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Hebbiantheory
FromWikipedia,thefreeencyclopedia

Hebbiantheoryisatheoryinneurosciencethatproposesanexplanationfortheadaptationofneuronsinthebrain
duringthelearningprocess.Itdescribesabasicmechanismforsynapticplasticity,whereanincreaseinsynaptic
efficacyarisesfromthepresynapticcell'srepeatedandpersistentstimulationofthepostsynapticcell.Introduced
byDonaldHebbinhis1949bookTheOrganizationofBehavior,[1]thetheoryisalsocalledHebb'srule,Hebb's
postulate,andcellassemblytheory.Hebbstatesitasfollows:
Letusassumethatthepersistenceorrepetitionofareverberatoryactivity(or"trace")tendstoinduce
lastingcellularchangesthataddtoitsstability.WhenanaxonofcellAisnearenoughtoexcitea
cellBandrepeatedlyorpersistentlytakespartinfiringit,somegrowthprocessormetabolicchange
takesplaceinoneorbothcellssuchthatA'sefficiency,asoneofthecellsfiringB,isincreased.[1]
ThetheoryisoftensummarizedbySiegridLwel'sphrase:"Cellsthatfiretogether,wiretogether."[2]However,
thissummaryshouldnotbetakenliterally.HebbemphasizedthatcellAneedsto"takepartinfiring"cellB,and
suchcausalitycanonlyoccurifcellAfiresjustbefore,notatthesametimeas,cellB.Thisimportantaspectof
causationinHebb'sworkforeshadowedwhatisnowknownaboutspiketimingdependentplasticity,which
requirestemporalprecedence.[3]ThetheoryattemptstoexplainassociativeorHebbianlearning,inwhich
simultaneousactivationofcellsleadstopronouncedincreasesinsynapticstrengthbetweenthosecells,and
providesabiologicalbasisforerrorlesslearningmethodsforeducationandmemoryrehabilitation.Inthestudyof
neuralnetworksincognitivefunction,itisoftenregardedastheneuronalbasisofunsupervisedlearning.

Contents
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Hebbianengramsandcellassemblytheory
Principles
Generalizationandstability
Exceptions
Hebbianlearningaccountofmirrorneurons
Seealso
References
Furtherreading
Externallinks

Hebbianengramsandcellassemblytheory
Hebbiantheoryconcernshowneuronsmightconnectthemselvestobecomeengrams.Hebb'stheoriesontheform
andfunctionofcellassembliescanbeunderstoodfromthefollowing:"Thegeneralideaisanoldone,thatanytwo
cellsorsystemsofcellsthatarerepeatedlyactiveatthesametimewilltendtobecome'associated',sothatactivity
inonefacilitatesactivityintheother."[1]:70Healsowrote:"Whenonecellrepeatedlyassistsinfiringanother,the
axonofthefirstcelldevelopssynapticknobs(orenlargesthemiftheyalreadyexist)incontactwiththesomaof
thesecondcell."[1]:63

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GordonAllportpositsadditionalideasregardingcellassemblytheoryanditsroleinformingengrams,alongthe
linesoftheconceptofautoassociation,describedasfollows:
Iftheinputstoasystemcausethesamepatternofactivitytooccurrepeatedly,thesetofactive
elementsconstitutingthatpatternwillbecomeincreasinglystronglyinterassociated.Thatis,each
elementwilltendtoturnoneveryotherelementand(withnegativeweights)toturnofftheelements
thatdonotformpartofthepattern.Toputitanotherway,thepatternasawholewillbecome'auto
associated'.Wemaycallalearned(autoassociated)patternanengram.[4]:44
Hebbiantheoryhasbeentheprimarybasisfortheconventionalviewthat,whenanalyzedfromaholisticlevel,
engramsareneuronalnetsorneuralnetworks.
WorkinthelaboratoryofEricKandelhasprovidedevidencefortheinvolvementofHebbianlearningmechanisms
atsynapsesinthemarinegastropodAplysiacalifornica.
ExperimentsonHebbiansynapsemodificationmechanismsatthecentralnervoussystemsynapsesofvertebrates
aremuchmoredifficulttocontrolthanareexperimentswiththerelativelysimpleperipheralnervoussystem
synapsesstudiedinmarineinvertebrates.Muchoftheworkonlonglastingsynapticchangesbetweenvertebrate
neurons(suchaslongtermpotentiation)involvestheuseofnonphysiologicalexperimentalstimulationofbrain
cells.However,someofthephysiologicallyrelevantsynapsemodificationmechanismsthathavebeenstudiedin
vertebratebrainsdoseemtobeexamplesofHebbianprocesses.Onesuchstudyreviewsresultsfromexperiments
thatindicatethatlonglastingchangesinsynapticstrengthscanbeinducedbyphysiologicallyrelevantsynaptic
activityworkingthroughbothHebbianandnonHebbianmechanisms.

Principles
Fromthepointofviewofartificialneuronsandartificialneuralnetworks,Hebb'sprinciplecanbedescribedasa
methodofdetermininghowtoaltertheweightsbetweenmodelneurons.Theweightbetweentwoneurons
increasesifthetwoneuronsactivatesimultaneously,andreducesiftheyactivateseparately.Nodesthattendtobe
eitherbothpositiveorbothnegativeatthesametimehavestrongpositiveweights,whilethosethattendtobe
oppositehavestrongnegativeweights.
ThefollowingisaformulaicdescriptionofHebbianlearning:(notethatmanyotherdescriptionsarepossible)

where istheweightoftheconnectionfromneuron toneuron and theinputforneuron .Notethatthisis

patternlearning(weightsupdatedaftereverytrainingexample).InaHopfieldnetwork,connections aresetto
zeroif
(noreflexiveconnectionsallowed).Withbinaryneurons(activationseither0or1),connections
wouldbesetto1iftheconnectedneuronshavethesameactivationforapattern.
Anotherformulaicdescriptionis:

where istheweightoftheconnectionfromneuron toneuron , isthenumberoftrainingpatterns,and

the thinputforneuron .Thisislearningbyepoch(weightsupdatedafterallthetrainingexamplesare
presented).Again,inaHopfieldnetwork,connections aresettozeroif
(noreflexiveconnections).
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AvariationofHebbianlearningthattakesintoaccountphenomenasuchasblockingandmanyotherneural
learningphenomenaisthemathematicalmodelofHarryKlopf.Klopf'smodelreproducesagreatmanybiological
phenomena,andisalsosimpletoimplement.

Generalizationandstability
Hebb'sRuleisoftengeneralizedas

orthechangeinthe thsynapticweight isequaltoalearningrate timesthe thinput timesthe

postsynapticresponse .Oftencitedisthecaseofalinearneuron,

andtheprevioussection'ssimplificationtakesboththelearningrateandtheinputweightstobe1.Thisversionof
theruleisclearlyunstable,asinanynetworkwithadominantsignalthesynapticweightswillincreaseordecrease
exponentially.However,itcanbeshownthatforanyneuronmodel,Hebb'sruleisunstable.[5]Therefore,network
modelsofneuronsusuallyemployotherlearningtheoriessuchasBCMtheory,Oja'srule,[6]ortheGeneralized
HebbianAlgorithm.

Exceptions
DespitethecommonuseofHebbianmodelsforlongtermpotentiation,thereexistseveralexceptionstoHebb's
principlesandexamplesthatdemonstratethatsomeaspectsofthetheoryareoversimplified.Oneofthemostwell
documentedoftheseexceptionspertainstohowsynapticmodificationmaynotsimplyoccuronlybetween
activatedneuronsAandB,buttoneighboringneuronsaswell.[7]ThisisduetohowHebbianmodification
dependsonretrogradesignalinginordertomodifythepresynapticneuron.[8]Thecompoundmostcommonly
identifiedasfulfillingthisretrogradetransmitterroleisnitricoxide,which,duetoitshighsolubilityand
diffusibility,oftenexertseffectsonnearbyneurons.[9]Thistypeofdiffusesynapticmodification,knownasvolume
learning,counters,oratleastsupplements,thetraditionalHebbianmodel.[10]

Hebbianlearningaccountofmirrorneurons
Hebbianlearningandspiketimingdependentplasticityhavebeenusedinaninfluentialtheoryofhowmirror
neuronsemerge.[11][12]Mirrorneuronsareneuronsthatfirebothwhenanindividualperformsanactionandwhen
theindividualsees[13]orhears[14]anotherperformasimilaraction.Thediscoveryoftheseneuronshasbeenvery
influentialinexplaininghowindividualsmakesenseoftheactionsofothers,byshowingthat,whenaperson
perceivestheactionsofothers,thepersonactivatesthemotorprogramswhichtheywouldusetoperformsimilar
actions.Theactivationofthesemotorprogramsthenaddsinformationtotheperceptionandhelpspredictwhatthe
personwilldonextbasedontheperceiver'sownmotorprogram.Achallengehasbeentoexplainhowindividuals
cometohaveneuronsthatrespondbothwhileperforminganactionandwhilehearingorseeinganotherperform
similaractions.
ChristianKeysersandDavidPerrettsuggestedthat,whileanindividualperformsaparticularaction,theindividual
willsee,hear,andfeelhimselfperformtheaction.Thesereafferentsensorysignalswilltriggeractivityinneurons
respondingtothesight,sound,andfeeloftheaction.Becausetheactivityofthesesensoryneuronswill
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consistentlyoverlapintimewiththoseofthemotorneuronsthatcausedtheaction,Hebbianlearningwouldpredict
thatthesynapsesconnectingneuronsrespondingtothesight,sound,andfeelofanactionandthoseoftheneurons
triggeringtheactionshouldbepotentiated.Thesameistruewhilepeoplelookatthemselvesinthemirror,hear
themselvesbabble,orareimitatedbyothers.Afterrepeatedexperienceofthisreafference,thesynapses
connectingthesensoryandmotorrepresentationsofanactionwouldbesostrongthatthemotorneuronswould
startfiringtothesoundorthevisionoftheaction,andamirrorneuronwouldhavebeencreated.
Evidenceforthatperspectivecomesfrommanyexperimentsthatshowthatmotorprogramscanbetriggeredby
novelauditoryorvisualstimuliafterrepeatedpairingofthestimuluswiththeexecutionofthemotorprogram(for
areviewoftheevidence,seeGiudiceetal.,2009[15]).Forinstance,peoplewhohaveneverplayedthepianodonot
activatebrainregionsinvolvedinplayingthepianowhenlisteningtopianomusic.Fivehoursofpianolessons,in
whichtheparticipantisexposedtothesoundofthepianoeachtimehepressesakey,sufficestolatertrigger
activityinmotorregionsofthebrainuponlisteningtopianomusic.[16]Consistentwiththefactthatspiketiming
dependentplasticityoccursonlyifthepresynapticneuron'sfiringpredictsthepostsynapticneuron'sfiring,[17]the
linkbetweensensorystimuliandmotorprogramsalsoonlyseemtobepotentiatedifthestimulusiscontingenton
themotorprogram.

Seealso
Dale'sprinciple
Coincidencedetectioninneurobiology
Leabra
Metaplasticity
Tetanicstimulation
Synaptotropichypothesis

References
1.Hebb,D.O.(1949).TheOrganizationofBehavior.NewYork:Wiley&Sons.
2.SiegridLwel,GttingenUniversityTheexactsentenceis:"neuronswiretogetheriftheyfiretogether"(Lwel,S.and
Singer,W.(1992)Science255(publishedJanuary10,1992)"SelectionofIntrinsicHorizontalConnectionsintheVisual
CortexbyCorrelatedNeuronalActivity".UnitedStates:AmericanAssociationfortheAdvancementofScience.pp.209
212.ISSN00368075.
3.CaporaleNDanY(2008)."Spiketimingdependentplasticity:aHebbianlearningrule".AnnualReviewofNeuroscience.
31:2546.doi:10.1146/annurev.neuro.31.060407.125639.PMID18275283.
4.Allport,D.A.(1985)."Distributedmemory,modularsystemsanddysphasia".InNewman,S.K.EpsteinR.(Eds.).
CurrentPerspectivesinDysphasia.Edinburgh:ChurchillLivingstone.ISBN0443030391.
5.Euliano,NeilR.(19991221)."NeuralandAdaptiveSystems:FundamentalsThroughSimulations"(PDF).Neuraland
AdaptiveSystems:FundamentalsThroughSimulations.Wiley.Archivedfromtheoriginal(PDF)on20151225.
Retrieved20160316.
6.Shouval,Harel(20050103)."ThePhysicsoftheBrain".TheSynapticbasisforLearningandMemory:Atheoretical
approach.TheUniversityofTexasHealthScienceCenteratHouston.Archivedfromtheoriginalon20070610.
Retrieved20071114.
7.Horgan,John(May1994)."Neuraleavesdropping".ScientificAmerican.270:16.doi:10.1038/scientificamerican059416.
8.Fitzsimonds,ReikoMuMingPoo(January1998)."RetrogradeSignalingintheDevelopmentandModificationof
Synapses".PsychologicalReviews.
9.Lpez,PC.P.Araujo(2009)."Acomputationalstudyofthediffuseneighbourhoodsinbiologicalandartificialneural
networks"(PDF).InternationalJointConferenceonComputationalIntelligence.
10.Mitchison,GN.Swindale(October1999)."CanHebbianVolumeLearningExplainDiscontinuitiesinCorticalMaps?".
NeuralComputation.11:15191526.doi:10.1162/089976699300016115.
11.KeysersCPerrettDI(2004)."Demystifyingsocialcognition:aHebbianperspective".TrendsinCognitiveSciences.8
(11):501507.doi:10.1016/j.tics.2004.09.005.
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12.Keysers,C.(2011).TheEmpathicBrain.
13.GalleseVFadigaLFogassiLRizzolattiG(1996)."Actionrecognitioninthepremotorcortex".Brain.119(Pt2):593
609.doi:10.1093/brain/119.2.593.PMID8800951.
14.KeysersCKohlerEUmiltaMANanettiLFogassiLGalleseV(2003)."Audiovisualmirrorneuronsandaction
recognition".ExpBrainRes.153(4):628636.doi:10.1007/s0022100316035.PMID12937876.
15.DelGiudiceMManeraVKeysersC(2009)."Programmedtolearn?Theontogenyofmirrorneurons".DevSci.12(2):
350363.doi:10.1111/j.14677687.2008.00783.x.
16.LahavASaltzmanESchlaugG(2007)."Actionrepresentationofsound:audiomotorrecognitionnetworkwhile
listeningtonewlyacquiredactions".JNeurosci.27(2):308314.doi:10.1523/jneurosci.482206.2007.
17.BauerEPLeDouxJENaderK(2001)."FearconditioningandLTPinthelateralamygdalaaresensitivetothesame
stimuluscontingencies".NatNeurosci.4(7):687688.

Furtherreading
Hebb,D.O.(1961)."Distinctivefeaturesoflearninginthehigheranimal".InJ.F.Delafresnaye(Ed.).Brain
MechanismsandLearning.London:OxfordUniversityPress.
Hebb,D.O.(1940)."HumanBehaviorAfterExtensiveBilateralRemovalfromtheFrontalLobes".Archives
ofNeurologyandPsychiatry.44(2):421438.doi:10.1001/archneurpsyc.1940.02280080181011.
Bishop,C.M.(1995).NeuralNetworksforPatternRecognition.Oxford:OxfordUniversityPress.ISBN0
198538499.
Paulsen,O.Sejnowski,T.J.(2000)."Naturalpatternsofactivityandlongtermsynapticplasticity".Current
OpinioninNeurobiology.10(2):172179.doi:10.1016/S09594388(00)000763.PMC2900254 .
PMID10753798.Citeusesdeprecatedparameter|coauthors=(help)

Externallinks
Overview(http://icwww.epfl.ch/~gerstner//SPNM/node71.html)
HebbianLearningtutorial(Part1:NoveltyFiltering(http://blog.peltarion.com/2006/05/11/thetalenteddrhe
bbpart1noveltyfiltering),Part2:PCA(http://blog.peltarion.com/2006/06/20/thetalenteddrhebbpart2p
ca/))
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