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HarmonicResonanceTheory:AnAlternativeto

the"NeuronDoctrine"Paradigmof
NeurocomputationtoAddressGestalt
propertiesofperception
StevenLehar
slehar@cns.bu.edu
SubmittedtoPsychologicalReviewJuly1999
RejectedNovember1999
SubmittedtoBehavioral&BrainSciencesSeptember2000
Notacceptedforreview
becauseapparentlyonlyonepaperisallowedperauthoratatime!
ResubmittedtoBehavioral&BrainSciencesMarch2004
NotacceptedforreviewMarch2004
Author'sFormalComplaintMarch2004
PaulBloomRespondsMarch2004
Author'sLastWord!March2004
SubmittedtoJournalofIntegrativeNeuroscienceAugust2004
Revise&ResubmitApril2005
Author'sResponseMay2005
StephenKercelRespondsMay2005

ShortAbstract
Theconventionalviewofneuroscience,knownastheneurondoctrine,isbasedontheassumption
thatneurocomputationinvolvesdiscretesignalscommunicatedalongfixedtransmissionlinesbetween
discretecomputationalelements.Thisconceptisshowntobeinadequatetoaccountforinvariancein
recognition,aswellasfortheholisticglobalaspectsofperceptionidentifiedbyGestalttheory.A
HarmonicResonancetheoryispresentedasanalternativeparadigmofneurocomputation,that
exhibitsboththepropertyofinvariance,andtheemergentGestaltpropertiesofperception,notas
specialmechanismscontrivedtoachievethoseproperties,butasnaturalpropertiesoftheresonance
itself.

LongAbstract
Theconventionalviewofneuroscience,knownastheneurondoctrine,isbasedontheassumption
thatneurocomputationinvolvesdiscretesignalscommunicatedalongfixedtransmissionlinesbetween
discretecomputationalelements.Thisatomisticconceptisshowntobeinadequatetoaccountfor
invarianceinrecognition,andperceptualcompletionphenomena.Itisalsoinconsistentwiththe
holisticglobalaspectsofperceptionidentifiedbyGestalttheory.Aharmonicresonancetheoryis
presentedasanalternativeparadigmofneurocomputation,inwhichelectrochemicalstandingwaves
intheneuralsubstrateareproposedastheprinciplepatternformationmechanisminthebrain,to
replacethetemplatelikeconceptofthespatialreceptivefield.Theprincipleofharmonicresonance
exhibitsboththepropertyofinvariance,andtheemergentGestaltpropertiesofperception,notas
specialmechanismscontrivedtoachievethoseproperties,butasnaturalpropertiesoftheresonance
itself.

1Introduction
Neuroscienceisinastateofseriouscrisis.Foralthoughourknowledgeoftheneurophysiologicaland
biomolecularpropertiesofthebraincontinuestoadvancebyleapsandbounds,therehasbeennoreal
progressinourunderstandingofthemostfundamentalquestionsofinformationrepresentationand
processinginthebrain.Infactwehavenoideawhatthecorrectlevelofdescriptionshouldbeto
capturetheessentialaspectsofneurocomputation,becausethereisnouniversallyacceptedtheoryof
howthebraincodesinformation,aproblemthatSearle(1997,p.198)hascalled"thedirtysecretof
contemporaryneuroscience".NeurosciencethereforeremainsinwhatKuhncalledapreparadigmatic
state,withnorealconsensusonthefoundationalassumptionsorelementalprinciplesofthescience.
Curiouslythereisalwaysagreatdealofresistanceinneurosciencetodiscussionofalternative
paradigmatichypotheses.Thisconservativetendencystemsfromageneralmisunderstandingofthe
roleoftheoriesasopposedtoparadigmsinscientificinvestigation.Forwhiletheoriesareacceptedor
rejectedusingthewellestablishedproceduresofscientificevidence,debatesoveralternative
paradigmsrequireamoregeneralhandling,asdiscussedbyKuhn(1970).Thereasonforthisisthat
theparadigmrepresentsthephilosophicalinfrastructure,orsetofinitialassumptionsuponwhich
theoriesarebuilt.Thereforedebatesbetweencompetingparadigmscannotberesolvedbythenormal
rulesofevidence,becausetheinterpretationoftheevidenceitselfdependsonone'sparadigmatic
assumptions(Kuhn1970).Furthermore,theolderparadigmwhichischallengedbythenew
hypothesisisitselfmerelyahypothesiswhichwasneverprovenbeyondareasonabledoubt,and
thereforethenewproposalshouldnotbeheldtoahigherstandardofproofmerelybecauseitis
proposedatalaterdate.Incontemporaryneurosciencethisdefaultconceptofneurocomputationisa
setofassumptionswhichhascometobeknownastheneurondoctrine(Barlow1972,1995,Shepard
1991).Whiletheneurondoctrineisbynomeansuniversallyaccepted,andalternativeparadigmshave
beenproposed,itremainsneverthelessthedominantparadigminthesensethatauthorswhoadoptits
tenetsasinitialassumptionsarenotchallengedtojustifythatchoice.Howevertheonlyreasonthe
neurondoctrinehasachievedthisstatusismoreamatterofhistoricalprecedentandabsenceofviable
alternatives,ratherthanduetoanyintrinsicmeritsofitsown.Althoughtheneurondoctrineserves
adequatelyasamodeloftheindividualneuron,thisparadigmismuchmoreproblematicasageneral
theoryofneurocomputationandrepresentationinthebrain.
InthispaperImakethecaseforanalternativeparadigmintheformofaharmonicresonancetheory,
inwhichelectrochemicalstandingwavesintheneuralsubstrateareproposedasthemechanismthat
encodesspatialpatternsinthebrain.WhatIproposeisnotaspecifictheoryofdetailed
neurophysiologicalfunction,butaparadigmintheformofanovelprincipleofcomputationand
representation.Whetherornotthebrainactuallymakesuseofthisprincipleisasubjectforfuture
experimentalinvestigation,andforreexaminationofexistingdatainthelightofthenewhypothesis.
AlthoughIdonotpresentincontrovertibleevidencetoprovethatharmonicresonanceisemployedin
thebrain,thereisconsiderablesupportiveevidencefromawidevarietyofdiversesources.For
exampleharmonicresonanceoffersanexplanationfortheglobalsynchronyobservedin
electroencephalogram(EEG)recordings,aswellasanalternativeinterpretationofthesynchronous
firingobservedinretinalandcorticalneurons,bothofwhichcanbeseenasdirectevidencefor
resonancephenomenainthebrain.Harmonicresonancealsooffersanexplanationforavarietyof
symmetriesandperiodicitiesobservedinmanyaspectsofperceptionandbehavior,propertieswhich
alsohappentobecharacteristicofstandingwaves.Buttheprincipalfocusofthepresentpaperisnot
somuchonthestrengthofthatcorroborativeevidenceasaproofofharmonicresonanceinthebrain,
butratheronthepropertiesofharmonicresonanceasapossibleprincipleofcomputationand
representationinthebrain.Forharmonicresonanceexhibitscertainuniquepropertiesthatarequite
unlikeanyotherproposedprincipleofcomputationeitherbiologicalorartificial.Inparticular,
harmonicresonanceexhibitstheholisticGestaltcharacterobservedinmanyperceptualphenomena,
whichcannotbemeaningfullyexpressedinanequivalentTuringmachinedescription.Harmonic
resonancealsooffersaspatialrepresentationschemethatexhibitsmanyoftheinvariancesobservedin
perception,inamannerthatisuniquetoharmonicresonance,andwhichareverydifficulttoaccount

forinconventionalneuralnetworkterms.Finally,harmonicresonanceofadifferentformhasalready
beenidentifiedinbiologicalsystemsinthefieldofembryologicalmorphogenesis,wheretheprinciple
ofreactiondiffusionhasbeenidentifiedasthesystemthatdefinesthespatialstructureofthe
developingembryobywayofchemicalstandingwavepatterns.Thefactthatnatureemploysa
standingwaverepresentationinthisotherunrelatedbiologicalfunctionoffersanexistenceproofthat
harmonicresonancebothcananddoesserveasaspatialrepresentationinbiologicalsystems,andthat
representationhappenstoexhibitsthesameholisticGestaltpropertiesthathavebeenidentifiedas
prominentpropertiesofperceptionandbehavior.

2TheNeuronDoctrine
EversinceRamonyCajaldemonstratedthatthenervoussystemiscomposedofdiscretecellsrather
thanacontinuousnetwork,theneurondoctrine(Barlow1972,1995,Shepard1991)hasemergedas
thedominantparadigmofneurocomputation.Therapidpropagationofionsthroughtheintracellular
fluid,incombinationwiththerelativelyslowtransmissionacrossthechemicalsynapsesuggestthat
theneuronbehavesasaquasiindependentprocessorthatrespondstotheinputsignalsreceived
throughitsdendritestoproduceanoutputsignalthroughtheaxonanditscollaterals.Hodgkin&
Huxley(1939)demonstratedthatthefrequencyofactionpotentialscorrelateswiththeinputappliedto
thecellpresynaptically.Thisinturnsuggestsaratecode,inwhichthesignificantsignalisnotcarried
bytheindividualactionpotentials,ashadbeenpreviouslyassumed,butbythefrequencyoftheir
occurrence.
2.1FeatureDetectorHierarchy
NeuroscienceexperiencedsomethingofarevolutionwiththediscoverybyHubelandWieselofcells
inthevisualcortexthatrespondtoparticularfeaturespresentedatspecificlocationsinthevisual
field.Somecellswerefoundtorespondtosimplefeaturessuchasalocaledgeofaparticularlocation
andorientation,whileothercellshadmorecomplexresponsefunctions,asifinresponsetospatial
combinationsofsimplecellresponses.Eventuallywholehierarchiesofcellswereidentifiedinvarious
regionsofthevisualcortex,withcellsinthehighercorticalareasrespondingtoevermorecomplex
combinationsoflowerlevelprimitives.Thisdatasuggeststhatthecortexhasahierarchical
organizationthatencodesthepresenceofparticularpatternsinthevisualfieldbytheactivationof
lowerandhigherordercellstunedspecificallytothosepatterns.
Thecomputationalprinciplebehindthesecorticalfeaturedetectorcellshasalsobeenproposed.Hubel
(1988)suggeststhatlowerlevelfeaturedetectorsaretriggeredbyvisualedgesbythesameessential
principleasthatusedinedgedetectorsemployedincomputerimageprocessing,i.e.eachcellis
equippedwithareceptivefieldwhosespatialpatternofexcitatoryandinhibitorysynapsesmatchthe
spatialfeaturethatthecellistunedtodetect.Inessencetheselowerlevelfeaturedetectorsperforma
localtemplatematchtothepatternofactivationdetectedintheirinputfield,withthepatternof
synapsesinthedendriticfieldactingasthespatialtemplate.Higherordercellsrespondtomore
complexcombinationsoflowerlevelfeaturesbybeingconnectedtothecorrespondinglowerorder
featurecellsbywayoftheappropriatepatternofexcitatoryandinhibitoryconnections.
Atthehighestlevelsofthecortexinformationispresumedtobeencodedbyamassively
interconnectednetworkofcells,eachcellrepresentingacomplexcombinationoflowerlevelfeatural
primitives(Barlow1972).Thehighercorticalneuronseachrepresentsomeaspectofthecomplex
perceptualandcognitiveexperience,andconversely,anyparticularexperienceisrepresentedbya
characteristicconstellationofinnumerableactivationsofhighercorticalneurons.
2.2AbstractionandCompression
Oneessentialaspectofthisparadigmofrepresentationisthatextendedelementsofperceptual

experience,suchastheperceptionofwholeobjectsinascene,areencodedinacompressedmannerin
highercorticalcentersbytheactivationofasinglecellorsmallsetofcellsdedicatedindividuallyor
collectivelytotherepresentationofthoseobjects.Thenumberofcellsrequiredtoencodethatobject
atthehigherlevelthereforeismuchsmallerthanthenumberoflowerlevelcellsthatencodethe
object'scomponentfeatures.Inotherwordsinformationisprogressivelyreducedorabstractedasit
progressesupthroughthecorticalhierarchy,fromthemoreexplicitprimaryareastothemoreabstract
associationareas.Thereisasequentialprogressionimpliedinthisparadigmofrepresentation,with
informationflowingbottomupfromprimarytohigherareas,althoughreciprocalfeedbackpathways
implicatesomekindoftopdownprocess,presumablyforthepurposesofcognitiveexpectationand
perceptualcompletion.
2.3FragmentedArchitecture
Anotheressentialaspectoftheneurondoctrineisitsfragmentaryordistributednature.Foralthough
theprimarycorticalareasrevealtopographicalmapsofthesensoryworld,asintheprimaryvisualand
somatosensorycortices,highercorticalareasarefragmentedintomultiplecorticalmapsofthosesame
sensoryareas,asinthesecondaryvisualandsomatosensoryareas.Eachofthesemultiplecopiesof
thesamesensoryfieldappearstobespecializedfortherepresentationofparticularaspectsor
modalitiesofthatsensoryexperience,suchascolor,shape,motion,andbinoculardisparity.Even
withineachofthesemaps,thefeaturesarerepresentedinfragmentaryform,withseparatecells
dedicatedtoencodingfeaturesofdifferentorientations,shape,binoculardisparities,color,directions
ofmotion,etc.ateverylocationwithinthatvisualmap.Thisissuggestiveofananalytical
representationalstrategyinwhichthesensoryworldisbrokendownintoitscomponentfeatures,each
ofwhichisrepresentedbydistinctcorticalmechanismstunedtodetectthosefeatures.
Theneurondoctrineisbynomeansuniversallyacceptedinneuroscience.Alternativeparadigmshave
beenproposed,suchasKhler&Held'selectricfieldtheory(Khler&Held1947),Pribram's
holographictheory(Barrett1969,Pribrametal.1974,Pribram1999),DeValois&DeValois'theory
ofFouriercoding(DeValois&DeValois1979,1988),vonderMalsburg'stemporalcorrelation
hypothesis(vonderMalsburg&Schneider1986,vonderMalsburg1987),Penrose'stheoryof
quantumconsciousness(Penrose1989,1994),Harrison'sandSmythies'theoryofconsciousnessin
hiddendimensions(Harrison1989,Smythies&Beloff1989,Smythies1994),tonameafew.
Howevernoneoftheseparadigmshaseverbeenworkedoutinenoughdetailtospecifyexactlyhow
perceptualinformationisencodedorprocessedinthebrain.Bycontrast,theneurondoctrinehasthe
distinctmeritofbeingclearandexplicitlydefined,andthereforeamenabletoquantitativecomputer
simulations.Thereforeintheabsenceofamoreviablealternative,theneurondoctrineremainstothis
daythedominantparadigmofneurocomputation,andmuchofthecontemporaryresearch,and
discussionoftheresultsofthatresearch,isbasedimplicitlyorexplicitlyontheassumptionsofthis
paradigm.Theneurondoctrineisalsofairlyconsistentwithcontemporaryunderstandingofthe
neuronatthecellularlevel.Howeverthisconceptofneurocomputationhassomeserious
shortcomingsthatcometolightwhenconsideringalargersystemslevelofanalysisofbrainfunction.

3ProblemswiththeNeuronDoctrine
Inthefirstplacethereisaproblemwiththenotionofthevisualneuronasafeaturedetectorthat
operatesbywayofahardwiredreceptivefieldofexcitatoryandinhibitorysynapsesanchoredtothe
tissueofthebrain.Forthisconceptisnodifferentthanatemplatetheory,thelimitationsofwhichare
wellknown.Atemplateisaspatialmapofthepatterntobematched,whichisinherentlyintolerantto
anyvariationinthestimuluspattern.Forexampleamismatchwillberecordedifthepatternis
presentedatadifferentlocation,orientation,orspatialscalethanthatencodedinthetemplate.
Thesolutiontotheproblemofinvariancecommonlyproposedinneuralmodelingisafeaturebased
approach,i.e.tobreakthepatternintoitscomponentfeatures,anddetectthoselocalfeatures

independentlyofthewhole(Selfridge1959,Marr1982,Biederman1987).Verysimplefeaturessuch
asorientededges,bars,orcorners,aresufficientlyelementalthatitwouldnotbeprohibitiveto
providetemplatesforthemateverylocationandorientationacrossthevisualfield.Inthepurestform
ofthisconcept,thespatialmatchrepresentedbythesingleglobaltemplateisreplacedbyan
enumerativematchthattalliesthenumberandtypeoflocalfeaturespresentinsomeregionofthe
visualfield,andmatchesthislistagainstthelistoffeaturescharacteristicoftheglobalform.For
exampleasquaremightbedefinedbythepresenceoffourcorners,eachofwhichmightbedetected
byalocalcornerdetectorappliedateverylocationthroughoutalocalregionoftheimage.The
enumerativelistingoffourcornerfeatureswouldbethesameforsquaresofdifferentrotations,
translations,andscales,andthereforethefeaturelistasarepresentationisinvarianttorotation,
translation,andscale.
Despitethecurrentpopularityofthefeaturedetectorconceptinneuralnetworkmodels,the
fundamentallimitationsofthisapproachtoperceptionwerepointedoutdecadesagobyGestalt
theory.Inthefirstplace,localfeaturescannotbereliablyidentifiedintheabsenceoftheglobal
context.Forexampleacornerdetectorincomputersimulationswilltypicallygeneratecountless
cornerresponsesinanaturalscene,onlyasmallfractionofwhichwouldbeidentifiedaslegitimate
cornerfeaturesintheglobalcontext.Anotherproblemwiththefeaturebasedapproachisthatinthe
tallyofdetectedfeatures,itisimpossibletodeterminereliablywhichfeaturesbelongtowhich
objects.Whateverlocalregionisselectedforthetallyofdetectedfeatures,mightjustaswellinclude
featuresfromseveraldifferentobjectstoconfoundthefeaturelist,andconversely,theobjectcentered
onthatregionwilloftenextendoutbeyondtheregion,andtherebylosecriticalfeaturesfromits
featurelist.Apurefeaturebasedsystemwouldalsobeeasilymisledbyspatialocclusions,which
occurcommonlyinvisualscenes,butappeartoposenoseriousproblemtovisualrecognition.
Hybridsolutionshavealsobeenproposedinwhichtheobjecttemplateisdefinedasapatternof
regions,eachofwhichrepresentsanapproximatelocusforaparticularfeature(Selfridge1959,
Biederman1987).Forexampleasquaremightbedefinedasfourcircularregionsaroundacenter,
eachofwhichdefinesthepossiblerangeofacornerfeatureatthatpoint,whichwouldbesearchedout
bycornerdetectorsappliedthrougharangeoforientationsthroughouteachofthoseregions.The
positionalandorientationaltoleranceaffordedbythisschemeallowsamultitudeofdifferent
variationsofasquaretostimulatethesamesquaretemplate.Whiletheobjecttemplateisthus
renderedsomewhatrotationandscalesensitive,thetoleranceallowedinitscomponentfeatures
permitsasmallernumberofobjecttemplatesthanwouldberequiredinasimpletemplatemodelto
recognizeallpossiblevariationsofthesquare.
Howeverthehybridschemeisalsofundamentallyflawed,becausethefeaturebaseddetectionofthe
individualcomponentfeaturessuffersthesameproblemsasapurefeaturebasedscheme,beingeasily
confusedbyextraneousfeaturesineachregionofinterest,whilethetemplatelikedetectionofthe
globalconfigurationofthoselocalfeaturessuffersalltheproblemsinherentinatemplatebased
scheme,withhardlimitstotherangeofvariabilityofeachcomponentfeature.Amorefundamental
problemwiththisconceptisthattherangeoflegitimatelocationsandorientationsforanycomponent
featurecannotbedefinedintheabstract,butonlyrelativetotheotherfeaturesactuallypresent.For
exampleifonecornerofasquareisdetected,theexactlocationandorientationofthatcorner
constrainsthepermissiblelocationsoftheotherthreecornersmuchmorepreciselythanwouldbe
encodedintheobjecttemplate.Thereforetherearemanypossibleconfigurationsofcornersthat
wouldregistertothehybridmodelasasquare,onlyasmallfractionofwhichwouldcorrespondto
legitimatesquares.
Thefeaturebasedapproachtovisualrecognitioncanbeimplementedrelativelyeasilyincomputer
algorithms(Ballard&Brown1982,Marr1982).Howeverdespitedecadesofthemostintensive
research,noalgorithmhaseverbeendevisedthatcanperformreliablyexceptinthemostcontrolled
visualenvironments.Theproblemwithboththefeaturebasedmodelandwiththehybridmodelisthat
theyconfuseinvariancetostimulusvariationwithablindnesstothosevariations.Forthehybrid

squaredetectorthatrespondstoasquareknowslittleabouttheexactconfigurationofthecornersof
thatparticularsquare,andtheenumerativefeaturedetectorknowsevenless.Thisisincontrasttoour
subjectiveexperienceinwhichtheregionofthevisualfieldthatisrecognizedasbelongingtoa
squareisperceivedtoashigharesolutionastheedgesofthesquareitself,evenwhenthoseedgesare
notactuallypresentinthestimulus,liketheillusorysidesofaKanizsafigure.Furthermore,wecan
easilyindicatewhereanoccludedormissingcornerofasquareortriangleoughttobelocated,based
ontheconfigurationoftherestofthefigure.
Theproblemsinherentintemplateandfeaturebaseddetectionapplynotonlytoinvarianceinthe
perceptionofsimpleobjectsandtheircomponentfeatures,buttothewholeconceptofafeatural
hierarchy,extendinguptohigherordercomplexobjectsorconcepts.Fortheprincipleofinvariance
impliesamanytoonerelationbetweenthemanypossiblestimulusvariationsthatallindicatetheone
recognizedobject.Whatisrequiredisakindoftopdowncompletionthatmakesuseofthehigher
levelrecognitionoftheobjecttodeterminewhatitsexpectedcomponentpartsshouldbe.Butthis
feedbackiscomplicatedbythemanytoonerelationinthebottomupdirection,becauseasimplistic
topdownfeedbackfromtheinvariantrecognitionnodewouldinvolveaonetomanyrelationto
activateeverypossiblecombinationoflocalfeaturenodesthatcanevertriggerthatinvariantnode.If
ontheotherhandthetopdownfeedbackisonlydirectedtofeaturenodeswhichhaveactually
detectedsomefeature,thiswouldprecludetheperceptualfillinginoffeaturesabsentfromthe
stimulus,andtherebydefeatthewholepurposeofthefeedback.

4AdaptiveResonanceTheory
Althoughtheideaofvisualprocessingasafeedforwardprogressionthroughahierarchical
architecturerepresentsthemostdirectorsimplisticformoftheneurondoctrine,therehasbeena
growingawarenessoftheneedforsomekindofcomplementarytopdownprocessingfunction,both
onperceptualgroundstoaccountforexpectationandperceptualcompletionasseenintheKanizsa
figure,andonneurophysiologicalgroundstoaccountforthereciprocalfeedbackpathwaysidentified
neurophysiologically,runningfromhighertolowercorticalareas.Severaltheoristshaveproposed
neuralnetworkmodelstogreaterorlesserdegreeofcomputationalspecificitythatincorporatesome
kindoffeedbackfunction(Fukushima1987,Carpenter&Grossberg1987,Grossberg&Mingolla
1985,1987,Damasio1989).Unfortunatelythesemodelshavebeenpersistentlyhandicappedbythe
templatelikeconceptoftheneuralreceptivefieldinherentintheneurondoctrine,whichmakesit
impossibleforthemtoprovideanadequateaccountofthejointpropertiesofinvariancein
recognition,andspecificityincompletionphenomena.
PerhapsthemostexplicitmodelofneuralfeedbackisseenintheAdaptiveResonanceTheory(ART,
Carpenter&Grossberg1987).TheprincipalfocusofARTisonthemannerinwhichaneuralnetwork
modeldetectsnoveltyinastreamofinputpatterns,andusesthatinformationtocategorizetheinput
patternsonthebasisofnovelty.Thesignificantpropertyofthismodelinthepresentcontextisnotin
thedetailsofitslearningmechanism,butinthemannerinwhichbottomupinformationismixedwith
topdowninformationstoredinthelearnedsynapticweights,asamodelofcognitiveexpectationor
perceptualcompletion.ThesignificantfeatureoftheARTmodelisthatthepatternrecognitionnodes
inwhatiscalledtheF2layer,areequippednotonlywithbottomupreceptivefieldsforpattern
recognition,butalsowithprojectivefieldsthatpropagatetopdownbacktotheinputorF1layer,and
thepatternofsynapticweightsintheseprojectivefieldsgenerallymatchthebottomupweightsused
forrecognition.If,afterlearningiscomplete,apartialorincompletepatternispresentedattheinput,
thatpatternwillstimulatetheactivationofthesingleF2nodewhosesynapticweightsbestmatchthe
inputpattern.TopdownfeedbackfromthatF2nodewillinturnimpingeitspatternbackontheF1
layer,fillinginorcompletingeventhemissingportionofthepattern,inamannerthatissuggestiveof
perceptualcompletionofmissingoroccludedportionsofarecognizedobject.ThefactthattheF2
nodesencodewholecategoriesofsimilarpatterns,ratherthanexactsinglepatterns,embodiesakind
ofinvarianceinthemodeltothevariationsbetweenpatternsofthesamecategory.

4.1ProblemswithAdaptiveResonance
Theinvarianceembodiedintheprincipleofadaptiveresonancediffersfundamentallyfromthe
invarianceobservedinperception,becausethesynapticweightsoftheF2nodeafterlearningseveral
patterns,encodeonlyasinglepatternatafixedlocationinthemodel,andthatpatternisakindof
average,orblurringtogetherofallofthepatternsthatbelongtothatcategory.Inotherwordsthe
systemwouldbehavemuchthesameifallofthepatternsofaparticularcategorywerefirstaveraged
togetherandthenlearnedasasinglepattern,ratherthanpresentedinsequenceasvariationsona
centraltheme.Thisimposesasevererestrictiononthekindofvariationthatcanbetoleratedwithina
category,foritrequiresasignificantoverlapbetweenpatternswithinaparticularcategory,otherwise
theaverageofthepatternsinthatcategorywouldproduceonlyafeaturelessblur.Asamodelof
learningandcategorizationthisisnotnecessarilyafatalproblem,aslongasthefeaturesrepresented
bytheF1nodesarepresumedtoalreadybeinvarianttostimulusvariation,i.e.thattheyencode
significantandstablecharacteristicsofthestimuluspattern,andthereforesignificantlysimilar
patternswouldbeexpectedtohaveconsiderableoverlapintheirF1featurerepresentation.However
theprincipleofadaptiveresonanceisinadequateasageneralmodeloftopdownfeedbackfor
perceptualcompletionacrossaninvariancerelation,becausethefeedbackinthismodelcanonly
completeasinglevariantoftherecognizedpatterninarigidtemplatelikemanner,andthatpatternis
nomorethanablurredtogetheraverageofallofthepatternsofthatparticularcategory.
Considerbycontrast,thepropertyofspatialinvarianceinvisualrecognition.Aspatialpattern,for
exampletheshapeoftheletterE,hasverylittleoverlap,pointforpoint,withvariationsofthatpattern
atdifferentorientations.Andyetthoserotatedpatternsarenotperceivedasapproximateorimperfect
letterE'swithdiminishedrecognitionconfidence,buteachoneisperceivedasaperfectEshape,
althoughitisalsoperceivedtoberotatedbysomeangle.Ifontheotherhandthepatternistruly
incomplete,liketheshapeoftheletterFconsideredasanincompleteEshape,thisdoesindeed
registerperceptuallyasapartialorimperfectmatchtotheshapeoftheletterE.Furthermore,
identificationoftheFshapeasanincompleteEimmediatelyhighlightstheexactmissingsegment,
i.e.thatsegmentisperceivedtobemissingfromaveryspecificportionofthefigure,andtheexact
locationofthatmissingsegmentvarieswiththelocation,orientation,andscaleoftheFstimulus.This
isaverydifferentandmorepowerfulkindofinvarianceandcompletionthanthatembodiedinthe
ARTmodel.Andyetitisexactlythekindofinvariancetostimulusvariationthatwouldberequiredin
theF1noderepresentationtomaketheARTmodelatallviableasamodelofrecognition.
Theproblemcanbetracedtothecentralprincipleofrepresentationinthemodel,whichisaspatial
templatethatisanchoredtothetissueofthebrainintheformofafixedreceptivefield.This
mechanismisthereforehardwiredtorecognizeonlypatternsthatappearatexactlythesamephysical
locationasthattemplateinthebrain.TheproblemofinvarianceintheARTmodelbecomes
abundantlyclearwhenattemptingtoapplyitsprincipleofinvariancetothespatialvariationsof
rotation,translation,andscale.LearningrotationinvarianceintheARTmodelforapatternlikeE
wouldbeequivalenttolearningthesinglepatternconstructedbythesuperpositionofEsatall
orientationssimultaneously,whichcreatesnothingbutacircularblur.Andthemodelaftertraining
wouldrespondmorestronglytothiscircularblurthantoanyactualletterE.Addingtranslationand
scaleinvariancetothesystemwouldinvolvelearningthesuperpositionofeveryrotation,translation
andscaleofthelearnedpatternacrossthevisualfield,whichwouldproducenothingbutauniform
blur.
TheinvarianceembodiedintheARTmodeltovariationsinthepatternswithinaparticularcategoryis
notreallyaninvariance,butismoreofablindnesstothosevariations,becausewhendetectinga
patternintheinput,theF2recognitionmechanismcannotdeterminewhichoftheallowablevariations
ofthepatternareactuallypresentontheinput.Whatisrequiredtoaccountforinvariancein
perceptionisasystemthatcandetectthecharacteristicpatternoftheinputdespitestimulusvariations,
andyethaveacapacitytocompleteapartialpatternwithrespecttothespecificvariationofthe
patternpresentontheinputfieldinotherwords,invarianceinrecognition,butspecificationin

completion.Thefactthatthisfunctionalityisinprinciplebeyondthecapacityoftheneuralreceptive
fieldwasrecognizedalreadybyLashley(1942),andwasacentralthemeofGestalttheory.

5TheBindingProblem
Thesubjectiveconsciousexperienceexhibitsaunitaryandintegratednaturethatseemsfundamentally
atoddswiththefragmentedarchitectureidentifiedneurophysiologically,anissuewhichhascometo
beknownasthebindingproblem.Fortheobjectsofperceptionappeartousnotasanassemblyof
independentfeatures,asmightbesuggestedbyafeaturebasedrepresentation,butasanintegrated
whole,witheverycomponentfeatureappearinginexperienceintheproperspatialrelationtoevery
otherfeature.Thisbindingoccursacrossthevisualmodalitiesofcolor,motion,form,andstereoscopic
depth,andasimilarintegrationalsooccursacrosstheperceptualmodalitiesofvision,hearing,and
touch.Thequestioniswhatkindofneurophysiologicalexplanationcouldpossiblyofferasatisfactory
accountofthephenomenonofbindinginperception?
Onesolutionistoproposeexplicitbindingconnections,i.e.neuronsconnectedacrossvisualor
sensorymodalities,whosestateofactivationencodesthefactthattheareasthattheyconnectare
currentlyboundinsubjectiveexperience.Howeverthissolutionmerelycompoundstheproblem,forit
representstwodistinctentitiesasboundtogetherbyaddingathirddistinctentity.Itisadeclarative
solution,i.e.thebindingbetweenelementsissupposedlyachievedbyattachingalabeltothemthat
declaresthatthoseelementsarenowbound,insteadofactuallybindingtheminsomemeaningfulway.
VonderMalsburgproposesthatperceptualbindingbetweencorticalneuronsissignalledbywayof
synchronousspiking,thetemporalcorrelationhypothesis(vonderMalsburg&Schneider1986).This
concepthasfoundconsiderableneurophysiologicalsupport(Eckhornetal.1988,Engeletal.1990,
1991a,1991b,Grayetal.1989,1990,1992,Gray&Singer1989,Stryker1989).Howeveralthough
thesefindingsaresuggestiveofsomesignificantcomputationalfunctioninthebrain,thetemporal
correlationhypothesisasproposed,islittledifferentfromthebindinglabelsolution,theonly
differencebeingthatthelabelisdefinedbyanewchannelofcommunication,i.e.bywayof
synchrony.Ininformationtheoreticterms,thisisnodifferentthansayingthatconnectedneurons
possestwoseparatechannelsofcommunication,onetotransmitfeaturedetection,andtheotherto
transmitbindinginformation.Thefactthatoneofthesechannelsusesasynchronycodeinsteadofa
ratecodeshedsnolightontheessenceofthebindingproblem.Furthermore,asShadlen&Movshon
(1999)observe,thetemporalbindinghypothesisisnotatheoryabouthowbindingiscomputed,but
onlyhowbindingissignaled,asolutionthatleavesthemostdifficultaspectoftheproblem
unresolved.
Iproposethattheonlymeaningfulsolutiontothebindingproblemmustinvolvearealbinding,as
impliedbythemetaphoricalname.Agluethatissupposedtobindtwoobjectstogetherwouldbemost
unsatisfactoryifitmerelylabeledtheobjectsasbound.Thesignificantfunctionofglueistoensure
thataforceappliedtooneoftheboundobjectswillautomaticallyactontheotheronealso,toensure
thattheboundobjectsmovetogetherthroughtheworldevenwhenone,orbothofthemarebeing
actedonbyforces.Inthecontextofvisualperception,thissuggeststhattheperceptualinformation
representedincorticalmapsmustbecoupledtoeachotherwithbidirectionalfunctionalconnections
insuchawaythatperceptualrelationsdetectedinonemapduetoonevisualmodalitywillhavean
immediateeffectontheothermapsthatencodeothervisualmodalities.Theonedirectionalaxonal
transmissioninherentintheconceptoftheneurondoctrineappearsinconsistentwiththeimmediate
bidirectionalrelationrequiredforperceptualbinding.Eventhefeedbackpathwaysbetweencortical
areasareproblematicforthisfunctionduetothetimedelayinherentintheconceptofspiketrain
integrationacrossthechemicalsynapse,whichwouldseemtolimitthereciprocalcouplingbetween
corticalareastothosewithinasmallnumberofsynapticconnections.Thetimedelaysacrossthe
chemicalsynapsewouldseemtoprecludethekindofintegrationapparentinthebindingofperception
andconsciousnessacrossallsensorymodalities,whichsuggeststhattheentirecortexisfunctionally
coupledtoactasasingleintegratedunit.

6Atomisticv.s.HolisticPrincipleofComputation
Theneurondoctrineincorporatesanatomisticassumption,thatneurocomputationinvolvesdiscrete
signalscommunicatedalongfixedtransmissionlinesbetweendiscretecomputationalelements.The
westernscientifictraditionhasaparticularbiasinfavorofanatomisticview,notonlyof
neurocomputation,butoftheprinciplesofcomputationingeneral.Forthereductionistapproach
favoredbywesternscienceinvolvesbreakingcomplexproblemsintosimplerpieceswhichcanthen
beanalyzedindividually.ThatiswhythediscoverybyCajalofthediscretecellularstructureofthe
nervoussystemtriggeredanintensiveprogramofinvestigationofthepropertiesoftheindividual
neuron,inthehopethatthiswouldshedlightontheoperationofthelargernervoussystemmadeup
ofthoseelementaryunits.Therearecertainclassesofphysicalsystemsforwhichthisanalytical
approachworkswell,specifically,forsystemswhosecomponentelementsoperaterelatively
independently,i.e.whentheprocessesandmechanismsactivewithintheelementaremoreclosely
coupledthanthosethatoperatebetweenelements.Howeverthisatomisticviewofneuroscienceis
nothingmorethananoptimisticassumption,becauseatomisticsystemsareincomparablyeasierto
studyandtocharacterizemathematicallythanareholisticorwidelycoupledanddynamicfeedback
systems.Butalthoughitwouldbeveryconvenientforneuroscienceifthebrainworkedonan
atomisticprinciple,Gestalttheoryhasdemonstratedwithagreatvarietyofdifferentphenomenathat
thisisinfactnotthecase,andthatthebrainoperatesonaholistic,emergentprincipleofcomputation.
Furtherevidenceforholisticprocessesinthebraincomesfromelectroencephalogram(EEG)
recordingswhichrevealglobalelectricaloscillationsthatpervadetheentirecortex.Thisglobal
resonanceisnowbeginningtomanifestitselfalsoinneurophysiologicalrecordingsintheformof
synchronousoscillations.Infactthesynchronousspikingofremotecorticalneuronsisdifficultto
accountforinconventionalneuralterms,becausethephaseofthespikingsignalshouldbecome
scrambledasitpropagatesdowntheaxoncollateralsanddendritesofthepreandpostsynapticcells
respectively,duetotherandomthicknessesandirregularpathlengthsofthosemanyparallelbranches.
Thesynchronyshouldbefurtherdisruptedbythespiketrainintegrationacrossthechemicalsynapses,
eachofwhichactsasalowpassfilter,blurringthesharpspikeofthepresynapticactionpotential
intoasmoothriseanddecayinthepostsynapticcell.Thefactthatahighresolutiontemporal
synchronyisobservedacrossremotecorticalareasconnectedbycountlessparallelpathsthrough
countlesssynapticjuncturessuggeststhatthissynchronyisactuallytransmittedbysomeothermeans.
Infactithasbeenshown(Pribram1971,Blandetal.1978)thatthediscretespikingoftheaction
potentialsissuperimposedonamoresubtlegradedpotentialoscillation,andPribram(1971)showed
thatthegradedpotentialoscillationpersistsevenwhenthespikingdischargefallsbelowthreshold.
Thissuggeststhatthespikingdischargeisnotthecausaloriginoftheneuralsignal,butmerelythe
overtmanifestationofamoresubtleunderlyingelectricaloscillation,likethewhitecapsonocean
waves,andthatoscillationseemstopervadetheneuraltissueunrestrictedbytheboundariesofthe
cell.Thewholeconceptoftheneurondoctrinehasblindedneurophysiologiststothepossibilityof
significantsignalsthatpervadetheextracellularmatrix,foritisassumedthatsignalswhichareneither
channeledbythecellwall,norgatedbythechemicalsynapse,cannotpossiblytakepartinmeaningful
computation.TheGestaltperspectiveontheotherhandsuggeststhatitisjustthatkindofholistic
fieldlikeprocesswhichmustbesoughtouttoaccountforthemostsignificantandinterestingaspects
ofneurocomputation.
TheprincipalreasonforthedemiseoftheGestaltmovementwasitsfailuretospecifythevague
holisticaspectsofperceptionthatitidentifiedinmorerigorousquantitativeterms,inamannerthat
relatestoknownneurophysiology.Inanotherpaper(Lehar2000)Ihavespecifiedtheelusiveholistic
Gestaltprinciplessomewhatmorepreciselyastheprinciplesofemergence,reification,multistability,
andinvariance.InthatpaperIproposedacomputationalmodelofperceptiontodemonstratehow
thosesameGestaltprinciplescanserveausefulcomputationalfunctioninperception.Howeverthat
modelwasexpressedintermsthatareindependentofanyneurophysiologicalassumptions.The
objectiveofthepresentpaperistoproposehowthoseGestaltaspectsofperceptioncanberelatedto
ourunderstandingofneurophysiology,inordertodevelopaneurophysiologicallyplausibleGestalt

theoryofneurocomputation.
6.1Emergence
ThemostsignificantgeneralpropertyofperceptionidentifiedbyGestalttheorywasthepropertyof
emergence,wherebyalargerpatternorstructureemergesunderthesimultaneousactionof
innumerablelocalforces.Koffka(1935)suggestedaphysicalanalogyofthesoapbubbleto
demonstratetheoperationalprinciplebehindemergence.Thesphericalshapeofasoapbubbleisnot
encodedintheformofasphericaltemplateorabstractmathematicalcode,butratherthatform
emergesfromtheparallelactionofinnumerablelocalforcesofsurfacetensionactinginunison.The
finegrainedandcontinuouscharacterofemergenceacrossbothspaceandtimeisfundamentallyat
oddswiththeatomisticnotionofneurocomputationembodiedintheneurondoctrine.
6.2Reification
Reificationistheconstructive,orgenerativeaspectofperceptionidentifiedbyGestalttheory.
ReificationisseeninvisualillusionsliketheKanizsafigure,wherethesubjectiveexperienceofthe
illusionencodesmoreexplicitspatialinformationthanthestimulusonwhichitisbased.Specifically,
illusoryedgesareseeninplaceswheretherearenoedgesinthestimulus,andthoseedgesbounda
continuoussurfaceperceptwhoseillusorybrightnesspervadestheentireillusorysurfaceasaspatial
continuum.Reificationinperceptionindicatesthatperceptionisnotmerelyapassiveprocessof
recognitionoffeaturesinthevisualinput,assuggestedintheneurondoctrine,butthatperception
createstheperceivedworldasaconstructiveorgenerativeprocess.
6.3Multistability
Multistabilityisseeninavarietyofvisualillusions,includingtheNeckercube,andRubin'sfigure/
vaseillusion.Thesignificancefortheoriesofperceptionisthatitrevealsperceptionasadynamic
systemwhosestablestatesrepresentthefinalpercept.Multistabilityandreificationworkhandin
hand,becauseeachperceptualstateisreifiedasafullsurfaceorvolumeperceptineachofits
alternatestates,i.e.thesubjectivereversalofafigureliketheNeckercubeisnotexperiencedasa
changeinacognitiveinterpretation,ortheflippingofasinglecognitivevariable,butisvividly
experiencedasaninversionofaperceptualdatastructure,changingtheperceiveddepthofeverypoint
intheperceivedstructure.
6.4Invariance
AcentralfocusofGestalttheorywastheissueofinvariance,i.e.howanobject,likeasquareora
triangle,canberecognizedregardlessofitsrotation,translation,orscale,orwhateveritscontrast
polarityagainstthebackground,orwhetheritisdepictedsolidorinoutlineform,orwhetheritis
definedintermsoftexture,motion,orbinoculardisparity.Invarianceisalsoseenintheperceptionof
colorandbrightness,wherethecolorofanobjectisgenerallyjudgedindependentofthecolorofthe
lightfallingonit.Recognitionisalsoinvarianttoelasticdeformationofnonrigidobjects,for
exampleanimalbodiesarerecognizedindependentoftheirposturalconfiguration,andfacesare
recognizeddespitedistortionsimposedbyfacialexpressions,orevenmoreextremedistortionsoften
observedincaricatures.Evennormallyrigidobjectslikehousesorcarsarerecognizedindeformed
form,aswhenseenthroughdistortingmirrorsorlenses,orasoftendepictedincartoonrenditions.
Althoughisolatedcounterexamplesexist,forexampletherecognitionofcomplexfiguresandof
facesisnotcompletelyrotationinvariant,thefactthatinvarianceisobservedthroughsomany
stimulusvariationsandacrosssuchawidevarietyofperceptualmodalitiessuggeststhatinvarianceis
fundamentaltoperception,andthereforereflectsafundamentalcharacteristicofthemechanismof
biologicalcomputation.

6.5BrainAnchoring
Oneofthemostdisturbingpropertiesofthephenomenalworldformodelsoftheperceptual
mechanisminvolvesthesubjectiveimpressionthatthephenomenalworldrotatesrelativetoour
perceivedheadasourheadrotatesrelativetotheworld,andthatobjectsinperceptionareobservedto
translateandrotatewhilemaintainingtheirperceivedstructuralintegrityandrecognizedidentityin
theirmotionsthroughtheperceivedworld.Ifweassumethatthestructuralperceptoftheworldis
representedbyaspatialpatternofactivationofsomesortinthetissueofthebrain,thissuggeststhat
theinternalrepresentationofexternalobjectsandsurfacesisnotanchoredtothetissueofthebrain,as
suggestedbycurrentconceptsofneuralrepresentation,butisfreetorotateandtranslatecoherently
relativetotheneuralsubstrate,assuggestedinKhler'sfieldtheory(Khler&Held1947).Inother
wordstheperceptualpictureoftheworldcanmoverelativetotherepresentationalsubstrate,and
discretepatternsofperceptualstructurecanmoverelativetothatbackgroundwhilemaintainingtheir
perceptualintegrityandrecognizedidentity.
6.6TheEnigmaofGestalt
Itissmallwonderthatinthefaceofthisformidablearrayofmostenigmaticproperties,theoriesof
visionhavegenerallybeenrestrictedtosimplisticmodelsofisolatedaspectsoftheproblemina
piecemealmanner.ThisdoesnothoweverinanywayjustifythefactthattheGestaltpropertiesof
perception,discoveredandidentifiedalmostacenturyago,aresounderrepresentedincontemporary
theoriesofneurocomputation.OurfailuretofindaneurophysiologicalexplanationforGestalt
phenomenadoesnotsuggestthatnosuchexplanationexists,onlythatwemustbelookingforitinthe
wrongplaces.TheenigmaticnatureofGestaltphenomenaonlyhighlightstheimportanceofthe
searchforacomputationalmechanismthatexhibitsthesesameproperties.Infact,anymodelthatfails
toaddresstheGestaltphenomenaofperceptionisworsethannomodelatall,foritisadiversionfrom
therealissuesofperception.

7HarmonicResonanceTheory
ThepropertiesofperceptionasobservedphenomenallyandasdescribedbyGestalttheoryaretruly
bafflingwhenitcomestoproposingacomputationalmechanismtoaccountforthoseproperties.
Howeverthereisonephysicalphenomenonthatexhibitsexactlythosebafflingpropertiesobservedin
perception,andthatisthephenomenonofharmonicresonance,ortherepresentationofspatial
structureexpressedaspatternsofstandingwavesinaresonatingsystem.Thistantalizingsimilarity
cannotbecoincidental,consideringthatnootherphysicalmechanismorphenomenonhaseverbeen
identifiedthatexhibitsthesesameenigmaticproperties.Themostremarkablepropertyofharmonic
resonanceisthesheernumberofdifferentuniquepatternsthatcanbeobtainedineventhesimplest
resonatingsystem.Apioneeringstudyofmorecomplexstandingwavepatternswaspresentedby
Chladni(1787)whodemonstratedtheresonantpatternsproducedbyavibratingsteelplate.The
techniqueintroducedbyChladniwastosprinklesandontopoftheplate,andthentosettheplateinto
vibrationbybowingwithaviolinbow.Thevibrationoftheplatecausesthesandtodanceabout
randomlyexceptatthenodesofvibrationwherethesandaccumulates,therebyrevealingthespatial
patternofnodes.ThistechniquewasrefinedbyWaller(1961)usingapieceofdryicepressedagainst
theplate,wheretheescapinggasduetothesublimationoftheicesetstheplateintoresonance,
resultinginahighpitchedsquealastheplatevibrates.Figure1(adaptedfromWaller1961P.69)
showssomeofthepatternsthatcanbeobtainedbyvibratingasquaresteelplateclampedatits
midpoint.Thelinesinthefigurerepresentthepatternsofnodesobtainedbyvibrationatvarious
harmonicmodesoftheplate,eachnodeformingtheboundarybetweenportionsoftheplatemoving
inoppositedirections,i.e.duringthefirsthalfcycle,alternatesegmentsdeflectupwardswhile
neighboringsegmentsdeflectdownwards,andthesemotionsreverseduringthesecondhalfcycleof
theoscillation.ThedifferentpatternsseeninFigure1canbeobtainedbytouchingtheplateata
selectedpointwhilebowingattheperipheryoftheplate,whichformsanodeofoscillationatthe

dampedlocation,aswellasattheclampedcenterpointoftheplate.Theplateemitsanacousticaltone
whenbowedinthismanner,andeachofthepatternsshowninfigure1correspondstoaunique
temporalfrequency,ormusicalpitch,thelowesttonesbeingproducedbythepatternswithfewerlarge
segmentsshownattheupperleftoffigure1,whilehighertonesareproducedbythehigherharmonics
depictedtowardsthelowerrightinthefigure.Thehigherharmonicsrepresenthigherenergiesof
vibration,andareachievedbydampingclosertothecentralclamppoint,aswellasbymorevigorous
bowing.Therearemanymorepossiblepatternsinasquareplatethanthosedepictedinfigure1,
whichwouldberevealedbysuspendingtheplatewithoutclamping,allowingpatternswhichdonot
happentoexhibitanodeatthecenterofthesquare,andofcoursetherearemanymorepatterns
possibleinplatesofdifferentshapes(Waller1961),andmanymorestillinvolumetricresonant
systemssuchasavibratingcubeorsphere,whichdefinethreedimensionalsubdivisionsofthe
resonatingvolume,althoughthesehavenotreceivedmuchattentionduetothedifficultyinobserving
thestandingwavepatternswithinasolidvolumeorvolumetricresonantcavity.Faraday(1831)
extendedChladni'sphenomenonbyobservingstandingwavesonthesurfaceofliquids,which
producesgeometricalarraysofstandingwavesonthesurfaceofthefluidintheformofconcentric
rings,parallelridges,gridandcheckerboardpatterns,arraysofconelikepoints,andevenbrick
patterns.ThisworkhasbeenextendedmorerecentlybyCristiansenetal(1992),Kumar&Bajaj
(1995),Kudrolli&Gollub(1996),Kudrollietal.(1998)andothers,whohavedemonstratedpatterns
ofequilateraltriangles,regularhexagons,superlattice,andquasicrystalarraypatternsbydrivingthe
oscillationofthefluidlayerwithacontrolledwaveform.Figure2showssomeofthepatterns
producedbyKudrollietal.Itseemsthatthisworkisonlytouchingthesurfaceofthefullpotentialof
thisphenomenonforproducingcomplexgeometricalpatternsbyrelativelysimpledrivingoscillations.

Figure1
Chladnifiguresforasquaresteelplate(adaptedfromWaller1961)demonstratesthefantastic
varietyofstandingwavepatternsthatcanarisefromasimpleresonatingsystem.Asquaresteel
plateisclampedatitsmidpointandsprinkledwithsand.Itisthensetintovibrationeitherby
bowingwithaviolinbow,orbypressingdryiceagainstit.Theresultantstandingwavepatterns
arerevealedbythesand,thatcollectsatthenodesoftheoscillationwherethevibrationis
minimal.

Figure2
Variouspatternsofstandingwavesonfluidsurfacesgeneratedbyvibratingthecontaining
vesselswithvariousdrivingfrequencies,producingA:hexagonal,B:rectangularlattice,C:
quasicrystalpatterns,amongmanymore.D:Thispatternisdefinedmorebytheshapeofthe
wallsofthecontainerratherthanbythedrivingwaveform,showinghowinterferencepatterns
intheresonatingsystemtendtosubdividetheresonatingsystemintoperiodicandsymmetric
subpatternsinanessentiallyGestaltmanner.
7.1ReactionDiffusionSystems
Theutilityofstandingwavepatternsasarepresentationofspatialformisdemonstratedbythefact
thatnaturemakesuseofaresonancerepresentationinanotherunrelatedaspectofbiologicalfunction,
thatofembryologicalmorphogenesis,orthedevelopmentofspatialstructureintheembryo.Afterthe
initialcelldivisionsfollowingfertilization,theembryodevelopsintoanellipsoidofessentially
undifferentiatedtissue.Then,atsomecriticalpointaperiodicbandedpatternisseentoemergeas
revealedbyappropriatestainingtechniques,showninfigure3A.Thispatternindicatesanalternating
patternofconcentrationofmorphogens,i.e.chemicalsthatpermanentlymarktheunderlyingtissue

forfuturedevelopment.Thispatternissustaineddespitethefactthatthemorphogensarefreeto
diffusethroughtheembryo.Themechanismbehindtheemergenceofthisperiodicpatternisa
chemicalharmonicresonanceknownasreactiondiffusion(Turing1952,Prigogine&Nicolis1967,
Winfree1974,Welshetal.1983)inwhichacontinuouschemicalreactioninvolvingamorphogenP
catalyzestheproductionofmoremorphogenPaswellasofamorphogenS,buttheconcentrationof
morphogenSinturninhibitsproductionofmorphogenP(seeGilbert1988pp655661fora
summary).Theresultofthiscircularreactionisanalogoustotheperiodicpatternsofaresonating
steelplate.Thechemicalharmonicresonanceintheembryocantherebydefineaspatialaddressing
schemethatidentifieslocalcellsintheembryonictissueasbelongingtooneoranotherpartofthe
globalpatternintheembryobywayoftherelativeconcentrationofcertainmorphogens.Perhapsthe
mostvisibleexampleofthekindsofpatternsthatcanbedefinedbyreactiondiffusionsystemsare
thoseseeninanimalmarkings,suchasthestripesofthezebraorthespotsoftheleopardwhichhave
alsobeenattributedtoreactiondiffusionprocesses(Murray1981,1988).Mostofthemarkingson
animalskinsareforthepurposeofcamouflage,andthereforethosepatternsaregenerallysomewhat
irregularforthatreason.Howevertherearecaseswhereanimalandplantmarkingsareintendedto
attractattention,andinthosecasesthetruepotentialofmorphogenicprocessesasarepresentationof
geometricalformisdemonstrated.Thisisseenforexampleinpoisonousanimals,likethecaterpillar
showninfigure3B,aswellasinavarietyofpoisonoussnakesthatshowperiodicgeometrical
patternsofrings,diamonds,orstripes,aswellasinanimalsthatdisplayformatingpurposes,likethe
plumageofapeacock,andofvariousbirdsofparadise,andintheplantkingdomitisseeninthe
formsofflowers.Thesymmetryandperiodicityobservedintheseexamplescanbeseenasboth
evidenceofthepropertiesofmorphogenesis,andatthesametime,aspropertiesofbiologicalvision,
whoseparticularsensitivitytoperiodicityandsymmetryareexploitedbythosemorphogenic
markingsinordertoattractvisualattention.
Inthecaseofanimalcoatmarkingsthechemicalpatternsonlydefinedifferentpatternsofcoloration.
Butthesamespatialaddressingschemeisalsoresponsiblefordefiningthepatternoftissuetypesin
theembryo,forthepatternofconcentrationofthesemorphogensduringacriticalperiodof
developmenthasbeenshowntoberesponsibleforpermanentlymarkingthetissueforsubsequent
developmentintoboneversusmuscletissueetc.Thisthereforeexplainssomeofthegeometrical
regularitiesobservedintheshapeofthemuscles,bones,andinternalorgansofthebody.The
periodicityinmorphogenesisisultimatelyresponsiblefortheperiodicsegmentsobservedinthe
bodiesofwormsandinsects,inthevertebraeofvertebrates,andsimilarresonanceshavebeen
implicatedinmanyothersymmetriesandperiodicitiesinplantandanimalforms,includingthe
bilateralsymmetryofthehumanbody,thepentalateralsymmetryofthestarfish,theangularandradial
periodicityofthebonesinthehumanhandandfingers,andthegeometricalformsobservedinplant
andflowerstructures.Murray(1988)makestheconnectionbetweenchemicalandvibrational
standingwaves,showinghowavarietyofdifferentanimalcoatpatternscanbeproducedasstanding
wavesinasteelplatecutintheshapeofananimalskin,showninfigure3C.

Figure3
A:Aperiodicbandedpatternrevealedbychemicalstainingemergesinadevelopingembryo,
duetoachemicalharmonicresonancewhosestandingwavesmarktheembryonictissuefor
futuregrowth.B:Thischemicalharmonicresonancehasbeenidentifiedasthemechanism
behindtheformationofpatternsinanimalskins,aswellasfortheperiodicityinthevertibraeof
vertibrates,thebilateralsymmetryofthebodyplan,aswellastheperiodicityofthebonesinthe
limbsandfingers.C:Murrayshowstheconnectionbetweenchemicalandvibrationalstanding
wavesbyreplicatingthepatternsofleopardspotsandzebrastripesinthestandingwave
resonancesinavibratingsteelsheetcutintheformofananimalskin.
7.2PropertiesofHarmonicResonance
Therehasbeenmuchinterestrecentlyinthephenomenonknownaschaostheory(Gleik1987)in
whichspatialortemporalpatternisseentoemergeunexpectedlyinotherwisedisorganizedsystems.
Chaostheoryhasevenbeenimplicatedintheoriesofneurocomputation(Freeman1995).However
chaosissomewhatofamisnomer,sincethephenomenonisbetterdescribedasunexpectedorder
ratherthandisorder.Furthermore,manyofthephenomenathatfallundertherubricofchaostheory,
suchasperioddoublingandlimitcycles,areactuallymanifestationsofharmonicresonance,an
orderlyratherthanachaoticorganizationalprinciple.
Thereareseveralpropertiesoftheharmonicresonancemodelthataresuggestiveofhuman
recognition.Unlikearigidtemplate,thepatterndefinedbyastandingwaverepresentationiselastic
andadaptive.Thiscanbeseeninthemannerthatthespatialpatternsofanimalskinsaredefined.The

parametersofthereactiondiffusionthatdistinguishbetweenthespotsandstripesofthetiger,zebra,
leopard,andgiraffeareencodedasgeneralrulesforthegenerationofthosepatternsratherthanasa
spatialtemplateofanyonesuchpattern.Forexampleifaspotorstripeweretobefixedatonepoint
asthepatternwasemerging,therestofthepatternwouldredistributeitselftoadapttothatfixed
featurewhilemaintainingthegeneralcharacteroftheencodedpattern.Thisinvarianceinthe
representationallowsonesetofparameterstogenerateaninfinitevarietyofexemplarsofany
particularpatterntype,ortoadaptmostflexiblytoanyfixedconstraintsorboundaryconditions.Itis
harmonicresonancethatexplainstheadaptivenessofthebodyplaninmorphogenesistovariationsin
thegeometryoftheembryonictissue.Thisadaptivenessisseeninmostdramaticforminthe
body(ies)ofsiamesetwins,wherethebodyplanisobservedtosplitlikeamirrorreflection,
bifurcatingeverybone,muscle,tendon,andbloodvesselasifitwerepartoftheoriginalplan.This
kindofinvariancetodistortionisaprominentcharacteristicofhumanrecognitionalso,asseenfor
exampleintheeasewithwhichwerecognizewildlydistortedcaricaturesoffamiliarfaces,orthe
distortedreflectionsinacurvedmirror.
7.3EmergenceinHarmonicResonance
AsoapbubbleistheclassicalphysicalanalogyusedinGestalttheory(Koffka1935p.107,Attneave
1982)toexemplifytheprincipleofemergence.Howeverthestandingwaveoffersanevenmore
dramaticexampleoftheprincipleofemergence,andthekindofholisticprocessesidentifiedby
Gestalttheory.Likethesoapbubble,astandingwavepatterninaflute,forexample,istheresultof
relativelysimplelocalinteractionsatthemolecularlevel.Yettheeffectsoftheseinteractionsareas
globalastheresonatingsystemitself.Unlikethesoapbubble,theresonancecandefineawholesetof
uniquepatternscorrespondingtothefundamentalanditshigherharmonics,asseenintheChladni
figures.Theemergentpatternsdefinedbythoseharmonicsarenotrigid,ortemplatelike,butmore
likeanelastictemplatethatautomaticallyconformstoirregularitiesintheresonatingcavity.For
exampleifafluteiscurved,orflaredattheendlikeatrumpet,orbulginginthemiddlelikeabarrel,
theperiodicpatternoftheresonancewillbecorrespondinglydeformed,definingperiodicsegmentsof
equalvolume,althoughofunequalgeometry.Inotherwords,theresonancedefinesthetopology,
ratherthanthetopographyoftheencodedforms.Resonancesinconnectedsystems,suchasthe
engineandchassisofyourcar,haveanaturaltendencytosynchronizeorcouplewitheachother
(Dewan1976,Strogatz&Stewart1993)soastoproduceasingleemergentoscillationwhichexhibits
higherharmonicswhicharecharacteristicofeachcomponentresonator,embeddedinafundamental
waveformthatcapturestheresonanceofthesystemasawhole.
7.4HarmonicResonanceintheBrain
Oscillationsandtemporalresonancesarefamiliarenoughinneuralsystemsandareobservedatevery
scale,fromlongperiodcircadianrhythms,tothemediumperiodrhythmicmovementsoflimbs,allthe
waytotheveryrapidrhythmicspikingofthesinglecell,orthesynchronizedspikingofgroupsof
cells.Harmonicresonanceisalsoobservedinsinglecelledorganismsliketheparameciuminthe
rhythmicbeatingofflagellainsynchronizedtravellingwaves.Similarwavesareobservedin
multicellularinvertebrates,suchasthesynchronizedwavelikeswimmingmovementsofthehydra
andthejellyfish,whosedecentralizednervoussystemsconsistofadistributednetworkoflargely
undifferentiatedcells.Themuscleoftheheartprovidesperhapstheclearestexampleofsynchronized
oscillation,fortheindividualcellsofthecardiacmuscleareeachindependentoscillatorsthatpulseat
theirownrhythmwhenseparatedfromtherestofthetissueinvitro.Howeverwhenconnectedto
othercellstheysynchronizewitheachothertodefineasinglecoupledoscillator.Thefactthatsuch
unstructuredneuralarchitecturescangiverisetosuchstructuredbehaviorsuggestsalevelof
computationalorganizationbelowthatoftheswitchingandgatingfunctionsofthechemicalsynapse.
Vertebratestooexhibitprominentrhythmicmotions,mostevidentinsimplervertebratessuchas
caterpillarsandcentipedes,butevenlargervertebratessuchassnakes,lizards,fish,andeelsexhibita
gracefulundulatingmotionsuggestiveofanunderlyingwavelikecomputationalmechanism.Strogatz

&Stewart(1993)haveevencharacterizedthemovementoflargermammalsintermsofoscillations,
showingforexamplehowthevariousgaitsofahorse(trot,canter,gallop,etc.)correspondtothe
variousmodesofoscillationoffourcoupledoscillators.Theideaofoscillationsinneuralsystemsis
notnew.Howevertheproposaladvancedhereisthatnaturemakesuseofsuchnaturalresonancesnot
onlytodefinerhythmicpatternsinspaceandtime,butalsotodefinestaticspatialpatternsintheform
ofelectricalstandingwaves,forthepurposethatiscommonlyascribedtospatialreceptivefields.
Whilethespecificneurophysiologicalevidenceforthispatternformationdeviceremainstobesought
outandidentified,Iwillshowthatasaparadigmfordefiningspatialpattern,thestandingwaveoffers
agreatdealmoreflexibilityandadaptivenesstolocalconditionsthanthealternativereceptivefield
model,andthatasingleresonatingsystemcanreplaceawholearrayofhardwiredreceptivefieldsin
aconventionalneuralmodel.
7.5PhysiologicalandPsychophysicalEvidence
Theneurophysiologicalbasisforastandingwavetheoryofneuralrepresentationissupportedbythe
observationthatblocksofneuraltissuethatareconnectedbyelectricalsynapses,orgapjunctions,to
formaneuralsyncytiumhavebeenshowntoexhibitsynchronousspikingactivity(Kandell&
Siegelbaum1985).Dermietzel&Spray(1993)haveidentifiedgapjunctionsincorticaland
subcorticaltissue,andshowthatgapjunctionsareubiquitousinthebrainandnervoussystem.
Peinadoetal.(1993)alsoimplicategapjunctionsinlocalcorticalcircuits.Howeverharmonic
resonancetheorydoesnotstrictlyrequiregapjunctions,forthecellmembranethatactsasaninsulator
fortheflowofdirectcurrentdoesnotinsulateanalternatingcurrent(AC)orvoltagefluctuations,
whicharefreetopassfromcelltocelllikeanACcurrentthatcrossestheinsulatingdielectricofa
capacitor.Thatiswhy,forexample,itispossibletomeasureneuralactivitywithextracellularaswell
asintracellularelectrodes.Bremer(1953)observedelectricaloscillationsinthecatspinalchord,that
maintainsynchronizationfromoneendofthechordtotheother,evenwhenthechordisseveredand
reconnectedbycontactalone.Bremerobservesthattheelectricalsynchronyalongthespinalchord
propagatesfasterthanelectricalimpulses.Gerard&Libet(1940)publishsimilarobservationsforthe
rabbit.Inmorerecentliteratureagreatvarietyofdifferentkindsofoscillationshavebeenassociated
withthevisual,somatosensory,andmotorcortices,suggestingthattheymaymediatefunctional
integrationofsensoryormotorinformationprocessinginthebrain.(Eckhornetal.1988,Nicoleliset
al.1995,Murthy&Fetz1992,Sompolinskyetal.1990).Hashemiyoon&Chapin(1993,1994)report
retinallyderiveddarkspontaneousfastfrequencyoscillationsthroughoutthesubcorticalvisualsystem
ofratswhicharesuppressedbytoniclightstimulation.Theyremainremarkablyphasecoherentwhile
fluctuatingbetweenmultiplefrequenciesapproximating10,20,and40Hz.Grayetal.(1989)report
stimulusinduced~40Hzoscillationsinthevisualcortexandsuggestafunctionalroleforthese
oscillationsinvisualprocessing.Severalresearchershaveproposedthatsuchsynchronousoscillations
arerelatedtotheintegrationoftheconsciousexperience(vonderMalsburg&Schneider1986,von
derMalsburg1987,Edelman1987,Llins1983,Cricketal.1990,Singeretal.1993,Eckhornetal.
1988,Zeki1993,Bressleretal.1993).
Psychophysicalevidencehasalsobeenreportedinsupportofsomekindofoscillationsinperceptual
processing.Lehar(1994)presentsaharmonicresonancetheorytoaccountfortheformationofalarge
varietyofillusorycontoursindifferentspatialconfigurations,demonstratingtheflexibilityof
harmonicresonanceasanalternativetoareceptivefieldmodel,anddemonstratingsomeofthe
functionalpropertiesofharmonicresonanceinvisualprocessing.Kristofferson(1990)showshowthe
linearplotoftheWeberfunctionforatemporaldiscriminationtaskbecomesastepfunctionafter
extensivepractice,whenthetaskbecomesoverlearned.Thestepfunctioniscomposedofaseriesof
flatplateauswithinwhichthediscriminationthresholdremainsconstant,separatedbysudden
increasesindiscriminationthreshold,sothatthestepfunctionstraddlesbackandforthacrossthe
nonlinearfunctionpredictedbyWeber'slaw.Aperioddoublingisobservedbetweensuccessive
plateaus,i.e.eachplateauisdoublethelengthofthepreviousplateau,asdiscussedbyGeissler
(1997).InotherwordsthemicrostructureofWeber'slawisnotasmoothlogarithmicfunction,buta
discontinuousstepfunctionrisinginoctaveslikethoseofamusicalscale.Geissler(1987,1998)

interpretsthisphenomenonasevidenceforaphaselockingbetweencoupledoscillatorsofarangeof
differentfrequencies.
7.6PatternRecognitionbyTunedResonators
Thestandingwaveandthepatternedreceptivefieldsharethepropertythattheyeachdefineaspatial
patternintheneuralsubstrate.Intheconventionalneuralnetworkparadigm,thecellbodyactsasthe
focalpoint,whosestateofelectricalactivityrepresentsthepresenceorabsenceofacorresponding
patternofactivationsampledinthecell'sreceptivefield.Thestandingwavepatternontheotherhand
appearsatfirstsighttobeamoredistributedrepresentation,inthatthepresenceofsuchapatternin
theneuralsubstrateestablishesapatternedinteractionbetweentheresonatingneurons,i.e.the
activationofanyonecelldependsonthetotalpatternofactivationinallneighboringcells,butthere
isnosinglecellthatisactiveinthepresenceofaparticularpattern,andinactiveintheabsenceofthat
pattern,asisthecasewiththecellbodyintheneuralnetworkparadigm.Thisfocalpointiscrucial,
sinceitwouldseemtobetheveryessenceofarecognitionsystemtoreduceacomplexspatialmatch
toasinglematchvalue.Thereishoweveraquantitythatisassociatedwitheachpatternofresonance.
InthecaseoftheChladnifiguresdepictedinFigure1,eachindividualpatternisassociatedwitha
uniquetemporalfrequencyofoscillation,oraudibletone.Thereforeanaudiorecordingofthe
vibratingsteelplatecouldbeusedtouniquelyidentifywhichpatternwaspresentontheplateduring
recording.Furthermore,anaudioplaybackofthatfrequencyinthepresenceoftheplatewouldhave
theeffectofregeneratingthatsamepatternofresonancebackontheoriginalplate.Theaudiotonecan
thereforebeconsideredasanabstractedrepresentation,orreduceddimensionalityencodingofthe
spatialpatternontheplate.Thereforematchingthetonegeneratedbyavibratingplatetoatonestored
inmemorycorrespondstoarecognitionofthatspatialpattern,justastheactivationofacellbodyina
receptivefieldmodelrepresentsarecognitionofthespatialpatternpresentinitsinputfield.Theitem
intheresonancemodelcorrespondingtothecellbodyinthereceptivefieldmodelcanbeenvisaged
assomekindoftunedresonator,perhapsacellwithanaturaltendencytospikeatacharacteristic
frequency.
TheprinciplebehindthisconceptofrecognitioncanbedemonstratedusingaChladniplatecoupledto
anarrayofacousticalresonatorstunedtothespecificharmonicfrequenciesofthatplate.Figure4A
showsthreeresonatorscoupledtoaChladniplate,tunedtothreeselectedpatternsofstandingwaves
fromfigure1.Forconveniencethesewillbecalledthe"+","x",and"/"patterns.Whentheplateisset
intovibration,thestandingwavepatternthatappearsontheplateactivatestheresonatortunedtothat
patternassuggestedforeachpatterninfigure4A.Thesystemautomaticallyincorporatesreification
withrecognition,forjustasthevibrationoftheplatewithaparticularstandingwavepatternactivates
itscorrespondingresonator,soalsodoesthevibrationoftheresonatoratitscharacteristicfrequency
automaticallyregenerateitscharacteristicpatternbackontheplate,assuggestedinfigure4B.

Figure4
A:Abankoftunedresonatorstunedtothefrequencyofthreespecificstandingwavepatterns
arecoupledtotheplatetobehaveasfeaturedetectors,thatbecomeactivewhenevertheir
patternofstandingwavesispresentontheplate.B:Thesystemautomaticallyperforms
reificationwithrecognition,fortheactivationofanyoftheresonatorsregeneratesits
characteristicstandingwavepatternbackontheplate.C:Iftheplateisrestingonrubber
ridgesintheformofoneofthestandingwavepatterns,theridgesbehaveasaninputpattern,
forcingtheresonanceontheplatetoconformtothepatternoftheinput.
7.7InputPatternAppliedbyDamping
Waller(1961)describeshowdifferentpatternsofstandingwavesareproducedonasteelplate.In
Waller'stechniquetheresonanceisenergizedbyapieceofdryicepressedagainsttheplate,wherethe
gasgeneratedbysublimationoftheiceproducesagapbetweentheiceandtheplatethatopensand
closesperiodicallyasthegasescapes.Thesignificantpropertyofthismethodofenergizingthe
oscillationsisthatitdoesnotforcetheoscillationsatanyparticularfrequency,butallowsthenatural
resonanceoftheplatetodeterminetheperiodofthevibration.Asimilarprincipleisseeninmusical
instruments.Forexamplethesoundofatrumpetisenergizedbythepulsingofthetrumpeter'slips.
Howeverthatpulsingistriggeredbythesoundwavesreflectedbackfromthefarendofthetrumpet,

sothefinalresonanceisdeterminednotbythelocaldynamicsofthelips,asmuchasbytheresonance
ofthetrumpet/lipsystemasawhole,witheverypartoftheresonantcavitycontributingtothefinal
oscillation.Wallerdescribeshowhigherharmonicpatternsareachievedbypressingthedryiceharder
againsttheplate,justasatrumpetercanjumptoahigheroctavebypursinghislipsandblowing
harder,whichinturndoublesthespatialfrequencyofthestandingwavepatterninthebodyofthe
trumpet.ThefullrangeofChladnifiguresisobtainedbydampingtheplateatvariouspoints,either
withthetouchofafingerorbyrestingtheplateonrubberstudsorridges.Thedampedpointsrestrict
thestandingwavepatternstothosethatexhibitstationarynodescoincidentwiththedampedpoints.
Thisisthesameprincipleusedinaflute,whereanopenholedampsthevibrationofairatthatpoint
inthetube,allowingonlystandingwavepatternstoemergethathaveanodeatthelocationofthe
openhole.Thepatternofdampingcanthereforebeseenascorrespondingtotheinputpatterninthe
receptivefieldmodel,becausethepatternofdampingcallsupacorrespondingpatternofstanding
wavesthatmatchesthepatternoftheinput,assuggestedinfigure4C,wheretheplateisshown
restingonrubberridgesthatmatchthenodesofthethreestandingwavepatternscorrespondingtothe
threeresonators.Anonspecificenergizationoftheplatewhilerestingontheseridgesthereforecalls
upthestandingwavepatternthatmatchesthepatternofdamping,andthatstandingwaveinturn
activatestheresonatortunedtoitscharacteristicfrequency,whichrepresentstherecognitionofthe
patternofdampingpresentontheplate.
7.8Emergence,Reification,Multistability,andInvariance
Thereareseveralsignificantdifferencesbetweenthereceptivefieldortemplatemodelofrecognition
andthatoftheHarmonicResonancemodel.FortheHarmonicResonancemodelautomatically
exhibitstheGestaltpropertiesofemergence,reification,multistability,andinvariance,notas
specializedcircuitsormechanismscontrivedtoproducethoseproperties,butasnaturalpropertiesof
theresonanceitself.EmergenceintheHarmonicResonancemodelisseeninthefactthatthereisno
needforasetofspecializedspatialreceptivefieldsdevisedtomatchallofthepatternstowhichthe
systemistuned,becausethespatialpatterningmechanisminharmonicresonanceoccursinemergent
fashioninthenaturalvibrationalmodesofahomogeneoussteelplate.Thisdefinesabasissetof
geometricalpatternsinanascendingorderofcomplexity,correspondingtotheeigenfunctionsofthe
plate.Thetunedresonatorswhichrespondtothesevibrationalmodesareverysimpledevices,which
couldeitherbetunedindividuallytotheharmonicsoftheplate,ortheycouldbedevisedtoadjust
theirowntuningadaptivelytomatchthetemporalfrequenciescommonlyproducedbytheplateto
whichtheyarecoupled.Emergencethereforeoffersasimplemechanismwhosedynamicbehavioris
verymuchmorecomplexthanitsarchitecturewouldsuggest.Forthesimpleplateandresonator
systemisfunctionallyequivalenttoamuchmorecomplexneuralnetworkmodelwithspecial
patternedreceptivefieldstunedtodetecteverypatterntowhichthesystemresponds.Itisthe
emergentnatureofthiscentralmechanismofpatterndetectionthataccountsfortheotherGestalt
propertiesofthesystemi.e.reification,multistability,andinvariance.
Reificationoccursautomaticallywithnoadditionalmechanismrequired,forthevibrationofthe
resonatoratitscharacteristicfrequencyautomaticallyregenerateitscharacteristicpatternbackonthe
plate.Thereforeifanoisyorirregularorincompletepatternofdampingispresentedontheplate,the
resonanceresultingfromthatinputpatternwillsetupthenearestmatchingstandingwavepattern,
whichinturnwillactivatethecorrespondingresonator.Thevibrationofthatresonatorinturnwill
reifyorcompleteitspatternbackontheplate,automaticallyfillinginanymissingfeatures,as
suggestedinfigure5A.ThispropertyoftheHarmonicResonancemodelcorrespondstothe
perceptualtendencytoperceivecompleteobjectsevenwhenportionsofthemareoccluded.

Figure5
A:Inthecaseofnoisyorincompleteinputpatterns,thesystemautomaticallyfillsinor
completesthemissingportionsofthepattern,whileregisteringtheidentityoftherecognized
patternintheresonatorresponse.B:Inthecaseofambiguouspatternsthesystembecomes
multistablebetweenalternativeinterpretations.C:Evenforsimpleinputsthesystemcanbe
multistablewhenstimulatedtohigherenergies,interpretingtheinputwithanystandingwave
thatcontainstheinputasasubset.Thisexcludesthepatternontheright.D:Thesystemis
invarianttoelasticdeformationsoftheinputpattern,aslongastheglobalgestaltispreserved.
MultistabilityisalsoanintrinsicpropertyoftheHarmonicResonancemodel,whichnotonlymakes
themodelconsistentwiththepropertiesofperception,butthemultistabilityitselfprovidesinteresting
functionalpropertiestotherecognitionsystem.Figure5Bdepictsadampinginputpatternwhich
mightbeinterpretedaseitherhalfofan"X"pattern,orthecentraldiagonalofthe"/"pattern.Inthe
presenceofthisinputthereforethesystembecomesbistablebetweenthesetwostandingwave
interpretations.Theactivationofthetworesonatorswillalsoalternateinsynchronywiththe
alternationofpatternsontheplate,i.e.therecognitionoftheidentityofthepatternsalternateswith
thepatternsthemselves.Anyadditionalevidencepresentintheinputwillbiasthiscompetition.For
exampleifoneoftheundampedcornersoftheplateislightlydampedwiththetouchofafingertip,
thiswillimmediatelyfavorthe"X"featureinterpretationwhichisdampedalongthatdiagonal,and
thattinybiasfactorresultsinthecomplete"X"featurepatternbeingreifiedontheplate.Thisis

consistentwiththeobservedpropertiesofbistablestimulisuchastheNeckercube,inwhichevery
portionoftheperceptappearstoinvertwiththeinversionofthespatialinterpretationofthestimulus,
suggestingacompleteinversionofahighresolutionlowlevelrenditionoftheperceptualexperience,
ratherthantheflippingofapairofhigherlevelabstractfeaturerecognitionnodes.TheNeckercube
canalsobestabilizedinoneortheotherstatebyprovidingtinygapsinthelineswheretheycross,
suggestingoneedgeoccludinganother,andthistinyvariationinonesmallportionofthefigurewill
stabilizetheentireperceptualstructure.
Multistabilityhasfurtherimplicationsforthepropertiesofharmonicresonanceasarecognition
system.Forinfacttheresponseofthesystemtoagiveninputisnotconfinedtoasinglestanding
wavepattern,butasWallerexplains,higherharmonicsofthatpatterncanalsobeevokedbypressing
thedryiceharderagainsttheplate,i.e.drivingthesystemtohigherenergies,justasatrumpeter
jumpstohigheroctavesbypursinghislipsandblowingharder.Forexampleiftheplateisdampedin
thepatternofan"X"assuggestedinfigure5C,thelowestenergystandingwaveinresponsetothis
inputwouldbetheonewiththe"X"shapedpatternofnodesasdescribedabove.Howeveriftheplate
isexcitedtohigherenergystatesbypressingtheiceharderagainsttheplate,thenpatternsofhigher
harmonicstandingwavescanemergeinresponsetothesamestimulus,includingthefourpatterns
depictedtotherightinfigure5C(amongothers),becauseallofthesepatternsshareincommonthe
"X"shapednodalpatternacrossthediagonalsofthesquare.Thestandingwavepatterndepictedon
thefarrightinfigure5Contheotherhandcouldnotappearinthepresenceofthe"X"shapedinput,
becausethatpatternrequiresthefreedomtooscillatealongoneofthediagonalswhicharedampedby
theinputpattern.Theinputpatternthereforedoesnotcalluptheoneandonlyencodedpatternto
whichitmatchesbestasinthecaseofthetemplatemodel,butratherthesystemtendstocallupany
ofthemanyencodedpatternsofwhichtheinputisasubset.Inthepresenceoftheinputthereforethe
systemismultistable,withthelowestenergystaterepresentingthesimplestencodedpatternthat
matchestotheinput,butwithadditionalenergythesystemcanbemadetomatchmorecomplex
patternstotheinput,aslongasthosepatternscontaintheinputpatternasasubset.Thiskindof
systemcanthereforebemadeto"searchthrough"itscatalogofencodedpatternsforanygiveninput,
byenergizingtheoscillationstogreaterorlessermagnitude,correspondingtopressingtheiceharder
orsofteragainsttheplate.Aseachpatternismatchedagainsttheinput,thatpatternisreifiedonthe
plateinfullspatialform,andasthatpatternappearsontheplate,itscorrespondingresonatorbecomes
energized,therebylabelingthespatialpatternwiththeidentityencodedbytheresonator.Thelowest
energypatternsarerepresentedbythelowerharmonics,whichdefinethesimplestorleastelaborate
patterns.Thesimplestpatternsarethemoststable,andthereforearethemostlikelytobeperceived,
correspondingtotheGestaltnotionofprgnanz,or"Gestaltgoodness".
Invarianceisalsoanaturalpropertyofaharmonicresonancerepresentation,whichisadirect
consequenceoftheemergentnatureoftheencoding.Forthespatialpatternisencodednotasarigid
template,butasacomplexdynamicinteractionbetweenelementsinacontinuousresonatingsystem.
Recognitionisthereforeinvarianttoelasticdeformationsofthepatternontheplate,aslongasthey
maintaintheiressentialglobalstructure.Forexamplethestandingwavepatternsshowninfigure5D
subdividetheplateintothesamenumberofregionswithequalareaasthoseinfigure5A,and
thereforetheseresonanceswillactivatethesamefeaturedetectorsdespitethedistortioninthepattern.
Theissueofinvarianceinrecognitionisoftenconfusedintheoreticaldiscussionswithablindnessto
variation,i.e.asystemisconsideredinvarianttoacertainstimulusvariationifitsresponseisidentical
acrossthatvariation.Howeverinvarianceinperceptiondoesnothavethatcharacter,forapatternthat
variesforexamplethroughrotation,translation,orscale,isrecognizedasthecharacteristicpattern,
andatthesametimeitisperceivedtoberotated,translated,orscaled.Althoughtherecognitionitself
isinvarianttothesetransformations,theperceptisobservedtoreflectthosestimulusvariations.This
suggestsatwolevelresponse,withinvarianceatthehigherlevelthatisneverthelesscoupledtoa
lowerlevelreifiedrepresentationinwhichthosevariationsareplainlyevident.Thisisexactlywhat
occursintheHarmonicResonancemodel,becausethepatterncompletionoccursinthereified
representationwithrespecttothedeformationsofthepatternasseeninfigure5D,whilethehigher
levelrecognitionrepresentedbytheresonatorresponseremainsinvarianttothosedistortions.This

uniquepropertyofharmonicresonance,soclearlymanifestinthesubjectiveexperienceofvision,is
virtuallyimpossibletoaccountforwithaneuralnetworkmodelduetotherigidtemplatelikeproperty
oftheneuralreceptivefield.
7.9RotationInvarianceinRecognitionandCompletion
Invarianceofanevenmoreimpressiveformisobservedinstandingwavesgeneratedinacircular
symmetricresonator.Figure6A(adaptedfromWaller1961pp.9and27)depictssomeofthe
standingwaveChladnifiguresthatcanbeproducedonacircularsteelplateclampedatitsmidpoint,
sortedbythenumberofdiametersandconcentriccirclespresentinthepattern.(Thepatterns[0,0]and
[0,1]expressedintermsof[diameters,circles]arenotactuallypossibletoproduceonasteelplatefor
technicalreasons,andaredepictedherefortheoreticalcompletenessoftherepresentation.)These
patternsareshownwithonenodelineorientedvertically.Howeverasaconsequenceofthesymmetry
oftheplate,thesesamepatternscanactuallyoccuratanyorientation,whilemaintainingthesame
frequencyofvibration.Figure6B(adaptedfromWaller1961p.9)plotsthevibrationfrequencyof
someofthesepatternsasafunctionofthenumberofdiametersinthepattern(correspondingtothe
columnsinfigure6A),andthenumberofconcentriccircles(correspondingtotherowsinfigure6
A).Thefrequenciesareexpressedasmultiplesofafundamentalfrequency.Forexamplethe"X"or
"+"shapedpattern[2,0]withtwodiametersandnocircles,hasavibrationfrequencyof1,whereasthe
"asterisk"pattern[3,0]hasavibrationalfrequencybetween2and3.Wallernotesthatnotwo
frequenciesareexactlyequal,andthatthereforethefrequencyuniquelyencodestherangeofpossible
patterns.Thediagramcanbeextendedindefinitelytogreaternumbersofdiametersandcircles,
althoughthehigherharmonicsrequireeverincreasingenergyofvibration.

Figure6

A:Chladnifiguresforacircularplate,sortedbythenumberof[diameters,circles]ineach
pattern.Thesepatternscanappearatanyorientationontheplate.Eachdistinctpatternhasa
uniquevibrationfrequency,plottedinB.Thevibrationfrequencythereforeoffersarotation
invariantrepresentationofthepatternpresentontheplate.
Therotationinvariancerelationbetweenthespatialpatternofstandingwavesandthecorresponding
vibrationfrequencyissignificant.Foritmeansthatinthepresenceofan"inputpattern",i.e.apattern
ofdamping,theplatewillresonateatthefrequencycorrespondingtothatpattern,andthatfrequency
willremainunchangedastheinputpatternisrotatedtoanyangle.Abankofresonatorstuned
specificallytothefundamentalfrequenciesofvibrationoftheplatethereforeencodearotation
invariantrepresentationofthosepatterns.Theuniquepropertyofharmonicresonanceisinthe
couplingbetweenthefrequencyanditspattern.Forifaresonatorisactivatedtopdown,i.e.inthe
absenceofanyinputstimulus,theactivationofthatresonatorinthevicinityoftheplatewilltendto
regenerateitscorrespondingpatternontheplate.Iftheresonatorisactivatedataloweramplitudein
theabsenceofanyinputstimulus,thiswillresultinanindeterminatepatternontheplate,sincethe
patternisattemptingtoreifyitselfatallorientationssimultaneously,assuggestedinfigure7A.
Howeverifthetopdownactivationoftheresonatorisappliedatsufficientlyhighamplitude,the
patternwillemergeontheplateatarandomorientation,andsincetheorientationoftheemergent
patternisunconstrainedbythetopdownsignal,thereifiedpatternwillbefreetospinontheplatelike
acompassneedleassuggestedinfigure7B.

Figure7
Reificationthroughrotationinvariance.A:Withaweaktopdownprimingofa"+"feature,the
reificationisindeterminate,asthepatternattemptstoreifyitselfatallorientations
simultaneously.B:Withastrongerprimethepatternisreifiedatsomearbitraryorientation,
butremainsfreetospinlikeacompassneedle.C:Atouchontheplatewithafingerbreaksthe
symmetryandlocksthereifiedpatterntoanorientationthatmatchesthedampedpoint.D:A
noisypatternisreifiedandcompletedattheorientationthatbestmatchestheprimedpattern.
Inthepresenceofaweakorpartialinputappliedsimultaneouslytoamoderatetopdownprime,the
inputwillanchortheorientationofthepatternprojectedtopdown,resultinginabottomup/top

downresonance,inwhichtheinputpatternisinterpretedandfilledininthecontextofthecurrent
topdownprime.Forexampleatopdownprimingofthecircularplatewithan"X"featureoscillation
couldpotentiallyregeneratethat"X"patternatanyorientation.Atouchofafingerontherimofthe
plateduringthisprimingwouldfixthat"X"patternatanorientationwhereonenodelinecoincides
withthedampedpoint,assuggestedinfigure7C.Thestandingwavemodelalsomirrorsseveral
propertiesofrecognitioninthebottomupmode.Intheabsenceofatopdownprimingsignal,anoisy
orambiguousinputpatternwillstimulatetheappearanceofthenearestmatchingpatternencodedin
theharmonicrepresentation,andthatbestmatchingpatternwillbereifiedontheplate,withany
missingdetailsfilledin,assuggestedinfigure7D.Itisthispropertyofbottomuptopdown
matchingacrossaninvariancerelationthatrepresentthegreatestpromiseoftheHarmonicResonance
theoryasarepresentationofperceptualprocesses,foritnowprovidesamechanismfortheinvariance
observedinperception.
7.10DynamicPatternFormation
Oneofthemostcompellingfeaturesofaharmonicresonancerepresentationisthatthesame
mechanismthatdefinesstaticpatternsofstandingwavesisalsocapableofgeneratingcomplex
dynamicpatternsofthesortthataremostlikelyresponsibleforthesinusoidaloscillationsinfishand
snakes,andtheperiodiccyclicpatternofmotionofthecentipede'sfeet.Forsuchcomplexpatterns
canbegeneratedbyinterferencebetweenspatialwavesdrivenatslightlydifferentfrequencies,
resultinginacyclicrotationofthestandingwavepatternataratethatisproportionaltothefrequency
differencebetweenthecomponentwaveforms.Inotherwordstheorientationandrateofrotationofa
spatialstandingwavepatterncanbecontrolledbythephaseshiftbetweencomponentdriving
oscillations,inthemanneroftherotatingLissajoufiguresonanoscilloscope,generatedbyplotting
twosinusoidsagainsteachotherinxandyaxesonthescope.AswiththeLissajoufigures,complex
dynamicfigurescanbegeneratedbyadditionofmorecomponentwaveformswhichcanresultin
endlesscombinationsfromasmallsetofpatternprimitives.
7.11HarmonicResonanceandBinding
TheHarmonicResonancetheoryfinallyprovidesapromisingcomputationalprincipletoaccountfor
theunityoftheconsciousexperience,foritisintheverynatureofresonancesindifferentresonators
tounitewhentheresonatorsarecoupled,toproduceasinglecoherentcoupledoscillationofthe
systemasawhole.Theindividualoscillatorsthatmakeupthecoupledsystemhaveamutual
influenceoneachother,eachoneinducingtheotherstomatchtoitsownoscillation,resultingina
singlecoherentglobaloscillationstate,asdescribedbyDewan(1976)andbyStrogatz&Stewart
(1993).
Whentheoscillatorswhicharebeingcoupledareintheformofspatialresonators,likeaChladni
plate,thecouplingbetweendifferentplateswillnotonlybindtheminsynchronousoscillation,but
thatsynchronyinturnwillgeneratesimilarpatternsonthecoupledplates.Thespatialresonatorin
effecttransformsthespatialpatternofitsstandingwaveintoahighlystructured,butonedimensional
temporalwaveform,whichiscommunicatedtotheotherresonatorsthroughthecoupling.Theother
resonatorsinturntransformthattemporalsignalbackintoaspatialpatterndefinedbystandingwaves.
Buttheinfluenceisreciprocalthroughthecoupling,sotheresonanceintheotherplatesalsoget
communicatedbacktothefirst,lockingthespatialpatternsonthecoupledplatestoeachother.Ifthe
perceptualinformationindifferentcorticalmapsisexpressedintermsofstandingwavepatterns,the
couplingbetweenthesemapswouldsynchronizethespatialpatternswithinthemlikethepictureson
anarrayoftelevisionsetsinashopdisplaywhicharealltunedtothesamechannel,exceptthatthe
synchronyofthecorticalmapsisnotduetothefactthattheyareallpresentingidenticalcopiesofthe
samesignal,butbecauseeachcontributesitsowninfluencetoapatternofresonancethatissharedin
commonamongthosemaps.

Itisalsointhenatureofharmonicresonancetoformmultipleidenticalorsimilarcopiesofa
particularwaveform,replicatedinmirrorreversedsymmetry,asseeninmanyofthepatternsinfigure
1.Forapatternwhichissymmetricalaboutadiagonal,forexample,thepatternsofresonanceon
oppositesidesofthediagonalcanbeconsideredastwoseparatecopiesofthesameresonanthalf
patternthatarecoupledtoeachotherinasingleresonance.Neurophysiologicalmappingofthe
somatosensorycortex,whichwasoriginallythoughttodefineasimplesomatotopicmap,hason
closerexaminationrevealedmultiplecopiesofthebodymapofteninmirrorreversedpatterns(Kolb
&Whishaw1980,p.176)suggestiveofastandingwaverepresentation.Nowidenticalcopiesofa
corticalmaparenotveryuseful.Howeverifthedynamicpropertiesoftheresonatingsubstratewere
slightlydifferentindifferentcorticalregions,specializeddynamicpropertiesofeachdifferentcortical
areawouldenhanceoremphasizespecificaspectsoftheresonanceinthoseareas,whilemaintaininga
dynamiccouplingtootherareaswithslightlydifferentdynamicproperties.Howeverthespecial
featuresenhancedinonebrainareawouldnotremainisolatedtothatarea,butrathertheeffectsofany
resonanceinanybrainareawouldimmediatelymodulatetheresonanceinallotherareas
simultaneously,asseeninthephenomenonofaudiofiltering,aswhenvariousacousticalcavitiesare
coupledtoeachother.ThisprincipleisalsoobservedinanalogRFcircuits,wheretheadditionof
differentcomponentsatdifferentpointsinthecircuitdoesnotprocessthesignalinaninputoutput
manner"downstream"oftheaddedcomponentasindigitalcircuitry,butratheritmodulatesthesignal
inthecircuitasawhole,enhancingcertainaspectsofthewaveforminallofthecomponentsofthe
circuitsimultaneously,althoughtheexactwaveformobservedwithineachcomponentexhibitssubtle
variations.

8Conclusion
Theharmonicresonancetheoryisnotafullyspecifiedtheoryofneurocomputation,butaparadigm,
i.e.asetofassumptionsastothefundamentalprinciplesbehindbiologicalcomputation,asa
challengetothemoreconventionalconceptofneurocomputationembodiedintheneurondoctrine.
Whetherornotthisprincipleisactuallyoperationalinthebrainisasubjectforfutureinvestigation,
andforreexaminationofexistingdatainthelightofthenewhypothesis.Infacttheremayalreadybe
considerableevidenceavailabletootherresearcherswhichmighteithersupportorrefutethe
hypothesis,butthatevidenceremainsunpublishedbecauseitssignificanceisonlyrevealedinthelight
ofthenewhypothesis.Itisforthisreasonthateveryparadigmatichypothesisdeservesatthevery
leasttobeexposedtothelargercommunity,aslongasitcanbeshowntobetrulyuniqueandoriginal,
andnoteasilyrefutedbytrivialarguments.
Agreatdealofworkremainstobedonebeforetheharmonicresonancetheorycanbeconsidereda
realneurophysiologicaltheorythatmakestestablepredictions.Inthefirstplacethemodelwillhaveto
bespecifiedingreaterdetail,todefinethespecificrepresentationalcodeofvisualneurons,whichcan
bematchedagainstactualneurophysiologicalrecordings.Fortheharmonicresonancetheorysuggests
thatthesignificantsignalinthebrainisnotcarriedbythepowerfulspikesoftheactionpotential,but
byamoresubtleandpervasivegradedpotentialoscillation,asrecordedbyPribram(1971)andBland
etal.(1978).Thetheorywillhavetobeelaboratedtoidentifyexactlywhichsystemsorcircuitsare
resonatinginthebrain,howthoseresonancesaresustained,howtheyaremodulatedbysensoryinput,
andhowthesubjectivevariablesofperceivedcolor,shape,andformareencodedinthestandingwave
representation.Theharmonicresonancetheorywillalsohavetoexplainwhycertaincorticalneurons
seemtobehaveasiftheywerefeaturedetectors,andtoprovideanalternativefunctionalexplanation
forthatobservedbehavior.
Evenasaparadigmatichypothesis,thepresentationoftheprinciplesofharmonicresonanceinthis
papermightseemunsatisfactorytoscientistsaccustomedtoamorerigorousmathematical
specificationofthefunctionalprinciplesofaproposedmodel.Howevertherearecertainclassesof
physicalsystemswhichsimplydonotsuccumbtomathematicalanalysisbecausethephenomenain
questionarealreadythesimplestmodelofthemselves,i.e.thereisnowaytoreducethephenomenato

simplermathematicaltermswithoutsettingprohibitivelyrestrictiveconstraintsontheparametersof
thesystem.Inallbutthesimplestcases,harmonicresonancehasexactlythisproperty.Anditisthat
verypropertythatrepresentsthemostinterestingaspectsofharmonicresonanceasarepresentationin
thebrain.Chladni(1787)andWaller(1961)bothreferenceworkbyvariousauthorswhoprovide
mathematicalsolutionstotheequilibriumstatesofvariousstandingwavepatternsfoundonsteel
plates.Buttheseanalysesareanabstractionorreductionistdescriptionoftheactualmechanismof
harmonicresonance,whichisactuallyafinegrainedprocessinvolvingmolecularinteractions
throughouttheresonatingsystemresultinginglobaleffects.Themathematicalsolutionsforsteel
plateshaveonlybeendevelopedtoaccountforthesimplestofthesepatterns,likethoseforacircular
orsquareplate.Thiskindofanalysisbecomesintractableinthecaseofirregularorarbitraryshaped
steelplates,orplatesofnonuniformthickness.Andyetitisthisveryflexibilitythatrepresentsthe
mostinterestingaspectoftheresonance.Ifthediscussionwerelimitedtothesimplecasesforwhich
analyticalsolutionshavebeenfound,thetheorywoulddegeneratetoatemplatetheory,andthereby
loseitsmostappealingproperties.
Physicalsystemsthatdefymathematicalcharacterizationareoftenaddressednumerically,using
computersimulations.Thisistheapproachusedforexampletomodelthebehavioroftheatmosphere,
whichisapproximatedbyquantizationofthesysteminspaceandtimetotinylocalelementswhich
aresimpleenoughtobetreatedasasinglepoint.Howeverthisapproachtoohasitslimits,forthe
quantizationinspaceandtimeinevitablyintroducesinhomogeneitiesintothesystem.Thecaseof
harmonicresonanceisevenmoreproblematicfornumericalsimulationsthanamodelofthe
atmosphere,becauseatmosphericparameterssuchastemperature,pressure,andhumiditytendto
diffuseisotropicallyintoadjacentregionsinarelativelysimplemanner,sothattheseparameterscan
becomputedfairlyaccuratelybyconsideringonlynearestneighboringregions.Inthecaseofstanding
wavepatterninaresonatingsystemontheotherhand,thelocalvalueofthepatterndependsonthe
configurationoftheentireresonatingsystemasawhole,andcannotbeevenapproximatedfromthe
valuesofadjacentregionsofthesystem,forresonancedoesnotsimplydiffusebetweenadjacent
pointsinthesystem,butisinfluencedbytheentireconfigurationofthesystemasawhole,as
evidencedbythefactthatatouchoftheplateatanypointwillcompletelychangethestandingwave
patternacrossthewholeplate.
TheveryfactthatharmonicresonanceexhibitsthisGestaltlikenature,andthefactthatthesekindsof
systemsaresodifficulttocharacterizebothmathematicallyandcomputationally,providesallthe
morereasontoinvestigatethesephenomenaasapossiblepropertyofbiologicalcomputation,to
accountforexactlythoseaspectsofperceptionwhichhavedefiedcharacterizationinmorerigorous
analyticalterms.Ifthisworkmustbedone"byanalogy"usingvibratingsteelplates,thatdoesnotin
anywayinvalidatetheresults,forthevibratingsteelplateisitselfacomputationalmechanism,albeit
onewhoseoperationalprinciplesareradicallydifferentfromanyknowncomputationaldevice,and
thereforethe"output"ofthesteelplateisnodifferentinprinciplefromtheoutputofacomputer
simulation,aslongasthesamephenomenacanbereplicatedbydifferentinvestigators.Thefactthat
resonanceexhibitssimilarpropertieswhetherexpressedasaphysical,electrical,chemical,oracoustic
resonanceshowsthatresonanceisageneralprinciplethattranscendsanyparticularphysical
instantiation,andtherebyrepresentsahigherorderorganizationalprincipleofphysicalmatter.
Consideredasacomputationalparadigm,harmonicresonancehasuniqueemergentpropertiesthat
cannotbemeaningfullyreducedtoanequivalentTuringmachinedescription.ThemessageofGestalt
theoryisthatitisexactlythiskindofunconventionalenigmaticphysicalphenomenonwhichshould
besoughtoutasthefunctionalprinciplebehindbiologicalcomputation.
Finally,theharmonicresonancetheoryalsooffersanexplanationforoneofthemostenduring
mysteriesofhumanexperience,whichisthequestionofwhyresonancesinmusicalinstrumentsand
therhythmicbeatingofdrumshavesuchapowerfulabilitytoevokethedeepestemotionalresponse
inthehumansoul.Iproposethatthemusicalinstrumentrepresentsman'sfirstmodestsuccessat
replicatingthephysicalprinciplebehindbiologicalcomputation,andthestrongemotionalresponse
evokedbytheseinanimateresonancesreflectsanunconsciousrecognitionoftheessentialaffinity

betweenmindandmusic.

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