Functional Clusters for Shape, Texture, and Motion Encoding in Macaque V2

Macaque primary visual cortex (V1) exhibits exquisite columnar organization, while midlevel area V4 does not. Here we investigated the functional organization and representational bases of intervening area V2 in three macaques (one male, two females) with high-density Neuropixels recordings and a variety of visual stimuli-shape, texture, drifting grating, and translational motion patches. We observed dense clusters of similarly tuned neurons often spanning ∼500 µm, consistent with a columnar structure. In terms of representational bases, V2 responses were largely explained by stimulus features based on local image statistics: shape tuning is well modeled by a linear combination of orientation filters, and direction selectivity is stronger with surface compared with object motion, in striking contrast to V4. Overall, our results support the progression from columns to sparse clusters as neuronal representations transform from encoding local features and feature conjunctions in V1/V2 to a high-dimensional object-based code in V4.