Maxim I. Stamenov, Vittorio Gallese. Mirror Neurons and the Evolution of Brain and Language


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Tableofcontents
Introduction
I.Mirrorneuronssystem
Past,present,andfutureofadiscovery
Theneuralcorrelatesofactionunderstandinginnon-humanprimates
LeonardoFogassiandVittorioGallese
Themirrorsysteminhumans
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
II.Furtherdevelopmentsinthestudyofmirrorneuronssystem
Tableofcontents
Mirrorneuronsandtheselfconstruct
KaiVogeleyandAlbertNewen
Behavioralsynchronizationinhumanconversationalinteraction
151
JenniferL.RotondoandStevenM.Boker
Tableofcontents
Lookingforneuralanswerstolinguisticquestions
323
BernardH.Bichakjian
Mirrorneuronsandculturaltransmission
IndiaMorrison
IV.Applications
Mirrorneuronsandtheneuralbasisforlearningbyimitation:
Computationalmodeling
343
AudeBillardandMichaelArbib
Mirrorneuronsandfeedbacklearning
353
SteveWombleandStefanWermter
Aconnectionistmodelwhichuniesthebehavioraland
thelinguisticprocesses:Resultsfromrobotlearningexperiments
363
YuuyaSugitaandJunTani
Nameindex
377
Subjectindex
385
Introduction
readyexistingneuralnetworks.WhatisreallychallenginginconsideringMirror
NeuronsSystem(MNS)inthisrespectisthatthere-organizationanddevelopment
ofnewfunctionssupportedbycorrespondingbraincircuitsseemstoin
uencethe
wayofperformanceofthesestructuresnotjustatthemacroscale,butalsoatthe
microscaleofsingleneuronsperformance.Herethelatestreportofthestudyof
mirrorneurons(cf.Fogassi&Gallese,thisvolume)speaksforitselfinshowingthe
scaleoffunctionalspecializationinthewayofperformanceofsingleneuronsin
differentareasofmonkey
sbrain.Althoughitisimpossibletostudyhumanswith
suchdirect-invasionmethods,onecouldinferthatMNSfunctions
both
thesame
way(asisthecasewithmonkeys)aswellasinamore
exibleway(asinthecase
ofhumans).Ifthisisthecase,theuniquenessofhumanMNSconsistsinitsca-
pacitytofunctionasacomponentstructureofacognitivemoduleandacentral
cognitivesystem.Thechallenge,then,wouldbefurthertodevelopnewtechno-
logicalmeansforanon-invasive
invivo
studyofsingleneuronfunctioninginthe
brain.Ininvestigatingthisaspectofbrainperformancewewilladdmore
esh
fromtheneurophysiologicalpointofviewtothepointmadebyWeigand(thisvol-
ume)thathumansarecomplexadaptivesystemsonascaleunprecedentedinthe
biologicalkingdom.Thecomparativestudyofthebehaviorofmirrorneuronsin
humansandmonkeyspromisestobecome,inthenearfuture,anevenmorechal-
lengingandcontroversialenterpriseforthewayweenvisagehowmentalfunctions
areimplementedbyneurophysiologicalprocesses.
TurningtothementalaspectofMNSdiscovery,we
ndnolesssurprising
consequencesofitforourunderstandingofconsciousandunconsciousfunctions
Introduction
Inthe
rstpartthediscoverersofmirrorneuronsaregiventhe
oortopresent
astate-of-the-artreportaboutthestatusofMNSinthebrainofmonkeys(Fogassi
&Gallese)andhumans(Rizzolatti,Craighero,&Fadiga).FogassiandGalleseillus-
tratethefunctionalpropertiesofmirrorneurons,discussapossiblecorticalcircuit
thataco-evolutionoflanguageandWMcapacityhastakenplaceinthehu-
manbrain.
Senkforfurtherdevelopsthepointaboutthe
wide-scalere-wiringofthebrain
circuitsinhumans
.Shereportsresultsofexperimentsshowingthatdifferentbrain
circuitsareengageddependingonthenatureofthepriorexperienceinperform-
inganaction,inwatchingtheexperimenterperformanaction,inimaginingan
action,orinacognitivecontroltaskofcostestimation.Innon-invasivescalpERP
Introduction
modulateandbemodulatedbymirror-likeactivationsinordertocoordinateself-
andother-generatedactions.
Usingfunctionalmagneticresonanceimaging(fMRI),McGlone,Howardand
Robertsinvestigatedtheneuralbasisofaproposedmirror(i.e.observation-action)
systeminhumans,inwhichparticipantsobservedanactorusingherrighthandin
graspingactionsrelatedtotwoclassesofobjects.Action-speci
cactivationwas
Thethirdpartofthisvolumeincludesarticlesdealingwiththeevolutionof
brain,languageandcommunication.Intheirarticle,LiandHombertofferan
overviewofthelast6millionyearsofhominidevolutionandasketchofadiverse
arrayofinformationfromdifferentdisciplinesthatarerelevanttotheevolution-
aryoriginoflanguage.They
rstdistinguishtheproblemoftheoriginoflanguage
fromtheproblemsassociatedwiththestudyofevolutionoflanguage.Theformer
isanenterpriseconcernedwiththeevolutionofthecommunicativebehaviorof
ourhominidancestors,nottheevolutionoflanguage.Thislatterconcernslinguis-
ticchangeandisthesubjectmatterofdiachroniclinguistics.Thusthestudyofthe
evolutionarychangeofcommunicationisnotastudyoflinguisticchange(within
certainhumanlanguagethatalreadyhasthecriticalfeaturesqualifyingitassuch
).LiandHombertdiscussasetoffundamentalproblemsrelatedtotheemer-
genceoflanguagecapacity,e.g.,theemergenceoflanguageandtheemergenceof
anatomicallymodernhumans,thefourevolutionaryprocessesleadingtotheemer-
genceoflanguage(thereductionofthegastrointestinaltract,theenlargementof
thevertebralcanal,thedescentofthelarynx,andtheincreaseofencephalization),
aswellasthethreeevolutionarymechanismsunderlyingtheemergenceoflan-
guage(theduplicationofHometicgenes,thechangeofthedevelopmentalclock,
andthecausalroleofbehaviorinevolution).Onthebasisofthisbroadbiologi-
calbackground,theyproceedwiththeconsiderationofthefoundationalaspects
ofsymboliccommunicationassuch.Heretheyintroduceacoreexplanatorycon-
cept
thatofcognitivereserve,bywhichtheymeanthecognitivecapabilitythatis
notfullyutilizedormanifestedinthenormalrepertoireofbehaviorofamammal.
LiandHombertalsodiscusssomeimportantstepstowardthe
crystallization
languageduringthehominidevolution.
Studdert-Kennedymakesthepointthattheunboundedsemanticscopeofhu-
manlanguagerestsonitsdualhierarchicalstructureofphonologyandsyntax
Introduction
loguefacialimitation(alsouniqueamongprimatestohumans)todifferentiation
ofthevocaltractanddigital(particulate)vocalimitation.
AtthebeginningofhercontributionWeigandmakestheimportantclaimthat
mirrorneuronsarenot
simplecomponents
butthemselvesimplementcomplex
unitsintegratingseveraldifferentdimensionsofneuralandcognitiveprocessing.
Itisfromacomplexintegratedwhole,theMNS,thattheevolutionofthelan-
guagefacultyhasstarted.Theconsequencesofpositingthishypothesisareex-
plicitlypointedoutanddiscussed.Thebasicpointtobemadeisthatthebasis
forlanguageemergenceshouldhavebeenagrammarofaratherdifferenther-
adialoguegrammar.Weigandoffersadescriptionofthecriterialfeatures
ofsuchadialoguegrammar,includingthoseofintentionality,socialmotivation
suggestthatverbalintelligenceandspeecharticulationaretwodifferentthings,and
theriseofthelatterdoesnotexplainthedevelopmentoftheformer.
Anderson,Koulomzin,BeebeandJaffereportthatself-groomingmanualac-
tivityamongfour-month-oldscorrelateswithincreaseddurationofattentivegaze
xationonthemother
sface.Eightfour-month-oldinfantswereselectedfrom
alargerstudyandwerecodedsecond-by-secondforinfantgaze,headorienta-
tionandself-touch/mouthingbehaviorduringface-to-faceplaywiththemother.
Amongtheseinfantsitwasfoundthatattentivehead/gazecoordinationiscontin-
gentuponself-touch/mouthingbehavior.Episodesofmutualgazeareespecially
prominentuptotheageof4months,beforewhichinfantsexhibitanobligatory,
automatictendencytoremaintotallygaze-lockedonthematernalface.Regular
Introduction
theambientlanguage.Oneofthecriticalsteps,inthisrespect,islearningtorepre-
sentspatialcategories(relatedamongothertoself-andother-enactedbehavioral
actions).McCunehasinmindhere,amongothers,thelearningandrepresent-
ingofthecognitivestructureassociatedwiththerelationalwordslikeverbsand
prepositionsinnaturallanguage.TothedegreeMNSsupportsand
represents
the
controlstructureofabehavioralaction(likegraspingasmallobject),itmayhave
SugitaandTanipresenttheirnovelconnectionistmodeldevelopedforthelin-
Mirrorneuronssystem
Past,present,andfutureofadiscovery
Theneuralcorrelatesofaction
understandinginnon-humanprimates
LeonardoFogassiandVittorioGallese
IstitutodiFisiologiaUmana,Universit
diParma,Italy
Introduction
Ineverydaylifewearecommonlyexposedtoactionsperformedbyotherindividu-
als.Dependingonthecontextorthecircumstances,wemaybewitnessingdifferent
LeonardoFogassiandVittorioGallese
theactionisnotovertlyexecuted,butsimplyimagined.Inotherwords,thesecir-
cuitscouldcontainthe
representation
ofthoseactionsthat,whentheinternaldrive
andthecontextaresuitable,areovertlyexecuted.Ifsuchmotor
representations
arepresentinthemotorsystem,theycouldbeautomaticallyretrievednotonly
whenweexecuteormentallyrehearseaspeci
caction,butalsowhenweobserve
thesameactionperformedbyotherindividuals.Thismechanismmayconstitute
thebasisforactionunderstanding.
Aneuralmechanismforunderstandingtheactionsmadebyothersisanec-
essaryprerequisitealsofornon-humanprimatessuchasmonkeys.Particularly
soforthoselivinginlargesocialgroups,inwhichindividualsneedtorecognize
Theneuralcorrelatesofactionunderstanding
Figure1.
Lateralviewofthelefthemisphereofastandardmacaquemonkeybrain.
LeonardoFogassiandVittorioGallese
Inthefollowingsectionswewillsummarizethevisualandmotorproperties
ofF5mirrorneurons.
VisualpropertiesofF5mirrorneurons
Mirrorneuronsdischargewhenthemonkeyobservesanotherindividual(ahuman
beingoranothermonkey)performingahandactioninfrontofit(seeFigure2).
Differentlyfromcanonicalneurons,theydonotdischargetothesimplepresen-
tationoffoodorofotherinterestingobjects.Theyalsodonotdischarge,ordis-
Figure2.
ExampleofthevisualandmotorresponsesofaF5mirrorneuron.Thebehav-
ioralsituationduringwhichtheneuralactivitywasrecordedisillustratedschematically
intheupperpartofeachpanel.Inthelowerpartrastersandtherelativeperistimulus
responsehistogramsareshown.
:Atraywithapieceoffoodplacedonitwaspre-
sentedtothemonkey;theexperimentergraspedthefoodandthenmovedthetraywith
thefoodtowardthemonkey,whichgraspedit.Astrongactivationwaspresentduring
observationoftheexperimenter
sgraspingmovementsandwhilethesameactionwas
performedbythemonkey.Notethattheneuraldischargewasabsentwhenthefood
waspresentedandmovedtowardthemonkey.
:As
,exceptthattheexperimenter
graspedthefoodwithpliers.Notethatonlyaweakdischargewaselicitedwhentheob-
servedactionwasperformedwithatool.Rastersandhistogramsarealigned(vertical
bar)withthemomentinwhichtheexperimentertouchedthefood.Abscissae:time.
Ordinate:spikes/bin.Binwidth:20ms.(Modi
Theneuralcorrelatesofactionunderstanding
A
B
20
20
LeonardoFogassiandVittorioGallese
Table1.
Mirrorneuronssubdividedaccordingtotheobservedhandactionseffective
inactivatingthem
Observedhandactions
No.ofneurons
Grasping
Manipulating
Handsinteraction
Holding
Grasping/Placing
Grasping/Manipulating
Grasping/Handsinteraction
Grasping/Holding
Grasping/Graspingwiththemouth3
Placing/Holding
Handsinteraction/Holding
Grasping/Placing/Manipulating1
Grasping/Placing/Holding
Total
92
totheobservationoftwoormoreactions.Observationofgraspingaction,alone
orassociatedtootheractions,isbyfarthemosteffectiveindrivingtheneurons
discharge.Amongneuronsrespondingtotheobservationofgraspingactionthere
aresomeveryspeci
Theneuralcorrelatesofactionunderstanding
monkey)orthemonkeyitselfexecutedaprecisiongriptograspasmallpieceof
food.Strictlycongruentneuronsrepresentabout30%ofallF5mirrorneurons.
broadlycongruent
wede
nedthoseneuronsinwhichthecodedobserved
actionandthecodedexecutedactionaresimilarbutnotidentical.Forexamplea
neuroncoulddischargewhenthemonkeyexecutedagraspingactionandwhen
itobservedanexperimentergraspingandtakingawayapieceoffood.Insome
casesthereiscongruenceaccordingtoalogicalor
causal
sense:forexample,a
neuronrespondedwhenthemonkeyobservedanexperimenterplacingapieceof
foodonatrayandwhenthemonkeygraspedthesamepieceoffood.Thetwo
actionscanbeconsideredtobepartofalogicalsequence.Broadlycongruentneu-
ronsrepresentabout60%ofallF5mirrorneurons.Finally,inabout10%ofF5
LeonardoFogassiandVittorioGallese
Mirrorneuronsandactionunderstanding
Thetriggeringfeaturethatevokesthemirrorneurons
dischargeisthesightof
ahand-objectinteraction.Formostmirrorneuronstheresponseisindependent
fromthehandusedbytheobservedagenttoperformtheactionandalsofromthe
orientationoftheobservedhand.Thedischargeispresentbothwhentheagent
handexecutingtheactionisseenfrontallyorfromasideview.Whatmattersisthat
40 cm
A
B
C
D
Theneuralcorrelatesofactionunderstanding
Figure3.
ExampleofaF5mirrorneuronrespondingtoactionobservationinFull
visionandinHiddencondition.
Thelowerpartofeachpanelillustratesschematicallytheexperimenter
sactionas
observedfromthemonkey
svantagepoint:theexperimenter
shandstartingfroma
xedposition,movingtowardanobjectandgraspingit(panelsAandB),ormimicking
grasping(panelsCandD).Thebehavioralparadigmconsistedoftwobasicconditions:
Fullvisioncondition(A)andHiddencondition(B).Twocontrolconditionswerealso
performed:Mimickinginfullvision(C),andMimickinghidden(D).Intheselasttwo
conditionsthemonkeyobservedthesamemovementsasinAandB,butwithoutthe
LeonardoFogassiandVittorioGallese
isgoingtopickupabook,evenifthebookisnotvisible.Fullvisualinformation
aboutanactionisnotnecessarytounderstanditsgoal.
Ifmirrorneuronsareindeedtheneuralsubstrateforactionunderstanding,
Theneuralcorrelatesofactionunderstanding
Acorticalcircuitforactionunderstanding
Mirrorneuronsareendowedwithbothvisualandmotorproperties.Whatisthe
LeonardoFogassiandVittorioGallese
The experimenter grasps (PG) and
releases the object with the left hand
The experimenter grasps (WH) and
releases the object with the left hand
The experimenter presents the object
with the left hand
The monkey grasps (PG) the object
with the hand
The monkey grasps (WH) the object
with the hand
The experimenter grasps (PG) and
releases the object with the right hand
Theneuralcorrelatesofactionunderstanding
PFmirrorneuronsrespondedduringtheexecutionofhand,mouth,orhand
Figure4.
ExampleofthevisualandmotorresponsesofaPFmirrorneuron.Thisneu-
ronstarted
ringabout300msbeforetheexperimenter
shandtouchedtheobject.The
dischargecontinueduntiltheexperimenter
shandtookpossessionoftheobject,ceased
duringtheholdingphase,andstartedagainduringthereleasingaction.Thisneuron
displayedaspeci
cityfortheobservedgrip:theobservationofgraspingachievedby
opposingtheindex
ngertothethumb(precisiongrip,PG),wasmuchmoreeffec-
tivethantheobservationofgraspingachievedby
exingall
ngersaroundtheobject
(wholehandprehension,WH).Thisselectivitywasreciprocatedbytheneuron
smo-
torselectivity:theneuron
sdischargewashigherwhenthemonkeygraspedtheob-
jectusingaprecisiongripthanwhenusingawholehandprehension.Notethatthe
LeonardoFogassiandVittorioGallese

Theneuralcorrelatesofactionunderstanding
ofahandandthemotorprogramcontrollingmouthgrasping.Oncethisequiva-
lenceisputinplace,amirrorsystemmatchinghandactionsobservationonmouth
actionsexecutioncanbeestablished.Sucha
primitive
matchingsystem,how-
ever,wouldbebene
cialalsoinadulthood,whenamoresophisticatedhand/hand
matchingsystemisdeveloped,inordertoprovidean
abstract
categorizationof
theobservedactions:whatisrecognizedisaparticularactiongoal,regardlessof
theeffectorenablingitsachievement.
ThirtypercentofPFneuronsrespondingtotheobservationofactionswere
devoidofmotorproperties(
actionobservationneurons
).Theirvisualresponse
wasverysimilartothatofPFmirrorneurons:theywereactivatedbytheobser-
vationofasingletypeoroftwoorthreetypesofhandactions.Theseneurons
areimportantbecausetheirhigherpercentagewithrespecttoF5(30%vs.22%),
probablyre
ectstheirproximitytoSTSaneuronssharingthesameproperties.
Inthelightofthese
ndingsweproposethatapossiblecircuitforactionun-
derstandingcouldberepresentedbythreecorticalareasofthreedifferentlobes:
LeonardoFogassiandVittorioGallese
Theneuralcorrelatesofactionunderstanding
theobject.Similarly,thevisualdischargeofmirrorneuronsisnotdirectlyfollowed
byamonkeyaction.Therefore,alsowhenanactionisnotdirectlyexecuted,thein-
ternalmotorcircuitgeneratesa
representation
ofit.Itisimportanttostressthatthis
LeonardoFogassiandVittorioGallese
newskillthatdevelopedbyexploitinginnewwaysresourcespreviouslyselected
formotorcontrol.
Theneuralcorrelatesofactionunderstanding
macaques,aschimpanzees,areabletofollowthemovementsofheadandeyes
andoftheeyesalone.Thesedatacorrelatewellwithphysiologicalstudiesshow-
LeonardoFogassiandVittorioGallese
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Themirrorsysteminhumans
GiacomoRizzolatti,LailaCraigheroandLucianoFadiga
IstitutodiFisiologiaUmana,Universit
diParma,Italy/Dipartimentodi
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
2020
Figure1.
Visualandmotorresponsesofarepresentativemirrorneuron.Testingcon-
ditionsareschematicallyrepresentedabovetherasters.Histogramsineachpanelrep-
resentthesumofeightconsecutivetrials.A,atraywithapieceoffoodispresented
tothemonkey,theexperimentergraspsthefood,putsthefoodagainonthetrayand
thenmovesthetraytowardthemonkeythatgraspsthefood.B,asabove,exceptthat
theexperimentergraspsthefoodwithpliers.C,activegraspingofthemonkeyinthe
dark.Thepresenceofadischargeduringthislastsituationdemonstratesthatthemotor
dischargeoftheneuronisnotduetomonkey
sobservationofitsownhand.
Themirrorsysteminhumans
1990).AlthoughthisissuewasnotsystematicallyaddressedSTSdonotappearto
dischargeduringactiveaction,oratleastthisphenomenon,ifpresent,isnotso
prominentasinF5.Regardlessofthisaspect,itisclearthatSTS,PFandF5forma
systemwherethebiologicalactionsaredescribedinvisualtermsandthenmatched
onmotorneuronscodingthesameaction.
Theaimofthepresentarticleistoreviewtheavailableevidenceontheexis-
tenceofamirrorsysteminhumans.Itisimportanttonotethatwhensingleneuron
recordingtechniqueisused,informationistypicallyobtainedconcerningasingle
brainareaorcenter.Thus,thefactthatuptonowonlyonemirrorneuroncircuit
hasbeende
nedinthemonkeydoesnotexcludetheexistenceofothermirrorneu-
roncircuits.Thispointisimportanttostressbecause,asitwillbeshownbelow,
circuitswithmirrorpropertiesappeartobemorewidespreadinhumansthenin
monkeys.Thisdifferencemaybeaspeciesdifference,butmostlikelyisaconse-
quenceofthedifferenttechniqueusedinmonkeys(singleneuronstudies)andin
humans(brainimagingtechniques).
Mirrorsysteminhumans:Neurophysiologicalevidence
rstsetofevidence,albeitindirect,infavorofamirrorsysteminhumanscomes
fromthestudyofthereactivityofthecerebralrhythmsduringmovementobser-
vation.TraditionalEEGstudiesdistinguishedtworestrhythmsbothinthealpha
range(8
13c/s):aposterioralpharhythmandcentralmurhythm.Thesetwo
rhythms,besidesdifferenttopography,havedifferentfunctionalsigni
cance.The
posterioralpharhythmispresentwhenthesensorysystems,thevisualoneinpar-
ticular,arenotactivated,anddisappearsatthepresentationofsensorystimuli.The
murhythmispresentduringmotorrestanddisappearsduringactivemovements
aswellasduringsomatosensorystimulation(seeChatrian1976).
InbothAandB,rastersandhistogramsarealignedwiththemomentatwhichthe
experimentertouchesthefoodeitherwithhishandorwiththepliers(verticalline).In
Crastersandhistogramsarealignedwiththemomentatwhichthemonkeytouches
thefood.Binwidth,20ms.Ordinates,spikes/bin;abscissas,time.
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
experimentsinwhichanon-biologicalmotion(e.g.awaterfall)wasshowntothe
recordedsubjectsdidnotdesynchronizetherhythms.Thustherhythmsthatare
Themirrorsysteminhumans
Therationaleoftheexperimentwasthefollowing:ifthemereobservationof
thehandandarmmovementsfacilitatesthemotorsystem,thisfacilitationshould
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
Mirrorsysteminhumans:Brain-imagingstudies
Theneurophysiologicalexperimentsdescribedabove,whilefundamentalinshow-
ingthatactionobservationelicitsaspeci
c,coherentactivationofmotorsystem,
donotallowthelocalizationoftheareasinvolvedinthephenomenon.Dataon
thelocalizationofhuman
mirrorsystem
havebeenobtained,however,using
brain-imagingtechniques.
rststudyinwhichthisissuewasaddressedwasratherdisappointing
Themirrorsysteminhumans
below)and,inthecaseofBroca
sarea,byitshomologywithmonkey
sareaF5.This
lastinferencewasrecentlystronglycorroboratedby
ndingsshowingthatBroca
areaisanareainwhichnotonlyspeechbutalsohandmovementsarerepresented
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
Figure2.
Lateralviewsofleftandrighthemispheresduringobservationofactions
madewiththemouth(square),hand(circle)andfoot(asterisk)aftersubtractionof
staticmouth,handandfootobservation,respectively.InAintransitiveactionsare
shown,inBobject-directed(transitive)actionsarerepresented.
observationofastaticface,astatichandandastaticfoot,respectively,ascontrol
conditions.TheresultsareshowninFigure2.
Duringnonobject-relatedaction(chewing)activationswerepresentinareas
6,44onbothsidesandinarea45intherighthemisphere.Righthemisphereacti-
vationswerelargerandstrongerthanlefthemisphereactivations.Duringobject-
relatedaction(biting)thepatternofpremotoractivationswassimilar,although
weaker,tothatfoundduringnonobject-relatedaction.Inaddition,twoactivation
Themirrorsysteminhumans
foundinthemouthmovementcondition.ThecaudalfocuswasagaininareaPG.
Thislastfocusconsiderablyoverlappedthatofmouthmovementcondition.
Duringtheobservationofmimickedfootactionstherewasanactivationofa
dorsalsectorofarea6.Duringtheobservationofobject-relatedactions,therewas
asintheconditionwithoutobjectanactivationofadorsalsectorofarea6.Inaddi-
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
B
F6
F2
F1
F4
F5
as
cs
cs
ips
PE
S1
VIP
LIP
AIP
PF
PFG
PG
Figure3.
Themirrorsysteminhumans
Figure4.
Lateralviewofhumanleftfrontallobe.Themultiplemovementsomatotopy
isindicatedbythenamesofinvolvedbodyparts.TheterminologyofFoerster(1936)
andVogtandVogt(1926)hasbeenadoptedtorepresentthecytoarchitectonicalsubdi-
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
Figure5.
LateralviewofhumanlefthemispheresshowingenlargedBrodmann
s(A)
andvonEconomo
Themirrorsysteminhumans
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
ization(Binkofskietal.1999).Duringcomplexobjectmanipulationanactivation
Themirrorsysteminhumans
Theassumptionatthebasisofthehypothesisthatmirrorsystemallowsaction
understandingisthatthesameneuronalpatternthatisendogenouslyactivatedfor
actionexecutionisalsoactivatedexogenouslyduringactionobservation.Thus,if
oneacceptsthenotionthattheactingindividual
knows
whatwillbetheresultsof
his/heraction,onehastoadmitalsothathe/shewillbeabletoknowtheoutcome
oftheobservedaction,beingthesamemechanisminvolvedinbothoccasions.
Theviewthatthemotorsystemplaysacrucialroleinactionunderstanding
doesnotdeny,ofcourse,thatactionsaredescribed,coded,and(possibly)recog-
nizedintheinfero-temporallobeand,especially,intheareaslocatedinthesu-
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
toemit,unspeci
cally,theresponsesappropriatetoit,imitationmayconcerna
movementorasequenceofmovements.Furthermore,imitationmaybepreceded
ornotbytheunderstandingoftheactionmeaning,itmaybeanapproximateor
aprecisereplicaoftheobservedactionand,
nally,itmayconcernasequenceof
Themirrorsysteminhumans
1999).Neurophysiologicalevidenceinfavorofthe
rsttypeofresonanceisscanty
inthemonkey.Someneurons,however,werefoundinPFthatdischargeduring
meaningless(intransitive)armmovementsaswellasduringobservationofsimilar
movements(seeGalleseetal.2002).Strongerevidenceinfavorofthismechanism
comesfromhumanexperiments.EEG,MEG,TMS,aswellasbrainimagingstud-
ies(seeabove),allshowthattheobservationofintransitivemeaninglessactions
activatemotorareas.
Responsefacilitationwithactionmeaningunderstanding.
Humanobserverstypi-
callyimitatemovementsmadebyotherindividuals,havinganunderstandingwhat
theotherindividualisdoing.Atthispoint,animportanttheoreticaldistinction
shouldbeconsidered:Leavingapartthesymbolicgesturesor
quasisymbolic
ges-
tures,suchasthearmmovementsinvitinganotherindividualtoapproachortogo
away,therearetwodifferenttypesofactionswithmeaning:
motoracts
actions
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
mostcircumstancesoflittleandinmanycaseevendangerousfortheobserv-
ingindividual.Imitationoccursthereforeforsocialreasonsorinordertolearn
fromothers.
Muchmorecomplexisthecapacitytoimitateamotoractionand,evenmore
so,asequenceofmotoractions.Aninterestinghypothesistoexplainhowthiscan
occurswasrecentlyadvancedbyByrne(inpress)inhisdiscussionofwhatherefers
toas
action-level-imitation
.Withthistermheindicatesthecopingofanaction
notpresentpreviouslyinthebehaviorrepertoireoftheobserver.Accordingtohis
suggestion,thisbehaviorcanbeimitatedbydissectingitintoastringofcompo-
nents,formedbysimplersequentialpartsthatarealreadyintheobserver
sreper-
toire.Speci
cally,thebehaviorobservedinanotherindividualcouldbeseenas
madeupofasequenceofsimpleelementsor,usingourterminology,ofmotor
acts.Thesystemofmirrorneuronswouldprovidetheneuralbasisforrecognizing
Themirrorsysteminhumans
bolicorspatialimperativestimulus.Furtherevidenceinfavorofthenotionthat
duringactionpreparationtheindividualgeneratesaninternalcopyofthevisual
GiacomoRizzolatti,LailaCraighero,andLucianoFadiga
Binkofski,F.,Buccino,G.,Posse,S.,Seitz,R.J.,Rizzolatti,G.,&Freund,H.(1999).A
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observedandexecuted
ngermovements:comparingsymbolic,spatialandimitative
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Zilles,K.,Rizzolatti,G.,&Freund,H.-J.(2001).Actionobservationactivatespremotor
andparietalareasinasomatotopicmanner:anfMRIstudy.
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edelectroencephalograpy.
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Furtherdevelopmentsinthestudyof
Isthehumanbrainunique?
GerhardRoth
BrainResearchInstitute,UniversityofBremen/
HanseInstituteforAdvancedStudy,Delmenhorst,Germany
Humansareproudoftheirbrainandtheircognitiveabilities,andmanyofusin-
cludingmanyneuroscientistsbelievethattheallegeduniquenessofhumannature
isduetotheuniquenessofthehumanbrain.Inthefollowing,Iwillbrie
ydiscuss
somepopularclaimsaboutthehumanbrainthatcanbefoundeveninthescien-
cliterature.Theseare:(1)Thehumanbrainingeneralisanatomicallyunique;
(2)Humanshavethelargestbraininabsoluteterms;(3)Humanshavethelargest
brainrelativetobodysize;(4)Humanshavethelargestcerebralcortex,particularly
prefrontalcortex;(5)Humanshavesomebraincentersorfunctionsnotfoundin
otheranimals.
Firstclaim
GerhardRoth
Toothed
Man
Ape
Hare
Armadillo
Figure1.
Seriesofmammalianbrains,alldrawntothesamescale.Evidently,manhas
neitherthelargestbrainnorthemostconvolutedcortex.Convolutionofthecortexas
wellasofthecerebellumincreasesmonotonicallywithanincreaseinbrainsize.
Isthehumanbrainunique?
Amoredif
cultproblemisthepresenceofstructureshomologoustothe
Brainweightinmammals
[gram]
Spermwhale8,500
Chimpanzee400
Elephant
Man
Dog
Horse
Cat
Gorilla
550
Rat
Cow
Mouse
Thirdclaim
Humanshavethelargestbrainrelativetobodysize.Thisiswrong,
too.Whilethehumanbrainoccupiesabout2%ofbodymass,inverysmallrodents
GerhardRoth
relativebrainsizegoesupto10%.However,againamongprimates,humanshave
thelargestrelativebrainsize.
0,0010,010,1110100
10 000100 000
body weight (kg)
brain weight (g)
Figure2.
Isthehumanbrainunique?
brain weight (g)
body weight (kg)
shrew
squirrel
rhesus monkey
chimpanzee
african elephant
sperm
hippopotamus
dog
rat
shrew
0.00010.01110010,0001,000,000
Figure3.
relative brain weight (percentage ofbody weight)
body weight (kg)
shrew
squirrel
rhesus
monkey
chimpanzee
african elephant
sperm
hippopotamus
dog
rat
shrew
0.00010.01
10010,0001,000,000
Figure4.
Brainweightasapercentageofbodyweightforthesame20mammalian
species.Double-logarithmicgraph.FromNieuwenhuysetal.(1998),modi
ed.
GerhardRoth
body weight (kg)
Birds
Mammals
Reptiles
Cyclostomes
Osteichthyes
Chondrichthyes
Amphibians
brain weight (g)
0.0010.01
Figure5.
Isthehumanbrainunique?
List2.
Encephalizationinmammals.
Encephalizationquotientinmammals
Man
7.4Marmot1.7Cat
Dolphin
5.3Fox
1.6Horse0.9
Chimpanzee2.5Walrus1.2Sheep0.8
Monkey
2.1Camel
1.2Mouse0.5
Elephant
1.9Dog
1.2Rat
Whale
1.8Squirrel1.1Rabbit0.4
(AfterJerison;Blinkov&Glesner)
sapiens
(Marino1998).WhilemanhasanEQofabout7,thedolphins
Sotalia
uvi-
atilis
Delphinusdelphis
Tursiopstruncatus
haveEQsof3.2,andthegreatapes
(exceptman)haveEQsaround2.Thus,humanshaveamuchlargerbrainthanex-
pectedamongprimates,buteveninthisrespecttheirbrainisbynomeansunique,
astheexampleofdolphinsshows.
Fourthclaim
Humanshavethelargestcerebralcortex,particularlyprefrontalcor-
tex.Thereareenormousdifferencesbothinabsoluteandrelativebrainandpal-
GerhardRoth
increaseinglialcellsandbloodvessels,largeisocorticeswouldprobablybeboth
Isthehumanbrainunique?
GerhardRoth
Homo habilis
Homo erectus
Homo sapiens
Hominids
Australopithecines
GreatApes
Gorilla
Orang
Chimpanzee
Bonobo
Australopithecus africanus
Australopithecus robustus
Australopithecus boisei
log body weight (kg)
log endocranial volume (cm)
304050
75100
1000
1250
Figure6.
Increaseinrelativebrainsizeinthegreatapesandinhominids.AfterPilbeam
andGould(1974),modi
ed.
andconsciousness.Giventhefactthat
Homosapiens
hasanabsolutelyandrela-
tivelylargebrainandcortex,itappearstobetheanimalwiththehighestnumber
ofcorticalneuronsand/orsynapses,probablywiththeexceptionoftheelephant.
Thus,inthisrespecthumansarenottrulyexceptional.Whatishighlyremarkable,
however,isthestrongincreaseinrelative(andabsolute)brainsizeinhominidevo-
lutionduringthelast3
4millionyears.Whileinnon-humanprimatesaswellas
inhominidsthatdidnotrepresentourancestors,brainsizeincreaseswithbody
sizetoapowerof0.33
0.34,inthelineageleadingto
Homosapiens
itincreasedtoa
powerof1.73,i.e.ina
positivelyallometricfashion
,whichmeansthatbrainsizein-
creasedfasterthanbodysize(Figure6).However,thereasonsforthisphenomenon
Isthehumanbrainunique?
importantbasisforanincreasedcapabilityoflearningandmemoryformation.
TheothertraitconcernsthepresenceoftheBrocaspeechcenterinthefrontal
loberesponsiblefortemporalaspectsoflanguageincludingsyntax,alongwiththe
Wernickespeechcenterinthetemporallobewhichisresponsibleforthemean-
ingofwordsandsentences(althoughmeaningislikewisedependentonsyntax
GerhardRoth
Isthehumanbrainunique?
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Theco-evolutionoflanguageandworking
memorycapacityinthehumanbrain
OliverGruber
MaxPlanckInstituteofCognitiveNeuroscience,Leipzig,Germany
Introduction
OliverGruber
tablishedtheviewthatthesepremotorbrainareassubservetheverbalrehearsal
Theco-evolutionoflanguageandworkingmemorycapacity
mechanism.Thus,memoryperformanceunderarticulatorysuppressionhastorely
onother,non-articulatoryphonologicaland/orvisualstoragemechanismswhich
couldbemoresimilartoworkingmemorymechanismsinnon-humanprimates.
Inthefollowingstudiesarticulatorysuppressionwasusedtodeprivehumansub-
jectsofspeci
cverbalstrategiesandtomakethustheresultsofthesestudiesmore
comparabletothe
ndingsinnon-humanprimates.
Duringthe
Figure1.
Experimentaldesign.Subjectsperformedblockwiseaverbalitem-
OliverGruber
resolutionstructuralscanforeachsubjectfollowedbythreerunsof518gradient
echo-planarimage(EPI)volumeseach(TR2-s,TE40ms,
ipangle90
;number
ofslices16,voxelsize3
5mm
,distancefactor0.2)thatweresynchronized
withstimuluspresentation.
Twodifferentbrainsystemsunderliephonologicalworkingmemory
inhumans
Asexpected,silentarticulatorysuppressionledtoasigni
cantreductionofmem-
oryperformance,whereasalternating
ngertappingshowednosuchinterference
effect(meanpercentageofcorrectresponsesduringsingle-taskcondition/silent
articulatorysuppression/alternating
ngertapping:93.2/77.6/91.1%;F=21.61,
p0.001).Inordertorevealbrainareasinvolvedinmemoryperformanceunder
thevarioussecondaryconditions,wecomparedbrainactivityduringeachmem-
Theco-evolutionoflanguageandworkingmemorycapacity
OliverGruber
tobedifferentiallydistributedalongthesecorticalstructures.Inparticular,while
thephonologicaltaskvariantyieldedstrongactivationsalongtheanteriorparts
Theco-evolutionoflanguageandworkingmemorycapacity
mans(Figure3;Gruber&vonCramon2001).Moreover,otherstudiesprovide
sensory
information
Figure4.
Anevolutionary-basedmodelofhumanworkingmemory.Verbalrehearsal
isconsideredtobethemostef
cientandpredominantworkingmemorymechanism
inhumanswhichcanbeaccessedviarecodingmechanismsandwhichoperatesinde-
pendentlyfromtheoriginalstimulusmodality.Itisneurallyimplementedbythebrain
OliverGruber
appearspromisinginthatitmayoffernewexplanationsformanybehavioral,
neuropsychologicalandneuroimaging
ndingsinhumansubjects.Themodel
alsopermitstheharmonizationofcon
ictingworkingmemorymodelsderived
fromhumanrespectivelyanimalresearch(e.g.,Baddeley&Hitch1974;Goldman-
Rakic1996).
Withregardtoourdiscussiononmirrorneuronsandtheroleofpremotorcor-
ticesinhumanevolution,the
ndingspresentedherestronglysuggestthatBroca
areaandotherpremotorcorticesconstitutenotonlyasophisticatedlanguagesys-
tem,butalsoaveryef
cientworkingmemorymechanism.Thewell-knowneffect
ofarticulatorysuppressiononmemoryperformancecanbetakenasanindica-
tionfortheclearlyhighercapacityofthismemorymechanismascomparedtothe
Theco-evolutionoflanguageandworkingmemorycapacity
Gruber,O.(2000).Twodifferentbrainsystemsunderliephonologicalshort-termmemory
inhumans.
Neuroimage,11
,S407.
Gruber,O.(2001).Effectsofdomain-speci
cinterferenceonbrainactivationassociated
withverbalworkingmemorytaskperformance.
CerebralCortex,11
,1047
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Cerebralcorrelatesofworkingmemoryfortemporalinformation.
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Gruber,O.,&vonCramon,D.Y.(2001).Domain-speci
cdistributionofworkingmem-
oryprocessesalonghumanprefrontalandparietalcortices:Afunctionalmagnetic
resonanceimagingstudy.
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,380
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,2526
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basedregionalorganizationinhumanprefrontalcortex.
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componentofworkingmemory.
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,342
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contributionstospatialandnonspatialvisualworkingmemory.
Neuroimage,11
,409
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(1996).LocalizationofgrasprepresentationsinhumansbyPET.1.Observationversus
execution.
ExperimentalBrainResearch,111
(2),246
Rizzolatti,G.,&Arbib,M.A.(1998).Languagewithinourgrasp.
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(5),188
Romanski,L.M.,Tian,B.,Fritz,J.,Mishkin,M.,Goldman-Rakic,P.S.,&Rauschecker,
J.P.(1999).Dualstreamsofauditoryafferentstargetmultipledomainsintheprimate
prefrontalcortex.
NatureNeuroscience,2
(12),1131
Schumacher,E.H.,Lauber,E.,Awh,E.,Jonides,J.,Smith,E.E.,&Koeppe,R.A.(1996).PET
evidenceforanamodalverbalworkingmemorysystem.
Neuroimage,3
,79
OliverGruber
Figure2.
Brainregionssubservingphonologicalworkingmemoryunderdifferentcondi-
tions.Greenindicatesmemory-relatedactivationsthatoccurredonlyinabsenceofarticu-
latorysuppressionduringbothsingle-andnon-interferingdual-task(ST/DT)conditions.
Redindicatesmemory-relatedactivationsthatoccurredonlyunderarticulatorysuppres-
sion(AS).Brownindicatesmemory-relatedactivationsthatwerepresentinallconditions
investigatedinthisstudy,i.e.independentfromarticulatorysuppression.Barsintheinserts
showthemeanpercentageofsignalchangesproducedbythememorytasksinrelationto
therespectivecontrolconditions(L,left;R,right;fromGruber2001).
Figure3.
Domain-speci
cdistributionofworkingmemoryprocessesalonghumanpre-
frontalandparietalcortices.Predominantactivationofthecortexalongtheanteriorparts
oftheintermediateandsuperiorfrontalsulciandoftheinferiorparietallobulebyphono-
logicalmemory(indicatedinyellowandred),andofthecorticesalongposteriorpartsofthe
samefrontalsulciandofthesuperiorparietallobulebyvisualworkingmemory(indicated
inblueandgreen).Eachtaskwasperformedunderarticulatorysuppression(fromGruber
&vonGramon2001).
Episodicactionmemory
Characterizationofthetimecourse
andneuralcircuitry
AvaJ.Senkfor
AvaJ.Senkfor
DAY1
DAY2
Study Item Test
(Objects) (Photos)
Study Source Test
(Objects) (Photos)
Perform
Cost
Perform
Cost
Perform
Cost
Perform
Cost
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
Figure1.
Duringthestudyphases,150realobjectsortoyversionsofrealobjects,are
presentedoneatatime.Precedingeachobjectisanencodingtaskcue
Perform
Cost
.After7secondsatonesignalsremovaloftheobjectandthenextcueand
objectoccurs4secondslater.Ondayone,thetestphaseincludesallstudiedobjects
(digitalphotographs)plusanequalnumberofnewphotographsofobjects(ItemTest).
ParticipantsmakeOld/Newjudgmentstoeachobject.Ondaytwo,thestudyphaseis
Episodicactionmemory
Table1.
Accuracies(standarderror)intheitemandsourcememorytests.
Itemtest
Sourcetest
Hit
93(0.8)
94(0.5)
Perform
94(0.9)
94(0.8)
Cost
92(1.2)
94(1.0)
Hit/Hit
90(3.0)
Perform
93(3.1)
Cost
87(3.2)
Correct
95(0.8)
96(0.6)
Rejection
arewellmatchedinseveralways:bothrequirethatanobjectbeidenti
ed,andboth
requireaself-initiatedstrategytoproduceanacceptableresponse;participantsse-
lectedtheirown
typical
action,orthebestbasisforacostestimate.However,
onlythePerformtaskrequiredanalysisoftheobject
ssomatomotorproperties
(size,shape,andweightrelativetohandapertureandmusculareffort),selection
ofanappropriatemotorprogram,andexecutionofthemotorprogram.Attest,
participantsviewedallofthestudiedobjectsintermixedwithanequalnumberof
unstudiedobjects,andresponded
new
toeachobject.Figure1shows
AvaJ.Senkfor
Figure2.
Episodicactionmemory
300msaftertheonsetofthedigitalobjectimages,studiedobjectselicitmorepos-
itiveERPsthannewobjects,andthisdifferencecontinuesfortheremainderof
the1300msepochinboththeitemandsourcememorytests.Agenerallysimilar
old/neweffect
AvaJ.Senkfor
Figure3.
Episodicactionmemory
motoricinteraction.Italsoshowsthatthesedifferencesareapparentonlywhen
relevanttoone
AvaJ.Senkfor
Study Source Test
(Objects) (Photos)
Perform
Watch
Imagine
Cost
.
.
.
.
.
.
.
.
.
.
.
Figure4.
216realobjects,ortoyversionsofrealobjects,arepresentedoneatatime,
eachprecededbyaspokenencodingtaskcuefromataperecorder
Perform
Watch
Imagine
,or
Cost
.After7secondsatonesignalstheremovaloftheobjectandan-
othercueandobjectoccurs4secondslater.Attest,digitalcolorphotographsofall
Encodingtask
Reactiontime
Accuracy
Perform
1546(47)
93(1.1)
Watch
1651(58)
88(1.7)
Imagine
2072(99)
82(2.5)
Cost
1762(59)
78(2.6)
Episodicactionmemory
Figure5.
GrandaverageERPsfromrightandleftprefrontalsites,rightandleftmedial
fronto-centralsites,andleftlateraltemporal,parietal,andoccipitalsiteselicitedby
correctlyrememberedobjectsencodedwithperform,watch,imagine,andcosttasks.
AvaJ.Senkfor
advanceoftheparticipantsovertbuttonpressresponses(averageRTwas1757ms).
WethushypothesizethatthethreepatternsofERPresponsesarecausallyrelatedto
anaccuratedecision,sothatreductionoreliminationofanyoneofthethreebinary
distinctionswouldbeaccompaniedbydistinctpatternsofmemoryconfusions.
Episodicactionmemory
Figure6.
Meanamplitudefrommedialsitesforcorrectlyrememberedobjectsencoded
withtherightversuslefthandduringstudyforeachofthefourencodingtasks
Perform,Watch,Imagine,andCost.
Summaryandcurrentdirections
rstexperimentshowsthatalthoughrecentlyexperiencedobjectselicitdif-
ferentbrainactivitythanunstudiedobjects,merepresentationofanobjectdoes
notnecessarilyevokespeci
cmemoriesofone
spriorinteractionswithit.Both
experimentsshowthatwhenone
sprioractivities
are
AvaJ.Senkfor
Episodicactionmemory
Senkfor,A.J.,&Kutas,M.(2000).Effectsofagingonepisodicactionmemory.
Journalof
CognitiveNeuroscience
,Supplement,29.
Senkfor,A.J.,&VanPetten,C.(1998).Whosaidwhat:Anevent-relatedpotentialinves-
tigationofsourceanditemmemory.
JournalofExperimentalPsychology:Learning,
Memory&Cognition,24
,1005
Senkfor,A.J.,VanPetten,C.,&Kutas,M.(1999).Episodicactionmemory:AnERPanalysis.
JournalofCognitiveNeuroscience
,Supplement,31.
Senkfor,A.J.,VanPetten,C.,&Kutas,M.(submitted).Asourceisasource?AnERPanalysis
ofsourceanditemmemory.
Senkfor,A.J.,VanPetten,C.,&Kutas,M.(2002).Episodicactionmemoryforrealobjects:
AnERPinvestigationwithperform,watch,andimagineactionencodingtasksversusa
non-actiontask.
JournalofCognitiveNeuroscience,14
,402
Smith,M.E.,&Halgren,E.(1989).Dissociationofrecognitionmemorycomponents
followingtemporallobelesions.
JournalofExperimentalPsychology:Learning,Memory
&Cognition,15
,50
VanPetten,C.,&Senkfor,A.J.(1996).Memoryforwordsandnovelvisualpat-
terns:Repetition,recognition,andencodingeffectsintheevent-relatedpotential.
Psy
chophysiology,33
,491
VanPetten,C.,Senkfor,A.J.,&Newberg,W.(2000).Memoryfordrawingsinlocations:
Spatialsourcememoryandevent-relatedpotentials.
Psychophysiology,37
,551
Theroleofobjectsinimitation
AndreasWohlschl
gerandHaroldBekkering
Max-Planck-Institutf
rpsychologischeForschung,M
nchen,Germany
Introduction
Imitationplaysanimportantroleinskillacquisition
andnotmerelybecauseit
avoidstime-consumingtrial-and-errorlearning.Observingandimitatingisalsoa
specialcaseofthetranslationofsensoryinformationintoaction.Theactormust
translateacomplexdynamicvisualinputpatternintomotorcommandsinsuch
away,thattheresultingmovementvisuallymatchesthemodelmovement.For
thatreason,imitationisoneofthemostinterestingexamplesofperceptual-motor
co-ordination.
Althoughhumansareverysuccessfulinimitatingmanycomplexskills,the
mechanismsthatunderliesuccessfulimitationarepoorlyunderstood.Thetransla-
tionproblemisparticularlyinterestinginchildren,becausetheymustperformthe
translationdespitetheobviouslygreatdifferencesinorientation,bodysize,limb
lengths,andavailablemotorskills.Additionally,thesedifferencesresultinverydif-
ferentdynamicproperties(Meltzoff1993).Nevertheless,childrenspontaneously
andcontinuouslytrytoimitatethecustomsandskillsmanifestedbytheadults
andpeers.
Basedonearlier
ndings(Meltzoff&Moore1977),MeltzoffandMoore
(1994)developedanin
uentialtheory
thetheoryofactiveinter-modalmap-
ping(AIM)
thatassumesasupra-modalrepresentationalsystemthatmergesthe
perceptualandtheactionsystems.Thissupra-modalrepresentationalsystemis
thoughttomatchvisualinformationwithproprioceptiveinformation.TheAIM
theoryisinlinewiththecommonviewthat
inimitation
perceptionandaction
arecoupledbymeansofadirectperceptual-motormapping(cf.e.g.,Butterworth
\f
AndreasWohlschl
gerandHaroldBekkering
observation-executionmatchingsystem,becausethey
rebothduringtheobserva-
tionandduringtheexecutionofparticularactions.Supportforasimilarsystemin
humanscomesfromthe
ndingofamotorfacilitationduringactionobservation
Theroleofobjectsinimitation
\f
agoal-directedimitation.Indeed,recentresearchshowedthatalready6-month-
oldinfantsselectivelyencodethegoalobjectofanobservedreachingmovement
(Woodward1998).Theseresultsdemonstratethatchildrenperceivethegoalsand
intentionsofothersfromaveryearlyageon.
Wetestedourhypothesisofgoal-directedimitationbyavariationofthehand-
to-eartaskthatallowedtheremovalofthegoalobjectsofthemodel
smovement.
Insteadoftouchingtheears,themodelnowcoveredoneoftwoadjacentdotsstuck
tothesurfaceofatablewitheithertheipsi-orthecontra-lateralhand.Resultswere
similartothoseofthehand-to-eartask.Childrenalwayscoveredthecorrectdot;
buttheyquiteoftenusedtheipsi-lateralhandwhenthemodelcoveredthedot
contra-laterally.However,whenthesamehand-movementswereperformedwith
thedotsremoved,childrenimitatedalmostperfectlyipsi-lateralwithipsi-lateral
andcontra-lateralwithcontra-lateralmovements.
Thus,itseemsthatinimitationthepresenceorabsenceofgoalobjecthasa
decisivein
uenceonimitationbehaviour.Goal-orientedmovementsseemtobe
imitatedcorrectlywithrespecttothegoal;butthemovementitselfisfrequently
ignored.Movementswithoutgoalobjectsorwithasingle,non-ambiguousgoal
objectareimitatedmoreprecisely.Itseemsthatifthegoalisclear(orabsent),then
thecourseofthemovementplaysamorecentralroleinimitation.Onemightalso
saythatthen,themovementitselfbecomesthegoal.
Agoal-directedtheoryofimitation
Basedontheseresults,wedevelopedatheoryofgoal-directedorientationthat
\f
AndreasWohlschl
gerandHaroldBekkering
Thegoal-directedtheoryofimitationdoesnotonlyexplaintherecentdataofimi-
tationresearch,butalsogivesimitationamorefunctionalnature.Directmapping,
ontheotherhand,hasaratherautomatictaste.Thegoal-directedtheoryofimita-
tionallowsimitatorstolearnfrommodelsevenifthedifferencesinmotorskillsor
inbodyproportionsaresohugethattheimitatorisphysicallyunabletomakethe
samemovementasthemodel.Whatevermovementtheimitatoruses,thepurpose
oflearningbyimitationcanberegardedasbeingful
lledassoonashereachesthe
samegoalasthemodel.
Experiments
Theseriesofexperimentspresentedhereprovidesfurtherevidenceforthethe-
oryofgoal-directedimitation.Experiment1teststhe
ideo-motorprinciple
inchil-
dren
simitationbehaviour.Experiment2and3testthe
generalvalidity
ofour
goal-directedtheoryofimitationbyusingadultsubjectsinsteadofchildren.Ex-
periment2replicatesthedotexperiment(seeabove)withadultsandthusteststhe
generalvalidity
ofthe
ideo-motorprinciple
.Experiment3triestoclarifythe
hierar-
chicalorganisation
byinvestigatingtheimitationofmorecomplexobject-oriented
actionsinadultsandthusteststhe
generalvalidity
selectionofgoalsaspects
imitation.
Experiment1
Accordingtothe
ideo-motorprinciple
,themovementselicitedbythegoalofanac-
tionarethosethataremoststronglyassociatedwiththeachievementofthegoal.
Wealreadyshowed(Bekkeringetal.2000)thatcontra-lateralmovementsarequite
frequentlyimitatedwithipsi-lateralones(so-calledcontra-ipsi-error).This
nd-
ingisinkeepingwiththe
ideo-motorprinciple
,becauseitisquitelikelythatthe
moredirect,ipsi-lateralmovementismorestronglyassociatedwithreachingfor
anobjectthantheindirect,contra-lateralone.Experiment1triestoshow,thatthe
contra-ipsi-errorisnottheonlymanifestationofthe
ideo-motorprinciple
inimi-
tation.Onewaytoshowthatthe
ideo-motorprinciple
isofmoregeneralvalidityin
Theroleofobjectsinimitation
\f
right hand
grasping
Figure1.
ResultsofExperiment1.Irrespectiveofthefactthatthemodelusedboth
handswithequalfrequency,childrenprefertousetherighthandwhenimitating
thegraspingofobjects.Nosuchhandpreferenceisobservedwhenchildrenimitated
pointingtowardstheobjects.
Wethereforeasked16childrentoimitatecontra-andipsi-lateralmovementsto-
wardsoneoftwoobjects(acombandapen)onthetable.Halfofthechildrenwere
showngraspingmovements,whereastheotherhalfobservedtheexperimenter
pointingtowardstheobjects.Inthegraspingcondition,thedominanthandwas
usedabouttwiceasoftenasthelefthand,whereasinthepointingconditionhand
usewasbalanced(seeFigure1).Interestingly,thepreferenceforthedominanthand
inthegraspingconditionledto17%ipsi-contra-errors(allofthemweremadewith
therighthand),anerrorthathardlyoccurredinpreviousexperiments.Nosuch
errorsoccurredinthepointingcondition.Insummary,theresultsofExperiment
1demonstratethegoal-directednessofimitation,thestrengthoftheideo-motor
principleinimitation,andthatnotonlytheobjectsidentity(combvs.pen),but
\f
AndreasWohlschl
gerandHaroldBekkering
tasksliketouchingthecontra-lateralear,wedon
texpectadultstoshowthesame
error-pronebehaviourthatwefoundinchildren.Nevertheless,ifthegoal-directed
theoryofimitationisgenerallyvalid,some(perhapsweaker)effectsinadult
simi-
Figure2.
StimuliusedinExperiment2.Theadultparticipantshadtoputtheirhands
inapositionsimilartothatdepictedinthestimuliphotographs.Notethattheonly
Theroleofobjectsinimitation
\f\b
Inordertobeabletomeasureresponsetimesprecisely,weslightlymodi
thetask.First,weused
ngermovementsinsteadofwholehandmovements.Sec-
ond,themodelmovementswerenotpresentedbytheexperimenterbutonacom-
puterscreen.Subjectswereinstructedtoputtheirhandsnexttoeachotheron
thetable,justasdepictedinthestimuli(seeFigure2),andtoimitatethedepicted
downward
ngermovementasquicklyaspossibleafterthepresentationofoneof
thestimuli.Asintheexperimentwithchildren,thereweretwoconditions.Inone
condition,thestimulicontainedtwodots,oneofwhichwascoveredbyoneofthe
ngersattheendofaneitheripsi-lateralorcontra-lateraldownwardmovement.
Intheothercondition,thestimulidepictedthesamemovements,buttherewere
nodotspresent.
Twelveadultsubjectswentthroughboth,thedotsandtheno-dotscondition,
inblocks.Resultsshowedthatalthoughadultsalmostmadenoerrors(0.6%),
thesefewerrorsmainly(77.8%)occurredwithstimulidepictingcontra-lateral
movementstowardsdots(contra-ipsierror).Second,RTwerefasterforipsi-lateral
movements,butonlyifdotswerepresent(seeFigure3).Theseresults,thatbasically
replicatethe
ndingsinchildren,showthatalsoinadultsdotsasactiongoalsare
activatingthedirect,ipsi-lateralmotorprogramme,whichleadstofasterresponses
contra
depicted movement type
RT/ms
finger errors
with object
Figure3.
ResultsofExperiment2.Imitatingcontra-lateralwasslowerandmore
errorsweremade,butonlyifdotswerepresent.
\f\t
AndreasWohlschl
gerandHaroldBekkering
Experiment3
Althoughadultsshowthesameeffectsaschildreninsimpleactions,morecomplex
actionsareneededtoinvestigatethe
generalvalidity
andthe
hierarchicalorganisa-
tion
ofourgoal-directedtheoryofimitation.Currently,wehaveonlydatafrom
actionsthatcomprisetwovariableaspects:thegoalobjectandtheeffector.Inthe
followingexperiment,weincreasedthenumbertofourvariableaspects:thegoal
object
,the
treatment
oftheobject,the
effector
,andthe
movementpath
.Giventhe
highercomplexity,weexpectedalsoadultstoshowasubstantialnumberoferrors
inimitation.Rankingthedifferenterrortypesaccordingtothenumberoferrors
shouldyieldinsightintothehierarchyofgoalaspects.Weexpectedthesubjectsto
showtheleastnumberoferrorsinchoosingtheobjectandthetreatment,whereas
thechoiceoftheeffectorandthemovementpathshouldbequiteerror-prone.
Theactionweusedwasmorecomplexbutneverthelessquitesimple.Itcon-
sistedofmovingapenupsidedownintooneoftwocups(
object
)ortouchingthe
shandlewiththepen
scap(
treatment
).Ineithercase,thepenhadtorotated
by180
.Theexperimenterservedasthemodelandheeitherusedhisrightorhis
lefthand(
effector
).Inaddition,heeitherturnedthepenclockwiseorcounter-
clockwise(
movementpath
)tobringitintoanupsidedownpositionattheendof
themovement(seeFigure4).
32adultsservedasparticipantsintheexperiment.Theywerekeptna
veabout
thepurposeoftheexperiment.Beforeshowingtheactiontothem,theyweresim-
plyasked
CanyoudowhatIdo?
Wewereinterestedin
spontaneousimitation
andthereforeweranonlyonetrialforeachsubject.
Theresultsshowedthatindeed,adultsproducedaconsiderableamountof
Theroleofobjectsinimitation
\f\n
Figure4.
ThreeframesofanimitationsequenceofExperiment3.Themodelonthe
rightusesthelefthandtoputthepenupsidedownintotherightcupbyturningit
counter-clockwise.Theimitatorusestherighthandandturnsthepenclockwisetoput
itintotheleftcup(notshown).Inthisexample,theimitatorperfectlymirroredthe
AndreasWohlschl
gerandHaroldBekkering
object
treatmenteffectormovement
action feature
% correct
objects differ by colour and location
objects differ by location only
Figure5.
ResultsofExperiment3.Itwasmainlythetreatmentoftheobject(penwent
intothecupvs.pentoucheditshandle)thatwasimitatedcorrectly.Thecorrect
wasonlyused,ifthecupsdifferedbycolour.Otherwise,thechoiceofthecupaswellas
thechoiceoftheeffectorandthemovementpathwerebasicallyatchancelevel.
Discussion
Inthischapterwereportedaseriesofexperimentsthatdemonstratetheimpor-
tanceofobjectsandtheirtreatmentsinhumanimitation,bothforchildrenand
adults.Theexperimentsshowedthatitisprimarilythetreatmentofanobjectthat
isimitatedinobject-orientedactions,whereasthechoiceoftheeffectorandthe
movementpatharefollowingtheso-called
ideo-motorprinciple
:Themotorpro-
grammemoststronglyassociatedwiththeachievementofthegoalisactivated
duringtheexecutionoftheimitativeactanditisprobablyalreadyexecuteddur-
Theroleofobjectsinimitation
ignoringthemotorpartoftheaction.Ofcourse,inmostcasesthemodelactsin
anef
cientanddirectwayontheobject.Iftheimitatorcopiestheactiongoaland
ifthisactiongoalinturnactivatesthemostdirectmotorprogrammeintheim-
itator,thenbothactionsresembleeachotherinallaspects,leadingtoanimpres-
sive,mirror-likebehaviour.Whenthereisnoobject,themovementsthemselves
becomethegoalandtheyarealsoimitatedinamirror-likefashion.Itisproba-
AndreasWohlschl
gerandHaroldBekkering
imitationof
ngermovements.Onemightspeculatethatduringtheevolution
ofspecies,
rstasystemforactionunderstandinghadtobeevolvedbeforethe
imitationofactiongoalscouldevolve.
Note
Correct
hastobeunderstoodinthemirror-sense,becausechildrenspontaneously
imitateipsi-lateralmovementsinamirror-fashion.
References
Bekkering,H.,Wohlschl
ger,A.,&Gattis,M.(2000).Imitationofgesturesinchildrenis
goal-directed.
TheQuarterlyJournalofExperimentalPsychology:SectionA:HumanEx
perimentalPsychology,53
,153
Butterworth,G.(1990).Onreconceptualisingsensori-motordevelopmentindynamic
systemsterms.InH.Bloch&B.I.Bertenthal(Eds.),
SensoryMotorOrganizationsand
DevelopmentinInfancyandEarlyChildhood
[NATOAdvancedScienceInstitutesseries,
D:BehaviouralandSocialSciences,56](pp.57
Theroleofobjectsinimitation
Wapner,S.,&Cirillo,L.(1968).Imitationofamodel
shandmovements:Agechangesin
transpositionsofleft-rightrelations.
ChildDevelopment,39
,887
Wohlschl
ger,A.,&Bekkering,H.(2002).Ishumanimitationbasedonamirror-neurone
system?Somebehaviouralevidence.
ExperimentalBrainResearch,143
,335
Woodward,A.L.(1998).Infantsselectivelyencodethegoalobjectofanactor
sreach.
Cognition,69
Themirrorsystemandjointaction
ntherKnoblichandJeromeScottJordan
MaxPlanckInstituteforPsychologicalResearch,Munich,Germany/
IllinoisStateUniversity,Normal,USA
Introduction
Themostexcitingaspectofthediscoveryofmirrorneuronsisthatpartsofthecog-
nitivesystemareentirelydevotedtotheprocessingofsocialinformation(Fadiga,
Fogassi,Pavesi,&Rizzolatti1995;Gallese,Fadiga,Fogassi,&Rizzolatti1996;Riz-
zolatti&Arbib1998).Forasinglebeinglikeamonkeyonmars,themirrorsystem
ntherKnoblichandJeromeScottJordan
formsofactioncoordination.Wethenproposeafunctionalmechanismforaction
coordinationthatextendsthefunctionalityofthemirrorsystembymodulatingthe
planningofone
sownactioninresponsetoperceivingtheoutcomesofsomebody
else
sactions.Althoughthismechanismisinherentlynon-linguistic,itmayalsobe
abridgingelementinthetransitionfromasystemthatproducesandunderstands
manualactionstoasophisticatedlanguagefaculty(Calvin&Bickerton2000;Riz-
zolatti&Arbib1998).Finally,wewillprovidesomeempiricalfortheexistenceof
suchamechanisminhumans.
Ego-centeredandgroup-centeredactionunderstanding
Themostobviousinterpretationofthefunctioningofthemirrorsystemisthatitis
specializedinprocessinginformationaboutobject-directedactionsofothersthat
matchactionsintheobserver
Themirrorsystemandjointaction
understandingprovidedbythemirrorsystemmaybepurelyegocentric,andits
mainfunctiontograspinteresting(eatable)objectsbeforetheyaregone.
However,therearesituationsinwhichapurelyegocentricperspectiveisnot
suf
certainsituationsjointaction(Clark1997)allowsonetoachievegoalsthatcould
notbeachievedotherwise.Forinstance,asharedgoalcouldbetoremoveaheavy
objectthatisblockingtheway,inordertoproceed.Ifoneindividualtriestomove
theobject,theeffectobtainedmightberathersmall.Iftwoindividualsjointheir
effortsandpushtheobjectsimultaneously,theeffectobtainedwillbemuchlarger.
Hence,thejointeffectwillbemuchlargerthanthesumoftheindividualeffects.Al-
ternatively,ifthegoalistojointlysteeracanoetowardsacertainlocation,agroup
ismorelikelytoachievethis,whenpaddlinginturn.Otherwise,itisverylikelythat
theactionsofoneindividualcounteracttheactionsoftheotherindividual,thereby
ntherKnoblichandJeromeScottJordan
Perceptual Input
JointAction System
(Joint Effects)
Mirror System
(IndividualAction Effects)
Motor System
––
Figure1.
Modelformodulationofthemirrorsystembyjointeffects.
Themirrorsystemandjointaction
thiscase,performancecanbeoptimizedwithouttheadditionalrepresentationof
jointeffects.Therefore,theindividualconditionprovidesabaselineforthegroup
conditions,becausetheactioncon
ictcanberesolvedwithouttheprocessingof
socialinformation(withouttakingtheother
sactionsintoaccount).Ourpredic-
tionisthatgroupslearntocoordinatetheiractionsaswellasindividualsifindivid-
ualandjointeffectscanbeclearlydistinguished,butperformworseifthatisnot
thecase.
Screen border
Screen border
Tracker
Figure2.
Illustrationoftrackingtask.
\f
ntherKnoblichandJeromeScottJordan
before
before
after
after
a) Effect ofa compensatory button press.
b) Effect ofan anticipatory button press.
Figure3.
Effectsofcompensatoryandanticipatorybuttonpresses.Thesmallsolidcir-
Themirrorsystemandjointaction

ipantwasprovidedwithaseparatecomputermonitor,andalleventstakingplace
duringtheexperiment(e.g.themovementsofthetrackerandthemovementsof
Block
Distance in mm
Joint
Joint +
Individual
Individual +
Figure4.

ntherKnoblichandJeromeScottJordan
acousticaldidnotmakeadifferenceintheindividualcondition,ithelpedgroupsto
Block
Individual +
Joint
Joint +
Individual
Anticipatory Brake Rate
Figure5.
Anticipatorybrakerateforindividualsandgroupswithandwithoutacous-
ticalsignalacrossblocks.
Themirrorsystemandjointaction

(Individual+)ordidnotreceive(Individual
)auditoryfeedback,andgroupswho
received(Joint+)anddidnotreceive(Joint
)auditoryfeedback.Theprogression
ofeachlinere
ectschangesinanticipatorybrakerateacrossblocks.
Duringtheinitialtrials(Block1),groupsandindividualswereclearlydifferent
andthepresenceoftheacousticalsignaldidnotmakearealdifferenceforeitherof
them.Individualsstartedoutatarelativelyhighlevelrightfromstartandtheinitial
anticipatorybrakeratewasclearlylowerinthegroupconditions.Inlaterblocks,
theanticipatorybrakerateincreasedforindividualsaswellasgroups.However,
whilethepresenceoftheacousticalsignaldidnotaffecttheincreaseinthean-
ticipatorybrakerateintheindividualcondition,itdidaffecttheincreaseinthe
groupcondition.Theanticipatorybrakerateincreasedsharplyfromthe
rsttothe
secondblockinthegroupconditioninwhichtheacousticalsignalwaspresent.
Inthe
nalblockitwasalmostashighintheindividualconditions.Incontrast,
theanticipatorybrakerateincreasedonlyslightlyforgroupswhodidnotreceive
theacousticalsignal,theextentoftheincreasebeingcomparabletotheindividual
conditions.
Theresultsfortheperformanceandthestrategymeasureareconsistentwith
thepredictions.Groupslearnedtocoordinatetheiractionsaswellasindividuals
whenindividualandjointeffectscouldbeclearlydistinguished,butperformed
worseifthatwasnotthecase,whereasindividualsdidnotbene
tfromthead-
ditionalacousticalactioneffect.Atpresent,thereareprobablyalternativewaysof
explainingtheresults.Furtherexperimentsareneededtounambiguouslyassess

ntherKnoblichandJeromeScottJordan
mightbemodulatedbyrepresentationofjointeffects.Afurtherimplicationofthe
presentstudyisthatitmightbeusefultolookatotherformsofcoordinatedaction
Brainactivationtopassiveobservation
ofgraspingactions
FrancisMcGlone,MatthewHoward,andNeilRoberts
CentreforCognitiveNeuroscience,UniversityofWales,Bangorand

FrancisMcGlone,MatthewHoward,andNeilRoberts
Theexistenceforasimilarsysteminmancouldhavebeeninferredfromthe
pioneeringworkofoneofthe
rstelectromyographers,E.Jacobson(the
rstscien-
tisttorecordeyemovementsduringsleep),whointheearlypartofthelastcentury
recorded
minute
voltagesfromsomaticmuscleswhenhissubjectsimaginedper-
formingspeci
carmmovements.Onlythemusclegroupsthatwouldhavegener-
atedthesemovementsproducedthesignals.Morerecently,employingtranscranial
Brainactivationtopassiveobservationofgraspingactions
\b
Methods
Subjects.
Writtenconsentwasobtainedfor13right-handedhealthysubjects(5
female;8male;meanage28years).Allsubjectshadnormalvisionorworecontact
lenses,andwerepaidfortheirtime.
ExperimentalDesign.
Foreachexperimentalcondition,100EPIbrainvolumes
werecollectedover300seconds.Thisacquisitionperiodwasdividedinto20
epochs,eachof
fteensecondsinlength.Subjectsperformedabaseline(OFF
event)inthe
rstepoch,followedbythetaskofinterest(ONevent).ThisON/OFF
boxcar
designwasrepeatedinsubsequentepochs.
EachsubjectunderwentfMRIscanningoftwodifferentexperimentalcondi-
tions.IntheONeventofeachcondition,subjectspassivelyvieweda15second
QuickTimemovie(AppleComputer,USA)ofactorsperformingastereotypical,
goaldirectedmotorbehaviour.FortheONevents,subjectsviewedacloseupofan
actor
shandperformingaprecisiongriptask,pickingupapenfromadesktop,or
anumberofsmallcoins.InallconditionstheOFFeventwasastaticpictureofthe
actorandobject(seeFigure1).
fMRIStimulusPresentation.
AllstimuliweregeneratedbyanApplePowerMa-
cintoshG3runningPsyscopesoftware.AnLCDprojector(EpsonLMP7300)was
\t
FrancisMcGlone,MatthewHoward,andNeilRoberts
(a)
Figure1.
Stillimageofcoins(a)andpen(b)shownduringOFFperiod.DuringON
periodobjectswerepickedupina15secondvideoclipback-projectedontoascreen
placedattheendofthescanner.Subjectsviewedtheimagesthroughprismspectacles.
tion:Eachofthefunctionalvolumesisrealignedwiththe
rstoneintheseries,
usingacubicsplineinterpolationtechnique.Thealgorithmalsoincorporatesspin-
historycorrectiontominimisemotioncorrelatedintensityvariations.(2)Spatial
normalisation:Thefunctionaldatawerespatiallytransformedtoastandardised
canonicalco-ordinateframeofreferencebaseduponthebicommisuralco-ordinate
systemofTalairachandTournoux(1988).Thisisachievedbytransformingthe3D
high-resolutionvolumetoaT1-weightedtemplate,initiallyusinga12parame-
teraf
Brainactivationtopassiveobservationofgraspingactions
\n
basisfunctiontodescribethedeformations.(3)Spatialsmoothing:Thestereotac-
ticallynormalisedfunctionalvolumesweresmoothedusinganisotropicGaussian
kernelof6mmfullwidthathalfmaximum.Thisallowedforinter-individualvari-
ationinthelocationoffunctionalactivity,andconditionedthedatasuchthatthey
conformedtotherandomGaussian
FrancisMcGlone,MatthewHoward,andNeilRoberts
Condition-speci
cactivations
Coingraspingcondition
.Twodistinctbandsofactivationextendingfromdorsal
toventralwerepresentinthiscondition(Figure2a).Severaldistinctclusters
ofactivationwereseenintheprecentralgyrus[BA4],conjoinedwithacti-
Figure2.
Averagedactivationpatternsfoundfromwatchingcoins(toprow)andpen
(bottomrow)beingpickedup.Thevolumerenderedimagesshowtheglobalactivation
producedbyobservingtheseactions,withmoreactivationbeinggeneratedbythecoin
condition.Thecoronalsectionsdemonstrateclearlythelackofactivityfoundinthe
pengraspingconditioninIFG.Datathresholdedatp0.05.
Brainactivationtopassiveobservationofgraspingactions
Table1a.
Areasofsigni
cantactivation(p0.05,corrected)forallconditions.
Region
R/LBAxyzZ-score
VisualAreaV5
R1945
6666.84
VisualAreaV5
L39
663Inf
SuperiorParietalLobule
R739
4557Inf
SuperiorParietalLobule
L7
5157Inf
InferiorParietalLobule
L40
27215.92
InferiorParietalLobule
R4051
39425.4
MiddleFrontalGyrus/SulcusL44/6
576365.92
MiddleFrontalGyrus
R6/8480546.16
SuperiorFrontalGyrus*
R633
6575.77
Superior/MiddleFrontalGyrus+L6
9637.75
InferiorFrontalGyrus*
R445715275.16
(*=Coinconditiononly,+=penconditiononly)
Table1b.
Fixedeffectsanalysis(Voxelsuniquelyactivatedincoincondition).
Region
R/LBAxyzZ-score
VisualAreaV5
R1948
75186.69
VisualAreaV5
L40/39
5766.04
SuperiorParietalLobuleR1924
78456.56
SuperiorParietalLobuleR7/4042
39575.2
InferiorParietalLobuleL4063
48335.75
Middle/InferiorFrontalGyrusR44/6516395.92
PrecentralGyrus
R463
15365.84
ExtrastriateCortex
R3948
6395.28
uniquelyactivatedinthecoinconditionwerefoundinthemiddleoccipitalgyrus,
FrancisMcGlone,MatthewHoward,andNeilRoberts
s(Jacobson1930,1932)wherehedescribes,inaseriesofpapers,theelectro-
physiologicalrecordingof
microscopic
contractionsinvoluntarymusclescon-
cordantwithsubjects
imaginingorobservingstereotypicalmovements,suchas
servingatennisball.Themorerecentobservationsofenhancedmotorevokedpo-
tentials(MEP
s)recordedduringTranscranialMagneticStimulationofmotorcor-
texwhilethesubjectobservedtheexperimentergraspingobjectsprovidesfurther
evidencethata
mirrorsystem
Brainactivationtopassiveobservationofgraspingactions
servationsareataverypreliminarystage,thenweshouldbeabletodemonstrate
thisbyinvestigatingtheeffectsofmanipulatingegocentricandallocentricspace
andthedependenceofmirroractivityonobserverrelevantactions.Wecarriedout
afurtherseriesofexperimentsinwhichweusedsubjectsover-trainedinaspeci
manualtask,namelyprofessionalmusicians(violinists),toestablishthedegreeto
whichobserverfamiliarityisrepresentedinMN
s(Howard2001).
Inhumans,MNactivitycouldbecovert,aswellasovert,theformerproviding
afeelingofempathyorfamiliarityintheobserverastheobservedactionhasto
bebyde
nitiononethatisintheobserversbehaviouralrepertoire
orimagined
repertoire(Grafton1996).Thismightunderpintheattractionofwatchingfootball
matches,boxing,orevocative
lmsinthattheobserveris
onthepitch
inthering
emotivelyengaged
whilstremainingmotoricallypassive.Ithasbeenpostulated
FrancisMcGlone,MatthewHoward,andNeilRoberts
Howard,M.A.,McGlone,F.,Tipper,S.,Brooks,J.C.W.,Sluming,V.,Phillips,N.,&Roberts,
N.(2001).ActionobservationinorchestralstringplayersusingeFMRI.
NeuroImage,
(6),1189.
Jacobson,E.(1930).Electricalmeasurementsofneuromuscularstatesduringmental
activities.
AmericanJournalofPhysiology,92
,567
Jacobson,E.(1932).Electrophysiologyofmentalactivities.
AmericanJournalofPsychology,
,677
Lhermitte.F.,Pillon,B.,&Serdaru,M.(1986).Humanautonomyandthefrontallobes1.
Imitationandutilizationbehavior
aneuropsychologicalstudyof75patients.
Annals
ofNeurology,19
(4),326
Maravita,A.,Spence,C.,Clarke,K.,Husain,M.,&Driver,J.(2000).Visionandtouch
throughthelookingglassinacaseofcrossmodalextinction.
Neuroreport,11
,3521
Passingham,R.(1993).
Thefrontallobesandvoluntaryaction
.Oxford:OxfordUniversity
Press.
Mirrorneuronsandtheselfconstruct
KaiVogeleyandAlbertNewen
InstituteofMedicine,ResearchCenterJ
lich,Germany/
DepartmentofPhilosophy,UniversityofBonn,Germany
Introduction
KaiVogeleyandAlbertNewen
theobservationofgoal-directedbehavioralsequencesofotheranimals.Inaddi-
Mirrorneuronsandtheselfconstruct
KaiVogeleyandAlbertNewen
Self-consciousnessandselfconstruct
Oneofthefocusesoftherecentdebateincognitiveneurosciencesistheconceptof
thehumanselfasamatterofempiricalneuroscience.Ifempiricalindicatorsfordif-
ferentdomainsofthehumanselfmodelcanbefound,thenanoperationalization
andamappingtoneuronalstructuresbecomespossible.
Classical
featuresofthe
selfdealtwithinthephilosophicalaswellaspsychologicaltraditionmaybethen
addressedempiricallywithrespecttotheirimplementationinspeci
cneuronal
networkarchitectures.
Consciousnessingeneralmaybede
nedastheintegratedinternalrepresenta-
tionoftheouterworldandourorganismbasedonactualexperiences,perceptions
andmemoriesprovidingre
ectedresponsestotheneedsofourenvironment.Con-
sciousnessisafundamentaltoolforourorientationintheworldandreliesupon
theintegrative,supramodal,sensory-independent,holisticrepresentationofthe
world.Thisworldmodelreferstodifferentcoordinatesystems,bothobject-and
viewer-centeredperspectivesinspacerepresentation,bothphysicalandsubjective
timescalesintimerepresentation.Thesereferenceframesareinturnbasedondata
ofthedifferentsensorysystems.Self-consciousnessincludesconsciousnessofones
ownmentalstates,suchasperceptions,attitudes,opinions,intentionstoact,and
soforth.Representingsuchmentalstatesintoonecombinedframeworkthatallows
Mirrorneuronsandtheselfconstruct
Itwaspostulatedthatthesebasicpropertiesareintegratedinapostulatedso-
selfmodel
asanepisodicallyactivecomplexneuralactivationpatternin
thehumanbrain,possiblybasedonaninnateand
hard-wired
\f
KaiVogeleyandAlbertNewen
withpbeingaphysicalevent),SELFinthiscontextreferstothespecialsituation,
inwhichIamtheagentmyself(e.g.
Mirrorneuronsandtheselfconstruct

Theory ofMind
TOM +TOM
SelfPerspective
Condition 3
TOM Stories
(T+S
Condition 2
Physical Stories
Condition 4
Selfand Other
Ascription Stories
(T+S+)
Condition 5
SelfStories
Ascription
Figure1.
Two-wayfactorialdesignofthestudy.Thisschemademonstratesthetwo-
wayfactorialexperimentaldesignapplied,inwhichbothfactorsTOMandSELFwere
variedsystematically.
ambiguoussituations.Thecorrectassignmentofanotherperson
smentalstatein
theTOMconditionswastestedbyaskingtheparticipantstoinferaspeci
cbe-
haviororattitudeofanotherpersoninthegivencontextofthestory,judgedas

KaiVogeleyandAlbertNewen
Exampleof
TOMstories
Aburglarwhohasjustrobbedashopismakinghisgetaway.Asheisrunninghome,
apolicemanonhisbeatseeshimdrophisglove.Hedoesn
tknowthemanisabur-
glar,hejustwantstotellhimhedroppedhisglove.Butwhenthepolicemanshouts
outtotheburglar,
Hey,you!Stop!
,theburglarturnsround,seesthepoliceman
andgiveshimselfup.Heputshishandsupandadmitsthathedidthebreak-inat
thelocalshop.
Question:Whydidtheburglardothat?
Exampleof
selfandotherascriptionstories
Mirrorneuronsandtheselfconstruct

ization,andstatisticalanalysiswasperformedusingStatisticalParametricalMap-
ping(SPM99,WellcomeDepartmentofCognitiveNeurology,London,UK).For
thefMRIgroupdataanalysis,allimagesofallsubjectswereanalyzedinonedesign
matrixina
xed-effectmodel.Thedatawereanalyzedbothwithrespecttothespe-
ceffectsofeachconditionagainstthebaseline(
unlinkedsentences
condition)
andwithrespecttothemaineffectsofTOMandSELF.Inaddition,thecontrastof
SELFrelativetoTOMwascalculatedtoassessthesigni
canceofthespeci
cdiffer-

KaiVogeleyandAlbertNewen
Mirrorneuronsandtheselfconstruct

Neuronalimplementationoftheselfmodel
Ourresultsdemonstratethattheabilitytoattributeopinions,perceptionsor
attitudestoothers,oftenreferredtoasTOMor
mind-reading
andtheabilityto
applySELFrelyonbothcommonanddifferentialneuralmechanisms.Thecere-
bralimplementationofTOMcapacityislocatedpredominantlyintheanterior
cingulatecortex.Thispartofthedescribedexperimentreplicatespreviousstudies
Figure2.
MaineffectsofTOMandSELFandtheirinteraction.(2a)(T+S+plusT+S
relativeto(T
S+plusT
).UnderthemaineffectofTOMthereissigni
cantac-
tivationintherightanteriorcingulatecortex,andleftsuperiortemporalcortex.(2b)
(T+S+plusT
S+)relativeto(T+S
plusT
).UnderthemainfactorSELFthereis
stillconsiderableactivationattheanteriorcingulatecortexandsigni
cantactivation

KaiVogeleyandAlbertNewen
abilitytoapplySELFisrequired.TakingSELFappearstoactivatetherightinferior
Mirrorneuronsandtheselfconstruct
\b
experiencesandproprioceptiveinformationisdisturbed,theresultwouldbethe
lossoftheexperientialperspectivity.
Conclusion
Theresultsofourstudydemonstratethattheabilitytoattributeopinions,percep-
tionsorattitudestoothersandtheabilitytoapplySELFrelyinpartondifferen-
tialneuralmechanisms.WhereasTOMispredominantlyassociatedwithincreased
activityintheanteriorcingulatecortex,thecapacitytotakeSELFispredomi-
\t
KaiVogeleyandAlbertNewen
cerebralimplementationofanimportantfeatureofself-consciousnessandhave
importantsigni
canceforcognitiveandneurophilosophicaltheoriesofconscious-
ness.
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Keenan,J.P.,Wheeler,M.A.,Gallup,G.G.Jr.,&Pascual-Leone,A.(2000).Self-recognition
andtherightprefrontalcortex.
TrendsinCognitiveScience,4
,338
Maguire,E.A.,Burgess,N.,Donnett,J.G.,Frackowiak,R.S.,Frith,C.D.,&O
Keefe,J.
(1998).Knowingwhereandgettingthere:Ahumannavigationnetwork.
Science
Melzack,R.,Israel,R.,Lacroix,R.,&Schultz,G.(1997).Phantomlimbsinpeoplewith
congenitallimbde
ciencyoramputationinearlychildhood.
Brain
,1603
Metzinger,T.(1993).
SubjektundSelbstmodell
.Paderborn:Sch
ningh.
Metzinger,T.(1995).Fasterthanthought.Holism,homogeneityandtemporalcoding.
InT.Metzinger(Ed.),
ConsciousExperience
(pp.425
461).Thorverton:Imprint
Academic.
Perner,J.,&Howes,D.(1992).
Hethinksheknows
:Andmoredevelopmentalevidence
againstsimulation(roletaking)theory.
MindandLanguage,7
,72
Premack,D.,&Woodruff,D.(1978).Doesthechimpanzeehavea
theoryofmind
BehavioralandBrainSciences,4
,515
Rizzolatti,G.,Fadiga,L.,Matella,M.,Bettinardi,V.,Paulesu,E.,Perani,D.,&Fazio,
F.(1996).Localizationofgrasprepresentationinhumansbypositronemission
tomography:1.Observationversusexecution.
ExperimentalBrainResearch
,246
Shallice,T.&Burgess,P.(1996).Thedomainofsupervisoryprocessesandtemporal
organizationofbehaviour.
PhilosophicalTransactionsoftheRoyalSociety:Biologic
Sciences
,1405
Shobris,J.G.(1996).Theanatomyofintelligence.
Genetic,Social,andGeneralPsychology
Monographs
(2),133
KaiVogeleyandAlbertNewen
Vallar,G.,Lobel,E.,Galati,G.,Berthoz,A.,Pizzamiglio,L.,&LeBihan,D.(1999).Afronto-
Behavioralsynchronizationinhuman
conversationalinteraction
JenniferL.RotondoandStevenM.Boker
AugustanaCollege,SiouxFalls,SouthDakota,USA/
UniversityofNotreDame,USA
Introduction
Intheir1998article,RizzolattiandArbibnotedthatneuronsintheF6regionof
theventralpremotorcortexareactivatedinprimateswhentheyobserveormimic
asimpleaction,suchasgraspingagrape.Theauthorssuggestedthatthereis,per-
JenniferL.RotondoandStevenM.Boker
thattheybegintodesynchronizeordecouplethemselvesfromoneanother.We
Behavioralsynchronizationinhumanconversationalinteraction
gazingmoreatotherindividuals),andpreferand/ortoleratecloserproximities
totheirconversationalpartnersthanmen(Dovidio,Ellyson,Keating,Heltman,&
Brown1988;Hall1984;Williams&Best1986).Womenareusuallytherecipients
ofsomeofthesebehaviorsalso,suchasbeinggazedatmoreoftenandbeingap-
proachedmorecloselybyothers(Hall1984).Incontrast,menstareawaymore,are
JenniferL.RotondoandStevenM.Boker
nancearethesametraitsascribedtomenordescribedascharacteristicofmas-
culinity.Inturn,manyofthesetraitsarefoundtobehighlyvaluedinAmericanso-
Behavioralsynchronizationinhumanconversationalinteraction
Apparatus
PhysicalmotionwasrecordedusingAscensionTechnologiesMotionStar16sensor
JenniferL.RotondoandStevenM.Boker
uenceanother
s,givingasenseofnonverbalconversational
ow.Usingthese
xedinterval,and
valuestotherightofsynchronyindicatecorrelationsobtainedwhenthewindow
wasaheadofsynchrony.Becauseofthisstructure,theseplotsareabletoshow
theprogressionofstronglyandweaklycorrelatedmovementacrosstimeaswellas
whichoftheparticipantsinitiatesthemovementtowhichtheyarebothstronglyor
weaklyresponding.Whenhighcorrelationsareononesideofsynchrony,person1
+1+2
Time
Lead
Figure1.
Behavioralsynchronizationinhumanconversationalinteraction
issaidtobeleadingthemovement;whenhighcorrelationsarefoundontheother
sideofsynchrony,person2issaidtobeleadingthemovement.
Suchgraphssuggestthatbehaviorsinitiatedbyoneindividualaresubsequently
matchedormirroredbytheother.Thereisalsoevidenceofsustainedbehavioral
coordinationintheareasofverticalhighcorrelation,whichusuallyareimme-
foreachconversation.
Toassesspatternsofsymmetryformationandbreakingforour4conversation
types(HDF/LDM,HDM/LDM,HDF/LDF,HDM/LDF),wethoughtofeachlag
andbininthedifferencematricesasacellandcalculatedmeanandstandarderror
JenniferL.RotondoandStevenM.Boker
.50.51.01.52.0
Figure2a.
Female
female
.50.51.01.52.0
Figure2b.
LDM
Behavioralsynchronizationinhumanconversationalinteraction
.50.51.01.52.0
Figure2c.
LDF
.50.51.01.52.0
Figure2d.
Male
\f
JenniferL.RotondoandStevenM.Boker
ahighdominantfemale
sbehaviorbyapproximatelythree-quartersofasecond.
Thesameistruewhenalowdominantmale
sbehaviorisdisplayed
rst.Forthis
particularpairingthereisevidenceofgreaterbehavioralsynchronyforheadmo-
tionsinitiatedbythelowdominantmale,somethingcontrarytothepersonality
perspective.
AnotherinterestingresultwasfoundfortheHDM/LDFdyads,whichdemon-
strateperiodicityinleadingandfollowing.Theresultssuggestthatheadmotions
performedbylowdominantfemaleswerematchedormirroredbytheirhighdom-
inantmalepartnersabout1.5secondslater;inaddition,highdominantmalehead
motionswerematchedormirroredbythelowdominantfemalesaboutahalf-
secondaftertheywereinitiated.Thisperiodicitywasnotduplicatedbyanyother
dyadcompositionandsuggeststhatfurtheranalysisofthisparticulardyadtype
mightleadtointerestinginsightsaboutthenonverbalcommunicationpatterns
displayedbythese
traditional
malesandfemales.
Discussion
Suchindicatorssuggestthatsymmetryformationandbreakinginconversation
followin
uencesfromcontextualcues,speci
callygenderandpersonality.Males
andfemalesappeartoresponddifferentiallybasedonpartnergender.Malesin
conversationwithoneanotherappearunaffectedbycoordinatedmovement;if
viewedalone,onemightconcludethatnonverbalcoordinationisanunnecessary
componentoftheirconversationalstyle.However,wheninconversationwithfe-
males,malesadapttotheirconversationalpartnersaswellasinitiatemovements
Behavioralsynchronizationinhumanconversationalinteraction

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Bargh,J.A.,Raymond,P.,Pryor,J.B.,&Strack,F.(1995).Attractivenessoftheunderling:
Anautomaticpower-sexassociationanditsconsequencesforsexualharassmentand
aggression.
JournalofPersonalityandSocialPsychology
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Benjamin,G.R.,&Creider,C.A.(1975).Socialdistinctionsinnonverbalbehavior.
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visualattention:Anintegratedanalysisofmovementandgazeinmixed-sexdyads.
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Bilous,F.R.,&Krauss,R.M.(1988).Dominanceandaccommodationintheconversational
behaviorsofsame-andmixed-genderdyads.
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Bull,P.E.(1987).
PostureandGesture
.NewYork:PergamonPress.
Burgoon,J.K.,Buller,D.B.,&Woodall,W.G.(1996).
NonverbalCommunication:The
unspokendialogue
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Burgoon,J.K.,&Dillman,L.(1995).Gender,immediacy,andnonverbalcommunication.
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Condon,W.S.,&Ogston,W.D.(1967).Asegmentationofbehavior.
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Mehrabian,A.(1981).
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Mitchell,G.,&Maple,T.L.(1985).Dominanceinnonhumanprimates.InS.L.Ellyson&
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Mulac,A.(1989).Men
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Rizzolatti,G.,&Arbib,M.A.(1998).Languagewithinourgrasp.
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259).Chicago:Nelson-
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StevenM.BokerandJenniferL.Rotondo
thatmovementwithherorhisleftarm,thenthisdyadexhibitedspatiotemporal
mirrorsymmetry.Wewillexplorespatiotemporalmirrorsymmetryindyadsas
onepersonmimicsanotherwhiletheystandfacetoface.
Symmetryformationinconversation
Therearemanyfeaturesofconversationalbehaviorthatgiverisetosymmetry.The
moststrikingofthese,obvioustoeventhecasualobserver,isthatpeopletend
tomimiceachother
spostureduringconversation(Lafrance1985).Thismirror
symmetryinconversants
Symmetryinsynchronizedmovement

increases.Inthiswaywecanconsidertheretobeanincreasedredundancyinthe

StevenM.BokerandJenniferL.Rotondo
Symmetryinsynchronizedmovement
\b
ingeachother.Priortoeachtrialtheywouldbeinstructedovertheheadphones
headpositionwererelativetothepositionofthetrunk.Thenthevelocityalong
eachaxisforeachsensorwascalculatedas
+3)
where
)isthevelocityatsample
forsensor
alongaxis
,and
3)isthe

Thustherootmeansquarevelocity
)givesanestimateofthetotalactivityfora
dancerattime
.Whilethisoverallestimateofvelocitydoesnotgiveanyestimateof
\t
StevenM.BokerandJenniferL.Rotondo
andtheinstructioncategory.Therewasnosigni
canteffectofsexontheoverall
RMSvelocityduringthedance.Therewasaneffectoflengthoftherhythm(p
0.01)suchthatrhythmswith8beatspermeasureproducedhighervelocitiesthan
rhythmsoflength7.
Cross-correlationoftwodancers
velocities
Symmetryinsynchronizedmovement
\n
middleofatrialinwhichbothparticipantswereinstructedtolead.Thestrong
verticaldarkgraybands,whichpeaksareseparatedbyapproximately1400msand
occurduringtheinterval15secondstoabout27secondsfromthebeginningof
Time (s)
Time (s)
Time (s)
Time (s)
Cross Correlation
Lag (s)Lag (s)
b.
Figure1.
Cross-correlationmatricesplottedfortwentysecondsfromfourexampletri-
als.Thescaleontherightdenotesthecolorassignedtoeachvalueofcrosscorrelation.
(a)Thebeginningofatrialinwhichbothsubjectswereinstructedtofollow.(b)Twenty
secondsfromthemiddleofanexampletrialinwhichbothsubjectswereinstructedto
lead.(c
d)Thebeginningoftwoexampletrialsinwhichonesubjectwasinstructedto
leadandtheothertofollow.
StevenM.BokerandJenniferL.Rotondo
Implicationsformirrorsystems
Theresultsofthecurrentexperimentdemonstratethatwhenindividualscoordi-
natetheirmovementsinalead-followorlead-leaddancetoarepeatingrhythm,the
overallvelocityofeachindividualentrainsreliablywiththelengthoftheperceptual
segmentationfromtheauditorystimulus.Thus,cyclicmovementiscreatedwitha
frequencythatmatchesthefrequencyoftherepeatingrhythmandstable,reliable,
Symmetryinsynchronizedmovement
References
Bavelas,J.B.,Black,A.,Chovil,N.,Lemery,C.R.,&Mullett,J.(1988).Formandfunction
inmotormimicry:Topographicevidencethattheprimaryfunctioniscommunicative.
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Bertamini,M.,Friedenberg,J.D.,&Kubovy,M.(1997).Detectionofsymmetryand
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Kubovy,M.(1994).Theperceptualorganizationofdotlattices.
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Lafrance,M.(1985).Posturalmirroringandintergrouprelations.
PersonalityandSocial
PsychologyBulletin,11
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Lindenmayer,A.,&Prusinkiewicz,P.(1990).
Thealgorithmicbeautyofplants
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SpringerVerlag.
Palmer,S.E.,&Hemenway,K.(1978).Orientationandsymmetry:Effectsofmultiple,
Mirrorneuronssystemandtheevolution
ofbrain,communication,andlanguage
Ontheevolutionaryoriginoflanguage*
CharlesN.LiandJean-MarieHombert
UniversityofCalifornia,SantaBarbara,USA/LaboratoireDynamique
duLangage,Lyon,France
Originvs.evolutionoflanguage
Inrecentyearstherehasbeena
urryofscholarlyactivitiesontheoriginoflan-
CharlesN.LiandJean-MarieHombert
languageemerged,whereasthestudyoftheoriginoflanguageendsatthepoint
intimewhenlanguageemerged.Thisdistinctiondoesnotbelittlethesigni
cance
oftheresearcheffortprobingintotheolderandolderlayersofhumanlanguage.
Nordoesitdismisstheimportanceoftheproto-humanlanguageifandwhenits
featurescanbeinferred.Thedistinctionmustbemadeforoneimportantrea-
son,andthatisthewayscasual,spokenlanguagechangesandthewayshominid
communicativebehaviorchangesarefundamentallydifferent.
Theevolutionarychangeofcommunicationvs.linguisticchange
Thefundamentaldifferenceliesinthefactthatearlyhominidcommunicativebe-
havior,nothumanlanguage,issubjecttotheconstraintsofDarwinianevolution.
Inotherwords,theevolutionofourhominidancestors
communicativebehavior
Ontheevolutionaryoriginoflanguage
ofsymbolseachofwhichrepresentsdirectly,consistentlyandexclusivelyanentity
intheworld.Theemergenceofthe
rstsymboliccommunicativesignalamong
hominidsisnotonlyanimportantevolutionarylandmarkbutalsorepresentsa
quantumleapfromnon-humanprimatecommunication.Priortothislandmark
development,thecommunicativesignalsofhominidsshouldnotbequalitatively
differentfromnon-humanprimatecommunicativesignals.Non-humanprimate
communicativesignalsarenotsymbolic.Theyhavefunctions,notmeanings.Con-
CharlesN.LiandJean-MarieHombert
locationoftheconversation.Thisisthe
displacement
featureofhumanlanguage
Ontheevolutionaryoriginoflanguage
World Population (In Millions)
Time (Years Before Present)
Figure1.
Estimatesofhumanpopulation
TheFrenchpaleo-demographer,Jean-NoelBiraben,independentlyarrivedata
similarconclusioninhis
Essaisurl
evolutiondunombredesHommes
,(1979).
AccordingtoBiraben
sestimate,theworldpopulationincreasedby500%around
40,000yearsago.
Thepopulationexplosionduringtheperiodof60,000
40,000yearsagoisalso
con
\t\f
CharlesN.LiandJean-MarieHombert
skillsbeforethecreationoftheGrotteChauvetpainting.Personalornamenta-
lationchange.Itshowsalongstasischaracterizedbyarelative
atlineun-
tiltheendoftheMiddlePaleolithicandthebeginningofUpperPaleolithic
whenthecurvebeginstoshootupvertically.Figure2ismodeledafterLewin
(1993:33).
TheUpperPaleolithictoolsincludehaftedbladesthatareatleasttwiceaslong
astheyarewideandnumeroustypesofhaftedsmallgeometricallyshapedtools
suchaschiselsand
lesforcarvingandmakingboneinstruments.Theyindicate
alevelofsophisticationinvolvingdesignandsymbolismpreviouslyunattainedin
hominidhistory.
Question:Whatisthereasonbehindthisexplosivedevelopmentoftools?
60
50
40
Million ofyears before present
Oldowan
Acheulian
Mousterian
Upper Paelolithic
present
Figure2.
Numberofdistinctstonetoolsinhominidhistory
Ontheevolutionaryoriginoflanguage
\t
4.ThecolonizationofAustraliaoccurredapproximately60,000yearsago.At
thetime,becauseofglaciation,Australia,PapuaNewGuineaandTasmania
formedonecontinuouslandmass,whilemanyofthepresentdayislandsof
theIndonesiaarchipelagowereconnectedwiththeMalaysiapeninsulaofAsia.
ReachingAustraliafromAsiaentailedthecrossingofdeep,fast-movingocean
waterofapproximately100kilometers.Suchsea-crossingrequiredsocialorga-
nization,collaborativeeffort,sophisticatedplanning,someskills,equipment
andknowledgeofnavigation.
Question:Whatenabledhumanstocrossdeep,fast-movingoceanwateratthat
timebutnotbefore?
Tosumup,thesefourpiecesofevidencecollectivelypointtoanewcognitive
capacityforsophisticatedcultureemergingduringtheperiodof80,000
yearsago.Wecannotattributethisnewcognitivecapabilitytoalargerbrain,be-
causehumancranialcapacity,ifanything,hasdecreasedsincethedawnofanatom-
icallymodernhumanataround150,000
120,000yearsago.Infact,thesigni
cant
\t
CharlesN.LiandJean-MarieHombert
minidcommunicativebehavior?Accordingtoourresearch,therearethreemecha-
nismsandfourprocessesofevolutionthatareespeciallyimportant.Wewillbrie
sketchthesethreemechanismsandfourprocesses.
Fourevolutionaryprocessesleadingtotheemergenceoflanguage
Reductionofthegastrointestinaltract
Thereductionofthegastrointestinaltractisanecessaryconcomitantdevelopment
oftheincreaseinencephalizationinhominidevolution.Thereasonisthatanen-
largedbrainconsumesanenormousamountofenergythathastocomeatthe
expenseofsomepartofahomeostaticsystemofthehominidanatomy.Thebrain
ofanewborninfant,forinstance,consumes60%oftheenergyittakesin.Leslie
Ontheevolutionaryoriginoflanguage
\t
Enlargementofthevertebralcanal
\t
CharlesN.LiandJean-MarieHombert
Iamhungry
,isliterally
Ihavehunger.
IfaSpaniardsaysinEnglish,
Ihave
hunger
tomean
Iamhungry
,theSpaniardhasnotmadeagrammaticalmis-
takeinEnglish.Hisutteranceissimplyunidiomatic,i.e.notinaccordwiththeway
nativespeakerssayit!
Mostpolyglotshavewitnessedinterestingandamusingexamplesofunid-
iomaticutterancesbynon-nativespeakers.Theimportantpointisthattheways
ofsayingthingstendtobeuniquetoalanguageoragroupofcloselyrelated
languages,andtheyarenotcon
nedtoafewspecialexpressions.
TheNewZealandlinguist,AndrewPawley,haswritteneloquentlyaboutthis
aspectoflanguage.Sohashisteacher,theAmericanlinguist,GeorgeGrace(1987).
IwillquotefromanarticlebyPawley(1991),
Alanguagecanbeviewedasbeing(amongotherthings)acodeforsaying
things.Thereareanumberofconventionsthatconstrainhowthingsshould
besaidinalanguagegenerallyorinparticularcontexts.HereIwillmention
onlythegeneralmaxim:
beidiomatic
.Thismeans,roughly,thatthespeaker
(author,translator,etc.)shouldexpresstheideaintermsthatnativespeakers
areaccustomedto.Forexample,ifyouaskmethetimeandmywatchshows
thelittlehandpointingjustpastthe5andthebighandpointingtothe2,an
idiomaticanswerwouldbe
stenpast
,or
veten
.Areplysuchas
veo
clockandonesixth
ve-sixthofanhourtosix
ssixless
fty
wouldnotcountasidiomatic.Tobreaktheidiomaticityconvention
istospeakunnaturally.(Pawley1991:433)
Theimplicationofthisimportantcharacteristicoflanguageisthatlinguisticbe-
haviorrequiresaprodigiousmemory.Theneocortexofourbrainmustbeableto
storeavastamountofknowledgeacquiredthroughlearning:thevocabulary,the
grammar,andthemyriadwaysofsayingthings.Wewishtoemphasizethatthis
Ontheevolutionaryoriginoflanguage
\t
ofthehumanbraininontogeny,whichchannelsthehumaninfant
sattentionto
thelinguisticandsocialinteractionofhis/herenvironmentandenablesthehu-
maninfanttolearnacomplexsymbolicbehaviorrequiring,amongotherthings,
aprodigiousmemory.Theacquisitionoflanguageis,then,acomplexinterplay
\t
CharlesN.LiandJean-MarieHombert
Amajorillustrationwouldbethedevelopmentofanotherwisenormalembryo
ofnewphenotypicmanifestationandpossiblyanewspecies.Theconsequenceof
duplicatingthegenesregulatingstructuraldevelopmentisthattheresultingstruc-
turesshouldalsoshowsignsofduplication.However,theduplicatedstructures
willbemodi
ediftheduplicatedgeneshaveundergonemutation.Anobviousex-
ampleofrepeatedstructuresinhumansarethevertebratecolumn.Thebrainalso
containsmanyrepeatedstructures,forexample,theradialunitsoftheembryoin
itsearlystageofdevelopment,whichareultimatelyresponsibleforthesizeandar-
chitectonicpatternoftheneocortex(Rakic1988).Theserepeatedstructurescould
Ontheevolutionaryoriginoflanguage
\t\b
Changeofthedevelopmentalclock
Thesecondevolutionarymechanismconcernsthechangeofthedevelopmental
clock.Developmentalclockdesignatesthelengthofontologicaldevelopmentofan
animaloranorganofananimal.
\t\t
CharlesN.LiandJean-MarieHombert
thedirectionoftheevolutionoftheirownspecies.Bynowweknowthatamong
Ontheevolutionaryoriginoflanguage
\t\n
wouldconferasigni
CharlesN.LiandJean-MarieHombert
languagemodule.Webelievethatitdoesnot.Insteadofpostulatingalanguage
moduleinmybrain,wewouldliketointroducetheconceptof
cognitivereserve
Cognitivereserve
Bycognitivereserve,wemeancognitivecapabilitythatisnotfullyutilizedorman-
ifestedinthenormalrepertoireofbehaviorofamammal.Thevariousprojects
traininggreatapestomanipulatelinguisticsymbolsareevidencefortheapes
cog-
nitivereservespeci
callyinthedomainofsymboliccommunication.Regardlessof
thecontroversysurroundingthedegreeofsuccessoftheseprojectsinvolvingthe
chimpsSarahandWasho,thebonoboKanzi,thegorrillaKoko,thereisnodoubt
thatthesegreatapesareabletoacquireandusesomelinguisticsignsafterextensive
andintensivetraining.Itistruethatintheirnaturalenvironment,apes
commu-
nicationisstrictlynon-symbolicandtheygivenoindicationofdevelopingsym-
boliccommunication.Itisonlythroughhumaninterventionthattheysucceedin
acquiringsomelinguisticsymbols.Theimportantpointisthattheyhavethecog-
nitivereserveforacquiringandusingsomelinguisticsymbolseveniftheprocess
ofacquisitionishighlyunnatural.Weknowmanymammalscanbetrainedbyhu-
Ontheevolutionaryoriginoflanguage
offashioningatwig/strawintoaprobefor
shingouttermitesbythechimpsof
Gombearealsostrokesofgeniusthatattesttotheexistenceofcognitivereserve.
Theimportanceoftheconceptofcognitivereserveandtheearlierdiscussion
CharlesN.LiandJean-MarieHombert
howtheythinktheyutterasentenceintheirownnativelanguage.Suchasentence
is,atbest,atokenoftheformalwrittenlanguageratherthancasualspokenlan-
guageformanyreasonsofwhichthemostimportantoneisthatsuchasentence
isindependentofanycommunicativecontext.Theentirecommunicativecontext,
linguisticornon-linguistic,visual,auditoryortactile,ofanycasualconversation
servesassucharichsourceofinformationtotheinterlocutorsthatrendersthe
grammar,thedictionandtheorganizationofcasualspeechsigni
cantlydifferent
fromthoseofwrittenlanguage.
Towardthecrystallizationoflanguage
rststageoftheevolutionarydevelopmenttowardcasualspokenlanguage
istheincreaseofcommunicativesymbolsforconcreteobjects,e.g.food,preda-
tor,objectsforlandmark,differentanimals,differentplants.Duringthisstage,the
creationofeachnewsymbolrepresentsastrokeofgeniusbyahominid,andthe
establishmentofeachnewlycreatedsymbolintherepertoireofthecommunica-
tivesignalsofthesocialgrouptowhichthecreatorbelongs,requiressocialand
culturaltransmission.Thesocialgroupmostlikelyconsistsofclosekininthebe-
ginningbeforeitextendstoamoredistantlyrelatedclan.Itisimportanttorealize
thattheentireprocessisanevolutionaryevent.Itdidnothappeneveryday.It
didnothappeneveryyear,anditprobablydidnothappenineverygeneration.
Wemustavoidunconsciouslyprojectingourframeofmindontotheevolutionary
sceneinvolvingourhominidancestors.Theyhadneitherthecognitivecapability
northeculturalenvironmentwehave.Theywereatthebeginningofalongevo-
lutionarypaththatultimatelyledtotheemergenceoflanguage.Theydidnothave
languageyet.Webelievethattheonsetofsymboliccommunicationbeganwith
Homoerectus.Thereareseveralreasonsforourbelief:(a)TheHomoerectusbrain
at800
950ccisconsiderablylargerthanthebrainofallearlierhominids,includ-
ingHomohabilis,the
rstspeciesoftheHomogenus.(b)TheHomoerectusbrain
showsanincreaseincerebralasymmetries.(c)Theyarethe
rsthominidswhich
migratedoutofAfricaandreachedasfarasAsiaandIndonesiaevidencedbythe
famousfossilsofPekingmanandJavaman.Themigrationsuggestsanexpanding
population,whichinturn,suggestsahigherlevelof
tness,probablycausedby
improvedcommunicativecapability.(d)AsHolloway(1995)pointsout,sincethe
timeoftheHomoerectus,theevolutionofthehominidbrainshowedagradual
increaseinvolume,re
Ontheevolutionaryoriginoflanguage
related.Inotherwords,theHomoerectusbrainevolvedexclusivelyforthepurpose
ofgreatercognitivecapacity.Webelievethatthisevolutionaryprocessofthebrain
iscorrelatedwiththegradualevolutionofthesymboliccommunicativebehavior.
Havingafewcommunicativesymbolsforconcreteobjects,however,
isnottan-
tamount
tobeingawareoftheabstractprincipleofassociatingsymboliccommu-
nicativebehaviorwithconcreteobjects,eventhoughthesymbolitselfisatoken
ofthisprinciple.Inotherwords,thereisasigni
CharlesN.LiandJean-MarieHombert
communicativesymbols,andbecausetheincreaseofmirrorneuronsimprovedthe
speedandcapacityforlearning.
Enhancementofculturewasnecessarybecause
itfacilitatedthespreadofnewlycreatedlinguisticsymbol.Growthofthesizeof
socialgroupandpopulationwasnecessarybecausethemorehominidsacquired
linguisticcommunicativesymbols,themorelikelyanewgeniuswouldemergeto
createanadditionallinguisticsymbolforanotherconcreteobjects.Theanatomi-
calinnovationsaswehavealreadypointedout,alsoinvolvethedecreaseoftheG.I.
tract,theexpansionofthethoracicnerves,thedecreaseinsomatizationandthe
descentofthelarynx.Thesechangesemergedthroughaco-evolutionaryprocess.
Theydidnotoccurinafewgenerations.Ontheonehand,thedevelopmentofho-
Ontheevolutionaryoriginoflanguage
activityoractionimpliestheexistenceofanagentoranactor.Ifahominidhad
awordforanaction,wecanassumethatthehominidalreadyunderstoodthat
anactionrequiredanactortoexecuteit.Asforstabilizingawordorder,itisa
socialconvention,negotiatedconsciouslyorunconsciouslybythemembersofa
community.
Whataboutalloftheothergrammaticalstructuresbeyondwordorderfound
incontemporaryspokenlanguages?
Wehavebynowhistoricallinguisticdatathataccountfortheemergenceof
nearlyallgrammaticalconventions,beitin
ection,derivation,subordination,
conjunction,interrogative,imperativeorsubjunctive.Linguistshavebeenableto
elucidatethepreciseprocessesandmechanismsbywhichsuchgrammaticalcon-
structsmayemergeinalanguage.Grammaticalizationisoneofthemostimportant
mechanismsintheemergenceofgrammar(Traugott&Heine1991).Grammatical-
izationbegantooccurasthehominidsstartedtolinksymbolicsignalsintolarger
communicativeunits.
Whataboutthenotionofgeneratingsentencesthatisthefoundationofgen-
erativegrammar?
Inourdiscussionofthede
ningcharacteristicsoflanguageatitscrystalliza-
tion,wedidnotmentionrecursivefunctionorgenerativity.YeteversinceChom-
sky
sfamouspublicationof
SyntacticStructures
,manyscholarsincludingmost
linguistsconsiderrecursivefunctiontheuniquede
ningfeatureofhumanlan-
guage,e.g.Pinker(1995).Indeed,ifonesurveystheliteratureonlanguage,one
cannotfailtonoticetheomnipresenceoftheconceptofrecursivefunctionorgen-
erativity.Itdepictsthespeaker
sabilitytogenerateaninde
nitelylargenumberof
sentencesfroma
nitevocabularywitha
CharlesN.LiandJean-MarieHombert
Nowsupposeinanarti
ciallanguagewehavetwore-writingruleswhichcan
beappliedrepeatedly:
Ifweapplythe
rstrule,weobtainasentenceconsistingofone
.Ifweapplythe
secondruleandthenapplythe
rstruletotheoutputofthesecondrule,whichis
,weobtainastringoftwo
s,namely,
.Ifweapplythesecondruletwice,
the
Ontheevolutionaryoriginoflanguage
writtenlanguage.
Inde
nitelylargenumberofsentences
nitelylongsen-
tences
are
theoretical
nitesetofpossiblehumpbackwhalecourtshipsignals,eachof
whichrepresentsapointalongthecontinuumofthemale
semotionalstateand
hisphysical
tness.Thisin
CharlesN.LiandJean-MarieHombert
tensingscontinuouslyforseveralhours.Thestrongerandhealthierthebird,the
Ontheevolutionaryoriginoflanguage
ofwhichareavailableinspeechinteraction.Writtenlanguageisnottheresultof
biologicalevolution.Aswehavepointedout,itisaculturalproduct.
Finallywewishtopointoutthatmuchofwhatgenerativelinguistsconsideras
canonicalgrammaticalconstructionsareformalizedorconventionalizedinwrit-
tenlanguage.Sincemostlinguistsspeakortrytospeakinthestyletheywrite,the
canonicalgrammaticalconstructionsaretransportedintotheirspokenlanguage.
Theymaybelievethatsuchgrammaticalconstructionsarethementalprototypes
ofthelanguage,andthedataofcasualspokenlanguagerepresentwhatChomsky
(1965)callsthe
degenerate
datathatarefragmentsofthefullforms.Thisbelief
isfurtherreinforcedbythefactthatformalwrittenlanguagecarriesgreatersocial
prestigethancasualspokenlanguagebecauseofthewrittengenre
sassociations
withliteracy,educationandsocialstatus.Thus,theperceptionthatthecanonical
grammaticalconstructionsofformal,writtenlanguagearethementalprototypes
oflanguageisbasedonasocialprescriptionenforcedthrougheducation,literacy
andvaluesystem.Fromtheperspectiveofevolution,languageis
rstandforemost
ahumancommunicativebehavior.Ifwearetostudythehumancommunicative
behavior,wemustbaseourstudyoncasual,spokenlanguagetranscribedwiththe
utmost
delityandnotviewedasfragmentsofstylisticconventions.Casualspoken
languagedoesnothavethegrammarofformalwrittenlanguage.Inaforthcom-
ingarticle,PaulHopper,afterdescribingandanalyzingdataonseveralsyntactic
constructionsfromaspokenEnglishcorpus,concluded,
Corpusstudiessuggestthatthe
degenerate
dataarethetruesubstanceof
naturalspokenlanguage,andthatwhatmygrammarsgivemethenarenor-
mativizedassembliesofthesefragmentsthattendtoimpressthemselveson
measmentalprototypesbecauseoftheirgreatersocialprestige.
(Hopper,forthcoming)
nditironicthatintheempiricalinvestigationoflanguage,whichisahuman
behaviorwithanevolutionaryhistory,itisnecessarytodefendtheimportance
ofauthentic,uneditedbehavioraldatacollectedfromcasual,spokenlanguage.
Obviouslyformanylinguists,suchdataarefragmentedandunimportant.Suchan
attitudehasimpededtheinvestigationoftheevolutionaryoriginoflanguage.
Inconclusion,thispaperrepresentsacondensationof6millionyearsofho-
minidevolutionandasketchofadiversearrayofinformationfrommanydisci-
plinesthatarerelevanttotheevolutionaryoriginoflanguage.Manyimportant
topicsareleftoutandmanyothersreceiveonlyabriefcursorypresentation.Asa
consequence,thepaperremindsmeofanoldChinesesaying:
Flowerappreciation
onagallopinghorse.
Forblurredimagesandobscurelandscape,weapologize.
However,itisimportanttonotethatduringthepastdecademajorcontributions
towardanunderstandingoftheoriginoflanguagehavecomeprimarilyfromthe
neurosciencesandpaleoanthropology.Wehopethatwearesuccessfulindemon-
\f\f
CharlesN.LiandJean-MarieHombert
stratingthatlinguisticscanalsocontributetowardanunderstandingoftheorigin
oflanguage,oncewemovebeyondthemistcreatedbythegenerativeparadigm.
Notes
*TheauthorsaregratefultoCNRSforsupportingtheirresearchontheoriginoflanguage.
WearealsogratefultoJoanBybee,NoamChomsky,PaulHopper,FritsKortlandt,Guido
Martinotti,AlainPeyraube,MaximStamenov,EddaWeigandandBruceWilcoxfortheir
invaluablecommentsandsuggestions.
Itwillbeclearlaterinthispaperthatthede
nitionoflanguageasthecasualspoken
languageofhumanbeingsisofextremeimportance.Iusetheterm
anatomicallymodern
humans
tocircumventtheconfusioncausedbyaproliferationoftaxonomictermssuchas
EarlyHomosapiens,ArchaicHomosapiens,Homosapiensapiens,etc.Comparedtothe
hominidsofthepast250,000years,anatomicallymodernhumanshaveagracileskeleton
characterizedbylongboneshape,aspeci
cdepthandextentofmuscleinsertion,athin
cranialwallandmandibularbody,ahigh,domedcranium,areducedjaw,andtheabsence
ofaprominentbrowbridgeovertheeyebrow,i.e.nosupraorbitaltorus.
Forasuccinctandcomprehensiveanalysisofthecurrenthominintaxonomy,seeWood
andCollard(1999).Anewdiscovery,however,posesadditionalchallengetothealready
controversialhominintaxonomy.OnDecember4,2000,FrenchandKenyanpaleoanthro-
pologistsannouncedthediscoveryof
MillenniumAncestor
(Orrorintugeensis)inthe
TugenhillsofKenya
sBaringodistrictintheGreatRiftValley.Thefossilremainsinclude
variousbodypartsbelongingto
Ontheevolutionaryoriginoflanguage
\f
Non-humanprimatecommunicativesignalsaretypicallymulti-modal,involvingvisual
aswellasauditory,andsometimestactilechannels.Buttheirvisual(suchasfacialex-
pression,bodypostures)andtactilecommunicativesignalsareevenmoretransparentas
manifestationsoftheiremotionalandmotivationalstates.
Thedeceptiveuseofcommunicativesignalsamongprimates,whichhasbeenobserved
amongseveralspecies,wouldinvolvecognitionbeyondtheinvoluntaryvocalizationstim-
ulatedbyanexternalcircumstance.Deception,however,isnotfrequentlyobservedamong
primates,eventhoughitsuggeststhattheuseofacommunicativesignalfordeceptionis
subvertedbytheneo-cortex.
Thereisanamusingincidenceinvolvingachimpdiscoveringacacheofdelectablefood
atJaneGoodall
scampinGombe.Itimmediatelywentbehindatreeandcovereditsmouth
sothatitsinvoluntaryfoodcallcannotbeheardanditsfacialexpressionofexcitement
cannotbeseenbyitscompanions.Thisepisodeissigni
cantbecauseitdemonstratesthat
(1)thechimpisawareofitsownemotionalreactiontothesuddendiscoveryofdelectable
food,and(2)throughitsneocortex,itistryingtoconcealitsemotionalstatesexpressedby
itscommunicativesignals.Similarincidencesinvolvingotherprimateshavebeenreported
byethologists,forexample,CheneyandSeyfarth
s(1990)accountofdeceptionbyvervet
monkeys.
Thechangeofsomeanimalcommunicativesignalsmaybeculturallytransmitted,e.g.
thecourtshipsongsofthewhite-crownedsparrowareknowntohavedialectaldifferences.
Insuchcasesofthechangeofanimalcommunicativebehavior,theclassicalDarwinian
evolutionaryprocessdoesnotapplyandthetimeofchangemaybeveryshort.
Iflanguagecrystallizedseveraltensofthousandsofyearsaftertheemergenceofanatom-
icallymodernhumans,thepolygenesisoflanguagewouldbepossible.Theissueofmono-
genesisvs.polygenesisoflanguageisbrie
ydiscussedinLi(2002,2003).
Them-DNAcontainsonly37genesand16569basepairs.Thesmallnumberofgenes
andbasepairsmakeiteasytoexaminethevariabilityofm-DNAindifferentindividuals.
Mostimportantofall,mitochondrialgenesarematernallytransmitted,althoughrecentin-
vestigationsshowthatrareleakageofpaternalm-DNAintoafertilizedovumispossible.
Ifthesourceofm-DNAisexclusivelymaternal,thenvariationofthem-DNAcanonlybe
causedbymutation.Thusamolecularclockbasedonanaveragemutationrateinthem-
DNAtendstobereliable.ForaninformativediscussionofthemitochondrialDNAandits
relevancetohumanevolution,seeCann(1995).
\f\r
BeforeTimWhiteunearthedthefossilsofAustrolopithecusGhariinEthiopiain1997,
theOldowaninKenyaistheoldestknownstonetooltechnology.TheOldowantoolsdate
from2.5to1.7millionyearsago,andtheyareassociatedwiththeemergenceofthegenus
Homo.However,AustrolopithecusGhari,whichisdated2.5millionyearsago,usedstone
toolswhichwerecarriedfromasitemorethan50milesawayfromthelocationoftheGhari
fossils.Thisdiscoverynulli
edthelong-standingbeliefthatstonetoolswereaHomoin-
vention.TheAcheuliantechnologyemergedwiththeHomoerectus.Themajordifference
betweentheOldowanandtheAcheulianistheadditionofthehandax,thecleaverand
thepickintheAcheuliantechnology.TheMousteriantechnologycontainedalargerrange
oftooltypesthantheAcheulian.However,theMousteriantechnology,associatedwiththe
Neanderthals,didnotexhibitmuchtechnologicalimprovementovertheAcheulian.
\f
CharlesN.LiandJean-MarieHombert
Foraninformativediscussionoftheevolutionofthehumanbrainandacomparisonof
thehumanbrainwithanimalbrains,seeFalk(1991)andRoth(2000).
\r
Someanimalshaveabettermemoryforcertainsensoryexperiencethanhumans.For
example,dogsandcatsarebetterthanhumansinrememberingolfactoryexperience.This
fact,however,doesnotimplythatdogsandcatshavealargercapacityforcognitivememory.
Theirolfactoryperceptionismuchmoreacutethanthatofhumanbeings.Theirgreater
abilitytoperceiveanddifferentiateodorsisconnectedtotheirbetterolfactorymemory.
Forasuccinctsummaryofchildren
sacquisitionofgrammar,seeBatesandGoodman
\r
WetakenoteofthefactthatChomsky
scurrenttheoreticalstanceisconsiderablydif-
ferentfromhis1986pronouncements.InhisnewMinimalistProgram(Chomsky1995),
grammarislargelyderivedfromthelexicon.Ifwearecorrectinassumingthatwhatiscon-
sideredinnatebyChomskyandhisfollowersisthenewestversionoftheso-called
Uni-
versalGrammar
,whichisaustereandminimal,theissueofrepresentationalinnatenessfor
languagebehaviorispracticallymoot.
Insomegrammars,one
ndssomesporadicdiscussionofsomeparticularwaysofsay-
ingthings.TypicallysuchagrammarconcernsalanguageunrelatedtotheIndo-European
languagefamily.Theauthorsaremotivatedtodiscusssomewaysofsayingthingsinthose
languagesbecausemanyofthesewaysofsayingarebizarrefromtheIndo-Europeanper-
spective.Forexample,inmostgrammarsofSub-SaharaAfricanandEastAsianlanguages,
serialverbconstruction
isusuallypresentedbecauseitisaconstructionthatdoesnotoccur
inIndo-Europeanlanguages.
\r
Agenecarriestheinformationforcodingaparticularprotein,andeachproteinplaysan
importantroleintheanatomyandphysiologyofananimal.Hencethemutationofagene,
Ontheevolutionaryoriginoflanguage
\f
intheset
canleadtoconfusion.This
totalnumber
isnolongeraninteger.Itisanin
nity,
ormoreprecisely,acountablein
nity.Thereareotherkindsofin
nitiesinmathematics.
Noneofthemisaninteger.
 \r
Therearebynowmanycorporaofcarefullytranscribeddataofcasualconversations.
OnecorpususedbymeistheSantaBarbaraCorpusofSpokenAmericanEnglishcreatedby
JohnDuBois(2000).
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Mirrorneurons,vocalimitation,
andtheevolutionofparticulatespeech
MichaelStuddert-Kennedy
HaskinsLaboratories,NewHaven,CT,USA
\f\t
MichaelStuddert-Kennedy
ception,foritisthere,notintheremoterecessesofthecognitivemachinery,that
thespeci
callylinguisticconstituentsmaketheir
rstappearance
(1996:31).What
followsisabriefaccountofwhat
Mirrorneurons,vocalimitation,andtheevolutionofparticulatespeech
\f\n
permutedandcombinedtoformanunlimitedvocabularyofmeaningfulunits
(morphemes,words);attheupperlevel,syntax,themeaningfulunitsarerepeat-
 \f
MichaelStuddert-Kennedy
TTCD
TTCL
GLO
glottis
velum
tongue tip
tongue
body
center
jaw
+ lower lip
TTCL
lip protrusion upper & lower lips,jaw
lip aperture
tongue tip constrict location
upper & lower lips,jaw
tongue tip,tongue body,jaw
tongue tip constrict degree tongue tip,tongue body,jaw
tongue body constrict location tongue body,jaw
tongue body constrict degree tongue body,jaw
velic aperture velum
glottal aperture glottis
TTCD
GLO
tract variable
coordinative structure
Figure1.
Tractvariablesandarticulatorsinamodelofspeechproduction.
Mirrorneurons,vocalimitation,andtheevolutionofparticulatespeech

temtheparametersspecifytheprimaryend-effectoroftheconstriction(lips,
 
MichaelStuddert-Kennedy
wide
wide
wide
wide
alveolar
alveolar
alveolar
alveolar
[nt]
[dnt]
narrow
uvular
narrow
uvular
VELIC
TB
TT
LARYNGEAL
LARYNGEAL
Figure2.
Gesturalscoresfortheadulttarget,
nt](above),andforthechild
erroneousattempt,[
nt](below)
tionofthesamearticulatorelicitedinstantresponsefromadifferentcooperating
articulator,asafunctionofgesturalgoal.
Abstractly,then,agestureisacoordinativestructurecontrollinganequiva-
lenceclassofarticulatormovements,whereequivalenceisde
nedbyfunctionor
goal,theachievementofacertainvocaltractcon
guration.Thus,wearriveatthe
rstrequirementofanyhypothesisconcerningtheroleofmirrorneuronsinlan-
Mirrorneurons,vocalimitation,andtheevolutionofparticulatespeech
 
Behavioralevidenceforgesturesfromspeecherrors
Adults:Randomspeecherrorsinthelaboratory
 
MichaelStuddert-Kennedy
the2nd
3rdyearoflife(Jaeger1992a,b).Whatweoftensee,then,inachild
searly
wordsareunsuccessfulattemptstoachievetheamplitude,duration,andrelative
phasingofgesturesforwhichthechildhassuccessfullyrecognizedboththeend-
effectors(lips,tonguebody,tonguetip,velum,larynx)and,fororalgestures,their
roughlocusofconstriction.
WedrawourexamplesfromastudyofaNorthAmericanchild,
Emma
,con-
ductedovera4monthperiodaroundhersecondbirthday(Studdert-Kennedy&
Goodell1995).Noneofthewordsdiscussedbelowimmediatelyfollowedanadult
modelutterance;allwordswerethereforespontaneousdelayedimitationsdrawn
fromthechild
slongtermmemory.Emmahadalreadydevelopedafavoriteges-
turalroutine,a/labial-consonant/vowel/alveolar-consonant/vowel/sequence,that
Mirrorneurons,vocalimitation,andtheevolutionofparticulatespeech
 
Table1.
Sometargetwordsandatwo-yearoldchild
sattemptstosaythem,illustrat-
ingparadigmaticandsyntagmaticgesturalerrors(fromStuddert-Kennedy&Goodell
Targetword
ConsonantgesturesinadultformChild
sattempts
raisin
rezn]
syllableemergedas[d
nt].Theswitchfrom[n
t]to[d
nt]evidentlyfollowsfrom
asimpleerroroftiming:velicopeningslidesawayfromsyllableinitial[n](sothat
alveolarclosureandreleasenowyield[d])intoalignmentwithalveolarclosure
(butnotrelease)for
nal[t],yielding[nt].
Thenextthreewordsaremorecomplicated.Theperhapsunexpectedwords,
elephant
hippopotamus
,werenamesforpicturesinabook.Forbothwordsthe
childadoptedatacticthatshefavoredforwordsorphrasesofthreeormoresyl-
lables:sheloweredherjawandsubstitutedthewidepharyngealgestureofvocalic
]asasortofplace-holderfortheinitialsyllableorsyllables.Here,
veattempts
at-
nt])allincludealabialgesture,butonly[

n]achievesthecorrect
labiodental[f],only[

bin]and[

pin]therequiredaccompanyingglottalabduc-
 
MichaelStuddert-Kennedy
tion.Otherattemptsarevoicedthroughoutandincludeanintrusivehighfront
vowel,[i],harmonizingwithalveolarclosurefor[n];intwoattempts,[
bin]
and[

min],velicloweringfor[n]slidesintoalignmentwithlabialclosurefor[f],
yielding[m].
For
hippopotamus
,Emmasubstitutedheridiosyncratic[
]forthe
rsttwo
syllables.Shethencompressedtheremainingthreesyllables,-
potamus
,intoone,
nz],builtaroundherlabial-alveolarroutine.Figure3illustratesthesurprisingly
simpleprocessbywhichshemayhaveaccomplishedthetransformation.Shecor-
rectlyexecutedlabialclosureandglottalabductionfor[p]andalveolarclosurefor
[t],thuslinkingthelipandtonguetipgesturesofherfavoredroutine,asindicated
bythearrowsinthe
gure.Sheomittedlabialclosurefor[m]andglottalabduc-
tionfor[t]and[s];sheomittedthelowbacktonguebodygesturesforthevowels,
roughlyharmonizingthevocalicnucleustothefollowingalveolarclosure;andshe
allowedvelicloweringforomitted[m]toslideintoalignmentwithalveolarclosure
wide
wide
wide
wide
wide
alveolar
crit
alveolar
crit
alveolar
alveolar
...potamus
[pnz]
narrow pharyngeal
VELIC
TB
TT
LARYNGEAL
LARYNGEAL
Figure3.
Gesturalscoresforthe
Mirrorneurons,vocalimitation,andtheevolutionofparticulatespeech
 \b
for[t],yielding[n].Theoutcomeofthesemaneuverswas[p
nz],asyllablecom-
 \t
MichaelStuddert-Kennedy
Table2.
Initialconsonantsinearlywordsof4English-learningchildreninthreehalf-
hoursessionsduringtransitionfrombabblingtoworduse(13
16monthsapproxi-
mately):TabulationofdatafromStanfordChildPhonologyProjectinAppendixCof
Vihman(1996).
Bilabial[p,b,m,w]883463
Alveolar[t,d,n,r,l,s,z]4614251
86
Velar[k,g]
27490
40
Total
16121804
%oferrors
20764
accuracy.Errorsinglottalandvelicgesturestendtobeerrorsofphasingratherthan
omission;andwhileerrorsofpreciselocationdooccurfororalgestures(e.g./s/for
/,/p/for/f/),theend-effector(lips,tonguetip,tonguebody)andsotherough
locusofconstriction(i.e.placeofarticulation)tendstobecorrect.Inotherwords,
gesturalerrorsonconsonantstendtobeerrorsofamplitudeorphasingratherthan
ofomissionorend-effector.Table2lendsfurthersupporttothisclaimwithdata
from4childrenintheStanfordChildPhonologyProject(Vihman1996,Appendix
C):80%ofsinglegestureerrorsonvoicing(glottalaction)ormanner(primarily,
gesturalamplitude),20%onplaceofarticulation.
Mirrorneurons,vocalimitation,andtheevolutionofparticulatespeech
 \n
Howmightsomatotopicrepresentationofthevocaltracthavearisenevolu-
MichaelStuddert-Kennedy
routinestoimitatenewwords,demonstratesthegenerativecapacitynecessaryfor
buildingalargevocabularyfromasmallrepertoireofgestures.
Mirrorneurons,vocalimitation,andtheevolutionofparticulatespeech
nativestructurethatcontrolsexecutionofthatmovementorgesture.Ifwefurther
assume,withRizzolattiandArbib(1998),thata
closedsystem
ofexpressivefa-
cialcommunicationwasanevolutionaryprecursorof
theopen[i.e.particulate
andcombinatorial]vocalizationsystemweknowasspeech
(p.192),howmight
thetransitionfromclosedanalogfacialcon
gurationtoopendigitalvocalgesture
havecomeabout?
Fromanalogfaceandcrytoparticulatespeech
Bothhumanandnon-humanprimatesexpressandcommunicatetheiremotions
bybothfaceandvoice(Darwin1872/1965;Hauser1996).Indeed,thetwomodes
aresointricatelyrelatedthatweoftencannottell
MichaelStuddert-Kennedy
Mirrorneurons,vocalimitation,andtheevolutionofparticulatespeech
indeedperhapsoftenstilltoday,brachio-manualimitationmighthavebeenmedi-
atedthroughamirrorsystemforrepresentingaholisticfunction,suchasthrowing
astone,graspingabranch,seizingfood.Suchactswouldhavebeenmotivatedby
recognizingthegoalsofconspeci
cacts.Theformoftheimitatedactswouldthen
havebeenshapedautomaticallybytheactor
smorphologyandbythedemands
ofthephysicalcontext.As
ngers,thumbs,wristscametobeengagedinmore
andmorediversephysicalacts,theywouldhavebecomeincreasinglydifferentiated
andcapableofindependentaction,freetoengageinarbitrarynon-iconicconven-
MichaelStuddert-Kennedy
(MacNeilage&Davis2000).Earlyhominidsthusfoundthemselveswithasurfeit
ofsyllablesinsearchofmeaningsratherthanofmeaningsinsearchofanexpres-
sivemode.Differentiationofthesyllableintoitscomponentgestureswouldhave
followedunderpressureforspeedandeconomy,againmuchasweobservethe
processinachild
searlywords,describedabove.
Beallthisasitmay,whatevertheoriginoftheshiftfromiconictoarbi-
trarysymbolicreference,priorevolutionofparticulatedmirrorsystemsforhand
andfacewouldhavefacilitatedevolutionofvocalimitation,andsoofparticulate
speech,byofferinganeuralmechanismthatthevocaltractcouldcoopt.
Conclusion
Theproposedevolutionarypathfrommanualtofacialtovocalimitationrests
onthehypothesisthatallthreemodesofimitationaremediatedbymirrorneu-
ronsystemsofrepresentation.ThehypothesisisconsistentwithAlvinLiberman
Mirrorneurons,vocalimitation,andtheevolutionofparticulatespeech
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Constitutivefeaturesofhuman
dialogicinteraction
Mirrorneuronsandwhattheytell
usabouthumanabilities
EddaWeigand
ArbeitsbereichSprachwissenschaftWestf
lischeWilhelms-Universit
nster,Germany
Mirrorneuronsandhumanabilities
Thediscoveryandexperimentalproofofso-calledmirrorneuronsseemtofocus
theinterestofvarioussciencesonthe
centralquestionofhumanabilities
(Rizzolatti
&Arbib1998).Humanbehaviouristhejointinterestofallofuswho,comingfrom
differentdisciplines,areparticipatinginthisconferenceonmirrorneuronsandthe
evolutionofbrainandlanguage.Humanbehaviourfromalinguisticpointofview
canbeconsideredtobehumandialogicinteraction.In20thcenturylinguistics
therearemainly
twoconceptsoflanguage
:thearti
cialconceptoflanguageasa
signsystemandthenaturalconceptoflanguage-in-use.Ifweaccepttheviewthat
inlanguageusewearedialogicallyinteracting,thenaturalconceptoflanguage
usecanbeidenti
edashumandialogicinteractionoras
parole
performance
\f
EddaWeigand
Inscience,thechallengeisthesamefordifferentdisciplines.Atthebeginning
ofthenewmillennium,weare
nallyabletoexpressthischallengemoreprecisely:
itisthe
challengeofaddressingthecomplex
.Everydisciplineselectsapartofthe
complexwhichsurroundsusasitsobjectofstudy.Classicaltheorizingtoldusnot
toaddressthecomplexmixoforderanddisorderdirectlybuttoabstractfromdis-
orderandtoestablishalevelbelow,inlinguistics
lalangue
or,withsomemodi
cation,
biological
physiological structure
cognitive function
perceptual function
Figure1.
Mirrorneuron
Thereisnotasimpleuniformunit
attheoutset,buttheintegrationofdifferent
dimensions
,differentabilities,thematerialaspectsofbiologyandtheimmaterial
functionsofintentionandperception.Referringtothebasicconceptofconscious-
ness,MaximStamenov(1997:278)problematizesthebeliefinasingleandreason-
ablywell-de
Constitutivefeaturesofhumandialogicinteraction

metaphor
Recentresearchinneurosciencescon
thatdifferenthumanabilities
suchasrationality,emotion,perceptionareinterrelatedandcannotbeseparated
frombiological-physiologicalstructure(Damasio1994;Schumann1999).Itisa
complexintegratedwhole
,themirrorneuron,fromwhichtheevolutionoflanguage
hasstarted.Matterandenergyorfunctionareperhapsoneandthesamephe-
nomenon.Theconsequencesforhumanabilitiestobedrawnfrommirrorneurons
arefar-reaching:
First,
integration
ofdifferentabilitiesisabasiccharacteristic.
Second,themirrorneuronscharacterize
humanbeings
attheverybeginning
asdirected
invariousrespects:
socially,dialogically,purposefully,intentionally,
interactively,cognitively
Third,
humanbeingsarepartoftheworld
.Theycanperceivetheworldonly
insofarastheirabilitiesallowit.Thereisnorealityassuch,onlyrealityas
perceivedbyhumanbeingsthroughthe
lteroftheirabilities.
Integrationinvariousrespectscharacterizesourstartingpoint.Italsoentersthe
staircasemodelofthedifferentdisciplines
developedbyGell-Mann(1994:111f.).At
rstsightthelistingofdisciplinesmightseemasanattempttodistinguishdifferent
levelsbutitshouldbereadasalistingofdifferentintegrateddimensions,orasGell-
Mannputsit:
...whilethevarioussciencesdooccupydifferentlevels,theyform
partofasingleconnectedstructure.

Figure2.
Staircasemodel
Forlinguistics,wecandrawdecisiveconclusionsfromthisbasicfactofintegra-
tion.Itreveals
orthodoxlinguisticmodelsandclassicalmethodologyastheorymyth

EddaWeigand
Constitutivefeaturesofhumandialogicinteraction

Constitutivefeaturesoftheobject
humandialogicinteraction

EddaWeigand
TheyareinterrelatedintheminimalculturalunitoftheActionGamewithin
Constitutivefeaturesofhumandialogicinteraction

plexityoftheactiongame.Actionsdonotrepresent
xedpatternsappropriatefor
speci
cconditionsbutdependonindividualreasoning.Inordertounderstandthe
sampledialogue,Imustdescribethesituationbecausethetextisnotautonomous
initselfbutacomponentintheactiongame.Inmostcases,itisnotsuf
cientto
relyonlyonempiricallyregistrablemeansasanobserveraswedowhenanalysing
authentictextsfromtextcorpora.Weinsteadhavetorefertotheactinghumanbe-
ings,wehavetounderstandtheirindividualevaluationofactionconditionsandwe
havetoperceivewhatisgoingoninthesituation.Theseareexactlytheprerequisites
neededalsoforthemirrorneuron-in-function.

EddaWeigand
(2)MotherYouareplayingthepianoagain.
DaughterShallIstopit?
MotherNo,itdoesn
tmatter.I
mgoingtoworkoutside.
rstutteranceofthemotherpointstothefactthatwedonotexpressour
claimsverballybyreferencetorulesonly.
Youareplayingthepianoagain
canbeun-
derstoodasarepresentativespeechact.Butwhatdoesthismean?Doesthemother
appreciatethatthedaughterisplayingthepianoorissheangry?Intonationisnot
soclearastodecidethisquestion.Everythingislefttosupposition.Eventhedaugh-
terwhoknowshermotherwellisuncertainofwhatshemeantinthisindividual
situation.Shereferstohermother
spreferenceofworkinginsilencewithoutbeing
disturbedbythepianoandunderstandstheutteranceasareproach.Inthispartic-
ularsituationhoweverthispreferenceisnotvalid.With
ShallIstopit
?thedaughter
becomesthevictimofamisunderstandingwhichhoweverisimmediatelyclari
bythenextutteranceofhermother.
Thisexampledemonstratesthatthereareinevitablyopenpointsindialogue
whichcannotbeavoided.Noteverythingissaidexplicitly;otherwisewecouldnot
starttalkingbecausewewouldhavetore
ectontheessentialpointstobeexpressed
inordertoavoidmisunderstandings.Howcouldanativespeakernottrainedin
linguisticssucceed?Wheregeneralizedrulescometotheirlimits,
suppositions
have
toaccountforindividualityandchance.
Meaningisnotde
Orthodoxlinguisticsmeanstheorizingonthebasisof
xedcodes.Communica-
tioninthissenseisthoughtofasthetransferenceof
xedpatterns,withmeanings
alreadyde
nedinadvance,equallyvalidforbothsides.Thefollowingauthentic
examplesdemonstratethatsuchaviewisanillusion.
(3)Ifyouarehomeless,youwill
ndahomeinHongKongbecausethereall
arehomeless.(heardonGermantelevision,translatedintoEnglish)
(4)Changeistheonlyconstantinthelifeofacompany.
TheEconomist
,March25th
31st,2000,p.115)
Ifwereallyinteractedonthebasisofsigntheorywewouldhavetorejecttheseex-
amplesasnonsensebecausetheycontaincontradictions.Inperformancehowever
webehaveascomplexadaptivesystemsandaccepttheseexamples.Wenegotiate
theirmeaninginawaythat
change
canbeunderstoodas
constant
oraperson
being
homeless
cannevertheless
haveahome
.Examplesofthistypemakeveryclearthat
wehavetoabandontheviewoflanguageasasignsystem.Inlanguageuse,thereare
Constitutivefeaturesofhumandialogicinteraction
\b
nosigns,nosymbolshavingmeaningindependentofthespeaker.AsWittgenstein
toldus,wordsinordinarylanguageuse
havemeaningonlyinthestreamoflife
Dialogicinteractionthereforeisbasicallycharacterizedbythe
Principleof
MeaningUncertainty
.Theso-calledexactnaturalscienceshavealreadyintroduced
aPrincipleofUncertaintyinquantumphysics.Itisthispointofmeaninginde-
terminacy,or
structuresasLi(2003)callsit,whichwehavetoacceptin
theoryandaddressasacomplexmixoforderanddisorder,ofgeneralizedrules
andchance.
OneofthemajorissuesDeacon(1997:39
46)isconcernedwithinhisbook
thesymbolicspecies
referstothequestionofwhythereareno
simplelan-
guagesusedbyotherspecies
.Suchsimplesymboliclanguagesconsistingofa
very
limitedvocabularyandsyntax
arearti
cialsystems,each
\t
EddaWeigand
Letushavealookatashortadvertisingtextfrom
TheEconomist
demonstrateshumanbeingsaspurposefulbeingswhokeeptheirrealpurposes
Constitutivefeaturesofhumandialogicinteraction
\n
standing,usingthetextasverbalmeansintheactiongameandintegrallyusing
othermeans,cognitiveandperceptualones,toachievetheirpurposeseffectively.
Fundamentalsofatheoryofthedialogicactiongame
Letusnowaddresstheessentialquestionofhowwecancopewiththesefeatures
inatheoryofhumandialogicinteraction.Thechallengeistoaddressthecomplex,
\f
EddaWeigand
Principlesofprobability:Basicandcorollaryones
TheTheoryoftheDialogicActionGameisbasedonthreebasicprinciplesandaset
dialogic purpose (state ofaffairs)
dialogic means
action function (proposition)
reference predication
Figure3.
Dialogicaction
Assumingthatcommunicationevolvedfromgestures,wecansayitisthedialogic
meansofagesturewhichsignalsadialogicclaim.Thequestionariseswhattypes
ofdialogicpurposesorclaimswehavetodistinguishininteraction.Idealtwith
Constitutivefeaturesofhumandialogicinteraction

willrestrictmyselftobrie
yindicatetheclaims.Inmyopinion,ourspeechacts
arepredominantlybasedontwobasicclaims:aclaimtotruthandaclaimto
volition.Bothclaimscorrespondexactlytothebasicmentalstatesofbeliefand
desire.Theactionfunctionbeingbasedeitheronaclaimtotruthoraclaimto
volitionreferstothepropositionalfunction,whichconsistsofreferenceandpred-
Directive [help (KP,Sp)]
gesture
claim to volition
Figure4.
Directiveaction(example)

EddaWeigand
Function (proposition)
communicative means
action reference predication (speaking,thinking,
perceiving)
M EA N I N G P O S I T I O N S
pragmatic claims identifying characterizing
to truth and volition objects
C O G N I T IV E BA S E
mental states of cognitive and physical abilities
beliefand desire
Figure5.
Thespeechact
Figure5startsfrom
thecomplex
phenomenologicallevelofthedialogicallyori-
entatedspeechact,identi
thesimple
withregardtomeaningpositions,onthe
onehand,andindicatestheintegrationofmeans,ontheother.Meaningposi-
tionsareconsideredtobecognitive-socialpositionsdevelopedfromacognitive
base.Thementalstatesandabilitiesofthecognitivebasecanbeunderstoodasthe
brain-in-function.
AfurtherquestiontobeaddressedconcerningtheActionPrincipleturnsout
tobethemostimportant:itisthequestionofhowtoreadthe
arrow
,inFigure5
thehorizontalonewhichcorrelatesfunctionswithmeans.Inorthodoxmodels
xedcodesthearrowisreadasconventionalcorrelation.Itishoweversimply
notthecasethatutterancesarerelatedtospeechactfunctionsbya
xedgener-
alizedcodeinrelationtospeci
Constitutivefeaturesofhumandialogicinteraction

Letmeillustratethetechniqueofnegotiationbyanexamplewhichwehave
alreadydiscussedaspartofExample(2):
(6)Youareplayingthepianoagain.
YouareplayingtheGame-boyagain.
Game-boy
isatradenameforahand-heldcomputergame)
Orthodoxlinguisticsassumes
xedcodessuchasthecodeofsymbolizing.Utter-
ancesliketheExamples(6)cannothoweverbedecodedby
xedcodes.Toacertain
degree,theycanbeunderstoodbyreferencetorulesasdeclarativesentenceswhich

EddaWeigand
reaction
making a claim fulfilling the claim
Figure6.
Complementarysequelofactionandreaction
Oneofthecharacteristicsofdialogicactionisthatthe
singlespeechactcannotbe
consideredcommunicativelyautonomous
.Communicativeactionsarealwaysdia-
logicallyorientated.ActionandReactionarenotarbitrarilycombinedbutinterre-
latedbythesamedialogicclaim.Theinitiativeactionmakestheclaim,thereactive
actionful
lsitinapositiveornegativesenseorleavesthedecisionopen.Aristotle
principleoftheautonomyofthesingleactmayrelatetopracticalactsbutnotto
communicativeones.
Themirrorneuronscon
rmnotonlytheActionPrinciplebutalsotheDia-
logicPrincipleproper.Theysignaleitheractionorreactionbythesamemeansof
agesture.Theprecisemeaningandunderstandinghastobenegotiatedbycogni-
tiveandfurtherspeci
cperceptualmeans.Thus,togiveoneexample,adirective
speechactclaiminghelpaimsatareactivespeechactofconsent.Asitisprimar-
ilyimportanttodistinguishapositivereactionfromanegativeone,twosimple
differentgestureshaveevolved:noddingandshakingheads.
DIRECTIVE [help (KP,Sp)]
claim made claim fulfilled
gesture gesture
Figure7.
Directiveinteraction(example)
Fromthegeneralinteractivefunctionofnegotiationoroftryingtocometoan
understanding,speci
cspeechacttypesandsub-typesaretobederivedbyusing
functionalcriteria,forinstance,criteriawhichdifferentiatethepragmaticclaims
orintroducepropositionalcriteria(Weigand1989,1991).Dialogicinteractionas
Constitutivefeaturesofhumandialogicinteraction

consequentlyistobereadasprincipleofprobability.Itistheinitiativespeechact
itselfwhichtellsusviarationalreasoningwhichreactivespeechactisexpected.Ra-
tionality,however,representsalimitedconceptinperformance.Language-in-use,
beingpartofourdailylife,referstolifeconcepts,whichareconceptsofprobability
ageacultureattributestohumanbeings,eitherwithrespecttothefreedomof

EddaWeigand
accordingtoregularities.Whereregularitiesendweusesuppositionsandother
means.Themodelwhichsubstitutestheorthodoxviewthereforeis
anopenmodel
whichacceptsmisunderstandingsandconsiders
humanbeingsascomplexadaptive
systems
Constitutivefeaturesofhumandialogicinteraction
\b
Humanbeingsareguidedintheirbehaviourbyintentionalityandotherper-
ceptualandcognitiveabilitieswhicharenotidenticalforeveryhumanbeing.They
\t
EddaWeigand
Roth,G.(2000).GeistohneGehirn?HirnforschungunddasSelbstverst
ndnisdes
Menschen.
Forschung&Lehre,5
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Stamenov,M.I.(1997).Grammar,meaningandconsciousness:Whatsentencestructure
cantellusaboutthestructureofconsciousness.InM.I.Stamenov(Ed.),
Language
Structure,DiscourseandtheAccesstoConsciousness
(pp.277
342)[Advancesin
ConsciousnessResearch,12].Amsterdam&Philadelphia:JohnBenjamins.
Weigand,E.(1984).LassensichSprechaktegrammatischde
nieren?InG.Stickel(Ed.),
PragmatikinderGrammatik.Jahrbuch1983desInstitutsf
rdeutscheSprache
(pp.65
91)[SprachederGegenwart,60].D
sseldorf:Schwann.
Weigand,E.(1989).
SprachealsDialog.SprechakttaxonomieundkommunikativeGrammatik
[LinguistischeArbeiten,204].T
bingen:Niemeyer.
Weigand,E.(1991).Thedialogicprinciplerevisited:Speechactsandmentalstates.In
S.Stati,E.Weigand,&F.Hundsnurscher(Eds.),
DialoganalyseIII.Referateder3.
Arbeitstagung,Bologna1990
,Bd.I(pp.75
104)[Beitr
gezurDialogforschung,1].
bingen:Niemeyer.
Weigand,E.(1998a).Contrastivelexicalsemantics.InE.Weigand(Ed.),
ContrastiveLexical
Semantics
(pp.25
44)[CurrentIssuesinLinguisticTheory,171].Amsterdam&
Philadelphia:JohnBenjamins.
Weigand,E.(1998b).Emotionsindialogue.InS.
Cmejrkov
,J.Hoffmannov
,O.M
llerov
&J.Sv
Somefeaturesthatmakemirrorneurons
andhumanlanguagefacultyunique
MaximI.Stamenov
Georg-August-Universit
ttingen,Germany/
BulgarianAcademyofSciences,So
a,Bulgaria
Thepuzzleofhumanlanguagefaculty
Onecanapproachtheproblemabouttheuniquenessofhumanlanguagefaculty
fromdifferentperspectives:
1.Onecanpointoutthathumanshaveareferentialpotentialofsymbolicunits
(standardlyidenti
ableas
words
)whichoutnumbersthecapacitiesofthe
otherbiologicalspeciesbyseveralordersofmagnitude.Whileotherspecies
havecommunicativecalls(symbols)bydozens,humanshavethemindozens
ofthousands.Inpsychologicalterms,humanLong-TermMemory(LTM)
musthaveaqualitativelydifferentorganizationcomparedtoLTMsoftheother
species;
2.Onecanpointouttheuniquedevelopmentofhumansintermsoftheneuro-
physiologysupportingtheperformanceofarticulatespeechwithhighspeed
andreliability.Thecomplexityandspeedinthemotorperformancerequired
fortheimplementationofhumanspeechamountstoamostcomplexmo-
torperformanceeverynormalhumanindividualcanroutinelyexecute.Ifwe
comparelanguagecapacitywithmusicalcapacityandperformance,eachnor-
malhumanindividualaroundtheworldwithoutformaltrainingwhatsoever
spontaneouslyperformsonthelevelofamusicalvirtuosofromthepointof
viewofthecomplexityofskillsinvolvedandspeedofperformance;
3.Onecanpointoutalsotherelatedfeatofbeingcapabletodevelopaphono-
logicalsystemforhigh-speedreliablecategorizationofthesoundsofarticulate
speechinspeechperception;
4.Onecanpointouttheuniquedevelopmentofformalstructureoflanguage,
namelysyntax.Afrequentlymadeclaimisthatthecoreoflanguageasasep-
aratecapacityofhumanspecies(i.e.,asdifferentfromothercognitivecapaci-
MaximI.Stamenov
ties)isidenticaltosyntax.Syntaxisauniquefacultyofthemind,onewithout
asecond.Thisisthecase,amongothers,becausesyntacticcomputationisref-
erentially
blind
.Becomingfreefromthesnaresofreference,itmanagedto
optimizeitselfascloseaspossibletothelevelofformalperfection;
5.Onecanpointoutthatlanguagefacultydemandedforitsdevelopmentspe-
cevolutioninthecapacitytounderstandconspeci
csviatherequirement
oftakingstrategicallytheperspectiveoftheotherinterlocutor(i.e.,empathiz-
ingtoher/him)inordertobeableproperlytointerpretthecontentofher/his
message.Thiscapacityintentionallyto
transcend
sownegocentricper-
spectiveapparentlybecomespossibleonlywiththeadventoflanguage.
Thelistofthesefeatures,de
nitely,couldbefurtherdevelopedandelaborated.In
tryingtotracetheoriginoflanguage,attemptsweremadetoidentifythepointof
departureforthedevelopmentofhuman-likelanguagetakingasingleonefrom
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
i.e.,wehavenoaccessinprincipletothewayofrepresentationandcomputationof
languagestructure.ThisisessentiallythepositionofNoamChomsky(1993,1994,
1995).Itisbotharadicalandsafeinitsagnosticismposition.
Correspondingly,foralinguist,themostconsequentandtheleasttrouble-
somewayofdealingwithMirrorNeuronsSystem(MNS)anditspotentialpurport
MaximI.Stamenov
awareoftherealnatureandspeci
cityofMNS.Inotherwords,beforetryingto
constructscenariosoflanguageoriginandevolutionbasedonMNSwemusttake
caretoanalyseproperlythenatureofMNSitself.Thiscanbeachievedbestby
comparingitwiththemostadvancedstructurallyandfunctionallymentalrep-
resentationsofbehavioralactionsweareinpossessionof
thelanguage-speci
mentalrepresentations.
WhatmakesMNSpeculiarfromlinguisticpointofview
AtthebeginningofthissectionIwillrepeatonceagainthebasicfeaturesofMNS.
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
c.theMNSdoesnotsupportcognitive(language-like)mentalre-presentations
ofpropositionalformat.Correspondingly,itdoesnotinvolve
understanding
interpretation
whatsoeveroftheechoed(internallymimicked)action.It
justlookscompatibleinitsstructuretoabehavioralsituationdescribablebya
transitivesentencestructureinnaturallanguage.Butthepossibilitytodescribe
somebehaviorbysomelanguage-speci
cstructureisbynomeanssupposed
toimplythatthesituationitselforthebraincircuit
representing
itpossesses
anisomorphicstructure;
d.theMNSisalow-level
modular
(inthesenseofFodor1983)mechanism,
whiletheintroductionoflinguisticstructure/meaningdistinctionrequiresthe
emergenceandthedevelopmentofatleasttwodistinct
centralsystems
(inthe
senseofFodor1983)processingstructureandmeaningandsolvingproblems
intheWorkingMemory(WM);
e.theMNSislockeddeicticallytotheimmediatepresent,itisenactedinresponse
toanactuallyobservedhere-and-nowbehavior,i.e.itapparentlydoesnotneed
thesupportfromtheLTMand/orofageneral-purposecognitivesystemlike
WMforthesakeofperformingoff-linecognitivecomputations(cf.however
Fogassi&Gallese,thisvolume,whereanexperimentisreportedwhereadelay
ofupto1,5secdoesnotpreventamonkeyfromreactingwithitsMNS).
Myconclusionisthefollowingone:asmuchasMNSlookscompatibleinitsper-
formancetolanguage-speci
csemanticstructure(cf.Rizzolatti&Arbib1998)in
possessingapatternthatseemsisomorphictoatransitiveactioninitiatedbyan
agentwhomanipulatesanobject(aninanimatepatient),thereallychallenging
aspectsofthissystememergewhenonestudieshowitdiffersfromlanguage.
MNSisnotintersubjective(interpersonal)butinter-embodied
MaximI.Stamenov
WhyMNSlooksinterpersonallyblindfrompsychologicalandlinguisticpoint
ofview?Thisseemstobethecasebecause:
1.itistriggeredautomatically;theveryvisualavailabilityofathirdpersonma-
nipulatinganobjectwouldbeenoughtotriggertheresponse;
2.itisenactedinanunconsciousway(andcannotbepotentiallydeautomatized
andmanipulatedstrategically);
3.itisdueneithertoamappingnortoanidenti
cationoftheobserverwiththe
agentoftheaction.Thisisthecasebecauseitdoesnotrequirecommunicative
(dialogical)
rst-persontosecond-personbindingeveninitsminimal(root)
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
icalanalysisofthestructureofobservationandperformanceandhasnoevident
correlatesintheactualneuralimplementationofMNSitself.
IthinkthatMNS,asamatteroffact,implementsa
direct
resonance-basedat-
tunement
toaclassofactions(theresonance-likefunctioningofMNSissuggested
MaximI.Stamenov
8.input(tooutput)systemsexhibitcharacteristicandspeci
cbreakdownpat-
terns(inourcasethesecouldbeecholaliaandechopraxiainhumans,inthe
caseofmonkeysandprimatesthepathologyofMNSremainstobeinvesti-
gated);
9.theontogenyofinput(tooutput)systemsexhibitsacharacteristicpaceand
sequencing(whichinthecaseofMNSremainstobestudied)(cf.Fodor
1983:47
Thus,notonlytheperipheralfacultiesofperceptionbutalsosomebasictypes
ofbehavioralactionimportantforthesurvivalofthecorrespondingspeciesmay
turnouttobeinheritedhardwiredandperformasautonomousbraincircuits.The
fragmentariness(modularity)ofbrain
smindappliesonabroaderscalethanit
waspreviouslyenvisaged.Wemaynowclaimthatthemindisalsomodularinim-
plementingspeci
cactionclassesasspeci
csub-circuitsofthebodyschema.The
highspecialization(andcorrespondingfragmentariness)ofthemultiplepercep-
tualandactioncircuitscanserveasareasononitsownforcallingintoexistence
centralsystems
likelanguageandconsciousness.Theiraim,fromthispointof
view,wouldbetouniteintoasinglegestaltthefragmentarybodyimage.
TheMNSAgentcomparedtothelinguisticself
Whatweseeinthedemonstrationsofastandardexperimentinthisparadigmis
asfollows.Weseeamonkeyreaching,e.g.,forabanana.TheMNs
re.Inthenext
conditionoftheexperimentweseeamonkeyobservinganexperimenterreaching
forabananaoutsideofthescopeofherownreach.TheMNs
reasifthemon-
keyitselfisreachingtograspthefruit.Thus,weareledpracticallyautomatically
toinferthatwehaveasortofa
projection
oftheobserver-monkeytotheagent-
experimenterleadingtoanidenti
cationoftheformerwiththelatter.Thestruc-
tureofthesituation,asseenandverballydescribedbyahumanmind,isprojected
asthementalandneuralstructureimplementedinthemonkey
smindandbrain.
Inthisway,wearriveatthescenariothatpresupposestheexistenceoftwosepa-
ratementalre-presentationswithatransitivestructure(subjectplusdirectobject)
thatincludeanobserverintheperceptualsystemandofanagentinthepre-motor
systemandtheir
matching
(mapping)asanoutcomeofMNSactivation:
Themeaningoftheobservedactiondoesnotresultfromtheemotionit
evokes,butfromamatchingoftheobservedactionwiththemotoractivity
whichoccurswhenindividualperformsthesameaction.[...]Whatisim-
portanttostresshereisthattheproposedmechanismisbasedonapurely
observation/executionmatchingsystem.Theaffectivevalenceofthestimuli,
evenifpossiblypresent,doesnotplayaroleinthis
understanding
system.
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
Whatwehaveashardevidence,however,isthatitistheactionitselfthatistracked
rstbothinperceivingandinpreparingforaction.Wehavenoevidencewhatso-
everthatthey(agentandobserver)are:
rstdistinguished,
b.inorderafterwardstobeidenti
edwitheachother.
Mypoint,inotherwords,isthatthisdistinctionissimplyanartefactofthewaythe
referredtoexperimentswerere-presentedanddescribed.Withthiswayofconcep-
tualisation,theMNSautomaticallybecomesadirectforerunnerandprerequisite
ofallhigh-levelsocialskillsinmonkeys,primatesandhumans.
Whattheavailableexperimentalstudiesactuallyshowisthecon
uenceof
theperceptualandmotorpropertiesinthebrainrepresentationoftheobject(cf.
Gallese2000b;Fogassi&Gallese,thisvolume).Onemayobjectthatthisappliesfor
theobjectofactionbutnotforthesubjectofperception-cum-actioncontinuum.
However,a
lonely
motorrepresentationofhowtohandleanobjectseemstome
highlyimplausible.Thissamespeci
cationmustnecessarilycodethesubject
sca-
pacitiestomanipulatetheobject.Theverycharacteristicsoftheembodimentof
theagentarealreadynecessarilyimprintedinthemotorspeci
cation.Thisis,sim-
ply,theothersideofthesamecoinofthe
instruction
how-to-handle-an-object.
Inthisway,thelatterspeci
cationcodesthepropertiesofboth
agent
MaximI.Stamenov
iorarefunctionalcomponentsofthehumanWMandpresupposetheavailability
ofcentralsystemswhileMNS
agent
ismodularandboundtoacoupleofaction
classes.Thus,whatlooksas
thesame
agentrolefromsemanticpointofviewturns
outtobeimplementedinhumansinthreedifferentways
asaMNSagent,asan
implicitcentralexecutiveandtheexplicit(self-conscious)self.Ifthereisalink
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
animateentity(individualofthesamespecies).Theobjectisjustthethingtobe
achieved.Thethingtendstobeofasizecapableofbeingfreelymanipulated,e.g.,
byasinglehand.Ifthisisthecase,thereisnopossibilitytointerprettheobject
asthesecondinterlocutorininterpersonalinteraction.Forsomesituationtobe
MaximI.Stamenov
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
AsIpointedoutintheSection2.1.above,anotherpossibilityistorepresent
thiscircuitastheonecontaininga
MaximI.Stamenov
body,unlikethephysicalone,isnotunitedintoasinglegestalt
itisa
mosaic
(modular)oneandonethatisnot
owned
byaself,butscandalouslysharedonan
online-servicebasis.
Therepresentationoftheembodiedagent-selfinlanguage,ontheotherhand,
appearstofunctiononaratherdifferentbasis.Itisremarkabletonoticethatitis
partofthelanguage-speci
csemanticrepresentationtotalkaboutthewholebody
doingdifferentthingsintheworld,butnotaboutalienated(modular)orshared
bodypartsactingasautonomousagentsonitsownaccount.Forexample,itisquite
normaltosay(1)andanomalous(atleastinEnglishandallotherIndo-European
languagesIamawareof)toutter(2),whileitwouldbestrictlyanomalous(in
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
Itwouldbenotsuper
uousatthispointtoaddthatthewayoftalkingaboutthe
embodiedselfasauni
edanduniqueentityisspeci
ctolanguage,andnotnec-
essarilytothewayoneconsciouslycanimagineoneselfactingintheworld.For
example,ifIamaskedtoimaginethe
meaning
of(6),Iwilltendtoimagine
myown
ngersasthepro
ledmemberofmybodydoingtheaction,whileas
abaseofthepro
ledbodymemberImayseemyownarm(Iusehere
pro
base
asconceptsfromthedescriptiveapparatusofthecognitivegrammarof
Langacker1987):
(6)Iampluckingtherose.
PleasenotethatIwillnotimaginemyselfasawholehumanagentinthewayI
canactuallyseeorimagineathird-personagentdoingthissameaction.Thus,the
language-speci
crepresentationoftheselfdoesnotnecessarilycoincidewiththe
structureoftheselfcapableofbeingvisualized(imagined).Inotherwords,the
linguisticself-consciousnessdoesnotnecessarilycoincide,i.e.,itnotidenticalin
itsstructure,withthespatial-cognition-basedself-consciousness.Thelinguistically
basedself-consciousnessappearstobethemostuni
edone,itisthebestcandidate
forfunctioningas
thecenterofautobiographicalgravity
(theother
theimagin-
ableself
looksaslessuni
edmentalrepresentationsoftheselfasanagentinthe
world).Inthisway,
oneandthesame
,presumably,psychologicalfunction(self-
consciousnessasobserveroragent)isimplementedinaatleastpartiallydifferent
wayinlanguageandexplicitspatialcognition.
InthecaseofMNS,asamatteroffact,thereisnoagentwhatsoeverand,
therefore,thereisnogroundforaself-likepsychologicalstructureandintersub-
jectivityeither.Correspondingly,theidenti
cationinMNSofacognitivepattern
containinganagentoreventheanalogybothinstructureandfunctionmaybe
MaximI.Stamenov
WhatmakesMNSchallengingfromlinguisticpointofview:
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
Therearemanyquestionsthatremaintobeexploredintryingto
ndout
aplaceforMNSinthescenariosoftheoriginanddevelopmentofthelanguage
faculty,e.g.:
a.istheBroca
sareatheonlyplaygroundofaMN-likesystem?
b.ifthereareotherMNSsinotherbrainareasrepresentingtheactionpotentialof
otherbodymembers,dotheytendtocon
atedifferentclassesofaction(from
structuralandfunctionalpointofview),asisthecasewiththe
rstonetobe
discovered?
c.arethereotherMN-likesystemsthatareJanus-like,i.e.,pointingtomorethan
onecognitivemechanism(languageandspeechinourcase)?
d.canthetypesofactiontowhichtheotherMN-likesystemsseemcompatible
beofdifferentlevelofcognitiveabstraction(asisthecasewithspeechand
languageinthecaseofMNS)?
Theactionstructureasrepresentedinlanguage
Belowforthebene
tofnonlinguistreadersIwillprovideashortpresentation
whatmakesthehumanlanguagefacultyuniqueonthesyntacticandsemanticlevel
takingasexamplethecognitivepatternofanagentmanipulatinganobject
the
onethatlooksbestcomparabletoMNS.
Firstofall,itisimportanttobecomeawarethatlanguage-speci
crepresenta-
tionsofbehavioraleventshaveatleastthreedifferentsourcesofsystematicvari-
ationinthecontributingcognitivepatterns.Allofthem,unlikeMNS,arehighly
abstractintheircognitiveformat.Onlythemappingofthesethreetiersofcog-
nitivepatternformationcanleadtowhatweexperienceasasingleholisticactof
meaningfulexperiencesupportedbylanguagestructure.
Thethreetiersinquestionareimplementedbydifferentcentralsystemsin
WM(e.g.,byspatialcognition,abstractlanguageofthought,andlanguage-speci
syntax).EachofthemrequiresaccesstoLTM,aswellasaccesstothemotorroutines
involvedintheexecutionandcontrolofinnerandouterspeech.Languageisnot
supportedbyaclosedmodularsinglebraincircuit,asisthecasewithMNS,butby
MaximI.Stamenov
thefollowingfourtypesofverbmeaning(
Actionsart
)areusuallydistinguished
(accordingtoascaleofinnercomplexityasestablishedbylogicalanalysis):
a.a
state
isatypeofpredicate(thecorrelateoftheverbinlogic)thatdenotes
propertiesornon-dynamiccircumstances,forexample
besad
beafraid
(ifwe
speakabouthumanstates);
b.an
activity
isatypeofpredicatethatindicatesadynamiceventinwhichthere
isnochangeofstatesuchasin
see
,or
c.an
achievement
isatypeofpredicatethatindicatesachangeinastateor
dynamiccircumstancessuchas
break
,or
d.an
accomplishment
isatypeofpredicatethatischaracterizedbyacausative
semantics,e.g.,
causetodie
(cf.Vendler1967).
Theverb-speci
cmeaningprovidesuswiththepossibilitiestonameandconcep-
tualizedifferentbroadclassesofstaticanddynamicsituationsintheexternaland
internalworld.Thelanguagealsoprovidesuswiththemeanstodiscussverbs
representationalpotentialanddevelopclassi
cationsoftheirstructurewithlay-
erednestingofactors(arguments)fromonetofour.Theprototypicalconstruction
amongthemisthatofanagentactingintheworldofconcreteobjectsasexpressed
intransitiveverbs.
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
Theimportantpointtorealizeinanalysingthelistofsemanticrolesgivenabove
inourcontextisthattheselfcanbeprojected
strategically
inanyofthesemantic
MaximI.Stamenov
suadeyou
Youunderstandme
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
aimsattheattunementofabodymemberofamonkeyoraprimatetoactingin
theworldhere-and-nowwiththeshortestpossiblerouteinthelocalneuralcir-
cuitry.Thisattunementisnumbanddumb,unconscious,sel
essandagentless.
Whocouldhaveenvisagedonly10yearsagothatsuchasystemwaitsforitsdis-
coverersburiedsomewhereinthemonkey
s,primate
sandpotentiallyman
sbrain?
InmonkeysandprimatesMNScannotsupportunderstanding,imitationofinten-
tionalaction,languageandroleplayingintheirhumansense.Theactualstructure
ofMNSseemstoconsistminimallyoftwocomponents(howtheobjectlooksand
howitlookswhenhandled)whichareanatomicallylocalizedintwoorthreediffer-
entbrainregions(cf.Fogassi&Gallese,thisvolume).The
rstcomponentisasort
representation
offeaturesofsubjectandobjectofactionintheformofasort
ofa
\b\f
MaximI.Stamenov
encapsulationoftheserialcomponentofMNSonanevolutionaryscaleandthe
generalizationofitsapplicationtothenascentmechanismsofspeechandlanguage.
Thisscenarioisaverychallengingoneasitwouldchangethewayweenvisagethe
Somefeaturesthatmakemirrorneuronsandhumanlanguagefacultyunique
\b
Ramachandran,V.(2000).Mirrorneuronsandimitationlearningasthedrivingforcebeh-
thegreatleapforward
inhumanevolution.http://www.edge.org/documents/
archive/edge69.html
Rizzolatti,G.,&Arbib,M.A.(1998).Languagewithinourgrasp.
TrendsinNeuroscience
Rizzolatti,G.,Fadiga,L.,Gallese,V.,&Fogassi,L.(1996).Premotorcortexandthe
recognitionofmotoraction.
CognitiveBrainResearch,3
,131
Rizzolatti,G.,Fogassi,L.,&Gallese,V.(2001).Neurophysiologicalmechanismsunderlying
theunderstandingandimitationofaction.
NatureReviewsNeuroscience,2
,661
Stamenov,M.I.(1997).Grammar,meaning,andconsciousness:Whatcanthesentence
structuretellusaboutthestructureofconsciousness?InM.I.Stamenov(Ed.),
LanguageStructure,Discourse,andtheAccesstoConsciousness
(pp.277
Amsterdam&Philadelphia:JohnBenjamins.
Stamenov,M.I.(forthcoming).Languageandself-consciousness:Modesofself-
presentationinlanguagestructure.InT.Kircher&A.David(Eds.),
TheSelfin
NeuroscienceandPsychiatry
.Cambridge:CambridgeUniversityPress.
Trabant,J.,&Ward,S.(Eds.).(2001).
NewEssaysontheOriginofLanguage
.Berlin:Mouton
deGruyter.
Vendler,Z.(1967).Verbsandtimes.In
LinguisticsinPhilosophy
(pp.97
121).Ithaca,NY:
CornellUniversityPress.
Weigand,E.(2000).Thedialogicactiongame.InM.Coulthard,J.Cotterill,&F.Rock(Eds.),
DialogueAnalysisVII.Workingwithdialogue
(pp.1
18).T
bingen:Niemeyer.
Altercentricperceptionbyinfants
andadultsindialogue
Ego
svirtualparticipationinAlter
complementaryact
SteinBr
ten
UniversityofOslo,Norway
Introduction:Thequestions
Recentinfancyresearchrevealsinborncapacityforattunementtoothers
gestures
andforreciprocalface-to-faceinterplay.Humaninfantscanengageinprotocon-
versationwiththeircaregiversinthe
rstweeksoflife,andcanreciprocatecaregiv-
\b
SteinBr
ten
intheseterms:Ego
sspeakingismonitoredbypredictivesimulationofAlter
slis-
tening,andEgo
slisteningbypostdictivesimulationofAlter
sspeaking(Br
ten
1973,1974).
Alikelysupportofsuchsimulation-of-mindabilityisthepreverbalcapacity
uncoveredininfantsforwhatIterm
altercentric
perception,entailingvirtualpar-
ticipationinwhatAlterisdoingasifbeingjointlydonefromAlter
sstance(Br
ten
1996,1998a,c).Thispermitsareply,then,tothesecondquestionintermsofinfant
learningbyother-centredparticipantobservation.Trevarthen(1998:16)
ndsthat
theef
Altercentricperceptionbyinfantsandadultsindialogue
\b
Figure1.
Infantgirl(11days)onthenursingtableinmutuallyattunedinterplaywith
hermother(Br
ten1998c:29).
Dialoguepartnerssimulatingoneanother
sverbalproduction
andunderstanding
Aconversationmodel
Drawing
interalia
\b
SteinBr
ten
withMead
s(1934)theoryofanticipatoryresponsebyperspective-taking,andthe
latterwithLiberman
s(1957)originalmotortheoryofspeechperception,towhich
:the speakers virtual
participation in the listener process
by predictory simulation ~
with the utterance candidate Fas
input and simulated content
Cas output compared to the
intended content C:
ifC= C then F:= F
else modify Ffor another trial
(Left circuit*)
cosim
cosim
Speaker:
Listener:
comprehended content
cosim
formulation
uttered
Legend broken lines circuits:
(Right circuit**)
:the listener
virtual participation in the speaker
production by postdictory
simulation ~ with the comprehend
candidate Cas input and F
as output compared to the
utterance F as heard:
ifF= F then C:= C
else modify Cfor another trial
coco
intended content
Figure2.
Aconversationmodelofspeaker
sandlistener
svirtualparticipationinone
another
sactivitybytheirsimulatingthecomplementaryprocessintheother
smind.
Whileproducinganutterance,thespeakerregulatesownencodingbysimulatingthe
listener
sdecoding,andwhileprocessingtheutterance,thelistenerchecksowndecod-
ingbysimulatingthespeaker
sencoding(Br
ten1973,1974).(Inthelegendspeci
ca-
tionofsuchmentalsimulationloops,thelogicalbecome-operator
meansthatthe
valueofthevariabletotherightoftheoperatorisassignedtothevariabletotheleft).
Altercentricperceptionbyinfantsandadultsindialogue
\b\b
Thereby,areplyisimpliedtothe
\b\t
SteinBr
ten
Figure3.
Reciprocatinghermother
sspoon-feeding,aninfant(111/2months)
demonstrateslearningbyaltercentricperception:hervirtualparticipationinthecare-
givers
previousspoon-feedingactshasleftherwithaproceduralmemoryofhaving
beenavirtualco-authoroftheirenactments(asiftheyhadbeenhand-guidingher)
andwhichguidesherreciprocalre-enactment(fromBr
ten1998c:15;cf.alsoFigure4
(bottom)).
Altercentricperceptionbyinfantsandadultsindialogue
\b\n
\t\f
SteinBr
ten
of-motion
andthefolksense
beingmovedby
).Whenvirtuallymovingwiththe
facingotherasiffacingthesamedirectionandbeinghand-guided,altercentric
co-ordinatesareentailedbyone
smovingfromthebodilycenteroftheother,not
sown,andwhichthenhavetobereversedtoallowforexecutedre-enactment
inabody-centredframe.
Theabovegivesthede
ningcharacteristicsoflearningbyaltercentricpartici-
pation,permittingthisexplanatoryreplytothesecondquestion(Q2):infantslearn
tore-enactfromaltercentricperceptionoftheircaregivers
enactmentasifthey
hadbeenhand-guidedfromthecaregivers
stance.Whenthisaltercentriccapac-
ityisbiologicallyimpaired,themirrorreversalrequiredbyface-to-facesituations
willpresentlearningproblems,forexample,tosubjectswithautism.Theirpre-
dictedandrevealeddif
cultiesingesturalimitationinface-to-facesituationshave
beenexplainedbytheirattributedinabilitytotranscendownbody-centredview-
point(Br
ten1993,1994).
Whenaskedto
DoasIdo
,theyhavedif
cultiesin
revertingandmatchingthemodel
sgesture,suchasgraspthumb,orpeek-a-boo,or
evensimplyraisingarms(cf.Br
ten1998a:115
118;Ohta1987;Whiten&Brown
1998:267
Feeder
smouthmovementsrevealtheiraltercentricperception
oftherecipient
sact
Altercentricperceptionbyinfantsandadultsindialogue
\t
Figure4.
Exposuretomanualactsinviting,respectively,unilateralandreciprocal
matchingresponses.
(Top)
Whenthemacaquemonkeyobservesthegraspingofapieceoffoodand
whengraspingthefoodbyitself,thereisagrasp-speci
cpremotorneurondischarge
(drawingadaptedfromDiPellegrinoetal.1992).
(Middle)
Atoddler(18months)watchestheexperimenter
sfailedefforttopullthe
dumbbellapartandthen,whenhandedtheobject,pullsitapart(Meltzoff1995;cf.also
accountsinBr
ten(Ed.)1998:47
62,112
(Bottom)
Infant(113/4months),reciprocatinghissister
sspoon-feedingisdemon-
stratingbyhismouthmovementshisaltercentricperception:heisopeninghisown
mouthasheputsthespoonintohermouth,revealinghisvirtualparticipationinher
preparing-to-eatact(drawingaftervideorecordbyBr
ten1996:2).
inanAmazonatribeEibl-Eibesfeldt(1979:15)picturesaYanomamibaby(11to12
months)who,whileheldbythebigsister,extendsamorseltothesister
smouth,
openingownmouthintheprocessandtighteningownlipsasthesister
smouth
\t
SteinBr
ten
closesonthemorsel.Whenfeedingababyorasickperson,adultcaregiversin
Westernculturesexhibitthesametendencytobeopeningtheirmouthastheyput
thespoonintherecipient
smouth,therebyunwittinglymanifestingthattheirown
executedspoon-feedingactisaccompaniedbyvirtualparticipationintherecipi-
ent
scomplementaryactofreceivingthefoodbythemouth.Ofcourse,motiva-
tionmakesadifferencehere;nosuchovertmanifestationcanbeexpectedofthe
unmotivated,disengagedobserver.
Parallelstoconversationalef
ciency
Whenmouthmovementsofthefeeder
infantoradult
ectthecorrespond-
ingmouthmovementsoftheonebeingfedwemayseeaparallelherealsoto
thevirtualparticipationexhibitedbypartnersinverbalconversation(Figure2),
co-enactingoneanother
Altercentricperceptionbyinfantsandadultsindialogue
\t
InacontrolofthedumbbellexperimentportrayedinFigure4(middle)theex-
perimenterremainspassivewhileamachinewiththedumbbellinitsclaws
topullitapartwithoutsuccess.Thatdidvirtuallynotelicitanyre-enactmentby
thewatchingtoddlers.AccordingtoMeltzoffandMoore(1998:52)theywerede-
niedtheopportunitytoreadanyintentionand,asIseeit,theinanimateobject
\t
SteinBr
ten
dicatewhysuchanadaptationmayhavebeencriticaltohumanevolution,taking
ustothequestion(Q3).
Inwhichevolutionaryconditionswouldaltercentricperceptionhave
affordedthemostcriticalselectiveadvantage?
Based
interalia
oncomparativestudiesofinfant-adultrelationsinhumansand
chimpanzees,myspeculativereplyisthis:Compensatingforthelostprotective
andnurturingadvantageofoffspringsclingingtotheirmother
sbodywhichmay
beattributedtoMioceneapes,suchanadaptedmirrorsystemwouldhavemadea
criticaldifferencetoPliocenehominids,andtoearlyHomoerectusbeforethein-
ventionofbaby-carriers(Figure5(right))whichwouldhaverestoredsomeofthe
lostadvantagecontinuedtobeenjoyedbymodernapes(Figure5(left)).
Deprivedofthisprotectiveandlearningbody-clingingadvantage,withchil-
drenhavingtobeleftholdof(inatreeorontheground)whenbipedalparents
neededtousebotharms,andgivenincreasedinfanthelplessnessandprolonged
childhood,thoseHominidae/Homospeciesparentsandchildrenwouldhavehad
acriticalselectiveadvantage,Isubmit,whoseresonantmirrorsystemhadbeen
adaptedtosubservealtercentricperceptionandvirtualparticipationinothers
ac-
Altercentricperceptionbyinfantsandadultsindialogue
\t
Figure5.
Clingingtothemother
sbody,offspringsofgreatapes,likethisinfant(51/2
months)ofachimpanzeemotherintheKristiansandZooandWildlifePark(whoalso
\t
SteinBr
ten
totherelativelyhighface-to-facefrequencyinhumaninfant-adultrelationsand,
hence,affectsthenatureofculturallearning.
Whileinfantsofgreatapesenjoytheadvantageofclingingtotheirparentsfor
safetyandinsilencebyanaturalmodeofbodilyattachment,hominidinfantshave
hadtoresorttoasortofearly
mental
clinging
andtolearnfromengagementsin
face-to-facepedagogy.Thiswouldentailanevolvedadaptivementalarchitecture
forinterpersonalconnectivitythatbecomesoperativeeveninface-to-facerelations
atsomedistance,andwhichwouldhavebeenmostcriticalbeforetheinventionof
baby-carryingbagsattributedbyRichardLeakey(1995:97)toHomoerectus.If
andwhentheymigratedfromAfrica,theycouldnothavedonesowithoutsuch
aninvention.
InsomecontemporaryAfricancultures,unlikemanyWesterncultures,such
baby-bagsarestillinuse,forexample,intheGusiiculture,whereoftenasibling
oftheinfantisassignedthetaskofcarryingtheinfantontheback(LeVine&
LeVine1988),andwhereface-to-faceinterplaywithinfantsbylookingortalking
isrelativelyrare:
Virtuallyallof[theGusiimothers
]interactionwiththebabiesincludeshold-
ingorcarrying,andtheyoftenrespondtoinfantvocalorvisualsignalswith
physicalcontactratherthanreciprocaltalkingorlooking.Bycontrast,the
Bostonmothers,whoholdtheirbabiesinlessthanathirdoftheirinterac-
tionsfrom6monthsonward,seektoengagetheirinfantsinvisualandverbal
communicationatadistance.
Altercentricperceptionbyinfantsandadultsindialogue
\t\b
Symboliccommunication
(tertiary
intersubjectivity)byconversational
speechandself-reflectivenarratives~
fromabout24months~andsecond-
orderunderstandingofothers(theory
or simulation ofmind) from 3
6 years
Object-orientedinterpersonalcommunion
(secondary
intersubjectivity) about jointly attended objects and states ~ from 9
monthsorearlier~imitativelearningbyaltercentricparticipation
Figure6.
(Abovetheline)
Ontogeneticlayersorstepsofintersubjectiveattunement
inearlychildhoodwitheachlower-orderlayercontinuingthroughoutlifetosupport
higher-orderlayers(cf.Br
ten&Trevarthen1994;Br
ten(Ed.)1998;Stern1985).
(Below)
\t\t
SteinBr
ten
articulatingothers,anemergingambientspeechlanguage,perhapsaccompanied
byaspeech-mediatedpedagogy,wouldhaveaffordedopportunitiesfortheirbe-
ginningtoattunethemselvesbyaltercentricspeechperceptiontotheprosodyand
rhythmoftheemerginglanguagecultureintowhichtheywereborn.
Stepsofintersubjectiveattunementtowardsspeechinearlychildhood
Inearlyontogeny,wehavedistinguishedprecursoryandsupportivestepstowards
speechanddialogueintermsofthesedifferentlayersordomainsofintersubjec-
tiveattunementandunderstanding(cf.Br
ten(Ed.)1998),succinctlyindicatedin
Figure6(abovetheline):
Primaryintersubjectiveattunement
inadyadicreciprocalformatofprotocon-
versation(Trevarthen1974,1992,1998),preparingforandsupportinghigher-
orderabilitieslaterinlife.Forexample,Kuguimutzakis(1998)havevideo-
recordsof45-minutes-oldsattemptingtoimitatehisuttering/a/,andfrom
herstudiesofspeechperceptionKuhl(1998)
ndsthatby6monthsinfants
have
pruned
soundsfromtheirperceptualspacethatmakenosenseinthe
ambientlanguage.
Secondaryintersubjectiveattunement
wheninfantsjoinothersinsharedatten-
tionaboutobjects(Trevarthen&Hubley1978),learningbyimitationtoma-
nipulateobjects(Meltzoff1988),andtoreciprocatecaregivers
acts(Br
ten
1996).Suchobject-orientedculturallearninginatriangularformatopensfor
semanticlearning(Akhtar&Tomasello1998;Hobson1998).
Tertiaryintersubjectiveunderstanding
inconversationalandnarrativespeech,
entailingpredication(Akhtar&Tomasello1998)andasenseofverbalor
narrativeselfandother(Stern1985).Understandingofothers
mindsand
emotion(Dunn1998)opensforperspective-takingandemotionalabsorp-
tion,evenin
ctionalothers(Harris1998),andforsimulationofconversation
partners
minds(cf.Figure2).
Thus,thepreverbalcapacitiesandopportunitiesforculturallearninginearlyin-
fancy,includingthecapacityforaltercentric(speech)perceptionandperspective-
taking,nurtureandsupportthehigher-orderlevelabilitiesforconversationaland
narrativespeech.Forexample,inanOxfordstudy,RallandHarris(2000)
thatwhen3-and4-year-oldsareaskedtoretellfairytales,sayaboutCinderella,
theymanagebestwhentheverbsinthestorieslistenedtoareconsistentwiththe
stanceoftheprotagonistwithwhomtheyidentify,invitingtheiraltercentricpar-
ticipationin
Cinderella
sslippers
,asitwere.Thechildrenhavetroublewhenthe
Altercentricperceptionbyinfantsandadultsindialogue
\t\n
verbsinthestoriestoldareusedfromthereverseperspective,atoddswiththeir
perspective-taking.
\n\f
SteinBr
ten
communityevolvingasaco-creatednoveltyandasenvironmentallygiveninon-
togenytothosebornintothespeechcommunity,nurturingtheirimpressivespeech
learningcapacity.Thiswouldnothavebeenasef
cient,wemaynowclaim,were
itnotsupportedbyaninnate,preverbalvirtual-othermechanismenablingal-
Altercentricperceptionbyinfantsandadultsindialogue
\n
spooninownhand
tofeedthecaregiverinasemblantmanner(accompaniedbythebaby
virtualparticipationinthecaregiver
sfood-intake).
Ifthisistenable,andthereismatchingmirrorsystemsupport,weshouldexpect,for
example,thatinhumans
give
-mirrorneuronsbeactivatedduringowngivingandwhile
watchingtheothergive,andthat
grasp
-mirrorneuronsbeactivatedduringowngrasping
andwhilewatchingtheothergrasp.(PresentedbytheauthorataseminarwiththeInstitute
ofHumanPhysiologyfaculty,UniversityofParma,4May2000).
Thispredictionwasstimulated
interalia
bythediscoveryofallocentric
place
neurons
inanimals(cf.O
Keefe1992)andbymyuncoveringaltercentricperceptionininfants.
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Visualattentionandself-groomingbehaviors
amongfour-month-oldinfants
Indirectevidencepointingtoadevelopmentalrole
formirrorneurons
SamuelW.Anderson,MarinaKoulomzin,BeatriceBeebe,
andJosephJaffe
NewYorkStatePsychiatricInstituteandDepartmentofPsychiatry,
ColumbiaUniversity,NewYork,USA
Introduction
Oralandmanualself-groominggesturesobservedinfour-month-oldinfantsdur-
ingthecourseofmother-infantface-to-faceplayarefoundtocoincidesigni
cantly
withprolongationofvisualattentiontomother
sface.Ithaslongbeenbelieved
thatitisthevisualpresentationofanimatedfacialdisplaysalonethatattractsand
maintainswhatisregardedasanessentiallypassiveinterestonthepartofthein-
fant(Tronick1989).Butalthoughthebehavioralrepertoireoffour-month-oldsis
\n
SamuelW.Anderson,MarinaKoulomzin,BeatriceBeebe,andJosephJaffe
activatecorrespondingbehaviorintheinfant.Whiletheexactfunctionalroleof
mirrorneuronsinareaF5isnotyetclear,weproposethatconcentrationofvisual
attentiontothespatialregionofthemother
sfaceisinitiatedandmaintainedby
premotorintermodallydrivenneuronensemblesthatbothrecognizethematernal
facialdisplaysandgroomingactivityandenabletheinfanttoenactanapproxi-
Visualattentionandself-groomingbehaviorsamongfour-month-oldinfants
\n\b
Codingofinfantbehavior
GazeDirectionandHeadOrientationcodeswereconceptualizedintwodimen-
sionalrotationalspace,categorizedaccordingtothreecategoriesofangularro-
tationineachdimension,expressedasdeparturesfromthefrontalalignmentof
motherandinfantestablishedbytheirseatingarrangement(seeFigure1).Vertical
codesare
Level
(zerodegrees),
(+10degrees)and
Down
10degrees);horizon-
60
30
0
30
60
90
10
0
Figure1.
Codedrotationalheadpositions.
VerticalRange:plusorminus10deg.
HorizontalRange:plusorminus90deg.
(Solidellipseidenti
esrangeofheadpositionsconsistentwithsustainedfrontal
attention).
\n\t
SamuelW.Anderson,MarinaKoulomzin,BeatriceBeebe,andJosephJaffe
talcodesare
Enface
(zeroto30degrees),
MinorAvert
Visualattentionandself-groomingbehaviorsamongfour-month-oldinfants
\n\n
notedthateachofthepartitionspreservesarealtimepatternofpairwisese-
quentialeventswithinitjustasdotheone-stepconstraintsthatcharacterize
thetransitionmatrixasawhole.
TotestforListing
sLaw,correspondingrowsofthestablesubmatrices
arecomparedbytheNeyman-Pearsonlikelihoodratiotest,predictingthat
oncethesystemachievesattentionalcompliance,continuedgazingdirectly
atmotherwillconstrainheadmovementtonomorethanminorvariations
aroundthe
Enface
position
aconstraintexpectedtobeabsentwhenthe
sattentionisdirectedelsewhere;
3.Thematrixanalysisdescribedin1and2wasperformedseparatelyforeye/head
coordinationdatacollectedduringactiveself-grooming(showninTable1)
andduringperiodswhennosuchoralormanualactivityoccurred,repeating
theNeyman-Pearsontest.Summariesofthesedataareshownintheformof
simpli
ed4-statefrequencytables,inwhichnonzerocellfrequenciespredicted
byListingareidenti
ed,asseeninTables2and3.
Table1.
Normalized12-statetransitionprobabilitymodel
GAZEON
GAZEOFF
EFLEFUEFDAVLAVUAVDEFLEFUEFDAVLAVUAVD
EFL.940.000.003.000.000.000.041.000.003.013.000.000
EFU.000.000.000.000.000.000.000.000.000.000.000.000
EFD.000.000.778.000.000.000.000.000.222.000.000.000
AVL.000.000.000.000.000.000.000.000.000.000.000.000
AVU.000.000.000.000.000.000.000.000.000.000.000.000
AVD.000.000.000.000.000.000.000.000.000.000.000.000
EFL.122.000.000.000.000.000.816.000.041.020.000.000
EFU.000.000.000.000.000.000.000.000.000.000.000.000
EFD.044.000.011.000.000.000.033.000.912.000.000.000
AVL.133.000.000.000.000.000.100.000.067.700.000.000
AVU.000.000.000.000.000.000.000.000.000.000.000.000
AVD.000.000.000.000.000.000.000.000.000.000.000.000
HeadCodeAbbreviations
EFL=EnFace-LevelEFU=EnFace-UpEFD=EnFace-Down
AVL=Avert-LevelAVU=Avert-UpAVD=Avert-Down
SamuelW.Anderson,MarinaKoulomzin,BeatriceBeebe,andJosephJaffe
Table2.
Cumulativetransitionfrequencymatrix,selfstimulation*
ON
OFF
299
0
12
H Off/On
8
H On/On
13
0
H On/On
80
H Off/On
6
H On/Off
1
H Off/Off
7
H On/Off
0
H Off/Off
1
5
H Off/Off
2
H On/Off
6
106
Listing Ratio:425/546 = 0.778
* Underlined cells predicted to accumulate all entries according to Listing
s Law.
Results
AcomparisonofRows1and7inthe1stand4thquadrantsofTable1yieldsa
signi
cantNeyman-Pearsonchisquare:=13.823(p.025).Noresultsweresig-
cantapartfromthesixcellcomparisonsonjusttheserows(DF=5),indicating
thatgaze-dependentheadstabilityoccursonlyduringhead
Level
andinthe
Enface
orientation,aspredictedbythemodel.Thisresultshowsuponlyduringactiveoral
ormanualself-stimulation,however.Thecorrespondingchisquarecomputedfor
eye-headcoordinationintheabsenceoforalormanualactivityis9.918,whichdoes
notreachsigni
canceatthe5%level.Tables2and3summarizethesecontrasting
effectsineyeandheadbehaviorsunderthetwoconditions.
TheListingRatio
therelativefrequencyofpredictedcellentries
is0.778
duringselfgroomingactivity,whileduringtheabsenceoforalandmanualactivity
itis0.640.
Visualattentionandself-groomingbehaviorsamongfour-month-oldinfants
Table3.
Cumulativetransitionfrequencymatrix,noselfstimulation*
ON
OFF
72
0
12
H Off/On
3
H On/On
12
0
H On/On
65
H Off/On
7
H On/Off
0
H Off/Off
0
H On/Off
0
H Off/Off
0
4
H Off/Off
0
H On/Off
7
71
Listing Ratio:162/253 = 0.640
* Underlined cells predicted to accumulate all entries according to Listing
s Law.
Discussion
Ladavasetal.(1998)haveproposedthategocentricvisualspaceisfunctionally
dividedintodifferentregionsfordifferentbehavioralactivitiesinbothmonkeys
andhumans.ThereisevidencethatareaF5containscircuitsforrepresentingvisual
spaceneartheface,whichshehastermed
peripersonalspace.
Cross-modalstudiesinpatientswithunilaterallesionsindicatethatsomeof
theF5neuronshavebimodalreceptive
eldsforbothvisualandtactileinputssuch
thatstimulationineithermodalitycanfacilitateorinhibitperceptualextinction,
butonlyprovidedthestimuliaredeliveredneartheface.Thegeometricbound-
SamuelW.Anderson,MarinaKoulomzin,BeatriceBeebe,andJosephJaffe
Butitshouldbenotedthatadistanceof60cmpermitsthemothertoreach
outandtouchtheinfant,butnottheconverse,afactthatmayexplainwhyour
groomingeffectdidnotholdfor
otherdirected
manualcontact,whichrequires
themothertoleanforwardormakeothermovesnotundertheinfant
scontrol.
Suchmovesmakeherrelativelyunavailabletotheinfant,whereastheinfant
sown
peripersonalspaceisalwaysavailable.
Anotherpossibilityisthatthisunidirectionaleffect,whichwehavefoundonly
amongA-infants(Koulomzin1993),mightbeduetoatendencyamong
avoidant
A-infantmothershabituallytospendlesstimeinclosetactileproximity,i.e.,more
timeoutsideofperipersonalspace,thandothemothersof
securelyattached
infants.
Conclusion
Wehavefoundasigni
canttendencyforinfantstomaintainfocalattentiononthe
mother
sactivefacewhileperformingoralandmanualtactilegesturesduringface-
to-faceplay.Thisapparentindicationofdividedattention,fromtheviewpointof
Visualattentionandself-groomingbehaviorsamongfour-month-oldinfants
60degreesfromthelineofsight,preservingastableenfaceheadorientationfor2
secondsormore,consistentwithListing
sLawwhichpredictsthatsteadyattention
willproducejustsuchatendencyforheadorientationtolineupwithgazedirec-
tion.Amongtheseinfantsitwasfoundthatattentivehead/gazecoordinationwas
showntobecontingentuponself-touch/mouthingbehavior.
Itisknownthatmothersandtheirveryyounginfantsspendmuchtimegazing
intoeachother
sfaces,duringwhichtimeoral-to-skincontactandmanualgroom-
ingoftheinfantoftenoccur.Episodesofmutualgazeareespeciallyprominent
uptotheageof4months,beforewhichinfantsexhibitanobligatory,automatic
SamuelW.Anderson,MarinaKoulomzin,BeatriceBeebe,andJosephJaffe
Koulomzin,M.(1993).Attention,affect,self-comfortandsubsequentattachmentinfour-
month-oldinfants.DoctoralDissertation,YeshivaUniversity,NewYorkCity.
Koulomzin,M.,Beebe,B.,Anderson,S.,Jaffe,J.,Feldstein,S.Crown,C.(2002).Infantgaze,
head,faceandself-touchatfourmonthsdifferentiatessecurevs.avoidantattachment
atoneyear.
AttachmentandHumanDevelopment
(inpress).
Ladavas,E.,Zeloni,G.,&Farne,A.(1998).Visualperipersonalspacecenteredonthefacein
humans.
Brain
(12),2317
Meltzoff,A.(1999).Originsoftheoryofmind,cognitionandcommunication.
Journalof
CommunicationDisorders
(4),251
Posner,M.I.,&Rothbart,M.K.(1998).Attention,selfregulationandconsciousness.
PhilosophicalTransactionsoftheRoyalSocietyofLondon
SeriesB:
BiologicalSciences
(1377),1915
Rizzolatti,G.,Fadiga,L.,Fogassi,L.,&Fadiga,L.(1999).Resonancebehaviorsandmirror
neurons.
ArchivesItaliennesdeBiologie
3),85
Rizzolatti,G.,Fogassi,L.,&Gallese,V.(1997).Parietalcortex:Fromsighttoaction.
Current
OpinioninNeurobiology,7
(4),562
Tronick,E.(1989).Emotionsandemotionalcommunicationininfants.
American
Psychologist
(2),112
Tronick,E.,&Weinberg,K.(1990).
InfantRegulatoryScoringSystem(IRSS)
.Boston,MA
USA:TheChildDevelopmentUnit,Children
sHospital.
Tweed,D.,Haslwanter,T.,&Fetter,M.(1998).Optimizinggazecontrolinthreedimensions.
Science
(5381),1363
Theroleofmirrorneuronsinthe
ontogenyofspeech
MarilynMayVihman
UniversityofWales,U.K.
Introduction
Ithasbeensuggestedthatchildren
sownvocalpatternsplayakeyroleinthedevel-
opmentofsegmentalrepresentationsofadultwords(Vihman1991,1993;Vihman
&DePaolis2000).Thediscoveryofmirrorneuronsprovidesaneurophysiologi-
calmechanismforsuchan
articulatory
lter
.Assumingthatonly
withinreper-
toire
behaviorscanelicitmirrorresponses,childproductionofadult-likesyllables
wouldbeaprerequisiteforthiskindofmatchingor
ltering.Thispaperwillout-
linethedevelopmentalshiftinperceptionfromprosodictosegmentalprocessing
overthe
rstyearoflifeandrelatethatshifttothe
rstmajormaturationalland-
markinvocalproduction,theemergenceofcanonicalsyllables.Wespeculatethat
itistheactivationoftherelevantmirrorneurons,consequentuponthatmatura-
tionalchange,thatmakespossibletheuniquelyhumanshifttosegmentallybased
responsestospeechandthento
rstwordproduction.
Advancesinspeechperception:Fromprosodictosegmentalpatterns
Overthepastdecadeorsoexperimentalworkininfantspeechperceptionhasin-
creasinglyturnedfromtheearlyfocusoninfantcapacityfordiscriminationbe-
MarilynMayVihman
Table1.
Advancesinperceptionandrepresentationofthenativelanguageinthe
rstyear
Childattendsmoreto...
ProsodicpatternsSegmentalpatterns
Atbirth...
nativelanguage
(vs.prosodically
Theroleofmirrorneuronsintheontogenyofspeech
Table1.
continued
Childattendsmoreto...
Prosodicpatterns
Segmentalpatterns
By11mos
wordformsfamiliarfrom
everydayexperience
(Hall
&Boysson-Bardies1994)
uninterruptedwords
[but
notwhenlow-pass
ltered]
(Myersetal.1996)
familiarwords
[evenwith
reversal
oftheaccen-
tualpattern
with
changetotheonsetCofthe
accentedsyllable](Vihman
etal.2000)
underlieafamiliarityresponsetospeech.Exceptionalevidenceofveryearly(holis-
tic)responsetosegmentalpatternsinvolvesstimulithatcanbeassumedtobeim-
buedwithstrongaffector
personalrelevance
forinfants(e.g.,theinfant
sown
name:Mandel,Jusczyk,&Pisoni1995,andfamilytermsfor
Tincoff&Jusczyk1999;forelaborationofthenotionof
personalrelevance
,see
VanLancker1991).
Infants
apparentlygreaterearlymemoryforprosodicpatternsfollowsnatu-
MarilynMayVihman
frombroadly
universal
tonative-languageonly:Werker&Tees1984;Best1994).
Firstperceptualrepresentationsofspeechforms
Despiteintensiveexperimentalworkoninfantresponsestospeechforovertwenty
years,Hall
andBoysson-Bardies(1994)wasthe
rststudytoexamineinfantre-
sponsesto
untrained
speechforms.Theseauthorstested11-month-oldFrenchin-
fantsonwordpatternsexpectedtobefamiliarfromeverydayexposure.Thewords
werechosenfromthoseproducedearlyinthesecondyearbyFrenchinfantsinan
Theroleofmirrorneuronsintheontogenyofspeech
familiarwordsversusthesamefamiliarwordsunderalteredstress(e.g.,
).Nosigni
cantdifferencewasfound,suggestingthatthestresspatterndid
notconstituteanessentialpartoftheinfants
lexicalrepresentations.Wevalidated
this
ndinginanadditionalexperimentinwhichwecontrastedthelistofaltered
familiarwordsusedinthepreviousexperimentwithalistofaltered
unfamiliar
words;undertheseconditionstheformerreceivedsigni
cantlylongerlooks(p
.001),againdemonstratingthattheinfantscould
listenthrough
thestresspattern
torecognizefamiliarwords.
MarilynMayVihman
ing.Unfortunately,however,infantspeechperceptionstudiestodatehaveincluded
neithercanonicalbabblingstatusforinfantparticipantsnorindividualpercep-
tualresponses,sowelacksomuchasacorrelationalstudyshowingtheonsetof
CVproductioninrelationtochangesinattentiontosegmentalspeechpatterns
inperception.
Theemergenceofeasilyrecognizedbabbledsyllableswithadult-liketiming
inthemiddleofthe
rstyearappearstobematurationallybasedand
tsinto
abroaderframeworkofrhythmicmotoricadvancesthatoccuraroundthatage
Theroleofmirrorneuronsintheontogenyofspeech
tatedthevowels(Kuhl&Meltzoff1988).KuhlandMeltzoffproposedthat
fantsmakeanintramodalauditory-auditorymatch;andsecond,theydevelopa
MarilynMayVihman
Table2.
10Months
/be
bi/[p
daddy
di/[d
[ha:i]
mommy
14Months
TemplateschemaCVCۈV;埂i00;i
/be
bi/[bebi]
bottle
/ba
[ba
daddy
di/
[t
hiya
ha:ji]lady
iji]mommy
Alice
rstspontaneouswords,recordedat9
10months,arelistedhereinfull.
Theroleofmirrorneuronsintheontogenyofspeech
Fernald,A.(1989).Intonationandcommunicativeintentinmothers
speechtoinfants.
ChildDevelopment
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Fernald,A.(1992).Humanmaternalvocalizationstoinfantsasbiologicallyrelevantsignals.
InJ.H.Barkow,L.Cosmides,&J.Tooby(Eds.),
TheAdaptedMind
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Oxford:OxfordUniversityPress.
Friederici,A.D.,&Wessels,J.M.I.(1993).Phonotacticknowledgeofwordboundariesand
itsuseininfantspeechperception.
Perception&Psychophysics
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Hall
,P.,&deBoysson-Bardies,B.(1994).Emergenceofanearlylexicon.
InfantBehavior
andDevelopment
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Hall
,P.,&deBoysson-Bardies,B.(1996).Theformatofrepresentationofrecognizedwords
ininfants
earlyreceptivelexicon.
InfantBehaviorandDevelopment
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Hepper,P.G.,Scott,D.,&Shahidullah,S.(1993).Newbornandfetalresponsetomaternal
voice.
JournalofReproductiveandInfantPsychology
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Jusczyk,P.W.,&Aslin,R.N.(1995).Infants
detectionofthesoundpatternsofwordsin
uentspeech.
CognitivePsychology
Jusczyk,P.W.,Cutler,A.,&Redanz,N.J.(1993).Infants
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stresspatternsofEnglishwords.
ChildDevelopment
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Jusczyk,P.W.,&KemlerNelson,D.G.(1996).Syntacticunits,prosody,andpsychological
realityduringinfancy.InJ.L.Morgan,&K.D.Demuth(Eds.),
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Jusczyk,P.W.,KemlerNelson,D.G.,Hirsh-Pasek,K.,Kennedy,L.J.,Woodward,A.,&
Piwoz,J.(1992).Perceptionofacousticcorrelatesofmajorphrasalunitsbyyoung
CognitivePsychology
,252
MarilynMayVihman
Oller,D.K.,&Eilers,R.E.(1988).Theroleofauditionininfantbabbling.
ChildDeve
lopment,59
,441
Querleu,D.,Renard,X.,Versyp,F.,Paris-Delrue,L.,&Cr
pin,G.(1988).Fetalhearing.
EuropeanJournalofObstetricsandReproductiveBiology
,191
Schwartz,R.G.(1988).Phonologicalfactorsinearlylexicalacquisition.InM.D.Smith,&
J.L.Locke(Eds.),
TheEmergentLexicon
(pp.185
222).NewYork:AcademicPress.
Stark,R.E.(1980).Stagesofspeechdevelopmentinthe
rstyearoflife.InG.Yenikomshian,
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Stemberger,J.P.,&Bernhardt,B.H.(1999).Theemergenceoffaithfulness.InB.Mac-
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ErlbaumAssociates.
Thelen,E.(1981).Rhythmicalbehaviorininfancy.
DevelopmentalPsychology,17
,237
Tincoff,R.,&Jusczyk,P.W.(1999).Somebeginningsofwordcomprehensionin6-month-
olds.
PsychologicalScience,10
,172
VanLancker,D.(1991).Personalrelevanceandthehumanrighthemisphere.
Brainand
Cognition,17
,64
Vihman,M.M.(1991).Ontogenyofphoneticgestures.InI.G.Mattingly&M.Studdert-
Kennedy(Eds.),
ModularityandtheMotorTheoryofSpeechPerception
(pp.69
Hillsdale,NJ:LawrenceErlbaumAssociates.
Vihman,M.M.(1993).Variablepathstoearlywordproduction.
JournalofPhonetics,21
Vihman,M.M.,&DePaolis,R.A.(2000).Theroleofmimesisininfantlanguage
development.InC.Knight,J.Hurford,&M.StuddertKennedy(Eds.),
TheEvolutionary
EmergenceofLanguage
.Cambridge:CambridgeUniversityPress.
Vihman,M.M.,Macken,M.A.,Miller,R.,Simmons,H.,&Miller,J.(1985).Frombabbling
tospeech.
Language,61
,395
Vihman,M.M.,Nakai,S.,&DePaolis,R.A.(2000).Theroleofaccentualpatternin
earlylexicalrepresentation.Posterpresentedat12thInternationalconferenceonInfant
Studies,Brighton,UK.July.
Vihman,M.M.,&Velleman,S.L.(2000).Theconstructionofa
rstphonology.
Phonetica,
,255
Werker,J.F.,&Tees,R.C.(1984).Cross-languagespeechperception.
InfantBehaviorand
Development,7
,49
Wood,N.,&Cowan,N.(1995).Thecocktailpartyphenomenonrevisited.
Journalof
ExperimentalPsychology:Learning,MemoryandCognition
,255
Mirrorneurons
registration
ofbiologicalmotion
Aresourceforevolutionofcommunication
andcognitive/linguisticmeaning
LoraineMcCune
RutgersUniversity,NewBrunswick,NJ,USA
Mirrorneurons
registrationofbiologicalmotion:Aresourcefor
evolutionofcommunicationandcognitive/linguisticmeaning
Languageisasymbolicorrepresentationalprocess.Meaningsinalanguagemap
theworldmentallyandaresharedamongalinguisticcommunitythroughsomeex-
LoraineMcCune
(imaginal,representational)states,adistinctiondescribedbySartre(1948)and
Mirrorneurons
registrationofbiologicalmotion
areashowsgreaterdendriticbranchingthantheareafororalmotorcontrolforthe
mutualin
LoraineMcCune
win1872/1965).Laryngealrepresentationinthemirrorsystemcouldprovidea
mechanismforsuchrecognition.
Mirrorneurons
registrationofbiologicalmotion
Relationalwords
\f
LoraineMcCune
inrelationtotheground),deixis(directionwithrespecttothespeaker),man-
ner(e.g.,
walked
ran
rolled
vs.
)andcause(e.g.,transitivevs.intransitive)
(Talmy1975).
Motioneventsalsoprovidethereferencepointfortheinitialrelationalmean-
ingsreportedforyoungchildren.They,alongwithgenerictransitiveverbs,form
thecoreofchildren
Mirrorneurons
registrationofbiologicalmotion

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Lookingforneuralanswers
tolinguisticquestions
BernardH.Bichakjian
UniversityofNijmegen,TheNetherlands
Towardaconcertedapproach
Longleftalmostexclusivelytolinguists,languageisattractingmoreandmoreat-
tentionfromthevariousbranchesofbiology.Forthecollectiveefforttobearfruit,
theremustbeaminimumofsharedknowledge.Ontheonehand,linguistswill
havetolearntoputasidetheirprejudicesaboutevolution,while,ontheother

BernardH.Bichakjian
deep-rooted,butiftheyaretocontributetotheunderstandingoflanguage,lin-
guistsmustlearntoworkwiththeevolutionaryparadigm,becauseif,asDobzhan-
skypointedout,
nothinginbiologymakessenseexceptinthelightofevolution
(quotedfromMayr1997:178),notrueunderstandingoflanguagecanbeachieved
exceptinthelightofevolution.
Sayingthatthebrainofapesismoredevelopedthanthatofdogsisnotsome
formofracismagainstcanines;arguingthatwarm-bloodednessconfersselective
advantagesisnotasignofcrueltyagainstcrocodiles;pointingoutthebene
tsof
thedomesticationofplantsandanimalsisnotanactofdiscriminationagainst
hunter-gatherers;andrecognizingthegreatereffectivenessof
rearmsoverbows
andarrowsisnotahumanrightsviolation.Likewise,itmustbepossibleforlin-
guiststoassessandcomparetheproductioncostandthefunctionalyieldoflin-
guisticitemsandconcludethatitemXismoreadvantageousthanitemYwithout
theirbeingaccusedofunethicalbehavior.
Eveninbiology,whereitisfullyaccepted,
evolution
isadif
cultconcepttode-
ne.Thegradualincreaseinsizeandcomplexityofthebrainisanunquestionable
caseofevolution,andsoarethechangesundergonebyseals,whoseancestors
ter-
restrialfeatureshavebeenmodi
edintoalternativessuitableforasemi-aquaticlife.
Inbothcases,thechangesareadaptive.Evolutionistswouldalsoarguethatsome
changescanbetotallyneutral,but,byandlarge,evolutionisseenastheaccumula-
tionofmutationsthatconfertheirbearersselectiveadvantagesandeventuallylead
totheformationofanewspecies(Mayr1963:621&1988:253).
Inlinguistics,ifagivenfeature
thatis,aspeechsound,agrammaticaldistinc-
tion,orasyntacticstrategy
hasbeenconsistentlyreplacedwithanewalternative,
itwillbethetaskofthelinguisttotracktheselectiveadvantagesthattheincom-
ingfeaturehasoveritsantecedent.Aneasyexampleiswordorder.Theancestral
wordorderwashead-lastorSOV,themodernoneishead-
rstorSVO.Thisperva-
siveshiftneedsanexplanation,anditisincumbentuponlinguiststouncoverthe
selectiveadvantagesofthemodernwordorder.
Thepatternofevolutionmayseemconfusing.If,forinstance,wefocusonthe
evolutionofclassesfrom
shestomammals,theprocesslookslinearandunidi-
rectional
shesgaverisetoamphibians,amphibianstoreptilians,andreptilians
tomammals,andneverwastheprocessreversed.Ontheotherhand,ifthefocus
isonthespecializedlegandheadanatomyofwoodpeckers,the
sh-likefeatures
ofwhalesanddolphinsortheatrophiedwingsofkiwistheobviousconclusionis
thatdiversityorevencoursereversalispossibleinthecaseofecologically-adapted
features.Evolutionthereforeneverimpliesaneatlylinearpattern,nordoesitmean
thetotalextinctionofancestralspecies.Thesurvivaloftoday
sreptiliansdoesnot
belietheirbeingancestraltomammals,northefactthatmammalshaveselective
advantagesoverreptilians.Likewise,returningtothewordorderexample,theex-
Lookingforneuralanswerstolinguisticquestions

istenceofextantSOVlanguagescannotbeusedtodisputeSOVbeingancestralto
SVO,northefactthatSVOhasselectiveadvantagesoverSOV.
Whatevolutionistsshouldknowaboutlanguage
Ifitisnecessaryforlinguiststolosetheirmisgivingsaboutevolutionaryreasoning,
evolutionists,andindeedotheradvocatesofinnatemodelswouldmakecontribu-

BernardH.Bichakjian
mergedpart,butwecansurmiseitsexistenceanditsbeingidenticalinnaturewith
thevisiblepart.Wehavenopossibilityoftracingthedevelopmentsthatledtothe
featuresfoundatthedawnoftheempiricalperiod,butwehavenoreasontodoubt
thattheyweretheproductofsteadydevelopmentsofwhichtheobservedonesare
thecontinuation.
Scientistswhoaretryingonthebasisoffossilindicationstodeterminewhen
inthehominidlineofevolutionmodernlikespeechbecamepossible,mayrightly
arguethatintheabsenceofagivenindicator
largercanalsfortheinnervationof
thetongueorlargerductsfortheinnervationofthethoracicmuscles
oral
Lookingforneuralanswerstolinguisticquestions
\b
triguing,butbeforeexaminingitspossiblesigni
cance,Ishalldiscusstworadical
changes
onethathasplayedamajorroleintheevolutionofsyntaxandasimilar
oneinthedevelopmentalhistoryofwriting.
Whywastheoriginalwordorderheadlast?
Insyntax,theoriginalorderwasheadlast,
i.e
.,verbscameaftertheirobjects,auxil-
iariesafterparticiples,prepositionsaftertheirobjects,adjectivesandgenitivesbe-
foretheirmodi
\t
BernardH.Bichakjian
anas-you-goprocessing,itisonlylogicalfortheoriginalwordordertohavebeen
headlast,andforthemodernonetobehead
rst.
Thisindeedislogicalreasoningdoneonthebasisofwhatisknownofthetwo
cerebralhemispheres(Levy1974:167;Posner&Raichle1994:94&162;Gazzaniga
salofwordorderisaportentousandheuristicallyimportantphenomenonwhich
cannotbebrushedasidewithadhocexpedientsandrelativisticplatitudes.Itis
incumbentuponlinguists,alongwithneuroscientists,toexplainwhytheoriginal
wordorderwasheadlastandwhyitwasreversed.
Whywasthedirectionofwritingreversed?
Sincespacedoesnotallowevenanoutlineofthedevelopmentofwritingfrom
(vanSommers1991:5,10
11,20),buttherewasperhapsanotherimportantfactor
onwhichneurologycanshedlight.
Ifwebearinmindthatpictorialobjectsaregenerallyperceivedbytheright
hemisphere,andthattherighthemispherenormallyscansleftward,whilethelin-
Lookingforneuralanswerstolinguisticquestions
\n
Thesigni
canceofmirrorneuronsforlanguageorigin
ringofneuronsinthesimianhomologueofBroca
sarea,bothwhenthemon-
keyisgraspinganobjectandwhenitseesthegraspingoftheobject,hasprompted
observerstohypothesizethattheperformanceand/orthewitnessingofanaction
couldhavebeenthestimulusthatledtotheemergenceofasystemofcommu-
Figure1.
BernardH.Bichakjian
cortexofthelefthemisphere
withtheactivationpattern
closelycorresponding
thatofthespatialintelligencetest,whichalsodisplaysanactivationpatterninthe
Lookingforneuralanswerstolinguisticquestions
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n,T.(1979).
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vocalbehavior.
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J.GrahamBeaumont(Eds.),
HemisphereFunctionintheHumanBrain
London:ElekScience.
Lieberman,P.(1998).
EveSpoke
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MacLarnon,A.M.,&Hewitt,G.P.(1999).Theevolutionofhumanspeechtheroleof
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Science,288
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AnimalSpeciesandEvolution
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Mayr,E.(1988).
TowardaNewPhilosophyofBiology
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Mayr,E.(1997).
ThisisBiology
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proto-world
wordorder.InC.Knight,
M.Studdert-Kennedy,&J.Hurford(Eds.),
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(pp.372
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UniversityPress.
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Acognitive-anatomicalanalysis.
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cAmerican
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vik(Eds.),
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GraphicSkills:Researchperspectivesandeducationalimplications
(pp.3
38).London:
AcademicPress.
Mirrorneuronsandculturaltransmission
IndiaMorrison
DepartmentofHumanities,UniversityofSk
vde,Sweden
IndiaMorrison
inthesharingofmentalrepresentations
essentialforthetransmissionofcultural
informationfromonepersontothenext.
HereIshallfocusontworathermorequirkyaspectsofhumanculturalbe-
havior,
catchymemes
migratorymannerisms.
Catchymemesarethose
Mirrorneuronsandculturaltransmission
ofthetermmemeshouldbetakensomewhatindependentlyofthissense;
meme
isausefulexistingwordforcertainphenomenathatarefamiliartoeveryone.
Oneofthehopesofacognitiveneuroscienceofculturaltransmissionistoil-
luminatemechanismsofculturaltransmissionwhilealsoprovidinganadaptive
explanationforcatchymemesandmigratorymannerismsthatdoesnotnecessar-
ilystipulatetheinterestsofentitiesapartfromhumans.(Butthisdoesnotpre-
cludethepossibilitythatcultureisneverthelessaDarwinianevolutionarysystem,
averyimportantpossibilitythatcanalsobeexploredinlightofneurophysiological
mechanisms.)Agoodwayofbeginningthisendeavorwouldbetoask,
Whatneu-
rophysiologicalfactorscanin
uencethelikelihoodthatanobservedactionwillbe
repeatedbytheobserver?
Thereisahostofmechanismsthatcanbesweptunder
thisbroadheading,butthreeofthemostpivotalarethoseof:
1.Memory;
2.Patternsofexcitationandinhibition;
3.Emotion.
Theroleofmirrorneurons,orthehumanequivalent,wouldbepervasiveamong
thesein
uentialfactors.Inmemory,theywouldcomeintoplayinthepercep-
tionofbehaviorthatisrememberedandlaterperformedbytheobserver.Their
IndiaMorrison
Mirrorneuronsandculturaltransmission
IndiaMorrison
interlocutor
smeaning,intention,andsocialpositionduringconversation,there
ishighinformationvaluenotonlyinthemovementsthemselvesbutintheway
theyareorganizedinthesyntaxofconversationalmovement.Socialinteractions
enlistasuiteofcognitivemechanisms.Forinstance,complexexecutiveprocessing
occursinthefrontallobe,andemotionalprocessingisdistributedthroughoutthe
brain.Emotionin
uencesfuturebehaviorbycouplingperceptionwithdisposition
toact(e.g.LeDoux1998).Ratherthanhavinganimmediatein
uenceonfuture
behavior,mirrorperceptionmorelikelycontributesatarelativelyearlystagetoa
cascadeofresponseswhichcoupleperceptionwithaction,andactiondisposition
withmemory.Brothers(1995)hasproposedthatcomplexshadesofperceptual-
emotionalresponseinsocialinteractionsarere
ectedbycorrespondinglycomplex
pro
lesofactiondispositioninthebrain.ExpandingDamasio
sterm,shesug-
geststhatemotionalactiondispositionsarethus
somaticallymarked
(Damasio
Mirrorneuronsandculturaltransmission
IndiaMorrison
Plotkin,H.(1997).
Evolutioninmind
.London:PenguinBooks.
Rizzolatti,G.etal.(1996).Premotorcortexandtherecognitionofmotoractions.
Cognitive
BrainResearch,3
,131
Sperber,D.(1996).
ExplainingCulture:Anaturalisticapproach
.Oxford:Blackwell
Publishers.
Wong,W.,&Barlow,H.(2000).Tunesandtemplates.
Nature,404
,952
Applications
Mirrorneuronsandtheneuralbasis
forlearningbyimitation
Computationalmodeling
AudeBillardandMichaelArbib
UniversityofSouthernCalifornia,LosAngeles,USA
Introduction
Nothingissocontagiousasanexample,andweneverdosuchgoodactsand
suchbadactsthatwedonotproducesimilarones.Weimitatethegoodactions
byemulation,andthebadonesbythemalignityofournature,thatshamekept
AudeBillardandMichaelArbib
Whatisimitation?
Thereisstilldebateconcerningwhatbehaviorstheterm
imitation
referstoandin
whichspeciesitisexhibited(Byrne&Whiten1988;Tomasello1990).
True
imita-
tioniscontrastedtomimicry.Imitationismorethanthemereabilitytoreproduce
others
actions;itistheabilitytoreplicateandlearnskillswhicharenotpartofthe
susualrepertoiresimplybytheobservationofthoseperformedbyothers.
Thecurrentagreementisthatonlyhumansandpossiblyapesareprovidedwiththe
abilityfortrueimitation,althoughrecentdata(Myowa-Yamakoshi&Matsugawa
1999)suggestthatimitationofmanipulatoryactionsinchimpanzeesisquitelim-
itedcomparedtothatofhumans.Typically,chimpanzeestook12trialstolearnto
imitate
abehavior,andindoingsopaidmoreattentiontowherethemanipulated
objectwasbeingdirected,ratherthanactualmovementofdemonstrator.
Simplerformsofimitationandmimicryhave,however,beenshowninrats
(Heyes1996),parrots(Moore1996),mynahbirds(Nottebohm1976),dolphins
(Hermann2000),andmonkeys(Kawamura1963;Visalberghi1990).Imitationis
interestingtodevelopmentalpsychologistsbecauseitunderliesthehumanchild
growingcapacityforrepresentationandsymbolization.Thedevelopmentofcom-
pleximitationabilitiesinchildrenaccompaniesthegrowthofthechild
scommu-
Mirrorneuronsandtheneuralbasisforlearningbyimitation
Acomputationalmodeloflearningbyimitation
Themodel(seeFigure1)isbiologicallyinspiredinitsfunction,asitscompo-
nentmoduleshavefunctionalitiessimilartothoseofspeci
cbrainregions,and
initsstructure,asthemodulesarecomposedofarti
cialneuralarchitectures
AudeBillardandMichaelArbib
Learning System
Cerebellum module
Learning oftemporal
sequences ofmovement
PMd module
Learning ofcoordinated
activation ofnodes in M1
Figure1.
Themodelconsistsofthreepartsforvisualrecognition,motorcontroland
learningandiscomposedofbiologicallyinspiredmodules,namelythetemporalcortex
(STS),thespinalcord,theprimarymotorcortex(M1),thepremotorcortex(PM)and
thecerebellum.Thebottomleftpanelindicatesthetakingofdatafromvisualinput
(extractingthearmandbodymovementsofahuman);therighthandpanelshowsan
avatarimitatingtheobservedmovement.
Mirrorneuronsandtheneuralbasisforlearningbyimitation
Cerebellum
DRAMA Networks
Attentional Mechanism
Leg
Elbow
STS
Frame ofReference Transformation
Video input
Spinal Cord
Mechanical Simulation 65 DOF 2 muscles ext-flexor per DOF
Somatotopic Body Map
Flexor
Extensor
Figure1.
(continued)
Neuralmechanismsbehindimitation
Theconfusionbehindthedifferentde
nitionsofimitationresultsinpartfroma
lackofinformationconcerningtheneuralmechanismsatthebasisofthisabilityin
animals.Motorskillimitationreliesontheabilitytorecognizeconspeci
actions
andtotransformvisualpatternsintomotorcommands.Whilethereisanimpor-
tantbodyofliteratureonvisualrecognitionofmovementsandonmotorlearn-
AudeBillardandMichaelArbib
Mirrorneuronsandtheneuralbasisforlearningbyimitation
Mirrorneurons,imitationandlanguageacquisition
Westartedthisbriefarticlebymentioningthatimitationunderliescognitivepro-
cessesfundamentaltosocialcognition.Toconcludethisarticle,webrie
ysum-
marizeanimportantgoalofourresearch,namely,tolinktheabilitytoimitateto
thatforlanguage.Thisrelatesdirectlytotheconferenceon
Mirrorneuronsand
theevolutionofbrainandlanguage
fromwhichthebookfollows.Theconference
itselfwasinspiredinpartbythepaperbyRizzolattiandArbib(1998).Theob-
servationthattheareaF5(whichcontainsthemirrorneuronsystem)inmonkeys
couldcorrespondtoBroca
sareainthecorrespondingarea6ofthehumanmotor
cortexledtheauthorstoproposethat
humanlanguage[...]evolvedfromabasic
mechanism[themirrorneuronsystem]thatwasnotoriginallyrelatedtocommu-
nication
.Arbib(2001)extendedthisideabypostulatingthattheabilitytoimitate
providedacrucialbridgefromthemonkeymirrorneuronsystemtothehuman
abilityforlanguage.
Itisinterestingtorelatetheseideastothepsycholinguisticliterature(Gar-
AudeBillardandMichaelArbib
Figure2.
Thedoll-shaperobotmirrorsthemovementsofthearmandheadofthe
humandemonstrator.Whiledirectingtherobot,thedemonstratorinstructstherobot
(sayinge.g.
Youmoveyourrightarm
).Throughtrainingtherobotlearnstodissociate
themeaningofeachwordinthedemonstrator
ssentencesandtoassociateitwithits
perceptions.
monkeysandchimpanzees,beforetheycouldbefullyexpressed,asitisthecase
inhumans.
Ourfutureworkwilladdressthesedifferentissues.Inpreviouswork(Billard
2000;Billard&Hayes1999),wefollowedanengineeringapproach,exploitingthe
imitationgametoenhancetherobustnessofthesystemandexploitingthefact
thatimitationandlearningofaregularlanguagecouldbeprogrammedusingthe
samearti
Mirrorneuronsandtheneuralbasisforlearningbyimitation
Arbib,M.A.,Billard,A.,Iacoboni,M.,&Oztop,E.(2000).Syntheticbrainimaging:Grasp-
ing,mirrorneuronsandimitation.
NeuralNetworks,13
,975
Billard,A.(1998).DRAMA,aconnectionistmodelforrobotlearning:Experimentson
groundingcommunicationthroughimitationinautonomousrobots.PhDthesis,Dept.
ofArti
cialIntelligence,UniversityofEdinburgh,U.K.
Billard,A.(2002).Imitation:Ameanstoenhancelearningofasyntheticproto-languagein
anautonomousrobot.InC.Nehaniv,&K.Dautenhahn(Eds.),
ImitationinAnimals
andArtifacts
(pp.281
310).Cambridge,MA:MITPress.
Billard,A.(1999).Learningmotorskillsbyimitation:abiologicallyinspiredroboticmodel.
AudeBillardandMichaelArbib
Meltzoff,A.N.(1990).Thehumaninfantasimitativegeneralist:a20-yearprogressreport
oninfantimitationwithimplicationsforcomparativepsychology.InHeyesC.M.,&
B.G.Galef(Eds.),
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(pp.347
370).New
York:AcademicPress.
Meltzoff,A.N.,&Gopnik,A.(1989).Onlinkingnonverbalimitation,representationand
language:Learninginthe
rsttwoyearsoflife.InG.E.Speidel,&K.E.Nelson(Eds.),
TheManyFacesofImitationinLanguageLearning
(pp.23
52).Berlin:SpringerVerlag.
Meltzoff,A.N.,&Moore,M.K.(1977).Imitationoffacialandmanualgesturesbyhuman
neonates.
Science,198
,75
Moore,BruceR.(1996).Theevolutionofimitativelearning.InC.M.Heyes,&B.G.
Galef(Eds.),
SocialLearninginAnimals:Therootsofculture
(pp.245
265).NewYork:
AcademicPress.
Myowa-Yamakoshi,M.,&Matsuzawa,T.(1999).Factorsin
uencingimitationofmanip-
ulatoryactionsinchimpanzees(
Pantroglodytes
JournalofComparativePsychology,
,128
Nadel,J.,Guerini,C.,Peze,A.,&Rivet,C.(1999).Theevolvingnatureofimitationasa
formatforcommunication.InJ.Nadel,&G.Butterworth(Eds.),
ImitationinInfancy
(pp.209
234).Cambridge:CambridgeUniversityPress.
Nottebohm,F.(1976).Vocaltractandbrain:asearchforevolutionarybottlenecks.
Annals
ofNewYorkAcademyofSciences,280
,643
Oztop,E.,&Arbib,M.A.(2002).Schemadesignandimplementationofthegrasp-related
mirrorneuronsystem.
BiologicalCybernetics
(toappear).
Piaget,J.(1962).
Play,DreamsandImitationinChildhood
.NewYork:Norton.
Rizzolatti,G.,&Arbib,M.(1998).Languagewithinourgrasp.
TrendsinNeurosciences,21
Rizzolatti,G.,Fadiga,L.,Gallese,V.,&Fogassi,L.(1996).Premotorcortexandtherecog-
nitionofmotoractions.
CognitiveBrainResearch,3
,131
Rizzolatti,G.,Fadiga,L.,Matelli,M.,Bettinardi,V.,Perani,D.,&Fazio,F.(1996).
Localizationofgrasprepresentationsinhumansbypositronemissiontomography:1.
Observationversusexecution.
ExperimentalBrainResearch,111
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Speidel,G.S.(1989).Imitation:Abootstrapforlearningtospeak.InG.E.Speidel&K.E.
Nelson(Eds.),
TheManyFacesofImitationinLanguageLearning
(pp.151
180).Berlin:
SpringerVerlag.
Tomasello,M.(1990).Culturaltransmissioninthetooluseandcommunicatorysignaling
ofchimpanzees.InS.T.Parker&K.R.Gibson(Eds.),
LanguageandIntelligencein
MonkeysandApes:Comparativedevelopmentalperspectives
(pp.274
311).Cambridge:
CambridgeUniversityPress.
Trevarthen,C.,Kokkinaki,T.,&Fiamenghi,Jr.G.A.(1999).Whatinfants
communicate:Withmothers,withfathersandwithpeers.InJ.Nadel&G.Butterworth
(Eds.),
ImitationinInfancy
(pp.61
124).Cambridge:CambridgeUniversityPress.
Visalberghi,E.,&Fragaszy,D.(1990).Domonkeyape?InS.T.Parker&K.R.Gibson(Eds.),
LanguageandIntelligenceinMonkeysandApes:Comparativedevelopmentalperspectives
(pp.247
273).Cambridge:CambridgeUniversityPress.
Mirrorneuronsandfeedbacklearning
SteveWombleandStefanWermter
CentreofInformatics,UniversityofSunderland,U.K.
Introduction
Theneuroscienceevidencereviewedin(Rizzolatti&Arbib1998)suggeststhatmir-
rorneuronsareinvolvedinthecomparationof
goaldirectedactions
andtheper-
SteveWombleandStefanWermter
ofrecognitionofcorrectconstructions,andthisadditionalknowledgecanbeinte-
gratedintotheproductionprocess.Secondly,wecanmeasurehowwellthesystem
estimatesithasalreadystoredtheinformationcontainedintheexamplepresented.
Itcanthenmodifythedegreetowhichitadaptsitselftooptimisefornovelinfor-
mation.Notethatthismeasuredoes
not
requirefeedbackitselftobecalculated.It
issimplyanestimategeneratedbythelearneronitsownuseofacquiredknowl-
edge.Itdoesnotmeasureifthatknowledgeisna
veorincorrectonlythedegreeto
whichitisused.Howeverasthelearner
sacquiredknowledgeimprovesthisutil-
isationrepresentationmapsintoacon
dencemeasure.Thiscanbe
quanti
feedback.
AhighlevelrepresentationofourmodelisgiveninFigure1.Thepartsin
thesquareboxesrepresentthe
mirrorneuronsystem
thatwehavedevelopedour
abstractmodelof.Inourmodelthelearnerexaminesanewlygeneratedrepresen-
tationanddeemstheproducedsequencetobeworthyofproductiononlyifthe
utilisationmeasureissuf
cientlyhighforallpartsofthesequence
thelearner
knowsthatitsknowledgeisgood.Thisrequiresthecalculationofathresholdfor
lter.Thisiscalculatedinacomputationallyef
cientnon-neuralmanner.Itrep-
resentsaminimum
ringratenecessaryforallneuronstobe
ringatinorderto
driveamirrorneuron.
EXTERNAL
EXTERNAL
FEEDBACK
KNOWLEDGE
SYSTEM
UTILISATION/
REPRESENTATION
MIRROR
NEURONS
EXTERNAL
TENTATIVELY
PRODUCED
Figure1.
Highlevelrepresentationofmodelmirrorsystem.
Mirrorneuronsandfeedbacklearning
Experimentalneuroscienceresultsindicatethatmirrorcellsarehighlyselec-
tive,only
ringwhentheirspeci
cassociatedgoaldirectedactionoccurs(Rizzo-
latti&Arbib1998).Inordertofacilitatethiswithinourmodelwerequirethatin
additiontoinputfromtheutilisationsystemthemirror
eldreceiveshighlevel
inputfromboththesequenceproductionmechanismandfromthecentralexec-
utive
orexternalinput
elds.Theformerselectivelystimulatesindividualmirror
neuronsatasubcriticallevelallowingforfullactivitytooccurwhenadditional
stimulationisreceivedfromtheutilisationsystem.Thelatterisnecessarytoform
T
X
X
V
T
V
S
S
P
P
1
2
3
4
5
6
E
Figure2.
TheDSFSGdevisedbyReber.
SteveWombleandStefanWermter
backpropagationfortraining.Trainingsequencesweregeneratedstochasticallydi-
rectlyfromtheDSFSG.Wesummederrorsoverasequencebeforeupdatingthe
Mirrorneuronsandfeedbacklearning
SteveWombleandStefanWermter
Results
The1-normerrorwascarefullycalculatedusingthe755mostfrequentstringsgen-
eratedbytheDSFSG,witheachstringweightedaccordingtotheasymptoticfre-
quencydistribution.Forthisvaluealittleover97%oftheasymptoticfrequency
distributionofthein
nitesetofReberstringsisspanned.Thisisadeepsearch,
withtheprobabilityoftheleastfrequentofthesestringsoccuringnaturallybe-
ingonly2
1in32000.Wefounditquiteeasytoreducethe1-normerror
perelementinasequence,perstringtobelow0.1,andwithalittlemoreworkto
Training
1-norm
bias
bias
Filtered
ParadigmError
Mean
SD
Success
BP&Filter0.0173
22.87e-061.65e-06100.0%
MNSystem0.0085
12.32e-061.04e-06100.0%
DSFSG
4.904e-061.35e-06100.0%
IncorrectUn
lteredMin
Max
RejectionsSuccess
Pass
Fail
BP&Filter0.00%
0.49050.0795
MNSystem0.00%
0.86460.1361
DSFSG
1.00000.0000
Theresultsquotedarebasedon5000selfgeneratedteststringsforallmeasures
basedonsystemproduction,andwererepeated10timesforthecalculationof
bias
meansandstandarddeviations.
Mirrorneuronsandfeedbacklearning
Behavioural Results for Basic SRN
Mean SRN
Mean SRN +/
Mean SRN +/
Mean DSFSG
Mean DSFSG +/
Mean DSFSG +/
Difference Measure
50100150200250300350400450500
Figure3.
Plotofdifferencemeasureforminimum1-normerrorfoundusingthebasic
\f
SteveWombleandStefanWermter
Behavioural Results for Full Mirror System
Mean SRN
Mean SRN +/
Mean SRN +/
Mean DSFSG
Mean DSFSG +/
Mean DSFSG +/
Figure4.
Plotofdifferencemeasureforminimum1-normerrorfoundusingtheab-
stractedmirrorsystem.ThedifferencemeasurefortheDSFSGisgivenforreference.
Themeanandstandarddeviations(SD)werecalculatedfrom20trialsforeachsizeof
Mirrorneuronsandfeedbacklearning

techniquewehaveintroducedinthefeedbackmechanism.Usingtheutilisation

SteveWombleandStefanWermter
stractedmirrorneuronsystemtobothanalysetentativeproductionandtomodify
thelearningprocesstheproductionperformanceofalearningagentonthesyn-
taxacquisitiontaskpresentedbytheReberGrammarcanbecomebehaviourally
Aconnectionistmodelwhichuni
esthe
behavioralandthelinguisticprocesses
Resultsfromrobotlearningexperiments
YuuyaSugitaandJunTani
BrainScienceInstitute,RIKEN/GraduateSchoolofArtsandSciences,
UniversityofTokyo
Introduction
Howcanrobotscommunicatewithhumansortheotherrobotsusingnaturallan-
guage?Toachievethis,itisnecessaryforrobotstounderstandthemeaningofsen-
tences.However,whatdoes
themeaningofsentences
mean?Weinvestigatethis
problemfromtheviewpointofembodimentandintersubjectivity.Inotherwords,
weconsiderthatnotonlythesymbolicinteractionbutalsothebehavioralinterac-
tionamonghumansandrobotsareindispensableintheprocessesofco-acquisition
oflanguageandbehavior.
Tounderstandtheprocessesofco-acquisitionoflanguageandbehavior,we

YuuyaSugitaandJunTani
theactiveinvestigationoflanguagebasedontheAIapproachaimingatunderstant-
ingtheintelligentbehaviororconstructingandconstruingintelligence.Howevera
systemequivalenttoAIwhichcankeepsustainablecommunicationwithhuman-
beingshasnotbeenachievedyetsincethesymbolspre-assignedinthesystemsof-
tencauseaframeproblemaswellasasymbol
groundingproblem(Harnad1990,
Toavoidtheseproblems,thereareroughlytwostrategies:(1)denyingthe
necessityoftheinternalrepresentationandavoidanceofsymbols,and(2)au-
tonomouslygeneratingthesymbolswhichwouldbeusedforinternalrepresen-
tation.Intherangeoftheroboticsresearch,itisnotablethatthedescriptionof
theenvironmentshouldbeconstructedfromtherobot
sownview.Thatistosay,
Amodeltounifybehaviorandlanguage

Inthischapter,weproposeanovelmodelwhichrepresentsthelinguisticpro-
cessandthebehavioralprocessasco-dependentdynamicalsystems.Theessential
argumentsinourmodelingaresummarizedasfollows.
Behaviordenotesthestructureofthesensory-motorsequenceswhicharise

YuuyaSugitaandJunTani
hidden
input
hidden
input
context
context
(Sensory-Motor Module)
(Linguistic Module)
sensory-motor image
word
prediction ofthe next sensory-motor image
prediction ofthe next word
Figure1.
OurproposedRNNarchitecture:TwoRNNsareconstrainedeachother
throughapartoftheircontextunits.
isforbehavior,and
isforlanguage.
EachRNNmodulepredictstheinputimageofthenexttimestepfromthecurrent
inpututilizingthecontextunitsactivations.
Amodeltounifybehaviorandlanguage
\b
thesystemcanbeidenti
ednotbyexternalinputvaluesbutbycontextactivation
values.
Aschemecalled
context-binding
isdevelopedfororganizingthemany-to-
\t
YuuyaSugitaandJunTani
Figure2.
Ourvision-basedmobilerobot.
Setting
Figure3showstheenvironmentwheretherobotlearningexperimentswerebeing
conducted.Therearetwoobjectsintheenvironment
aredpostboxandablue
block.Aftereachcommandsentenceisgiventotherobot,therobotistrainedto
generateanappropriatesequenceofactionsinasupervisedlearningmanner.
Therobothasthevisionsensorwhichrecognizesthecolorandthesizeofthe
objects.Therobotcanmaneuverbychangingtherotationspeedofleftandright
wheels.Itcanattendtotheobjectsbyrotatingthecameraheadhorizontallyand
vertically.Itcanalsopushtheobjectsforwardwithitssimplearmequippedinthe
frontside.Intheenvironment,theblueblockisplacedrightinfrontofthewall
Amodeltounifybehaviorandlanguage
\n
Figure3.
Theenvironmentforourexperiment:Ablueblock(left)andaredpostbox
(right)havebeenplaced.
totheseactionmodules.Italsohas
vesensoryinputunitswhichrepresentvisual
information(colorandsizeoftheobjectsstared).Thesensoryinputsarefedinto
YuuyaSugitaandJunTani
(b)
Figure4.
Anexampleofarobot
sbehaviorintheenvironment.
Results
Theco-learningexperimentswereconductedusing60sequencesofsentencesand
theircorrespondingbehavioralsequences.Thelearningprocesstook400,000iter-
ativestepsbeforetheconvergence.
Thenthephaseplotdiagramusingthecontextunitsactivationsaregener-
atedbothforthebehavioralRNNandthelinguisticRNN.Thephaseplotswere
generatedbyiterativelyactivatingtheRNNsintheclosed-loopmodewithinputs
comprising300stepsofpossibleactionsequences.Thegeneratedsequencesofthe
contextunitsactivationareplottedinthetwodimensionalphasespaceasshown
inFigure5.
Inthis
Amodeltounifybehaviorandlanguage
1.0
0.0
(b)
1.0
Figure5.
ThephaseplotdiagramsforthebehavioralRNNontheleftsideandthe
linguisticRNNontherightsideafter400,000iterativestepsofco-learning.
1.0
1.0
Figure6.
ThephaseplotdiagramforthelinguisticRNNafter400,000iterativestepsof
co-learning.Thecontextvaluesofthebindingstepsareplotted.
Wealsostudiedhowthecontextstatestransitamongtheseclustersastherobot
YuuyaSugitaandJunTani
1.0
1.0
blockpostbox
frontrear
robot
(2)
(3)
(4)
(5)
Figure7.
Amodeltounifybehaviorandlanguage
1.0
1.0
Figure8.
Thephaseplotsgeneratedafter200,000iterativestepsinindependent
learning.
Inthecaseoftheindependentlearningoflinguisticsequences,theattractorap-
pearsscatteredlikeclouds,andnoclusterstructurescanbeobserved.Thisoutcome
impliesthatindependentlinguisticlearningresultsinrote-learningofthetraining
dataratherthangeneralizedlearningwithextractingstructures.
Theseresultsareduetothefactthatthesequencesofsentencesgiventothe
robotarenotmerelyrandom,buttheyaregiveninacontext-dependentman-
YuuyaSugitaandJunTani
Amodeltounifybehaviorandlanguage
YuuyaSugitaandJunTani
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Nameindex
Agayoff,J.316
Aglioti,S.139,146
Aiello,L.182
Ainsworth,M.296
Aitken,P.G.177
Akhtar,N.288
Allman,J.M.186
Altschuler,E.L.40
Andersen,R.A.49
Anderson,S.W.8
Arbib,M.4,9,29,77,115,116,123,
125,151,165,202,208,220,221,
223,229,251,253,283,284,289,
290,273,274,315,326,329,348,
349,353,355,374
Aslin,R.N.306,307
Awh,E.78
Baddeley,A.78,84
Bailey,P.47
Baldissera,F.41
Baldwin,J.M.187
Balter,M.179
Bard,K.A.285
Bargh,J.A.153
Barlow,H.336
Baron-Cohen,S.30,139,258,264
Barresi,J.116
Bavelas,J.B.164
Becker,J.T.78
Beebe,B.8,296
Beer,R.D.364
Bekkering,H.4,102,104,106,115
Bell,B.D.29
Bell,M.A.316
Benjamin,G.R.152
Bente,G.153,154
Benton,A.L.102
Berger,L.182
Berlucchi,G.139,146
Bernhardt,B.H.311
Bert,J.39
Bertamini,M.163
Best,D.L.153
Best,C.T.308
Bichakjian,B.7,325,326
Bickerton,D.116,222,325
Billard,A.9,345,349,350,365
Binkofski,F.48
Biraben,J.-N.179
Blackmore,S.334
Bloom,P.325
Boker,St.5,151,155,167
Bolton,D.146
Bottini,G.145
Bowerman,M.319
Bowlby,J.285
Boysson-Bardies,B.307,308
Brain,W.R.29
Brass,M.54
ten,St.7,273
276,278,
282,287
290,298
Brodmann,K.47
Brooks,R.364
Brookshire,R.H.145
Brothers,L.338
Browman,C.P.209,210,214
Brown,J.D.280
Brownell,H.H.145
Bryan,K.L.145
Nameindex
Buccino,G.29,43,46,50,132
Bucher,M.A.327
Buchtal,F.317
Bull,P.E.152
Burgess,P.145
Burgoon,J.K.152,153
Butterworth,G.101
Byrne,R.54,284,344
Call,J.19
Calvin,W.H.116
Campos,J.319
Cann,R.L.201
Carey,D.P.37,51
Carruthers,P.140
Carstairs-McCarthy,A.223
Cavada,C.23
Cavalli-Sforza,L.L.333
Chatrian,G.E.39
Cheney,D.L.177,201
Choi,S.319
Chomsky,N.184,199,202,209,251
Cirillo,L.102
Cleeremans,A.353,355,361
Cochin,S.40
Cohen-Seat,G.39
Clark,H.4,117,123
Colby,C.I.49
Collard,M.200
Condon,W.S.151,152
Cooper,R.P.306
Corballis,M.C.125
Courtney,S.M.78
Cowan,N.310
Craighero,L.3,55
Creider,C.A.152
Damasio,A.R.139,231,247,338
Darwin,C.52,221,223,317,323
Davis,B.214,224,326
Dautenhahn,K.365
Dawkins,R.334
DeCasper,A.J.307
DeRenzi,E.E.49
Deacon,T.W.73,74,222,237,326,
Decety,J.42,93
DeGusta,D.326
Dennett,D.1,334
DePaolis,M.214,218,305
Dillman,L.153
Dimberg,U.52,337
DiPellegrino,G.101,111,281,310,
Donald,M.209,221,222,274,286,
Dovidio,J.F.153
DuBois,J.203
Duffy,J.R.29
Duhamel,J.-R.49
Duncan,S.D.153
Dunkan,J.329,330
Dunn,J.288
Edgar,B.181
Eibl-Eibesfeldt,I.281
Ekman,P.152
Elman,J.185,356,364
Engelkamp,J.89
Faaborg-Anderson,K.L.317
Fadiga,L.3,15,40,41,102,110,
115,126,222,224,251,255,302,
310,347
Fagg,A.H.348
Falk,D.202
Farwell,L.213,311
Feldman,M.W.333
Ferguson,C.A.213,311
Fernald,A.307
Ferrari,P.F.30,264
Fifer,W.P.307
Fink,G.R.145,146
Fiske,D.W.153
Nameindex
Fodor,J.7,253,255,256
Foerster,O.47
Fogassi,L.1,3,23,26,37,49,51,
115,257,269
Foldi,N.S.145
Fowler,C.A.209,211,220
Fox,N.A.316
Fragaszy,D.19
Friederici,A.D.306,307
Friesen,W.V.152
Friston,K.J.127
Fromkin,V.213
Fuster,J.M.146
Gainotti,G.29
Gallagher,H.L.140,145
Gallagher,S.140
Gallese,V.1,3,15,16,18,22,23,25,
27,28,37,51,53,115,116,135,
136,146,165,251,255,253,257,
260,261,269,275,289,295,303,
326,347,355,374
Garey,L.J.70
Garton,A.F.349
Gastaut,H.J.39
Gattis,M.115
Gazzaniga,M.S.328
Gell-Mann,M.230,231,247
Gentilucci,M.14,53,125
Gertsman,L.296
Gibson,K.193
n,T.327
Gold,E.M.353,361
Goldberg,M.E.49
Goldman,A.136,146,165,253,
275,289,374
Goldman-Rakic,P.S.23,78,84
Goldstein,L.209,210,214,224
Goodell,E.W.214,215
Goodwin,C.337
Gopnik,A.140,344,349
Gould,S.J.72,73
Grace,G.184
Grafton,S.T.42,77,126,132,133,
136,316,318
Gray,J.T.101
Graziano,M.S.A.23
Greenwald,A.G.54
zes,J.42,45
Gross,C.G.23
Gruber,O.3,79,81
n,O.70
Halgren,E.87
Hall,J.A.153
Halle,M.209,223
Hall
,P.307,308
Hallet,M.93
Ham,R.19
Happ
,F.G.E.140,141,145
Hari,R.40,43
Harnad,S.357,364
Harris,P.L.275,288
Harris,R.232,233,237
Hassan,F.178
Hauser,M.D.221,250
Haxby,J.V.78
Hayes,G.345,349,350,365
Head,H.102
Heilman,K.M.29
Heimann,M.280
Heine,B.195
Hemenway,K.163
Henley,N.M.153,154
Henson,R.N.A.82
Hepper,P.G.307
Hewitt,G.183,326
Heyes,C.334,344
Hill,J.146
Hitch,G.J.84
Hobson,R.P.288
Hockett,C.178,209
Hofman,M.A.73
Holloway,R.192
Hombert,J.-M.6,250
Hommel,B.116
Hop
eld,J.J.345
Nameindex
Hopper,P.199
Hoshi,E.53
Howard,M.A.5,133
Howes,D.140
Hubley,P.288
Humphrey,N.275
Hurford,J.R.222,250
Hyv
rinen,J.23,49
Iacoboni,M.42,43,45,46,77,111,
115,318,335,347,348
Ingold,T.193
Jacobson,E.132
Jaeger,J.J.214,224
Jakobson,R.209
Jaffe,J.8
James,W.50
Jeannerod,M.49,335,336
Jellema,T.31
Jerison,H.J.66,68
Johanson,D.181
Johnson,M.H.317
Jordan,J.S.4
Jordan,M.I.364,366,367
rgens,U.73
Jusczyk,P.W.306
Kaas,J.H.65,71
Kaplan,F.365
Karten,H.J.65
Kawamura,S.344
Kawato,M.364
Kay,R.F.326
Keenan,J.P.146
Kelso,J.A.S.211
KemlerNelson,D.G.306,308
Kendon,A.151,152,337
Kephart,N.C.102
Kermoian,R.319
Kirchner,J.A.317
Klein,R.181
Knight,C.222,223,250
Knoblich,G.4
Koulomzin,M.8,296,302
Krams,M.347,348
Kubota,K.23
Kubovy,M.163,167
Kugiumutzakis,G.288
Kuhl,P.288,311
Kutas,M.88,93,98
Ladavas,E.301
Lafrance,M.164
Lakoff,R.153
Langacker,R.263
Leakey,R.286
LeDoux,J.338
Leinonen,L.23,49
Lemmo,M.S.29
Leslie,A.254,264
Leuba,G.70
LeVine,R.286
LeVine,S.286
Levy,J.328
Lewin,R.180
Lhermitte,F.133
Li,Ch.6,201,233,237,243,250
Liberman,A.M.207,220,224,264,
Lieberman,P.326
Lindblom,B.306,309
Lindenmayer,A.163
Linkey,H.E.153
Lock,E.D.349
Luppino,G.29
Lyons,J.250
McCune,L.8,9,311,316
318,320
McCune-Nicolich,L.318,320
McGlone,F.5
MacKay,I.R.A.213
MacKay,W.A.49
Nameindex
MacLarnon,A.183,326
MacNeilage,P.F.214,224,326
MacPhail,E.M.65
Maguire,E.A.146
Mandel,D.R.306,307
Maple,T.L.153
Maravita,A.132
Marino,L.69
Mataric,M.345
Matelli,M.14,15,23,46
Matsumura,M.23
Mattingly,I.G.220
Mayr,E.188,324
Matsuzawa,T.344
Mead,G.H.276
Mehler,J.306
Mehrabian,A.153
Meltzoff,A.N.52,101,102,219,
220,223,280
283,288,295,296,
311,344,349
Melzack,R.139
Menn,L.213
Metzinger,T.139
Mitchell,G.153
Moon,C.306
Moore,B.C.344
Moore,C.116,219,220,223
Moore,M.K.52,101,283,344
Morris,D.223
Morris,R.G.78
Morrison,I.9
Mountcastle,V.B.49
Mowrey,R.A.213
Mrzljak,L.73
Muakkassa,K.F.23
Mulac,A.153
Munhall,K.209
Murata,A.15
Myers,J.306
Myowa-Yamakoshi,M.344
Nadel,J.344,349
Newberg,W.88
Newen,A.5
Newmeyer,F.J.327
Nicholas,C.E.145
Nieuwenhuys,R.67
Ninio,A.319
Nishitani,N.43
Northcutt,R.G.65
Nottebohm,F.344
Nyman,G.23,49
Nystrom,L.E.78
Connell,J.F.182
Ogston,W.D.151,152
Ohta,M.280
Keefe,J.291
Oller,D.K.306,309
Oztop,E.348
Palm,G.70
Palmer,S.E.163
Pandya,D.N.15,23,37,49,177
Passingham,R.125
Paulesu,E.78
Pause,M.41,49
Pawley,A.184
Payne,R.197
Pellegrino,G.di101,111,281,310,
Perner,J.140
Perrett,D.I.23,31,37,51,126
Petrides,M.23
Piaget,J.316,344
Pilbeam,D.72,73
Pinker,St.184,237,325
Plotkin,H.C.188,333
Polack,J.B.364
Poranen,A.49
Posner,M.I.303,328
Postle,B.R.78
Pouplier,M.213
Povinelli,D.J.263
Premack,D.30,139
Preuss,T.M.73
Nameindex
Prigogine,I.247
Prinz,W.54,116
Prusinkiewicz,P.163
Querleu,D.307
Quine,W.V.O.208
Raichle,M.E.328
Rakic,P.186
Rall,J.288
Ralph,J.318
Ramachandran,V.251
Reber,A.S.353,355
Redlich,N.A.165
Regan,D.87
Restak,R.M.323
Rizzolatti,G.3,4,14,15,22,23,26,
27,29,37,42,45,49,52,53,77,
93,115,116,123,125,126,132,
136,151,165,202,208,220,221,
223,229,251,253
256,258,273,
274,283,284,289,290,296,315,
316,326,329,333,336,347
353,355,374
Roberts,A.C.71
Roberts,N.5
Rockel,A.J.70
Roland,R.E.93
Romanski,L.M.83
Rommetveit,R.274,275
Roth,G.3,202,232,233
Rothi,L.J.29
Rotondo,J.5,151,155
Rubin,S.91
Ruff,H.319
Rummelhart,D.364,366,367
Sakata,H.29,49
Salenius,S.40
Salmelin,R.40
Sartre,J.316
Saltzman,E.209
Schleicher,A.323
Schnitzler,A.40
Scho
eld,W.N.102
Schumacher,E.H.84
Schumann,J.H.231
Sch
z,A.70
Schwartz,R.G.311
Searle,J.315
Seitz,R.J.49
Seltzer,B.15,23,37,49
Senechal,M.163
Senkfor,A.4,88,91,93,98
Seyfarth,R.M.177,201
Shallice,T.145
Shannon,C.E.165
Shattuck-Hufnagel,S.213
Sherry,S.T.179
Shimizu,T.65
Shipman,P.178
Sinclair,H.319
Smith,M.E.87
Snyder,L.H.49
Sommers,P.van328
Speidel,G.S.344
Spence,K.W.51
Sperber,D.333
Spergel,D.N.327
Stamenov,M.7,230,254,268
Stark,R.E.306,309
Steels,L.365
Stemberger,J.P.311
Stern,D.N.278,287,288
Strafella,A.P.41
Strick,P.L.23
Studdert-Kennedy,M.6,209,214,
215,250
Sugita,Y.10,364
Swanson,R.102
Swettenham,J.258,264
Talairach,J.128
Talmy,L.319,320
Tani,J.10,364,374
Nameindex
Tanji,J.53
Tees,R.C.306,308
Thelen,E.310
Thorne,B.154
Thorpe,W.H.51,52
Tinbergen,N.52
Tincoff,R.306,307
Tipper,S.132
Tomasello,M.19,284,288,344
Tournoux,P.128
Trabant,J.250
Traugott,E.195
Trevarthen,C.274,287,288,344,
Tronick,E.295,296
Turvey,M.T.211,220
Tweed,D.298
Umilt
,M.21,22
Uylings,H.B.M.71
Vallar,G.146
VandenBroeke,M.P.R.224
VanLancker,D.307
VanPetten,C.88,91,93
Velleman,S.L.311
Vendler,Z.266
Vihman,M.8,214,218,305,307,
308,310,311,318,320
Visalberghi,E.19,344
Vogeley,K.5,138,139,145,146
Vogt,C.47
Vogt,O.47
VonBonin,G.47
VonEconomo,C.44,47
VonFersen,L.70
VonGramon,D.Y.82
Walker,A.178
Wapner,S.102
Ward,S.250
Watkins,L.B.29
Weaver,W.165
Weigand,E.7,232,235,240
245,247,253
Weinberg,K.296
Weisfeld,G.E.153
Weitz,S.153
Wellmann,140
Werker,J.F.306,308
Wermter,St.9
Wessels,J.M.I.306,307
Weylman,S.T.145
Whalen,D.H.264
Wheeler,P.182
White,R.180
Whiten,A.19,280,344
Williams,J.E.153
Wohlschl
ger,A.4,102,106,115
Womble,St.9
Wong,W.336
Wood,B.200
Wood,N.310
Woodruff,G.30,139
Woodward,A.L.103
Worsley,K.J.129
Wullimann,M.F.63
Zetterstr
m,R.306,309
Zimmermann,E.74
Subjectindex
accentualpatternofFrench308
action(s)88,93,316
articulatory209
coordination117,118
encoding88
goaldirected353,355,362
imagined93
joint4,116,117,123
mechanismfor14
memoryfor87,91
motor53,54
other-generated123
partiallyhidden22
performed93
representation3,28
self-generated123
structure265
understanding3,13,14,21,50,
51,268,269
watched93
actiongame
culturalunitofthe229,234
dialogic229,239
TheoryofDialogic239
actionprinciple240,242
actionunderstanding50,51,116,
adult-infantrelation(s)284
affectattunement287
affordance282
agent194,257
ofaction2
agranularfrontalcortex
inthemonkeys45
Human45
altercentricframeofreference283
altercentricperception273,278,280
articulatory
lter8,305,310,311
articulatorysuppression3,78,80,
attachmenttheory286
autism280
babbling
canonical309,310
rhythmic311
behavior
communicative189,199
inter-embodied254
symbolic188
behavioralre-enactment282
bifurcation102
bipedalism200
body
image139
schema263
brain
allometry66,68,71
evolution71
growthperiod73
human45,63,77,133
maturationof73
monkey45
tetrapod64
brainsize
absolute65,68
relative66,68
Subjectindex
Broca
scenter/area/region(left
interiorfrontalgyrus)42,73,
74,77,84,125,188,265,
318,326,329
canonicalsyllables305
childdevelopment274
chimpanzee(s)201,284
cognitivereserve190,191
CoherencePrinciple240,245
communication
hominid194
linguistic10
symbolic6,176,177,178,188,
competence-in-performance232,
complementaryact(s)273,282
complementaryprocesses273
consciousness72,138,230,246,315
kindsof71
representational8,318
seatof71
self-138,257
statesof68,70
context366,367
conversation
proto-273
face-to-face273
cortex69,70,72
cerebral69
prefrontal68,71
premotor125,130,132
culturaltransmission333,338
deception201
developmentalclock187
dialogicprinciple240,243,244
dialogue273
grammar7
partners275
efferencycopysignal29
encephalization183,185,188,193
error(s)
contra-ipsi-104
gesture213
ipsi-contra-105
speech213
Event-relatedpotentials(ERPs)4,
87,89,91,93
evolution187,190,286,289
hominid6,73,199
Human84
ofcommunication6
ofthehumanbrain202
oflanguage(
see
Language
evolution)
ofparticulatespeech207
ofrhythmicbehavior336
evolutionarymechanisms185
evolutionaryoriginoflanguage199
evolutionaryperspective230
familiarityeffect308,309
frontallobe45
functionalMagneticResonance
Imaging(fMRI)5,29,43,78,
79,82,84,111,126,127,
132,140,142
gesture(s)19,210,233,234,240,
241,244
dynamic6
emotionalfacial285
phonetic217
goal(s)103,110
common116
detection21,22
social337
grammar194,195,202
graspact283
grasping124,129,130
Subjectindex
grunt317
hand-to-eartask103
hominids175,176,188,221
Homoerectus
192,200,201,222,
284,286
Homohabilis
Homosapiens
200,222
human(s)49,63,65,68,101,102,
111,118,175,198,202,221,
284,301
abilities229
being(s)233,234
anatomicallymodern175,178,
181,200
ideo-motorprinciple104,110
imitation4,9,19,51,52,101,222,
223,311,344,349
creativityin111
facial208,219,220,224
gestural280
goal-directed4,103
goal-directedtheoryof103,
infant278
learningby344,345
neonatal287
roleofobjectsin101
program-level284
vocal208,219,220,224
individual(s)235,245
infant(s)8,219,278,286,296,301,
302,311,315,317,326
integrationallinguistics232
intention275
conscious151
interaction
complementaryfeaturesof
interpersonal283
conversational151
dialogic229,232,233,244
infant-adultchimpanzee285
interpersonal274
preverbal273
intermediatefrontalsulcus81
intersubjectiveattunement287,288
intersubjectivity
primary287
secondary287
language
acquisition328,349,353
crystallizationof192
embodied374,375
emergenceof178,180,182
evolutionof6,8,77,124,175,
faculty249
originof6,175,329
proto-human176
spoken191
written191
larynx317
learning
byaltercentricparticipation
byaltercentricspeechperception
byimitation288
cultural287
face-to-face287
feedback353,360
imitative280
infant274
piecemeal311
robot363,366,368
task356
leftinteriorfrontalgyrus(
see
Broca
region)
linguisticpatient258
Macaquemonkey(s)116,208,281,
283,335
Subjectindex
Subjectindex
primate(s)151,201,221
Non-human13
principalsulcus82
principlesofemotion,ofrhetoric,of
politeness245
principlesofprobability239
processingofsocialinformation116
productionmilestone310
prosodicpattern307
prosody177
ofspeech305
reciprocalgive-and-takesituation
reciprocalmatchingresponses281
reciprocalplay285
reciprocatecaregivers
acts288
recursivefunction195,198
rehearsal3,78,84
relationalwords319
representation14,28,29
forspeech305
lexical309
mental316
segmental310
rhythmicmotoricadvances310
robots10,349
segmentalpatterning305,307
selectiononphonologicalgrounds
311,312
selectiveadvantage284
self
construct138
embodied261
linguistic258
perspective139,140,145
self-grooming296
sentencecompletion277
SharedAttentionMechanism(SAM)
258,264
simulation
children
s289
ofconversationpartners
minds
predictive274
simulation-of-mind274,275
simulationtheory135
singleneuronrecordingtechnique
skillacquisition101
speech
centers73
ontogenyof305
vs.language264
speechact(s)235,236,241,242,
244,271,273,275
speechperception208,288
altercentric288
infant305,310
motortheoryof276
structure
semantic266
syntactic267
superiortemporalsulces(STS)23
supra-modalrepresentationalsystem
symbolgroundingproblem364
symbolicspecies237,246
symmetrybreaking151,154,157,
symmetrybuilding163
symmetryformation151,152,154,
157,164
TheoryofMind(TOM)30,136,
139,140,145
147,258,264,275,
theorytheory136,137
trail-and-errorlearning101
Subjectindex
vocalmotorscheme(s)311
Wernicke
scenter(area)73,74
within-repertoiremotorbehavior
wordtemplates311
WorkingMemory(WM)3,335
capacity77,104
evolutionary-basedmodelof
functionalneuroanatomyof78
modelsof78
multimodal82
phonological80
writing
directionof328
evolutionof328
wordorder327
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