A Multi-Step Pathway for the Establishment of Sister Chromatid Cohesion 英文参考文献.docVIP

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A Multi-Step Pathway for the Establishment of Sister Chromatid Cohesion 英文参考文献.doc

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A Multi-Step Pathway for the Establishment of Sister Chromatid Cohesion 英文参考文献

AMulti-StepPathwayfortheEstablishmentof SisterChromatidCohesion Mark Milutinovich1,2,3,Elc?in Unal¨ 1,2,3,Chris Ward1,2,4,Robert V.Skibbens5,Douglas Koshland1,2,3* 1 Howard Hughes Medical Institute, Carnegie Institution, Baltimore, Maryland, United States of America, 2 Department of Embryology, Carnegie Institution, Baltimore, Maryland, United States of America, 3Department ofBiology, Johns Hopkins University, Baltimore, Maryland, United States ofAmerica, 4 Ingenuity Program, Baltimore Polytechnic Institute, Baltimore, Maryland, United States of America, 5 Department of Biological Sciences, Lehigh University, Bethlehem, Pennsylvania, United States of America The cohesion of sister chromatids is mediated by cohesin, a protein complex containing members of the structural maintenance of chromosome (Smc) family. How cohesins tether sister chromatids is not yet understood. Here, we mutate SMC1, the gene encoding a cohesin subunit of budding yeast, by random insertion dominant negative mutagenesistogenerateallelesthatarehighlyinformativeforcohesinassemblyandfunction.Cohesinsmutatedin theHingeorLoop1regionsofSmc1bindchromatinbyamechanismsimilartowild-typecohesin,butfailtoenrichat cohesin-associatedregions(CARs)andpericentricregions.Hence,theHingeandLoop1regionsofSmc1areessential for thespecific chromatin binding of cohesin. This specific binding and asubsequent Ctf7/Eco1-dependent step are both required for the establishment of cohesion. We propose that a cohesin or cohesin oligomer tethers the sister chromatidsthroughtwochromatin-bindingeventsthatareregulatedspatiallybyCARbindingandtemporallybyCtf7 activation,toensurecohesinscrosslinkonlysisterchromatids. Citation:MilutinovichM,U¨nalE,WardC,SkibbensRV,KoshlandD(2007)Amulti-steppathwayfortheestablishmentofsisterchromatidcohesion.PLoSGenet3(1):e12. doi:10.1371/journal.pgen.0030012 that have been observed in preparations of puri?ed Smc complexes[13]. Introduction Proper transmission of eukaryotic chromosomes during ce