Ratliff F, Knight B W, Graham N
Proc Natl Acad Sci U S A. 1969 Mar;62(3):733-40. doi: 10.1073/pnas.62.3.733.
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.
神经网络中的侧向抑制通常会减弱对正弦刺激（包括空间和时间刺激）的响应幅度。对于具有突然开始和指数时间衰减以及高斯空间分布（理论计算中常假定的形式）的抑制性影响，在低时间和空间频率下衰减最大。随着频率增加，衰减减小，直到最终被抑制响应的幅度变得等于但从不超过未被抑制响应的幅度。然而，对于在时间上延迟到最大值或在空间上具有偏心最大值的抑制性影响，对某些中间频率的被抑制响应的幅度可能大于未被抑制响应的幅度。这种“放大”的产生是因为延迟和空间分离将网络“调谐”到特定的时间和空间频率；抑制在响应的波谷开启，在波峰关闭，从而倾向于产生尽可能大的幅度。在一种神经网络——鲎侧眼视网膜中观察到了这种放大现象。这些基本原理具有普遍性，并且在任何具有负反馈的系统中都可能出现这些效应。
