Equation-Free Analysis of Two-Component System Signalling Model Reveals the Emergence of Co-Existing Phenotypes in the Absence of Multistationarity 英文参考文献.docVIP

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Equation-Free Analysis of Two-Component System Signalling Model Reveals the Emergence of Co-Existing Phenotypes in the Absence of Multistationarity 英文参考文献.doc

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Equation-Free Analysis of Two-Component System Signalling Model Reveals the Emergence of Co-Existing Phenotypes in the Absence of Multistationarity 英文参考文献

Equation-FreeAnalysisofTwo-ComponentSystem SignallingModelRevealstheEmergenceofCo-Existing PhenotypesintheAbsenceofMultistationarity RebeccaB.Hoyle1*,DanieleAvitabile2,AndrzejM.Kierzek3 1Department of Mathematics, University of Surrey, Guildford, United Kingdom, 2School of Mathematical Sciences, University of Nottingham, Nottingham, United Kingdom,3DivisionofMicrobialSciences,UniversityofSurrey,Guildford,UnitedKingdom Abstract Phenotypicdifferencesofgeneticallyidenticalcellsunderthesameenvironmentalconditionshavebeenattributedtothe inherent stochasticity of biochemical processes. Various mechanisms have been suggested, including the existence of alternative steady states in regulatory networks that are reached by means of stochastic fluctuations, long transient excursionsfromastablestatetoanunstableexcitedstate,andtheswitchingonandoffofareactionnetworkaccordingto theavailabilityofaconstituentchemicalspecies.Hereweanalyseadetailedstochastickineticmodeloftwo-component systemsignallinginbacteria,andshowthatalternativephenotypesemergeintheabsenceofthesefeatures.Weperforma bifurcationanalysisofdeterministicreactionrateequationsderivedfromthemodel,andfindthattheycannotreproduce thewholerangeofqualitativeresponsestoexternalsignalsdemonstratedbydirectstochasticsimulations.Inparticular,the mixedmode,wherestochasticswitchingandagradedresponseareseensimultaneously,isabsent.However,probabilistic and equation-free analyses of the stochastic model that calculate stationary states for the mean of an ensemble of stochastictrajectoriesrevealthatslowtranscriptionofeitherresponseregulatororhistidinekinaseleadstothecoexistence ofanapproximatebasalsolutionandagradedresponsethatcombinetoproducethemixedmode,thusestablishingits essentialstochasticnature.Thesametechniquesalsoshowthatstochasticityresultsintheobservationofanall-or-none bistableresponseoveramuchwiderrangeofexternalsignalsthanwouldbeexpectedondeterministicgrounds.Thuswe demonstratetheapplicationofnumericalequation-freemethodsto