Evolutionary Genomics of a Temperate Bacteriophage in an Obligate Intracellular Bacteria (Wolbachia) 英文参考文献.docVIP

EvolutionaryGenomicsofaTemperateBacteriophagein anObligateIntracellularBacteria(Wolbachia) BethanyN.Kent,LisaJ.Funkhouser,ShefaliSetia,SethR.Bordenstein* DepartmentofBiologicalSciences,VanderbiltUniversity,Nashville,Tennessee,UnitedStatesofAmerica Abstract Genome evolution of bacteria is usually influenced by ecology, such that bacteria with a free-living stage have large genomesandhighratesofhorizontalgenetransfer,whileobligateintracellularbacteriahavesmallgenomeswithtypically low amounts of gene exchange. However, recent studies indicate that obligate intracellular species that host-switch frequentlyharboragentsofhorizontaltransfersuchasmobileelements.Forexample,thetemperatedouble-strandedDNA bacteriophageWOinWolbachiapersistentlytransfersbetweenbacterialcoinfectionsinthesamehost.Hereweshowthat despitethephage’srampantmobilitybetweencoinfections,theprophage’sgenomedisplaysfeaturesofconstraintrelated to its intracellular niche. First, there is always at least one intact prophage WO and usually several degenerate, independently-acquiredWOprophagesineachWolbachiagenome.Second,whiletheprophagegenomesaremodularin compositionwithgenesofsimilarfunctiongroupingtogether,themodulesaregenerallynotinterchangeablewithother unrelated phages and thus do not evolve by the Modular Theory. Third, there is an unusual core genome that strictly consistsofheadandbaseplategenes;othergenemodulesarefrequentlydeleted.Fourth,theprophagerecombinasesare diverseandthereisnoconservedintegrationsequence.Finally,themolecularevolutionaryforcesactingonprophageWO arepointmutation,intragenicrecombination,deletion,andpurifyingselection.Takentogether,theseanalysesindicatethat whilelateral transferofphage WOispervasive betweenWolbachia withoccasional newgeneuptake,constraintsofthe intracellularnicheobstructextensivemixturebetweenWOandtheglobalphagepopulation.AlthoughtheModularTheory haslongbeenconsideredtheparadigmoftemperatebacteriophageevolutioninfree-livingbacteria,itappearsirrelevantin phagesofo