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Anatomy/Physiology of Binocular Vision • Goals – Follow the M and P pathway out of primary visual cortex – Answer where binocularly and disparity driven cells appear – Learn a bit about stereopsis – Answer (partially) how an oculocentric neuronal organization gives rise to an egocentric visual perception Parallel Pathways: Magnocellular (M) and Parvocellular (P) • Each pathway is sensitive to specific visual stimuli • Each pathway has its own timing characteristics • Each pathway is NOT strictly parallel! – More of a “Bob ‘N Weave” pathway arrangement Magnocellular (M-pathway) The Table Setter • • • • • Coarse visual form Moving (or modulating)target Processing time: rapid Peripheral fusion Coarse stereopsis Parvocellular (P-pathway) The Details • • • • • Spatial detail Chromatic detail Stationary (or moving slowly) target Processing time: slow Fine stereopsis Parallel Pathways On the Move • Lateral geniculate nucleus • Segregation of P and M pathways into layers (1-2 Magno.; 3-6 Parvo.) • LGN serves as a relay station to primary visual cortex (18) – Where vision will become a conscious event – Where stereopsis and fusion takes place – Where visual and cognitive processing take place Primary Visual Cortex (V1) • Located along calcarine sulcus • M and P pathways continue in different paths as they reach layer 4 of V1 – M pathway to layer 4 Ca – P pathway to layer 4Cb and layer 4A • Organized into ocular dominance zones – Monocular cells in layer 4C – Binocular driven cells outside of layer 4C Parallel Pathways in V1 • M pathway: – From 4Ca to layer 4B in same vertical column (1 mm wide) – From 4B to layers 2/3 in same vertical column (1 mm wide)and neighboring columns Parallel Pathways in V1 • P pathway: – From 4Cb to layers 4A and 3 in same vertical column (1 mm wide) – In layer 3, cytochrome oxidase, a metabolic marker, has dense staining in layer 2/3; absent in layer 4 – Called “blobs” – Although considered “P-cells only”, a significant M-pathway input exists Parallel Pathways in V1 • Blob and interblob regions:a split in the parvocellular pathway • Blob regions are situated in the center of ocular dominance columns – Blob regions: color opponency, low contrast and spatial frequency, not orientation selective – Interblob regions: little color opponency, high contrast and spatial frequency, very orientation selective M and P Pathways In V2 • V2 has areas of high cytochrome oxidase activity in form of thick and thin stripes • M pathways project to thick stripes • P pathway – Blob cells: thin stripes – Interblob cells: inter stripes Other Visual Areas • V2: in area 18, flanking V1 – Thin/inter stripe regions (P pathway) projects to V4 – Thick stripe (M pathway) projects to V3 and MT – Some overlap in response characteristics in V2 due to “cross-talk” between M and P at blob region Other Visual Areas • V3: in area 18 flanking V2 – Receives M pathway input – Output to middle temporal area (MT) – Also output to V4!?! • V4 – Receives P-pathway input from thin/inter stripe regions of V2 – Receives strong Minput Vision Association Areas • Area MT – In parietal lobe – M-pathway input – Output to parietal areas and V4 – Sensitive to motion – Some areas have receptive fields in head-centric coordinates, NOT oculocentric Vision Association Areas • Inferotemporal cortex • Posterior parietal cortex • P-input (V4): fine stereopsis, color vision, fine pattern vision • M-input (MT/V4): coarse stereopsis, low spatial freq., fast flicker and motion • Complex object recognition: faces • Spatial position and object motion Final Words About M/P Pathways • Significant cross-talk in V1, V4 and beyond • Ultimately, these two independent, yet overlapping streams must converge to form unitary perceptions of objects • We do not process the world like a poorly printed photograph, with the colors offset Ocular Dominance Columns • Vertical columns that respond most strongly to one eye • Extends through the full thickness of V1 • Absent in areas outside V1 • Binocular cells outside layer 4C respond predominantly to one eye over the other Orientation Columns • If ocular dominance columns are loaves of bread, orientation selective columns are slices (parallel to pia) • Orientation selectivity is interrupted by blobs Binocular Cells and Stereopsis • Binocular cells in V1 receptive fields for each eye share most characteristics – Corresponding retinal loci – Latency – Size/shape of receptive field Binocular Cells and Stereopsis • Binocular cells in V1 receptive fields for each eye share most characteristics – Corresponding retinal loci – Latency – Size/shape of receptive field • If perfect overlap of receptive fields exist, it argues for a creation of an EGOCENTRIC PERCEPTION early in visual processing • It cannot explain, however, why we are sensitive to binocular disparity (stereopsis) Binocular Disparity • Results from different perspective of each eye to a particular visual target • Neurons tuned to disparity have been found in V1 • Receptive fields for each eye do not PERFECTLY overlap • More prevalent in V2 (75% cells tuned to disparity) • 4 main classifications of disparity tuned cells – Near cells/ Far cells – Excitatory cells tuned to zero disparity – Tuned excitatory – Tuned inhibitory Profiles of Disparity Tuned Cells • Near cells: resp. to targets closer than fixation distance • Far cells: resp. to targets farther than fixation distance • Excitatory cells tuned to zero disparity: narrow peak responses around zero disparity Profiles of Disparity Tuned Cells • Tuned excitatory: stim. by stimuli near zero disparity BUT ON EITHER SIDE/ suppressed by uncorrelated images • Tuned inhibitory: suppressed by stimuli near zero disparity BUT ON EITHER SIDE / stim. by uncorrelated images QuickTime™ and a Video decompressor are needed to see this picture.