北大神經(jīng)生物學(xué)課件神經(jīng)系統(tǒng)對(duì)運(yùn)動(dòng)的調(diào)節(jié)

CentralNervousSystemControlofMovementXING,GUOGANG(邢國(guó)剛),MD.Ph.D.北京大學(xué)神經(jīng)科學(xué)研究所NeuroscienceResearchInstitute,PekingUniversity第一頁(yè)，共一百一十二頁(yè)。1Introduction生命在于運(yùn)動(dòng)……運(yùn)動(dòng)是動(dòng)物維系個(gè)體生存和種族繁衍的基本功能之一第二頁(yè)，共一百一十二頁(yè)。2Howdoesacoordinatedmovementachieved?第三頁(yè)，共一百一十二頁(yè)。3Thementalbodyimageseemstobegeneratedbysomatosensory,proprioceptive,andvisualinputstotheposteriorparietalcortex(area5,area7)Abaseballpitcherplanningapitch…第四頁(yè)，共一百一十二頁(yè)。4Thehighestlevel－

strategyRepresentedbytheassociationareasofneocortexandbasalgangliaoftheforebrainIsconcernedwithstrategy:ThegoalofthemovementandthemovementstrategythatbestachievesthegoalThemotorcontrolhierarchyhavethreelevels第五頁(yè)，共一百一十二頁(yè)。5Themiddlelevel－tacticsRepresentedbythemotorcortexandcerebellumIsconcernedwithtacticsThesequencesofmusclecontractionsArrangedinspaceandtimeRequiredtosmoothlyandaccuratelyachievethestrategicgoalThemotorcontrolhierarchyhavethreelevels第六頁(yè)，共一百一十二頁(yè)。6Thelowestlevel－

execution

RepresentedbythebrainstemandspinalcordIsconcernedwithexecutionActionofthemotorneuronandinterneuronpoolsthatgeneratethegoal-directedmovementandmakeanynecessaryadjustmentsofpostureThemotorcontrolhierarchyhavethreelevels第七頁(yè)，共一百一十二頁(yè)。7Overview脊髓內(nèi)的“下運(yùn)動(dòng)神經(jīng)元(Lowerneurons)”，除了受到脊髓內(nèi)局部環(huán)路的影響外，還受到大腦皮層運(yùn)動(dòng)區(qū)及腦干中許多“上運(yùn)動(dòng)神經(jīng)元(Upperneurons)”的支配和協(xié)調(diào)基底神經(jīng)節(jié)和小腦則向那些上運(yùn)動(dòng)神經(jīng)元提供某種感覺、認(rèn)知或感性的信息，使運(yùn)動(dòng)更加精確和協(xié)調(diào)第八頁(yè)，共一百一十二頁(yè)。8第九頁(yè)，共一百一十二頁(yè)。9SensorimotorcortexBasalgangliaCerebellumBrainstemSpinalcordInterneuron-MotorneuronMotorunit(finalcommonpathway)Receptorsmuscle,skin,joint12345Theseprogramsareaccessed,executed,andmodifiedbydescendingcommandsfromthebrainThebrain’scommandandcontrolofthemotorprogramsinthespinalcordThemotorsystemconsistsofallourmusclesandtheneuronsthatcommandthemThespinalcordcontainscertainmotorprogramsforthegenerationofcoordinatedmovementsThemotorcontrolcanbedividedintotwoparts:Thespinalcord’scommandandcontrolofcoordinatedmusclecontraction第十頁(yè)，共一百一十二頁(yè)。10Part1

SpinalControlofMovement第十一頁(yè)，共一百一十二頁(yè)。11Introduction“runningaroundlikeachickenwithitsheadcutoff”

Therhythmicmovementscouldbeelicitedinthehindlegsofcatsanddogslongaftertheirspinalcordshadbeenseveredfromtherestofthecentralnervoussystem–CharlesSherington&GrahamBrown(English)Theimportanceofcircuitrywithinthespinalcordforthecoordinatedcontrolofmovements第十二頁(yè)，共一百一十二頁(yè)。12Spinalcircuitry

controlofmovement第十三頁(yè)，共一百一十二頁(yè)。13Spinalmotorneurons

Alphamotorneurons

Innervateskeletalmuscle(extrafusalmuscle)

Function:directlycommandmusclecontractGammamotorneurons

Innervatemusclespindle(intrafusalmuscle)

Function:regulatingthemusclespindleInterneurons

AllowscoordinatedmotorprogramstobegeneratedFinalcommonpathway(directlycommandmusclecontract)Compare:

UppermotorneuronsLowermotorneurons第十四頁(yè)，共一百一十二頁(yè)。14TheLowerMotorNeurons

－AlphamotorneuronsAlphamotorneuronsaredirectlyresponsibleforthegenerationofforcebymuscleMotorunit:onealphamotor

neuronandallofthemusclefibersitinnervatescollectivelymakeupmotorunit,theelementarycomponentofmotorcontrolMotorneuronpool:Thecollectionofalphamotorneuronsthatinnervatesasinglemuscle第十五頁(yè)，共一百一十二頁(yè)。15ThemotorunitisanalphamotorneuronandallofthemuscleAmotorunitandmotorneuronpoolThemotorneuronpoolisallofthealphamotorneuronsthatinnervateonemuscle第十六頁(yè)，共一百一十二頁(yè)。16Muscleinnervationbylowermotorneurons30mixedspinalnervescervical1-8thoracic1-12lumbar1-5sacral1-5TheventralhornofthespinalcordcontainsmotorneuronsthatinnervateskeletalmusclefibersVentralroots+Dorsalroot=Spinalnerve第十七頁(yè)，共一百一十二頁(yè)。17Thecervicalenlargementofthespinalcordcontainsthemotorneuronsthatinnervatethearmmuscles

Thelumbarenlargementcontainsneuronsthatinnervatethemusclesoftheleg

支配上肢的神經(jīng)元集群位于頸膨大處，支配下肢的則在腰膨大處ThedistributionofmotorneuronsinthespinalcordThemotorneuronsthatinnervatedistalandproximalmusculaturearefoundmainlyinthecervicalandlumbar-sacralsegmentsofthespinalcordWhereasthoseinnervatingaxialmusculaturearefoundatalllevelsSegmentsC3-T1haveaswollenventralhornthatinnervateskeletalmusculatureinarmsSegmentsL1-S3haveaswollenventralhornthatinnervateskeletalmusculatureinlegs第十八頁(yè)，共一百一十二頁(yè)。18Motorneuronscontrollingflexorsliedorsaltothosecontrollingextensors

Motorneuronscontrollingaxialmusclesliemedialtothosecontrollingdistalmuscles支配軀干部肌肉的神經(jīng)元位于脊髓前角灰質(zhì)最內(nèi)側(cè)，由此向外排列的神經(jīng)元?jiǎng)t支配肢體由近及遠(yuǎn)分布的肌肉Thedistributionoflowermotorneuronsintheventralhorn第十九頁(yè)，共一百一十二頁(yè)。19Muscleweaknessandparalysis－所支配的骨骼肌癱瘓、肌張力下降、腱反射消失(軟癱)肌萎縮、纖維顫動(dòng)或肌束顫動(dòng)見于如肌萎縮側(cè)索硬化癥(amyotrophiclateralsclerosis,

ALS)：Selectivedamagetoalphamotorneurons(degeneration)－脊髓-運(yùn)動(dòng)神經(jīng)元的進(jìn)行性潰變?yōu)橹飨逻\(yùn)動(dòng)神經(jīng)元的損傷表現(xiàn)第二十頁(yè)，共一百一十二頁(yè)。20LouGehrigFarewellSpeech:"Fans,forthepasttwoweeksyouhavebeenreadingaboutthebadbreakIgot.YettodayIconsidermyselftheluckiestmanonthefaceofthisearth.……SoIcloseinsayingthatImayhavehadatoughbreak,butIhaveanawfullottolivefor."LouGehring,astarbaseballplayerwithNewYorkYankees,whodiedofALS(amyotrophiclateralsclerosis)in1936第二十一頁(yè)，共一百一十二頁(yè)。21InputstoalphamotorneuronsThisinputisimportantfortheinitiationandcontrolofvoluntarymovementThisinputmaybeexcitatoryorinhibitoryandispartofthecircuitrythatgeneratesthespinalmotorprogramsThisinputprovidesfeedbackaboutmusclelength第二十二頁(yè)，共一百一十二頁(yè)。22SPINALCONTROLOFMOTORUNITSHowtheactivityofthemotorneuronisitselfcontrolled?第二十三頁(yè)，共一百一十二頁(yè)。23ReflexReceptorEffectorSpinalMotorNeuron(center)SkinMuscleJointMusclecontractionHigherCenterVoluntarymovementsNociceptivestimulusStretchmusclesPosturemaintenance第二十四頁(yè)，共一百一十二頁(yè)。24Themyotaticreflex(stretchreflex)Twotypesofmyotaticreflex

－TendonreflexandmuscletonusA.Tendonreflex

—快速牽拉肌腱而發(fā)生的牽張反射Clinicapplication：了解脊髓不同節(jié)段的功能狀態(tài)

單突觸反射潛伏期很短，約0.7s只夠一次突觸傳遞時(shí)間延擱第二十五頁(yè)，共一百一十二頁(yè)。25+第二十六頁(yè)，共一百一十二頁(yè)。26B.Muscletonus

肌肉受到緩慢而持續(xù)的牽拉而發(fā)生的收縮－使骨骼肌能保持一定的肌肉張力意義:維持身體的姿勢(shì)(posture)，而不表現(xiàn)明顯的動(dòng)作表現(xiàn)：extensor(伸肌)和flexor(屈肌)都發(fā)生肌緊張直立時(shí)，以伸肌緊張為主，因重力作用于關(guān)節(jié)，使關(guān)節(jié)趨向彎曲，伸肌受到牽拉，引起肌緊張反射，肌緊張度增加以對(duì)抗關(guān)節(jié)屈曲因重力持續(xù)作用于關(guān)節(jié)，肌緊張也就持續(xù)發(fā)生。使直立姿勢(shì)得以維持多突觸反射潛伏期較長(zhǎng)，經(jīng)過多個(gè)突觸傳遞第二十七頁(yè)，共一百一十二頁(yè)。27

Musclespindles:ConsistsofspecializedskeletalmusclefibersInthismiddleregion,groupIa

sensoryaxonswraparoundthemusclefibersofthespindleThespindlesandtheirassociatedIaaxons,specializedforthedetectionofchangesinmusclelength(stretch):proprioceptorsandpropriceptionIaaxonsarethethickestmyelinatedaxonssotheyconductactionpotentialsveryrapidlyIaaxonsenterthespinalcordviathedorsalroots,branchrepeatedly,andformexcitatorysynapsesuponbothinterneuronsandalphamotorneuronsoftheventralhornsProprioceptionfrommusclespindlesMusclespindle

是一種可感受肌肉長(zhǎng)度變化或感受牽拉刺激的本體感受器梭內(nèi)肌感受部裝置位于中間,收縮成分位于兩端,梭內(nèi)肌收縮時(shí)或牽拉梭外肌時(shí),感受裝置對(duì)牽拉敏感性增高

第二十八頁(yè)，共一百一十二頁(yè)。28+第二十九頁(yè)，共一百一十二頁(yè)。29GammamotorneuronsExtrafusalfibersareinnervatedbyalphamotorneuronsIntrafusalfibersreceivetheirmotorinnervationbyanothertypeoflowermotorneuroncalledagammamotorneuronGammamotorneuronsinnervatetheintrafusalmusclefiberatthetwoendsofthemusclespindleActivationofthesefiberscausesacontractionofthetwopolesofthemusclespindle,therebypullingonthenon-contractileequatorialregionandkeepingtheIaafferentsactive第三十頁(yè)，共一百一十二頁(yè)。30ThefunctionofgammamotorneuronsActivationofalphamotorneuronsshortenstheextrafusalmusclefibers.Ifthemusclespindlebecomesslack,itgoes“offtheair”andnolongerreportsthelengthofthemuscleActivationofgammamotorneuronscontractsthepolesofthespindle,keepingit“ontheair”第三十一頁(yè)，共一百一十二頁(yè)。31TheGammaloopChangingtheactivityofthegammamotorneuronschangesthesetpointofthemyotaticfeedbackloopTheGammaloopGammamotorneuronintrafusalmusclefiberIaafferentalphamotorneuronextrafusalmusclefibersAlphaandgammamotorneuronsaresimultaneouslyactivatedbydescendingcommandsfromthebrain第三十二頁(yè)，共一百一十二頁(yè)。32ProprioceptionfromGolgitendonorgansGolgitendonorganActslikeastraingauge,itmonitorsmuscletension,ortheforceofcontractionLocatedatthejunctionofthemuscleandthetendonandareinnervatedbygroupIbsensoryaxonsthatareslightlysmallerthantheIaaxonsinnervatingthemusclespindles第三十三頁(yè)，共一百一十二頁(yè)。33GolgitendonorgansrespondtoincreasedtensiononthemuscleandtransmitthisinformationtothespinalcordviatypeIbsensoryafferents.Becausetheactivatedmuscledoesnotchangelength,theIaafferentsremainsilentinthisexample當(dāng)肌肉等長(zhǎng)收縮時(shí),腱器官傳入沖動(dòng)頻率,肌梭傳入沖動(dòng)頻率不變;當(dāng)肌肉等張收縮時(shí),腱器官傳入沖動(dòng)頻率不變,肌梭傳入沖動(dòng)頻率Golgitendon

organslieinseriesbetweenthemusclefibersandtheirpointsofattachmentMusclespindlesarearrangedparalleltotheextrafusalfibers第三十四頁(yè)，共一百一十二頁(yè)。34TheIbafferentsenterthespinalcord,branchrepeatedly,andsynapseoninterneuronsintheventralhornSomeoftheseinterneuronsforminhibitoryconnectionswiththealphamotorneuronsinnervatingthesamemuscle.ThisisthebasisforthereversemyotaticreflexCircuitryofthereversemyotaticreflex＋-Musclespindlesituatedinparallelwiththemusclefibers,GolgitendonorgansaresituatedinseriesIaactivityfromthespindleencodesmusclelengthinformationWhileIbactivityfromtheGolgitendonorganencodesmuscletensioninformation腱器官是張力感受器,其傳入沖動(dòng)對(duì)同一肌肉運(yùn)動(dòng)神經(jīng)元起抑制作用肌梭是長(zhǎng)度感受器,其傳入沖動(dòng)對(duì)同一肌肉運(yùn)動(dòng)神經(jīng)元起興奮作用當(dāng)肌肉被動(dòng)牽拉時(shí),二者傳入沖動(dòng)頻率均：首先興奮肌梭發(fā)動(dòng)牽張反射,肌肉收縮,牽拉力大到一定程度,興奮腱器官而抑制牽張反射牽張反射的特點(diǎn)：反射弧簡(jiǎn)單，感受器和效應(yīng)器在同一塊肌肉第三十五頁(yè)，共一百一十二頁(yè)。35SignificanceofthereversemyotaticreflexInextremecircumstances,thisreflexarcprotectsthemusclefromoverloadThenormalfunctionistoregulatemuscletensionwithinanoptimalrangeAsmuscletension

increases,theinhibitionofthealphamotorneuronslowsmusclecontractionAsmuscletensionfalls,theinhibitionofthealphamotorneuronisreduced,andmusclecontractionincreasesThistypeofproprioceptivefeedbackisthoughttobeparticularlyimportantfortheproperexecutionoffinemotoracts,suchasthemanipulationoffragileobjectswiththehands,whichrequireasteadybutnottoopowerfulgrip第三十六頁(yè)，共一百一十二頁(yè)。36SpinalinterneuronsMostoftheinputtothealphamotorneuronscomesfrominterneuronsofthespinalcordInterneuronsreceivesynapticinputfromprimarysensoryaxons,descendingaxonsfromthebrain,andcollateralsoflowermotorneuronaxonsTheinterneuronsarethemselvesnetworkedtogetherinawaythatallowscoordinatedmotorprogramstobegeneratedinresponsetotheirmanyinputs第三十七頁(yè)，共一百一十二頁(yè)。37Reciprocalinhibitionofflexorsandextensorsofthesamejoint+++-InhibitoryinputContractionofonesetofmusclesaccompaniedbytherelaxationoftheantagonistmusclesiscalled

reciprocalinhibitionReciprocalinhibitionisalsousedbydescendingpathwaystoovercomethepowerfulmyotaticreflex第三十八頁(yè)，共一百一十二頁(yè)。38(A

&Cfibers)ExcitatoryinputFlexorreflex:usedtowithdrawalimbfromanaversivestimulus,suchasthewithdrawalofyourfootfromthethumbtackThepainfibersenteringthespinalcordbranchprofusedlyandactivateinterneuronsinseveralspinalsegments.Thesecellseventuallyexcite

thealphamotorneuronsthatcontrolalloftheflexormusclesoftheaffectedlimb(andinhibitoryinterneuronsarealsorecruitedtoinhibitthealphasthatcontroltheextensors)++++Circuitryofthepolysynapticflexorreflex第三十九頁(yè)，共一百一十二頁(yè)。39Circuitryofthecrossed-extensorreflexSpinalcordIpsilateralextensorsinhibitedContralateralextensorsactivated(contract)Ipsilateralflexorsactivated(contract)ContralateralflexorsinhibitedIpsilateralkneejointclosesuptoremovefootfromtackContralateralkneejointopensuptosupportgreaterweight第四十頁(yè)，共一百一十二頁(yè)。40ThegenerationofspinalmotorprogramsforwalkingHeadlesschickens;behaviorAcompletetransectionofacat’sspinalcordatthemid-thoraciclevelleavesthehindlimbscapableofgeneratingcoordinatedwalkingmovementsWhenyouwalk,youalternatelywithdrawandextendyourtwolegs

Whatisthemechanismtocoordinatethetiming?Thiscouldbedescendingcommandsfromuppermotorneurons?No!Thiscontrolisexertedfromwithinthespinalcord?Yes!第四十一頁(yè)，共一百一十二頁(yè)。41第四十二頁(yè)，共一百一十二頁(yè)。42CentralpatterngeneratorswithinthespinalcordThecircuitforthecoordinatedcontrolofwalkingmustresidewithinthespinalcordIngeneral,circuitsthatgiverisetorhythmicmotoractivityarecalledcentralpatterngenerators

Howdoneuralcircuitsgeneraterhythmicpatternsofactivity?Thesimplestpatterngenerators,aresingleneuronswhosemembranepropertiesendowthemwithpacemakerproperties(pacemakerneurons)第四十三頁(yè)，共一百一十二頁(yè)。43Rhythmicactivityinaspinalinterneuron(Pacemakerneuron)SomeneuronsrespondtotheactivationofNMDAreceptorswithrhythmicdepolarization(a)Intherestingstate,theNMDAreceptorchannelsandthecalcium-activatedpotassiumchannelsareclosed(b)GlutamatecausestheNMDAreceptorstoopen,thecellmembranetodepolarize,andCa2+ionstoenterthecell(c)Theriseinintracellular[Ca2+]causestheCa2+-activatedpotassiumchannelstoopen.K+ionsleavetheneuron,hyperpolarizingthemembrane.ThehyperpolarizationallowsMg2+ionstoenterandclogtheNMDAchannel,arrestingtheflowofCa2+(d)As[Ca2+]falls,thepotassiumchannelsclose,resettingthemembraneforanotheroscillation第四十四頁(yè)，共一百一十二頁(yè)。44Apossiblecircuitforrhythmicalternatingactivity(walking)Walkingisinitiatedwhenasteadyinputexcitestwointerneuronsthatconnecttothemotorneuronscontrollingtheflexorsandextensors,respectively.Theinterneuronsrespondtoacontinuousinputbygeneratingburstsofoutputs.Theactivitiesofthetwointerneuronsalternatebecausetheyinhibiteachotherviaother(inhibitory)interneurons.Thus,aburstofactivityinoneinterneuronstronglyinhibitstheother(Pacemakerneuron)(Pacemakerneuron)第四十五頁(yè)，共一百一十二頁(yè)。45Part2

BrainControlofMovement第四十六頁(yè)，共一百一十二頁(yè)。46Howdoesthebraincommunicatewiththemotorneuronsofthespinalcord?第四十七頁(yè)，共一百一十二頁(yè)。47Overview來自高級(jí)中樞的下行投射對(duì)腦干和脊髓環(huán)路的運(yùn)動(dòng)控制主要影響隨意運(yùn)動(dòng)和朝向目標(biāo)的運(yùn)動(dòng)發(fā)出下行投射的中樞位于腦干某些部位以及大腦皮質(zhì)的運(yùn)動(dòng)區(qū)域腦干內(nèi)的兩個(gè)主要結(jié)構(gòu)－前庭核和網(wǎng)狀結(jié)構(gòu)對(duì)維持姿勢(shì)很重要皮質(zhì)主要運(yùn)動(dòng)區(qū)和次要運(yùn)動(dòng)區(qū)(運(yùn)動(dòng)前區(qū))主要對(duì)運(yùn)動(dòng)進(jìn)行計(jì)劃并使運(yùn)動(dòng)高效率執(zhí)行，皮質(zhì)的這種作用是通過對(duì)腦干內(nèi)的某些高級(jí)中樞以及腦干、脊髓中運(yùn)動(dòng)神經(jīng)元和環(huán)路中間神經(jīng)元的控制來完成的第四十八頁(yè)，共一百一十二頁(yè)。48Thedescendingspinaltractscontrolofmovement第四十九頁(yè)，共一百一十二頁(yè)。49ThedescendingspinaltractsControlpostureandunderbrainstemcontrolContolvoluntary

movementsofthedistalmusculatureandunderdirectcorticalcontrol第五十頁(yè)，共一百一十二頁(yè)。50(Pyramidaltract)ControlfinemovementsofthearmsandfingersTheeffectsoflateralpathwaylesions(inmonkeys)－byDonaldLawrence&HansKuypers(inthelate1960s)TheLateralPathwaysXJoinincorticospinaltractinthelateralcolumnofthespinalcordTerminateinthedorsolateralregionoftheventralhornsandintermedialgraymatterWheremotorneuronsandinterneuronscontrolthedistalmuscles,particularlyflexorsInputfromfrontalcortexX第五十一頁(yè)，共一百一十二頁(yè)。51TheVentromedialPathwaysContainfourdescendingtracts

thevestibulospinaltractthetectospinaltractthepontinereticulospinaltractthemedullaryreticulospinaltract

Oringinateinthebrainstem

TerminateamongthespinalinterneuronsControllingproximalandaxialmusclesUsesensoryinformationaboutbalance,bodyposition,andthevisualenviromenttoreflexivelymaintainbalanceandbodyposture第五十二頁(yè)，共一百一十二頁(yè)。52Thevestibulospinalandtectospinaltractskeeptheheadbalanceontheshouldersasthebodymovesthroughspace,andtheyturntheheadinresponsetonewsensorystimuli(midbrain)RecivedirectinputfromretinaReciveprojectionsfromvisualcortex,aswellasafferentscarringsomatosensoryandauditoryinformationConstructamapofthewordaroundusThemotionofthefluidinthislabyrinth,whichaccompaniesmovementsofthehead,activateshaircellsthatsignalthevestibularnucleiviacranialnerveVIIIOne:ProjectsbilaterallydownthespinalcordandactivatesthecervicalspinalcircuitsthatcontrolneckandbackmusclesandguidheadmovementRelysensoryinformationfromthevestibularlabyrinthintheinnerearAnother:ProjectsipsilaterallyasfardownasthelumbarspinalcordIthelpsusmaintainanuprightandbalancedposturebyfacilitatingextensormotorneuronsofthelegsStimulationatonesiteinthismapLeadtoanorientingresponsethatdirectstheheadandeyestomove…第五十三頁(yè)，共一百一十二頁(yè)。53ThepontinereticulospinaltractenhancetheantigravityreflexesofthespinalcordByfacilitatingtheextensorsofthelowerlimbsHelpsmaintainastandingposturebyresistingtheeffectsofgravityThemedullaryreticulospinaltracthastheoppositeeffect:itliberatestheantigravitymusclesfromreflex

controlActivityinbothreticulospinaltractsiscontrolledbydescendingsignalsfromthecortexControlpostureofthetrunkandtheantigravitymusclesofthelimbs第五十四頁(yè)，共一百一十二頁(yè)。54AsummaryofthemajordescendingspinaltractsandtheirpointsoforiginMotorcortexdirectlyactivatesspinalmotorneuronsandliberatesthemfromreflexcontrolbycommunicatingwiththenucleioftheventromedialpathwaysSo,thecortexiskeyforvoluntarymovementandbehaviorTheventromedialpathwaysoriginatefromseveralregionsofthebrainstemandparticipatemainlyinthemaintenanceofpostureandcertainreflexmovementsInitiationofavoluntarymovements,requiresinstructionsthatdescendfromthemotorcortexalongthelateralpathways第五十五頁(yè)，共一百一十二頁(yè)。55ourattention第五十六頁(yè)，共一百一十二頁(yè)。56MotorCortexMotorCortex(Primarymotorcortex)(Premotorarea)(lateralregion)(Supplementarymotorarea)(medialregion)AreasofneocortexintimatelyinvolvedintheplanningandinstructionofvoluntarymovementThemotorcortexisacircuscribedregionofthefrontallobeArea4liesjustanteriortothecentralsulcusontheprecentralgyrusArea6liesjustanteriortoarea4第五十七頁(yè)，共一百一十二頁(yè)。57第五十八頁(yè)，共一百一十二頁(yè)。58Asomatotopicmap

ofthehumanprecentralgyrus運(yùn)動(dòng)皮層對(duì)軀體運(yùn)動(dòng)的調(diào)控特點(diǎn)：交叉性：但頭面部肌肉的支配是雙側(cè)性的有精細(xì)的功能定位：大體呈身體倒影:在4區(qū)內(nèi)側(cè)近中線部位是下肢代表區(qū)，向外側(cè)依次為軀干、前臂、手指，最外側(cè)靠近外側(cè)溝處為面部和舌代表區(qū)頭面代表區(qū)內(nèi)部為正立的運(yùn)動(dòng)精細(xì)部位其代表區(qū)大：如手指和面部第五十九頁(yè)，共一百一十二頁(yè)。59Area5PosteriorparietalcortexPrefrontalareas:inhumansarethoughttobeimportantforabstractthought,decisionmaking,andanticipatingtheconsequencesofactionArea6and4togethercontributemostoftheaxonstothedescendingcorticospinaltractArea6liesatthejunctionwheresignalsencodingwhatactionsaredesiredareconvertedintosignalsthatspecifyhowtheactionwillbecarriedoutPrefrontalareas,alongwiththeposteriorparietalcortex,representthehighestlevelsofthemotorcontrolhierarchy,wheredecisionsaremadeaboutwhatactionstotakeandtheirlikelyoutcomeBoththeprefrontalandtheparietalcortexsendaxonsthatconvergeoncorticalarea6Area5,receivesinputsfromprimarysomatosensorycorticalarea3,1,2Area7,receivesinputsfromhigher-ordervisualcorticalareas(MT)ThecontributionsofposteriorparietalandprefrontalcortexPrimarysomatosensorycorticalarea3,1,2Higher-ordervisualcorticalareas(MT)Area7PrefrontalareasForabstractthought,decisionmaking,andanticipatingtheconsequencesofactionArea6Area4Descendingcorticospinaltracts第六十頁(yè)，共一百一十二頁(yè)。60Theinput-outputorganizationofM1PyramidalcellIncorticallayerV(Motorcortex,M1,Area4)Area6(anterior)Area3,1,2(posterior)Lowermotorneuronsofspinalcord

Thalamus(VLc)CerebellumBrainstemBasalgangliaThalamus(VLo)Somatosensoryinputs第六十一頁(yè)，共一百一十二頁(yè)。61Higher-ordermotorplanningUsingPET(positronemissiontomography)tomonitorchangesinthepatternsofcorticalactivationthataccompanyvoluntarymovementsWhenthesubjectswereaskedtoperformaseriesoffingermovementsfrommemory,thefollowingregionsofcortexshowedincreasedbloodflow:thesomatosensoryandposteriorparietalareas,partsoftheprefrontalcortex(area8),area6,andarea4－PlayaroleingeneratingtheintentiontomoveandconvertingthatintentionintoaplanofactionWhenthesubjectswereaskedonlytorehearsethemovementmentallywithoutactuallymovingthefinger,area6remainedactivebutarea4didnot－PerRolandandhiscolleagues(Danishneurologist)第六十二頁(yè)，共一百一十二頁(yè)。62Thedischargeofacellinthepremotorarea(PMA)beforeamovementArea6(SMAandPMA)playsanimportantroleintheplanningofmovement,particularlycomplexmovementsequencesofthedistalmusculature(a)Ready:Amonkeysitsbeforeapaneloflights.Thetaskistowaitforaninstructionstimulusthatwillinformhimothemovementrequiredtoreceiveajuicereward,thenperformthemovementwhenatriggerstimulusgoeson.TheactivityofaneuroninPMAisrecordedduringthetask(b)Set:theinstructionstimulusoccursatthetimeindicatedbytheupwardarrow,resultinginthedischargeoftheneuroninPMA(c)Go:shortlyafterthemovementisinitiated,thePMAcellceasesfiring第六十三頁(yè)，共一百一十二頁(yè)。63初級(jí)運(yùn)動(dòng)皮層和前運(yùn)動(dòng)區(qū)運(yùn)動(dòng)皮層(motorcortex)初級(jí)運(yùn)動(dòng)皮層(primarymotorarea)，即4區(qū)次級(jí)運(yùn)動(dòng)區(qū)或稱前運(yùn)動(dòng)區(qū)輔助運(yùn)動(dòng)區(qū)(supplementarymotorarea)，位于初級(jí)運(yùn)動(dòng)皮層前面的6區(qū)皮層的內(nèi)側(cè)部分，大部分在大腦半球的內(nèi)側(cè)面外側(cè)前運(yùn)動(dòng)區(qū)(lateralpremotorarea)或稱前運(yùn)動(dòng)皮層(premotorcortex)，位于6區(qū)的外側(cè)部分第六十四頁(yè)，共一百一十二頁(yè)。64皮層神經(jīng)元運(yùn)動(dòng)皮層中的神經(jīng)細(xì)胞可分為兩大類錐體細(xì)胞，特征是具有向皮層表面伸展的頂樹突。其軸突離開運(yùn)動(dòng)皮層到其他皮層或皮層下結(jié)構(gòu)，是主要的傳出神經(jīng)元非錐體細(xì)胞，包括星形細(xì)胞、籃狀細(xì)胞和顆粒細(xì)胞，多數(shù)屬于抑制性神經(jīng)元第六十五頁(yè)，共一百一十二頁(yè)。65第Ⅵ層的錐體細(xì)胞投射至丘腦，它們也有上行軸突側(cè)支至皮層的各層

皮層各層中錐體細(xì)胞的投射第II、III層中的錐體細(xì)胞投射至其他皮層區(qū)，位置較淺的細(xì)胞投射至同側(cè)皮層(如輔助運(yùn)動(dòng)區(qū)、前運(yùn)動(dòng)皮層、中央溝后的感覺皮層)，較深的投射至對(duì)側(cè)皮層大多數(shù)向皮層下結(jié)構(gòu)的投射起源于第Ⅴ層的錐體細(xì)胞，皮層脊髓神經(jīng)元在第Ⅴ層的深部，其中包括最大的錐體細(xì)胞即Betz細(xì)胞。較淺的第Ⅴ層錐體細(xì)胞則投射至延髓、橋腦和紅核，最淺的第Ⅴ層細(xì)胞投射至紋狀體第六十六頁(yè)，共一百一十二頁(yè)。66皮層脊髓束和皮層延髓束大腦皮層通過皮層脊髓束和皮層延髓束控制運(yùn)動(dòng)皮層延髓束終止于延髓的顱神經(jīng)運(yùn)動(dòng)核，控制面部肌肉的活動(dòng)皮層脊髓束則支配脊髓運(yùn)動(dòng)核，控制軀體肌肉的活動(dòng)第六十七頁(yè)，共一百一十二頁(yè)。67皮層脊髓束和皮層延髓束走行皮層脊髓束和皮層延髓束一起經(jīng)內(nèi)囊下行到中腦腹側(cè)，分散地穿過橋腦核，在延髓集合成錐體繼續(xù)下行至延髓和脊髓交界處，約3/4的纖維交叉至對(duì)側(cè)而在脊髓的背外側(cè)束下行，稱為外側(cè)皮層脊髓束，另1/4纖維不交叉而在脊髓腹側(cè)下行，稱為腹側(cè)皮層脊髓束第六十八頁(yè)，共一百一十二頁(yè)。68皮層脊髓束和皮層延髓束走行外側(cè)皮層脊髓束主要終止于脊髓腹角外側(cè)的運(yùn)動(dòng)神經(jīng)元和中間區(qū)內(nèi)的中間神經(jīng)元，腹側(cè)皮層脊髓束則主要投射至雙側(cè)腹角腹內(nèi)側(cè)的運(yùn)動(dòng)神經(jīng)元和中間區(qū)內(nèi)側(cè)的中間神經(jīng)元，支配軀干中線肌肉和肢體近側(cè)肌肉的皮層延髓束終止于腦干的顱神經(jīng)感覺和運(yùn)動(dòng)核第六十九頁(yè)，共一百一十二頁(yè)。69

運(yùn)動(dòng)皮層通過兩條獨(dú)立的通路分別影響脊髓運(yùn)動(dòng)神經(jīng)元一條經(jīng)過錐體束的直接通路直達(dá)脊髓灰質(zhì)外側(cè)和內(nèi)側(cè)的運(yùn)動(dòng)神經(jīng)元一條間接通路，即經(jīng)過紅核和網(wǎng)狀結(jié)構(gòu)的接替第七十頁(yè)，共一百一十二頁(yè)。70皮層脊髓纖維和運(yùn)動(dòng)神經(jīng)元的聯(lián)系皮層脊髓神經(jīng)元對(duì)脊髓的運(yùn)動(dòng)神經(jīng)元有直接突觸聯(lián)系，起到強(qiáng)烈的直接興奮性影響皮層脊髓神經(jīng)元間接影響運(yùn)動(dòng)神經(jīng)元皮層脊髓束的主要功能是控制肢體遠(yuǎn)端肌肉的活動(dòng)，特別是手指的活動(dòng)通過腦干神經(jīng)元間接地控制脊髓運(yùn)動(dòng)神經(jīng)元。在初級(jí)運(yùn)動(dòng)皮層、前運(yùn)動(dòng)皮層和輔助運(yùn)動(dòng)區(qū)，均有神經(jīng)元投射至腦干的網(wǎng)狀脊髓神經(jīng)元及其他下行神經(jīng)元而間接影響脊髓運(yùn)動(dòng)神經(jīng)元皮層脊髓神經(jīng)元經(jīng)過上頸段脊髓中的脊髓固有神經(jīng)元，再影響位于頸膨大中的支配前臂肌肉的運(yùn)動(dòng)神經(jīng)元經(jīng)過抑制性中間神經(jīng)元以施加抑制性影響第七十一頁(yè)，共一百一十二頁(yè)。71次級(jí)運(yùn)動(dòng)區(qū)的運(yùn)動(dòng)功能初級(jí)運(yùn)動(dòng)皮層主要和運(yùn)動(dòng)的執(zhí)行有關(guān)次級(jí)運(yùn)動(dòng)區(qū)主要與運(yùn)動(dòng)的準(zhǔn)備（策劃）有關(guān)使用PET影像研究局部腦血流與運(yùn)動(dòng)的關(guān)系：執(zhí)行簡(jiǎn)單動(dòng)作如單指按壓彈簧，則4區(qū)(S1)的手代表區(qū)血流增加；執(zhí)行復(fù)雜動(dòng)作如輪排手指，則輔助運(yùn)動(dòng)區(qū)(S2)的血流亦增加；若只默想復(fù)雜動(dòng)作，則僅有輔助運(yùn)動(dòng)區(qū)的血流增加第七十二頁(yè)，共一百一十二頁(yè)。72次級(jí)運(yùn)動(dòng)區(qū)的運(yùn)動(dòng)功能次級(jí)運(yùn)動(dòng)區(qū)是選擇正確的運(yùn)動(dòng)戰(zhàn)略所必需和初級(jí)運(yùn)動(dòng)皮層相比，損毀次級(jí)運(yùn)動(dòng)區(qū)(前運(yùn)動(dòng)皮層、輔助運(yùn)動(dòng)區(qū)和后頂葉皮層區(qū))引起更為復(fù)雜的運(yùn)動(dòng)異常，主要影響制定正確運(yùn)動(dòng)策略的能力一側(cè)的輔助運(yùn)動(dòng)區(qū)和前運(yùn)動(dòng)皮層被損毀后，猴不會(huì)使用對(duì)側(cè)前臂繞過擋在前面的透明塑料板，并通過側(cè)面的洞去取所看見的食物，相反，猴總是試圖直接去取食物而一再撞到塑料板上這種運(yùn)動(dòng)缺損和人的前額葉聯(lián)絡(luò)皮層或后頂葉皮層受損時(shí)引起的失用癥(apraxia)相似。這種病人能正常進(jìn)行簡(jiǎn)單的運(yùn)動(dòng)，但喪失了做需要有次序地收縮肌肉的較復(fù)雜的動(dòng)作的能力，例如刷牙和梳頭等第七十三頁(yè)，共一百一十二頁(yè)。73Thebasalgangliacontrolofmovement第七十四頁(yè)，共一百一十二頁(yè)。74基底神經(jīng)節(jié)(basalganglia)是從端腦衍生的一些皮層下神經(jīng)核團(tuán)的總稱，位于大腦兩半球的深部?；咨窠?jīng)節(jié)中與運(yùn)動(dòng)功能有關(guān)的主要是紋狀體(striatum)尾核與殼核是基底神經(jīng)節(jié)的主要輸入核(inputnuclei)，蒼白球的內(nèi)側(cè)部、腳內(nèi)核和黑質(zhì)網(wǎng)狀部是基底神經(jīng)節(jié)的主要輸出核(outputnuclei)，后兩者的結(jié)構(gòu)和機(jī)能也很相似第七十五頁(yè)，共一百一十二頁(yè)。75Asummaryoftheloopfromthecortextothebasalganlgliatothethalamusandbacktoarea6Cerebralcortex(SMA)CaudatenucleusPutamenNewStriatumGlobuspallidus(OldStriadum)StriadumThalamus(theventrallateralnucleus,VLo)Frontalcortexneuron(Input)(Output)紋狀體的主要傳入沖動(dòng)來自大腦皮層廣泛的區(qū)域(運(yùn)動(dòng)區(qū)、體感區(qū)、聯(lián)合區(qū)、邊緣區(qū)甚至頂葉)傳出沖動(dòng)經(jīng)過丘腦返回皮層，主要是輔助運(yùn)動(dòng)區(qū)(supplementarymotorarea)和運(yùn)動(dòng)前皮層(premotorcortex)，而與脊髓沒有直接的聯(lián)系第七十六頁(yè)，共一百一十二頁(yè)。76基底神經(jīng)節(jié)與大腦皮層的纖維聯(lián)系，與運(yùn)動(dòng)有關(guān)的主要為三條回路皮層新紋狀體(尾、殼核)蒼白球(內(nèi)側(cè)部)丘腦皮層皮層新紋狀體(尾、殼核)蒼白球(外)丘腦底核蒼白球(內(nèi))丘腦皮層皮層新紋狀體(尾、殼核)黑質(zhì)丘腦皮層基底神經(jīng)節(jié)與與大腦皮層之間主要的回路聯(lián)系第七十七頁(yè)，共一百一十二頁(yè)。771.皮層新紋狀體蒼白球(內(nèi)側(cè))丘腦皮層回路大腦皮層(包括運(yùn)動(dòng)區(qū)、體感區(qū)、聯(lián)合區(qū)、邊緣區(qū)甚至頂葉)

新紋狀體(包括尾核和殼核，即紋狀體的輸入核)

蒼白球的內(nèi)側(cè)部

丘腦(包括腹前核、腹外側(cè)核、內(nèi)側(cè)背核和中央中核)

大腦皮層(主要是運(yùn)動(dòng)區(qū)和輔助運(yùn)動(dòng)區(qū)皮層)第七十八頁(yè)，共一百一十二頁(yè)。782.皮層新紋狀體蒼白球(外)丘腦底核蒼白球(內(nèi))丘腦皮層回路大腦皮層相當(dāng)廣泛的區(qū)域

新紋狀體

蒼白球的外側(cè)部

丘腦底核蒼白球的內(nèi)側(cè)部

丘腦

皮層運(yùn)動(dòng)區(qū)和運(yùn)動(dòng)前區(qū)第七十九頁(yè)，共一百一十二頁(yè)。793.皮層新紋狀體黑質(zhì)丘腦皮層回路大腦皮層相當(dāng)廣泛的區(qū)域(包括運(yùn)動(dòng)區(qū)、體感區(qū)、聯(lián)合區(qū)、邊緣區(qū)甚至頂葉)

新紋狀體

黑質(zhì)網(wǎng)狀部

丘腦的腹前核和腹外側(cè)核

大腦皮層(運(yùn)動(dòng)區(qū)和輔助運(yùn)動(dòng)區(qū))紋狀體(尾殼核)黑質(zhì)間存在具有局部定位特征的往返纖維聯(lián)系從黑質(zhì)的致密部有纖維投向新紋狀體，它組成多巴胺能神經(jīng)元系統(tǒng)；但黑質(zhì)除有纖維至上丘和腳橋被蓋核外，沒有其他向下投射的纖維 所以，基底神經(jīng)節(jié)的運(yùn)動(dòng)功能是通過大腦皮層中與運(yùn)動(dòng)控制有關(guān)的區(qū)域而間接實(shí)現(xiàn)的第八十頁(yè)，共一百一十二頁(yè)。80直接通路易化運(yùn)動(dòng)直接通路包括通路1和3從皮層到新紋狀體(輸入核)的遞質(zhì)是谷氨酸，是興奮性的；從新紋狀體到蒼白球內(nèi)側(cè)部及黑質(zhì)網(wǎng)狀部(輸出核)的遞質(zhì)是GABA和P物質(zhì)，是抑制性的；從這些輸出核到丘腦的遞質(zhì)也是GABA，也是抑制性的；再由丘腦返回到皮層的輔助運(yùn)動(dòng)區(qū)，則是興奮性的。當(dāng)皮層有興奮沖動(dòng)下行到紋狀體的輸入核使它興奮時(shí)，就會(huì)使輸出核中的抑制細(xì)胞抑制，從而激活(去抑制)了丘腦的神經(jīng)元，再通過輔助運(yùn)動(dòng)區(qū)和運(yùn)動(dòng)前區(qū)激活運(yùn)動(dòng)皮層和向腦干及脊髓的投射神經(jīng)元，從而起易化運(yùn)動(dòng)的效果第八十一頁(yè)，共一百一十二頁(yè)。81間接通路抑制運(yùn)動(dòng)間接通路包括通路2從皮層到新紋狀體(輸入核)的遞質(zhì)是谷氨酸，是興奮性的；從紋狀體輸入核到蒼白球外側(cè)部的遞質(zhì)是GABA和腦啡肽，是抑制性的；從蒼白球外側(cè)部到丘腦底核的遞質(zhì)是GABA，也是抑制性的；從丘腦底核到蒼白球內(nèi)側(cè)部(輸出核)的遞質(zhì)是谷氨酸，是興奮性的；從蒼白球內(nèi)側(cè)部到丘腦的遞質(zhì)是GABA，抑制性的；再由丘腦返回到皮層的輔助運(yùn)動(dòng)區(qū)，則是興奮性的當(dāng)皮層與紋狀體輸入核興奮而使蒼白球外側(cè)部抑制時(shí)，轉(zhuǎn)而使丘腦底核產(chǎn)生興奮(去抑制)，也就興奮了紋狀體的輸出核，抑制了丘腦，從而降低了輔助運(yùn)動(dòng)區(qū)的興奮性，產(chǎn)生抑制運(yùn)動(dòng)的效果第八十二頁(yè)，共一百一十二頁(yè)。82黑質(zhì)－紋狀體多巴胺能投射易化運(yùn)動(dòng)從黑質(zhì)中的多巴胺能神經(jīng)元發(fā)出纖維到新紋狀體，易化運(yùn)動(dòng)興奮直接通路而易化運(yùn)動(dòng)，也就是使新紋狀體通過GABA與P物質(zhì)等抑制紋狀體的輸出核，興奮丘腦，最后起易化運(yùn)動(dòng)的作用。抑制間接通路而易化運(yùn)動(dòng)，就是使新紋狀體抑制，致使GABA與腦啡肽的抑制作用降低，因而興奮了蒼白球的外側(cè)部，抑制丘腦底核，從而抑制紋狀體輸出核，興奮了丘腦，最后也起易化運(yùn)動(dòng)的作用。所以，多巴胺能投射通過直接通路或間接通路，最后對(duì)運(yùn)動(dòng)都起易化作用第八十三頁(yè)，共一百一十二頁(yè)。83+-Parkinson’sdiseaseHuntington’sdisease-第八十四頁(yè)，共一百一十二頁(yè)。84Thecerebellum

controlofmovement第八十五頁(yè)，共一百一十二頁(yè)。85Overview小腦(cerebellum)是中樞神經(jīng)系統(tǒng)中最大的運(yùn)動(dòng)結(jié)構(gòu)主要作用是維持軀體平衡、調(diào)節(jié)肌肉張力和協(xié)調(diào)隨意運(yùn)動(dòng)小腦并不直接發(fā)起運(yùn)動(dòng)和指揮肌肉的活動(dòng)，而是作為一個(gè)皮層下的運(yùn)動(dòng)調(diào)