Principled Neural Partitions Among Aspects of Artifact and Action Concepts The Harvard ...

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Principled Neural Partitions Among Aspects of
Artifact and Action Concepts
Citation
Leshinskaya, Anna. 2015. Principled Neural Partitions Among Aspects of Artifact and Action
Concepts. Doctoral dissertation, Harvard University, Graduate School of Arts & Sciences.
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http://nrs.harvard.edu/urn-3:HUL.InstRepos:17467200
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Principled neural partitions among
aspects of artifact and action concepts
Adissertationpresented
by
AnnaLeshinskaya
to
TheDepartmentofPsychology
Inpartialfulfillmentoftherequirements
forthedegreeof
DoctorofPhilosophy
inthesubjectof
Psychology
HarvardUniversity
Cambridge,Massachusetts
May2015
©2015–AnnaLeshinskaya
allrightsreserved.
Thesisadvisor: ProfessorAlfonsoCaramazza AnnaLeshinskaya
Principled neural partitions among aspects of artifact
and action concepts
Abstract
Theresearchinthisdissertationinvestigatestheneuralimplementationofconcep-
tualknowledge:alevelofrepresentationcharacterizedbybroadgeneralityandinde-
pendencefromallsensoryormotorchannels.Decadesofneuropsychologicalwork
haveestablishedthatconceptualknowledgeisnotrepresentedinasingleneurallocus,
butratheracrossmultipleneuralloci,andthattheselociareoftencontent-selective:
theyareresponsibleforcertainaspectsofconceptualknowledge(e.g.,animals;color)
morethanothers.Onwhatbasisarethesepartitionsdelineated,andwhatdetermines
theircorticallocations?Thepresentworkattemptstounravelsomeoftheseprinciples
byidentifyingneurallocithatrepresentnon-concreteattributesofactionandartifact
concepts,suchasfunctions,intentions,andoutcomes.Asinvestigatedhere,theseas-
pectsofconceptsarenotreducibletoanyparticularsensoryormotorqualities.This
isbeneficialintwoways.First,itpresentsanessentialtestcaseforadominanttheory
ofneuralorganizationofconcepts,whichproposesthatconceptualknowledgeisor-
ganizedbythesensorymodalitythoseconceptsreferto.Second,ithelpsovercomea
methodologicalissueinfunctionalmagneticresonanceimaging:thedifficultyofdis-
sociatingconceptualrepresentationsfromlower-level/sensory-motorassociates.The
focusonnon-concretepropertiesenabledustoovercomethischallenge.Ourfind-
ingsrevealthatdistinctportionsoftheinferiorparietalloberepresentfunctionsof
artifactsandintentions/outcomesofactions.Thissuggeststhatdifferentaspectsof
non-concreteconceptualknowledgearerepresentedinspatiallydistinctneuralareas,
implyingthatneuralorganizingfactorscanbeabstract.Furthermore,thecorticallo-
cationsoftheseaspectsofconceptswereadjacenttocomputationallyrelatedcogni-
tivesystems:outcomesofactionswerenearanareainvolvedintheoryofmind,and
iii
Thesisadvisor: ProfessorAlfonsoCaramazza AnnaLeshinskaya
functionswerenearanareapreviouslyshowntobeinvolvedinphysicalmanipulation
knowledgeoftools.Theseobservationssuggestthatconceptualknowledgeisneurally
localizedaccordingtocomputationalconsiderations:conceptsarelocalizednearto
cognitivecomponentsthatrelyonthoseconceptsfortheiroperation
iv
Contents
1 GeneralIntroduction 1
1.1 Overview:Guidingquestionsaboutthesemanticsystem . . . . . . . . 1
1.2 Evidencefromneuropsychology:Whatarethepartitionsinsemanticmem-
ory? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.3 Themethodologicalchallenge:targetingsemanticrepresentationswith
fMRI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
1.4 Approach:usingnon-concreteattributestostudysemanticorganization 7
1.5 Theimportanceofnon-concreteattributesforcognition . . . . . . . . 8
1.6 Focus:functionsandintentionsastypesofnon-concreteattributes . . 10
1.7 Hypothesisspace:whatcanlocationtellusaboutorganizingprinciples? 12
1.8 Nextsteps:whatisunknownabouttheneurallocationsoffunctionsand
intentions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
1.9 Summaryofthegeneralintroduction. . . . . . . . . . . . . . . . . . 25
2 Non-motoraspectsofactionconcepts 27
2.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
2.2 Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
2.3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
2.4 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
3 Abstractcategoriesoffunctionsinanteriorparietallobe 60
3.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
3.2 Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
3.3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
3.4 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
4 Aspectsofactionandartifactconcepts 93
4.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
4.2 Experiment1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
4.3 Experiment2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
4.4 Experiment3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
v
4.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
5 GeneralDiscussion 148
5.1 Kindsofpartitions . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
5.2 Topographyofpartitions . . . . . . . . . . . . . . . . . . . . . . . . 150
5.3 Implicationsforcognitivearchitecture . . . . . . . . . . . . . . . . . 152
5.4 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
vi
Acknowledgments
Thebiggestpartofthiswork–thelearningprocess–mightnotbereadilyvisiblein
theproductthatemergedfromit.Theseinternaltransitionsinmyownthinkingand
abilities,andtheexperiencesandtransitionsthemselves,arewhatIammostgrateful
for.AndforthisIhavetothankmanypeople.Mostgratitudeisowedtomyadvisor,
AlfonsoCaramazza.Takingmeonasastudent,inwhatcouldonlyhavebeenaleapof
faith,iswhatallowedmetostartthisjourney;andallowedmetobelievethatIcouldbe
capableofit.Hethenletthereignslooseenoughtoallowmetopursuethequestions
Iwantedtoask.Herepeatedlywarnedmeaboutthepossibleperilsofthisenterprise,
butthisonlystokedmydeterminationtodoit:itforcedmetosummoneveryounce
ofdrivethatIcouldharness.Andateveryturn,Ihadtobesureofmyfooting.When-
everIcametoAlfonsoataforkintheroad,he’dsitbackandask,“well,whatisthe
experimentyouwanttodo?”IappreciatedtheownershipAlfonsoallowedmetotake
overmywork.Whetheroutoffaithorfatigue,itiswhathelpedmegetpastthefail-
uresalongtheway.Onecan’tacquirelearnedhelplessnesswhentheprocessisinone’s
control.SoIlearnedperseveranceinstead.Mostlythough,whatIlearnedfromAl-
fonsowashowtothinkabouttheoriesandtheprocessoftestingthem,andhowtoask
questions–particularly,thosethatpokeattheflooronwhichonebuildstheirtheories
andexperiments;thatis,thethefloorofassumptionsandinheritedideas,whichoften
bypassscrutiny.IlearnedequallymuchfromthemembersoftheCognitiveNeuropsy-
chologyLab.Inmyfirstyear,IwasluckyenoughtoworkwithMariusPeelen(now
amemberofthethesiscommittee),whopatientlyguidedmeinmylearningofdata
analysisandexperimentaldesign.Hegavemeacognitivetool-kitforthinkingabout
neuroimagingdataandformakingmethodologicaldecisionsinprincipledways;he
taughtmetoreasonaboutmethods,ratherthanlookforpre-definedsolutions.This
servedmethroughtherestofmygraduatecareer.Iwasalsoluckyenoughto‘growup’
vii
inthelabwithStefanoAnzellotti.Ilearnedanincredibleamountfromhim,andwith
him,duringourtimetogether.HavingapeertowhomIcouldbringmybiggestuncer-
tainties,andwhosereasoningIquicklycametotrustfully,wasawonderfulsourceof
support.IappreciatemostofallhowStefanoalwaysunderstoodevenmymostpoorly
communicatedthoughtsinlabmeetingandtranslatedthemtoeveryonewithgreater
eloquenceandinsightthatIcouldhaveevermustered.Ihavemanylabmembersto
thankfortheirmentorship:JorgeAlmeida,TomMcKeeff,LiubaPapeo,Domenica
Romagno,AngelicaLingnau,NickOosterhofandTaliaKonkle.Theywerealwaysa
sourceofinspirationforme,andhelpedmesolvemanyproblemsalongtheway.In
particular,IappreciatedTalia’sconstantwillingnesstoofferguidance,totalkthrough
complexanalysisproblems,andtoaskexcitingandconstructivequestions.Herun-
abatedenthusiasmandencouragementofeveryoneinthelabwasagreatsourceof
supportandlabspirit.IalsowanttothankNickOosterhofformuchmethodological
guidance—evengoingasfarasoptimizingmycodetoruntentimefasteronaweek-
end;andLiubaPapeoforbeingarolemodel,mentor,hotelroommate(eveninthe
worstofconditions),andfriend.Frommysecondyearonwards,Ialsohadthepleasure
ofSusanCarey’smentorshipandencouragement.Herenthusiasm,andbeliefinthe
importanceofunderstandingconceptualknowledge,wereessentialmypursuitofthis
lineofresearchandanywill-powertogetthroughtimesofdiscouragement–Iwouldgo
toSusanintheworstoftimesandcomeoutfeelingexcitedagain.Herthinkingabout
conceptsisthemostprecisethatIhavefoundinthefield.Learningfromher—inpar-
ticular,throughtheDevelopmentalProseminarcourse–hasbeenahugetreasure.I
alsolearnedanenormousamountfrommypeersandcollaborators.Istillremember
thedaythatJuanManuelContrerassatmedowntolunchtotalkaboutpatternanal-
ysismethods,andwecameoutwithanexperimentideathatwecarriedthroughto
completion(thoughnotinthisthesis).However,throughthatcollaborationwashow
Imadethemostgreatleapsinmytechnicalabilitiesandcreativity,andIthankJuan
Manuelgreatlyforpushingustoalwaysbebetter—cleanerdata,fastercode,betterde-
signs,andbolderideas.Isoenjoyedthejourneywetooktogether.I’malsogratefulto
AlekChakroffforthemanytheoreticalandmethodologicaldiscussionswehadinany
andeverycontext(car,bus,seaplane,kayak,bar,island,beach,hikingtrail),andfor
themuch-neededproof-reading,practice-talkfeedback,andlistening.Ialsowantto
thankAmySkerry,DonalCahill,RomanFeiman,StevenFrankland,JorieKoster-Hale,
viii
MarkThornton,MichaelCohen,MaryamVaziriPashkam,ElinorAmit,andBriaLong
forstimulatingdiscussionsanddebatesoncognitivesciencethatwereaspleasurable
astheywereinsightful.Ialsowanttothankthemanyotherprofessorswhoservedas
mentorsthroughouttheseyears.RebeccaSaxetaughtmehowtothinkcriticallyyet
constructivelyabouttheneuroscienceofhigherlevelcognition,andwasawonderful
sourceoffeedbackformywork.IlearnedahugeamountfromherandNancyKan-
wisher’scoursesonneuroimaging,bothaboutmethodologybutmoreimportantly,
aboutthemeta-questionsinourfield,aboutwhichquestionsareexciting,andinwhich
directionweneedtopushourmethodstobeabletoanswerthem.IalsothankJason
Mitchellforhisguidanceandsupportduringourhighlyproductivecollaboration.
AndIthankJoshGreeneforservingonthecommittee,andofferinghisinvaluable
feedbackandwarmencouragement.Fromthepragmaticside,noneofthisworkwould
havebeenpossiblewithouttheexcellentneuroimagingstaff,researchassistants,and
administratorswhosupportedit.TammyMoranandRossMairrunprobablythe
mostefficientscanningcenteranywhere,andtheydideverythingtheycouldtoensure
thatscanningwasalwayssmoothandsafe.MyfavoritemomentwaswatchingTammy
changetheprojectorlight-bulbinthemiddleofascan,anddespiteneverhavingdone
sobefore,andwithsuchspeedandefficiencythatoursubjectneverhadtoleavethe
scanner.Ialsogreatlybenefitedfromwonderfulresearchassistantsandstaff:PetraPa-
jtas,SilviaUbaldi,ValentinaBrentari,andEvaNaldiniStrausberg,whoalwaysworked
relentlesslytokeepthelabrunning.IalsothankCeliaRaiafororchestratingthegradu-
ateprogramwithefficiencyandgrace.Iworkedwithmanytalentedundergraduatesin
thelab,includingAndrewMessing,JoyceZhou,IsobelGreen,AlanRozet,andGabbie
Guigliano.Itwasapleasuretoworkwithsuchamotivatedandtalentedsetofpeople!
Finally,IthanktheNaturalSciencesandEngineeringCouncilofCanada,theMind,
Brain,andBehaviorInterfacultyInitiative,andtheNormalAndersonFundatHarvard
Universityforfundingthiswork.Onthepersonalside,Iwanttothankmymother,
NinaLeshinskaya,forgivingmeeverything.Ican’tthinkofathingshedidn’tsacri-
ficeformewhenshecould,sothatIcouldhavethebestopportunities.Shetaughtme
tovalueknowledgeandlearning,andalwaysencouragedmetopursuewhatismost
intellectuallyvaluable–notjustwhatismostprofitableormostfashionable.Ioweto
herallofthesuccessandhappinessthatIhavefoundinlife.IalsowanttothankJon
Leckmanforloving,enduring,andlettingmego.Tomyincrediblefriendsandcom-
ix