How do neuronal populations represent concurrent stimuli? We measured population responses in cat primary visual cortex (V1) using electrode arrays. Population responses to two superimposed gratings were weighted sums of the individual grating responses. The weights depended strongly on the relative contrasts of the gratings. When the contrasts were similar, the population performed an approximately equal summation. When the contrasts differed markedly, however, the population performed approximately a winner-take-all competition. Stimuli that were intermediate to these extremes elicited intermediate responses. This entire range of behaviors was explained by a single model of contrast normalization. Normalization captured both the spike responses and the local field potential responses; it even predicted visually evoked currents source-localized to V1 in human subjects. Normalization has profound effects on V1 population responses and is likely to shape the interpretation of these responses by higher cortical areas.
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