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ExcitableCells

ExcitableCells

Indextothispage
TheRestingPotential
IonicRelationsintheCell
Depolarization
ActionPotentials
MyelinatedNeurons
Hyperpolarization
IntegratingSignals

Excitablecellsarethosethatcanbestimulatedtocreateatinyelectriccurrent.
Musclefibersand
nervecells(neurons)areexcitable.
Thecolorphoto(courtesyofJulieH.SandellandRichardH.Masland)isofasingleinterneuronin
theretinaofarabbit.Thecellhasbeeninjectedwithafluorescentdyetorevealallitsbranches.Each
ofthesmallknobsatthetipsofthebranchesmakesasynapsewithanothercellintheretina.
Theelectriccurrent
inneuronsisusedtorapidlytransmitsignalsthroughtheanimal.
inmusclesisusedtoinitiatecontraction.[Link]

TheRestingPotential
Allcells(notjustexcitablecells)havearestingpotential:anelectricalchargeacrosstheplasma
membrane,withtheinteriorofthecellnegativewithrespecttotheexterior.Thesizeoftheresting
potentialvaries,butinexcitablecellsrunsabout70millivolts(mv).
Therestingpotentialarisesfromtwoactivities:
Thesodium/potassiumATPase.Thispumppushesonlytwopotassiumions(K+ )intothecell
foreverythreesodiumions(Na+ )itpumpsoutofthecellsoitsactivityresultsinanetlossof
positivechargeswithinthecell.
Somepotassiumchannelsintheplasmamembraneare"leaky"allowingaslowfacilitated
diffusionofK+ outofthecell(redarrow).

IonicRelationsintheCell
Thesodium/potassiumATPaseproduces
aconcentrationofNa+ outsidethecellthatissome
10timesgreaterthanthatinsidethecell
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aconcentrationofK+ insidethecellsome20times
greaterthanthatoutsidethecell.
Theconcentrationsofchlorideions(Cl )andcalciumions
(Ca2+ )arealsomaintainedatgreaterlevelsoutsidethecell
EXCEPTthatsomeintracellularmembraneenclosed
compartmentsmayalsohavehighconcentrationsofCa2+
(greenoval)

Depolarization
Certainexternalstimulireducethechargeacrosstheplasmamembrane.
mechanicalstimuli(e.g.,stretching,soundwaves)activatemechanicallygatedsodiumchannels
certainneurotransmitters(e.g.,acetylcholine)openligandgatedsodiumchannels.
Ineachcase,thefacilitateddiffusionofsodiumintothecellreducestherestingpotentialatthatspot
onthecellcreatinganexcitatorypostsynapticpotentialorEPSP.
Ifthepotentialisreducedtothethresholdvoltage(about50mvinmammalianneurons),anaction
potentialisgeneratedinthecell.

ActionPotentials
Ifdepolarizationata
spotonthecell
reachesthe
thresholdvoltage,
thereducedvoltage
nowopensup
hundredsofvoltage
gatedsodium
channelsinthat
portionofthe
plasmamembrane.
Duringthe
millisecondthatthe
channelsremain
open,some7000
Na+ rushintothe
cell.Thesudden
complete
depolarizationofthe
membraneopensup
moreofthevoltage
gatedsodium
channelsinadjacent
portionsofthe

Thenerveimpulse.Intherestingneuron,theinterioroftheaxonmembraneis
negativelychargedwithrespecttotheexterior(A).Astheactionpotential
passes(B),thepolarityisreversed.ThentheoutflowofK+ ionsquickly
restoresnormalpolarity(C).Attheinstantpicturedinthediagram,themoving
spot,whichhastracedthesechangesontheoscilloscopeastheimpulseswept
pasttheintracellularelectrode,isatpositionC.

membrane.Inthisway,awaveofdepolarizationsweepsalongthecell.Thisistheactionpotential.(In2/5

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membrane.Inthisway,awaveofdepolarizationsweepsalongthecell.Thisistheactionpotential.(In
neurons,theactionpotentialisalsocalledthenerveimpulse.)

Therefractoryperiod
Asecondstimulusappliedtoaneuron(ormusclefiber)lessthan0.001secondafterthefirstwillnot
triggeranotherimpulse.Themembraneisdepolarized(positionB),andtheneuronisinitsrefractory
period.Notuntilthe70mvpolarityisreestablished(positionC)willtheneuronbereadytofire
again.
Repolarizationisfirstestablishedbythefacilitateddiffusionofpotassiumionsoutofthecell.Only
whentheneuronisfinallyrestedarethesodiumionsthatcameinateachimpulseactivelytransported
backoutofthecell.
Insomehumanneurons,therefractoryperiodlastsonly0.0010.002second.Thismeansthatthe
neuroncantransmit5001000impulsespersecond.

Theactionpotentialisallornone
Thestrengthoftheactionpotentialisanintrinsicpropertyofthecell.Solongastheycanreachthe
thresholdofthecell,strongstimuliproducenostrongeractionpotentialsthanweakones.However,
thestrengthofthestimulusisencodedinthefrequencyoftheactionpotentialsthatitgenerates.

MyelinatedNeurons
Theaxonsofmanyneuronsareencasedinafattysheathcalledthemyelinsheath.Itisthegreatly
expandedplasmamembraneofanaccessorycellcalledtheSchwanncell.Wherethesheathofone
Schwanncellmeetsthenext,theaxonisunprotected.Thevoltagegatedsodiumchannelsof
myelinatedneuronsareconfinedtothesespots(callednodesofRanvier).

Theinrushofsodiumionsatonenodecreatesjustenoughdepolarizationtoreachthethresholdofthe
next.Inthisway,theactionpotentialjumpsfromonenodetothenext.Thisresultsinmuchfaster
propagationofthenerveimpulsethanispossibleinnonmyelinatedneurons.

Multiplesclerosis
Thisautoimmunedisorderresultsinthegradualdestructionofmyelinsheaths.Despitethis,
transmissionofnerveimpulsescontinuesforaperiodasthecellinsertsadditionalvoltagegated
sodiumchannelsinportionsofthemembraneformerlyprotectedbymyelin.

Hyperpolarization
Despitetheirname,someneurotransmittersinhibitthetransmissionofnerveimpulses.Theydothis
byopening
chloridechannelsand/or
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potassiumchannelsintheplasmamembrane.
Ineachcase,openingofthechannelsincreasesthemembranepotentialby
lettingnegativelychargedchlorideions(Cl )INand
positivelychargedpotassiumions(K+ )OUT
Thishyperpolarizationiscalledaninhibitorypostsynapticpotential(IPSP)becauseitcounteracts
anyexcitatorysignalsthatmayarriveatthatneuron.Althoughthethresholdvoltageofthecellis
unchanged,itnowrequiresastrongerexcitatorystimulustoreachthreshold.
Example:Gammaaminobutyricacid(GABA).Thisneurotransmitterisfoundinthebrainand
inhibitsnervetransmissionbybothmechanisms:
bindingtoGABAAreceptorsopenschloridechannelsintheneuron.
bindingtoGABABreceptorsopenspotassiumchannels.
More

IntegratingSignals
Asingleneuron,especiallyoneinthe
centralnervoussystem(seecolorphotoat
top),mayhavethousandsofotherneurons
synapsingonit.Someoftheserelease
activating(depolarizing)
neurotransmittersothersreleaseinhibitory
(hyperpolarizing)neurotransmitters.
Thereceivingcellisabletointegratethese
signals.Thediagramshowshowthis
worksinamotorneuron.
1.TheEPSPcreatedbyasingle
excitatorysynapseisinsufficientto
reachthethresholdoftheneuron.
2.EPSPscreatedinquicksuccession,
however,addtogether
("summation").Iftheyreach
threshold,anactionpotentialis
generated.
3.TheEPSPscreatedbyseparateexcitatorysynapses(A+B)canalsobeaddedtogethertoreach
threshold.
4.Activationofinhibitorysynapses(C)makestherestingpotentialoftheneuronmorenegative.
TheresultingIPSPmayalsopreventwhatwouldotherwisehavebeeneffectiveEPSPsfrom
triggeringanactionpotential.
Normally,thenumberofEPSPsneededtoreachthresholdisgreaterthanshownhere.
Onemightexpectthatdepolarizationatonepointontheplasmamembranewouldgenerateanaction
potentialirrespectiveofinhibitorysignalselsewhere.However,thisisavoidedinmanyneuronsbythe
axonhillockandtheaxoninitialsegment(theAIS).Thisistheregionwheretheaxonemergesfrom
thecellbodyandisunmyelinated.Theportionoftheplasmamembraneinthisregionhas
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fewornosynapsesofitsownand
alowerthresholdthanelsewhereonthecell.
[Neuronscanestablishsuchdistinctivedomainsontheirplasma
membranebyanchoring(withactinfilaments)transmembrane
proteinsasbarrierstoblockthefreediffusionofmembrane
proteinsfromthecellbodytotheaxon.]
Theactionpotentialisusuallygeneratedintheaxoninitial
segment.Havingneitherexcitatorynorinhibitorysynapsesofits
own,itisabletoevaluatethetotalpictureofEPSPsandIPSPs
createdinthedendritesandcellbody.
Onlyif,overabriefinterval,thesumofdepolarizingsignals
minusthesumofthehyperpolarizingsignalsexceedsthe
thresholdoftheaxoninitialsegmentwillanactionpotentialbe
generated.
Thismethodfortheneurontoevaluateamixofpositiveandnegativesignalsoccursrapidly.Itturns
out,however,thatneuronsalsohavealongtermwaytointegrateamixofpositiveandnegative
signalsconvergingonthem.Thislongtermresponseinvolveschangesingeneactivityleadingto
changesinthenumberandactivityofthecell'smanysynapses.
Linktodiscussionsoflongtermfacilitationandlongtermdepression.
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