Neuronal Operations in the Visual Cortex

1 The Visual System of Cat and Monkey Compared.- 1.1 The Basic Layout of the Visual System in Cat, Owl Monkey, and Rhesus Monkey.- 1.1.1 The Retina.- 1.1.2 The Optic Chiasm and Optic Tract.- 1.1.3 The Dorsal Lateral Geniculate Nucleus (dLGN).- 1.1.4 Visual Cortex.- 1.1.5 Pulvinar.- 1.1.6 Callosal Connections.- 1.2 Quantitative Aspects of the Retino-Geniculo-Cortical Projections.- 1.2.1 The Overall Numbers of Cells in the Visual Pathway.- 1.2.2 Distribution of Retinal Cell Populations.- 1.2.3 Magnification Factors.- 1.3 Conclusion.- 2 The Visual Cortical Areas of the Cat.- 2.1 Description of the Visual Cortical Areas.- 2.1.1 Area 17: The Prototype of Visual Cortical Areas.- 2.1.2 Areas 18 and 19.- 2.1.3 The Lateral Suprasylvian Areas.- 2.1.4 Areas 20 and 21.- 2.1.5 Additional Visual Areas?.- 2.2 The Levels of Processing in the Visual Cortical System of the Cat.- 2.3 Additional Observations on the Retinotopic Organization in the Primary Complex.- 2.3.1 Variability of the 3 Cortical Maps.- 2.3.2 RF Scatter.- 2.3.3 The 17-18 Border and the Question of the Naso-Temporal Overlap.- 2.3.4 The 18-19 Border and the Question of the Visual Field Islands.- 2.4 Conclusion.- 3 Afferent Projections to Areas 17, 18, 19 of the Cat: Evidence for Parallel Input.- 3.1 The Relay of Retinal Afferents: The Dorsal Lateral Geniculate Nuclear Complex.- 3.2 The Geniculocortical Projection.- 3.3 Functional Streams in the Retino-Geniculocortical Projection.- 3.3.1 Functional Properties of Retinal and Geniculate X, Y, W Cells.- 3.3.2 Correlation with Retinal Morphology.- 3.3.3 Separation of Functional Streams at LGN Level.- 3.3.4 Correlation with LGN Morphological Types.- 3.3.5 Distribution of Functional Streams in dLGN Nuclear Complex.- 3.3.6 Input to Different Areas of Primary Visual Complex.- 3.4 Physiological Identification of the Functional Type of Afferents to Areas 17, 18 and 19.- 3.5 The Termination of Geniculate Afferents in the Visual Cortex.- 3.6 Other Subcortical Afferents: Pulvinar-Lateralis Posterior Complex, Intralaminar Nuclei, Claustrum, and Brainstem.- 3.7 The Ipsilateral Corticocortical Connections.- 3.8 The Connections Through the Corpus Callosum.- 3.9 Conclusion.- 4 Receptive Field Organization in Areas 17, 18 and 19 of the Cat.- 4.1 Twenty Years with the Simple-Complex-Hypercomplex Scheme.- 4.2 Criteria for Classifying Cortical RFs.- 4.2.1 The ON-OFF Overlap or the Parcellation of the RF into Subregions.- 4.2.2 Position Test.- 4.2.3 RF Dimensions.- 4.2.4 End-Stopping or the Hypercomplex Property.- 4.3 The A, B, C, S Scheme.- 4.3.1 Properties and Distribution of Cell Types.- 4.3.2 The S and A Families.- 4.3.3 Responses to Other Stimuli.- 4.4 Correspondence of the A, B, C, S Scheme with Other Classification Schemes.- 4.5 Conclusion.- 5 Parameter Specificity of Visual Cortical Cells and Coding of Visual Parameters.- 5.1 The Tuned Cells as Bandpass Filters: The Multichannel Representation of a Parameter.- 5.2 Are All Tuned Cells Simple (Passive) Bandpass Filters or Are Some of Them Active Filters?.- 5.3 Cells with Thresholds as High-Pass Filters: Single or Multichannel Representation of a Parameter.- 5.4 Conclusion.- 6 Influence of Luminance and Contrast on Cat Visual Cortical Neurons.- 6.1 Contrast-Response Curves Obtained with Sinusoidal Gratings.- 6.2 Contrast-Response Curves Obtained with Slits.- 6.3 The Extreme Contrast Sensitivity at the 18-19 Border.- 6.4 Influence of Contrast and Luminance on Other Response Properties.- 6.5 Conclusion.- 7 Coding of Spatial Parameters by Cat Visual Cortical Neurons: Influence of Stimulus Orientation, Length, Width, and Spatial Frequency.- 7.1 Orientation Tuning of Cortical Cells.- 7.1.1 Definitions and Criteria.- 7.1.2 Quantitative Determinations: Orientation-Response Curves.- 7.1.3 Qualitative Determination: Hand-Plotting.- 7.1.4 Distribution of Preferred Orientations.- 7.1.5 Orientation Columns.- 7.1.6 Conclusion.- 7.2 Influence of Stimulus Length on Cortical Cells.- 7.3 Selectivity of Cortical Neurons for Spatial Frequency and Stimulus Width.- 7.3.1 Selectivity for Spatial Frequency.- 7.3.2 Spatial Frequency and Coding of Stimulus Dimensions.- 7.3.3 Linearity of Cortical Cells.- 7.3.4 The Visual Cortex as a Fourier Analyzer.- 7.3.5 Spatial Frequency: Conclusion.- 7.4 Spatial Parameters: Conclusion.- 8 Coding of Spatio-Temporal Parameters by Cat Visual Cortical Neurons: Influence of Stimulus Velocity Direction and Amplitude of Movement.- 8.1 Influence of Stimulus Velocity.- 8.2 Influence of the Direction of Movement.- 8.3 Influence of Stimulus Movement Amplitude.- 8.4 Conclusion.- 9 Binocular Interactions in Cat Visual Cortical Cells and Coding of Parameters Involved in Static and Dynamic Depth Perception.- 9.1 The Binocularity of Cortical Cells and the Ocular Dominance Scheme.- 9.2 Position Disparity Tuning Curves and the Coding of Static Depth.- 9.3 Orientation Disparity, Another Mechanism for Static Depth Discrimination?.- 9.4 Neuronal Mechanisms Underlying Dynamic Depth Perception (Motion in Depth).- 9.5 Conclusion.- 10 The Output of the Cat Visual Cortex.- 10.1 The Projections of Layer V to the Superior Colliculus, Pons, Pretectum, and Pulvinar-LP Complex.- 10.2 The Projections of Layer VI to the dLGN and the Claustrum.- 10.3 The Commissural Projections.- 10.4 The Associative Corticocortical Projections.- 10.5 Conclusion.- 11 Correlation Between Geniculate Afferents and Visual Cortical Response Properties in the Cat.- 11.1 Electrical Stimulation of the Visual Pathways.- 11.2 The Question of ON or OFF Cell Input to Cortical S Cells.- 11.3 Other Attempts to Identify the LGN Input to Cortical Cells.- 11.4 Conclusion.- 12 Intracortical Mechanisms Underlying Properties of Cat Visual Cortical Cells.- 12.1 The Role of Intracortical Inhibition.- 12.1.1 Orientation Selectivity.- 12.1.2 Direction Selectivity.- 12.1.3 End-Stopping.- 12.1.4 Ocular Dominance.- 12.1.5 Velocity Upper Cut-Off.- 12.1.6 Absence of Response to Two-Dimensional Noise.- 12.2 Properties of the Intracortical Inhibitions.- 12.3 The Structural Counterpart of Inhibitions.- 12.4 Conclusion.- 13 Non-Visual Influences on Cat Visual Cortex.- 13.1 Non-Visual Sensory Inputs to the Visual Cortex.- 13.2 Influence of Eye Movements on Visual Cortical Cells.- 13.3 The Influence of Sleep and Anesthesia.- 14 Response Properties of Monkey Striate Neurons.- 14.1 Retinotopic Organization of Area 17.- 14.2 The Input-Output Relations of Monkey Striate Cortex.- 14.3 Receptive Field Organization and Size.- 14.4 Color Specificity in Monkey Striate Cortex.- 14.5 Influence of Light Intensity and Contrast on Monkey Striate Neurons.- 14.6 Influence of Spatial Parameters.- 14.7 Influence of Spatio-Temporal Parameters.- 14.8 Ocular Dominance Distribution and Depth Sensitivity.- 14.9 Columnar Organization and Functional Architecture of Striate Cortex.- 14.10 Correlation Between Response Properties and Afferent Input.- 14.11 Conclusion.- 15 Conclusion: Signification of Visual Cortical Function in Perception.- 15.1 Operating Principles in Cat Visual Cortex.- 15.1.1 Retinotopic Organization.- 15.1.2 Filtering.- 15.1.3 "Columnar" Organization.- 15.1.4 Distributed Processing in the Primary Complex.- 15.1.5 Changes with Eccentricity.- 15.1.6 Parallel Streams Within each Area.- 15.2 The Cat and Monkey Visual Cortex as a Model: The Question of the Relationship Between Animal Physiology and Human Visual Perception.- 15.3 The Role of the Primary Visual Cortex in Visual Perception: The Significance of Parameter Specificities for Object Recognition.- References.