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.Key words: color; monkeys; population coding; primary visual cortex; surfaces; VSDI Introduction "… space and color are not distinct elements but, rather, are interdependent aspects of a unitary process of perceptual organization." (Kanizsa, 1979).The above quotation from Kanizsa's (1979) book guides our work on the neural basis of color perception. The brain needs to construct a color signal to recover the reflective properties of surfaces. Therefore, the neural mechanisms of color perception must make comparisons of the color signals from different locations in the visual image; they must take into account the spatial layout of the scene (Delahunt and Brainard, 2004; Shevell and Kingdom, 2008). It is not known yet in detail how the brain integrates form and color but many scientists who investigated the problem concluded that the primary visual cortex (V1) plays an important role (Johnson et al., 2001(Johnson et al., , 2008 Friedman et al., 2003;Wachtler et al., 2003; Hurlbert and Wolf, 2004).Many investigators have reported the existence of colorresponsive neurons in V1 of macaque monkeys (Thorell et al., 1984; Victor et al., 1994; Leventhal et al., 1995;Johnson et al., 2001; Friedman et al., 2003). Most of the color-sensitive neurons in V1 are double-opponent cells; they are orientation-tuned and respond best to intermediate spatial frequency gratings or to sharp edges in the visual image (ϳ30 -40% of V1 cells). Doubleopponent cells were shown to be sensitive to achromatic luminance patterns as well as to color patterns. Single-opponent cells Received March 29, 2015; revised July 18, 2015; accepted July 22, 2015. Author contributions: S.Z., R.S....