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2024年3月30日发(作者:)
CerebralCortexOctober2005;15:1602--1608
doi:10.1093/cercor/bhi038
AdvanceAccesspublicationFebruary9,2005
PicturesofAppetizingFoodsActivate
GustatoryCorticesforTasteandReward
Increasingresearchindicatesthatconceptsarerepresentedas
distributedcircuitsofpropertyinformationacrossthebrain’s
modality-specifirentstudyexaminesthedistributed
representationofanimportantbutunder-exploredcategory,foods.
Participantsviewedpicturesofappetizingfoods(alongwithpictures
oflocationsforcomparison)edto
locationpictures,foodpicturesactivatedtherightinsula/operculum
andtheleftorbitofrontalcortex,bothgustatoryprocessingareas.
Foodpicturesalsoactivatedregionsofvisualcortexthatrepresent
ertheseareascontributetoadistributedneural
ydoesthiscircuit
becomeactiveduringthetastingofactualfoods,italsobecomes
processofpattern
completion,foodpicturesactivategustatoryregionsofthecircuit
tentwith
theoriesthatgroundknowledgeinthemodalities,theseinferences
ariseasreenactmentsofmodality-specificprocessing.
Keywords:concepts,fMRI,insula/operculum,knowledge,orbitofrontal
cortex
Introduction
Howareconceptsforeverydayobjectsrepresentedinthe
brain?Basedonaccumulatinglesionandneuroimagingevi-
dence,anobjectconceptisrepresentedasadistributedcircuit
ofpropertyrepresentationsacrossthebrain’smodality-specific
areas(Martin,2001;MartinandChao,2001;Thompson-Schill,
2003).Onencounteringaphysicalobject,relevantmodalities
bjectis
processed,associationareaspartiallycapturepropertyinforma-
tiononthesemodalities,sothatthisinformationcanlaterbe
reactivatedduringconceptualprocessing,whentheobjectis
absent(DamasioandDamasio,1994;SimmonsandBarsalou,
2003).Althoughtheseconceptualreenactmentsshareimport-
antcommonaltieswithmentalimagery,therearealsoimportant
imagerytypicallyresultsfromdeliberate
attemptstoconstructconsciousvividimagesinworking
rast,theperceptualreenactmentsthatunderlie
conceptualprocessingoftenappeartolieoutsideawareness,
resultinginsteadfromrelativelyautomaticandimplicitpro-
aryinterest,thesereenactmentsoccurin
responsestowordsandothersymbols,andplaycentralroles
intherepresentationofconceptualknowledge(Barsalou,1999,
2003a,b;Barsalouetal.,2003a,b).
Thecategoryoftoolsillustratesthedistributionofproperty
representationsacrossmodality-specifi
peopleuseahammer,adistributedsetofbrainareasbecomes
activetorepresentthehammer’sproperties,includingitsvisual
form(ventraloccipitotemporalcortex),thephysicalactions
Óhtsreserved.
Forpermissions,pleasee-mail:sions@
mmons
1
,AlexMartin
2
ou
1
1
DepartmentofPsychology,
2
EmoryUniversity,Atlanta,
GA30322,USAandCognitiveNeuropsychologySection,
LaboratoryofBrainandCognition,NationalInstituteof
MentalHealth,Bethesda,MD,USA
usedtomanipulateit(ventralpremotorcortexandintraparietal
sulcus),andthevisualmotionthatresults(middletemporal
gyrus)(Beauchampetal.,2002;Chaoetal.,1999;Chaoand
Martin,2000;Damasioetal.,2001;Graftonetal.,1997;Handy
etal.,2003;Johnson-Frey,2004;Martinetal.,1995;Peranietal.,
1995).Asjustdescribed,thebrain’sassociationareascapture
thisdistributedsetofmodality-specificstatesforlaterconcep-
equentoccasions,whennohammersare
present,reenactmentsofthesestatesrepresenthammerscon-
ceptually(languagecomprehensionandthought).
Intheexperimentreportedhere,weexploredthedistributed
onstitute
acentralcategoryforhumans,notonlyinperceptionandaction,
butinhighercognition(RossandMurphy,1999).Previous
researchonfoodconceptshasaddressedthevisualpropertiesof
fruitsandvegetables,relativetothevisualpropertiesofother
objectcategories(McRaeandCree,2002).Here,wefocus
insteadonthetastesofhigh-caloric,high-fatprocessedfoods,
suchascheeseburgersandcookies(seeFig.1).Wefocusontaste
propertiesbecausethetastesoffoodsareatleastasimportantas
sonprocessedfoodsbecause
theyarecentraltothemoderndietandbecausetheyare
associatedwithstronggustatoryandappetitiveresponsesthat
underliehowpeopleselectandconsumethem.
Ifadistributedcircuitofpropertyinformationrepresents
foodknowledge,thenviewingafoodpictureshouldnotonly
activatebrainareasthatrepresentvisualpropertiesofthe
picturedfood,butshouldalsoactivatebrainareasthatrepresent
howthefoodislikelytotasteandhowrewardingitwouldbeto
epartofthedistributedcircuitbecomesactiveby
viewingthepicture,theremaindershouldbecomeactivevia
theconceptualinferenceprocessofpatterncompletionacross
hecentralrolethatsuchinferencesplayin
normalfoodselectionandconsumption,itisessentialto
rmore,giventhe
extensivenessofeatingdisorders,obesityandotherfood-
relatedproblems,itisimportanttounderstandhowpeople
generatetasteandrewardinferencestothebroadarrayoffood
representationsavailableinmodernculture.
Wepresentedpicturesoffoodandnon-foodentities(loca-
tionpictures)tosubjectsundergoingevent-relatedfMRIand
predictedthatadistributedcircuitofbrainareaswouldbecome
activetorepresentthevisualandgustatorypropertiesofthe
ingthevisualpropertiesoffoods,alarge
literaturedemonstratesthatventraltemporalregionsunderlie
therepresentationofobjects’visualformproperties(Ishaietal.,
1999,2000).Thus,weexpectedregionsoftheinferiortemporal
andfusiformgyritorespondtothedistinctivevisualproperties
ously,locationpicturesshould
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esoflocation,food,andscrambledimagestimuli.
activateparahippocampalgyrus,giventhatthisregionresponds
tothevisual-spatialpropertiescharacteristicofbuildingsand
landmarks(Aguirreetal.,1998;EpsteinandKanwisher,1998;
Epsteinetal.,1999).
Mostimportantly,thecurrentstudyattemptedtodemonstrate
thatpicturesofvisualobjects,inthiscasefoods,canproduce
istributedaccountofconceptrepresen-
tationiscorrect,thenmultiplemodality-specificregionsshould
onlyshouldvisualareasbecomeactivetorepresentafood’s
uniquevisualproperties,gustatoryareasshouldbecomeactive
opleaccessknowledge
forapicturedfood,aninferenceisproducedabouthowittastes.
Eventhoughpeoplearenotactuallytastingthefood,their
gustatorysystembecomesactivetorepresentthisinference.
Specifically,wepredictedthatsimplyviewingpicturesof
appetizingfoods(relativetolocations)shouldactivatetwo
brainregionsthatcommonlyrespondtoactualtastestimuliin
psychophysicalneuroimagingstudies(Francisetal.,1999;de
Araujoetal.,2003a,b;O’Dohertyetal.,2001b).Thefirstarea,
aregionintheinsula/operculum,isknowntorepresenthow
foodsactuallytaste(Rollsetal.,1988;RollsandScott,2003;
Scottetal.,1986).Thesecondarea,aregioninorbitofrontal
cortex(OFC),isknowntorepresenttherewardvaluesoftastes
(Gottfriedetal.,2003;Rollsetal.,1989).Herewedemonstrate
thatsimplyviewingpicturesofprocessedfoodsactivatesboth
brainregionsinmuchthesamewaythattastestimulantsdoin
psychophysicalstudies.
MaterialsandMethods
Subjects
Nineright-handed,native-English-speakingvolunteersfromtheEmory
Universitycommunityparticipatedinthescanningstudy(sixfemaleand
threemale;agerange,18--45years).Allparticipantscompletedahealth
questionnairepriortoscanningandnoneoftheparticipantsindicated
rdancewithprotocols
prescribedbyEmoryUniversity’sInstitutionalReviewBoard,allpartici-
pantsreadandsignedaninformedconsentdocumentdescribingthe
proceduresandpossiblerisks.
Sixteennative-English-speakingvolunteersfromtheEmorycommu-
nityparticipatedinthestimulusselectionstudy(tenfemaleandsix
male;age19--46years).Noneofthesevolunteersparticipatedinthe
theimagingparticipants,all
participantsreadandsignedaninformedconsentdocumentdescribing
theproceduresandpossiblerisksinaccordancewithprotocols
prescribedbyEmoryUniversity’sInstitutionalReviewBoard.
ExperimentalDesign
Beforebeginningthebrainimagingphaseofthestudy,32typesoffoods
foods(burger,spaghetti,cookie,etc.)inthelistwerechosen
addition,onlyprocessedfoodsthatarerelativelyhighinfatandcalories
house,mall,school,etc.)inthelistwerechosenbecausetheyareall
typesofplacesthatparticipantsinthestudymightvisitfrequently.
Thefoodsandlocationswereequatedforfamiliaritybyhaving
volunteers(noneofwhomparticipatedinthebrainimagingexperi-
ment)providefamiliarityratingsforthe35typesoflocations,and32
sweremadeona1--7scale,with1indicatingthat
atypeoffoodorlocationwascompletelyunfamiliarand7indicating
ntheseratings,15foodand15
locationtypeswereselectedsuchthatnoreliablefamiliaritydifferences
nsixandtenpictures
foreachtypeoffoodandlocationwerethencollected.
Agroupof16participantsviewedall259picturesandratedeachfor
swere
madeona1--7scale,with1indicatingthatapicturewasnotatalltypical
eachtypeoffoodorlocation,thethreemosttypicalpictureswere
selectedforuseintheimagingstudy,thusyieldingatotalof90picture
stimuli(45foods,45locations)hefoodand
locationpicturesdepictednon-uniqueentitiesthatwouldnotbe
y,23location
picturesand22foodpictureswererandomlyselectedtocreate
phase-scrambledimagesthatwerepresentedduringscanningasfiller
items(seeFig.1).
Duringscanning,participantsviewedfood,locationandscrambled
hpicture,participantsusedaresponsepadtoprovide
yes/nojudgmentsastowhetheritwasthesameordifferentasthe
tureswerepresentedinthecenterofthe
persedamongpicturepresentationswere
variable(‘jittered’)interstimulusintervals(mean
=
5.7s,range
=
2--20s)
thatwereincludedtooptimizeestimationoftheevent-relatedfMRI
theseinterstimulusintervals,participantssawafix-
ipantswere
instructedthatwhentheysawthefixationcrosstheyshouldcontinue
attendingtothescreenandprepareforthenextpicturepresentation.
Priortobeginningdatacollection,participantsperformedanabbre-
viatedpracticeruntoinsurethattheyunderstoodthetaskinstructions.
allistsfor
each
run,npicture
presentationsfromeachcategorywerenovelpictures,whileone
picturewasrepeatedtomaintaintheparticipants’attentiontothe
rwords,onelocationpicture
the
threescanningruns,eachsubjectsawthreefoodpicturerepetitionsand
tsweretoldinadvancethat
gthisandgiven
thattherepeatedstimulioccurredinfrequently,thistaskrequires
subjectstopaycloseattentiontoeachpicturepresentationtoinsure
afromtherepetition
trialsineachrunwerenotanalyzedgiventhattheywereonlyincluded
tswere
highlyaccurateatrepetitiondetection(Meancorrect
=
98.8%,SD
=
0.94).Each5min8srunconsistedof4min48softherepetition
detectiontask,followedbyanadditional20srestperiod.
CerebralCortexOctober2005,V15N10
1603
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