT gnraion

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CSE SEGEGN N SS A ESS 299

hey drew he folowing onusions

1. Aahmen o he kineohores in he P orienaion eads o hegeneraion of P fore and he moemen of he kineohores owards he

poe o whih he miroubue is aahed P moemen does no requireAP and an our in he abene of miroubue 42 43)

2 Aahmen of he ineohore o miroubue in he AP orienaioneads o he generaion ofAP fore and he moemen of he kineohoreaway from he pinde poe o whih he miroubule is aahed AP

moemen requires ATP and ina miroubules )3. n promeaphae he iniia moemen of he kineohores is aways

poeward hi ugge ha if he ineohore remain aahed omiroubues during inerphase hey do so in he P orenaion 4)

Aahmen of boh kineohores in he P orienaion o opposie poesofhe pinde doe no neeariy peify he poiion of he hromoomeon he pinde The hromoome wi ome o re only when he foreappied from opposie direion are equal Osergren ) proposed ha ifhe fore on a kineohore were proporiona o is disane from hepinde poe hen hromosomes would ie preisey beween he poe a

meaphae Thi idea wa eed by examining he posiion of riaensfomed in permaoye from graopper whih had been reaed wih

Xray) a meaphase of meiosis 5) A equiibrium he wo kineohoreshafae one poe mus oleiey generae no more fore han he ingeineohore ha faes he oher poe The kineohoreopole disane forhe single kineohoe wa wie ha for he wo kineohores ha faed heoppoie poe as prdied by Osergren 4). is diul o see how foregeneraed a eiher he kineohore or he spindle poe ould be aeed byhe diane beween hem This suggess eiher ha P fore i generaedalong he engh of e miroubues or alernaiely a i rpresens someeasi omponen o whih he kineohore is aahed wih he miro-

ubule aing as he guide raher han he agen ha ommuniaes fore ohe ineohore he eney independene and he irouue indepen-dene of P moemen make he aer possibiiy araie

An example of an eai proein i he onraie proein pamin foundin erain roifers. he roifers are arried on onraie saks omposedargey of pamin whih hange from a highyexended o a gobuaronformaion in repone o an inreae in he aium onenraion. Thionformaiona hange produe a rapid onraion of he ak 46)Poibly a imiar eai proein generae he Pfore in he spinde

ICNITIN F ETSE SES he abiiy o miro-manipuae hromoome ha made poie o direy e he ee o

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3 Y & SZOSK

ehing he inekineohoe inkage on homoome oienaion andegegaion So fa hee expeimen have been poibe ony in meioi

e n an eegan eie of expeimen Nika and hi oeaguemiomanipuaed he homoome of gahoppe pmay pemao-ye in meaphae of meioi 13 1447) The fomaion of hiamaa inkhe kineohoe of a pai of homoog wih he homoome am fominghe inkage By deahing one of he kineohoe of a pai of homoog

fom he pinde and moving ino he oppoie haf of he pinde bohkineohoe wee aahed o he ame pinde poe unipoa oienaion)Afe hi opeaion boh kineohoe moved owad he poe o whihhey wee aahed uni one kineohoe deahed fom he pinde. Thedeahed kineohoe woud eaah a andom o be fom eihepinde poe. Reaahmen ha mainained he unipoa oenaion haeued fom he oigina manipuaion wa unabe. Ony when hekineohoe eaahed o he poe fom whih i had oiginay been

deahed did he wo kineohoe mve owad oppoie poe and hehomoome oienaion beome abe. Suh eoienaion even wee noe ikey u nea he pinde pe han a he equa 1 4)

he eoienaion ud be pevened if he neede ued fo momanipu

aion wa ineed beween he homoog and ued o appy foe owadhe pinde poe ha wa no aahed o he kineohoe 15) The impeinepeaion of hi eu i ha pinde aahmen beome abe onywhen he inkage beween he kineohoe i ehed. The high fequenyof eoienaion een in pemaoye eoveing fom od eamen

oud ao be pevened by eing he inkage beween he kineohoe47) n addiion univaen eihe ex homoome in he heeogameiex o auoome ha have no hiamaa) have been obeved o eoien

oninuay duing he meioi diviion 4).The aniion fom meaphae o anaphe aied o ou in e inwhih one homoome had been deahed fom he pinde 13) Thiugge ha he e i abe o monio he aahmen of kineohoe ohe pinde. A one exampe he kineohoe oud have a poein kinaeaiviy ha phophoyae and heeby inaivae opoiomeae Thaiviy migh be expeed ony in mioi and be inhibied when hekineohoe i aahed o he pinde in he P oienaion hen a hekineohoe beome aahed in he P oienaion he inhibiion of opo

iomeae aiviy woud be aboihed and deaenaion of ie homa-id woud aompany he aniion fom meaphae o anaphae

NPHSE n ou mode anaphae i iniiaed by he deaenaon of heie homaid whih aow eah homaid o move poewad a aeu of P foe ndie agumen ugge ha he magniude of hepoewad foe ineae duing anaphae Dieni homoome ae

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CSE SEGEGN N SS ND ESS 0

boken in anaphase if the two kinetohoes ae attahed to opposite poesof the spindle 49). In ontast ing hoosoes that appea to beatenated in etaphase ae not boken whih suggests that the agnitudeof the poewads foes is sale than in anaphase At anaphase these ing

hoosoes appea to pass though eah othe without beakage anobseation that stongly suggests the ation of DNA topoisomeases 50)The fate of dientis suggests that hoosoes inked by atenationwould be boken if els ade the tansition to anaphase withoutdeatenating siste hoatids. The foe puling siste homatids apatoud inease as a esut of an inease in the agnitude of he P foestansitted by te kinetohoetopoe io tubues o the eongationof the spindle ediated by inteations between the poetopole io

tubues o both.Foe on the kinetohoes does not see to be equied fo the initiass o sis kinocho spion nphs. Sisr chroidsepaation ous in els teated with olhiine 5), in aent fagents5) and in onopoa spindes 52). These obseations stongy suggestthat soe linkage between siste hoatids is destoyed as els pogessthough mitosis Ou odel postulates that the sepaation of siste ho-atids is due to the atiation of type II topoisoease at the begin-ning of anaphase

hat ediates the tansition between etaphase and anaphase and theatiation of topoisoease? One attatie andidate is destution ofatuation pooting fato MPF) an atiity that was st identied inXenpu by the ability of ytopas fom homonaly atiated ooytes toindue nonhooneteated ooytes to pass though eiosis I and yiedatue eggs eiewed in 5). Sine then MPF atiity has been found in awide aiety of itoti and eioti els hen injeted into es of ealyXenpu ebyos MPF auses hoosoe ondensation, nuea en-

eope beadown and spindle foation 54) els eain in etaphaseuntil the ee of MPF deays 55). Eithe MPF o soe oponentequied fo its atiation is destoyed at the end of itosis sine newpotein synthesis is equied fo the geneation of MPF atiity in the nextell yle 56) The osillation in MPF ees duing the ell ye pobablyepesents a oponent of the el yle ok that is atiated by fetiliation in enpuseggs 57).

One way in whih MPF oud poong etaphase is by ating eithediety o indietly to inhibit topoisoease II atiity Pehaps attah-

ent of all he inetohoes to the spindle in the P oientation atiates asyste that destoys MPF atiity

F The aailability of loned epliatos (teloees 59 60) and entoees 6 62) in yeast has opened up a new

Annu.Re


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302 MUY & SZSK

appoah to homosome stutue an behaio the onstution in itoof moeues that ontain a the eements known to be equie fo nomal

homosome funtion. hese atiial homosomes ae then intoueino yeast els to examine thei segegation in mitosis an meiosis (3, 4.he st linea atiial homosomes wee 1015 kiobases (kb in

ength (3, 4. Supisingy, they wee pesent in many opies pe e, anwee muh ess mitotialy stabe than iula entomei plasmis, whihwee pesent at one opy pe e an wee ost about one in eey hunemitoses (1, 2, 4. he behaio of these inea pasmis was onsistenwith anom segegation at mitoss, athe than the oee segegation ofsiste moeues haateisti of iula entomei pasmis o natual

homosomes he efet in segegation is not ue to mutations in theentomee (4, intefeene between the entomei an teomeihomatin stutues (4, o the faiue of the entomees to attah to themitoti spine. he last point was emonstate by onstuting ineapasmis that ontaine two entomees. Like ienti iua pasmis(, these inea moleules ae stutuay unstabe, an they yieeaangements that elete one ofhe two entomees (Muay & Sostak,unpuse. u eetons ae te esut meana eaage at

ous ae the two kinetohoes of a ienti moeue hae attahe toopposite spine poes (49 Sine one of the entomees was ony 2.5 kbfom the en of the inea ienti plasmi, it seems unikey that thee is aminimum teomeeentomee sepaation that must be exeee befoendl aachmn can cc

he shot inea entomei pasmis ae ess than 5% of the ength ofthe smaest yeast homosome homosome is 300 kb ( Lageatiia homosomes (55 kb show eiene of nomal siste homati

segegation thei opy numbe is one, an thei mitoti loss fequeny isabout 10 (4. A futhe inease in length to 1 04 kb ineases the stabiityby a fato of e, but suh moeues ae stil at least one hunefo essstabe than natua homosomes (Muay & Sostak, unpubishe. he55 kb atiia homosomes pai an segegate fom eah othe nomalyin meiosis, although again the eity of segegation is substantially essthan that of natual homosomes (4.

DEIVIVES F NUL CHMSS he abiity to eate iete

ateations in the stutue of the yeast genome (7, has mae it possibleto test the eet of aious stutual hanges on homosome behaio.

he st type of ateation is eate by integating a inea teomee-beaing pasmi into a yeast homosome (Figue 4 (Muay & Sostak,unpublishe. Plasmi integation is iete by eaage with a estition

enyme that uts in a egion of he plasmi that is homologous to the

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A EAKAGE

0! Itegi1Relui

!AcetC

B) SHORTENING

Transfomao

RecobaoFgur Manipulation of chromosome srucure. (a) A romosome can be broken and a new eomere generated by e iree inrdcon of apasmid arrying an invered repea of eomeric sequens Dark bas repesen homoogous seences on e pasmid and hromoome; arowseomei DN seqens (b) A romosome arm can be shoren by ransformaion wih a inea DNA fraen hose ends are moogos oisan sies on he homosome Open bars epresen fraen sequences a are no omoogous o he cromosome

(

i

Iz

C

!C

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04 M Y & SZSK

dsird sit of itgrtio. Th chromosom is bro ito two frgmts tth st of itgrtio. Th frgmt tht crris th ctrmr is

mitoticy stb d is rtid whi th ctric frgmt is ridy ost.Th xrimts r coductd i dioid stris so tht th oss of chromosom frgmt is ot th

To tst th ct of gth o chromsom bhvior w itroducdtomrs t vrious oits o yst chromosom Ctromric

chromosom frgmts of 42 d 0 kb r ost mitoticy t frqucis of10 d 1 0 rsctivy, i comriso to chromosom (0 kb (66}

which is ost t frqucy of 1 05. Simir rsuts hv b obtid by

Ty & Surosy (rso commuictio. For th smr frgmtdigr ysis hs show tht 70% of th oss vts occur byodisjuctio [th sistr of th c tht cs th chromosom frt

crris two cois (Murry & Szost uubishd]. This is i mrdcotrst to th bhvior of circur ctromric smids i which oy2% of th vts r du to odisjuctio (69 70 Th rmiig vtshv oy o coy i th sistr c tht ihritd th smid, whichsuggsts tht th smid fid to rict i th c cyc rcdig ossor tht o coy of th smid ws dstroyd r rictio. crigth siz of circur ctromric smids from to 100 b cuss tfoddcrs i mitotic oss frqucy from 1 0 to 10 (69 Uik irctromric smids th coy umbr of th circur ctromrcmocus is o, irrsctiv of thir siz. A circur vrsio of chromosom o frqy of10-3 69, 71, d my o h ts r d tosistr chromtid xchg tht crts dictric dimric chromosomSic sistr chromtid xchg dos ot ct th structur of irchromosoms, this my hv b o of th sctiv forcs fvorig th

voutio of ir chromosoms s gom siz icrsd durigvoutio

A umbr of rsuts suggst tht th gth f DNA mocus is o ofth mi dtrmits of mitotic stbiity:

1 . Th mioic oss frqucis of cromosom frgms d rticichroosoms of simir sizs r withi fctor of of ch othr dsitth fct tht most of th DNA o th rtici chromosoms is rocryotic

(Murry & Szostk uubishd.2. Frts of chromosom of dcrsig siz show grduydcrsig stbiity which dmostrts tht thr is o sig mt(rt from th ctromr whos rsc is rquird for high v ofstbiity (R. Surosy & B.K. T, rso commuictio Murry &Szostk uubishd

Th yst mutt hl icrss th sotous frqucy of

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OO SEGEGnO OSS D OSS

chromosom loss (7 this mutt th frqucy of chromosom lossdcrss with icrsig chromosom siz (7

4 yst zygots whr uclr fusio is rvtd by th kmuttio chromosoms r trsfrrd btw th two ucli. Th

frqucy of chromosom trsfr dcrss with icrsig chromosomsiz (7

Th scod ty of chromosom miultio is th crtio of lrgitrl dltios by trsformig with DA frgmt o of whos dsis homologous to sit o th chromosom d th othr d of which ishomologous to som distt chromosoml sit (Figur 4 R. urosky &B.K. Ty rsol commuictio Murry & zostk ublishd A

rcombitio vt t both ds of th frgmt rlcs ll thchromosoml iformtio tht lis btw th two sits of homology withthe sequenes of the fragment his tehniue has been used to podue an

lmost full lgth vrsio of chromosom whos ctromr is oly kb from th rst tlomr. Th tloctric chromosom i t tims lssstbl th th orml mtctric chromosom (Murry & zostkuublishd. Th tloctric chromosom is still o thousd tims morstbl th th 115 kb lir ctromric lsmids; this dmostrtstht th rimry cus of th lsmids' istbility is ot th closss of

thir ctromrs to thir tlomrs.Th rsults of studis o both rticil chromosoms d drivtivs of

url yst chromosoms suggst tht th fudmtl dtrmit ofmitotic stbility is th lgth of lir DA molculs This is cosisttwith th id tht cttio is sstil rrquisit for rorchromosom sgrgtio f th roducts of DA rlictio r cttdcirculr molculs thy r toologiclly likd to ch othr. cotrstshort lir molculs r ot likd: Th itrtwiigs c b rsolvd

simly by twistig o molcul bout th othr which xlis why thybut ot circulr ctromric lsmids sgrgt rdomly t mitosis. Toccout for th ordrd sgrgtio of log rticil chromosoms wsuggst tht thy c bcom likd o ch othr durig rlictio. uchikg rquirs th xistc of toologicl brrirs tht costri thcttio grtd by rlictio Ths brrirs could b ithrttchmt of th chromosom to th uclr mtri or th formtio ofchromosoml loos by DAbidig rotis which would ocovltlycrosslik th DA uch domis of surcoilig hv b s i

ucryotic chromosoms d it is itriguig tht thy r bout kb log(4 th siz of th smllst rticil chromosom tht shows ordrdsgrgtio t mitosis

Th lgth of DA molculs my ls ct thir bility to bcom



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06 MUY & SZOSK

cd du o h collso of rlco fors. O crculr molcu sgl org of rlco sucs o gv co bcus wo forsrsg from hs org wll m o h oos sd of h crc. For r mocu h fors w m oly f h smd s s

wo rco rgs d boh orgs r bfor hr hs b rcd by h for mg from h ohr org. Th chc of fulgh scod crro crss s h dsc bw h orgs crss. rcur lr molculs wh sgl rlco orgshoud b cb of grg rco d should show vry hgh frqu

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07

Ahogh irig rs o rcd rcombiio d rcombiiois dissbl i cri orgisms [ml Drosophila rviwd i fmBombyx], mbr o is o vidc sggs h rcombiio isormy ivovd i srig homolog sgrgio Rcombiio

rsls i h formio of chism h c s ois of lig bwhomologs As rsl h iochors of ir of homologs r hsicllid b c b widl srd lss h chism i cos o hcromr I boh Droophl (8 d s Lmbi & S odr,rsol commicio mioic rcombiio r h cromr issrssd I s movig h cromr of chromosom III from isorml ocio r h LEU2 g o h H4 g icrss h

frc of rcombiio r LEU2 d dcrss i r H4; his

dmosrs h h cromr isf is rsosibl for srssigrcombiio (E. mbi S. Ror rso commicioO irrio of his srssio of rcombiio is h i srs

h h iochors r o oo ighl lid d ms i sy or hwo iochors of ir of homologs o cr microbs fromoosi os of h mioic sidl. To chiv sbl bior oriio ch iochor ms microbl wih olriyoosi h o o which is sisr is chd A xibl iriochorig will llow h chd iochor o sc lrgr cr

volm h if i wr rigidl chd o is sisr.hil i sms cr h chism i homologos chromosoms h

hv combid h rof h ig is ow I mi hr is xc corrio bw h mbr of chism d h mbr ofrcirocl xchgs rviwd i 82 This sggss h rcombiiovs h r rsolvd s crossovrs gv rs o chism whil hosh r rsolvd s g covrsios do o This obsrvio imis hh sisr chromids of homologs r chd o ch ohr log hir

lgh Ths crossovr will v h rcombi chromids cdo hir sisrs cromr roxim o h oi of xchg, d o hirosisrs cromr disl Figr . G covrsio which mrlyrlcs h iformio o o chromid wih shor srch from isosisr wold o ld o his lig

As h homologs sr h chism mov owrd h omrs Thlig of osisr chromids b cio xlis his homoTooisomrsmdid rmovl of irwiig h chism owsh homologs o mov r dr h ic of h owrds forc

xrd h iochors Howvr rmovl of irwiigs sisohr h h chism fils o ld o srio bcs h homoogsrmi lid h chism Ths h chism mov rogrssivlyowrds h ds of h chromosoms s if ooisomrs c frilly h chsm



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308 MY & SZOSK

YES MEOC MNS The mutaton p13 blocks the st meiotcdvson, and dloid soes ae oduced 83. These soes show nomaleves of ecombnaton, whch demonstates that ths mutant has theexected henotye fo a faue of the "exta SPB dulcation nvoked inthe mode fo the meiotc cel cyce descibed evousy The dulcaton ofSPBs n the mtotic ce cyce is deendent on the CC3 gene oduct To

eain the behavo of thesp13mutant we ostuate that

1. The CC31 gene oduct can nduce both SPB and kinetochoeducaton while the SO 13 gene oduct can nduce SPB but ntknetochoe ducation.

2. The noma mitotic cel cyce incooates a feedback mechanism thatevents the SPBs fom beng dulcated twc between ound of chomo-some segegation

. The acivity o the SO 13 and CC31 gene oducts ae eguated in amutuay eusive fashion CDC3 s active in mtotcay gown ces andin meosis whie S013s active ony dung meioss I.

4 Dung meioss the contnued actvty of the SO13 gene oductis deendent on maintenance of the conditions equied to inducesouation.

The eesson of the SO 13 gene oduct s known to be induced by

soulaton medum (R. . sosito, esona communcationAccoding to ou scenaio, cells statng the ce cyce in soulaton

meda wil have hgh leves ofS013 gene oduct and w duicate theiSPBs but not thei knetochoes whch wil ead to meiosis The

cometion of meiosis I w aow the CDC31function to become actve,knetochoe and SPB ducaton w occu, and meosis I which s

mechanisticaly identica to mitosis wil folow. f cels ae etuned togowth condtions befoe meosis I occus the CC31functon w become

active the knetochoes wi be dulcated and meosis w be eaced bya mitotic dvision. The dioid oducts of this dvson wl have sueedmeiotic eves of ecombination. Such ces ae obseved when souatingcultues ae etuned to vegetative gowth 2 Thus ou model exainswhy ces cannot ente the meiotic ce cyce ae SPB ducation and howthey can escae fom the meiotc ce cyce befoe meosis

The eects of the cdc31 and dc1 mutants on meioss have beenexamned When stans mutant fo ethe gene ae ut though meiosis atthe nonemissve temeatue two diloid soes ae ecoveed, and

genetic analyss indcates that homoogous chomosomes have seaatedat meoss but that meosis and the segegaton of sste chomatds has

faied to occu 30, J Thomas & D. Botstein, esonal communcation.Ths behavio stongy suggests that meosis s functionay equvalent to

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CHOMOSOM SGGATION N MTOSS AND MOSS 309

mtoss and s consstnt wth th da that ths functons a ud fokntocho dupcaton, and that kntocho duplcaton n moss dosnot occu unt aft th compton of moss I

S03 (wdtyp) clls hat contan th combnaon dcnt muta-

tons p 5, o 52 poduc nabl spos (vwd n 9 15)Ectn mcoscopc studs show that th 5 mutant lacks syn-aptonmal complx wh p posssss t (B Bys R E. Espostopsona communcaton) whch dmonstats that th fomaton of S snot sucnt to nsu homoog dsjuncton n yast. In th cas ofpI asma numb of vabl spos can b covd and ths hav notundgon motc combnaton (84) Th p p13 doub mutantpoducs nomal vls of vab dpod spos that hav not undgon

combnaton (84) Th psnc ofpI ncass spo vablty np 3dpods whch suggsts that motc combnaton vnts can ntfth chomosom sggaton n th scond (mtossk) dvson ().haps th lnkag of homologs ntfs wth th coct ontaton onh maphas pa o wth h sggaon of ss chomads atanaphas. Th p 3 5 doubl mutant aso poducs vab butun combnd spos ( 5) Ths suggsts that both RAD and SI gnpoducts a qud to ntat mtoc combnaton and that th fauto do so lads to th fau of homoog dsjuncton at moss I and thus to

nvab spos

n p a numb of motc mutantsthat poduc nvab and anupod gamts hav bn shown to bdcnt fo motc combnaton vwd n . In fmal Dpthos homoogs (o unpad chomosoms) that hav not combndnt a scond pang systm th dstbutv pang pool n whch chomo-soms dson fom ach oth on th bass of sz (vwd n 85) Th ds-tutv pang mchansm s appanty sucnt to da wth aout ouunpad chomosoms gh numbs ad to ts bakdown and vyhgh funcs of nondsuncton (3. In ma Dp whch lackhomoogous pang combnaton and dstbutv pang th homo-logs appa to pa va spcc pang sts n th htochomatn(vwd n 3

h Dpmotc mutantplads to chomosom loss n mossand chomosom loss n th aly dvsons of th ygot (86). p acts

moss ony n mas and th chomosoms that a lost fom th ygot

nucus and ts pogny a always th panal chomosoms (86). In onstock th X chomosom was much mo sstant to nducd oss ncosss ths sstanc to oss maps to th cntomc htochomatn ofth X (86) W suggst that a wdtyp poduct of th p gn acts to



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30 MY & ZOTK

surss kintoor assm for miosis I in mals T mutant allsaow kintoor uliation to our for miosis ; trfor, tnw assml kintoors ar fti aus som sustan oronition rquir for kintoor assml is asnt at tis sta of

miosis n miosis tis ou a to romosoms wit on norma anon fti kintoor an a i frun of nonisuntion.

Simultanous oss of ot romosoms X an 4 is mor frqunt tant from t rout of t frqunis of tional sration foritr on alon 86 T aior of t two romosoms is orrat nt as of ar mitoti oss ot romosoms ar amost awas ost fromt sam o in iision 86. Tis aior is rminisnt of t astndcmutant in tat t romosoms tat nonisoin al arri at t sam ol

in mitosis an miosis. owr in t ndcmutant itr a or non of tromosoms in a nonisoin an srat to a sin o, wi int mutant som of t romosoms isoin an otrs sratnorma T orrat aior of romosoms ou ain ift fti kintoors assm in t mioti l awas fat sam o in susqunt iisions. On ay in wi su oorintationan ai is to atta a t kintoors of t sam a to t samo f our intrrtation is orrt t aior of ts mutants arustat intoors of t sam a ar a atta to on o in miosis

as w as in mitosis.Gostin 87 as sown tat in wilt mals at romtaas of

miosis t kintoor of a omoo is a sin misriastrutur By at mtaas t kintoor as mtamoros into aou strutur tat onsists of two iss lin si si 87 Wits two iss ou funtional innnt t a t samorintation an tus wi atta to mirotuuls of t sam olarity sotr wi no tnn of sistr romatis to sarat. W rit attron miroso of miosis inpmals wou islos t ourrnof kintoor uiation for miosis

Ar tr miosissi ns tat rnt sistr kintoor saration for mtaas of miosis ? fts in su ns wou -t to i ris to roios saration of sistr romatis, iwou sor as nonisuntion at itr miosis or miosis nin on wn saration ourr. Two mutants wit tis rorta n sri in Dh mS22 88 an 8. Ty aruniqu amon mioti mutants in tat ty at in ot mas an fmas,

an ot a n tooa osr to aus t roioussaration of sistr romatis in ma miosis 0 T fration ofnonisuntion nts tat ours at miosis is % an 30% for m-S22an , rstil For mS22 t frqun of amts tat ar



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CHR GRGA I D 3 1 1

n o copies of a given chromosome is much greater than that of gametes thacary two copies of the same chromosome, which suggests that somechromosomes are not incuded in the nuclei that form after the completionof meiosis.

R Sister chomatids separate from each other nmitosis and meoss II but not in meiosis I. Does this reect dierences inthe structure of the meosis I and I spindes or is t a reection of theorganization of the chromosomes themselves? One test between theseaenatives is vided b an eeant exment o Ncklas (16) Heinduced primary and secondary spematoctes to fuse wth each other toproduce a sine ce that had both meiosis and meisis sindes Hethen transferred ether a bivalent from the meisis to the meoss II spinde

or a pair of sister chromatids from the meiosis to the meiosis sindle. nbth cases the transferred entities behaved as the d have dne n hesindle the rinal beoned t : The bivalent searated nt thmoogs whle the sster chromatids from meosis II searated from eachother (6. his resut is entire consistent with our argment thatinetochore duication does not occur until the interva between meiosis Iand . omologs transferred from the meiosis sinde would have a sngeinetochore, while those transferred from the meiosis II to meiosis I spindewould have dulicated inetochores alowing the sister chromatids to

disjon from each other.

EVOLUTION OF MIOSIS

Our model rovides a unied view ofhe mitotic and meiotc ce ccles andsuggests how meiosis evoved from mitosis. In the prmitive forms ofmeiosis, disjunction was probably ensured by synaptonemal compefomation without recombination Athough in mitoss the chromosomeswere ebly attached to each other b intercatenation the rgd association

of homo ogs in meiosis medated b the SC probably led to a high frequencyof nondisnction. The introdctin of recombinatin t create a eiblelinage between homoogs woud have increased the deity of chromo-some segregation Ths the nitial seection for the evoution of meioticrecombination ma have been for a decrease in nondisjnction rather thanan increase in the otentia for enetic diversity in the succeedngeneratin.

KWDG

Because of space imitations we were unabe to quote every reference to agven tic To thse whose names are not mentioned we oer ur

Annu.Rev.Cell.Biol.1985.1:289315

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31 MURAY & SZOSTAK

aooges We ae eemey gaeu o a hose who ovie nomaonbeoe ubicaion We han D. Dawson, P Szaue, n B. Abes ohei heu omens on he anui an aus o hooahy

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