A neural model of how horizontal and interlaminar connections of visual cortex develop into adult circuits that carry out perceptual grouping and learning.

A neural model suggests how horizontal and interlaminar connections in visual cortical areas V1 and V2 develop within a laminar cortical architecture and give rise to adult visual percepts. The model suggests how mechanisms that control cortical development in the infant lead to properties of adult cortical anatomy, neurophysiology and visual perception. The model clarifies how excitatory and inhibitory connections can develop stably by maintaining a balance between excitation and inhibition. The growth of long-range excitatory horizontal connections between layer 2/3 pyramidal cells is balanced against that of short-range disynaptic interneuronal connections. The growth of excitatory on-center connections from layer 6-to-4 is balanced against that of inhibitory interneuronal off-surround connections. These balanced connections interact via intracortical and intercortical feedback to realize properties of perceptual grouping, attention and perceptual learning in the adult, and help to explain the observed variability in the number and temporal distribution of spikes emitted by cortical neurons. The model replicates cortical point spread functions and psychophysical data on the strength of real and illusory contours. The on-center, off-surround layer 6-to-4 circuit enables top-down attentional signals from area V2 to modulate, or attentionally prime, layer 4 cells in area V1 without fully activating them. This modulatory circuit also enables adult perceptual learning within cortical area V1 and V2 to proceed in a stable way.

[1]  W. Hardy The Conduction of the Nervous Impulse , 1918, Nature.

[2]  V. Mountcastle Modality and topographic properties of single neurons of cat's somatic sensory cortex. , 1957, Journal of neurophysiology.

[3]  D. Hubel,et al.  Receptive fields, binocular interaction and functional architecture in the cat's visual cortex , 1962, The Journal of physiology.

[4]  D. Hubel,et al.  Shape and arrangement of columns in cat's striate cortex , 1963, The Journal of physiology.

[5]  S Grossberg,et al.  Some nonlinear networks capable of learning a spatial pattern of arbitrary complexity. , 1968, Proceedings of the National Academy of Sciences of the United States of America.

[6]  D. Hubel,et al.  Receptive fields and functional architecture of monkey striate cortex , 1968, The Journal of physiology.

[7]  S. Grossberg On learning and energy-entropy dependence in recurrent and nonrecurrent signed networks , 1969 .

[8]  B. Julesz Foundations of Cyclopean Perception , 1971 .

[9]  J. Pokorny Foundations of Cyclopean Perception , 1972 .

[10]  S. Grossberg Contour Enhancement , Short Term Memory , and Constancies in Reverberating Neural Networks , 1973 .

[11]  S Grossberg,et al.  Some developmental and attentional biases in the contrast enhancement and short term memory of recurrent neural networks. , 1975, Journal of theoretical biology.

[12]  V S Ramachandran,et al.  Global Grouping Overrides Point-to-Point Disparities , 1976, Perception.

[13]  C. Malsburg,et al.  How patterned neural connections can be set up by self-organization , 1976, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[14]  D. Hubel,et al.  Ferrier lecture - Functional architecture of macaque monkey visual cortex , 1977, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[15]  T. Wiesel,et al.  Functional architecture of macaque monkey visual cortex , 1977 .

[16]  Paul C. Letourneau Chemotactic response of nerve fiber elongation to nerve growth factor. , 1978, Developmental biology.

[17]  S. Grossberg Decisions, patterns, and oscillations in nonlinear competitive systems with applications to Volterra-Lotka systems. , 1978, Journal of theoretical biology.

[18]  Stephen Grossberg,et al.  Communication, Memory, and Development , 1978 .

[19]  S. Grossberg Competition, Decision, and Consensus , 1978 .

[20]  Roman Bek,et al.  Discourse on one way in which a quantum-mechanics language on the classical logical base can be built up , 1978, Kybernetika.

[21]  T. Wiesel,et al.  Morphology and intracortical projections of functionally characterised neurones in the cat visual cortex , 1979, Nature.

[22]  R. W. Gundersen,et al.  Neuronal chemotaxis: chick dorsal-root axons turn toward high concentrations of nerve growth factor. , 1979, Science.

[23]  D. Frost,et al.  Effects of visual experience on the maturation of the efferent system to the corpus callosum , 1979, Nature.

[24]  R. W. Gundersen,et al.  Characterization of the turning response of dorsal root neurites toward nerve growth factor , 1980, The Journal of cell biology.

[25]  N. Swindale A model for the formation of ocular dominance stripes , 1980, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[26]  S. Grossberg Biological competition: Decision rules, pattern formation, and oscillations. , 1980, Proceedings of the National Academy of Sciences of the United States of America.

[27]  S. Grossberg,et al.  How does a brain build a cognitive code? , 1980, Psychological review.

[28]  D. Purves,et al.  Elimination of synapses in the developing nervous system. , 1980, Science.

[29]  G M Innocenti,et al.  Growth and reshaping of axons in the establishment of visual callosal connections. , 1981, Science.

[30]  D. Pollen,et al.  Phase relationships between adjacent simple cells in the visual cortex. , 1981, Science.

[31]  L. Palmer,et al.  Receptive-field structure in cat striate cortex. , 1981, Journal of neurophysiology.

[32]  P. Schiller,et al.  Effect of cooling area 18 on striate cortex cells in the squirrel monkey. , 1982, Journal of neurophysiology.

[33]  Stephen Grossberg,et al.  Studies of mind and brain , 1982 .

[34]  N. Swindale,et al.  A model for the formation of orientation columns , 1982, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[35]  Stephen Grossberg,et al.  Absolute stability of global pattern formation and parallel memory storage by competitive neural networks , 1983, IEEE Transactions on Systems, Man, and Cybernetics.

[36]  John H. R. Maunsell,et al.  Hierarchical organization and functional streams in the visual cortex , 1983, Trends in Neurosciences.

[37]  S. Grossberg The quantized geometry of visual space: The coherent computation of depth, form, and lightness , 1982, Behavioral and Brain Sciences.

[38]  A. Rosenfeld,et al.  A Theory of Textural Segmentation , 1983 .

[39]  John H. R. Maunsell,et al.  The visual field representation in striate cortex of the macaque monkey: Asymmetries, anisotropies, and individual variability , 1984, Vision Research.

[40]  R. von der Heydt,et al.  Illusory contours and cortical neuron responses. , 1984, Science.

[41]  J J Hopfield,et al.  Neurons with graded response have collective computational properties like those of two-state neurons. , 1984, Proceedings of the National Academy of Sciences of the United States of America.

[42]  S. Grossberg Outline of A Theory of Brightness, Color, and form Perception , 1984 .

[43]  D Ferster,et al.  Synaptic excitation of neurones in area 17 of the cat by intracortical axon collaterals of cortico‐geniculate cells. , 1985, The Journal of physiology.

[44]  G. Blasdel,et al.  Intrinsic connections of macaque striate cortex: axonal projections of cells outside lamina 4C , 1985, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[45]  S Grossberg,et al.  Neural dynamics of perceptual grouping: Textures, boundaries, and emergent segmentations , 1985, Perception & psychophysics.

[46]  G. Kanizsa Seeing and thinking. , 1985, Acta psychologica.

[47]  M. Stryker,et al.  Binocular impulse blockade prevents the formation of ocular dominance columns in cat visual cortex , 1986, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[48]  R Linsker,et al.  From basic network principles to neural architecture: emergence of orientation-selective cells. , 1986, Proceedings of the National Academy of Sciences of the United States of America.

[49]  R Linsker,et al.  From basic network principles to neural architecture: emergence of spatial-opponent cells. , 1986, Proceedings of the National Academy of Sciences of the United States of America.

[50]  Stephen Grossberg,et al.  Neural dynamics of surface perception: Boundary webs, illuminants, and shape-from-shading , 1987, Comput. Vis. Graph. Image Process..

[51]  P. C. Murphy,et al.  Corticofugal feedback influences the generation of length tuning in the visual pathway , 1987, Nature.

[52]  P. C. Murphy,et al.  Cerebral Cortex , 2017, Cerebral Cortex.

[53]  D. Ferster Spatially opponent excitation and inhibition in simple cells of the cat visual cortex , 1988, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[54]  Kevan A. C. Martin,et al.  A Canonical Microcircuit for Neocortex , 1989, Neural Computation.

[55]  R. von der Heydt,et al.  Mechanisms of contour perception in monkey visual cortex. I. Lines of pattern discontinuity , 1989, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[56]  R. von der Heydt,et al.  Mechanisms of contour perception in monkey visual cortex. II. Contours bridging gaps , 1989, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[57]  T. Wiesel,et al.  Columnar specificity of intrinsic horizontal and corticocortical connections in cat visual cortex , 1989, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[58]  R. Kalil,et al.  Synaptic connections between corticogeniculate axons and interneurons in the dorsal lateral geniculate nucleus of the cat , 1989, The Journal of comparative neurology.

[59]  E. L. Schwartz,et al.  A parametric model for synthesis of cortical column patterns , 1989, International 1989 Joint Conference on Neural Networks.

[60]  H. Ritter,et al.  A principle for the formation of the spatial structure of cortical feature maps. , 1990, Proceedings of the National Academy of Sciences of the United States of America.

[61]  E. Callaway,et al.  Emergence and refinement of clustered horizontal connections in cat striate cortex , 1990, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[62]  Richard Durbin,et al.  A dimension reduction framework for understanding cortical maps , 1990, Nature.

[63]  M. Stryker,et al.  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 , 1991, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[64]  P. Goldman-Rakic,et al.  Preface: Cerebral Cortex Has Come of Age , 1991 .

[65]  R. Born,et al.  Single-unit and 2-deoxyglucose studies of side inhibition in macaque striate cortex. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[66]  C. Gilbert,et al.  Synaptic physiology of horizontal connections in the cat's visual cortex , 1991, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[67]  E. Callaway,et al.  Effects of binocular deprivation on the development of clustered horizontal connections in cat striate cortex. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[68]  D. J. Felleman,et al.  Distributed hierarchical processing in the primate cerebral cortex. , 1991, Cerebral cortex.

[69]  D Sagi,et al.  Where practice makes perfect in texture discrimination: evidence for primary visual cortex plasticity. , 1991, Proceedings of the National Academy of Sciences of the United States of America.

[70]  Stephen Grossberg,et al.  Synchronized oscillations during cooperative feature linking in a cortical model of visual perception , 1991, Neural Networks.

[71]  T. Wiesel,et al.  Targets of horizontal connections in macaque primary visual cortex , 1991, The Journal of comparative neurology.

[72]  J. Lund,et al.  Local circuit neurons of macaque monkey striate cortex: III. Neurons of laminae 4B, 4A, and 3B , 1997, The Journal of comparative neurology.

[73]  W. Singer,et al.  Selection of intrinsic horizontal connections in the visual cortex by correlated neuronal activity. , 1992, Science.

[74]  K. Miller Development of orientation columns via competition between ON- and OFF-center inputs. , 1992, Neuroreport.

[75]  E. Kandel,et al.  Are adult learning mechanisms also used for development? , 1992, Science.

[76]  D. V. van Essen,et al.  Neuronal responses to static texture patterns in area V1 of the alert macaque monkey. , 1992, Journal of neurophysiology.

[77]  K. Obermayer,et al.  Statistical-mechanical analysis of self-organization and pattern formation during the development of visual maps. , 1992, Physical review. A, Atomic, molecular, and optical physics.

[78]  J. Kaas,et al.  Rapid reorganization of cortical maps in adult cats following restricted deafferentation in retina , 1992, Vision Research.

[79]  E R Kandel,et al.  Serotonin-mediated endocytosis of apCAM: an early step of learning-related synaptic growth in Aplysia. , 1992, Science.

[80]  E. Kandel,et al.  Modulation of an NCAM-related adhesion molecule with long-term synaptic plasticity in Aplysia. , 1992, Science.

[81]  P. Kellman,et al.  Strength of visual interpolation depends on the ratio of physically specified to total edge length , 1992, Perception & psychophysics.

[82]  C. Gilbert Horizontal integration and cortical dynamics , 1992, Neuron.

[83]  Patrick Cavanagh,et al.  Depth capture and transparency of regions bounded by illusory and chromatic contours , 1992, Vision Research.

[84]  T. Wiesel,et al.  Receptive field dynamics in adult primary visual cortex , 1992, Nature.

[85]  D. Pollen,et al.  Interneuronal interaction between members of quadrature phase and anti-phase pairs in the cat's visual cortex , 1992, Vision Research.

[86]  L C Katz,et al.  Development of local circuits in mammalian visual cortex. , 1992, Annual review of neuroscience.

[87]  T Poggio,et al.  Fast perceptual learning in visual hyperacuity. , 1991, Science.

[88]  U. Eysel,et al.  Network of GABAergic large basket cells in cat visual cortex (area 18): Implication for lateral disinhibition , 1993, The Journal of comparative neurology.

[89]  Ennio Mingolla,et al.  The role of edges and line-ends in illusory contour formation , 1993, Vision Research.

[90]  Mark W. Cannon,et al.  Spatial interactions in apparent contrast: Individual differences in enhancement and suppression effects , 1993, Vision Research.

[91]  M. Stryker,et al.  Development of orientation selectivity in ferret visual cortex and effects of deprivation , 1993, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[92]  U. Polat,et al.  Lateral interactions between spatial channels: Suppression and facilitation revealed by lateral masking experiments , 1993, Vision Research.

[93]  E. Kandel,et al.  Structural changes accompanying memory storage. , 1993, Annual review of physiology.

[94]  David J. Field,et al.  Contour integration by the human visual system: Evidence for a local “association field” , 1993, Vision Research.

[95]  Michael J. Hawken,et al.  Macaque VI neurons can signal ‘illusory’ contours , 1993, Nature.

[96]  M P Stryker,et al.  Rapid remodeling of axonal arbors in the visual cortex. , 1993, Science.

[97]  D. Heeger Modeling simple-cell direction selectivity with normalized, half-squared, linear operators. , 1993, Journal of neurophysiology.

[98]  K. Obermayer,et al.  Geometry of orientation and ocular dominance columns in monkey striate cortex , 1993, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[99]  George L. Gerstein,et al.  Feature-linked synchronization of thalamic relay cell firing induced by feedback from the visual cortex , 1994, Nature.

[100]  R. Frostig,et al.  Cortical point-spread function and long-range lateral interactions revealed by real-time optical imaging of macaque monkey primary visual cortex , 1994, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[101]  U. Polat,et al.  The architecture of perceptual spatial interactions , 1994, Vision Research.

[102]  L. Optican,et al.  Cortical feedback increases visual information transmitted by monkey parvocellular lateral geniculate nucleus neurons , 1994, Visual Neuroscience.

[103]  K. H. Britten,et al.  Neuronal plasticity that underlies improvement in perceptual performance. , 1994, Science.

[104]  T. Bonhoeffer,et al.  Relationship Between Lateral Inhibitory Connections and the Topography of the Orientation Map in Cat Visual Cortex , 1994, The European journal of neuroscience.

[105]  S Grossberg,et al.  3-D vision and figure-ground separation by visual cortex , 2010, Perception & psychophysics.

[106]  J. C. Anderson,et al.  Polyneuronal innervation of spiny stellate neurons in cat visual cortex , 1994, The Journal of comparative neurology.

[107]  M. Weliky,et al.  Functional mapping of horizontal connections in developing ferret visual cortex: experiments and modeling , 1994, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[108]  Stephen Grossberg,et al.  Rules for the cortical map of ocular dominance and orientation columns , 1994, Neural Networks.

[109]  KD Miller A model for the development of simple cell receptive fields and the ordered arrangement of orientation columns through activity-dependent competition between ON- and OFF-center inputs , 1994, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[110]  M. Dalva,et al.  Rearrangements of synaptic connections in visual cortex revealed by laser photostimulation. , 1994, Science.

[111]  J. Donoghue,et al.  Long-term potentiation of horizontal connections provides a mechanism to reorganize cortical motor maps. , 1994, Journal of neurophysiology.

[112]  C. Gilbert,et al.  Axonal sprouting accompanies functional reorganization in adult cat striate cortex , 1994, Nature.

[113]  Gerald Sommer,et al.  Pattern Recognition by Self-Organizing Neural Networks , 1994 .

[114]  G Westheimer,et al.  A quantitative measure for short-term cortical plasticity in human vision , 1994, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[115]  R. Reid,et al.  Specificity of monosynaptic connections from thalamus to visual cortex , 1995, Nature.

[116]  S. Grossberg,et al.  Cortical Dynamics of 3-D Surface Perception: Binocular and Half-Occluded Scenic Images , 1995 .

[117]  Stephen Grossberg,et al.  Cortical Dynamics of 3-D Figure-Ground Perception of 2-D Pictures , 1995 .

[118]  S. Nelson,et al.  An emergent model of orientation selectivity in cat visual cortical simple cells , 1995, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[119]  H. Jones,et al.  Visual cortical mechanisms detecting focal orientation discontinuities , 1995, Nature.

[120]  Victor A. F. Lamme The neurophysiology of figure-ground segregation in primary visual cortex , 1995, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[121]  Alan N. Gove,et al.  Brightness perception, illusory contours, and corticogeniculate feedback , 1995, Visual Neuroscience.

[122]  C. Koch,et al.  Recurrent excitation in neocortical circuits , 1995, Science.

[123]  M Stemmler,et al.  Lateral interactions in primary visual cortex: a model bridging physiology and psychophysics. , 1995, Science.

[124]  C. Gilbert,et al.  Long-range horizontal connections and their role in cortical reorganization revealed by optical recording of cat primary visual cortex , 1995, Nature.

[125]  C. Gilbert,et al.  Improvement in visual sensitivity by changes in local context: Parallel studies in human observers and in V1 of alert monkeys , 1995, Neuron.

[126]  L C Katz,et al.  Development of horizontal projections in layer 2/3 of ferret visual cortex. , 1996, Cerebral cortex.

[127]  Michael Bach,et al.  The abutting grating illusion , 1996, Vision Research.

[128]  K. Martin,et al.  Excitatory synaptic inputs to spiny stellate cells in cat visual cortex , 1996, Nature.

[129]  W. Singer,et al.  The origin and topography of long-range intrinsic projections in cat visual cortex: a developmental study. , 1996, Cerebral cortex.

[130]  M. Rosa,et al.  Visuotopic reorganization in the primary visual cortex of adult cats following monocular and binocular retinal lesions. , 1996, Cerebral cortex.

[131]  D. Fitzpatrick The functional organization of local circuits in visual cortex: insights from the study of tree shrew striate cortex. , 1996, Cerebral cortex.

[132]  J. Bolz,et al.  Specification of layer-specific connections in the developing cortex. , 1996, Progress in brain research.

[133]  Victor A. F. Lamme,et al.  Contextual Modulation in Primary Visual Cortex , 1996, The Journal of Neuroscience.

[134]  J. Bullier,et al.  Functional interactions between areas V1 and V2 in the monkey , 1996, Journal of Physiology-Paris.

[135]  P. C. Murphy,et al.  Functional morphology of the feedback pathway from area 17 of the cat visual cortex to the lateral geniculate nucleus , 1996, The Journal of neuroscience : the official journal of the Society for Neuroscience.

[136]  B. Finlay,et al.  Cortical target depletion and the developing lateral geniculate nucleus: implications for trophic dependence. , 1996, Cerebral cortex.

[137]  M. Stryker,et al.  The Role of Activity in the Development of Long-Range Horizontal Connections in Area 17 of the Ferret , 1996, The Journal of Neuroscience.

[138]  M. Sur,et al.  Orientation Maps of Subjective Contours in Visual Cortex , 1996, Science.

[139]  S. Grossberg,et al.  Contour Integration Across Polarities and Spatial Gaps: From Local Contrast Filtering to Global Grouping , 1997, Vision Research.

[140]  Stephen Grossberg,et al.  Cortical dynamics of three-dimensional surface perception: Binocular and half-occluded scenic images , 1997, Neural Networks.

[141]  M. Dalva,et al.  Reciprocal interactions between layer 4 and layer 6 cells in ferret visual cortex , 1997 .

[142]  K. Martin,et al.  Map of the synapses onto layer 4 basket cells of the primary visual cortex of the cat , 1997, The Journal of comparative neurology.

[143]  W Singer,et al.  The Perceptual Grouping Criterion of Colinearity is Reflected by Anisotropies of Connections in the Primary Visual Cortex , 1997, The European journal of neuroscience.

[144]  W. D. Ross,et al.  Visual brain and visual perception: how does the cortex do perceptual grouping? , 1997, Trends in Neurosciences.

[145]  S. Grossberg Cortical dynamics of three-dimensional figure-ground perception of two-dimensional pictures. , 1997 .

[146]  S. Grossberg,et al.  Cortical Synchronization and Perceptual Framing , 1997, Journal of Cognitive Neuroscience.

[147]  C. Koch,et al.  Constraints on cortical and thalamic projections: the no-strong-loops hypothesis , 1998, Nature.

[148]  C. Gilbert,et al.  Attention and Perceptual Learning Modulate Contextual Influences on Visual Perception , 1998, Neuron.

[149]  S. Grossberg,et al.  Texture segregation, surface representation and figure–ground separation , 1998, Vision Research.

[150]  J. Alonso,et al.  Functional connectivity between simple cells and complex cells in cat striate cortex , 1998, Nature Neuroscience.

[151]  S. Zucker,et al.  Evidence for boundary-specific grouping , 1998, Vision Research.

[152]  Pieter R. Roelfsema,et al.  Object-based attention in the primary visual cortex of the macaque monkey , 1998, Nature.

[153]  Haim Sompolinsky,et al.  Chaotic Balanced State in a Model of Cortical Circuits , 1998, Neural Computation.

[154]  E M Callaway,et al.  Prenatal Development of Layer-Specific Local Circuits in Primary Visual Cortex of the Macaque Monkey , 1998, The Journal of Neuroscience.

[155]  S. Miyauchi,et al.  Attention-regulated activity in human primary visual cortex. , 1998, Journal of neurophysiology.

[156]  D. Field,et al.  The role of “contrast enhancement” in the detection and appearance of visual contours , 1998, Vision Research.

[157]  U. Polat,et al.  Collinear stimuli regulate visual responses depending on cell's contrast threshold , 1998, Nature.

[158]  L. Finkel,et al.  Extraction of perceptually salient contours by striate cortical networks , 1998, Vision Research.

[159]  W. Newsome,et al.  The Variable Discharge of Cortical Neurons: Implications for Connectivity, Computation, and Information Coding , 1998, The Journal of Neuroscience.

[160]  Marius Usher,et al.  Visual synchrony affects binding and segmentation in perception , 1998, Nature.

[161]  E. Todorov,et al.  A local circuit approach to understanding integration of long-range inputs in primary visual cortex. , 1998, Cerebral cortex.

[162]  Zhaoping Li,et al.  A Neural Model of Contour Integration in the Primary Visual Cortex , 1998, Neural Computation.

[163]  Stephen Grossberg,et al.  A neural network model for the development of simple and complex cell receptive fields within cortical maps of orientation and ocular dominance , 1998, Neural Networks.

[164]  J. M. Hupé,et al.  Cortical feedback improves discrimination between figure and background by V1, V2 and V3 neurons , 1998, Nature.

[165]  S. Grossberg,et al.  Self-Organization of Binocular Disparity Tuning by Reciprocal Corticogeniculate Interactions , 1998, Journal of Cognitive Neuroscience.

[166]  S. Grossberg How does the cerebral cortex work? Learning, attention, and grouping by the laminar circuits of visual cortex. , 1999, Spatial vision.

[167]  J A Marshall,et al.  Models of receptive-field dynamics in visual cortex , 1999, Visual Neuroscience.

[168]  S. Grossberg,et al.  Contrast-sensitive perceptual grouping and object-based attention in the laminar circuits of primary visual cortex , 2000, Vision Research.

[169]  S. Grossberg,et al.  Context-sensitive binding by the laminar circuits of V1 and V2: A unified model of perceptual grouping, attention, and orientation contrast , 2001 .