Contrast gain control in the lower vertebrate retinas.

Control of contrast sensitivity was studied in two kinds of retina, that of the channel catfish and that of the kissing gourami. The former preparation is dominantly monochromatic and the latter is bichromatic. Various stimuli were used, namely a large field of light, a spot-annulus configuration and two overlapping stimuli of red and green. Recordings were made from horizontal, amacrine, and ganglion cells and the results were analyzed by means of Wiener's theory, in which the kernels are the contrast (incremental) sensitivity. Modulation responses from horizontal cells are linear, in that the waveform and amplitude of the first-order kernels are independent of the depth of modulation. In the N (sustained) amacrine and ganglion cells, contrast sensitivity was low for a large modulation input and was high for a small modulation input, providing an example of contrast gain control. In most of the cells, the contrast gain control did not affect the dynamics of the response because the waveform of the first-order kernels remained unchanged when the contrast sensitivity increased more than fivefold. The signature of the second-order kernels also remained unchanged over a wide range of modulation. The increase in the contrast sensitivity for the second-order component, as defined by the amplitude of the kernels, was much larger than for the first-order component. This observation suggests that the contrast gain control proceeded the generation of the second-order nonlinearity. An analysis of a cascade of the Wiener type shows that the control of contrast sensitivity in the proximal retinal cells could be modeled by assuming the presence of a simple (static) saturation nonlinearity. Such a nonlinearity must exist somewhere between the horizontal cells and the amacrine cells. The functional implications of the contrast gain control are as follows: (a) neurons in the proximal retina exhibit greater sensitivity to input of lower contrast; (b) saturation of a neuronal response can be prevented because of the lower sensitivity for an input with large contrast, and (c) over a large range of modulation depths, the amplitude of the response remains approximately constant.