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4.3.144BrainAsymmetry
Brainasymmetryorcerebralasymmetryreferstoanatomical,physiologicalorbehavioral
differencesbetweenthetwocerebralhemispheres.Thehemispherethatislarger,moreactive,or
superiorinperformanceisdominant.Thescientificstudyofcerebraldominanceisrecentand
datesbacktoPaulBrocasdiscoveryin1865,basedonobservingacquiredlanguagedeficit
(aphasia)followinglefthemisphere(LH)stroke,thattheleftcerebralhemisphereofright
handedpeopleisdominantforlanguage.Untilthemid1940s,thegeneralneurological
consensuswasthattheLHofrighthandersisdominantforallhigherfunctionsandthattheRH
isdominantinlefthanders.Europeanneurologistsandneuropsychologists,suchasHecaen,
Piercy,McFieandZangwill,thennotedthattheLHisdominantforlanguageandplanned
movements(praxis),whereastheRHisdominantforvisuospatialfunctions.Thisledto
replacingthedogmaofexclusiveLHspecializationbythetheoryofcomplementaryhemispheric
specialization(Denes&Pizzamiglio1999,Heilman&Valenstein1993).
Beginninginthe1960s,researchonVogelandBogenssplitbrainpatientsinSperrys
labatCaltechconfirmeddramaticallythemodelofcomplementaryhemisphericspecialization
bycomparingthepositivecompetenciesofthetwohemispheresinsteadofinferringthemfrom
deficit.Moreimportantly,splitbrainresearchshowedthateachdisconnectedcerebral
hemisphereconstitutedacompletecognitivesystemwithitsownsensations,perceptions,
memory,evenlanguage,personalityandconsciousness(Zaideletal.inpress).Thetwo
hemispherescanprocessinformationsimultaneouslyandindependently.Thisisthethesisof
hemisphericindependence(Zaideletal.1990).
Splitbrainresearch,inturn,gaveanimpetustoresearchonasymmetricfunctionsinthe
normalbrain,notablybyKimuraandBrydenatMcGill.EarlierstudiesbyShephardIvoryFranz
in1933oncomplementarynormalhemisphericspecializationforreadingandnonsenseshape
recognitionwereignoredbecauseofhisuntimelydeathandduetohisownantilocalizationist
stanceonmind/brainrelations.Curiously,bothFranzsworkandsubsequentworkinHebbslab
atMcGillintheearly1950sweremotivatedbyissuesoftransferoftraining(acrosstheretinaor
acrossthehands)andwereconfoundedbyissuesofattentionalscanningfromlefttoright.
Today,cerebralasymmetryremainsacornerstoneofhumanneuropsychologyandserves
asamodelsystemforafundamentalquestionincognitiveneuroscience:Howdoseparate
subsystemsofthemind/brainmaintaintheirindependence,ontheonehand,andinteract,onthe
other?
Inthisarticle,thefocusisonasymmetriesinthebrainsofnormalrighthanders.
Asymmetriesobservedafterunilateralbraindamageorinthesplitbrainarediscussedelsewhere
inthisbook.Alsoomittedarediscussionsofindividualdifferencesincerebralasymmetry
associatedwithlefthandednessandwithsex.Finally,thecoverageexcludesabnormal
asymmetriesassociatedwithcongenitaldisorders,suchasdyslexia,andwithpsychopathology,
suchasschizophrenia.
(Thenamesofscientistsassociatedwithspecificresultsaresometimesindicatedin
parentheses.)
1.AnatomicalAsymmetries

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Inaseminalpaperpublishedin1969,GeschwindandLevitskydescribedadramatic
anatomicalasymmetry:Theleftplanumtemporale,underlyingthesylvianfissurenear
Wernickesarea,waslargerthantherightin70%(really80%)ofrighthanders.
Correspondingly,thesylvianfissureislargerontheleftin70%ofrighthandersandthisissaid
tobeassociatedwithLHspecializationforlanguage.Theassociationisimprecise:Lesion
studiessuggestthatabout98%ofrighthandershavelanguagespecializedintheLH.Asimilar
anatomicalasymmetryisalreadypresentinthefetusby31weeksofgestation.These
asymmetriesoccurwhethertheareaisdefinedbyanatomicallandmarksorcytoarchitectonically.
Theyappeartobereducedinlefthanders.Lesion,PETandfMRIstudiessuggestthatplanum
temporaleasymmetryisrelatedtophonologicaldecodingandauditorycomprehension.
GeschwindandGalaburdaalsofoundthatparsopercularis(Brocasarea)islargeronthe
left.Moregrossly,thefrontalregioniswiderandlargerintheRHwhereastheoccipitalregionis
widerandlargerintheLH.
Scheibeldescribedanasymmetryindendriticorganizationintheleftandright
operculum:Higherorderdendriticbranchingwasgreaterontheleft,whereaslowerorder
dendriticbrancheswerelongerontheright.ThiswasattributedtoquickerearlyRHdevelopment
butgreaterLHdevelopmentafterthefirstyearoflife.Thisdynamicepigeneticpatternof
changinghistologicalasymmetriescontrastswiththestablegrossasymmetriesdescribedabove.
2.NeurochemicalAsymmetries
Theneurotransmitterdopamineisinvolvedwithcontrolofmotorbehaviorandhigher
integrativefunctions,andislateralizedtotheLH.Bycontrast,noradrenergicinnervationandthe
neurotransmittersnorepinephrine,aswellasserotonin,areassociatedwithupregulationand
downregulationofautonomicandpsychologicalarousal,respectively,andarelateralizedtothe
RH.ThisissaidtoconferLHdominancetotheactivationregulatorysystem,butRH
dominancetothearousalregulatorysystem(Tucker).Activationreferstoreadinessfor
action,andleadstomotorfunctions.Arousalreferstoorientationtoinputstimuliandleads
toattentionalcontrol.These,inturn,areconjecturedtodeterminehemisphericspecializationfor
perception,action,emotionandcognition.
Therearealsoasymmetriesinneuroendocrineactivity.Thus,thereisasymmetryin
cerebralcontrolofcortisolsecretionduringemotionalsituations(WittlinginDavidson&
Hugdahl1995).Thereisalsoasymmetryinneuroimmunologicactivity.LHactivationseemsto
beimmunoenhancing,whereasRHactivationisimmunosuppressive(Kang,Amassian).
Fluctuatinglevelsofsteroidhormones,suchaschangesofestrogenduringthemenstrual
cycle,mayaffectLHfunctionbymodulatingregionaldopamineandothercatecholamines.The
corpuscallosumisfurtherbelievedtomediatedopaminergicasymmetry.Thus,increased
callosalfunctionmaybeassociatedwithreduceddopaminergic,andconsequentlyfunctional,
asymmetry.Itisthereforepostulatedthatstablesteroidhormones,notablytestosterone,affectthe
organizationofhemisphericasymmetryprenatally,whereasfluctuatinghormones,notably
estrogen,affecttheactivationofneurotransmittertractsandselectivelychangecallosalfunction
(Weekes).
Asymmetriesinbrainreceptors,suchasnicotinicacetylcholinergicreceptors,may
underlieasymmetrichemisphericactivation,predominantlyontheright,duringuptakeof

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recreationaldrugs,includingalcohol,caffeine,andespeciallynicotine.Asymmetrichemispheric
effectshavealsobeenshownforpharmacologicalagents,suchaslithiumbicarbonate,
methylphenidate,andchlorpromazine.Thisisimportantforbiologicalpsychiatry.Forexample,
changesinbehavioralhemisphericasymmetryfollowingfloxetinetreatmentsuccessfully
predictstherapeuticresponseindepression(Bruder).
3.PhysiologicalAsymmetries
3.1.PETandfMRI
Thesetechniquesmeasurehemodynamicresponseandhaveagoodspatialresolution(~
4mm)butpoortemporalresolution(severalseconds).Cabeza&Nyberg(2000)providea
valuablereviewofimagingduringcognition.Ingeneral,thesestudiesdisplaybilateral
activationsthatcontrastwiththeasymmetriceffectoflateralizedlesions.
Attention:Sustainedattention(vigilance)tasksyieldprefrontalandparietalactivation,
preferentiallyintheRH.Spatialcompatibility(Stroop)tasksselectivelyengagetheanterior
cingulateandleftprefrontalcortex.Orientingofvisuospatialattentionselectivelyengagesright
posteriorparietalcortex.Dividedattentiontasksselectivelyactivateleftprefrontalcortex.Thus,
attentioninvolvesabilaterallydistributednetworkwhosecomponentsarelateralizedtooneor
theotherhemisphere.
Motor:Unimanualchoicereactiontimeselectivelyactivatesleftprefrontalintraparietal
areascomparedtosimplereactiontime,consistentwithLHspecializationforaction(Schultera).
Spatialcompatibilitytasksactivatedorsalpremotorandsuperiorparietalareas,predominantlyon
theleft(Iacoboni).Imitationofhandmovements(Iacoboni)andofobjectmanipulation
(Buccino)activatebilateralinferiorfrontalareas,moreontheleft,andbilateralparietalareas,
moreontheright(mirrorneurons).
Perception:Perceptionofobjectsandfacesactivatesthesameventralpathwaybutthere
isgreaterrightlateralizationforfacesthanforobjects.Perceptionofsmellactivatesolfactory
orbitofrontalcortex,especiallyontheright.Rightoccipitotemporal(fusiform)cortexis
selectivelyinvolvedinprocessingspecificvisualforminformation(sameviewofsameobject),
whereastheleftismoreinvolvedinprocessingcategoricalinformation(differentexemplars).
Musicperceptionengagesrightsuperiortemporalcortexformelodies,butleftBrocasarea/
insulaforrhythms.Recognitionofpicturesofselfselectivelyactivatesrightprefrontalcortex.
Imagery:Generationofimagesfromwordsactivatesleftinferiorposteriortemporallobe,
althoughmorecomplexvisualimageryselectivelyengagesrighttemporalcortex.Imageryof
mapsengagesrightsuperioroccipitalcortex,whereasmentalnavigationengagestheleftmiddle
occipitalgyrus.
Language:Spokenwordcomprehensionactivatesbilateralmedialandsuperiortemporal
gyrusbutactivationduringwrittenwordrecognitionisleftlateralized.Surprisingly,left

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prefrontalcortexisactivatedequallyduringwrittenwordrecognitionwithorwithoutspoken
responses.Writtenwordsalsoselectivelyactivateleftposteriortemporalcortex.Thereisa
progressionofhemisphericspecialization:Featureconjunctioninlettersoccursinmedial
extrastriatecortexcontralateraltothestimulusvisualfield(VF).Lettershapesactivateposterior
fusiformgyrusoppositetheprocessinghemisphere,whereasletternamesactivateanteriorand
posteriorfusiformgyrusbilaterally(Pollmann).Letterstringsactivateleftposteriorfusiform
area,whereaslegalwords(visualwordforms)activateleftanteriorfusiformareawithstimuliin
eitherVF(Cohen).Lexicalaccessviathevisualwordformactivatestheposteriortemporallobe
bilaterally,whereasphonologicalrecodingactivatesaleftlateralizednetworkofposterior
(fusiformgyrus,presumablyinvolvedinsublexicalgraphemicprocessing)andanterior(inferior
frontalgyrus,presumablyinvolvedinphonologicalrecoding)structures(Indefrey).Thissupports
thehemisphericdualroutmodel:Bothhemisphereshavelexicalroutes(sightreading),but
onlytheLHhasanonlexical(phonetic)route.
DeafnativeusersofAmericanSignLanguageshownormalLHactivationduring
languageprocessing.Inaddition,theyshowrecruitmentofRHfrontotemporalparietalregions
presumablyforprocessinglocationandmotioninformationpresentinASL.ThisRHrecruitment
maybesubjecttoacriticalperiod(Neville).
Listeningforthemoralofafableselectivelyactivatedfrontalandtemporalregionsofthe
RH.
Memory:Thephonologicalloopcomponentofworkingmemorydistinguishesbetweena
phonologicalstore,associatedwithleftparietalactivation,andarehearsalbuffer,associatedwith
activationinBrocasarea.Workingmemoryengagesprefrontalandparietalregions.Theseare
lateralizedbytypeofmaterial:Brocasareaandleftparietalareasareactivatedfor
verbal/numericaltasks;activationsfornonverbalmaterialtendtobebilateral.
Tulvingshemisphericencoding/retrievalasymmetry(HERA)modelpositsthat(1)the
leftprefrontalcortexisselectivelyinvolvedinsemanticmemoryretrieval,(2)theleftprefrontal
cortexisselectivelyinvolvedwithepisodicmemoryencoding(personallyexperiencedpast
events),and(3)therightprefrontalcortexisselectivelyinvolvedinepisodicmemoryretrieval.
However,recentdatashowmaterialspecificlaterality:LHactivationduringbothencodingand
retrievalofwords,andRHactivationduringbothencodingandretrievalforpatterns,faces,
scenes,ornonverbalsounds.
Differentsemanticdomainsengagedifferentcorticalregions.Forexample,retrievalof
animalinformationengagesleftoccipitalregions,perhapsreflectingprocessingofphysical
features,whereasretrievaloftoolinformationengagesleftprefrontalregions,perhapsreflecting
processingoflinguisticormotorinformation.Similarly,generatingactionwordsengagesleft
temporooccipitalregions,closetomotionperceptionregions.Inotherwords,knowledgeabout
objectattributesappearstobestoredclosetotheregionsinvolvedinperceivingthoseattributes.
Theremaybeadissociationinrightprefrontalcortexbetweendorsalcortex,associated
withfreerecall,andventrolateralcortex,associatedwithcuedrecall(Fletcher).Finally,
autobiographicalretrievalisassociatedwithactivationinarightfrontotemporalnetwork.
Emotions:Activationduringperceptionofdifferentemotionsconvergesintheleft
prefrontalcortex.Dolan(inGazzaniga2000)reportslateralizedrightamygdalaactivationduring

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masked(unconscious)presentationofconditionedfearfulfaces,butleftlateralizedamygdala
activationduringunmasked(conscious)presentation.
3.2.Theelectroencephalogram(EEG)
TheEEGhasagoodtemporalresolutionbutpoorspatialresolutionand,therefore,it
providesarelativelycrudemeasureofhemisphericasymmetry.Itisnowbeginningtobeused
togetherwithimagingtechniqueslikefMRI,thathaveagoodspatialresolutionbutpoor
temporalresolution.EEGmeasurestendtofocusonongoingchangesinthefrequencydomain
whereaseventrelatedpotential(ERP)measuresfocusonstimulusyokedchangesinthevoltage
wavepatternoverasecondorso.DifferentfrequencyrangesoftheEEGhavedifferent
functionalsignificance.EarlyworkbyGalinandOrnsteininthe1970sfocusedontheband
(812cyclespersecondorHz),whosepowerisinverselyrelatedtocognitiveengagement.They
foundalargerrighttoleftpowerratioduringverbal(LH)tasksandasmallerratioduringspatial
(RH)tasks.Theyalsofounddifferentialstableasymmetriesindifferentprofessionals,lawyers
withlargerR/Lratios(moreLHactivity),andartistswithlargerL/Rratios(moreRHactivity).
Morerecently,focushasshiftedtoasymmetriesinhigherfrequencybandsoftheEEG.
Forexample,thegammaband(2535Hz)showedadissociationbetweenwordandnonword
readingorlisteningintheLHbutnotintheRH(Lutzenbeger,Pullvermuller).Alsopromising,
aremeasuresofintrahemisphericandinterhemisphericconnectivityandcoherencefromlarge
(128)electrodearraysthatpermitbetterspatiallocalizationandprovideindicesofnetwork
dynamicsduringcognition(Gevins).
Davidson(inDavidson&Hugdahl1995)usedEEGmeasuresofsuppressionto
characterizeanteriorhemisphericemotionalasymmetriesandindividualdifferencesinemotional
reactivity,moodandtemperament.Approachrelatedpositiveaffectwasassociatedwithleft
anterioractivation,whereaswithdrawalrelatednegativeaffectwasassociatedwithrightanterior
activation.
3.3EventRelatedPotentials(ERP)
SeparatecomponentsoftheERPareassociatedwithsemanticandsyntacticcomponents
ofpostlexicalintegrationinlanguagecomprehension(Brownetal.inGazzaniga2000).Theleft
perisylvianareaiscriticalforsyntacticprocessing,aswellasforaspectsofhigherorder
semanticprocessing.Within200msecafterstimulation,processesrelatedtolexicalmeaningand
integrationemergeintheERPwaveform.(1)Atransientnegativityoverleftanteriorelectrode
sites(leftanteriornegativity,LAN)emerges200500msecafterwordonset.TheLANis
associatedwithinitialparsingandprocessingofsyntacticwordcategoryinformation.(2)A
transientbilateralnegativity(N400)developsbetween200and600msecafterwordonset.The
N400isassociatedwithcontextualeffectsofsemanticprocessing(e.g.,semanticunexpectedness
ofanouninasentenceordiscourse).(3)Atransientbilateralpositivitydevelopsbetween500
and700msec,calledthesyntacticpositiveshift(SPS)ortheP600,andisassociatedwith
syntacticcorrectionoperations,suchasarisefromsyntacticviolations.(4)Aslowpositiveshift
overthefrontofthebraindevelopsacrossthesentence,andisassociatedwithitsoverall
meaning.Theslowshifthasseparateleftandrightlateralizedcomponents.

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AnteriorLHspecializationforgrammarbutnotforsemanticsnormallydevelopsatage
3642monthsandisabsentinlaterlanguageacquisition(Neville).
3.4.PsychophysiologicalAsymmetries
Consistentwiththethesisofhemisphericindependence,brainasymmetryappliesto
cognition,emotion,andregulationofautonomicphysiologicalprocesses.
Neurotransmitter,neuroendocrine,immunomodulatory,cardiovascular,and
electrodermalactivity,skintemperature,andvasomotoractivityareallasymmetric.
Neuralcontrolofcardiovascularfunctionislateralized.Peripheralautonomicchronotopic
controlofcardiacactivity(heartrate)iscontrolledbyrightlateralizedsympatheticand
parasympatheticpathways.Bycontrast,AVconductionandcardiaccontractivityareleft
lateralized.EmotionalstimuliintheLVF,yieldlargerheartrateandbloodpulsevolumechanges
thanstimuliintheRVF(Wittling,Spence).Theseperipheralasymmetriesarenotassociatedwith
correspondingcognitiveasymmetries.Finally,theremaybeanasymmetryinpainsensitivity,
greaterintheRH.
Insum,theLHappearstospecializeformotorfunctionsandintheregulationofthe
bodysdefenseagainstinvadingagents,whereastheRHappearsspecializedforcontrolofvital
functionssupportingsurvivalandcopingwithstress.
4.BehavioralAsymmetries
4.1.ModelsofBehavioralAsymmetry
Behavioralasymmetries,orlateralityeffects,inthenormalbrainaremeasuredby
restrictinginputstimuliorbehavioralresponsesorbothtoonesensoryormotorside.Byfarthe
mostcommonexperimentalparadigmishemifieldtachistoscopywithunimanualresponses.
Stimuliareflashedforlessthan180msectoonevisualhemifield(projectingtotheopposite
hemisphere)topreventinvoluntarysaccadestowardsthestimulus,andresponsechoicesare
signaledbydifferentfingersofonehand,againcontrolledbytheoppositehemisphere.Forthe
sakeofsimplicity,thefollowingmethodologicaldiscussionreferstoahemifieldexperiment,but
similarcommentsapplytoexperimentsinothermodalities.
Splitbrainresearchmotivatestwolimitedcasemodelsofbehaviorallateralityeffectsin
thenormalbrain.Thefirstistheexclusivespecializationmodel,whereonlyonehemisphere,say
theLH,canprocessthestimuli,sothatstimuliintheleftvisualfield(LVF)mustberelayedfrom
theRHtotheLHthroughthecorpuscallosumpriortostimulusprocessing.Thismodeliscalled
thecallosalrelay(CR)model.Thesecondistherelativespecialization,orhemispheric
independence(ofrepresentationandstrategy)model,whereeachhemispherecanprocessthe
stimuli,thoughusuallynotequallywell,andtheinputVFdeterminestheprocessinghemisphere.
Thisiscalledthedirectaccess(DA)model.Considertheexampleofalateralizedlexical
decisiontaskwithorthographicallylegalnonwordsandunimanualword/nonwordresponses.A
callosalrelaymodelpredictsamaineffectofinputVF(RVFadvantageorRFVA)aswellasa
maineffectofanindependentstimulusvariable(say,wordness)butnosignificantinteraction
betweenthetwo(assumingthatwordsandnonwordsareequallycomplexandequally

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susceptibletodelayordegradationincallosalrelay).Bycontrast,amaineffectofVF(RVFA)
andasignificantinteractionofVFandanindependentvariable,saywordness,impliesthatthe
DAmodelholds.
Somebehaviorallateralitytasks,likephoneticperceptionindichoticlisteningto
nonsenseconsonantvowel(CV)syllables,areCR,exclusivelyspecializedintheLH.
Surprisinglymany,likelateralizedlexicaldecision,areDA.Forreasonsstillunclear,itappears
thattheVFofthetarget,ratherthantheresponsehand,determinesthehemisphereincontrolof
processing.EvenwhenDAobtains,theremaybeimplicitinterhemisphericprimingeffectsand
resourceborrowing,leadingtotheconceptofdegreesofhemisphericindependence.Many
tasksprobablyengagebothDAandCRcomponents(Zaidel).
4.2.Methodology
Signaldetection.Forreasonsstillunclear,behaviorallateralityeffectsaremorelikelyto
occurwithaccuracythanwithlatencyasthedependentvariable.Thesignaldetectionmeasureof
bias,,oftenshowsamorenegativebiasintheLVFthanintheRVF.Aspeedaccuracytradeoff
analysisofsomeDAtaskssuggeststhatthetwohemisphereshavesimilarspeedsensitivity
functions,suggestingsimilardecisionstrategies,butdifferentspeedbiasfunctions,implying
hemisphericdifferencesattheresponsechoicestage.Thishighlightstheneedtoapplysignal
detectionmeasurestobehaviorallateralityexperiments.Thatisrarelydone.
Previoustrialeffects.Commonlateralitytestsshowprevioustrialeffectsinvolvingthe
correctness,theVFandthetargetidentityoftheprevioustrial.Theyexhibitaselectiveimplicit
RHerrormonitoring,andsameVForsametargetprimingeffectsacrosstrials,indicatingthe
importanceofcontexteffectsandtheubiquitousnessofhemisphericmomentumeffectsintime.
Bilateralityeffect.Bilateraldisplays,withtargetsinoneVFanddistractorsintheother,
increasehemisphericindependenceandthebehavioralasymmetry(Boles).
Reliabilityandvalidity.Thisisasorelyneglectedtopic.Astandardmeasureofthe
lateralityindexis(LR)/(L+R),whereL=accuracyintheLVF,etc.Voyer(1998)carriedouta
metaanalysisof88experimentsandfoundanaveragetestretestorsplithalfreliabilityofthe
lateralityindexof.68.DichoticlisteninghadgreaterreliabilitiesthanVFstudies,andauditory
verbaltestshadgreaterreliabilitiesthannonverbaltests.Verbaldichotictestshadanaverage
reliabilityof.70.Dichoticlisteningtostopconsonantvowel(CV)syllablepairsshowedthe
highestreliability,.80.Accuracyyieldedhigherreliabilitiesthanlatency.Numberofitems
affectedsplithalfbutnottestretestreliability.Splithalfreliabilitieswerehigherthantestretest
reliabilities.ExtensivetestingwithdichoticCVsyieldedreliabilitiesaround.83andtestingwith
lateralizedlexicaldecisionincludinglateralizeddistractorsyieldedreliabilitiesaround.81
(Zaidel).
ThereliabilityofthedifferencebetweenthetwoVFscoresismaximizedwhentheyare
negativelycorrelatedwitheachother,butwheneachisbasedonalargenumberoftrialsthatare
positivelyintercorrelated.Indeed,thecorrelationbetweenthetwoearscoresindichoticlistening

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variesfrom.1to.72andtheuseofbilateralstimuliinVFstudiesreducesthecorrelation
betweenthetwoVFs(Zaidel).
Voyersestimateofcriterionvalidityfromcrosscorrelationsoflateralitymeasures
betweentestsyieldedalowvalidityof.26.However,theprocedureassumedatrueglobal
measureofhemisphericspecialization.Thisisunlikely.Differentfunctionsevenwithinthesame
domain,suchaslanguage,arelateralizedtodifferentdegrees.Thus,phoneticperceptionappears
tobeexclusivelylateralizedwhereasvisualandespeciallyauditorywordrecognitionaremore
bilaterallyrepresented.Instead,onecanusethesplitbrainasestablishingconstructvalidity.By
administeringthesamebehaviorallateralityteststosplitandnormalbrains,itispossibleto
establishtheunderlyingdegreeoflateralizationoftherelevantfunctions(Zaideletal.1990).
Thisprocedurerevealsthatlateralizedlexicaldecisionmeasuresrelativehemispheric
specialization,butdichoticlisteningtononsenseCVsyllablesisexclusivelyspecializedinthe
LH.Consequently,individualdifferencesintherightearadvantagerevealindividualdifferences
inLHcompetenceorincallosalrelay(fromRHtoLH),orboth.
Lateralityandability.Thereisanothersorelyneglectedaspectofthevalidityof
behaviorallateralitytests:Thedegreetowhichlateralizedperformancepredictstheunderlying
orassociatedabilityinfreevision.ItispossiblethataRVFAreflectsbetterLHperformance
overallandthusbetterscoresonanytestpredominantlycontrolledbytheLH.Thisconceptis
capturedbyLevysindividualdifferencesinstablehemisphericarousalasymmetry.Such
characteristicasymmetryissaidtoaccountforhalfofthevarianceinasymmetryscoresacross
tasks,anditcontributesmoretoasymmetryintaskswithbilateralstimuli.However,no
relationshipwasfoundbetweencharacteristicarousalasymmetryandperformanceonvarious
testsofcognitiveability.
Thequestionremainswhetherperformanceinalateralizedversionofataskpredictsfree
visionperformanceonthesametask?Itdoes,butsurprisinglyweakly.Forexample,therewasan
unexpectedRVFAinatestforrecognizingfacialemotionsandamoderatecorrelation(r=.433)
betweentheRVFAandperformanceonafreevisionversionofthetest(Kee).Inalateralized
lexicaldecisiontask,therewastheexpectedRVFAbutfreevisionmeasuresofvocabulary
correlatedpositivelywithaccuracyineitherVF(LVF:r=.524,RVF:r=.59),andreading
comprehensioncorrelatedonlywithLVFaccuracy(r=.385).Voyeralsoobservedapositive
correlationbetweentheRVFAinalateralizedmentalrotationtestandfreevisionperformance
onatestofspatialability.Sincethecorrelationsarerelativelysmall,itismisleadingtoinfer
hemisphericspecializationofacognitivetestfromtheVFAinalateralizedversionofthesame
orasimilartest.
4.3.DomainsofSpecialization
Consistentwiththetheoryofhemisphericindependence,hemisphericdifferencescan
occuratanylevelofinformationprocessing.
Attention.Hemisphericengagementbylateralizedstimulicreatesanactivationalbiasto
theoppositehalfofspaceandenhancesprocessingofmaterialinthatlocation.Taskdemands,
suchasprocessingfacialemotions,mayselectivelyactivatethespecializedhemisphere.Clinical

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andimagingstudiessuggestthattheRHisspecializedforalerting,arousalandvigilance.Indeed,
testingofcovertorientingofspatialattentionusingthePosnerparadigmdisclosesagreatercost
fordetectingLVFtargetswithinvalidcuesintheRVFthanfordetectingRVFtargetswith
invalidcuesintheLVF.ThereissomesuggestionthattheRVFshowsselectivesensitivityto
objectbasedcueing,whereastheLVFisselectivelysensitivetolocationbasedcueing.
Perception.Olderviewsheldthathemisphericspecializationisdividedbymodality,
material,orstageofprocessing,withtheLHspecializedforauditory,verbaloroutput
processing;theRHforvisual,nonverbalorinputprocessing.Amorerecentviewisintermsof
informationprocessingstyles:theLHisanalyticandtheRHissynthetic(Levy),andarelated
currentviewholdsthattheLHislocalandtheRHisglobal,e.g.,inprocessinghierarchic
patterns(Robertson).AmoreprecisestatementofthelatterviewisthattheLHisspecializedfor
processingrelativelyhighspatialandtemporalfrequencieswhereastheRHisspecializedfor
relativelylowfrequencies,giventhecontext(Sergent,Hellige,Christman,andSeeIvry&
Robertson1998).Theevidenceforhemisphericspecializationinthenormalbrain,intermsof
relativefrequencies,isstillcontroversial,andtheevidencefromnormalsforhemispheric
specializationinglobal/localprocessingisweak.Theglobal/localhemisphericdissociationhas
bettersupportfromneuroimagingstudiesinnormalsandfrombehavioralstudiesinhemisphere
damagedpatients.
TheRHissaidtobespecializedforprocessingfacialemotionsandemotionalprosody.In
spacelocalization,theLHissaidtobespecializedforprocessingcategoricalrelations,suchas
onoff,inside/outside,above/below,ornear/far;theRHissaidtobespecializedforcoordinate,
metricrelations(Kosslyn).TheclaimappealstoselectiveRHinvolvementinmagnocellular
vision(Livingstone),stillacontroversialview.Theclaimreceivedmixedempiricalsupport
(Hellige1993,Sergent).
Imagery.Thecomponentialtheoryofvisualimagery(Kosslyn)distinguishesimage
generation,maintenance,scanning,andtransformation.ThereiscontroversialevidenceforLH
specializationforimagegeneration(Farah)andforRHspecializationinimagerotation
(Sergent).Kosslynbelievesthatcategoricalimagegeneration(bydescriptionofparts)is
specializedintheLH,whereasimagegenerationbystoredmetricmemoriesisspecializedinthe
RH.
Emotions.ThereisaLVFA(RHspecialization)forrecognizingand,perhaps,expressing
facialemotions,aswellasemotionalintonationandemotionalgestures.TheRHmayhavea
selectiveroleinexpressingnegativeemotionsandtheLHinexpressingpositiveones.Heller(in
Banich&Heller1998)distinguishesemotionalvalence(pleasant,unpleasant),controlledby
anteriorcorticalregions,(pleasantmoreontheleft,unpleasantmoreontheright),from
emotionalarousal(high,low)controlledbyrightposteriorregions.Helleralsodistinguishestwo
groupsofindividualswithhightraitanxiety:Thosewithanxiousapprehension(worry),and
increasedLHEEGactivation(suppression),andthosewithanxiousarousal(panic),and
increasedRHactivation.Pettigrewbelievesthatbipolardisorderistheresultofaslow
interhemisphericswitchingmechanismthatbecomesstuckontheleft(mania)orontheright
(depression).

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Language.Findingsfromneurologicalpatientsguidethedesignandinterpretationof
behaviorallateralityexperimentsinnormalsubjects:theanteriorLHisspecializedforphonology
andsyntax,moreposterior(temporal)languageareasintheLHcontrolsemantics,andtheRHis
specializedforpragmatics.
Dichoticlisteningstudiessuggestthatphoneticperception(e.g.,distinguishingstop
consonantvowelsyllablesbyplaceofarticulationandvoicing)isspecializedintheLH(Zaidel).
IvryfindsthattheperceptionofvoiceonsettimeisspecializedintheRH,whereastheperception
ofplaceofarticulationisspecializedintheLH,becausevoicingiscarriedbylowspeech
frequenciesandplaceiscarriedbyhighfrequencies.
WordrepetitionprimingiscaseandfontsensitiveintheLVFbutnotintheRVF
(Marsolek)andthewordsuperiorityeffect(detectingaletterisfasterinawordthanina
nonsensestring)isgreaterintheRVF,suggestingthatwordrecognitionintheLHisquickerand
deeper.However,formatdistortion,e.g.,aLtErNaTiNgcaseorverticalpresentations,selectively
impairswordrecognitionintheRVF,moreforwordsthanfornonwords(Zaidel,Arguin),
suggestingaselectivecontributionofpatternrecognitiontowordrecognitionintheLH.Word
lengthcanaffectwordrecognitioninbothhemispheres,moreintheRHthanintheLH.Both
hemispheresshowcomparablewordfrequencyeffects,buttheLHshowsaselectiveword
regularityeffect(betterrecognitionofwordswithregularthanwithexceptionalspellingto
soundcorrespondence).Theprecedingobservationsareconsistentwiththeviewthatboth
hemispherescanrecognizewordsviaalexicalroute(sightvocabulary)butthattheLHhas
selectiveaccesstoanonlexicalroute(spellingtosound,phonics).However,whentaxed,the
RHdoesshowregularityeffects,suggestingthatithaslatentcompetenceforspellingtosound
conversion.
Whenwefixateduringreading,welookahead,thusselectivelyattendingtotheRVFin
English(readlefttoright),buttotheLVFinArabic(readrighttoleft).Indeed,hemifield
tachistoscopicstudiesshowthatwhilebothtypesoflanguagesarespecializedintheLH,early
visualstagesofwordrecognitionengagetheRHmoreinFarsithaninEnglish(Melamed).
SemanticprimingexperimentssuggestthattheRHactivatesautomaticallymoredistant
semanticrelationshipsandmultiplemeaningsofambiguouswords(Beeman,Chiarello,
Burgess).TheLHmaybemorespecializedforfunctionwords(closedclassitems)thanfor
content(openclassitems).SentencesaremoreeffectiveprimesfortheLHthanfortheRH
(Faust),showingthattheLHisselectivelyspecializedforsyntax.Still,bothhemispheresshow
sensitivitytogrammaticalagreement(Zaidel,Chiarello),suggestingthattheRHdoeshavesome
grammaticalcompetence.Finally,RHactivationofmultipledistantmeaningsmayhelpitmake
predictiveinferencesindiscourse,especiallywhenbreaksincoherenceoccur(Beeman).
Basedonlateralizedprimingexperiments,Chiarelloconcludedthatlexicalorthographic
relationshipsaremoreavailabletotheRH,phonologicalrelationshipsaremoreavailabletothe
LH,andsemanticrelationshipsareavailabletoboth.Automaticlexicalsemanticsismore
availabletotheRH,whereascontrolled,strategiclexicalsemanticsismoreavailabletotheLH.
Olfaction.Unlikevision,hearing,andsomesthesis,theolfactorysystemismainly
uncrossed:leftnostrilLH,rightnostrilRH.Studiesofnormalsubjectssuggestthatthe
olfactorythresholdsofthetwohemispheresarethesame.Odordiscriminationandintensity

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judgmentsshowanadvantageintherightnostril(RH),butodornamingshowsaleftnostril
superiority.However,resultsareinconsistentandbehavioralimagingandlesionstudiesreach
conflictingconclusions.Nonetheless,thereisagreementthatinterhemisphericinteractionis
criticalforolfaction:birhinalpresentation(bothnostrils)increasesperceivedodorintensityand
facilitatesodornaming,althoughitdoesnotaffectdetectionthresholds(Doty).
5.Convergence
Therearesystematicdiscrepanciesbetweenprofilesofhemisphericspecialization
observedinthenormalbrain(reviewedhere,Bradshaw&Nettleton1983,Bryden1982),inthe
splitbrain(e.g.,Zaideletal.inpress),andinhemispheredamagedpatients(Denes&
Pizzamiglio1999,Heilman&Valenstein1993).Lateralityeffectsaretypicallysmallestinthe
formerandlargestinthelatter.Thus,unilaterallesionscanleadtodevastatingcognitivedeficits
notmirroredinthedisconnected,letalonethenormal,hemispheres(Benson&Zaidel1985).For
example,largeperisylvianLHlesionscanleadtoglobalaphasiaandyetthedisconnectedRHis
neitherworddeafnorwordblind.Similarly,rightposteriorlesionscanleadtoneglectoftheleft
halfofspaceanddenialofillness(anosognosia)andyetthedisconnectedLHdoesnotexhibit
neglectordenial.Ingeneral,itseemsthatunilaterallesionsincorporatelossoffunctionin
diseasedtissue,diaschisis,andpathologicalinhibitionofresidualcompetenceinthehealthy
hemisphere.Thoseeffectsareminimizedinthesplitbrainandabsentinthenormalbrain.
Consequently,thedisconnectedhemispheresexhibitgreaterfunctionalcompetencethaninferred
fromlesionstudies.Thenormalhemispheres,inturn,evinceevengreatercompetencethanthe
disconnectedhemispheres,presumablybecausetheycanborrowresourcesfromeachothereven
whentheyprocessinformationindependently.
Hemisphericrepresentationforlanguageisillustrative.Phonologicalandsyntacticdeficit
followexclusivelyLHlesions,semanticdeficitsfollowpredominantlyLHlesionsandpragmatic
deficitsfollowpredominantlyRHlesions.ButthedisconnectedRHhassubstantialauditory
languagecomprehension,includingsomegrammar,amodicumoflexical(butnotsublexical)
reading,andonoccasionevensomespeech(Zaideletal.inpress).Consequently,appealisoften
madetoRHlanguageintryingtoaccountforperplexingsymptomsinaphasicsyndromes,
includingcovertreadinginpurealexia,semanticerrorsindeepdyslexiaorgoodmiminginthe
presenceofmisnamingwrittenwordsinopticaphasia.ThenormalbrainsuggeststhatRH
languagecompetenceisevenricher,includinggraphemephonemecorrespondence.The
challengeforfutureresearchistoreleaselatentRHlanguagecapacitiesforclinicalrecoveryin
aphasia,andtomodulatedegreeofhemisphericspecializationinthenormalbrainforcognitive
optimization.

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HelligeJB,1993HemisphericAsymmetry:WhatsRightWhatsLeft?HarvardUniversityPress,
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ZaidelEClarkeJM&SuyenobuB1990Hemisphericindependence:Aparadigmcasefor
cognitiveneuroscience.InABScheibelandAFWechsler(eds.),NeurobiologyofHigher
CognitiveFunctionpp.297355.GuilfordPress,NY
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Heilman & E Valenstein, (eds.) Clinical Neuropsychology, 4th Edition. New York, Oxford
University Press.
Eran Zaidel
Psychology
UCLA
Los Angeles, CA 90095-1563
USA.
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E-mail: ezaidel@psych.ucla.edu
Telephone: (310) 825-4343
Fax:(310)2063655

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