age and culture modulate object processing and … · on physical features and categorical...
TRANSCRIPT
Copyright 2007 Psychonomic Society, Inc. 44
Extensivebehavioralstudiessuggestthatinvisualpro-cessing,collectivistexperiencesbiasEastAsianstoat-tendtocontextualinformation,whereasindividualisticexperiencesbiasWesternerstoprocessobjectspreferen-tially(Chua,Boland,&Nisbett,2005;Nisbett&Masuda,2003;Nisbett&Miyamoto,2005;Nisbett,Peng,Choi,&Norenzayan,2001).Theseeffectsofcultureoncognitivefunctionhavebeendemonstratedacrossmanydomains,includingperceptualprocessing,semanticorganization,memory,reasoning,andneuralfunction.Attheperceptuallevel,Chuaetal.(2005)foundthat,whenviewingcom-plexscenes,EastAsiansmademoresaccadestotheback-groundcontexts,whereasWesternersfixatedfasterandlongeroncentralobjects.Instudiesonsemanticorganiza-tion,EastAsianswerefoundtoassociateimagesofpeopleonthebasisoffunctionalrelationships(suchasgroupingtogetheramotherandherchildbecauseofthematernalrelationship),whereasWesternersbasedtheirassociationsonphysicalfeaturesandcategoricalmembership(suchasgroupingtogetherawomanandamanbecausetheywerebothadults)(Chiu,1972;Ji,Zhang,&Nisbett,2004).Inamemorystudy,MasudaandNisbett(2001)demonstrated
thatEastAsianswerelesslikelytorecognizetargetob-jectsthattheyhadpreviouslyencodediftheobjects’back-groundhadchanged,incontrastwithWesterners,whoseobjectmemorywaslessaffectedbybackgroundchanges.Most recently,Gutchess,Welsh,Boduroglu, andPark(2006),usingstimulisimilartothoseusedbyMasudaandNisbett,observedculturaldifferencesintheventralvisualcortexaswellasinareasassociatedwithsemanticprocessingofobjects.Westernerswhoencodedcomplexpicturescontainingacentralobjectagainstabackgroundshowedmoreengagementofbilateralmiddletemporal,rightsuperiortemporal,andleftsuperiorparietalregions(areasimportantforobjectandsemanticprocessing)thanEastAsians.Incontrast,EastAsians,whenprocessingbackgrounds,showedgreaterengagementofleftoccipi-talandfusiformareas,whichareimplicatedinstructural,perceptualanalyses(Joseph&Gathers,2003).
Inthepresentstudy,weusedtheadaptationparadigmdevelopedbyGohetal.(2004)toinvestigatehowculturemightinteractwithagedifferencesinprocessingobjectsandbackgroundsaswellascontextualbindingofob-jectstobackgrounds.InGohetal.’sstudy,youngadults
Age and culture modulate object processing and object–scene binding in the ventral visual area
Joshua o. Goh University of Illinois at Urbana-Champaign, Urbana, Illinois
Michael W. chee, Jiat choW tan, and Vinod VenkatraManCognitive Neuroscience Laboratory, SingHealth, Singapore
and
andreW hebrank, eric d. leshikar, lucas Jenkins, bradley P. sutton, anGela h. Gutchess, and denise c. Park
University of Illinois at Urbana-Champaign, Urbana, Illinois
Behavioraldifferencesinthevisualprocessingofobjectsandbackgroundsasafunctionofculturalgrouparewelldocumented.Recentneuroimagingevidencealsopointstoculturaldifferencesinneuralactivationpatterns.ComparedwithEastAsians,Westerners’visualprocessingismoreobjectfocused,andtheyactivateneuralstructuresthatreflectthisbiasforobjects.Inarecentadaptationstudy,EastAsianolderadultsshowedanabsenceofanobject-processingareabutnormaladaptationforbackgroundareas.Inthepresentstudy,75youngandoldadults(halfEastAsianandhalfWestern)weretestedinanfMR-adaptationstudytoexaminedifferencesinobjectandbackgroundprocessingaswellasobject–backgroundbinding.Wefoundequivalentbackgroundprocessingintheparahippocampalgyrusinallfourgroups,diminishedbindingprocessesinthehippocampusinelderlyEastAsiansandWesterners,anddiminishedobjectprocessinginelderlyversusyoungadultsinthelateraloccipitalcomplex.Moreover,elderlyEastAsiansshowedsignificantlylessadaptationresponseintheobjectareasthandidelderlyWesterners.Thesefindingsdemonstratethemalleabilityofperceptualprocessesasaresultofdifferencesincohort-specificexperiencesorinculturalexposureovertime.
Cognitive, Affective, & Behavioral Neuroscience2007, 7 (1), 44-52
D. C. Park, [email protected]
Culture, Age, And VisuAl ProCessing 45
werepresentedwithquartetsofpicturesinwhicheitherthecentralobjectorthebackgroundofthepicturevaried(Figure1),andtheattenuationofthebloodoxygenleveldependent(BOLD)signalthatoccurreduponrepetitionofelementsofthepictureswasmeasured(seeGrill-Spector&Malach,2001).Inconditionsinwhichtheobjectwasre-peatedandthebackgroundchanged,theBOLDresponsediminished,relativetowhenboththeobjectandtheback-groundwerechanged, in the lateraloccipitalcomplex
(LOC)inbothhemispheres(Grill-Spector,Kourtzi,&Kanwisher,2001;Malachetal.,1995).Thissuggeststhattheseareaswereengagedforprocessingobjectsandthattheyshowedanadaptedresponseastheobjectsrepeatedacrossthequartets.Similarly,whentheobjectchangedandthebackgroundwasheldconstant,bilateralparahip-pocampalplaceareas(PPA)showedadaptation,suggest-ingthattheseareaswerespecializedforbackgroundpro-cessing(Epstein,Graham,&Downing,2003;Epstein&
OO
ON
NO
NN
A
B
P1 P2 P3 P4
P1 P2 P3 P4
+ + + + +
SD IPI SD IPI SD SDIPI
Quartet
+ + + + +
IQI IQI IQI
Time
Figure 1. Hybrid block/event-related fMRI experiment consisting of quartets of picture stimuli. (A) The four quartet conditions: four repeated objects and scenes (OO: old object, old scene); four novel scenes with a repeated object (ON: old object, new scene); four novel objects within a repeated scene (NO: new object, old scene); and four novel objects with four novel scenes (NN: new object, new scene). (B) Picture stimulus duration (SD) was 1.5 sec, with an interpicture interval (IPI) of 250 msec and mean interquartet interval (IQI) of 9 sec. A fixation cross was shown during the inter-vals when no picture was displayed.
46 goh et Al.
Kanwisher,1998).Finally,Henke,Weber,Kneifel,Wie-ser,andBuck(1999)demonstratedthatbindingareasinbilateralparahippocampalgyrus(separatefromthePPA)andrighthippocampusshowedadaptationwhenbothele-mentswererepeatedbutnotwhenbothwerevaried,sug-gestingthattheseareaswereimportantforcontextuallybindingatargetobjecttoascene;thiswasthefirststudythat showedbinding-relatedprocessingwhensubjectspassivelyviewedscenesratherthanwhentheyactivelyattemptedtobindsceneelements.
Agingisconsistentlyassociatedwithpoorerepisodicmemoryrelatedtobindingdeficitsatencoding(Chalfonte&Johnson,1996;Mitchell,Johnson,Raye,&D’Esposito,2000;Mitchell,Johnson,Raye,Mather,&D’Esposito,2000;Naveh-Benjamin,Hussain,Guez,&Bar-On,2003;Park,Puglisi,&Sovacool,1984;Spencer&Raz,1995).Cheeetal.(2006)testedasampleofolderadultsusingtheGohetal. (2004)paradigmto investigate theneu-ralcorrelatesofthesebindingdeficitscharacteristicofaging.WhencomparedwiththedatafromyoungadultsinGohetal.’sstudy,thedatafromtheCheeetal.studyshowedgreatlyreducedbindingactivityinolderadultsinthemedialtemporalregionsand,surprisingly,thetotalabsenceofobjectprocessingadaptationintheLOC.Incontrast,thebackgroundprocessingareasintheparahip-pocampalregionsshowedsimilarmagnitudesofadapta-tionresponsesforoldandyoungadults.Infollow-upex-periments,objectprocessingadaptationintheLOCwasdemonstratedagainwhenolderadultswereinstructedtoattendtothecentralobjectinthescene,aswellaswhentheolderadultsviewedtheobjectalonewithoutaback-ground.ThusthenonrecruitmentoftheLOCwhentheolderadultspassivelyviewedcomplexpicturesappearstorepresentaperceptualbiasdrivenbyage-relatedchangesinvisualattention(Madden&Langley,2003;Maylor&Lavie,1998;McCarley,Mounts,&Kramer,2004;Milhametal.,2002;Pringle,Irwin,Kramer,&Atchley,2001).
It isnoteworthy that thesubjects inbothGohetal.(2004)andCheeetal.(2006)wereSingaporeansofChi-neseheritage.BothbehavioralandimagingdatasuggestabiasinWesternerstoprocessobjectswithintheback-ground,whichraisesthequestionofwhetherthelossofobjectsensitivityinolderChineseadultsreflectsacultur-allybiasedvisualperceptualprocessinginEastAsiansthatisexacerbatedbychangesinvisualattentionwithage.
Toevaluatetheneuralcorrelatesofculturaldifferencesinperceptualprocessingasafunctionofage,wetested38youngandelderlyWesternsubjects(notofAsiandescent)usingtheGohetal.(2004)paradigmandcontrastedthedatafromthesesubjectswiththedatafromtheyoungandelderlyEastAsianswhoparticipatedintheGohetal.andCheeetal.(2006)studies.Wehypothesizedthatduetoprolongedexperiencewithinanobject-biasedculture,elderlyWesternerswouldshowgreaterengagementofobject-processingareasthandidtheelderlyEastAsiansfromtheCheeetal.study.Specifically,weexpectedtheelderlyWesternerstoshowgreaterobject-processingac-tivityinthelateraloccipitalareasthantheelderlyEastAsians.Wealsopredicted,onthebasisofbehavioralfind-ingsofcross-culturaldifferences,thatolderWesterners
wouldshowlessactivityinthebackgroundprocessingareasthanolderEastAsians.Finally,withregardtotheex-tensiveliteraturedocumentingbehavioralandfunctionaldeficitsinbindingwithageinWesterners,weexpectedbothelderlyEastAsiansandelderlyWesternerstoshowdeficitsinbindingreflectedbyreducedbinding-relatedactivity,relativetoyoungadults,inthehippocampalandparahippocampalregions.
MeTHOD
SubjectsThirty-eightright-handedvolunteers,including19youngWest-
erners(12malesand7femalesranginginagefrom19to27withameanageof21.7years)and19elderlyWesterners(14femalesand5males,ranginginagefrom60to78withameanageof68.1years)fromtheUnitedStates,gaveinformedconsenttoparticipateinthisstudy.SubjectswerescreenedforsignificantillnessesandcontraindicationsforfMRIscanning.DatafromourpreviousstudyofSingaporeansubjectswereincludedforcomparisonacrosscul-tures.Thesesubjectsincluded20youngEastAsians(13femalesand7malesranginginagefrom20to24withameanageof21.3years)and17elderlyEastAsians(11femalesand6malesranginginagefrom60to75withameanageof66.7years).Allsubjectshadnormalvisionorvisioncorrectedtotheacuityof20/30ontheSnellenchart.Subjectsunderwentneuropsychologicaltesting(seeTable1),andtheyalsotooktheWAIS–RComprehensiontest,whichisaculturallyappropriatemeasureofverbalintelligencewithdiffer-entversionsforEastAsiansandWesterners(Gong,1983;Wechsler,1981).Therewerenosignificantdifferencesacrossage[F(1,69)50.003,n.s.]orculture[F(1,69)50.032,n.s.]insubjects’perfor-manceonthistest,indicatingthatsubjectswerecomparableinthismeasureofgeneralintelligence.
Therewere,however,significanteffectsofageinseveraltestsinvolvingspeedofprocessingandworkingmemory;thisresultisconsistentwiththenotionthatagingisassociatedwithslowing(Madden&Langley,2003;Salthouse,1996). Incontrast, therewasnoeffectofcultureonperformanceinthesespeededworkingmemorytests.TherewasaneffectofageontheMini-MentalStateExam(MMSE)[F(1,69)511.34,p,.01],butthemeanscoreswerestillwellwithinthepopulationnormforelderlysubjects,withalloftheoldersubjectsscoring27orgreater(Crum,Anthony,Bas-sett,&Folstein,1993).ItisimportanttonotethattheMMSEscoresdidnotdifferacrossculturesbetweensubjectswithinthesameagegroup.Therewasamaineffectofcultureinthepattern-matchingtask[F(1,69)54.29,p,.05]andaninteractionofagewithcul-tureinthedigitsymboltask[F(1,69)57.00,p,.05](seeHeddenetal.,2002).
StimuliInthisfMRIexperiment,fullcolorpicturesof200objectsand
200placesceneswereused(Figure1A;describedinmoredetailinGohetal.,2004)tocomposepicturestimuliofobjectsplacedwithincongruentbackgroundscenes.Pictureswerepresentedinquartets,whichresultedinfourexperimentalconditions:(1)fourrepeatedob-jectandscenepairs(OO:oldobject,oldscene);(2)repeatedobjectswithinfournovelscenes(ON:oldobject,newscene);(3)fournovelobjectswithinrepeatedscenes(NO:newobject,oldscene);and(4)fournovelobjectandscenepairs(NN:newobject,newscene).Theobjectssubtendedvisualanglesofapproximately0.5º31.0º(minimum)to2.5º35.5º(maximum)fromeachoftheircenters,whilebackgroundscenessubtendedavisualangleofapproximately4.6º36.3ºfromthefixationpoint.
Eachpicturewithinaquartetwaspresentedfor1.5secandsepa-ratedfromthenextpicturebyaninterpictureintervalof250msec(fixation;seeFigure1B).Quartetswerepseudorandomlypresentedsuchthatagivenconditiondidnotoccurmorethanthreetimescon-secutively.Quartetswerealsoseparatedbyinterquartetintervalsof
Culture, Age, And VisuAl ProCessing 47
6,9,or12secwithameanseparationof9sec.ThejitteredintervalswerenecessaryforeffectiveestimationofBOLDresponsestothestimuliinthisrapid,event-relatedfMRIdesign(Dale,1999).
Functionalbrainimageswereacquiredaseachsubjectviewedthepicturestimulioveracourseoffourexperimentalrunsthatlasted348seceach.Eachruncomprised20quartets(5fromeachcon-dition),whichwereprecededandfollowedbyperiodsoffixationthatlasted30sectoallowforbetterestimationofbaselineBOLDresponses.Eachsubjectthereforeviewedatotalof20quartetsofeachexperimentalconditionacrossthefourruns.
Imaging ProtocolfMRIexperimentswereconductedattheCognitiveNeuroscience
LaboratoryinSingaporeandtheUniversityofIllinoisatUrbana-Champaign.Bothsitesusedidentical3.0TAllegrascanners(Sie-mens)andpersonnelworkedclosely together tobecertain thatprotocolsatthetwositeswereidentical.CriticalanalysesoffMRIsignal,noise,andstabilitywereperformed;theyshowedahighreli-abilityacrosssites(Suttonetal.,2007).1Fortheexperimentaltask,116functionalscanswereacquiredineachrunusingagradient-echoEPIsequencewithTRof3sec,FOV19.2319.2cm,anda64364matrix.Thirty-sixobliqueaxialslices,3mmthick(0.3mmgap)andapproximatelyparalleltotheAC–PCline,wereacquired.High-resolutioncoplanarT2anatomicaland3-DMPRAGEanatomicalimageswerealsoacquiredforimagecoregistrationofthefunctionalslicesinto3-Dspace.Stimuliwereprojectedontoascreenatthebackofthemagnetwhileparticipantsviewedthescreenusingamirror.
Image Data AnalysisFunctionalimageswereprocessedusingBrainVoyager20004.9
andBrainVoyagerQX1.3(BrainInnovation,Maastricht)custom-izedwithin-housescripts.GaussiansmoothinginthespatialdomainwasappliedusingaFWHMkernelof8mm.Functionaldatawerethenresampledinto13131mmresolutionpervoxel.Foreachofthefoursubjectgroups,thedatawereanalyzedusingagenerallinearmodel(GLM)comprisingsevenfiniteimpulseresponsepredictorsforeachofthefourexperimentalconditions(OO,ON,NO,andNN).ThuswemodeledtheevolutionoftheBOLDresponsetimecourseover21sec(sevenscans)fromstimulusonset.Subsequentcontrastanalysesofimagingdataconsideredonlythefourthpredictor(9secfromonset)foreachcondition,asthiswasidentifiedasthepeakresponsewithintheestimatedBOLDresponsetimecourseacrossallfourgroupsofsubjects(dataavailableuponrequest).Thisanalysis
resultedinastatisticalmapcontainingparameterestimatesforeachpredictorineveryvoxelofthe3-Dfunctionalbraindata.
Conjunction AnalysisAsinGohetal.(2004),weusedaconjunctionanalysisalongwith
aregionofinterest(ROI)approachtoidentifyandevaluateBOLDresponsesinobject,background,andbindingprocessingregions.Fortheconjunctionanalysis,wefirstcomputedvoxelmapscon-tainingtvaluesofeachcontrast(seebelow)ofparameterestimatesobtainedfromtheGLManalysis.Then,foreachvoxel,wecom-paredtherelevantcontrastsandenteredtheleastsignificanttvalueintoanewstatisticalvoxelmaponlyifallthecontrastsinconsid-erationwerepositive.Usingthisapproach,wedefined(1)object-processingvoxelsasthosethatshowedadaptationresponseswhenobjectswererepeatedregardlessofbackgroundrepetition(OO,NN;OO,NO;ON,NN;ON,NO);(2)background-processingvoxelsasthosethatshowedadaptationresponseswhenbackgroundswererepeatedregardlessofobjectrepetition(OO,NN;OO,ON;NO,NN;NO,ON);and(3)binding-processingvoxelsasthosethatshowedadaptationresponsesonlywhenbothobjectandbackgroundwererepeated,withtheadditionalrequirementthattheadaptationresponsesbegreaterthanthesumofpartialadaptationtoeitherobjectorbackgroundrepetitionalone(OO,NN;OO,ON;OO,NO;[OO,NN],[ON,NN]1[NO,NN]).Notethatbecausewewereconsideringadaptationresponses,thecontrastsaredescribedasattenuationsinsignal,ratherthanincreases(asismoretypical).
Next,theobject,background,andbindingROIsweredefinedascontiguousvoxels(thesmallestROIclusterconsistedof105voxels)thatshowedtherespectivesignificantconjunctionsatastatisticalthresholdofp,.001(uncorrected),exceptwhentheROIwasinthehippocampus,whereareducedthresholdofp,.005wasused(congruentwithproceduresusedbyothers:Eldridge,Knowlton,Furmanski,Bookheimer,&Engel,2000;Ojemannetal.,1997).TheseROIswereidentifiedseparatelyforthedatafromtheEastAsiangroupandthatfromtheWesterngroup.ExaminationofthepeakTalairachcoordinatesofthesefunctionalROIsinyoungandelderlyWesterners(seeTable2)showedthattheywerecomparablewiththoseoftheEastAsians(Cheeetal.,2006).2
Adaptation Magnitude AnalysisTocharacterizetheeffectsofageandcultureonthevisualpro-
cessingofobjects,backgroundscenes,andcontextualbinding,weevaluatedgroupdifferencesinadaptationmagnitudeineach
Table 1 Means (and Standard Deviations) for Demographic Information and Neuropsychological Test Scores
of Young and elderly Westerners and east Asian Subjects
Westerners EastAsians
Young Elderly Young Elderly(12males, (5males, (7males, (7males, FValues7females) 14females) 13females) 10females) Age3
M SD M SD M SD M SD Culture Age Culture
Age(years) 21.73 1.98† 68.10 5.53 21.30 1.11 66.65 4.00 – 2,709.5**0 –Yearsofeducation 15.26 1.38† 15.75 2.97 14.00 1.45 12.50 2.54 17.9** – –Patternmatching 36.13 7.39† 21.15 4.78 39.94 5.50 22.70 4.40 4.29* ,154.94** –Dotcomparison 16.07 1.87† 9.40 2.64 16.61 3.01 7.80 3.24 – ,138.98** –WAIS–RDigit–Symbol 72.93 9.36† 53.10 11.41 82.72 8.66 50.00 11.21 – ,116.46** 7.00**
WAIS–RInformation 22.93 3.47† 20.20 4.03 20.22 4.39 17.70 4.68 6.90* , 7.02** –WAIS–RComprehension 22.93 4.04† 21.05 3.53 21.28 3.72 23.05 4.88 – – –Mini-MentalStateExam 29.60 0.51† 29.00 1.12 29.39 0.92 28.30 1.38 – , 11.34** –WMS–IIIForwardSpatialSpan 10.14 2.19‡ 8.45 1.76 10.22 1.80 8.30 2.11 – , 11.76** –WMS–IIIBackwardSpatialSpan 9.71 1.49‡ 7.45 1.64 9.67 1.68 7.45 1.85 – , 23.15** –WAIS–IIIForwardDigitSpan 11.00 1.63‡ 10.00 2.85 12.39 2.38 10.50 1.85 – , 5.15** –WAIS–IIIBackwardDigitSpan 9.71 1.38‡ 6.70 2.49 9.22 2.44 6.40 1.90 – , 23.25** –
Note—OnlysignificantFvaluesarereportedforthemaineffectsofcultureandage,andtheinteractionbetweenageandculture.TestsarecompleteforallelderlysubjectsandallyoungEastAsiansanddataaremissingforsomeyoungWesterners. †n515. ‡n57. *p,.05. **p,.01.
48 goh et Al.
ofthefunctionalROIsasdefinedabove(seeEpstein,Higgins,&Thompson-Schill,2005,forasimilarapproach).WereasonedthatthedifferenceinBOLDresponseselicitedbytherelevantpairsofconditionsforeachfunctionalROIwouldgiveameasureofthein-tegrityoffunctionofthatregion,withlargeradaptationindicatingbetterfunctionalintegritythanweakorabsentadaptation.Intheobject-processingregions,adaptationmagnitudewasindexedbythedifferencebetweenONandNNresponses;inthebackground-processingregions,bythedifferencebetweenNOandNNresponses;andinthebindingregions,bythedifferencebetweenOOandNNresponses.NotethattheseROIwerealreadyidentifiedasbeinginvolvedinobject,background,andbindingprocessing,respec-tively.Thus,theresultingcontrastvalueswithineachROIreflect
thedegreeofattenuationinresponsetotherelevantrepeatedpicturecomponent(object,background,ortheassociationsbetweenthem)relativetowhennocomponentwasrepeated(NN).TheROImasksthatcharacterizedthelocusofeachfunctionalregionforEastAsiansandWesternerswereappliedtoeachoftherespectiveindividualsubjects.TheindividualmeasuresofmagnitudeofadaptationforeachROIweredeterminedandthedataobtainedweresubsequentlygroupedaccordingtoageandculture.
ReSulTS
We characterized the interaction between age andculturegroupusinganANOVAoftheadaptationmag-nitudedatafromallfourgroupsforeachROI(seeFig-ure2).Intheobject–backgroundbindingregions,weob-servedmaineffectsofageintherightparahippocampalgyrus[F(1,71)59.86,p,.001]andrighthippocampus[F(1,71)54.13,p,.05](Figure2A).Therewasnosig-nificantinteractionofagewithculture,suggestingthatreductionincontextualbindingisanage-relatedchangethatisindependentofculture.
Inthebackground-processingregions,therewerenosignificantdifferencesinadaptationresponseacrossallfourgroupsineithertherightorleftparahippocampalgyrus,suggestingthatbackgroundprocessingwaspre-servedacrossbothageandculture(Figure2B).
Ofparticularinterest,theanalysisofobject-processingregions showed evidence for a reduction of responsewith age in both left [F(1,71)5 11.24, p, .01] andright[F(1,71)510.85,p,.01]lateraloccipitalregions(Figure2C).There was also a marginally significantinteractionofagewithcultureintherightlateraloccipi-talregion[F(1,71)53.22,p,.08].Thisinteractionwaspredicted,andaplannedcomparisonofthiseffectinthisROIrevealedahighlysignificantdifferenceinobject-processingadaptationmagnitudesbetweenyoungandelderlyEastAsians[t(35)53.65,p,.001]butnotinyoungandelderlyWesterners[t(36)51.11,n.s.].More-over,theelderlyEastAsiansshowedsignificantlylowerobject-processingadaptationthandidelderlyWesterners[t(34)52.86,p,.05],whereastherewasnosignificantdifferencebetweenyoungEastAsiansandyoungWest-erners[t(37)50.01,n.s.].ThesamecomparisonsintheleftlateraloccipitalregionyieldedsignificantdifferencesasafunctionofageforbothWesterners[t(36)51.99,p,.05]andEastAsians[t(35)52.76,p,.01],withnoevidenceofaninteraction[F(1,71)50.07,n.s.].Overall,theanalysissuggeststhatobjectprocessinginthelateraloccipitalregionsisattenuatedinelderlyWesternersintheleftbutnottherightLOC,andgreatlyattenuatedinel-derlyEastAsiansinbothhemispheres.
DISCuSSION
Thepresentstudymakesthreemajorpointswithre-specttoneurocognitiveprocessesassociatedwithagingandculture;eachpointaddressesadifferentareaofven-tralvisualcortex.Thepatternofresultsobserveddemon-stratesthat(1)decreasesinneuralbindingprocessesaremanifestedcross-culturallyinelderlyadults;(2)neural
Table 2 Peak Talairach Coordinates of the Object, Background, and
Object–Background Binding ROIs Identified using the Conjunction Analysis for Young and elderly Westerners
BrainRegion
Brodmann’sArea
x
y
z
tValue
YoungSubjects
ObjectProcessing(NN.OOandNN.ONandNO.OOandNO.ON) Rinferioroccipitalgyrus 19 ]30 ]85 ]2 4.96 Linferioroccipitalgyrus 19 ]42 ]73 ]5 4.64 Rfusiformgyrus 37 ]45 ]67 ]2 4.52 Lfusiformgyrus 20 ]39 ]40 ]14 5.16
BackgroundSceneProcessing(NN.OOandNN.NOandON.OOandON.NO) Rparahippocampalgyrus 36 ]18 ]39 ]4 6.79 Lparahippocampalgyrus 36 ]22 ]43 ]5 5.73 Rlingualgyrus 19 ]13 ]68 ]6 4.52 Llingualgyrus 19 ]12 ]71 ]11 4.07 Rposteriorcingulate 29 ]10 ]48 ]8 5.34 Lposteriorcingulate 29 ]11 ]48 ]7 6.47 Lcuneus 18 ]6 ]92 ]9 4.21
ObjectandBackgroundSceneBinding(NN.OOandNO.OOandON.OOand[NN2OO].[NN2ON]1[NN2NO]) Rhippocampus 35 ]33 ]19 ]11 4.00 Rparahippocampalgyrus 37 ]29 ]49 ]11 6.86 Lparahippocampalgyrus 36 ]27 ]27 ]14 4.66 Rlingualgyrus 18 ]12 ]88 ]2 4.17 Lfusiformgyrus 37 ]30 ]70 ]11 3.97 Roccipito-parietalsulcus 19 ]49 ]73 ]12 3.11 Loccipito-parietalsulcus 19 ]36 ]79 ]19 4.61
ElderlySubjects
ObjectProcessing(NN.OOandNN.ONandNO.OOandNO.ON) Linferioroccipitalgyrus 19 ]45 ]74 ]5 4.17 Rfusiformgyrus 19 ]48 ]65 ]2 3.72 Lfusiformgyrus 37 ]39 ]43 ]17 3.85
BackgroundSceneProcessing(NN.OOandNN.NOandON.OOandON.NO) Rparahippocampalgyrus 36 ]24 ]44 ]8 4.15 Rlingualgyrus 19 ]20 ]70 ]11 4.17 Llingualgyrus 19 ]21 ]67 ]11 4.31 Rmiddleoccipitalgyrus 18 ]30 ]80 ]16 4.28 Lmiddleoccipitalgyrus 18 ]36 ]80 ]10 4.64
ObjectandBackgroundSceneBinding(NN.OOandNO.OOandON.OOand[NN2OO].[NN2ON]1[NN2NO]) RfusiformgyrusI 37 ]33 ]37 ]13 4.56 RfusiformgyrusII 19 ]30 ]67 ]8 3.48 Lfusiformgyrus 37 ]36 ]43 ]15 4.32 Roccipito-parietalsulcus 19 ]33 ]73 ]22 3.45 Loccipito-parietalsulcus 19 ]33 ]70 ]28 3.44
Culture, Age, And VisuAl ProCessing 49
processingofbackgroundscenesincomplexpicturesisunaffectedbyageorculture;and(3)object-processingre-gionsdeclinewithage,disproportionatelyinEastAsians.Eachresultisdiscussedinturn.
Binding Mechanisms Across Age and Cultural Group
Thefindingthat,duringpassiveviewingofpictures,bothelderlyEastAsiansandelderlyWesternersshoweddecreasedbindingintherighthippocampusandrightpara-hippocampalgyruscomparedwithyoungadultssuggeststhatexperience(intheformofculturalexposure,forex-ample)mayplayonlyarelativelymodestroleinmoderat-ingthebindingprocessandthatbiologicalmechanismsassociatedwiththeagingprocessmayplayalargerroleindecreasingolderadults’abilitytoengagemedialtemporalstructuresforbindingtothesamedegreethatyoungadults
do.Studieshaveshownthatthehippocampusandentorhi-nalcortexundergoatrophywithageandthatthisatrophycanberelatedtopoorermemoryperformance(Rodrigue&Raz,2004;Rosenetal.,2003).Thelossofneuraltissueavailableforprocessingcontextualbindingmaydiminishthequalityoftherepresentationsofassociativeinforma-tionthatareencodedandsubsequentlyaccessed.Thefind-ingofthisreducedbindinginolderadultsreplicatesthere-sultsofMitchell,Johnson,Raye,Mather,andD’Esposito(2000),inwhichanintentionalencodingtaskwasused.TheresultsofthepresentstudyandthoseinCheeetal.(2006)extendthisfindingtoapassiveviewingtask.
Age, Cultural Group, and the Processing of Background Context
Wefoundlittleevidencethatneuralareasthatarespe-cializedforbackgroundsceneprocessingdifferasafunc-
AR Hippocampus0.50
0.40
0.30
0.20
0.10
0.00
Ad
apta
tio
n M
agn
itu
de
R Parahippocampal Gyrus1.50
1.00
0.50
0.00
Object–BackgroundBinding
BL Parahippocampal Gyrus1.00
0.75
0.50
0.25
0.00
Ad
apta
tio
n M
agn
itu
de
R Parahippocampal Gyrus1.00
0.75
0.50
0.25
0.00
Background SceneProcessing
z = –4
x = 30
CL Lateral Occipital Region1.00
0.80
0.60
0.40
0.20
0.00
Ad
apta
tio
n M
agn
itu
de
R Lateral Occipital Region1.00
0.80
0.60
0.40
0.20
0.00
Object Processing
z = –7
Young Americans Elderly Americans Young East Asians Elderly East Asians
Figure 2. Mean magnitude of adaptation in young east Asians, young Westerners, elderly east Asians, and elderly Westerners. (A) Responses in hippocampal (left panel) and parahippocampal (right panel) binding regions. (B) Re-sponses in left and right parahippocampal areas engaged in background processing. (C) Responses in left and right lateral occipital complex engaged in object processing. Standard error bars are shown.
50 goh et Al.
tionofageorculture,althoughsomefurtherexplorationresultedinsomemarginallysignificanteffectsintheex-pecteddirection.WenotedthatthebackgroundadaptationresponseintheleftparahippocampalgyruswasslightlylowerforelderlyWesternersthanforyoungWesterners[t(36)51.62,p5.06]andyoungEastAsians[t(37)51.37,p5.09],whereastherewasnosignificantcontrastbetweenelderlyEastAsianscomparedwithyoungWest-erners[t(34)51.02,n.s.]andyoungEastAsians[t(35)50.72,n.s.].ThisfindingisconsistentwiththenotionofpreservedbackgroundprocessinginelderlyEastAsians.That the relatively equivalent background processingacrossgroupsinthisstudymayresultfromeffectivepro-cessingofcontextualinformationincomplexscenesbyolderadults.
Thereisconsiderableevidencethatolderadultsbut-tresstheirmemoryforcomplexpicturesbyusingcon-textualinformationwhenitispresentinpicturesthataresufficientlyrichinmeaninganddetail(Park,Puglisi,&Smith,1986;Park,Smith,Morrell,Puglisi,&Dudley,1990;Smith,Park,Cherry,&Berkovsky,1990). It ispossiblethatfurtherexplorationofneuralprocessingofcontextwilldemonstrateagedifferencesinthismecha-nismincasesinwhichthecontextualinformationislessmeaningfulormorepoorlyintegratedwiththetarget.Inparticular,alargenumberoffindingssuggeststhatolderadultsremembercontextualinformationassociatedwithwordsorabstractpictureslesswellthanyoungadultsdo(Parketal.,1990;Smithetal.,1990).Itmayalsobethatpronouncedculturaldifferencesintheneuralprocessingofbackgroundinformationwillsimilarlyemergeundermoredemandingconditions,accentuatingtheexpectedbiasforEastAsianculturalgroupstoshowapreferen-tialprocessingofbackgrounddetailrelativetoWesterngroups.
Age, Culture, and Object ProcessingInthepresentstudy,wefoundevidencefordiminished
objectprocessingintheLOCinolderadultsfrombothcultures,asreportedinitiallybyCheeetal.(2006).Per-hapsthemostimportantfindingfromthepresentstudyisthatelderlyWesternersshowedsignificantlygreaterobject-processingadaptationintheLOCthandidelderlyEastAsians,whoshowedalmostnoadaptationwhatso-ever.ThisfindingprovidesneuroimagingevidenceforculturalbiasesinperceptualprocessingofobjectsandisinagreementwithGutchessetal.(2006),whoreportedgreaterneuralengagementforobject-processingregionsinWesternersthaninEastAsiansinapicturerecognitiontask,albeitforyoungadultsinbothcases.AlthoughthestimuliusedinthisstudyweresimilartothoseusedbyGutchessetal.,thepresentstudydifferedinthatweusedapassiveviewingtaskandanadaptationparadigmasop-posedtoadirectedincidentalencodingtask.
Perhapsbecausethepresentparadigmiscomparativelysubtle,culturaldifferencesbecameapparentonlyinoldersubjectswhohadhadmoreexposuretotheirrespectivecul-turalenvironmentsthantheyoungsubjectshadhad.Thisisaplausibleexplanation,sincethereissubstantialevidencethatneuro-anatomicalchangesinthebrainarerelatedto
thelengthoftimeindividualsspendbeingengagedinspe-cificbehavioralpracticesandsensoryenvironments.InastructuralMRIstudy,posteriorhippocampalvolumewaspositivelycorrelatedwithspatialnavigationexperienceinLondontaxicabdriversversuscontrols(Maguireetal.,2003).Recently,Schneideretal.(2005)alsoshowedthatthevolumeofHeschl’sgyruswaspositivelycorrelatedwithmusicalexperienceinprofessionalmusiciansversusnonmusicians.Likewise,compellingevidenceforfunc-tionalchangesinrelationtoexperienceisclearlyseeninthefunctionalspecializationofbrainregionsforletterandnumberrecognitioninhumansubjects(Polk&Farah,1995;Polketal.,2002;Puce,Allison,Asgari,Gore,&McCarthy,1996).Wepositthatculturallydistinctbehav-iorsandthoughtcanalsobeconstruedasdifferencesinspecificexperiencesthataffectneuralfunction.Analter-nateexplanationtotheculturalexperiencehypothesisisthatAsiansocietyischangingrapidlyandthattheyoungSingaporeans(allofChinesedescent)haveinternalizedWesternvaluestothepointthattheynolongerdisplaybehavioralpatternscharacteristicofAsiancultures.Evenifthisisthecase,theresultsclearlydemonstratesystem-aticdifferencesbetweenEasternandWesternsubjects,withabiastowardmoreprocessingofobjectinforma-tioninelderlyWesterners,afindinginagreementwiththecultural/cognitiveframeworkproposedbyNisbettandMasuda(2003).
ConclusionInsummary,thepresentfindingssuggestthatagealone
cannotexplainthereducedexpressionofobject-processingregionsinelderlyEastAsiansandthatthefunctionalen-gagementofneuralareas,suchastheLOC,canbemodi-fiedthroughexperience.Themostplausiblebasisforthedifference,basedontheburgeoningliteratureinculturalpsychology,appearstobethatvisualexperienceisbiasedbycultural factors.Futureresearch isneeded tomorespecificallydeterminedifferencesinneuralcircuitrythatvaryasafunctionofexperience,withculturaldifferencesplayingaplausibleroleinshapingprocessesthatarebothperceptual,asinthepresentstudyofobjectandscenepro-cessingduringpassiveviewing,aswellasstrategic,asreportedbyGutchessetal.(2006).
AuTHOR NOTe
ThisworkwassupportedbyBMRCGrant04/1/36/19/372toM.W.C.aswellasbyNationalInstituteonAgingGrantsR01AGO15047andR01AGO60625-15toD.C.P. E.D.L.wassupportedbyNationalInstituteofMentalHealthTrainingGrantT32MH19554.Correspondenceconcern-ingthisarticleshouldbeaddressedtoD.C.Park,BeckmanInstitute,405NorthMathews,Urbana,IL61801(e-mail:[email protected]).
ReFeReNCeS
Chalfonte, B. L., & Johnson, M. K. (1996).Featurememoryandbind-inginyoungandolderadults. Memory & Cognition, 24,403-416.
Chee, M. W., Goh, J. O., Venkatraman, V., Tan, J. C., Gutchess, A., Sutton, B., et al. (2006).AgerelatedchangesinobjectprocessingandcontextualbindingrevealedusingfMR-adaptation. Journal of Cognitive Neuroscience, 18,495-507.
Chiu, L. H. (1972).Across-culturalcomparisonofcognitivestylesinChineseandAmericanchildren. International Journal of Psychology, 7,235-242.
Culture, Age, And VisuAl ProCessing 51
Chua, H. F., Boland, J. E., & Nisbett, R. E. (2005).Culturalvaria-tionineyemovementsduringsceneperception. Proceedings of the National Academy of Sciences, 102,12629-12633.
Crum, R. M., Anthony, J. C., Bassett, S. S., & Folstein, M. F. (1993).Population-basednormsfortheMini-MentalStateExamina-tionbyageandeducationallevel. Journal of the American Medical Association, 269,2386-2391.
Dale, A. M. (1999).Optimalexperimentaldesignforevent-relatedfMRI. Human Brain Mapping, 8,109-114.
Eldridge, L. L., Knowlton, B. J., Furmanski, C. S., Bookheimer, S. Y., & Engel, S. A. (2000).Rememberingepisodes:Aselectiveroleforthehippocampusduringretrieval. Nature Neuroscience, 3,1149-1152.
Epstein, R. [A.], Graham, K. S., & Downing, P. E. (2003).Viewpoint-specificscenerepresentations inhumanparahippocampalcortex. Neuron, 37,865-876.
Epstein, R. A., Higgins, J. S., & Thompson-Schill, S. L. (2005).Learningplacesfromviews:Variationinsceneprocessingasafunc-tionofexperienceandnavigationalability. Journal of Cognitive Neuro-science, 17,73-83.
Epstein, R. [A.], & Kanwisher, N. (1998).Acorticalrepresentationofthelocalvisualenvironment. Nature, 392,598-601.
Goh, J. O., Siong, S. C., Park, D., Gutchess, A., Hebrank, A., & Chee, M. W. (2004).Corticalareasinvolvedinobject,background,andobject-backgroundprocessingrevealedwithfunctionalmagneticresonanceadaptation. Journal of Neuroscience, 24,10223-10228.
Gong, Y. (1983).RevisionofWechsler’sAdultIntelligenceScaleinChina.Acta Psychologica Sinica, 15,362-370.
Grill-Spector, K., Kourtzi, Z., & Kanwisher, N. (2001).Thelateraloccipitalcomplexanditsroleinobjectrecognition. Vision Research, 41,1409-1422.
Grill-Spector, K., & Malach, R. (2001).fMR-adaptation:Atoolforstudyingthefunctionalpropertiesofhumancorticalneurons. Acta Psychologica, 107,293-321.
Gutchess, A. H., Welsh, R. C., Boduroglu, A., & Park, D. C. (2006).Culturaldifferencesinneuralfunctionassociatedwithobjectprocess-ing. Cognitive, Affective, & Behavioral Neuroscience, 6, 102-109.
Hedden, T., Park, D. C., Nisbett, R. [E.], Ji, L.-J., Jing, Q., & Jiao, S. (2002).Culturalvariationinverbalversusspatialneuropsychologicalfunctionacrossthelifespan. Neuropsychology, 16,65-73.
Henke, K., Weber, B., Kneifel, S., Wieser, H. G., & Buck, A. (1999).Humanhippocampusassociatesinformationinmemory. Proceedings of the National Academy of Sciences, 96,5884-5889.
Ji, L.-J., Zhang, Z., & Nisbett, R. E. (2004).Isitcultureorisitlan-guage?Examinationoflanguageeffectsincross-culturalresearchoncategorization. Journal of Personality & Social Psychology, 87,57-65.
Joseph, J. E., & Gathers, A. D. (2003).Effectsofstructuralsimilarityonneuralsubstratesforobjectrecognition.Cognitive, Affective, & Behavioral Neuroscience, 3,1-16.
Madden, D. J., & Langley, L. K. (2003).Age-relatedchangesinselec-tiveattentionandperceptualloadduringvisualsearch. Psychology & Aging, 18,54-67.
Maguire, E. A., Spiers, H. J., Good, C. D., Hartley, T., Frackowiak, R. S., & Burgess, N. (2003).Navigationexpertiseandthehumanhippocampus:Astructuralbrainimaginganalysis. Hippocampus, 13,250-259.
Malach, R., Reppas, J. B., Benson, R. R., Kwong, K. K., Jiang, H., Kennedy, W. A., et al. (1995).Object-relatedactivityrevealedbyfunctionalmagneticresonanceimaginginhumanoccipitalcortex. Proceedings of the National Academy of Sciences, 92,8135-8139.
Masuda, T., & Nisbett, R. E. (2001).Attendingholisticallyversusana-lytically:ComparingthecontextsensitivityofJapaneseandAmeri-cans. Journal of Personality & Social Psychology, 81,922-934.
Maylor, E. A., & Lavie, N. (1998).Theinfluenceofperceptualloadonagedifferencesinselectiveattention. Psychology & Aging, 13,563-573.
McCarley, J. S., Mounts, J. R., & Kramer, A. F. (2004).Age-relateddifferencesinlocalizedattentionalinterference. Psychology & Aging, 19,203-210.
McGonigle, D. J., Howseman, A. M., Athwal, B. S., Friston, K. J., Frackowiak, R. S., & Holmes, A. P. (2000).VariabilityinfMRI:Anexaminationofintersessiondifferences.NeuroImage, 11,708-734.
Milham, M. P., Erickson, K. I., Banich, M. T., Kramer, A. F., Webb, A., Wszalek, T., & Cohen, N. J. (2002).Attentionalcontrolintheagingbrain:InsightsfromanfMRIstudyoftheStrooptask. Brain & Cogni-tion, 49,277-296.
Mitchell, K. J., Johnson, M. K., Raye, C. L., & D’Esposito, M. (2000).fMRIevidenceofage-relatedhippocampaldysfunctioninfeaturebindinginworkingmemory. Cognitive Brain Research, 10,197-206.
Mitchell, K. J., Johnson, M. K., Raye, C. L., Mather, M., & D’Esposito, M. (2000).Agingandreflectiveprocessesofworkingmemory:Bindingandtestloaddeficits. Psychology & Aging, 15,527-541.
Naveh-Benjamin, M., Hussain, Z., Guez, J., & Bar-On, M. (2003).Adultagedifferencesinepisodicmemory:Furthersupportforanassociative-deficithypothesis. Journal of Experimental Psychology: Learning, Memory, & Cognition, 29,826-837.
Nisbett, R. E., & Masuda, T. (2003).Cultureandpointofview. Pro-ceedings of the National Academy of Sciences, 100,11163-11170.
Nisbett, R. E., & Miyamoto, Y. (2005).Theinfluenceofculture:Ho-listicversusanalyticperception. Trends in Cognitive Sciences, 9,467-473.
Nisbett, R. E., Peng, K., Choi, I., & Norenzayan, A. (2001).Cultureandsystemsofthought:Holisticversusanalyticcognition. Psycho-logical Review, 108,291-310.
Ojemann, J. G., Akbudak, E., Snyder, A. Z., McKinstry, R. C., Raichle, M. E., & Conturo, T. E. (1997).Anatomiclocalizationandquantitativeanalysisofgradientrefocusedecho-planarfMRIsus-ceptibilityartifacts. NeuroImage, 6,156-167.
Park, D. C., Puglisi, J. T., & Smith, A. D. (1986).Memoryforpictures:Doesanage-relateddeclineexist? Psychology & Aging, 1,11-17.
Park, D. C., Puglisi, J. T., & Sovacool, M. (1984).Picturememoryinolderadults:Effectsofcontextualdetailatencodingandretrieval. Journal of Gerontology, 39,213-215.
Park, D. C., Smith, A. D., Morrell, R. W., Puglisi, J. T., & Dudley, W. N. (1990).Effectsofcontextualintegrationonrecallofpicturesinolderadults.Journal of Gerontology, 45,P52-P58.
Polk, T. A., & Farah, M. J. (1995).Lateexperiencealtersvision. Na-ture, 376,648-649.
Polk, T. A., Stallcup, M., Aguirre, G. K., Alsop, D. C., D’Esposito, M., Detre, J. A., & Farah, M. J. (2002).Neuralspecializationforletterrecognition. Journal of Cognitive Neuroscience, 14,145-159.
Pringle, H. L., Irwin, D. E., Kramer, A. F., & Atchley, P. (2001).Theroleofattentionalbreadthinperceptualchangedetection. Psycho-nomic Bulletin & Review, 8,89-95.
Puce, A., Allison, T., Asgari, M., Gore, J. C., & McCarthy, G. (1996).Differentialsensitivityofhumanvisualcortextofaces,letter-strings,andtextures:Afunctionalmagneticresonanceimagingstudy. Journal of Neuroscience, 16,5205-5215.
Rodrigue, K. M., & Raz, N. (2004).Shrinkageoftheentorhinalcortexoverfiveyearspredictsmemoryperformanceinhealthyadults. Jour-nal of Neuroscience, 24,956-963.
Rosen, A. C., Prull, M. W., Gabrieli, J. D. E., Stoub, T., O’Hara, R., Friedman, L., et al. (2003).Differentialassociationsbetweenento-rhinalandhippocampalvolumesandmemoryperformanceinolderadults. Behavioral Neuroscience, 117,1150-1160.
Salthouse, T. A. (1996).Theprocessing-speedtheoryofadultagedif-ferencesincognition. Psychological Review, 103,403-428.
Schneider, P., Sluming, V., Roberts, N., Scherg, M., Goebel, R., Specht, H. J., et al. (2005).StructuralandfunctionalasymmetryoflateralHeschl’sgyrusreflectspitchperceptionpreference. Nature Neuroscience, 8,1241-1247.
Smith, A. D., Park, D. C., Cherry, K., Berkovsky, K. (1990).Agedifferencesinmemoryforconcreteandabstractpictures.Journal of Gerontology, 45,P205-P209.
Spencer, W. D., & Raz, N. (1995).Differentialeffectsofagingonmemoryforcontentandcontext:Ameta-analysis. Psychology & Aging, 10,527-539.
Sutton, B. P., Goh, J., Hebrank, A., Colcombe, S., Welsh, R., Chee, M., & Park, D. C. (2007).Investigation and validation of in-tersite fMRI studies using same imaging hardware.Manuscriptinpreparation.
Wechsler, D. (1981).WAIS–R manual:Wechsler Adult Intelligence Scale–Revised. SanAntonio,TX:PsychologicalCorporation.
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NOTeS
1.Magnetcomparabilitywasassessedbyhumanandphantomexperi-ments.ComparabilityoftheBOLDresponseswasaddressedbyscan-ningtwosubjects(whowerenotpartofthemainstudy)repeatedlyatbothsitesonamotorandvisualtask(McGonigleetal.,2000).Fifteenrunsofeachtaskateachsitewereperformed.Voxel-by-voxelwholebrainANOVAandICCanalysesusingtask,subject,andscannersitesasfactorswereperformedonthedata;theseanalysesindicatedhighreli-abilityacrosssites.Specifically,taskandsubjectaccountedforamuchgreaterproportionofthevarianceinthedatathandidsite.
2.TheROIwereidentifiedseparatelyacrossgroups,sinceacom-plete,whole-groupGLMrequiredadatasetthatrequiredalargeamount
ofcomputermemory,whichthecurrentsoftwaredoesnotallow.WeidentifiedtheROIbyanalyzingtheEastAsiansandWesternersassepa-rategroups.Wealsoperformedtheanalysisbyconsideringfourgroupsacrossageandculture.Althoughtherewereslightdifferencesinpeakvoxellocations,theresultswerelargelysimilaracrossgroups.Wereporthereonlytheformeranalysis,sinceitallowsustoidentifyandexaminetheBOLDresponsesintheobject-processingROIintheelderlyEastAsians,whichisabsentwhenanalyzingthatdatasetalone.
(ManuscriptsubmittedMarch16,2006;revisionacceptedforpublicationAugust1,2006.)