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MOTORUNITS MotorFunctionsoftheSpinal Cord andCordReflexes

CharlieE.Labarda,MD asingle motoneuron andthemusclefibers thatitinnervates numberofmusclefibersinnervatedcanvary fromafewfiberstothousandsoffibers, dependingonthenatureofthemotor activity eyemovementsmotoneurons innervate onlyafewmusclefibers;posturalmuscles motoneurons innervatethousandsof musclefibers
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SizePrinciple

ORGANIZATIONOF MOTORFUNCTIONBY THE SPINALCORD

Statesthatasmoremotorunitsare recruited,progressivelylarger motoneurons areinvolvedandgreater tensionwillbegenerated Theforceofcontractionofamuscleis gradedbyrecruitment ofmotorunits E.g.smallvs largemotoneurons

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Smallmotoneurons innervateafewmusclefibers becausetheyhavethelowestthresholds, theyfirefirst alsogeneratethesmallestamountsof force Largemotoneurons innervatemanymusclefibers havethehighestthresholdstofireaction potentials;thus,theyfirelast theyalsogeneratethegreatestamounts offorce
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Motoneuron
innervateextrafusal skeletalmusclefibers Actionpotentialsleadtoactionpotentialsin theextrafusal musclefiberstheyinnervate, whichresultsincontraction

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Motoneurons

Typesof Motoneurons

innervatespecializedintrafusal musclefibers, acomponentofthemusclespindles functionofthemusclespindle:tosense musclelength; thefunctionofthemotoneurons innervatingthemistoadjustthesensitivityof themusclespindles(sotheyrespond appropriatelyastheextrafusal fiberscontract andshorten)
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Motoneurons andmotoneurons are coactivated (activatedsimultaneously)so thatmusclespindlesremainsensitiveto changesinmusclelengthevenasthemuscle contractsandshortens

Extrafusal fibers
constitutethemajorityofskeletalmuscle innervatedbymotoneurons areusedtogenerateforce

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Intrafusal fibers Typesof MuscleFibers

arespecializedfibersthatareinnervatedby motoneurons andaretoosmalltogenerate significantforce encapsulatedinsheaths,formingmuscle spindlesthatrunparalleltotheextrafusal fibers

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Intrafusal MuscleFibers ofMuscleSpindles MUSCLE SPINDLES

twotypesofintrafusal fiberspresentin musclespindles:
nuclearbagfibersarelarger,andtheirnuclei areaccumulatedinacentral("bag")region nuclearchainfiberssmaller,andtheirnucleiare arrangedinrows("chains")

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Generally,bothtypesoffibersarepresentin everymusclespindle,butnuclearchain fibersaremoreplentifulthannuclearbag fibers

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distributedamongtheextrafusal muscle fibers especiallyabundantinmusclesutilizedfor finemovements(e.g.,musclesoftheeye) spindleshapedorganscomposedof intrafusal musclefibersandinnervatedby sensoryandmotornervefibers areattachedtoconnectivetissueand arrangedinparallelwiththeextrafusal musclefibers
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Innervation of MuscleSpindles:both sensory(afferent)and motor(efferent)nerves

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SensoryInnervation
groupIa afferentnerve innervatesboththenuclearbagfibersand thenuclearchainfibers amongthelargestnervesinthebody amongthefastestconductionvelocities formprimaryendingsinaspiralshaped terminalaroundthecentralregionofthe nuclearbagandnuclearchainfibers
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MotorInnervation
consistsoftwotypesofmotoneurons: Dynamic Static Dynamicmotoneurons synapseonnuclear bagfibersin"plateendings." Staticmotoneurons synapseonnuclear chainfibersin"trailendings,"whichspread outoverlongerdistances
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groupIIafferentnerves innervateonlythenuclearchainfibers haveintermediatediametersand intermediateconductionvelocities formsecondaryendingsonthenuclear chainfibers

Motoneurons aresmallerandslowerthan themotoneurons thatinnervatethe extrafusal fibers thefunctionofthemotoneurons (either staticordynamic)istoregulatethe sensitivityoftheintrafusal musclefibersthey innervate

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Events that Occur when a Muscle is Stretched Extrafusal muscle

Muscle is stretched is lengthened Intrafusal muscle also lengthened

Functionof MuscleSpindles

detected by the sensory afferent fibers

Group Ia afferent fibers (nuclear bag and nuclear chain fibers) detect the velocity of length change
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Group II afferent fibers (nuclear chain fibers) detect the length of the muscle fiber

stretchreceptors Function:tocorrectforchangesinmuscle lengthwhenextrafusal musclefibersare eithershortened(bycontraction)or lengthened(bystretch) Thus,musclespindlereflexesoperateto returnmuscletoitsrestinglengthafterithas beenshortenedorlengthened Toillustratethefunctionofthemuscle spindlereflex:considertheeventsthatoccur whenamuscleisstretched
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Activation of Group I afferent fibers

stimulates motoneurons in the spinal cord

Motoneurons are coactivated

innervate extrafusal fibers in the homonymous (same) muscle

ensures that the muscle spindle will remain sensitive to changes in muscle length even during the contraction
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cause the muscle to contract (shorten)

the original stretch (lengthening) is opposed when the reflex causes the muscle to contract and shorten

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Stretch(Myotatic)Reflex SPINALCORDREFLEXES
thesimplestofallspinalcordreflexes havingonlyonesynapsebetweensensory afferentnerves(groupIa afferents)and motorefferentnerves(motoneurons) exemplifiedbythekneejerkreflex

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arestereotypical motorresponsestospecific kindsofstimuli,suchasstretchofthemuscle Theneuronalcircuitthatdirectsthismotor responseiscalledthereflexarc Thereflexarcincludes: thesensoryreceptors; thesensoryafferentnervescarry informationtothespinalcord theinterneurons inthespinalcord Motoneuronsdirectthemuscleto contractorrelax
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The Knee-Jerk Reflex

Tapping the patellar tendon synapse on and activate motoneurons in the spinal cord Quadriceps muscle stretch (muscle spindle stretched)

StepsintheGolgitendonReflex
1. Whenthemusclecontracts,theextrafusal musclefibersshorten,activatingtheGolgi tendonorgansattachedtothem.Inturn, thegroupIb afferentfibersthatsynapseon inhibitoryinterneurons inthespinalcord areactivated.Theseinhibitory interneurons synapseonthe motoneurons
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group Ia afferent fibers are stimulated

innervate and cause contraction of the quadriceps (the muscle that originally 9/8/2011 was stretched

Quadriceps contract and shortens

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Knee jerk

GolgiTendonReflex
adisynaptic spinalcordreflex alsocalledtheinversemyotatic reflex Golgitendonorgan
isastretchreceptorfoundintendons sensescontraction(shortening)ofmuscleand activatesgroupIb afferentnerves arearrangedinserieswiththeextrafusal muscle fibers(contrastingtheparallelarrangementof musclespindlesinthestretchreflex)
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GolgiTendonReflex

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2.Whentheinhibitoryinterneurons are activated(i.e.,activatedtoinhibit),they inhibitfiringofthemotoneurons, producingrelaxationofthehomonymous muscle(themusclethatoriginallywas contracted) 3.Asthehomonymousmusclerelaxes,the reflexalsocausessynergisticmusclestorelax andantagonisticmusclestocontract.

StepsintheFlexorWithdrawal Reflex
1.Whenalimbtouchesapainfulstimulus,flexor reflexafferentfibers(groupsII,III,andIV)are activated.Theseafferentfiberssynapseon multipleinterneurons inthespinalcord(i.e., polysynapticreflex). 2.Ontheipsilateral sideofthepainstimulus, reflexesareactivatedthatcauseflexor musclestocontractandextensormusclesto relax.Thisportionofthereflexproduces flexionontheipsilateral side
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FlexorWithdrawalReflex
apolysynapticreflexthatoccursinresponse toapainfulornoxiousstimulus Somatosensory andpainafferentfibers initiateaflexionreflexthatcauseswithdrawal oftheaffectedpartofthebodyfromthe painfulornoxiousstimulus Thereflexproducesflexion ontheipsilateral side(i.e.,sideofthestimulus)andextension onthecontralateral side
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FlexorWithdrawalReflex

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3.Onthecontralateral sideofthepain stimulus,reflexesareactivatedthatcause extensormusclestocontractandflexor musclestorelax.Thisportionofthereflex producesextensiononthecontralateral side andiscalledthecrossedextensionreflex. Thus,ifthepainfulstimulusoccursonthe leftside,theleftarmandlegwillflexor withdrawandtherightarmandlegwill extendtomaintainbalance.
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Type of Reflex (Example)

Number of Synapses

Stimulus for Reflex Stretch Ia (lengthening) of the muscle Contraction Ib (shortening) of the muscle Pain; temperature

Sensory Afferent Fibers

Responses Contraction of the muscle Relaxation of the muscle

Stretch reflex One (knee jerk) Golgi tendon Two reflex (clasp knife) Flexorwithdrawal reflex (touching a hot stove)
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Many

II, III, and IV Flexion on ipsilateral side; extension on contralateral side School of Health Sciences

4.Apersistentneuraldischarge,calledan afterdischarge, occursinthepolysynaptic reflexcircuits.Asaresultofthe afterdischarge,thecontractedmuscles remaincontractedforaperiodoftimeafter thereflexisactivated.

CONTROLOFPOSTUREAND MOVEMENTBYTHEBRAIN STEM

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Descendingmotorpathways(i.e.,those descendingfromthecerebralcortexand brainstem)aredividedamongthepyramidal tractandtheextrapyramidal tract Pyramidaltracts arecorticospinal and corticobulbar tractsthatpassthroughthe medullary pyramidsanddescenddirectly ontolowermotoneurons inthespinalcord Allothersareextrapyramidal tracts

Thepontine reticulospinal tract originatesinnucleiofthepons andprojects totheventromedial spinalcord Stimulationhasageneralizedactivating effectonboth flexorandextensormuscles, withitspredominanteffectonextensors

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Theextrapyramidal tractsoriginateinthe followingstructuresofthebrainstem:

Therubrospinal tract originatesintherednucleusandprojectsto motoneurons inthelateralspinalcord Stimulationoftherednucleusproduces activationofflexormusclesandinhibitionof extensormuscles

Themedullary reticulospinal tract originatesinthemedullary reticular formationandprojectstomotoneurons in thespinalcord Stimulationhasageneralizedinhibitory effectonboth flexorandextensormuscles, withthepredominanteffectonextensors

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Thelateralvestibulospinal tract originatesinthelateralvestibularnucleus (Deiters'nucleus)andprojectstoipsilateral motoneurons inthespinalcord Stimulationproducesactivationofextensors andinhibitionofflexors

Boththepontine reticularformationandthe lateralvestibularnucleushavepowerful excitatoryeffectsonextensormuscles Therefore,lesionsofthebrainstemabove thepontine reticularformationandlateral vestibularnucleus,butbelow themidbrain, causeadramaticincreaseinextensortone, calleddecerebrate rigidity Lesionsabove themidbraindonot cause decerebrate rigidity
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Thetectospinal tract originatesinthesuperiorcolliculus (tectum or"roof"ofthebrainstem)andprojectsto thecervicalspinalcord Itisinvolvedincontrolofneckmuscles

Decerebrate Rigidity: AbnormalExtensorReflex

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CEREBELLUM

3maindivisionsofthecerebellum: thevestibulocerebellum dominatedby vestibularinputandcontrolsbalanceandeye movements Spinocerebellum dominatedbyspinalcord inputandcontrolssynergyofmovement Pontocerebellum dominatedbycerebral input,viapontine nuclei,andcontrolsthe planningandinitiationofmovements
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"littlebrain,"regulatesmovementandposture andplaysaroleincertainkindsofmotor learning helpscontroltherate,range,force,and directionofmovements(synergy) Damageresultsinlackofcoordination locatedintheposteriorfossa justbelowthe occipitallobe connectedtothebrainstembythreecerebellar peduncles,whichcontainbothafferentand efferentnervefibers
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LayersoftheCerebellar Cortex
Thegranularlayer theinnermostlayer containsgranulecells,GolgiIIcells,and glomeruli Intheglomeruli,axonsofmossyfibersfrom thespinocerebellar andpontocerebellar tracts synapseondendritesofgranuleandGolgi typeIIcells
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LayersoftheCerebellar Cortex
ThePurkinjecelllayer themiddlelayer containsPurkinjecells,anditsoutputis alwaysinhibitory

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Themolecularlayer theoutermostlayer containsouterstellate cells,basketcells, dendritesofPurkinjeandGolgiIIcells,and axonsofgranulecells Theaxonsofgranulecellsformparallel fibers,whichsynapseonthedendritesof Purkinjecells,basketcells,outerstellate cells,andGolgitypeIIcells.

InputtotheCerebellar Cortex
Twosystemsprovideexcitatoryinput tothe cerebellar cortex:
theclimbingfibersystem themossyfibersystem

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Eachsystemalsosendscollateralbranches directlytodeepcerebellar nuclei,inaddition totheirprojectionstothecerebellar cortex Excitatoryprojectionsfromthecerebellar cortexthenactivatesecondarycircuits, whichmodulatetheoutputofthecerebellar nucleiviathePurkinjecells

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Climbingfibers
originateintheinferioroliveofthemedulla andprojectdirectlyontoPurkinjecells makemultiplesynapticconnectionsalong thedendritesofPurkinjecells,althougheach Purkinjecellreceivesinputfromonlyone climbingfiber Thesesynapticconnectionsarevery powerful!
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Mossyfibers
constitutethemajorityofthecerebellar input Thesefibersincludevestibulocerebellar, spinocerebellar,andpontocerebellar afferents Mossyfibersprojecttogranulecells,which areexcitatoryinterneurons locatedin collectionsofsynapsescalledglomeruli Axonsfromthesegranulecellsthenascendto themolecularlayer,wheretheybifurcateand giverisetoparallelfibers
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Asingleactionpotentialfromaclimbing fibercanelicitmultipleexcitatorybursts, calledcomplexspikes, inthedendritesofthe Purkinjecell Itisbelievedthatclimbingfibers"condition" thePurkinjecellsandmodulatetheir responsestomossyfiberinput Climbingfibersalsomayplayarolein cerebellar learning.

Parallelfibers fromthegranulecellscontactthe dendritesofmany Purkinjecells,producinga "beam"ofexcitationalongtherowofPurkinjecells Thedendritic treeofeachPurkinjecellmayreceive inputfromasmanyas250,000parallelfibers! Incontrasttotheclimbingfiberinputtothe Purkinjedendrites(whichproducecomplexspikes), themossyfiberinputproducessingleaction potentialscalledsimplespikes Theseparallelfibersalsosynapseoncerebellar interneurons (basket,stellate,andGolgiII).
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Interneurons oftheCerebellum
Thefunctionofcerebellar interneurons isto modulate Purkinjecelloutput Withtheexceptionofgranulecells,allofthe cerebellar interneurons areinhibitory Granulecellshaveexcitatoryinputtobasketcells, stellate cells,GolgiIIcells,andPurkinjecells Basketcellsandstellate cellsinhibitPurkinjecells (viaparallelfibers) GolgiIIcellsinhibitgranulecells,therebyreducing theirexcitatoryeffectonPurkinjecells
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