Neuronal mechanisms of color categorization in areas V1, V2 and V4 of macaque monkey visual cortex

A landmark study conducted by Berlin and Kay (Basic Color Terms, University of California Press, Berkeley, 1969, pp. 1-12) demonstrates that well-developed languages contain exactly 11 basic color terms. The basic colors (8 chromatic and 3 achromatic) are situated in specific locations of color space, suggesting a fixed relationship between specific hue and luminance. To determine the physiologic origins of the basic colors, we have studied the responses of cells in visual cortical areas V1, V2 and V4 of the behaving macaque monkey, using chromatic and achromatic stimuli of varying luminance. A total of 569 cells (291 from V1, 205 from V2, 73 from V4) were obtained, and classified as 'B' (bright; 43-50% of the total cells in each area), 'D' (dark; 6-12% of the total), and 'B/D' (bright/dark; 27-28% of the total) color or non-color cells according to each cell's color/luminance preference in relation to the neutral gray background. About two thirds of 'B' cells in each area were color specific, whereas the proportion of color cells in 'B/D' and 'D' categories was lower. In all three areas (V1, V2, V4), color cells with preferences for midspectral colors (such as yellow, lime and green) also preferred high luminance levels, while color cells with preferences for endspectral colors (such as red and blue) responded preferentially to luminance levels closer to background. The data provide evidence for categorical color perception within the visual system, as well as providing a physiological basis for the increased saliency of endspectral contours observed at equiluminance in psychophysical studies.