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CorticalNeuralSystemsandCognitiveOperations:ASelectiveAnatomicalReviewKangCheng1andKathleenS.Rockland2

1LaboratoryforCognitiveBrainMapping

2LaboratoryforCorticalOrganizationandSystematics

RIKENBrainScienceInstitute

2-1Hirosawa,Wako-shi351-0198,Japan

e-mail:kcheng@mailman.riken.go.jpStructureandFunctionSynapsesNeuronsModulesAreasConnectionsBrainContentsPierrePaulBroca(1824-1880)BrainofMonsieurLeborgne“Tan”BrocaandBroca’sAreaBroca’sareaWernicke’sareaWilderPenfield(1891-1976)PenfieldandElectricalStimulation

(TheMontrealProcedure)Mapsofexperientialresponses(PenfieldandPerot,1963)Bastian,1880BielschowskyandBrodmann,1905NeuronGolgi,1883RamonyCajal,1954SantiagoRamonyCajal(1852-1934)CamilloGolgi(1843-1926)ReticularTheoryNeuronDoctrineSirCharlesScottSherrington(1857-1952)SynapsesPetersandJones,1984Sheng,2001spineboutonNeuroanatomicalMethodsNervecellGolgistainIntracellularinjectionsoftracers(biocytin,neuronbiotin,horseradishperoxidase(HRP))EMforfinecellularandsubcellulardetailsNeuronalpopulationsNisslstainNewerimmunocytochemistrymethods,e.g.,antibodiesAgainsttransmitters(glutamate;GABA;dopamine)Againstcalcium-bindingproteins(calbindin;parvalbumin;calretinin)Againstsynapticcomponents(synaptophysin;synapsin)Againstreceptors(glutamate;GABA)Histocehmistrytechniques,e.g.,Acetylcholinesterasereactions(neuronspositiveforneurotransmitteracetylcholine)Cytochromeoxidase(highlevelofmetabolicactivity)ConnectivityRelativelywellunderstoodMoredetailsinthistalkMicrocircuitryDifficult/stillunderinvestigationFunctionalnetworks2-deoxyglucose(2-DG)autoradiographyEarlyimmediategenes(replacing2-DG)(c-fos;jun;zipf268)NueroanatomyinhumansDegenerationfibersinpostmortembrainsNoninvasiveneuroimagingtechniques(PET,fMRI)ModerntechniquesLaserscanningconfocalmicroscope(LSCM)2-photonlaserscanningmicroscopeInsituhybridizationhistochemistryGenemanipulation(“reportergenes”)Virusvectors(adenovirus;pseudorabies;herpessimplexvirus,etc)NisslStainIIIIIIIVAIVBIVCaVVIIVCb1mmV1V2V1V2IIIIIIIVVVIKorbinianBrodmann(1868-1918)Brodmann,1907BrodmannandBrodmann’sAreasAClassicAxon

TerminalsApicalDendritesBasalDendrites

Connectivity•degeneratedaxons(lesion)•[3H]prolin•WGA-HRP•biocytin•kidneybeanlectin•biotinylateddextranamino(BDA)•fluoro-ruby•PHA-LAnterogradeTracersCellbodytoterminations•horseradishperoxidase(HRP)•fastblue(fluorescentdye)•diamidinoyellow(fluorescentdye)•choleratoxin•PHA-L(rare,Golgi-like)RetrogradeTracersTerminationstocellbodyFeedforward(V1→V2)200µmFeedback(MT→V1)300µm50µm50µmCallosal100µmRocklandandPandya,1979

LaminarOriginsofCorticalProjectionsJones,1984

LaminarOriginsofEfferentProjectionsIntracorticalModulesJ.Neurophysiol.20:408-434,1957Favorov,1991ColumnarOrganizationinSomatosensoryCortexNote:ColumnarorganizationinmonkeysomatosensorycortexwasfirstreportedbyT.P.S.PowellandV.B.MountcastleinBull.JohnsHopkinsHosp.105;133-162,1959.J.Neurophysiol.20:408-434,1957Blasdel,1992OcularDominanceColumnsandOrientationColumnsinV1Tootelletal.,1988HubelandWiesel,1977Kennedyetal.,1976Tritiated([H3])prolinlabeling2-[14C]deoxyglucosemethodAutoradiographyandOcularDominanceColumnsSzentagothai,1978VerticallyDirectedLocalConnectionsPetersandSethares,1996AColumnContainsManyMinicolumnsRakic,1995ColumnarOrganizationandCorticogenesismigratingneuronsradialglialventricularzonecorticalplateAlbright,1984DirectionSelectiveColumnsinAreaMTAnatomicallynotyetshowninmacaquemonkey!Tanaka,1996Fujitaetal.,1992ColumnsinAreaTEWangetal.,1996Tanaka,1997ColumnsinAreaTESaleemetal.,1993ColumnarAfferent&EfferentConnectionsinAreaTEChengetal.,199750µmAmy=amygdalaPut=putamenCd=caudateCl=claustrumH=hippocampusrh=rhinalsulcusLV=lateralventricle

ColumnarAfferent&EfferentConnectionsinAreaTEColumnarAfferent&EfferentConnectionsinAreaTEChengetal.,1997Hortonetal.,1999[H3]prolin-IVcCOII+IIICytochromeOxidaseBlobsinV1&StripesinV2

SincichandHorton,20022mmCytochromeOxidaseBlobsinV1&StripesinV2

Gilbert,1992sameeyeblobs1mmHorizontalConnectionMalachetal.,1993200mmHorizontalConnectionDefiningACorticalArea1.Cytoarchitectonics2.Topographicorganization3.Anatomicalconnections4.Physiologicalproperties&Neuroimaging(inhuman)FellemanandVanEssen,1991ParallelandHierarchicalOrganization

VanEssenandGallant,1994Advice:DoNotAcceptEasily!ParallelandHierarchicalOrganization

Visionforaction?etcMeriganandMaunsell,Howparallelaretheprimatevisualpathways?Annu.Rev.Neurosci.16,369,1993.Lennie,Singleunitsandvisualcorticalorganization,Perception,27,889,1998.IfIStillHaveTime...Zekietal.,1991Adirectdemonstrationoffunctionalspecializationinhumanvisualcortex.J.Neurosci.11,641-64

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