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In natural images, some boundaries are de ned by luminance di erences; others are de ned by texture di erences. Most neurons in primary visual cortex (V1) are well-driven by luminance boundaries at the appropriate orientation1-2. Boundaries de ned by di erences in texture, however, are more e ective stimuli for neurons in the secondary visual cortex (V2)3-9. Since the larger receptive elds of V2 are produced by combining the output of V1 neurons10-12, the extraction of texture boundaries by V2 receptive elds must involve computations on its V1 inputs across space. These computations must accomplish a speci c goal – extraction of texture boundaries – while preserving the luminance-boundary information already extracted by V1.

Orientation change detection and orientation pooling in space and time performed by two subpopulations of neurons in V2