Neurons in primary visual cortex (V1) are commonly classified as simple or complex based upon their sensitivity to the sign of stimulus contrast. The responses of both cell types can be described by a general model in which the outputs of a set of linear filters are nonlinearly combined. We estimated the model for a population of V1 neurons by analyzing the mean and covariance of the spatiotemporal distribution of random bar stimuli that were associated with spikes. This analysis reveals an unsuspected richness of neuronal computation within V1. Specifically, simple and complex cell responses are best described using more linear filters than the one or two found in standard models. Many filters revealed by the model contribute suppressive signals that appear to have a predominantly divisive influence on neuronal firing. Suppressive signals are especially potent in direction-selective cells, where they reduce responses to stimuli moving in the nonpreferred direction.
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