The document discusses visual adaptation and retinal gain controls. It explains how retinal neurons can detect contrast through temporal modulation caused by eye movements, and how receptive field surrounds make neurons more sensitive to patterns with narrow spatial gradients. It also discusses how brightness constancy relies on local contrast near object borders rather than comparing light across the entire retina.
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Field ), I /I'RB LB Bockground
The document discusses visual adaptation and retinal gain controls. It explains how retinal neurons can detect contrast through temporal modulation caused by eye movements, and how receptive field surrounds make neurons more sensitive to patterns with narrow spatial gradients. It also discusses how brightness constancy relies on local contrast near object borders rather than comparing light across the entire retina.
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VISUAL ADAPTATION AND RETINAL GAIN CONTROLS 271
(a) expect deviations from brightness constancy.
/I-Ro = Lo Receptive~ This line of reasoning leads to the conclusion that Field~),i~ /I'RB=LB the detection of contrast depends on light adaptation. A different view is that the dependence Bockground of retinal responses on contrast is a consequence of the spatially antagonistic c e n t e r - s u r r o u n d organization of receptive fields (discussed below in Appendix 2), but we believe this view to be incorrect. If there are eye movements, spatial contrast will be " s e e n " by the highly localized (b) Receptive receptive fields of retinal neurons as successive Field,) contrast, or in other words as temporal modulation of the amount of light falling on those receptive Background fields (Helmholtz, 1909). As shown above, in this situation, the mechanisms of adaptation together with the compact nature of the receptive fields FIG. 3. The invariance of contrast with changes in level of produce a response dependent on contrast without illumination, and how this is sensed by a visual cell's receptive any requirement for a spatially antagonistic field. This picture shows the luminance profile of an object surround. The purpose of having receptive field (with reflectance R o) on a background (with reflectance RB) when both are illuminated uniformly, with illumination L surrounds probably is to make retinal neurons more The position of the receptive field of an individual retinal sensitive to patterns with narrow spatial gradients, cell is indicated by the shaded rectangle. In (a), the visual or high spatial frequencies, than to coarse patterns cell is "looking" at the background. In Co), an eye movement has carried the receptive field of the cell onto the object. or diffuse light. Put another way, with light Equation (5) in the text demonstrates that this cell will respond adaptation but without receptive field surrounds, to the Weber contrast of the object on the background, i.e. retinal neurons would respond to contrast. (Ro-RB)/RB, which is independent of illumination L As However, with receptive field surrounds but without explained in the text, in order for this calculation to work the cell must be influenced by a gain control which adjusts adaptation, the retina would be unable to respond the cell's gain to be reciprocal with background illumination. to contrast; it would saturate quickly (see Section 1.2.2.). This point of view is similar to that put forward by Whittle and Challands (1969). proportional to the reciprocal of the value of the illumination at which the cell's receptive field has Consideration of Mach's statement together with been " l o o k i n g " up to the moment it crosses the more recent work on brightness constancy under border. This inference is suggested by Weber's Law, conditions of non-uniform illumination leads to an equation (2). Thus the signal in our hypothetical interesting conclusion about the spatial spread of n e u r o n will equal the stimulus I'Ro-I'RB adaptation. Mach, and others after him (Hering, multiplied by the gain, K/(I. RB), where K is a pro- 1920; Helson, 1964), have supposed that the eye portionality constant. The conclusion is that what compared, "the quantity of light on the entire retina the visual system uses to characterize an object is and the image of the paper", in order to calculate a signal the brightness of the paper. This is not correct. The first correction is that brightness depends on a local S = K" (I'Ro-I" RB)/I" RB (5) phenomenon, the contrast near the border of an object, not on the quantity of light in the image of which is proportional to the Weber contrast. In this the object (see Fig. 2). Second, the retina does not case the neural response to the object is invariant compare the quantity of light " o n the entire retina" with respect to changes in the illumination, I. This with the amount of light in the object, as Mach chain of reasoning only holds if the gain of the suggested. As Davidson and Freeman (1965) and retina is reciprocal with background illumination, Land and McCann (1971) have shown, brightness in analogy with Weber's Law. When retinal gain is constant under conditions of spatially non- is not reciprocal with background, one should uniform illumination. We have argued that retinal