Cambridge University Press
9780521852173 - Barrel Cortex - by Kevin Fox and Thomas Woolsey
Index
Index
6A-99 and barrel cortex-specific gene expression 245
Ablating a single barrel (effect on plasticity expression) 185
Absorption spectra for hemoglobin 220
Acetylcholinesterase
barrel field visualization 8
and staining the barrel field 10
transient developmental expression 104
ACI (adenylyl cyclase type 1) 235
role in barrel formation 100
role in silent synapse conversion 102
Active touch 142–145, 227–234
Activity blockade and dendrite orientation 97
Adenosine
and cortical blood flow 223
and hemodynamic coupling 223
Adenylyl cyclase type 1 see ACI
Adjacent spared whiskers 174
Adolescence (in rodents) 182
Adult plasticity 182–183
Age effect on plasticity 180–183
American opossum 6
AMPA receptor
insertion during development 102
and spine movement 205
and synaptic transmission 61
and thalamic axon development 101–103
Anesthetic
effect on blood flow 222, 223
effect on receptive field 130
Angiogenesis 226–227
Anterior lateral barrel subfield 1, 3, 10
Anterior thalamic nucleus 21, 23
Antidromic activation (and cortical stimulation) 151
Apoptosis and nerve section 211
APV (2-amino-5-phosphonovaleric acid)
effect on cortex development 96
effect on lesion-induced plasticity 96
Australian opossum 6
Autophosphorylation, CaMKII and plasticity 196
Axon-targeting cells (and single-layer cortex) 75
Barrel
composition 6–10
dimensions in mouse 113
dimensions in rat 113
formation 79, 97–98
sub-barrels 30
Barrel-column 113
Barrel field
as phenotypic trait 243
size 241
Barrel formation 98
and competition for space 97
timing 97
Barrel pattern 2
and growth associated protein 14, 208
rotation in thalamocortical afferent trajectory 29
use in pathway tracing 14
Barrel wall 6
coronal section 9
Barrelette
anatomical orientation 18
definition and the trigeminal nuclei 17
and interpolaris neurons’ dendritic trees 12
Barrelless mouse
and ACI 235
AMPA insertion 156
and AMPA receptor insertion 102
and development of barrel field 98
Barreloids
caudal tail innervation 24
location 23
size and orientation 22, 24
Basket cell 55–56
and gap junctions 71
large 56
nest 56
postulated role in DSI 63
projections to somata 40
small 55
and VPm terminations 30
BDNF (brain-derived neurotrophic factor)
and CREB in plasticity 209
rescue of infraorbital nerve section 94
role in silent synapse conversion 102
and rescue of apoptosis 211
Bidirection synapses 157
Bipolar cells 57
Birthday of individual cortical layers 83
Bitufted cells 57
Blood flow 218–227
fluorescence methods 219
oxygenation levels 220
radiographic methods 218
stroboscopic methods 219
supply to the barrel 218
Brain-derived neurotrophic factor see BDNF
Caged glutamate 234
and depression in the layer IV to II/III pathway 192
Cajal–Retzius cells 58
in cortical development 83
Calbindin-positive interneurons 40
Calcium-calmodulin-dependent protein kinase II, see CaMKII
CaMKII (calcium-calmodulin-dependent protein kinase II)
autophosphorylation 196
experience-dependent plasticity 197
dependence of LTP 157
and LTP induction in cortex 161
molecular switch hypothesis 195
percentage of postsynaptic density 196
role in experience-dependent plasticity 195–196
Cannabinoid
receptors in layer V inhibitory cells 43
role in DSI 63
role in LTD 168
Capsaicin and plasticity 215
Carnivores 6
and whiskers 2, 7
Catecholamine receptors 43
Caudal nucleus (nomenclature) 17
Caudalis
facial nucleus projection 20
and fur innervation 19
and primary afferent termination 19
Cell density and barrel formation 97
Center receptive field origins 132
Centrolateral nucleus projections 44
Cerebral blood flow 219, 220
C-fibers and tonic inhibition 215
C-fos and plasticity 206
Chandelier cells 56
receptive fields in layers II/III 42
receptive field size 128
and VPm terminations 30
Chemical LTD 163
Chessboard deprivation pattern 174
and depression 179
and potentiation 179
Cholinergic input to cortex 43
Cholinergic receptors 44
Chinchilla 2, 7
Clonal relationship of cortical cells 81
Column (cortical) 112
Columnar architecture (exceptions in barrel cortex) 12
Columnar hypothesis 11, 112–113
exceptions in barrel cortex 113
and latency 117, 121
transmission 117, 120–122
Computer modeling 237–240
Connexins 72
Contact response during whisking 144
Convergence (of whisker information) 115, 120
Correlated activity (role in pattern formation) 87
Cortical activity block
effect on lesion-induced plasticity 210
Cortical circuits 69
Cortical column 11–12
and modeling 237
Cortical connections
with MI 45
with SII 45
Cortical origin
of depression 179
of plasticity 183–186
Cortical plate
in developing cortex 83
and layer development 83
Cortical projections
and back projections 45
and forward projections 45
to other cortical areas 45
Cortical rhythms 240
Corticothalamic cells (percentage of inputs from VPm) 30
CPG15 and CREB in plasticity 209
CREB (cAMP response element-binding protein)
knockouts and plasticity 209
and plasticity 207, 208
Critical period
comparing visual and somatosensory cortex 182
lack of for potentiation in layers II/III 182
for layer IV 181
LTP 154, 210
for nerve degeneration 211
Cross-whisker potentiation 177
Cytochrome oxidase (CO) 9
staining in oralis 19
and trigeminal nuclei 17
and VPm staining 22, 24
Cumulative plasticity and CREB 208
Databases for cortical information 240
Deep vibrissae nerve 15, 16
Dendrite-targeting cells (and single-layer cortex) 75
Dendrites (orientation within layer IV) 30
Dendritic plasticity 210
2-deoxyglucose
and follicle reinnervation 214
and single-whisker stimulation 173
uptake in trigeminal nuclei 19
Depolarization-induced suppression of inhibition see DSI
Depressing synapse
layer VI to layer IV RS 70
and multipolar cells 70
short-term dynamics 64
Depression magnitude with whisker deprivation 174
Deprivation patterns 174
Development
of intracortical connections 106–108
of receptive fields and role of activity 94
role of activity in 94
Diagonal pathways (for plasticity) 190
Diffusible factors in cortical development 89
DiI (1,1-dioctadecyl-3,3,3,3-tetramethylindocarbocyanine percholate)
barrel field visualization 9
Dimensions (of barrels) 10
Direction selectivity 135–138
cortical mechanisms 136–138
of primary afferents 135
proportion of cortical cells 136
Discrimination of horizontal position 142
Disinhibition (for induction of LTP) 151
Dopaminergic input to cortex 43
Double bouquet cells 57
DSI 63, 64
differences between layer Va and Vb 63
Dynamic receptive fields 131
Dysgranular zone (projection received from VPM(vl) 24
Electrical synapses 71–73
Electroencephalography (use in defining receptive field size) 130
Electromyograph during whisking 144
ENU (N-ethyl-N-nitrosourea) mutagenesis 242
EPSP
miniature EPSP and effect of deprivation 186
timecourse 59
Ex vivo studies of cortical plasticity 192
Excitatory postsynaptic potential see EPSP
Excitatory synaptic transmission 59–62
Experience-dependent depression (relationship to LTD) 168–169
Experience-dependent plasticity 172–210
and LTP 159–162
Facial motor nucleus and sensory–motor loops 147
superior colliculus input 148
Facilitating synapse
and bitufted cells 70
layer VI to LTS 70
and LTS cells 68
short-term dynamics 64
Facilitation and direction selectivity 139
of adjacent whisker responses 139
Far side of the barrel (response latency) 118, 122
Fast rhythmic bursting cell 239
Fast spiking cells
characteristics of 66
direction selectivity 136
receptive field size 127
Fast spike units 125
Feedback
connections to trigeminal nuclei and plasticity 215
inhibition 124
loop types 145
Feedforward inhibition (evidence for) 126
fibroblast growth factor 8 (role in providing position cue in development) 89
fifth cranial nerve 16
Firing rate in vivo during whisker deprivation 169
Follicle 14–16
ablation
effect on development 95
effect on intracortical axons in adult 213
effect on dendrites in adults 214
composition and blood vessel innervation 16
and plasticity 210–212
reinnervation following damage 213
sinus 4, 14, 15
Forepaw representation and dendritic orientation 212
Forward genetics 241
Free nerve endings (location) 16
Fur innervation (in the trigeminal nuclei) 19
GABA (gamma aminobutyric acid) 62
GABA receptors
GABAA 62
GABAB 62
GABAC 63
Gamma oscillations 239
and exploratory whisking 72
and whisker stimulation 232
Gamma-aminobutyric acid see GABA
Gap junctions 71–73
in adult cortex 71
between excitatory cells in neonates 101
between inhibitory interneurons 42
extent and range of network 71
postulated between excitatory cells 72
Gene expression and plasticity 206
Gene trapping 245
Gerbil 10
Gestation
in mice 80
in rats 80
Glassy membrane 14
GluRs 61
GluR1
knockouts and presynaptic plasticity 159
and experience-dependent depression 199
insertion 198
phosphorylation by PKA 198
and rectification 156, 236
role in experience-dependent plasticity 198–199
role in LTD 167
role in postsynaptic LTP 157
tagging with green fluorescent protein 156
and whisker trimming 236
GluR2 4, 104
lack of experience dependence 236
Glutamate
evidence for being a cortical transmitter 60
and spine movements 205–206
Glutamate receptors see GluRs;see also mGluRs
growth-associated protein-14, 208
H-2ZI and barrel-specific gene expression 244–245
Halothane and cortical blood flow 224
Handshake hypothesis 91
Hemodynamic coupling 223
and stroke 227
Heterosynaptic depression
depriving one whisker 178
in vivo 177
Heterosynaptic LTD 162
Hippocampus
and discovery of LTP 151
and LTD 163
and single-layer cortex 73
Homeobox genes (role in segmentation) 89
Homosynaptic depression
depriving all whiskers 178
in vivo 177
Homosynaptic LTD 163
Horizontal pathways 190
anatomical evidence 189
for excitatory transmission between barrels 123
for potentiation 188–191
and peripheral lesions 212
5HT (role in thalamocortical patterning) 100
5HT1B receptors 4, 104, 242
barrel field visualization 8
and staining the barrel field 9
5HT transporter 4, 104
role in thalamocortical patterning 100
Hypertensive endothelial nitric oxide synthase knockouts 224
Inducible cAMP-responsive early repressor and plasticity 209
Immediate early genes 206, 207
Infraorbital nerve 16
development of mystacial pad innervation 93
section
and barrel formation 211
effect on horizontal connections 108
intracortical connections 212
and pattern formation 93
Inhibition
development of 108–109
and direction selectivity 139–141
and dual-frequency response circuits 71
feedforward and feedback 120
latency and duration in layer IV 124
lateral 126
local in the absence of intracortical transmission 134
and modeling 238
reversal potential in development 108
role in plasticity 162
timing of onset 108
Inhibition in layer V
sources for layer Va and Vb 43
sources for regular spiking and intrinsic bursting cells 43
Inhibitory cells 55–58
origin in development 82
receptive fields 133–134
Inhibitory connections on spines 109
Inhibitory feedback within the barrel 115, 119
Inhibitory interneurons 40–43
calretinin staining 40
fast spiking cells 42
and gap junctions 42
general classification 40
in layers II/III and their projections 42
in layer IV
and projections 41–42
and thalamic efficacy 41
in layer VI 43
LTS cells 42
Martinotti cells 43
parvalbumin staining 40
Inhibitory postsynaptic potential see IPSP IPSP
Inhibitory synapses
effect of whisker trimming during development 109
timing of development 109
and transmission 62–63
and transmission in neonatal cortex 101
Innervation
of the follicle 2–3
levels in relation to size of cortical area 2
Intrinsic burster (IB) cell
and inhibitory input 37
and intracortical connectivity 37
and lack of thalamic input 30
morphology 53
and plasticity pathways 191
pyramidal cells 53
Intermediate progenitor cells 80
Interpolar nucleus (nomenclature) 17
Interpolaris
and cortical feedback 147
dendritic field relative to barrelettes 22, 24
and primary afferent termination 19
and receptive field size 22, 24–25
thalamic projection 19
Intracortical connections
excitatory 31–40
general overview 32
Intralaminar nuclei projections to cortex 31, 44
Intrinsic imaging of cortex during whisking 233
Ionotropic receptors 61
IPSP (inhibitory postsynaptic potential)
effect on EPSP 59
timecourse 59
Ischemia 225
JunB and plasticity 207
Kainate receptor
elimination during LTP 154
postsynaptic receptors in development 105
presynaptic receptors in development 105
quantal activation 235
role in development of thalamocortical transmission 105
KROX24 and plasticity 206
Labeled line
and context of information 114
salience of information 114, 117, 121
in sensory processing 113–114
Lanceolate endings
location 16
rapidly adapting receptors 16
Laser Doppler flow cytometry 219
Late-phase plasticity 199–201
Latency
of layer IV GABAergic cells 115, 120
of response in layers IV and Vb 118, 122
of response in whisking 143
of response to principal whiskers within the column 117, 121
of thalamic responses 115, 118
of thalamocortical input 115, 118
Lateral inhibition 127–129
anisotropy within receptive field 129
and cortical substrates 128
role in plasticity 180
size of in the cortex 128
Lateral and medial ganglionic eminence (cortical progenitors) 82
Layer II/III to Vb pathway for depression 192
Layer IV receptive field size 130
Layer V pathways for plasticity 191
Layer VI recurrent collaterals
frequency response 70
on to inhibitory cells 70
and plasticity 204
Lectin binding and map formation 91
Lemniscal pathway 19
Lidocaine and effect of peripheral nerve block on VPm 214
Long-term memory and neocortical plasticity 153
Long-term depression see LTD
Long-term potentiation see LTP
Loops
as anatomical pathways 145
sensory 146–147
as sensory–motor mechanisms 145
sensory–motor 147–148
Low threshold spiking see LTS
LTD
early phase 194
induction methods 163
timing window 164
LTP
critical period of thalamocortical synapse 154
early phase 194
effect of experience on probability 160
in layer II/III cells 157–159
methods of induction 151
LTS
cells and short-term dynamics 68
characteristics of 66
Magnetic resonance imaging
arterial spin 219
blood flow measurement 219
BOLD functional MRI 220
Marginal zone
and layer I 83
in cortical development 83
Marsupials 6
and evolution 6
Martinotti cell 57–58
as an analogue of the hippocampal OL-M cell 75
and cross-column inhibition 58
Merkel’s disc
location 16
slowly adapting receptors 16
Metabotropic glutamate receptors see mGluRs
mGluRs (metabotropic glutamate receptors) 61, 62
group I 62
mGluR1α 62
mGluR2/3 62
mGluR5 62
in development of thalamocortical transmission 105
role in barrel formation 99
MI projections from barrel cortex 45
Migration
of cells during development 84
period of in the rat 83
Miniature EPSPs and effect of deprivation 186
Minicolumns 112
Monoamine oxidase
knockout and thalamocortical afferent patterning 99
mutant mouse strain 242
Monocular deprivation in relation to barrel cortex plasticity 177
Motor cortex primary area see MI
Motor cortex and sensory–motor loops 148
Multiwhisker responses 138–141
integration in texture discrimination 142
summation in plasticity 180
Muscimol
and barrel inactivation 133
and preventing plasticity 184
Myomorphs 6
Nearside of the barrel and response latency 118–120, 122
Neighboring barrels and cross barrel-column transmission 117, 122–124
Nerve growth factor (overexpression in mystacial pad) 94
Network tension 238
NeuroD2 and plasticity 208
Neurogliaform cells 58
Neuropeptide
expression in GABAergic inhibitory cells 63
and inhibitory cells 55
Neurotrophins
and cell death in principalis 94
neurotrophin-3 rescue of infraorbital nerve section 94
and pattern formation 93
Nicotinic receptors 4, 104
modulation of glutamatergic transmission 105
and staining the barrel field 9
transient developmental expression 104
Nissl stain 9
barrel field visualization 9
Nitric oxide synthase
and cortical blood flow 223
and hemodynamic coupling 223–224
role in LTP 162, 198, 199
NMDA receptor
contribution to sensory transmission 61
dependence of LTD 166
dependence of LTP 157
and in vivo plasticity 195
and neonatal cortical transmission 101
presynaptic 167
role in barrel formation 99
role in detecting correlated activity 88
role in development of horizontal connections 108
role in LTD 166
subunits
NR1 knockout mice and somatotopic pattern 96
NR2A 4, 104
NR2B 4, 104
NR2C 61
and synaptic transmission 61, 194–195, 197–198
Noradrenergic input to cortex 43
Occlusion of LTD by whisker deprivation 169, 193
Ocular dominance plasticity and LTD 163
Oral nucleus (nomenclature) 17
Oralis and primary afferent termination 19
Optical imaging
of blood flow 221
spectroscopy 223
Parafascicularis projections 44
Parvalbumin
and connexin 10–11
staining in layer IV 41
Pattern formation 79
and acetylcholinesterase staining 91
and infraorbital nerve transection 92
Pattern information 98
and thalamic afferents 92
theories on where it is carried in development 89
Pattern of whiskers 2
Peripheral pattern as a source of central pattern 93
Phase comparator and whisking 145
Phase lock and whisking 148
Phase-locked loop and decoding contact 145
Phosphatase (role in LTD) 167
Photostimulation 234
and intracortical pathways 235
Pinipeds 6
PKA (protein kinase A)
and CREB in plasticity 208
and immature synapse potentiation 199
regulatory subunits and barrel field formation 242
role in reversal of LTD 166see also RIIβ-subunit of PKA
Phospholipase C β1-subunit role in barrel formation 98, 100
Point-spread function 117, 122–124
Polymerization of actin and spine movement 205
POm (posterior medial thalamic nucleus) 26–29
back projections from cortex 48
control of receptive fields 28
and cortical feedback 146
cortical input from layer V 28
influence of zona incerta 28
inputs from superior colliculus 28
inputs from trigeminal nuclei 27
projections from the trigeminal nuclei 24
projections to cortical layer IV 26
projections to cortical layers I to V 26
topography of receptive fields 28
and transcortical relay 28
Pontine nuclei (projections from barrel cortex) 47
Posterior medial barrel subfield 1, 3, 10
Posterior medial thalamic nucleus see POm
Posterior nucleus 21, 23
caudal division 22, 24
Postnatal plasticity 181
Poststimulus time histogram (measuring receptive fields) 130
Potentiation
in a deprived barrel 177
magnitude with whisker deprivation 174
timecourse 179
Premap hypothesis
theory (manifestation in the ventricular zone) 85
transplantation studies 86
Preplate (in cortical development) 83
Presynaptic
factors expressed on thalamocortical afferents 103
LTP in cortex 157
plasticity (experience-dependent) 203–205
Primary afferent activity during whisking 144
Principal nucleus (nomenclature) 17
Principalis
and lateral inhibition 127
and primary afferent termination 18
receptive field size 22, 23
thalamic projection 19
tonic and phasic responses 22, 23
Probability of LTD and effect of whisker deprivation 193
Progenitor cells (in the cortex) 80–83
Protein kinase A see PKA PKA
Protraction of whiskers (relation to cortical firing) 148
PSD-95 involvement in AMPA insertion 162
Purkinje cells and LTD 163
Pyramidal cells 51–55
axons
layer II cells 51
layer III cells 52
connectivity of layer V cells 37
corticothalamic cells of layer VI 38
dendrites
layer II cells 51
layer III cells 52
of layer II 36
of layer III 36
of layer V 37
of layer V 52–55
of layer Va 37
of layer Vb 37
of layer VI
axonal projections 55
polymorphic 55
of layer VIa and their connections 39
of layer VIb and their connections 39
methods of definition 51
recurrent collaterals 54
regular spiking and instrinsic bursting 37
Quantal analysis
and LTD 166
and presynaptic LTP 158
Quantitative trait analysis 241
Quiet synapses at thalamocortical connections 156
Rabbit 6
and studies of cortical inhibition 128
Radial glial cells
and production of cortical neurons 80
Rapidly adapting cells (and direction selectivity) 22, 24
Rapidly adapting fibers 21, 23
direction selectivity 135
Receptive field
in absence of intracortical transmission 132
development (effect of whisker trimming) 106
Receptive field size 129–130
estimates for layer IV 131
in layer IV cells 115, 119
in POm 117, 121
technical factors affecting measurement 129–130
in VPm 117, 120–122
Receptor insertion in adults 156
Rectification in GluR1 homomeric channels 156
Recurrent collaterals (as pathways for plasticity) 189
Reelin (role in cortical development) 83
Regular spiking 53
Regular spiking units 126
direction selectivity 136
Regular spiking cells
characteristics of 65
and inhibitory input 37
and intracortical connectivity 37
and plasticity pathways 191
pyramidal cells 53
Reptilian cortex
and single-layer cortex 73
synaptic physiology 73
Response latency of layer VI recurrent collaterals 70
Resonance of whiskers 228, 230
Rete ridge 16
Reticular nucleus of the thalamus 22, 23, 25–26
cortical projections 25
projections to POm 26
projections to thalamic nuclei 25
projections to VPm 25
and thalamic spindle oscillations 25
Reverse genetics 240
RIIβ-subunit of PKA
and barrel formation 99, 100
specificity of isoform effect on LTD 198, 199
ROCK-PIIa and spine movements 206
Row transmission (asymmetry) 123
Septae 6
Septal areas
columns (effect of deprivation during development) 193
connections (inputs from POm) 30
connections to MI and SII 45
neurons (receptive fields) 123
pathways (and LTP) 189
regions (connectivity) 10–11
tenascin staining 10
Serotonergic
input to cortex 43
receptors 44
role in thalamic axon development 99, 104
Short-term dynamics
between excitatory cells 66
changes during experience-dependent plasticity 159
effects of age 64
effects of deprivation 192
effects of plasticity 65
intrinsic diversity of connections 65
and release probability 67
SII projection from barrel cortex 45
Silent synapses
in development 101
and LTP 154
and NeuroD2 208
timecourse of elimination 102
Simulation of a cortical column 239
Single spared whisker 174
Single-layer cortex 73–75
relationship to layers II/III 75
relationship to layer IV 77
relationship to layer V 77
Slowly adapting fibers 21, 23
and direction selectivity 135
Somatosensory cortex
barrel cortex as part of 2
proportion devoted to barrel cortex 2
Somatosensory area I and II
see SI and SII, respectively
Somatostatin expression in Martinotti cell subtypes 58
Sonic hedgehog and spinal cord development 85
Spared whisker response spread 176
Spatial summation of cortical EPSPs 62, 151
Spider’s web cells 58
Spike pairing and presynaptic plasticity 157
Spike threshold (and effect on direction selectivity) 137
Spike timing
and induction of LTD 164–165
in vivo (effect of whisker deprivation) 169
Spinal V nuclei (nomenclature) 17
Spindle oscillations and the reticular nucleus 25
Spine
attractant and spine movements 205
behavior during whisker deprivation 186
classification 201
density 203
motility effect of whisker trimming 107
and plasticity 201–203
turnover 201
turnover and chessboard deprivation 202
Spiny stellate cell 49–50
axonal projections 50
columnar projections 34–36
dendrite orientation 50
septal connectivity 34
Stability
synaptic 193
of synapses (percentage) 201
of axonal arbors 204
Star pyramidal cells 50–51
apical dendrite 51
axonal projections 51
basal dendrites 51
intracortical connectivity 34
Striatal connections (septal diffuse pathway from cortex) 47
Striatum (projection from barrel cortex) 47–48
Stroke 225–226
and vascular ligation 225, 226
photothrombotic infarct 225
Strontium 234
Subcortical contribution to cortical surround receptive fields 132
Subcortical plasticity 183, 211, 213–215
in adults 213
Subcortical projections
to motor structures 46–48
to sensory structures 48
Subplate cells role in thalamocortical afferent in growth 91
Subventricular zone (in development) 83
Succinate dehydrogenase 9
Summation
role in multiwhisker plasticity 180
supralinear in whisker responses 139
Superficial vibrissae nerve 15, 16
Superior colliculus and sensory–motor loops 148
Suppression of interwhisker responses 139
Surround receptive field
and the cortical component 132
Symmetrical synapse increase during development 109
Synapse
density developmental timecourse 101
formation (timing in layers II/III) 106
synaptogenesis and late-phase plasticity 201
Synaptic depression and LTD at the thalamocortical synapse 165
Syndactyly 172
SynGAP knockout (and barrel formation) 99
Tabla rasa hypothesis
and transplantation studies 86
Temporal summation (and thalamocortical response transformation) 115, 118–119
Temporal synchrony (role in receptive field generation) 133
Tenascin
barrel field visualization 8
staining in the barrel field 10
Tetrodotoxin (blockade of infraorbital nerve activity during development) 94
Texture discrimination 142
Thalamic input
modeling 238
to fast spiking cells 42
Thalamocortical afferent
arbor development 96, 101–106
axon projection 29–31
connection on spiny stellate cells 29
lack of projections to LTS cells 42
location in barrel 6–9
and neonatal experience-dependent plasticity 204
percentage of synapses in layer IV barrels 29
percentage of synapses on inhibitory cells 30
pattern 91
and specificity of projection 115
synapse development 79
terminations on inhibitory cells 29
timing of ingrowth in development 91
Thalamocortical inputs
and short-latency responses 115, 118–119
and short-term dynamics 68
Thalamocortical response transformation 118–120, 122
Thalamus and direction selectivity 136
Three-eyed frogs (role of correlated activity in development) 88
Tonic inhibition 63
Transplantation
of cortical cells during development 86–87
visual to barrel cortex 86, 87
Trigeminal ganglion
and direction selectivity 136
number of cells per whisker 2
Trigeminal nuclei 17–22
back projections from cortex 48
and fur innervation 19
and primary afferent innervation 18–19
projections within the trigeminal nuclei 20
and receptive fields 20–22
and whisker pattern 17
Trimming whiskers see Whisker trimming
Tufted pyramidal cells (and lack of thalamic input) 30
Vasointestinal polypeptide in bipolar cells 57
Velocity sensitivity 134
of primary afferents 21, 23
and whisking movements 142
Ventrolateral nucleus 21, 23
Ventral posterior medial thalamic nucleus see VPm
Vertical pathways for LTD 191
vesicular glutamate transporter 1 (VGLUT1) and vesicle localization 61
vesicular monoamine transporter and pattern formation 100, 242
Visual processing (rerouted to somatosensory cortex) 86
Voltage-sensitive dyes (and horizontal transmission) 123
VPm
back projections from cortex 48
core projection to cortex 25
and lateral inhibition 127
plasticity in adults 185
projections from the trigeminal nuclei 24
projections to cortical layers I and III 31
thalamocortical axon trajectory 29
ventroposteriomedial thalamic nucleus 22, 24–25
VPm(vl)
cortical projection pattern 24
and interpolaris projections 24
Walrus 6
Whisk 6
Whisk cycle 143
amplitude control 148
and phase sensitivity of cortical neurons 143
and position encoding 144
Whisker
natural resonance properties 228
metal 230
multichannel stimulator 231
pattern 2
resonance 228, 230
single-spared 174
stimulator 232
summation of responses
role in plasticity 180
supralinear 139
suppression of interwhisker responses 139
texture discrimination 142
Whisker trimming
effect on septal and barrel pathways 236
and GluR1 insertion 236
and intracortical axons 213
and plasticity 172
spared whisker response spread 176
Whisking
bending moment 230
frequency 142
patterns 141
stick and slip movements 228
and texture discrimination 142
Whisk phase
and anticipatory responses 233
cortical responses 144
responses intracellular recording 233
Working memory 154
Zona incerta (influence on POm) 28
Zinc staining 175
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