Rapid Birth–Death Evolution Specific to Xenobiotic Cytochrome P450 Genes in Vertebrates 英文参考文献.docVIP

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Rapid Birth–Death Evolution Specific to Xenobiotic Cytochrome P450 Genes in Vertebrates 英文参考文献.doc

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Rapid Birth–Death Evolution Specific to Xenobiotic Cytochrome P450 Genes in Vertebrates 英文参考文献

RapidBirth–DeathEvolutionSpecific toXenobioticCytochromeP450Genes inVertebrates James H.Thomas* DepartmentofGenomeSciences,UniversityofWashington,Seattle,Washington,UnitedStatesofAmerica Genes vary greatly in their long-term phylogenetic stability and there exists no general explanation for these differences.ThecytochromeP450(CYP450)genesuperfamilyiswellsuitedtoinvestigatingthisproblembecauseitis largeandwellstudied,anditincludesbothstableandunstablegenes.CYP450genesencodeoxidaseenzymesthat functioninmetabolismofendogenoussmallmoleculesandindetoxificationofxenobioticcompounds.Bothtypesof enzymeshavebeenintensivelystudied.Myanalysisoftennearlycompletevertebrategenomesindicatesthateach genomecontains50–80CYP450genes,whichareaboutevenlydividedbetweenphylogeneticallystableandunstable genes.Thestablegenesarecharacterizedbyfewornogeneduplicationsorlossesinspeciesrangingfrombonyfishto mammals,whereasunstablegenesarecharacterizedbyfrequentgeneduplicationsandlosses(birth–deathevolution) evenamongcloselyrelatedspecies.AlloftheCYP450genesthatencodeenzymeswithknownendogenoussubstrates are phylogenetically stable. In contrast, most of the unstable genes encode enzymes that function as xenobiotic detoxifiers.NearlyallunstableCYP450genesinthemouseandhumangenomesresideinafewdensegeneclusters, formingunstablegeneislandsthatarosebyrecurrentlocalgeneduplication.Evidenceforpositiveselectioninamino acidsequenceisrestrictedtotheseunstableCYP450genes,andsitesofselectionareassociatedwithsubstrate-binding regionsintheproteinstructure.Theseresultscanbeexplainedbyageneralmodelinwhichphylogeneticallystable genes have core functions in development and physiology, whereas unstable genes have accessory functions associatedwithunstableenvironmentalinteractionssuchastoxinandpathogenexposure.Unstablegeneislandsin vertebrates share some functional properties with bacterial genomic islands, though they arise by local gene duplicationratherthanhorizontalgenetransfer. Citation:ThomasJH(2007)Rapidbirth