Epigenetically-Inherited Centromere and Neocentromere DNA Replicates Earliest in S-Phase 英文参考文献.docVIP

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Epigenetically-Inherited Centromere and Neocentromere DNA Replicates Earliest in S-Phase 英文参考文献.doc

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Epigenetically-Inherited Centromere and Neocentromere DNA Replicates Earliest in S-Phase 英文参考文献

Epigenetically-InheritedCentromereandNeocentromere DNAReplicatesEarliestinS-Phase AmnonKoren1,2¤,Hung-JiTsai1,ItayTirosh2,LauraS.Burrack1,NaamaBarkai2,JudithBerman1* 1Department of Genetics, Cell Biology, and Development, University of Minnesota, Minneapolis, Minnesota, United States of America, 2Department of Molecular Genetics,TheWeizmannInstituteofScience,Rehovot,Israel Abstract Eukaryotic centromeres are maintained at specific chromosomal sites over many generations. In the budding yeast Saccharomyces cerevisiae, centromeres are genetic elements defined by a DNA sequence that is both necessary and sufficientforfunction;whereas,inmostothereukaryotes,centromeresaremaintainedbypoorlycharacterizedepigenetic mechanisms in which DNA has a less definitive role. Here we use the pathogenic yeast Candida albicans as a model organismtostudytheDNAreplicationpropertiesofcentromericDNA.Bydeterminingthegenome-widereplicationtiming programoftheC.albicansgenome,wediscoveredthateachcentromereisassociatedwithareplicationoriginthatisthe firsttofireonitsrespectivechromosome.Importantly,epigeneticformationofnewectopiccentromeres(neocentromeres) was accompanied by shifts in replication timing, such that a neocentromere became the first to replicate and became associatedwithoriginrecognitioncomplex(ORC)components.Furthermore,changingthelevelofthecentromere-specific histoneH3isoformledtoaconcomitantchangeinlevelsofORCassociationwithcentromereregions,furthersupporting the idea that centromere proteins determine origin activity. Finally, analysis of centromere-associated DNA revealed a replication-dependentsequencepatterncharacteristicofconstitutivelyactivereplicationorigins.Thisstrand-biasedpattern isconserved,togetherwithcentromereposition,amongrelatedstrainsandspecies,inamannerindependentofprimary DNAsequence.Thus,inheritanceofcentromerepositioniscorrelatedwithaconstitutivelyactiveoriginofreplicationthat firesatadistinctearlytime.WesuggestamodelinwhichthedistincttimingofDNAreplicatio