{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,14]],"date-time":"2026-04-14T12:41:28Z","timestamp":1776170488419,"version":"3.50.1"},"reference-count":51,"publisher":"Elsevier BV","issue":"5","license":[{"start":{"date-parts":[[2003,6,1]],"date-time":"2003-06-01T00:00:00Z","timestamp":1054425600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2003,6,1]],"date-time":"2003-06-01T00:00:00Z","timestamp":1054425600000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2013,7,17]],"date-time":"2013-07-17T00:00:00Z","timestamp":1374019200000},"content-version":"vor","delay-in-days":3699,"URL":"http:\/\/www.elsevier.com\/open-access\/userlicense\/1.0\/"},{"start":{"date-parts":[[2003,6,1]],"date-time":"2003-06-01T00:00:00Z","timestamp":1054425600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2003,6,1]],"date-time":"2003-06-01T00:00:00Z","timestamp":1054425600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2003,6,1]],"date-time":"2003-06-01T00:00:00Z","timestamp":1054425600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2003,6,1]],"date-time":"2003-06-01T00:00:00Z","timestamp":1054425600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2003,6,1]],"date-time":"2003-06-01T00:00:00Z","timestamp":1054425600000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"content-domain":{"domain":["cell.com","elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Neuron"],"published-print":{"date-parts":[[2003,6]]},"DOI":"10.1016\/s0896-6273(03)00332-5","type":"journal-article","created":{"date-parts":[[2004,4,21]],"date-time":"2004-04-21T03:12:17Z","timestamp":1082517137000},"page":"689-699","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":219,"title":["Dynamics of Orientation Selectivity in the Primary Visual Cortex and the Importance of Cortical Inhibition"],"prefix":"10.1016","volume":"38","author":[{"given":"Robert","family":"Shapley","sequence":"first","affiliation":[]},{"given":"Michael","family":"Hawken","sequence":"additional","affiliation":[]},{"given":"Dario L.","family":"Ringach","sequence":"additional","affiliation":[]}],"member":"78","reference":[{"key":"10.1016\/S0896-6273(03)00332-5_BIB1","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1017\/S0952523899162114","article-title":"A model for the intracortical origin of orientation preference and tuning in macaque striate cortex","volume":"16","author":"Adorjan","year":"1999","journal-title":"Vis. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB2","first-page":"81","article-title":"Dynamic differentiation of GABAA-sensitive influences on orientation selectivity of complex cells in the cat striate cortex","volume":"104","author":"Allison","year":"1995","journal-title":"Exp. Brain Res."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB3","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1152\/jn.2000.84.2.909","article-title":"Orientation tuning of input conductance, excitation, and inhibition in cat primary visual cortex","volume":"84","author":"Anderson","year":"2000","journal-title":"J. Neurophysiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB4","doi-asserted-by":"crossref","first-page":"3844","DOI":"10.1073\/pnas.92.9.3844","article-title":"Theory of orientation tuning in visual cortex","volume":"92","author":"Ben-Yishai","year":"1995","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB5","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1017\/S0952523800004314","article-title":"Role of inhibition in the specification of orientation selectivity of cells in the cat striate cortex","volume":"2","author":"Bonds","year":"1989","journal-title":"Vis. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB6","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1038\/30735","article-title":"Visual input evokes transient and strong shunting inhibition in visual cortical neurons","volume":"393","author":"Borg-Graham","year":"1998","journal-title":"Nature"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB7","doi-asserted-by":"crossref","first-page":"47","DOI":"10.1146\/annurev.neuro.21.1.47","article-title":"Local circuits in primary visual cortex of the macaque monkey","volume":"21","author":"Callaway","year":"1998","journal-title":"Annu. Rev. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB8","doi-asserted-by":"crossref","first-page":"470","DOI":"10.1523\/JNEUROSCI.20-01-00470.2000","article-title":"Membrane potential and firing rate in cat primary visual cortex","volume":"20","author":"Carandini","year":"2000","journal-title":"J. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB9","doi-asserted-by":"crossref","first-page":"5251","DOI":"10.1523\/JNEUROSCI.13-12-05251.1993","article-title":"Development of orientation selectivity in ferret visual cortex and effects of deprivation","volume":"13","author":"Chapman","year":"1993","journal-title":"J. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB10","doi-asserted-by":"crossref","first-page":"1347","DOI":"10.1523\/JNEUROSCI.11-05-01347.1991","article-title":"Relation of cortical cell orientation selectivity to alignment of receptive fields of the geniculocortical afferents that arborize within a single orientation column in ferret visual cortex","volume":"11","author":"Chapman","year":"1991","journal-title":"J. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB11","doi-asserted-by":"crossref","first-page":"1177","DOI":"10.1016\/S0896-6273(00)80498-5","article-title":"Strength and orientation tuning of the thalamic input to simple cells revealed by electrically evoked cortical suppression","volume":"20","author":"Chung","year":"1998","journal-title":"Neuron"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB12","doi-asserted-by":"crossref","first-page":"2056","DOI":"10.1046\/j.1460-9568.1998.00218.x","article-title":"Evidence for a contribution of lateral inhibition to orientation tuning and direction selectivity in cat visual cortex","volume":"10","author":"Crook","year":"1998","journal-title":"Eur. J. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB13","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1016\/0042-6989(82)90112-2","article-title":"The orientation and direction selectivity of cells in macaque visual cortex","volume":"22","author":"De Valois","year":"1982","journal-title":"Vision Res.."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB14","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1523\/JNEUROSCI.20-01-00495.2000","article-title":"Spatial and temporal structure of receptive fields in primate somatosensory area 3b","volume":"20","author":"DiCarlo","year":"2000","journal-title":"J. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB15","doi-asserted-by":"crossref","first-page":"735","DOI":"10.1113\/jphysiol.1991.sp018733","article-title":"A functional microcircuit for cat visual cortex","volume":"440","author":"Douglas","year":"1991","journal-title":"J. Physiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB16","doi-asserted-by":"crossref","first-page":"981","DOI":"10.1126\/science.7638624","article-title":"Recurrent excitation in neocortical circuits","volume":"269","author":"Douglas","year":"1995","journal-title":"Science"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB17","doi-asserted-by":"crossref","first-page":"287","DOI":"10.1016\/S0896-6273(00)00103-3","article-title":"Adaptation-induced plasticity of orientation tuning in adult visual cortex","volume":"28","author":"Dragoi","year":"2000","journal-title":"Neuron"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB18","doi-asserted-by":"crossref","first-page":"1172","DOI":"10.1523\/JNEUROSCI.08-04-01172.1988","article-title":"Spatially opponent excitation and inhibition in simple cells of the cat visual cortex","volume":"8","author":"Ferster","year":"1988","journal-title":"J. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB19","doi-asserted-by":"crossref","first-page":"423","DOI":"10.1016\/S0079-6123(08)63625-0","article-title":"The synaptic inputs to simple cells of the cat visual cortex","volume":"90","author":"Ferster","year":"1992","journal-title":"Prog. Brain Res."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB20","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1038\/380249a0","article-title":"Orientation selectivity of thalamic input to simple cells of cat visual cortex","volume":"380","author":"Ferster","year":"1996","journal-title":"Nature"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB21","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1002\/cne.902890211","article-title":"Arborisation pattern and postsynaptic targets of physiologically identified thalamocortical afferents in striate cortex of the macaque monkey","volume":"289","author":"Freund","year":"1989","journal-title":"J. Comp. Neurol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB22","doi-asserted-by":"crossref","first-page":"1014","DOI":"10.1038\/nn731","article-title":"Dynamics of the orientation-tuned membrane potential response in cat primary visual cortex","volume":"4","author":"Gillespie","year":"2001","journal-title":"Nat. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB23","doi-asserted-by":"crossref","first-page":"106","DOI":"10.1113\/jphysiol.1962.sp006837","article-title":"Receptive fields, binocular interaction and functional architecture in the cat's visual cortex","volume":"160","author":"Hubel","year":"1962","journal-title":"J. Physiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB24","doi-asserted-by":"crossref","first-page":"215","DOI":"10.1113\/jphysiol.1968.sp008455","article-title":"Receptive fields and functional architecture of monkey striate cortex","volume":"195","author":"Hubel","year":"1968","journal-title":"J. Physiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB25","doi-asserted-by":"crossref","first-page":"253","DOI":"10.1146\/annurev.ne.11.030188.001345","article-title":"Anatomical organization of macaque monkey striate visual cortex","volume":"11","author":"Lund","year":"1988","journal-title":"Annu. Rev. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB26","doi-asserted-by":"crossref","first-page":"8087","DOI":"10.1073\/pnas.110135097","article-title":"A Neuronal Network Model of Sharpening and Dynamics of Orientation Tuning in an Input Layer of Macaque Primary Visual Cortex","volume":"97","author":"McLaughlin","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB27","series-title":"Statistics of Directional Data","author":"Mardia","year":"1972"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB28","doi-asserted-by":"crossref","first-page":"1645","DOI":"10.1073\/pnas.022638499","article-title":"Spatial frequency and orientation tuning dynamics in area V1","volume":"99","author":"Mazer","year":"2002","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB29","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1016\/S0896-6273(03)00064-3","article-title":"Orientation and direction selectivity of synaptic inputs in visual cortical neurons","volume":"37","author":"Monier","year":"2003","journal-title":"Neuron"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB30","doi-asserted-by":"crossref","first-page":"774","DOI":"10.1126\/science.8047882","article-title":"Orientation selectivity of cortical neurons during intracellularblockade of inhibition","volume":"265","author":"Nelson","year":"1994","journal-title":"Science"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB31","doi-asserted-by":"crossref","first-page":"260","DOI":"10.1152\/jn.1981.46.2.260","article-title":"Receptive-field structure in cat striate cortex","volume":"46","author":"Palmer","year":"1981","journal-title":"J. Neurophysiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB32","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1038\/378281a0","article-title":"Specificity of monosynaptic connections from thalamus to visual cortex","volume":"378","author":"Reid","year":"1995","journal-title":"Nature"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB33","doi-asserted-by":"crossref","first-page":"2455","DOI":"10.1016\/S0042-6989(96)00247-7","article-title":"A subspace reverse-correlation technique for the study of visual neurons","volume":"37","author":"Ringach","year":"1997","journal-title":"Vision Res."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB34","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1038\/387281a0","article-title":"The dynamics of orientation tuning in the macaque monkey striate cortex","volume":"387","author":"Ringach","year":"1997","journal-title":"Nature"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB35","doi-asserted-by":"crossref","first-page":"1018","DOI":"10.1152\/jn.00614.2001","article-title":"Suppression of neural responses to nonoptimal stimuli correlates with tuning selectivity in macaque V1","volume":"87","author":"Ringach","year":"2002","journal-title":"J. Neurophysiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB36","doi-asserted-by":"crossref","first-page":"5639","DOI":"10.1523\/JNEUROSCI.22-13-05639.2002","article-title":"Orientation selectivity in macaque v1","volume":"22","author":"Ringach","year":"2002","journal-title":"J. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB37","article-title":"Dynamics of orientation tuning in macaque V1","volume":"in press","author":"Ringach","year":"2003","journal-title":"J. Neurophysiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB38","doi-asserted-by":"crossref","first-page":"757","DOI":"10.1113\/jphysiol.1996.sp021530","article-title":"Mechanisms underlying orientation selectivity of neurons in the primary visual cortex of the macaque","volume":"494","author":"Sato","year":"1996","journal-title":"J. Physiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB39","doi-asserted-by":"crossref","first-page":"1320","DOI":"10.1152\/jn.1976.39.6.1320","article-title":"Quantitative studies of single-cell properties in monkey striate cortex. II. Orientation specificity and ocular dominance","volume":"39","author":"Schiller","year":"1976","journal-title":"J. Neurophysiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB40","doi-asserted-by":"crossref","unstructured":"Shapley, R.M. (1994). Linearity and non-linearity in cortical receptive fields. In Higher Order Processing in the Visual System, Ciba Symposium 184, 71\u201387, Wiley, Chichester.","DOI":"10.1002\/9780470514610.ch4"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB41","doi-asserted-by":"crossref","first-page":"512","DOI":"10.1126\/science.1065916","article-title":"Dynamics and constancy in cortical spatiotemporal patterns of orientation processing","volume":"295","author":"Sharon","year":"2002","journal-title":"Science"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB42","doi-asserted-by":"crossref","first-page":"97","DOI":"10.1023\/A:1015760707294","article-title":"Coarse-grained reduction and analysis of a network model of cortical response","volume":"12","author":"Shelley","year":"2002","journal-title":"J. Comput. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB43","doi-asserted-by":"crossref","first-page":"93","DOI":"10.1023\/A:1020158106603","article-title":"States of high conductance in a large-scale model of the visual cortex","volume":"13","author":"Shelley","year":"2002","journal-title":"J. Comput. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB44","doi-asserted-by":"crossref","first-page":"865","DOI":"10.1016\/0306-4522(93)90133-Z","article-title":"Dynamics of orientation tuning in the cat striate cortex neurons","volume":"56","author":"Shevelev","year":"1993","journal-title":"Neuroscience"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB45","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1113\/jphysiol.1975.sp011056","article-title":"The contribution of inhibitory mechanisms to the receptive field properties of neurones in the striate cortex of the cat","volume":"250","author":"Sillito","year":"1975","journal-title":"J. Physiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB46","doi-asserted-by":"crossref","first-page":"5448","DOI":"10.1523\/JNEUROSCI.15-08-05448.1995","article-title":"An emergent model of orientation selectivity in cat visual cortical simple cells","volume":"15","author":"Somers","year":"1995","journal-title":"J. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB47","doi-asserted-by":"crossref","first-page":"514","DOI":"10.1016\/S0959-4388(97)80031-1","article-title":"New perspectives on the mechanisms for orientation selectivity","volume":"7","author":"Sompolinsky","year":"1997","journal-title":"Curr. Opin. Neurobiol."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB48","doi-asserted-by":"crossref","first-page":"582","DOI":"10.1007\/BF00229326","article-title":"The effects of contrast on the linearity of spatial summation of simple cells in the cat's striate cortex","volume":"79","author":"Tolhurst","year":"1990","journal-title":"Exp. Brain Res."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB49","doi-asserted-by":"crossref","first-page":"5908","DOI":"10.1523\/JNEUROSCI.18-15-05908.1998","article-title":"Contrast-invariant orientation tuning in cat visual cortex","volume":"18","author":"Troyer","year":"1998","journal-title":"J. Neurosci."},{"key":"10.1016\/S0896-6273(03)00332-5_BIB50","doi-asserted-by":"crossref","first-page":"1943","DOI":"10.1126\/science.286.5446.1943","article-title":"Linking spontaneous activity of single cortical neurons and the underlying functional architecture","volume":"286","author":"Tsodyks","year":"1999","journal-title":"Science"},{"key":"10.1016\/S0896-6273(03)00332-5_BIB51","doi-asserted-by":"crossref","first-page":"5203","DOI":"10.1523\/JNEUROSCI.21-14-05203.2001","article-title":"How simple cells are made in a nonlinear network model of the visual cortex","volume":"21","author":"Wielaard","year":"2001","journal-title":"J. Neurosci."}],"container-title":["Neuron"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0896627303003325?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0896627303003325?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2025,10,12]],"date-time":"2025-10-12T06:27:24Z","timestamp":1760250444000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0896627303003325"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2003,6]]},"references-count":51,"journal-issue":{"issue":"5","published-print":{"date-parts":[[2003,6]]}},"alternative-id":["S0896627303003325"],"URL":"https:\/\/doi.org\/10.1016\/s0896-6273(03)00332-5","relation":{},"ISSN":["0896-6273"],"issn-type":[{"value":"0896-6273","type":"print"}],"subject":[],"published":{"date-parts":[[2003,6]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"Dynamics of Orientation Selectivity in the Primary Visual Cortex and the Importance of Cortical Inhibition","name":"articletitle","label":"Article Title"},{"value":"Neuron","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/S0896-6273(03)00332-5","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"converted-article","name":"content_type","label":"Content Type"},{"value":"Copyright \u00a9 2003 Cell Press. Published by Elsevier Inc. All rights reserved.","name":"copyright","label":"Copyright"}]}}