Visual cortical mechanisms of perceptual grouping: interacting layers, networks, columns, and maps

The visual cortex has a laminar organization whose circuits form functional columns in cortical maps. How this laminar architecture supports visual percepts is not well understood. A neural model proposes how the laminar circuits of V1 and V2 generate perceptual groupings that maintain sensitivity to the contrasts and spatial organization of scenic cues. The model can decisively choose which groupings cohere and survive, even while balanced excitatory and inhibitory interactions preserve contrast-sensitive measures of local boundary likelihood or strength. In the model, excitatory inputs from lateral geniculate nucleus (LGN) activate layers 4 and 6 of V1. Layer 6 activates an on-center off-surround network of inputs to layer 4. Together these layer 4 inputs preserve analog sensitivity to LGN input contrasts. Layer 4 cells excite pyramidal cells in layer 2/3, which activate monosynaptic long-range horizontal excitatory connections between layer 2/3 pyramidal cells, and short-range disynaptic inhibitory connections mediated by smooth stellate cells. These interactions support inward perceptual grouping between two or more boundary inducers, but not outward grouping from a single inducer. These boundary signals feed back to layer 4 via the layer 6-to-4 on-center off-surround network. This folded feedback joins cells in different layers into functional columns while selecting winning groupings. Layer 6 in V1 also sends top-down signals to LGN using an on-center off-surround network, which suppresses LGN cells that do not receive feedback, while selecting, enhancing, and synchronizing activity of those that do. The model is used to simulate psychophysical and neurophysiological data about perceptual grouping, including various Gestalt grouping laws.