Representation of Color Surfaces in V1: Edge Enhancement and Unfilled Holes.

The neuronal mechanism underlying the representation of color surfaces in primary visual cortex (V1) is not well understood. We tested on color surfaces the previously proposed hypothesis that visual perception of uniform surfaces is mediated by an isomorphic, filled-in representation in V1. We used voltage-sensitive-dye imaging in fixating macaque monkeys to measure V1 population responses to spatially uniform chromatic (red, green, or blue) and achromatic (black or white) squares of different sizes (0.5°-8°) presented for 300 ms. Responses to both color and luminance squares early after stimulus onset were similarly edge-enhanced: for squares 1° and larger, regions corresponding to edges were activated much more than those corresponding to the center. At later times after stimulus onset, responses to achromatic squares' centers increased, partially "filling-in" the V1 representation of the center. The rising phase of the center response was slower for larger squares. Surprisingly, the responses to color squares behaved differently. For color squares of all sizes, responses remained edge-enhanced throughout the stimulus. There was no filling-in of the center. Our results imply that uniform filled-in representations of surfaces in V1 are not required for the perception of uniform surfaces and that chromatic and achromatic squares are represented differently in V1. SIGNIFICANCE STATEMENT: We used voltage-sensitive dye imaging from V1 of behaving monkeys to test the hypothesis that visual perception of uniform surfaces is mediated by an isomorphic, filled-in representation. We found that the early population responses to chromatic and achromatic surfaces are edge enhanced, emphasizing the importance of edges in surface processing. Next, we show for color surfaces that responses remained edge-enhanced throughout the stimulus presentation whereas response to luminance surfaces showed a slow neuronal 'filling-in' of the center. Our results suggest that isomorphic representation is not a general code for uniform surfaces in V1.