Direction selectivity in simple cells of primary visual cortex, defined from their spike responses, cannot be predicted using linear models. It has been suggested that the shunting inhibition evoked by visual stimulation is responsible for the nonlinear component of direction selectivity. Cortical inhibition would suppress a neuron's firing when stimuli move in the nonpreferred direction, but would allow responses to stimuli in the preferred direction. Models of direction selectivity based solely on input from the lateral geniculate nucleus, however, propose that the nonlinear response is caused by spike threshold. By extracting excitatory and inhibitory components of synaptic inputs from intracellular records obtained in vivo, we demonstrate that excitation and inhibition are tuned for the same direction, but differ in relative timing. Further, membrane potential responses combine in a linear fashion. Spike threshold, however, quantitatively accounts for the nonlinear component of direction selectivity, amplifying the direction selectivity of spike output relative to that of synaptic inputs.
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