Nitric Oxide Regulates Input Specificity of Long-Term Depression and Context Dependence of Cerebellar Learning 英文参考文献.docVIP

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Nitric Oxide Regulates Input Specificity of Long-Term Depression and Context Dependence of Cerebellar Learning 英文参考文献.doc

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Nitric Oxide Regulates Input Specificity of Long-Term Depression and Context Dependence of Cerebellar Learning 英文参考文献

NitricOxideRegulatesInputSpecificity ofLong-TermDepressionandContext DependenceofCerebellarLearning Hideaki Ogasawara1,2*,Tomokazu Doi1,3¤,Kenji Doya1,4,Mitsuo Kawato1 1ATRComputationalNeuroscienceLaboratories,Seika,Kyoto,Japan,2NationalInstituteofInformationandCommunicationsTechnology,Seika,Kyoto,Japan,3Graduate School of Information Science, Nara Institute of Science and Technology, Takayama, Ikoma, Nara, Japan, 4 Initial Research Project, Okinawa Institute of Science and Technology,Uruma,Okinawa,Japan Recent studies have shown that multiple internal models are acquired in the cerebellum and that these can be switched under a given context of behavior. It has been proposed that long-term depression (LTD) of parallel fiber (PF)–Purkinjecell(PC)synapsesformsthecellularbasisofcerebellarlearning,andthatthepresynapticallysynthesized messengernitricoxide(NO)isacrucial‘‘gatekeeper’’forLTD.BecauseNOdiffusesfreelytoneighboringsynapses,this volume learning is not input-specific and brings into question the biological significance of LTD as the basic mechanismforefficientsupervisedlearning.TobettercharacterizetheroleofNOincerebellarlearning,wesimulated the sequence of electrophysiological and biochemical events in PF–PC LTD by combining established simulation modelsoftheelectrophysiology,calciumdynamics,andsignalingpathwaysofthePC.Theresultsdemonstratethat thelocalNOconcentrationiscriticalforinductionofLTDandforitsinputspecificity.Pre-andpostsynapticcoincident firingisnotsufficientforaPF–PCsynapsetoundergoLTD,andLTDisinducedonlywhenasufficientamountofNOis provided by activation of the surrounding PFs. On the other hand, above-adequate levels of activity in nearby PFs causeaccumulationofNO,whichalsoallowsLTDinneighboringsynapsesthatwerenotdirectlystimulated,ruining inputspecificity.ThesefindingsleadustoproposethehypothesisthatNOrepresentstherelevanceofagivencontext andenablescontext-dependentselectionofinternalmodelstobeupdated.WealsopredictsparsePFactivityinvivo because,o