Positive Selection Drives the Evolution of rhino, a Member of the Heterochromatin Protein 1 Family in Drosophila 英文参考文献.docVIP

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Positive Selection Drives the Evolution of rhino, a Member of the Heterochromatin Protein 1 Family in Drosophila 英文参考文献.doc

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Positive Selection Drives the Evolution of rhino, a Member of the Heterochromatin Protein 1 Family in Drosophila 英文参考文献

PositiveSelectionDrivestheEvolution ofrhino,aMemberoftheHeterochromatin Protein1FamilyinDrosophila Danielle Vermaak1,Steven Henikoff1,2,Harmit S.Malik1* 1FredHutchinsonCancerResearchCenter,Seattle,Washington,UnitedStatesofAmerica,2HowardHughesMedicalInstitute,BasicSciencesDivision,FredHutchinson CancerResearchCenter,Seattle,Washington,UnitedStatesofAmerica Heterochromatin comprises a significant component of many eukaryotic genomes. In comparison to euchromatin, heterochromatinis genepoor, transposon rich,and late replicating.Itserves many important biological roles, from genesilencingtoaccuratechromosomesegregation,yetlittleisknownabouttheevolutionaryconstraintsthatshape heterochromatin. A complementary approach to the traditional one of directly studying heterochromatic DNA sequence is to study the evolution of proteins that bind and define heterochromatin. One of the best markers for heterochromatinistheheterochromatinprotein1(HP1),whichisanessential,nonhistonechromosomalprotein.Here we investigate the molecular evolution of five HP1 paralogs present in Drosophila melanogaster. Three of these paralogshaveubiquitousexpressionpatternsinadultDrosophilatissues,whereasHP1D/rhinoandHP1Eareexpressed predominantly in ovaries and testes respectively. The HP1 paralogs also have distinct localization preferences in Drosophilacells.Thus,RhinolocalizestotheheterochromaticcompartmentinDrosophilatissueculturecells,butina patterndistinctfromHP1Aandlysine-9dimethylatedH3.Usingmolecularevolutionandpopulationgeneticanalyses, wefindthatrhinohasbeensubjecttopositiveselectioninallthreedomainsoftheprotein:theN-terminalchromo domain, the C-terminal chromo-shadow domain, and the hinge region that connects these two modules. Maximum likelihood analysis of rhino sequences from20 species of Drosophila reveals that asmall numberof residues ofthe chromo and shadow domains have been subject to repeated positive selection. The rapid and positive selection of rhinoishighlyunusualforageneenco