On tuning and amplification by lateral inhibition.

Lateral inhibition in a neural network generally attenuates the amplitudes of the responses to sinusoidal stimuli-both spatial and temporal. For an inhibitory influence with an abrupt onset and an exponential decay in time, and with a Gaussian distribution in space (the forms often assumed in theoretical calculations), the attenuation is greatest at low temporal and spatial frequencies. The attenuation diminishes with increasing frequencies until ultimately the amplitudes of inhibited responses become equal to, but never exceed, the amplitudes of the uninhibited. For an inhibitory influence with a delay to the maximum in time or with eccentric maxima in space, however, the amplitudes of inhibited responses to certain intermediate frequencies may be greater than those of the uninhibited respones. This "amplification" results because the delay and the spatial separation "tune" the network to particular temporal and spatial frequencies; the inhibition is turned on at the trough of the response and off at the crest, thus tending to produce the greatest possible amplitude. The amplification has been observed in one neural network, the retina of the lateral eye of Limulus. The basic principles are general, and the effects may be expected in any system with negative feedback.