Silencing Mediated by the Schizosaccharomyces pombe HIRA Complex Is Dependent upon the Hpc2-Like Protein, Hip4 英文参考文献.docVIP

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Silencing Mediated by the Schizosaccharomyces pombe HIRA Complex Is Dependent upon the Hpc2-Like Protein, Hip4 英文参考文献.doc

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Silencing Mediated by the Schizosaccharomyces pombe HIRA Complex Is Dependent upon the Hpc2-Like Protein, Hip4 英文参考文献

SilencingMediatedbytheSchizosaccharomycespombe HIRAComplexIsDependentupontheHpc2-LikeProtein, Hip4 HollyE.Anderson1,AlexanderKagansky2,JosephineWardle1,JuriRappsilber2,RobinC.Allshire2, SimonK.Whitehall1* 1InstituteforCellandMolecularBiosciences,NewcastleUniversity,Newcastle,UnitedKingdom,2WellcomeTrustCentreforCellBiologyandInstituteofCellBiology, UniversityofEdinburgh,Edinburgh,Scotland,UnitedKingdom Abstract Background:HIRA(orHir)proteinsareconservedhistonechaperonesthatfunctioninmulti-subunitcomplexestomediate replication-independent nucleosome assembly. We have previously demonstrated that the Schizosaccharomyces pombe HIRAproteins,Hip1andSlm9,formacomplexwithaTPRrepeatproteincalledHip3.Herewehaveidentifiedanewsubunit ofthiscomplex. Methodology/Principal Findings: To identify proteins that interact with theHIRA complex, rapid affinity purifications of Slm9wereperformed.MultiplecomponentsofthechaperonincontainingTCP-1complex(CCT)andthe19Ssubunitofthe proteasomereproduciblyco-purifiedwithSlm9,suggestingthatHIRAinteractswiththesecomplexes.Slm9wasalsofound tointeractwithapreviouslyuncharacterisedprotein(SPBC947.08c),thatwecalledHip4.Hip4containsaHRDdomainwhich is a characteristic of the budding yeast and human HIRA/Hir-binding proteins, Hpc2 and UBN1. Co-precipitation experimentsrevealedthatHip4isstablyassociatedwithalloftheothercomponentsoftheHIRAcomplexanddeletionof + hip4 resultedinthecharacteristicphenotypesofcellslackingHIRAfunction,suchastemperaturesensitivity,anelongated cellmorphologyandhypersensitivitytothespindlepoison,thiabendazole.Moreover,lossofHip4functionalleviatedthe heterochromatic silencingof reporter geneslocated inthe mating typelocus andcentromeres andwas associated with increasedlevelsofnon-codingtranscriptsderivedfromcentromericrepeatsequences.Hip4wasalsofoundtoberequired forthedistinctformofsilencingthatcontrolstheexpressionofTf2LTRretrotransposons. Conclusions/Significance: Overall,theseresultsindicatethatHip4isanintegralcomponentoftheHIR