11 8 23 Barlow 80 Journal of physiology Journal of neuroscience Cerebral Cortex Neurosicentist physiology introductory 1 3 4 recurrent 1 V1 STS David Van Essen 1990 32 V1 V2 V3 V4 STS STS 7A MT MST IT V1 parafoveal vision fovea 1
V1 V2 Baillarger Gennari 4 striate cortex 4B 4C 5 6 3 V1 V2 2 V1 1981 Hubel & Wiesel orientation UC Berkeley Ohzawa 240(beats/min) 38.5 39 4 4.5 V1 orientation selectivity orientation orientation preference direction selectivity bar contrast polarity dark bar bright bar direction selectivity grating non preferred direction curve 2
Ohzawa reverse correlation 20Hz 30Hz dark bar bright bar correlation bright bar dark bar 20 30 X XT dark bright dark bright dark bright grating on o dark bright region grating direction selectivity 3 Ohzawa reverse correlation bar X T T=0 bar bar 1 reverse correlation stationary bar 20 40 V1 Hubel & Wiesel orientation selectivity orientation tuning selectivity simple cell 4 on center - o surround align full drive V1 drive drive peak tuning EPSP IPSP EPSP tuning IPSP IPSP tuning peak cross-orientation inhibition EPSP IPSP optimal tuning drive drive EPSP optimal tuning IPSP EPSP peak 3
tuning cross-orientation inhibition Adam Sillito 17 complex cell bicuculline GABA major bicuculline GABA GABA cross-orientation inhibition cross-orientation inhibition orientation 17 trigger timing EPSP EPSP IPSP IPSP IPSP EPSP IPSP isolate EPSP IPSP in vivo intra bar IPSP average single sweep IPSP EPSP IPSP orientation tuning bload 45 orientation EPSP IPSP tuning sharp 45 orientation selectivity 45 C B 4
in vivo B C B cross-orientation inhibition shunting inhibition cross-orientation inhibition shunting inhibition spine spine dendrite spine spine shunting synapse conductance shunting synapse shunting synapse spine1 1 nonlinear tree spine neck David Ferster in vivo whole-cell recording shunting IPSP shunting conductance sharp record seal leak shunting IPSP whole-cell recording tight G seal current clamp o on subregion o on o on o on timing timing o on 1 1 1 2 3 4 o on simple cell o on o on 4 sub region test frequency shunting 5
conductance conductance conductance non preferred orientation shunting conductance conductance shunting IPSP shunting linear hyperpolarizing inhibition 3 V1 1 1 2 1 1 bicuculline GABA 3 orientation tuning histgram tuning curve orientation optimal orientation bicuculline GABA orientation tuning curve optimal non-optimal 4c parvo cellular stream magno cellular stream parallel rst stage magno 4b magno parvo 2-3 interblob blob blob interblob 2-3 2-3 2-3 5 5 6 6 4 6 focus feedback focus target V1 feedback 4b 2-3 V1 stage stage orientation tuning develop 2-3 4a 4b 6 4 orientation selective tuning property GABA tuning tuning 2-3 optimal peak 180 tuning bimodal tuning 4a 4b 6
orientation selective 180 direction selectivity optimal 4b set 6 6 tuning tuning tuningcurve V1 30 pool tuning tuning excitation average inhibition control tuning optimal peak optimal range response spare direction selectivity MT 4b direction selective bidirectional 2 1 A preferred direction preferred direction linear preferred direction non-preferred direction preferred direction non preferred direction EPSP threshold set direction selectivity direction selectivity directionality estimation estimation 200 uptakemechanism RI 200 dendrite 200 excitation back ground 7
cortex 1 2 layer control technical specic block 4 V1 blob blob 2-3 blob radial blob spot o suppression red on - blue o red on - green o green on - red o red on - blue o lter green emission red cone red green blue emission red cone overlap green green bicuculline on o contrast white convergence V blob red on-blue o on o phasic on o convergence on peak o peak broad pathway parallel segregate cortex V1 blob level 1 converge on o converge 8
tuning Sompolinsky orientation EPSP IPSP optimal tuning EPSP aerent EPSP cortex recurrent excitation EPSP net EPSP EPSP IPSP interaction optimal sharp tuning cortex recurrent excitation Ringach Shapley reverse correlation V1 orientation tuning recurrent connection tuning sharp stripe pattern channel channel texture pattern channel level Blasdel optical recording optical recording orientation map map map ocular dominance column ocular dominance column blob blob orientation orientation 200-300 orientation ocular dominance column B singularity 180 200-300 orientation orientation texture surface surface Gilbert orientation ocular dominance column pin wheel linear zone pin wheel linear zone pin wheel 9
5 V1 contextual modulation contextual modulation V1 Hubel & Wiesel story 40 orientation orientation orientation orientation correlate V1 1965 Hubel & Wiesel Modulate 19 optimal orientation 19 end stop inhibition tuning 19 grating grating modulation grating circular patch optimal orientation contrast orientation center surround hard edge orientation Cross orientation modulation modulation orientation post stimulus time histogram grating cicular patch 10
direction selectivity grating grating orientation 180 center optimal orientation 180 center patch surround 180 orientation surround orientation 0 orientation orientation orientation preference neuron domain 90 grating 180 90 high contrast high contrast contrast Somers contrast-response curve gain contrast high contrast orientation orientation optimal 0.1 0.2 0.3Hz 0.6Hz 0.2Hz optimal surround orientation 0.1 0.15 0.2Hz 0.3Hz 0.4 0.6Hz orientation domain center surround Hz 11
orientation layer orientation Anti-Hebb Somers intensity contrast activity gain saturate gain contrast saturate contrast linear contrast dominant high contrast saturate contrast range domain lateral connection orientation eld modulation full 15 15 7 6 7mm grating pattern center patch gray annulus annulus mask mask 1 7 7 activity cross-orientation 1 1.5 2 cross-orientaion surround orientation preference lateral exciation lateral inhibition drive lateral con- 12
nection lateral connection contrast Blakemore contrast response function center patch center patch contrast A sigmoid curve B C B low contrast C high contrast contrast center patch contrast response function low contrast facilitation high contrast suppression gain contrast contrast modulation low contrast facilitatory contrast low contrast facilitatory center contrast interaction 6 V1 adaptation adaptation 1979 Movshon & Lennie V1 contrast grating A B orientation grating 120 adaptation response adaptation adaptation orientation adaptation conductance Ferster low contrast adaptation high contrast adaptation contrast DC high contrast DCshift network F1 drive base level 20 adaptation adaptation 13
DC-shift 3 depression depression 20Hz voltage clamp EPSC IPSC EPSC IPSC IPSC EPSC 1 adaptation depression depression depression 2-3 horizontal connection distal dendrite 2 train depression 1 100 350 2 horizontal input source single cell single horizontal input 2 1 2 average 2 100msec paired pulse depression depression depression short distance depression 2 4 single input facilitation depression 7 horizontal depression 2 2 EPSP facilitation depression horizontal connection depression aerent 4 2-3 2-3 preference lateral connection center surround center lateral excitation saturate horizontal connection recurrent depression horizontal excitation depression high contrast orientation preference domain specic depression modulation contrast facilitatory adaptation averaging 14
deviation adaptation dynamic range domain network contrast pattern adaptation pattern adaptation Somers local center-surround physiology aerent 1 orientation domain domain singularity Hodgkin-Huxley adaptaion cunductance after hyper conductance Somers Nelson leader geniculate geniculate saturation contrast saturation adaptation geniculate high contrast top down top down Somers V1 V1 top down attention orientation orientaion 90 15
orientation orientation specic orientation orientation orientation cross correlation orientation preference functional domain specicity 300 specicity preference 4 4 adaptation grating population adaptation 4 16