Expression of Arf Tumor Suppressor in Spermatogonia Facilitates Meiotic Progression in Male Germ Cells 英文参考文献.docVIP

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Expression of Arf Tumor Suppressor in Spermatogonia Facilitates Meiotic Progression in Male Germ Cells 英文参考文献.doc

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Expression of Arf Tumor Suppressor in Spermatogonia Facilitates Meiotic Progression in Male Germ Cells 英文参考文献

ExpressionofArfTumorSuppressorinSpermatogonia FacilitatesMeioticProgressioninMaleGermCells MichelleL.Churchman1,2,IgnasiRoig3,4,MariaJasin5,ScottKeeney1,3,CharlesJ.Sherr1,2 * 1Howard Hughes Medical Institute, St. Jude Children’s Research Hospital, Memphis, Tennessee, United States of America, 2Department of Genetics and Tumor Cell Biology,St.JudeChildren’sResearchHospital,Memphis,Tennessee,UnitedStatesofAmerica,3MolecularBiologyProgram,MemorialSloan-KetteringCancerCenter,New York,NewYork,UnitedStatesofAmerica,4CytologyandHistologyUnit,DepartmentofCellBiology,Physiology,andImmunology,UniversitatAutonomadeBarcelona, Barcelona,Spain,5DevelopmentalBiologyProgram,MemorialSloan-KetteringCancerCenter,NewYork,NewYork,UnitedStatesofAmerica Abstract The mammalian Cdkn2a (Ink4a-Arf) locus encodes two tumor suppressor proteins (p16Ink4a and p19Arf) that respectively enforcetheanti-proliferativefunctionsoftheretinoblastomaprotein(Rb)andthep53transcriptionfactorinresponseto oncogenicstress.Althoughp19Arfisnotnormallydetectedintissuesofyoungadultmice,anotableexceptionoccursinthe malegermline,whereArfisexpressedinspermatogonia,butnotinmeioticspermatocytesarisingfromthem.Unlikeother contextsinwhichtheinductionofArfpotentlyinhibitscellproliferation,expressionofp19Arf inspermatogoniadoesnot interferewithmitoticcelldivision.Instead,inactivationofArftriggersgermcell–autonomous,p53-dependentapoptosisof primary spermatocytes in late meiotic prophase, resulting in reduced sperm production. Arf deficiency also causes premature, elevated, and persistent accumulation of the phosphorylated histone variant H2AX, reduces numbers of chromosome-associated complexes of Rad51 and Dmc1 recombinases during meioticprophase, andyields incompletely synapsed autosomes during pachynema. Inactivation of Ink4a increases the fraction of spermatogonia in S-phase and restoresspermnumbersinInk4a-Arfdoublydeficientmicebutdoesnotabrogatec-H2AXaccumulationinspermatocytes orp53-dependentapoptosisresultingfr