Mechanisms of Visual Sensitivity: Backgrounds and Early Dark Adaptation
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i,%on Res. Vol. 23. No. I?. pp. 1423-3432.1983 0042-6989X3 53.00 + 0.00 Printed III Great Britain. Ail rights reserved Copyright .c 1983Pergamon Press Ltd MECHANISMS OF VISUAL SENSITIVITY: BACKGROUNDS AND EARLY DARK ADAPTATION WILSON S. GEISLER Department of Psychology, University of Texas, Austin, TX 78712. U.S.A. (Received 2 October 1980; in raised form 14 June 1983) Abstract-There is substantial physiological and psychophysical evidence for an adaptation mechanism whose effect, under many circumstances, is equivalent to placing a neutral density filter in front of the eye. Furthermore. this mechanism is of sufficient strength to predict the generalized Weber‘s law for increment thresholds on steady backgrounds. However, it was shown that an additional transient mechanism (with a time-course of around 100 msec) is also needed to account for the inc~ment-threshold results. The effect of this mechanism on increment thresholds during early dark adaptation was parametrically examined. Several models for the transient mechanism were considered. The one best able to account for the results consists of a subtractive inhibitory stage operating prior to a saturating nonlinearity. Light adaptation Dark adaptation Increment thresholds Lateral inhibition Weber’s law INTRODUCTION adaptation” model. The evidence comes from experi- In the cone system, increment and decrement thresh- ments in which flashed-background increment- olds measured against steady backgrounds are ap threshold functions were measured in the fovea under proximately described by Weber’s law over a wide various fixed states of light adaptation. For example. range of background intensities and test-field param- Geisler (1981) measured flashed-background eters. Substantial exceptions to the law occur for test increment-threshold functions 500 msec after the stimuli that are very small and brief (Barlow, 1958; offset of adapting backgrounds. Figure I shows the Geisfer, 1979a) or of high spatial or temporal fre- pattern of results obtained in the study. The solid quencies (Kelly, 1972); but. even under these condi- curve labelled - co is the increment-threshold func- tions the thresholds converge toward Weber’s law as tion obtained for flashed backgrounds in the dark- background intensity is raised. The widespread valid- adapted fovea. The other solid curves represent ity of Weber’s law across both stimulus conditions and species may be due to its usefulness for pattern recognition. In particular, Weber’s law implies that a visual system exhibit pattern constancy-the prop- erty that the set of visible lines and edges within the visual field remains constant across all illumination levels (Hemila, 1977; Geisler. 1978a). Many models for the adaptation processes under- lying Weber’s law have been proposed. These include logarithmic response compression coupled with a constant criterion for detection (Cornsweet and Pin- sker, 1965), power-law response compression with increasing internal variability (Mansfield, 19X), linear transduction with quanta1 fluctuations and sources of correlated neural noise (Triesman, 1966), power-law transduction followed by a multiplicative adaptation factor (Thijssen and Vendrik. 1971), and 0 I 2 3 4 5 nonlinear transduction preceded by a multiplicative LO6 BICKMOUNO adaptation factor (Lythgoe. 1936: Craik, 1938; Geis- Fig. I. The pattern of increment-threshold functions ler. 1975: 1981: Dawis and Purple. 1980). Recently, reported in Geisler (1981). The solid curves are psychophysical evidence has been accumulating for washy-background inc~ment-th~hold functions obtained the latter type of model (Geisler. 197Xb. 1981: Hood, at various fixed levels of adaptation. The dashed curve is the optimal function that can be obtained with multiplicative 1978; Hood ef ui.. 1978). which will be referred to adaptation. The dotted curve shows the actual function that here as the “dark-glasses” or “multiplicative- is obtained. 1423 1424 WILSON S. GEISLEK flashed-background increment-thr~shoId functions steady backgrounds that would be obtained if only obtained for the adapting-background intensities in- multiplicative adaptation were operating. The dotted dicated on the left. The functions obtained at all curve shows the function that is obtained: it is 0.7-l .O adapting-background intensities are roughly identical log units lower than the dashed curve at moderate to in shape and continuously accelerating when plotted high background levels. Changes in the maximum of in log-log coordinates. Furthermore, a function ob- the intensity-response function, as least of the sort tained at any given adapting-background intensity observed in electrophysiological research. are also can be brought into coincidence with a function unable to account for the results. Typically. light obtained at any other intensity by translation along adaptation decreases the maximum of the inten- a 45 line. This is exactly the prediction made by the sity-response function which should cause increment simple multiplicative-adaptation model (Geisler, thresholds to increase rather than decrease. 1978b). There appears to be at least one other factor or Figure 1 is also useful for explaining how multi- mechanism responsible for the increment thresholds plicative adaptation accounts for Weber’s iaw. The obtained with steady adapting backgrounds. Experi- X’s indicate the point on each curve where the ments in which the adapting background is momen- minimum Weber fraction (AZ/l) is reached. AS can be tarily switched off and on just before presentation of seen. the points where the Weber fraction reaches its the increment field (Ikeda and Boynton. 1965: Gei- minimum trace out a straight line of slope one (the sler, 1981) provide further evidence for this additional dashed line). In the standard increment-threshold mechanism. These studies indicate that this mech- experiment the adapting-background intensity and anism has a relatively quick decay time---on the order the background intensity against which threshold is of 100 msec. The purpose of the present experiments measured are identical. Now, if the adapting back- was to further explore this mechanism which is grounds produce just enough multiplicative adapta- apparently involved in the very early phases of light tion for the flashed-background increment-threshold and dark adaptation. curves to shift so that their minimum Weber fractions occur at the adapting-background intensity, then the increment-thresholds must be given by the X’s. In THEOR\ other words. Weber’s law is predicted. The results in Electrophysiological studies. including those listed Geisler ( 198 1) confirm that the increment-threshold in the introduction. have often found that the curves shift sufficiently to yield Weber’s law. flash-response characteristics of retinal neurons in the There is also considerable electrophysiological evi- vertebrate retina are described by the equation dence for the multiplicative adaptation model (Naka I” and Rushton, 1966; Dowling and Ripps, 1971, 1972; R = R,,, -. I” + I,” Glantz. 1972: Grabowski ef ai., 1972; Baylor and Hodgkin. 1974: Norman and Werblin, 1974; Werblin, where I is the intensity of the flash, R,,, is the 1974: Kleinschmidt and Dowling, 1975; Green ef al., maximum electrical response of the neuron. I, is the 1975: Fain, 1976; Norman and Perlman, 1979; Dawis half-saturation constant. and n is an exponent with a and Purple. 1980; Valeton and Norren, 1983). Most value usually between 0.7 and 1.0. As mentioned of these studies measured the effects of either bleaches above, changes in the flash-response characteristic or backgrounds on the intensity-response functions due to bleaching and backgrounds can usually be of retinal neurons. especially receptors. Typically it described by changes in I, (that is. by multiplicative was found that most of the changes in the intensity- adaptation) and by changes in R,,, (the maximum response functions are attributable to multiplicative response). In a previous paper (Geisler. 1981). a adaptation (i.e. to changes in the half-saturation simple model based on equation (I) was shown to constant). It is also typically found that light adapta- account fairly well for the effects of bleaches and tion produces some relatively small decreases in the backgrounds on flashed-background increment- maximum of the intensity-response function. Psycho- threshoId functions. It was assumed that threshold is physically. there is little evidence of such decreases reached when the response R , to the background- (Geisler. 1981); however. one would not expect the plus-increment exceeds the response R to the back- small effects observed in most electrophysiological ground alone by some fixed criterion amount (5. That studies to have much influence on the psychophysical is. threshold is reached when results. Although the multiplicative adaptation observed in R, - R=ii. (2) electrophysiological recordings, and in the appropri- The arguments in equation (1) for R + and R are, ate psychophysical experiments, is sufficient to respectively account for Weber’s law. the actual increment- threshold function obtained for steady backgrounds AI + s’ I i- ;: I, (3) cannot be explained by multiplicative adaptation alone (Geisler. 1981). The dashed curve in Fig. 1 and shows the (optimal) increment-threshold function for Mechanisms of visual sensitivity: backgrounds and early dark adaptation 1425 where I, is the intensity of the adapting background, intensity-r~ponse functions ~flash-response charac- and < and 1’are parameters