A gain-control model relating nulling results to the duration of dynamic motion aftereffects.

Strength of the motion aftereffect (MAE) is most often quantified by its duration, a high-variance and rather 'subjective' measure. With the help of an automatic gain-control model we quantitatively relate nulling-thresholds, adaptation strength, direction discrimination threshold, and duration of the dynamic MAE (dMAE). This shows how the nulling threshold, a more objective two-alternative forced-choice measure, relates to the same system property as MAE-durations. Two psychophysical experiments to test the model use moving random-pixel-arrays with an adjustable luminance signal-to-noise ratio. We measure MAE-duration as a function of adaptation strength and compare the results to the model prediction. We then do the same for nulling-thresholds. Model predictions are strongly supported by the psychophysical findings. In a third experiment we test formulae coupling nulling threshold, MAE-duration, and direction-discrimination thresholds, by measuring these quantities as a function of speed. For the medium-to-high speed range of these experiments we found that nulling thresholds increase and dMAE-durations decrease about linearly, whereas direction discrimination thresholds increase exponentially with speed. The model description then suggests that the motion-gain decreases, while the noise-gain and model's threshold increase with speed.

Spatio-temporal tuning of motion coherence detection at different luminance levels.

We studied effects of dark adaptation on spatial and temporal tuning for motion coherence detection. We compared tuning for step size and delay for moving random pixel arrays (RPAs) at two adaptation levels, one light adapted (50 cd/m(2)) and the other relatively dark adapted (0.05 cd/m(2)). To study coherence detection rather than contrast detection, RPAs were scaled for equal contrast detection at each luminance level, and a signal-to-noise ratio paradigm was used in which the RPA is always at a fixed, supra-threshold contrast level. The noise consists of a spatio-temporally incoherent RPA added to the moving RPA on a pixel-by-pixel basis. Spatial and temporal limits for coherence detection were measured using a single step pattern lifetime stimulus, in which patterns on alternate frames make a coherent step and are being refreshed. Therefore, the stimulus contains coherent motion at a single combination of step size and delay only. The main effect of dark adaptation is a large shift in step size, slightly less than the adjustment of spatial scale required for maintaining equal contrast sensitivity. A similar change of preferred step size occurs also for scaled stimuli at a light-adapted level, indicating that the spatial effect is not directly linked to dark adaptation, but more generally related to changes in the available low-level spatial information. Dark-adaptation shifts temporal tuning by about a factor of 2. Long delays are more effective at low luminance levels, whereas short delays no longer support motion coherence detection. Luminance-invariant velocity tuning curves, as reported previously [Lankheet, M.J.M., van Doorn, A.J., Bouman, M.A., & van de Grind, W.A. (2000) Motion coherence detection as a function of luminance in human central vision. Vision Research, 40, 3599-3611], result from recruitment of different sets of motion detectors, and an adjustment of their temporal properties.

Slow and fast visual motion channels have independent binocular-rivalry stages.

We have previously reported a transparent motion after-effect indicating that the human visual system comprises separate slow and fast motion channels. Here, we report that the presentation of a fast motion in one eye and a slow motion in the other eye does not result in binocular rivalry but in a clear percept of transparent motion. We call this new visual phenomenon 'dichoptic motion transparency' (DMT). So far only the DMT phenomenon and the two motion after-effects (the 'classical' motion after-effect, seen after motion adaptation on a static test pattern, and the dynamic motion after-effect, seen on a dynamic-noise test pattern) appear to isolate the channels completely. The speed ranges of the slow and fast channels overlap strongly and are observer dependent. A model is presented that links after-effect durations of an observer to the probability of rivalry or DMT as a function of dichoptic velocity combinations. Model results support the assumption of two highly independent channels showing only within-channel rivalry, and no rivalry or after-effect interactions between the channels. The finding of two independent motion vision channels, each with a separate rivalry stage and a private line to conscious perception, might be helpful in visualizing or analysing pathways to consciousness.

Motion coherence detection as a function of luminance level in human central vision.

We studied the changes and invariances of foveal motion detection upon dark adaptation. It is well-documented that dark adaptation affects both spatial and temporal aspects of visual processing. The question we were interested in is how this alters motion coherence detection for moving random texture. To compare motion sensitivity at different adaptation levels, we adjusted the viewing distance for equal detectability of a stationary pattern. At these viewing distances we then measured velocity tuning curves for moving random pixel arrays (RPAs). Mean luminance levels ranged from 50 down to 0.005 cd m-2. Our main conclusion is that foveal velocity tuning is amazingly close to luminance-invariant, down to a level of 0.05 cd m-2. Because different viewing distances, and hence, retinal image sizes were used, we performed two control experiments to assess variations of these two parameters separately. We examined the effects of retinal inhomogeneities using discs of different size and annuli filled with RPAs. Our conclusion is that the central visual field, including the near periphery is still rather homogeneous for motion detection at 0.05 cd m-2, but the fovea becomes unresponsive at the lowest luminance level. Variations in viewing distance had marked effects on velocity tuning, both at the light adapted level and the 0.05 cd m-2 level. The size and type of these changes indicated the effectiveness of distance scaling, and show that deviations from perfect invariance of motion coherence detection were not due to inaccurate distance scaling.

Spatial structure, contrast polarity and motion integration.

It has been shown that in the initial stages of motion processing, the ON and OFF pathways stay more or less separated. There is evidence that this distinction between motion signals from opposite contrast polarities remains at least partly intact in the integration stage of local motion information. At the same time, interactions between the two systems are also apparent. Here we constructed stimuli that contained a constant number of moving checks. The checks were either assigned only one contrast polarity, or contrast polarity was distributed across the checks either randomly or evenly. We investigated how the spatial configuration of the moving stimulus affected direction discrimination thresholds for the different polarity distributions. Our results provide new evidence for contrast-sign-specific integration of local motion signals within areas of limited size, and inhibitory interactions between these separate ON and OFF motion sensor pools.

Motion detection from photopic to low scotopic luminance levels.

In this study we quantify the influence of adaptation luminance on the threshold for direction-detection in coherently moving random-pixel arrays (RPAs). Square RPAs of a constant rms-contrast (35%) were used and we determined their 'critical' or threshold-width Wc. Mean retinal illuminances were varied in 13 steps of 0.5 log unit from the low photopic range (screen luminance 0.3 cd/m2) down to 6 log units attenuation, which appeared to be about the absolute threshold of vision under the conditions of our experiment. Moving RPAs were presented at six retinal locations (0, 3, 6, 12, 24 and 48 degrees) from the fovea to the far periphery in the temporal visual field of the right eye of three experienced observers (the authors). In order to ensure an honest comparison between these very disparate conditions, the spatial dimensions (including speed) were scaled according to the acuity, as measured separately for each of the viewing-conditions and observers. Acuity scaling proves to equate the performance for all eccentricities and luminance levels rather well. The fovea is special, but only in the sense that the absolute threshold for light detection is reahed earlier than in peripheral regions. In all other respects foveal results follow the pattern found for peripheral locations. Two different regimes can be discerned in the data, one for high and one for low speeds. In the low speed range Wc is almost constant, regardless of luminance level or eccentricity. The critical 'crossing-time' Tc for any pixel starting at one end of the stimulus and leaving at the opposite end is therefore inversely proportional to velocity in the low-speed range (time-velocity reciprocity). At medium-to-high speeds Wc increases linearly with velocity, so Tc is constant. This constant (minimum) value of Tc differs between subjects, but in all subjects it increases somewhat with decreasing luminance level, even for our acuity-scaled stimuli. The different behaviour for low and high speeds [reported before for photopic viewing conditions by van de Grind, W. A., van Doorn, A. J., & Koenderink, J. J. (1983. Journal of the Optical Society of America, 73, 1674-1683) and van de Grind, W. A., Koenderink, J. J., & van Doorn A. J. (1986. Vision Research, 26, 797-810)] proves to hold from photopic to low scotopic luminance ranges, provided the stimuli are scaled according to acuity. We draw the general conclusion that movement detection is a very robust process that tolerates extremely low retinal illuminance levels. Moreover, the visual system appears to use the same processing principles in combination with an acuity-scaled architecture under all adaptation states and at all eccentricities.

Integration after adaptation to transparent motion: static and dynamic test patterns result in different aftereffect directions.

One of the many interesting questions in motion aftereffect (MAE) research is concerned with the location(s) along the pathway of visual processing at which certain perceptual manifestations of this illusory motion originate. One such manifestation is the unidirectionality of the MAE after adaptation to moving plaids or transparent motion. This unidirectionality has led to the suggestion that the origin of this MAE might be a single source (gain control) located at, or beyond areas that are believed to be responsible for the integration of motion signals. In this report we present evidence against this suggestion using a simple experiment. For the same adaptation pattern, which consisted of two orthogonally moving transparent patterns with different speeds, we show that the direction of the resulting unidirectional MAE depends on the nature of the test stimulus. We used two kinds of test patterns: static and dynamic. For exactly the same adaptation conditions, the difference in MAE direction between testing with static and dynamic patterns can be as large as 50 degrees. This finding suggests that this MAE is not just a perceptual manifestation of a passive recovery of adapted motion sensors but an active integrative process using the output of different gain controls. A process which takes place after adaptation. These findings are in line with the idea that there are several sites of adaptation along the pathway of visual motion processing and that the nature of the test pattern determines the fate of our perceptual experience of the MAE.

Integration and segregation of local motion signals: the role of contrast polarity.

In the initial stages of visual processing in primates, more or less separated ON and OFF pathways have been shown to exist. There is ample evidence, that this separation includes the initial stages of motion processing. In the present study, experiments were conducted to investigate whether this ON versus OFF distinction persists into the integration stage of local motion information. We constructed stimuli that consisted of clusters of checks with equal contrast polarity, which could be varied in size, and compared them to stimuli with a random polarity distribution. We found that the ON versus OFF distinction remains partly intact, while interactions between the two systems are also apparent. These interactions prove to be highly correlated with the spatial structure of the stimulus. We propose a mechanism of contrast-sign specific integration of local motion signals, after which these separate ON and OFF pools engage in mutually inhibitory interactions.

Motion aftereffect of combined first-order and second-order motion.

When, after prolonged viewing of a moving stimulus, a stationary (test) pattern is presented to an observer, this results in an illusory movement in the direction opposite to the adapting motion. Typically, this motion aftereffect (MAE) does not occur after adaptation to a second-order motion stimulus (i.e. an equiluminous stimulus where the movement is defined by a contrast or texture border, not by a luminance border). However, a MAE of second-order motion is perceived when, instead of a static test pattern, a dynamic test pattern is used. Here, we investigate whether a second-order motion stimulus does affect the MAE on a static test pattern (sMAE), when second-order motion is presented in combination with first-order motion during adaptation. The results show that this is indeed the case. Although the second-order motion stimulus is too weak to produce a convincing sMAE on its own, its influence on the sMAE is of equal strength to that of the first-order motion component, when they are adapted to simultaneously. The results suggest that the perceptual appearance of the sMAE originates from the site where first-order and second-order motion are integrated.

Phototaxic foraging of the archaepaddler, a hypothetical deep-sea species.

An autonomous agent (animat, hypothetical animal), called the (archae) paddler, is simulated in sufficient detail to regard its simulated aquatic locomotion (paddling) as physically possible. The paddler is supposed to be a model of an animal that might exist, although it is perfectly possible to view it as a model of a robot that might be built. The agent is assumed to navigate in a simulated deep-sea environment, where it forages for autoluminescent prey. It uses a biologically inspired phototaxic foraging strategy, while paddling in a layer just above the bottom. The advantage of this living space is that the navigation problem--and hence our model--is essentially two-dimensional. Moreover, the deep-sea environment is physically simple (and hence easy to simulate): no significant currents, constant temperature, completely dark. A foraging performance metric is developed that circumvents the necessity to solve the traveling salesman problem. A parametric simulation study then quantifies the influence of habitat factors, such as the density of prey, and body geometry (e.g., placement, direction and directional selectivity of the eyes) on foraging success. Adequate performance proves to require a specific body geometry adapted to the habitat characteristics. In general, performance degrades gracefully for modest changes of the geometric and habitat parameters, indicating that we work in a stable region of "design space." The parameters have to strike a compromise between, on the one hand, to "see" as many targets at the same time as possible. One important conclusion is that simple reflex-based navigation can be surprisingly efficient. Additionally, performance in a global task (foraging) depends strongly on local parameters such as visual direction tuning, position of the eyes and paddles, and so forth. Behavior and habitat "mold" the body, and the body geometry strongly influences performance. The resulting platform enables further testing of foraging strategies or vision and locomotion theories stemming either from biology or from robotics.

Spatial asymmetries in cat retinal ganglion cell responses.

Enroth-Cugell and Robson (1966) first proposed a classification of retinal ganglion cells into X cells, which exhibit approximate linear spatial summation and largely sustained responses, and Y cells, which exhibit nonlinearities and transient responses. Gaudiano (1992a, 1992b, 1994) has suggested that the dominant characteristics of both X and Y cells can be simulated with a single model simply by changing receptive field profiles to match those of the anatomical counterparts of X and Y cells. He also proposed that a significant component of the spatial nonlinearities observed in Y (and sometimes X) cells can result from photoreceptor nonlinearities coupled with push-pull bipolar connections. Specifically, an asymmetry was predicted in the ganglion cell response to rectangular gratings presented at different locations in the receptive field under two conditions: introduction/withdrawal (on-off) or contrast reversal. When measuring the response to these patterns as a function of spatial phase, the standard difference-of-Gaussians model predicts symmetrical responses about the receptive field center, while the push-pull model predicts slight but significant asymmetry in the on-off case only. To test this hypothesis, we have recorded ganglion cell responses from the optic tract fibers of anesthetized cat. The mean and standard deviations of responses to on-off and contrast-reversed patterns were compared. We found that all but one of the cells that yielded statistically significant data confirmed the hypothesis. These results largely support the theoretical prediction.

Local and global factors affecting the coherent motion of gratings presented in multiple apertures.

Using stimuli composed of two independent gratings viewed through multiple apertures, we investigate a number of parameters affecting the integration of locally ambiguous motions into globally coherent motion. In four experiments, we varied local factors (grating spatial frequency, speed, contrast, duty cycle, orientation) and global factors (degree of similarity and common fate between the gratings, and symmetry in the configuration of the grating pattern) and examined their effects on global motion coherence. Our results, confirming accounts offered by previous investigators, indicate that local competition between motion signals generated by contours (ambiguous) and their line terminations (unambiguous) is important in determining global motion coherence in multiple-aperture stimuli. Our results also indicate that global factors can affect perceived coherence independently of local motion signals, suggesting the involvement of higher-level motion areas and a role for non-motion processes such as those involved in pattern and form perception. Comparing motion coherence with other two-dimensional (2-D) stimuli (plaids) shows that 2-D multiple-aperture stimuli are not analogous and that coherence models derived from plaid stimuli do not account for the data.

Aftereffect of high-speed motion.

A visual illusion known as the motion aftereffect is considered to be the perceptual manifestation of motion sensors that are recovering from adaptation. This aftereffect can be obtained for a specific range of adaptation speeds with its magnitude generally peaking for speeds around 3 deg s-1. The classic motion aftereffect is usually measured with a static test pattern. Here, we measured the magnitude of the motion aftereffect for a large range of velocities covering also higher speeds, using both static and dynamic test patterns. The results suggest that at least two (sub)populations of motion-sensitive neurons underlie these motion aftereffects. One population shows itself under static test conditions and is dominant for low adaptation speeds, and the other is prevalent under dynamic test conditions after adaptation to high speeds. The dynamic motion aftereffect can be perceived for adaptation speeds up to three times as fast as the static motion aftereffect. We tested predictions that follow from the hypothesised division in neuronal substrates. We found that for exactly the same adaptation conditions (oppositely directed transparent motion with different speeds), the aftereffect direction differs by 180 degrees depending on the test pattern. The motion aftereffect is opposite to the pattern moving at low speed when the test pattern is static, and opposite to the high-speed pattern for a dynamic test pattern. The determining factor is the combination of adaptation speed and type of test pattern.

Responses of complex cells in cat area 17 to apparent motion of random pixel arrays.

The characteristics of directionally selective cells in area 17 of the cat are studied using moving random pixel arrays (RPAs) with 50% white and 50% black pixels. The apparent motion stimulus is similar to that used in human psychophysics [Fredericksen et al. (1993). Vision Research, 33, pp. 1193-1205]. We compare motion sensitivity measured with single-step pixel lifetimes and unlimited pixel lifetimes. A motion stimulus with a single-step pixel lifetime contains directional motion energy primarily at one combination of spatial displacement and temporal delay. We recorded the responses of complex cells to different combinations of displacement and delay to describe their spatio-temporal correlation characteristics. The response to motion of RPAs with unlimited lifetime is strongest along the preferred speed line in a delay vs displacement size diagram. When using an RPA with a single-step pixel lifetime, the cells are responsive to a much smaller range of spatial displacements and temporal delays of the stimulus. The maximum displacement that still gives a directionally selective response is larger when the preferred speed of the cell is higher. It is on average about three times smaller than the receptive field size.

Pitfalls in estimating motion detector receptive field geometry.

A number of psychophysical investigations have used spatial-summation methods to estimate the receptive field (RF) geometry of motion detectors by exploring how psychophysical thresholds change with stimulus height and/or width. This approach is based on the idea that an observer's ability to detect motion direction is strongly determined by the relationship between the stimulus geometry (height and width) and the RF of the activated motion detectors. Our results show that previous estimates of RF geometry can depend significantly on stimulus position in the visual field as well as on the stimulus height-to-width ratio. The data further show that RF estimates depend on the stimulus in a manner that is inconsistent with basic predictions derived from current motion detector models. Hence previous estimates of height, width, and height-to-width ratios of motion detector RFs are inaccurate and unreliable. This inaccuracy/unreliability is attributed to a number of sources. These include incorrect fixed-parameter values in model fits, as well as the confounding of physiological spatial summation area through combined use of contrast thresholds and Gaussian-windowed stimuli. A third source of error is an asymmetric variation of spatiotemporal correlation in the stimulus as either its height or width is varied (and the other dimension held constant). Most importantly, a fourth source of unreliability is attributed to the existence of a nonlinear, nonmonotonic distribution of motion detectors in the visual field that has been previously described and is a natural result of visual anatomy.

Spatial and temporal properties of cat horizontal cells after prolonged dark adaptation.

We studied the change of spatial and temporal response properties for cat horizontal (H-) cells during prolonged dark adaptation. H-cell responses were recorded intracellularly in the optically intact, in vivo eye. Spatial and temporal properties were first measured for light-adapted H-cells, followed by a period of dark adaptation, after which the same measurements were repeated. During dark adaptation threshold sensitivity was measured at regular intervals. Stable, long lasting recordings allowed us to measure changes of sensitivity and receptive field characteristics for adaptation periods up to 45 min. Although cat H-cells showed no signs of dark suppression or light sensitization, they remained insensitive in the scotopic range, even after prolonged dark adaptation. Absolute thresholds were in the low mesopic range. The sensitization was brought about by a shift from cone to rod input, and by substantial increases of both spatial and temporal integration upon dark adaptation. The length constant in the light-adapted state was on average about 4 deg. After dark adaptation it was up to a factor of three larger, with a median ratio of 1.85. Response delays, latencies and durations for (equal amplitude) threshold flash responses substantially increased during dark adaptation.

Gain control and hyperpolarization level in cat horizontal cells as a function of light and dark adaptation.

First a model is presented that accurately summarizes the dynamic properties of cat horizontal (H-) cells under photopic conditions as measured in our previous work. The model predicts that asymmetries in response to dark as compared to light flashes are flash-duration dependent. This somewhat surprising prediction is tested and confirmed in intracellular recordings from the optically intact in vivo eye of the cat (Experiment 1). The model implies that the gain of H-cells should be related rather directly to the sustained (baseline) membrane potential. We performed three additional experiments to test this idea. Experiment 2 concerns response vs intensity (R-I-) curves for various flash-diameters and background-sizes with background luminance varying over a 4 log unit range. Results support the assumption of a rather strict coupling between flash sensitivity (gain) and the sustained level of hyperpolarization. In Experiment 3 we investigate this relation for both dark and light flashes given on each of four background light levels. The results suggest that there are fixed minimum and maximum hyperpolarization levels, and that the baseline hyperpolarization for a given illumination thus also sets the available range for dark and light flash-responses. The question then arises whether, or how this changes during dark adaptation, when the rod contribution to H-cell responses gradually increases. The fourth experiment therefore studies the relationship between gain and hyperpolarization level during prolonged dark-adaptation. The results show that the rod contribution increases the polarization range of H-cells, but that the gain and polarization level nevertheless remain directly coupled. H-cell models relying on a close coupling between polarization level and gain thus remain attractive options.

Non-visual information in structure-from-motion.

We examined whether non-visual signals improve visual perception of three-dimensional structure-from-motion. Observers discriminated curvature in quadratic surfaces defined by random dot cinematograms with limited lifetime. They either explored visually a static surface by making head movements that were fed back to the display (HM condition) or they viewed statically the same surface which now rotated (NHM condition). Both conditions showed a clear build-up of performance as lifetime increases, but with different time constants for the HM and NHM condition. A second experiment showed that these differences could not be caused by differences in motion detection for the HM and NHM conditions. We suggest that non-visual information is combined with visual information at a high stage of visual processing, and that it does not mainly serve as input for a retinal stabilization process.

Horizontal cell sensitivity in the cat retina during prolonged dark adaptation.

The effects of dark adaptation on the response properties of ganglion cells have been documented extensively in the cat retina. To pinpoint the different retinal mechanisms that underlie these effects, we studied the response characteristics of cat horizontal (H) cells during prolonged dark adaptation. H-cell responses were recorded intracellularly in the optically intact, in vivo eye. To disentangle rod and cone contributions, sensitivity changes during dark adaptation were tracked with white light and with monochromatic lights that favored either rod or cone excitation. Stable, long-lasting recordings allowed us to measure changes of sensitivity for adaptation periods up to 45 min. Thresholds for white light and 503-nm monochromatic light decreased steadily and in parallel. The maximum increase of sensitivity, after extinguishing a photopic adaptation light, was 1.8 log units only, reached after about 35 min. Sensitivity for 581-nm lights also increased steadily, but at a shallower slope. The steady increase of sensitivity was concomitant with a linear shift in resting membrane potential and with an increase in relative rod contribution to the threshold responses. Even though small-amplitude responses were rod dominated after prolonged dark adaptation, sensitivity to rod signals remained relatively low, compared to sensitivity of cone responses or to the absolute sensitivity of ganglion cells. This suggests that the cone-H-cell pathway plays no role in the dark-adapted cat retina.