On visual adaptation: II. The electroretinogram and the bipolar cells

First a model for theb-wave of the electroretinogram is given. The essential part of the model is the diffusion into the rod-bipolar synapse of a transmitter substance, followed by the induction of an inhibitor. Making use of this model, adaptation to an illumination too weak to cause of significant decrease in the concentration of visual pigment is interpreted as due to a decreased effectiveness of the rod impulse in exciting the bipolar cell. The disparity between threshold changes for very small test spots and for relatively large spots is explained simply, without invoking any additional physiological mechanisms. This research was supported in part by the United States Air Force through the Air Force Office of Scientific Research of the Air Research Development Command under Contract No. AF(638)-414 and in part by the United States Public Health Service Training Grant 2G-833.     

On the retinal basis of visual adaptation

Summary Visual Adaptation poses numerous problems: in some situations bleached pigment and background lights exert equivalent effects on the visual threshold, while in other situations they do not. Adaptation in man spans a range of more that 10 log units, yet receptor polarization—if this be considered as a measure of the response—saturates over 3 or 4 log units in various animals, including primates. To resolve problems such as these, a model is proposed for the responses and interactions of retinal cells, especially receptor, horizontal, and bipolar cells. It is postulated that receptors can give an output either in terms of ionic flux and polarization changes or chemical synaptic activity. These can both contribute to retinal thresholds, which will also depend, however, on the level of bleached pigment in receptors, as well as on processes in the neural network of the retina. The horizontal cells transmit a signal only if the transmission line is opened at each end through synaptic activity. This opening of the line occurs in response to light stimuli, as distinct from those due to bleaching. Bipolar cells integrate the receptor output with the surround inhibition of the horizontal cells. The amacrine cells perform a second difference operation with respect to rates of change. It is argued that the model is consistent with experimental data on cellular and membrane responses in the retina and that various psychophysical data can be explained on this basis. The objective of this paper is to try to understand the responses of an organ in terms of cell responses and interactions. Specifically, the organ here is the eye and the cells are the neurons in the retina.     

Measurement of visual channels by contrast adaptation.

Inspection of a high-contrast grating pattern affects our ability to detect patterns that are similar. This technique can be used to infer the underlying mechanisms of the visual system. By using this technique, measurements of the bandwidth of orientation channels are taken for different levels of adapting contrast and adapting duration. If the threshold elevation is plotted as the difference between the unadapted and adapted threshold in decibels, then the orientation bandwidth is invariant if taken at some fraction of the maximum elevation. This results from the fact that, as the orientation difference between the adapting and test patterns increases, the function relating threshold elevation to adapting contrast reduces in slope. These data contradict the often-used 'equivalent contrast transformation' (in which the fall off in the adaptation effect with respect to orientation is expressed in terms of an equivalent reduction in adapting contrast) as this would produce quite different bandwidths at different adapting contrasts. The data also address the issue of the neuronal mechanisms of adaptation.     

On perceptual analyzers underlying visual texture discrimination: Part I

We have found a class of feature detectors, based on the quasi-collinearity of dots, which result in visual texture discrimination even when second order statistics are equal. This degenerate counterexample to the Julesz conjecture on effortless texture discrimination has supplied the key to a simple theory of texture discrimination. Accordingly, effortless texture discrimination is based on two classses of perceptual detectors: Class A, those that measure differences in second-order (dipole) statistics; Class B, those that can still detect statistical differences in some features when second-order statistics are kept identical; for instance, the quasi-collinearity of adjacent dipoles. The difference thresholds (tuning curves) for the perceptual dipole and quasi-collinearity detectors have been determined. These texture pairs were generated by a method that creates micropatterns with iso-dipole duals from 4 disks. The extension of this 4-disk method to 5 and more disks with iso-dipole duals permits the search for other kinds of perceptual detectors and will be discussed in Part II.     

Photochemistry of the primary event in short-wavelength visual opsins at low temperature.

Two short-wavelength cone opsins, frog (Xenopus laevis) violet and mouse UV, were expressed in mammalian COS1 cells, purified in delipidated form, and studied using cryogenic UV-vis spectrophotometry. At room temperature, the X. laevis violet opsin has an absorption maximum at 426 nm when generated with 11-cis-retinal and an absorption maximum of 415 nm when generated with 9-cis-retinal. The frog short-wavelength opsin has two different batho intermediates, one stable at 30 K (lambda(max) approximately 446 nm) and the other at 70 K (lambda(max) approximately 475 nm). Chloride ions do not affect the absorption maximum of the violet opsin. At room temperature, mouse UV opsin has an absorption maximum of 357 nm, while at 70 K, the pigment exhibits a bathochromic shift to 403 nm with distinct vibronic structure and a strong secondary vibronic band at 380 nm. We have observed linear relationships when analyzing the energy difference between the initial and bathochromic intermediates and the normalized difference spectra of the batho-shifted intermediates of rod and cone opsins. We conclude that the binding sites of these pigments change from red to green to violet via systematic shifts in the position of the primary counterion relative to the protonated Schiff base. The mouse UV cone opsin does not fit this trend, and we conclude that wavelength selection in this pigment must operate via a different molecular mechanism. We discuss the possibility that the mouse UV chromophore is initially unprotonated.     

Predictive properties of visual adaptation

What humans perceive depends in part on what they have previously experienced. After repeated exposure to one stimulus, adaptation takes place in the form of a negative correlation between the current percept and the last displayed stimuli. Previous work has shown that this negative dependence can extend to a few minutes in the past, but the precise extent and nature of the dependence in vision is still unknown. In two experiments based on orientation judgments, we reveal a positive dependence of a visual percept with stimuli presented remotely in the past, unexpectedly and in contrast to what is known for the recent past. Previous theories of adaptation have postulated that the visual system attempts to calibrate itself relative to an ideal norm or to the recent past. We propose instead that the remote past is used to estimate the world's statistics and that this estimate becomes the reference. According to this new framework, adaptation is predictive: the most likely forthcoming percept is the one that helps the statistics of the most recent percepts match that of the remote past.     

Motion adaptation distorts perceived visual position

After an observer adapts to a moving stimulus, texture within a stationary stimulus is perceived to drift in the opposite direction-the traditional motion aftereffect (MAE). It has recently been shown that the perceived position of objects can be markedly influenced by motion adaptation. In the present study, we examine the selectivity of positional shifts resulting from motion adaptation to stimulus attributes such as velocity, relative contrast, and relative spatial frequency. In addition, we ask whether spatial position can be modified in the absence of perceived motion. Results show that when adapting and test stimuli have collinear carrier gratings, the global position of the object shows a substantial shift in the direction of the illusory motion. When the carrier gratings of the adapting and test stimuli are orthogonal (a configuration in which no MAE is experienced), a global positional shift of similar magnitude is found. The illusory positional shift was found to be immune to changes in spatial frequency and to contrast between adapting and test stimuli-manipulations that dramatically reduce the magnitude of the traditional MAE. The lack of sensitivity for stimulus characteristics other than direction of motion suggests that a specialized population of cortical neurones, which are insensitive to changes in a number of rudimentary visual attributes, may modulate positional representation in lower cortical areas.     

Factors in visual acuity: I. Neural inhibition and the visual perception of contours

Interpretations of the mechanisms of perception of contours or of Mach bands have stressed either the role of various spatial derivatives of light intensity at the retina or the importance of various forms of inhibitory effects between neighboring retinal elements. Evidence is presented here in support of the latter type of interpretation. It is considered that the brightness contrast and perceived contours arise from neural elements, each of which is stimulated in proportion to the intensity of photo-receptor excitation at a point of the retina and inhibited in proportion to the mean intensity in some neighborhood of that point. The role of the spatial derivatives is best seen as a particular manifestation of the inhibitory mechanism. Predictions based upon this hypothesis appear to be consistent with experimentally observed evidence. This research was supported by the United States Air Force through the Air Force Office of Scientific Research of the Air Research and Development Command under Contract No. AF 18(600)-1454. Reproduction in while or in part is permitted for any purpose of the United States Government.     

Work of the human visual system. I. Adequate visual stimulus]

It is shown that the adequate stimulus permitting to detect the presence of colour differentiation in the visual field is the change of relative space-time differences of light actions in different retinal points. Differences only in space or only in time are not sufficient for perception.     

The dynamics of visual adaptation to faces

Several recent demonstrations using visual adaptation have revealed high-level aftereffects for complex patterns including faces. While traditional aftereffects involve perceptual distortion of simple attributes such as orientation or colour that are processed early in the visual cortical hierarchy, face adaptation affects perceived identity and expression, which are thought to be products of higher-order processing. And, unlike most simple aftereffects, those involving faces are robust to changes in scale, position and orientation between the adapting and test stimuli. These differences raise the question of how closely related face aftereffects are to traditional ones. Little is known about the build-up and decay of the face aftereffect, and the similarity of these dynamic processes to traditional aftereffects might provide insight into this relationship. We examined the effect of varying the duration of both the adapting and test stimuli on the magnitude of perceived distortions in face identity. We found that, just as with traditional aftereffects, the identity aftereffect grew logarithmically stronger as a function of adaptation time and exponentially weaker as a function of test duration. Even the subtle aspects of these dynamics, such as the power-law relationship between the adapting and test durations, closely resembled that of other aftereffects. These results were obtained with two different sets of face stimuli that differed greatly in their low-level properties. We postulate that the mechanisms governing these shared dynamics may be dissociable from the responses of feature-selective neurons in the early visual cortex.     

Texture density adaptation and visual number revisited

Burr and Ross [1] have recently proposed that the visual dimension of number is itself directly adaptable. The aftereffect they describe is one that my colleagues and I previously used to investigate the perception of texture density [2-4]. Burr and Ross [1] argue that the effect is new because it concerns visual number, rather than texture density, but they did not report critical tests to support this claim. Here, I shall briefly describe the striking similarity between our prior work and that of Burr and Ross [1], and discuss how some of our results rule out Burr and Ross's [1] interpretation that numerosity, and not density, is at play. I shall also provide a new demonstration that confirms that these effects are based on density, using a display that explicitly dissociates density from numerosity. Taken together, this line of arguments suggests that Burr and Ross's [1] recent study may best be thought of as replicating support within a well-established line of work on texture density.     

On the barn owl’s visual pre-attack behavior: I. Structure of head movements and motion patterns

Barn owls exhibit a rich repertoire of head movements before taking off for prey capture. These movements occur mainly at light levels that allow for the visual detection of prey. To investigate these movements and their functional relevance, we filmed the pre-attack behavior of barn owls. Off-line image analysis enabled reconstruction of all six degrees of freedom of head movements. Three categories of head movements were observed: fixations, head translations and head rotations. The observed rotations contained a translational component. Head rotations did not follow Listing’s law, but could be well described by a second-order surface, which indicated that they are in close agreement with Donder’s law. Head translations did not contain any significant rotational components. Translations were further segmented into straight-line and curved paths. Translations along an axis perpendicular to the line of sight were similar to peering movements observed in other animals. We suggest that these basic motion elements (fixations, head rotations, translations along a straight line, and translation along a curved trajectory) may be combined to form longer and more complex behavior. We speculate that these head movements mainly underlie estimation of distance during prey capture.