cerebral cortex 1
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1.Forebrain1.Forebrain : Telencephalon: Telencephalon1
NEUROLOGY
Dr.G.Bhanu Prakash
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Forebrain (prosencephalon) =Forebrain (prosencephalon) =??? + ?????? + ???
TelencephalonTelencephalon Largest Brain divisionLargest Brain division
Major Components:Major Components:
??? structures??? structures of the Telencephalonof the Telencephalon
e.g. Basal Forebrain, Amygdala, Basal Ganglia, Striatume.g. Basal Forebrain, Amygdala, Basal Ganglia, Striatum
?????? trackstracks
e.g.e.g. corona radiata, internal capsulecorona radiata, internal capsule
????????
e.g.e.g. primary motor cortex, Brocas areaprimary motor cortex, Brocas area2
Forebrain 1: TelencephalonForebrain 1: Telencephalon
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Forebrain (prosencephalon) =Forebrain (prosencephalon) =telencephalon + diencephalontelencephalon + diencephalonTelencephalonTelencephalon
Largest Brain divisionLargest Brain division Major Components:Major Components:
Subcortical structuresSubcortical structures of the Telencephalonof the Telencephalon
e.g. Basal Forebrain, Amygdala, Basal Ganglia, Striatume.g. Basal Forebrain, Amygdala, Basal Ganglia, Striatum
White matterWhite matter trackstracks e.g. corona radiata, internal capsulee.g. corona radiata, internal capsule
Cerebral CortexCerebral Cortex
e.g. primary motor cortex, Brocas areae.g. primary motor cortex, Brocas area
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Forebrain 1: TelencephalonForebrain 1: Telencephalon
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OverviewOverview
Cortical organization (lamina)Cortical organization (lamina)
Cell types/ Layers/ ConnectionsCell types/ Layers/ Connections
vertical organizationvertical organization Types of cortexTypes of cortex
Brodmanns AreasBrodmanns Areas
Cytoarchitecture and relation to functionCytoarchitecture and relation to function
Key Brodmanns AreasKey Brodmanns Areas
Major White Matter tracts of the Cerebral cortexMajor White Matter tracts of the Cerebral cortex
Internal CapsuleInternal Capsule
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Descriptive AnatomyDescriptive Anatomy
to function (overview)to function (overview) Histological staining (Cell Bodies)Histological staining (Cell Bodies)
DiscoveredDiscovered ?????? arrangement of the cortexarrangement of the cortex
?????? are defined byare defined by cell type, size andcell type, size anddensitydensity
Broadmans areasBroadmans areas Regions of the cortex haveRegions of the cortex have unique ???unique ???
GaveGave numbersnumbers to each of theseto each of these ??????(Broadmans areas)(Broadmans areas)
Broadmans areas and cortical functionBroadmans areas and cortical function Laminar layers haveLaminar layers have ?????? correlatescorrelates
Size of these layers are related to theSize of these layers are related to the ??????
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Descriptive AnatomyDescriptive Anatomy
to function (overview)to function (overview) Histological staining (Cell Bodies)Histological staining (Cell Bodies)
DiscoveredDiscovered laminarlaminar arrangement of thearrangement of thecortexcortex
LaminaLamina are defined by cell type, size andare defined by cell type, size and
densitydensity Broadmans areasBroadmans areas
Regions of the cortex haveRegions of the cortex have unique laminarunique laminarpatternspatterns
GaveGave numbersnumbers to each of theseto each of these laminarlaminar
patternspatterns (Broadmans areas)(Broadmans areas) Broadmans areas and cortical functionBroadmans areas and cortical function
Laminar layers haveLaminar layers have functionalfunctional correlatescorrelates
Size of these layers are related to theSize of these layers are related to theareas functionareas function
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Cerebral CortexCerebral Cortex the Basic Insthe Basic Ins
andO
utsandO
uts Afferent fibers enterAfferent fibers enter ??????
?????? process the afferentprocess the afferent
informationinformation
?????? fibers convey thefibers convey theprocessed information toprocessed information toanother siteanother site
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Cerebral CortexCerebral Cortex the Basic Insthe Basic Ins
andO
utsandO
uts Afferent fibers enterAfferent fibers enter
cortexcortex
Local circuitsLocal circuits process theprocess theafferent informationafferent information
EfferentEfferent fibers convey thefibers convey theprocessed information toprocessed information to
another siteanother site
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Interneurons???
output
Cell Types and laminar locationsCell Types and laminar locations: Local: LocalCircuit (interneurons)Circuit (interneurons) Local circuit neuronsLocal circuit neurons AKAAKA
interneurons, intrinsic neuronsinterneurons, intrinsic neurons
?????? do not leave the immediate region ofdo not leave the immediate region ofcell body, can cross into different laminarcell body, can cross into different laminarlayerslayers
Inhibitory, containInhibitory, contain ??? (1 exception)??? (1 exception)
FunctionFunction ?????? FormulateFormulate rhythms, coordinate outputsrhythms, coordinate outputs
InterneuronsInterneurons classified byclassified by ??????
?????? -- Laver IVLaver IV
?????? (have spines)(have spines) Receptive field IIReceptive field II IVIV
Excitatory (Glutamate)Excitatory (Glutamate) ??????(lack spines)(lack spines)
Receptive field mainly IVReceptive field mainly IV
?????? Layers III And VLayers III And V
Receptive field (all layers)Receptive field (all layers)
?????? Layer IIILayer III
Receptive field mostly layer IIIReceptive field mostly layer III
Aspiny Spiny
??????
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InterneuronsStellate
(granule) cells
output
Cell Types and laminar locationsCell Types and laminar locations: Local: LocalCircuit (interneurons)Circuit (interneurons) Local circuit neuronsLocal circuit neurons AKAAKA
interneurons, intrinsic neuronsinterneurons, intrinsic neurons AxonsAxons do not leave the immediate regiondo not leave the immediate region
of cell body, can cross into differentof cell body, can cross into differentlaminar layerslaminar layers
Inhibitory, containInhibitory, contain GABA (1 exception)GABA (1 exception)
FunctionFunction regulate inputs to and fromregulate inputs to and from
pyramidal cells (cortical output cells)pyramidal cells (cortical output cells) FormulateFormulate rhythms, coordinate outputsrhythms, coordinate outputs
InterneuronsInterneurons classified byclassified by morphologymorphology Stellate (granule)Stellate (granule) -- Laver IVLaver IV
SpinySpiny (have spines)(have spines) Receptive field IIReceptive field II IVIV
Excitatory (Glutamate)Excitatory (Glutamate)
AspinyAspiny (lack spines)(lack spines) Receptive field mainly IVReceptive field mainly IV
BasketBasket Layers III And VLayers III And V Receptive field (all layers)Receptive field (all layers)
ChandelierChandelier Layer IIILayer III Receptive field mostly layer IIIReceptive field mostly layer III
Aspiny Spiny
Basket
cellsChandelier
cells
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inputs
output
Pyramidal cells: most
???
???
???
inputs
output
inputs
output
Cell Types and laminar locations:Cell Types and laminar locations: PyramidalPyramidal
(output neurons)(output neurons)
Cortico-
cortical
terminas
Pyramidal cells size by lociPyramidal cells size by loci Layer VILayer VI ??????; layer II&III; layer II&III ??????; layer V; layer V ??????
Layer VI pyramidal cellsLayer VI pyramidal cells
InputsInputs --DendritesDendrites span layersspan layers ??????
O
utputsO
utputs ?????? Layer II & III pyramidal cellsLayer II & III pyramidal cells
InputsInputs DendritesDendrites span layersspan layers ??????
OutputsOutputs ??????
Layer V pyramidal cellsLayer V pyramidal cells
?????? (largest pyramidal neurons)(largest pyramidal neurons) primary ???primary ???(upper motor neurons)(upper motor neurons)
InputsInputs ??????
OutputsOutputs ??????,, ???, ???, ??????, ???, ???
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inputs
output
Pyramidal cells: most
small
Medium
large
inputs
output
inputs
output
Cell Types and laminar locations:Cell Types and laminar locations: PyramidalPyramidal
(output neurons)(output neurons)
Cortico-
cortical
terminas
Pyramidal cells size by lociPyramidal cells size by loci Layer VILayer VI smallsmall; layer II&III; layer II&III small tosmall to
mediummedium; layer V; layer V LargeLarge
Layer VI pyramidal cellsLayer VI pyramidal cells
InputsInputs --DendritesDendrites span layersspan layers IVIV VIVI
OutputsOutputs Thalamus, cortex, claustrumThalamus, cortex, claustrum Layer II & III pyramidal cellsLayer II & III pyramidal cells
InputsInputs DendritesDendrites span layersspan layers II IVIV
OutputsOutputs Cortex opposite hemisphereCortex opposite hemisphere(callosal)(callosal)
Layer V pyramidal cellsLayer V pyramidal cells Betz cellsBetz cells (largest pyramidal neurons)(largest pyramidal neurons)
primary motor cortexprimary motor cortex (upper motor neurons)(upper motor neurons)
InputsInputs all cortical layersall cortical layers
OutputsOutputs Spinal cord (lower motor neurons),Spinal cord (lower motor neurons),
thalamus, striatum cortex, brain stemthalamus, striatum cortex, brain stem
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Pia
surface
WM
???? organization of cortex???? organization of cortex
Cortex is arranged inCortex is arranged in ??????
Run verticallyRun vertically(perpendicular to surface)(perpendicular to surface)
~ 1 mm in diameter,~ 1 mm in diameter,
containing 300containing 300 600 cells600 cells Neurons in a columnNeurons in a column
synapse mostly withsynapse mostly with ??????
?????? do connect near bydo connect near bycolumnscolumns
?????? is comprised of billions ofis comprised of billions of
columnscolumns Cortical columns are quasiCortical columns are quasi--
independent processingindependent processingunits that canunits that can ??????
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Pia
surface
WM
Vertical organization of cortexVertical organization of cortex
Cortex is arranged inCortex is arranged infunctional columnsfunctional columns Run vertically (perpendicularRun vertically (perpendicular
to surface)to surface) ~ 1 mm in diameter,~ 1 mm in diameter,
containing 300containing 300 600 cells600 cells Neurons in a columnNeurons in a columnsynapse mostly withsynapse mostly with otherothercells in the columncells in the column
Axon collateralsAxon collaterals dodoconnect near by columnsconnect near by columns
CortexCortex is comprised ofis comprised ofbillions of columnsbillions of columns
Cortical columns are quasiCortical columns are quasi--independent processingindependent processingunits that canunits that can act in parallelact in parallel
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Radial Glia and formation of ????Radial Glia and formation of ????
corticocortical
bundles (CC)
thalamic
radiation (TR)nucleus
basalis (NB)
monoamine
centers (MA)
Radial Glia
marginal
zone
cortical
place
subplate
zone
intermediate zone
ventricular
zone
Migration
neuron
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Radial Glia and formation ofRadial Glia and formation of
Functional ColumnsFunctional Columns
corticocortical
bundles (CC)
thalamic
radiation (TR)nucleus
basalis (NB)
monoamine
centers (MA)
Radial Glia
marginal
zone
cortical
place
subplate
zone
intermediate zone
ventricular
zone
Migration
neuron
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Each column is aEach column is a ?????? ?????? specificity for each columnspecificity for each column
i.e. a column may receive sensoryi.e. a column may receive sensoryinput from rapidly adaptinginput from rapidly adaptingmechanoreceptor receptive fieldmechanoreceptor receptive field
from the middle left fingertip (viafrom the middle left fingertip (viaspinal and thalamic pathways)spinal and thalamic pathways)
Adjacent columns may pertain toAdjacent columns may pertain to ??????from the same regionfrom the same region
i.e. a column adjacent to the onei.e. a column adjacent to the one
mentioned above may receive inputmentioned above may receive inputfrom slowly adaptingfrom slowly adaptingmechanoreceptor receptive fieldmechanoreceptor receptive fieldfromfrom the same fingertipthe same fingertip
Columns and receptive fieldsColumns and receptive fields
C
olumn contains the ???
RA = rapidly adapting receptive fieldi.e. epicritic(small filled circles)
SA = slowly adapting receptive field
i.e. protopathic
(largeopen circles)
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Each column is aEach column is a single receptivesingle receptivefieldfield single modalitysingle modality Regional functionalRegional functional specificity forspecificity for
each columneach column i.e. a column may receive sensoryi.e. a column may receive sensory
input from rapidly adaptinginput from rapidly adapting
mechanoreceptor receptive fieldmechanoreceptor receptive fieldfrom the middle left fingertip (viafrom the middle left fingertip (viaspinal and thalamic pathways)spinal and thalamic pathways)
Adjacent columnsAdjacent columns may pertain tomay pertain todifferent modalitydifferent modality fromfrom the samethe sameregionregion i.e. a column adjacent to the onei.e. a column adjacent to the one
mentioned above may receive inputmentioned above may receive inputfrom slowly adaptingfrom slowly adaptingmechanoreceptor receptive fieldmechanoreceptor receptive fieldfromfrom the same fingertipthe same fingertip
Columns and receptive fieldsColumns and receptive fields
C
olumn contains the microcircuitry for representing a single receptive field
RA = rapidly adapting receptive fieldi.e. epicritic(small filled circles)
SA = slowly adapting receptive field
i.e. protopathic
(largeopen circles)
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Haines
Laminar organization:Laminar organization: AfferentConnectionsAfferentConnections??????GlutamatergicGlutamatergic
Predominantly terminate inPredominantly terminate in layer IVlayer IV??????
??????Glutamatergic axons from corticalGlutamatergic axons from corticalregions outside of local circuitregions outside of local circuitTerminate in layersTerminate in layers II VIVI, within a, within a
single cortical columnsingle cortical column??????Origins: NonOrigins: Non--specific thalamicspecific thalamicnuclei, basal nucleus of Meynert,nuclei, basal nucleus of Meynert,Brain stem nucleiBrain stem nucleiSubstrates: glutamatergic,Substrates: glutamatergic,
cholinergic, serotonergic,cholinergic, serotonergic,noradrogergicnoradrogergicTerminates inTerminates in layers Ilayers I VIVI, across, acrossmultiple cortical columnsmultiple cortical columnsRegulates overal level of corticalRegulates overal level of corticalexcitability, arousal, sleep,excitability, arousal, sleep,
wakefulnesswakefulness
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Laminar organization:Laminar organization: AfferentConnectionsAfferentConnectionsThalamocortical afferentsThalamocortical afferentsGlutamatergicGlutamatergic
Predominantly terminate inPredominantly terminate in layer IVlayer IV((thalamothalamo--receptive layer)receptive layer)Corticocortical afferentsCorticocortical afferentsGlutamatergic axons from corticalGlutamatergic axons from corticalregions outside of local circuitregions outside of local circuitTerminate in layersTerminate in layers II VIVI, within a, within asingle cortical columnsingle cortical columnDiffuse cortical afferentsDiffuse cortical afferentsOrigins: NonOrigins: Non--specific thalamicspecific thalamicnuclei, basal nucleus of Meynert,nuclei, basal nucleus of Meynert,Brain stem nucleiBrain stem nucleiSubstrates: glutamatergic,Substrates: glutamatergic,cholinergic, serotonergic,cholinergic, serotonergic,noradrogergicnoradrogergicTerminates inTerminates in layers Ilayers I VIVI, across, acrossmultiple cortical columnsmultiple cortical columnsRegulates overal level of corticalRegulates overal level of corticalexcitability, arousal, sleep,excitability, arousal, sleep,
wakefulnesswakefulness
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Laminar EfferentOrganization:Laminar EfferentOrganization:
Corticofugal/Corticothalamic EfferentsCorticofugal/Corticothalamic Efferents
Haines
Cortico-thalamic
Corticofugal EfferentsCorticofugal Efferents
Pyramidal cells ofPyramidal cells of??????
?????? targetstargets
i.e. Corticospinal,i.e. Corticospinal,corticostriate,corticostriate,
corticobulbar, corticocorticobulbar, cortico--
tectal, corticopontine, etc.tectal, corticopontine, etc.
Corticothalamic EfferentsCorticothalamic Efferents
?????? cells mainly in layercells mainly in layer ??????
Large projections in someLarge projections in somesensory systemssensory systems
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Laminar EfferentOrganization:Laminar EfferentOrganization:
Corticofugal/Corticothalamic EfferentsCorticofugal/Corticothalamic Efferents
Haines
Cortico-thalamic
Corticofugal EfferentsCorticofugal Efferents
Pyramidal cells ofPyramidal cells oflayer Vlayer V
NonNon--cortical (subcortical)cortical (subcortical)targetstargets
i.e. Corticospinal,i.e. Corticospinal,corticostriate,corticostriate,corticobulbar, corticocorticobulbar, cortico--tectal, corticopontine, etc.tectal, corticopontine, etc.
Corticothalamic EfferentsCorticothalamic Efferents
PyramidalsPyramidals cells mainly incells mainly inlayerlayer VI and VVI and V
Large projections in someLarge projections in somesensory systemssensory systems
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Laminar EfferentOrganization:Laminar EfferentOrganization:
Corticocortical EfferentsCorticocortical Efferents
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Cortico-cortical:
callosal &
association
Pyramidals cells mostly located in layersPyramidals cells mostly located in layers
??????
Axons project to distal cortical targetsAxons project to distal cortical targets
notnot ?????? Terminate in layersTerminate in layers ?????? of a singleof a single
cortical columncortical column
Cortical Targets in contralateralCortical Targets in contralateralhemisphere arehemisphere are ?????? connectionsconnections
Tend to project to homologous area inTend to project to homologous area inopposite hemisphereopposite hemisphere
Cortical targets within same hemisphereCortical targets within same hemisphere(ipsilateral) are(ipsilateral) are ??????
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Laminar EfferentOrganization:Laminar EfferentOrganization:
Corticocortical EfferentsCorticocortical Efferents
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Haines
Cortico-cortical:
callosal &
association
Pyramidals cells mostly located in layersPyramidals cells mostly located in layersII & IIII & II
Axons project to distal cortical targetsAxons project to distal cortical targets(not local projections)(not local projections)
Terminate in layersTerminate in layers II VIVI of a singleof a singlecortical columncortical column
Cortical Targets in contralateralCortical Targets in contralateralhemisphere arehemisphere are Collosal (commissural)Collosal (commissural)connectionsconnections
Tend to project to homologous area inTend to project to homologous area inopposite hemisphereopposite hemisphere
Cortical targets within same hemisphereCortical targets within same hemisphere(ipsilateral) are(ipsilateral) are Association ConnectionsAssociation Connections
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Majorcortical connection from a gyrusMajorcortical connection from a gyrus
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Majorcortical connection from a gyrusMajorcortical connection from a gyrus
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Martin
??? (95%)
??? (5%)
???
???
Up to3 layers
4-5 layers
???function
6 layers
1o sensory/motor, association area
Cognition, reasoning, perception,
speech/language
Memory (e.g., hippocampus)
Smell/Emotion (e.g., olfactory ctx)
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Cerebral Cortex Contains Regions Differing inCerebral Cortex Contains Regions Differing inPhylogenetic DevelopmentPhylogenetic Development
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Martin
Neocortex (95%)
Allocortex (5%)
Archicortex
Paleocortex
Up to3 layers
4-5 layers
Limbic function
6 layers
1o sensory/motor, association area
Cognition, reasoning, perception,
speech/language
Memory (e.g., hippocampus)
Smell/Emotion (e.g., olfactory ctx)
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Cerebral Cortex Contains Regions Differing inCerebral Cortex Contains Regions Differing inPhylogenetic DevelopmentPhylogenetic Development
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Martin
6 layers < 6 layers
AllocortexAllocortex
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Martin
???layers