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The C erebellum Learning Movement, Language, and Social Skills Dianne M. Broussard The Cerebellum Learning Movement, Language, and Social Skills The Cerebellum Learning Movement, Language, and Social Skills Dianne M. Broussard Thiseditionfirstpublished2014(cid:2)C 2014byJohnWiley&Sons,Inc. Editorialoffices: 1606GoldenAspenDrive,Suites103and104,Ames,Iowa50010,USA TheAtrium,SouthernGate,Chichester,WestSussex,PO198SQ,UK 9600GarsingtonRoad,Oxford,OX42DQ,UK Fordetailsofourglobaleditorialoffices,forcustomerservicesandforinformationabouthowtoapply forpermissiontoreusethecopyrightmaterialinthisbookpleaseseeourwebsiteat www.wiley.com/wiley-blackwell. Authorizationtophotocopyitemsforinternalorpersonaluse,ortheinternalorpersonaluseofspecific clients,isgrantedbyBlackwellPublishing,providedthatthebasefeeispaiddirectlytotheCopyright ClearanceCenter,222RosewoodDrive,Danvers,MA01923.Forthoseorganizationsthathavebeen grantedaphotocopylicensebyCCC,aseparatesystemofpaymentshasbeenarranged.Thefeecodes forusersoftheTransactionalReportingServiceareISBN-13:978-1-1181-2563-2/2014. Designationsusedbycompaniestodistinguishtheirproductsareoftenclaimedastrademarks.All brandnamesandproductnamesusedinthisbookaretradenames,servicemarks,trademarksor registeredtrademarksoftheirrespectiveowners.Thepublisherisnotassociatedwithanyproductor vendormentionedinthisbook. Thecontentsofthisworkareintendedtofurthergeneralscientificresearch,understanding,and discussiononlyandarenotintendedandshouldnotberelieduponasrecommendingorpromotinga specificmethod,diagnosis,ortreatmentbyhealthsciencepractitionersforanyparticularpatient.The publisherandtheauthormakenorepresentationsorwarrantieswithrespecttotheaccuracyor completenessofthecontentsofthisworkandspecificallydisclaimallwarranties,includingwithout limitationanyimpliedwarrantiesoffitnessforaparticularpurpose.Inviewofongoingresearch, equipmentmodifications,changesingovernmentalregulations,andtheconstantflowofinformation relatingtotheuseofmedicines,equipment,anddevices,thereaderisurgedtoreviewandevaluatethe informationprovidedinthepackageinsertorinstructionsforeachmedicine,equipment,ordevicefor, amongotherthings,anychangesintheinstructionsorindicationofusageandforaddedwarningsand precautions.Readersshouldconsultwithaspecialistwhereappropriate.Thefactthatanorganization orWebsiteisreferredtointhisworkasacitationand/orapotentialsourceoffurtherinformationdoes notmeanthattheauthororthepublisherendorsestheinformationtheorganizationorWebsitemay provideorrecommendationsitmaymake.Further,readersshouldbeawarethatInternetWebsites listedinthisworkmayhavechangedordisappearedbetweenwhenthisworkwaswrittenandwhenit isread.Nowarrantymaybecreatedorextendedbyanypromotionalstatementsforthiswork.Neither thepublishernortheauthorshallbeliableforanydamagesarisingherefrom. LibraryofCongressCataloging-in-PublicationData Broussard,DianneM.,author. Thecerebellum:learningmovement,language,andsocialskills/DianneM.Broussard. p.;cm. Includesbibliographicalreferencesandindex. ISBN978-1-118-12563-2(cloth:alk.paper)–ISBN978-1-118-73007-2(emobi)– ISBN978-1-118-73013-3–ISBN978-1-118-73025-6(epdf)–ISBN978-1-118-73034-8(epub) I.Title. [DNLM:1.Cerebellum. 2.ExecutiveFunction. 3.NeuralPathways–physiology. 4.PsychomotorPerformance–physiology. WL320] QP379 (cid:3) 612.827–dc23 2013027959 AcataloguerecordforthisbookisavailablefromtheBritishLibrary. Wileyalsopublishesitsbooksinavarietyofelectronicformats.Somecontentthatappearsinprintmay notbeavailableinelectronicbooks. Coverimages:Backgroundart:MargaretBelknap;Topright:DianneM.Broussard;Bottomleft: (cid:2)C istockphoto:CEFutcher;Bottomright:(cid:2)C istockphoto:PicturePartners CoverdesignbyModernAlchemyLLC Setin9.5/12.5ptPalatinobyAptara®Inc.,NewDelhi,India 1 2014 Contents Acknowledgments ix Introduction xi SectionI: TheNeuronalMachine 1 1 StructureandPhysiology 3 AnatomyoftheCerebellarCortex 3 PhysiologyoftheCerebellarCortex 6 SubdivisionsoftheCerebellum 10 TheGatekeepers:VestibularandDeepCerebellarNuclei 13 AfferentConnectionsoftheCerebellum 15 EfferentConnectionsoftheCerebellum 19 References 21 FurtherReading 25 2 OperatingtheMachine 27 LearningintheCerebellarCortex 28 PatternRecognitionbytheCerebellum 30 NeuralNetworks 32 TheCerebellumasPartofa“ControlSystem” 32 MultipleSitesforCerebellarLearning? 34 TheCerebellarClock 36 Conclusions 38 References 38 FurtherReading 40 3 PlasticityintheCerebellarCortex 41 CerebellarLong-TermDepression 41 TheCalciumTrigger 42 TheSynapticConversation 44 TheMemoryTrace 46 WhatAboutPotentiation? 48 OtherSitesofPlasticity 51 Interneurons 52 Conclusions 55 References 55 FurtherReading 59 v vi Contents 4 AdjustingtheMemoryTrace 61 ConsolidationMechanisms 61 MemoryTransferandSynapticPlasticity 63 Mossy-FiberCollateralsintheDCN 64 IntrinsicPlasticityintheDCN 66 OtherChangestotheMemoryTrace 67 References 68 SectionII: MotorLearning 71 5 LearningaNewMotorResponse 73 LearningintheCerebellarCortex 75 CerebellarLTDandLearning 76 TheEngramfortheNMR 79 Conclusions 82 References 82 FurtherReading 85 6 RecalibrationforFineMotorControl 87 AStablePlatformforVision 87 AdjustinganOrientingMovement 95 AdjustingaTrackingMovement 97 Conclusions 99 References 99 FurtherReading 102 7 PerfectingLimbMovementsbyMotorLearning 103 UpdatingDynamicModels 104 ThrowingandPointing 107 SequenceLearning 109 SteppingandChangestogait 111 TheAgileMouse 113 Conclusions 114 References 114 FurtherReading 117 SectionIII: PrecisionControl 119 8 Coordination 121 PreciseOcularCoordination 122 CoordinatingtheEyesandtheHead 126 ErrorCorrectionforLimbMovements 127 PlanningforMultipleJoints 129 InternalModelsRevisited 132 Contents vii Conclusions 134 References 134 FurtherReading 137 9 BalanceandLocomotion 139 CerebellarAtaxia 140 SignalsfromtheInnerEar 142 CoordinatingLocomotion 146 Navigation 148 Conclusions 150 References 150 FurtherReading 152 10 Timing 153 TimingUsingDischargeRates 154 TimingUsingSynchronousFiring 161 Conclusions 165 References 166 FurtherReading 168 SectionIV: InterpretingtheWorld 169 11 IntelligenceandLanguage 173 GeneralIntelligence 175 ExecutiveFunction 177 Problem-solving 181 SpeechandLanguage 183 PossibleMechanisms 187 Conclusions 188 References 188 FurtherReading 191 12 Sensing,Feeling,andInteracting 193 SensoryPerception 194 AttendingtotheWorld 198 Prediction 200 MentalImagery 202 SocialSkills 204 Conclusions 208 References 208 FurtherReading 211 Summary:Whatdoesthecerebellumdo? 213 Index 215

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