Probing Evolutionary Repeatability Neutral and Double Changes and the Predictability of Evolutionary Adaptation 英文参考文献.docVIP

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Probing Evolutionary Repeatability Neutral and Double Changes and the Predictability of Evolutionary Adaptation 英文参考文献.doc

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Probing Evolutionary Repeatability Neutral and Double Changes and the Predictability of Evolutionary Adaptation 英文参考文献

ProbingEvolutionaryRepeatability:NeutralandDouble ChangesandthePredictabilityofEvolutionary Adaptation ScottWilliamRoy* NationalCenterforBiotechnologyInformation,NationalLibraryofMedicine,NationalInstitutesofHealth,Bethesda,Maryland,UnitedStatesofAmerica Abstract Background:Thequestionofhoworganismsadaptisamongthemostfundamentalinevolutionarybiology.Tworecent studiesinvestigatedtheevolutionofEscherichiacoliinresponsetochallengewiththeantibioticcefotaxime.Studyingfive mutations in the b-lactamase gene that together confer significant antibiotic resistance, the authors showed a complex fitnesslandscapethatgreatlyconstrainedtheidentityandorderofintermediatesleadingfromtheinitialwildtypegenotype to the final resistant genotype. Out of 18 billion possible orders of single mutations leading from non-resistant to fully- resistantform,theyfoundthatonly27(1.561027%)pathwayswerecharacterizedbyconsistentlyincreasingresistance,thus onlyatinyfractionofpossiblepathsareaccessiblebypositiveselection.Ifurtherexplorethesedatainseveralways. Principal Findings: Allowing neutral changes (those that do not affect resistance) increases the number of accessible pathways considerably, from 27 to 629. Allowing multiple simultaneous mutations also greatly increases the number of accessible pathways. Allowing a single case of double mutation to occur along a pathway increases the number of pathwaysfrom27to259,andallowingarbitrarilymanypairsofsimultaneouschangesincreasesthenumberofpossible pathwaysbymorethan100fold,to4800.Iintroducethemetric‘repeatability,’theprobabilitythattworandomtrialswill proceedviatheexactsamepathway.Ingeneral,Ifindthatwhilethetotalnumberofaccessiblepathwaysisdramatically affectedbyallowingneutralordoublemutations,theoverallevolutionaryrepeatabilityisgenerallymuchlessaffected. Conclusions:Theseresultsprobetheconceivablepathwaysavailabletoevolution.Evenwhenmanyoftheassumptionsof the analysis of Weinreich et al. (2006) are relaxed, I find that evolution to more highly cef