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ofthegenodurgthecellcycle(oralesandlosada，2018uhlann，2016)fairetoestab
lishorresolvehesionatilyannerleadstogenoicstabilityandaneuloidysisterchro
atidhesionisdiatedbyhes，argshaedataseachethatnsistsofsc1a，
sc3，rad21，andeitherstag1orstag2huansoaticcells(haarhuisetal，2014
losadaandhirano，2005nasythandhaerg，2009onal，2008etersetal，2008
zhengandyu，2015)hesrgsologicallyentradnatogeneratehysicallkages
betatidsandenablehesionhesregutesotherchroosobasedro
cesses，suchasdnareair，transcrition，andchroosofoldg(rkenschgerandodo，
2013wuandyu，2012)theseotherfunctionsofheslikelyalsovolvetheologicalentra
ntofchroososorossiblytheextrionofdnaloos(barrgtoal，2017
davidsoal，2016haarhuisetal，2017)
hesisloadedontochroosostelohaseandg1bythes24lex(niblau2
huans)(ciosketal，2000gillesieandhirano，2hietal，2004tonketal，
2004osoboundhesisdynaicand
isactivelyreovedfrochroososbythehesreleasgfactroteds5aords5b(chaal，2012kuengetal，2006loezserraetal，
2013ouyangandyu，2017ouyangetal，2013ouyangetal，2016)durgdnarelication
shase，aoolofhesisnvertedtothehesivefor，ddiatessisterchroatidhesion(gerlichetal，2006kuengetal，2006)huan
cells，hesionestablishntreirestheacetytionofsc3bytheacetyltransferaseses1and
es2andsubseentrecruitntofsoror，osos(aloretal，2017houandzou，2005nishiyaaetal，2010ouyangetal，
2016ranketal，2005rolefbenshaharetal，2008rowndetal，2009unaletal，2008
zhangetal，2008a)
thecheckotkaserotesc1andrad53arereired
thebuddgyeast，saharoycescerevisiae，toatacell
viabilitytheresenceofdrugscagdaagetodnaor
arrestofdnarelicationforks1±3itisthoughtthattheyactby
hibitgcellcyclerogression，allogtifordnareairto
takecec1andrad53alsoslo
resonsetodnaalkytion4，althoughthechanisforthis
anditsretiveiortancerotectgcellsfrodnadaage
havenotbeendeteredhereaalkytg
agentthylthanesulhonate(s)rofoundlyreducesthe
rateofdnarelicationforkrogressionho
doesnotreirerad53orc1theaeleratedshase
checkotutants4，therefore，isriarilyanseenceof
aroriateitiationevents5±7arelicationtheresenceofsntrast，
relicationforkscheckotutantslseirreversiblyat
highratesoreover，thecytotoxicityofscheckot
utantsourssecicallyageth，reventgdaageduced
dnarelicationforkcatastroheseestobeariarychanis
bytsreserveviabilitythefaceofdna
toensurethataletesetoftheeukaryoticgenois
reciselydulicateddurgtheliitederiodofshase
everycellcycle，dnarelicationitiatesatanuberof
relicationorigsonchroosos(gilbert，2001belland
dutta，2002)aseachchroosoregionrelicatesa
secificeriodgoforigactivation
tberegutedalthoughgunderstandg
ofrotefactorsvolveditiationandelongationof
relication，thechanissoforigactivationatthechrooso
levelareyettobecrifieddetailth，itis
iortanttodeterelocationsofallrelicationorigson
chroososhoallnubersofrelication
origshavesofarbeenidentifiedostanissother
thanbuddgyeastsaharoycescerevisiae(acaleand
bell，2005)
therocessofitiationofrelicationatdividualrelication
origsisosedoftsgof
relicationorigsg1haseandactivationoftheorigs
shaseg1hase，rerelicativelexes(rercs)are
fordatrelicationorigs(bellanddutta，2ey
andtterill，2003)thisreiresbdgoftheorig
regnitionlex(orc)toarelicationorig，folloichroosoatenance
lex，deendgontheloadgfactors，cdc6cdc18and
cdt1(diffleyetal，1994bellanddutta，2002)althoughre
rcforationisessentialforitiationofrelication，itisnot
itselfsufficientorigactivationshaseisregutedby
onservedrotekases，cycldeendentrote
kase(cdk)andcdc7?dbf4rotekase(dbf4deendent
kase，ddk)thesekasesarereiredforasseblyof
severalotherrotefactors，cdgcdc45andgsonto
rercsthisayleadtoactivationofhelicaseand
origdnaunacheryis
establishedthroughasseblyofraanddnaolyrases
ontothesglestrandeddna(bellanddutta，2002)
althoughrotesvolveditiationofrelicationare
nservedaongeukaryotes，thenucleotideseencesof
relicationorigsareverydiverseaonganiss
(gilbert，2001)，alybecaeofdifferencesdnabdg
roertiesoforcsbuddgyeast，orcregnizesthe
secificseencecalledthearsnsensseence(acs)
ntrast，noclearnsensseencehasbeenfound
origsfissionyeast，schizosaharoycesobe(clyne
andkelly，1995dubeyetal，1996okunoetal，1999)，
althoughatrichseencestotially
bdsarereired(chuangandkelly，1999)reirents
forsecificseencesbeelessclearulticelr
anisssuchastazoans，andorcexhibitslittleseence
secificitydnabdgvitro(vasheeetal，2003
reetal，2004)therefore，itisiortanttodeterethe
locationsoforcbdganddnasynthesisexerintally
genorotesgdnaicroarrayshave
received:24july2006aeted:8january2007ublishedonle:
15fe
uary2007
rresondgauthordeartntofbiology，graduateschoolof
science，osakauniversity，11，achikaneyaacho，toyonaka，osaka
5600043，jaantel:t81668505432fax:t81668505440
eail:
5resentaddress:thescrisresearchstitute，10550northtorrey
esroad，jol，ca92037，a
6resentaddress:deartntofbiologicalcheistryandolecur
haralogy，harvarddicalschool，240longa02115，a
theebojoual(2007)26，1327?1339|a2007euroeanolecurbiologyanization|allrightsreserved0261418907
ebojoual
ebo
joual
the
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