Local velocity representation: evidence from motion adaptation

Adaptation to a moving visual pattern induces shifts in the perceived motion of subsequently viewed moving patterns. Explanations of such effects are typically based on adaptation-induced sensitivity changes in spatio-temporal frequency tuned mechanisms (STFMs). An alternative hypothesis is that adaptation occurs in mechanisms that independently encode direction and speed (DSMs). Yet a third possibility is that adaptation occurs in mechanisms that encode 2D pattern velocity (VMs). We performed a series of psychophysical experiments to examine predictions made by each of the three hypotheses. The results indicate that: (1) adaptation-induced shifts are relatively independent of spatial pattern of both adapting and test stimuli; (2) the shift in perceived direction of motion of a plaid stimulus after adaptation to a grating indicates a shift in the motion of the plaid pattern, and not a shift in the motion of the plaid components; and (3) the 2D pattern of shift in perceived velocity radiates away from the adaptation velocity, and is inseparable in speed and direction of motion. Taken together, these results are most consistent with the VM adaptation hypothesis.     

Perceived direction of plaid motion is not predicted by component speeds

It has been shown that the perceived direction of a plaid with components of unequal contrast is biased towards the direction of the higher-contrast component [Stone, L. S., Watson, A. B., & Mulligan, J. B. (1990). Effect of contrast on the perceived direction of a moving plaid. Vision Research 30, 1049-1067]. It was proposed that this effect is due to the influence of contrast on the perceived speed of the plaid components. This led to the conclusion that perceived plaid direction is computed by the intersection of constraints (IOC) of the perceived speed of the components rather than their physical speeds. We tested this proposal at a wider range of component speeds (2-16deg/s) than used previously, across which the effect of contrast on perceived speed is seen to reverse. We find that across this range, perceived plaid direction cannot be predicted either by a model which takes the IOC of physical or perceived component speed. Our results are consistent with an explanation of 2D motion perception proposed by [Bowns, L. (1996). Evidence for a feature tracking explanation of why Type II plaids move in the vector sum direction at short durations. Vision Research, 36, 3685-3694.] in which the motion of the zero-crossing edges of the features in the stimulus contribute to the perceived direction of motion.     

Modulatory effects of binocular disparity and aging upon the perception of speed

Two experiments investigated modulatory effects of a surround upon the perceived speed of a moving central region. Both the surround’s depth and velocity (relative to the center) were manipulated. The abilities of younger observers (mean age was 23.1 years) were evaluated in Experiment 1, while Experiment 2 was devoted to older participants (mean age was 71.3 years). The results of Experiment 1 revealed that changes in the perceived depth of a surround (in this case caused by changes in binocular disparity) significantly influence the perceived speed of a central target. In particular, the center’s motion was perceived as fastest when the surround possessed uncrossed binocular disparity relative to the central target. This effect, that targets that are closer than their background are perceived to be faster, only occurred when the center and surround moved in the same directions (and did not occur when center and surround moved in opposite directions). The results of Experiment 2 showed that the perceived speeds of older adults are different: older observers generally perceive nearer targets as faster both when center and surround move in the same direction and when they move in opposite directions. In addition, the older observers’ judgments of speed were less precise. These age-related changes in the perception of speed are broadly consistent with the results of recent neurophysiological investigations that find age-related changes in the functionality of cortical area MT.     

Extrinsic factors in the perception of bistable motion stimuli

When viewing a drifting plaid stimulus, perceived motion alternates over time between coherent pattern motion and a transparent impression of the two component gratings. It is known that changing the intrinsic attributes of such patterns (e.g. speed, orientation and spatial frequency of components) can influence percept predominance. Here, we investigate the contribution of extrinsic factors to perception; specifically contextual motion and eye movements. In the first experiment, the percept most similar to the speed and direction of surround motion increased in dominance, implying a tuned integration process. This shift primarily involved an increase in dominance durations of the consistent percept. The second experiment measured eye movements under similar conditions. Saccades were not associated with perceptual transitions, though blink rate increased around the time of a switch. This indicates that saccades do not cause switches, yet saccades in a congruent direction might help to prolong a percept because (i) more saccades were directionally congruent with the currently reported percept than expected by chance, and (ii) when observers were asked to make deliberate eye movements along one motion axis, this increased percept reports in that direction. Overall, we find evidence that perception of bistable motion can be modulated by information from spatially adjacent regions, and changes to the retinal image caused by blinks and saccades.     

Luminance texture increases perceived speed

Previous psychophysical experiments have demonstrated that various factors can exert a considerable influence on the apparent velocity of visual stimuli. Here, we investigated the effects of superimposing static luminance texture on the apparent speed of a drifting grating. In Experiment 1, we demonstrate that superimposing static luminance texture on a drifting luminance modulated grating can produce an increase in perceived speed. This supports the hypothesis that texture changes perceived speed by providing landmarks to assess relative motion. In Experiment 2, we showed that contrary to static luminance texture, dynamic luminance texture did not increase perceived speed. This demonstrates that texture must provide reliable spatial landmarks in order to generate an increase in perceived speed. The results of Experiment 3 demonstrate that perceived speed depends on the size of the area covered by texture. This suggests that luminance texture and the motion stimulus interacted with each other over a limited spatial scale and that these local responses are then pooled to determine the speed of the motion stimulus. In Experiment 4, we showed that static texture contrast could produce a greater effect than motion stimulus contrast on perceived speed and that these effects could still be observed at brief presentation times. We discuss these findings in the context of models proposed to account for phenomena in the perception of speed.     

Motion direction signals in the primary visual cortex of cat and monkey

When an image feature moves with sufficient speed it should become smeared across space, due to temporal integration in the visual system, effectively creating a spatial motion pattern that is oriented in the direction of the motion. Recent psychophysical evidence shows that such "motion streak signals" exist in the human visual system. In this study, we report neurophysiological evidence that these motion streak signals also exist in the primary visual cortex of cat and monkey. Single neuron responses were recorded for two kinds of moving stimuli: single spots presented at different velocities and drifting plaid patterns presented at different spatial and temporal frequencies. Measurements were made for motion perpendicular to the spatial orientation of the receptive field ("perpendicular motion") and for motion parallel to the spatial orientation of the receptive field ("parallel motion"). For moving spot stimuli, as the speed increases, the ratio of the responses to parallel versus perpendicular motion increases, and above some critical speed, the response to parallel motion exceeds the response to perpendicular motion. For moving plaid patterns, the average temporal tuning function is approximately the same for both parallel motion and perpendicular motion; in contrast, the spatial tuning function is quite different for parallel motion and perpendicular motion (band pass for the former and low pass for the latter). In general, the responses to spots and plaids are consistent with the conventional model of cortical neurons with one rather surprising exception: Many cortical neurons appear to be direction selective for parallel motion. We propose a simple explanation for "parallel motion direction selectivity" and discuss its implications for the motion streak hypothesis. Taken as a whole, we find that the measured response properties of cortical neurons to moving spot and plaid patterns agree with the recent psychophysics and support the hypothesis that motion streak signals are present in V1.     

Analysis of the motion of 2-dimensional patterns: evidence for a second-order process.

The sum of two differently orientated moving sinusoidal gratings of similar spatial frequency, contrast, and velocity appears as a single coherent "plaid" pattern. The visual system is thought to analyse the motion of plaids in two stages, first analysing the motion of the (1-D) components, and then calculating a speed and direction which is consistent with those 1-D motions. We studied the apparent direction of motion of plaids made by adding two components that had the same spatial frequency and contrast, and were symmetrically oriented about the vertical axis. The gratings moved in jumps, and we studied the effect of varying the size of the jump, the angle between the component gratings, and the temporal interval between the jumps, on the perceived direction of motion. When the size of the jumps was increased to 3/8 of their spatial period, the perceived direction of motion of the plaid pattern reversed, although if one component were presented alone, its direction of movement did not reverse. Reversed motion of this type was consistently obtained if the angle between the components was greater than about 140 degrees, if the interval between jumps was at least 25 msec, and if the spatial frequency of the component gratings was less than about 4 c/deg. When the angle between the components was smaller, or the time between jumps was greater, most observers saw normal motion in the direction predicted by the two-stage hypothesis. When the spatial frequency was raised, observers saw no consistent motion.(ABSTRACT TRUNCATED AT 250 WORDS)     

Two-stage analysis of the motion of 2-dimensional patterns, what is the first stage?

The sum of two differently orientated moving sinusoidal gratings of similar spatial frequency, contrast, and velocity appears as a single coherent "plaid" pattern. The visual system is thought to analyse the motion of plaids in two stages, first analysing the motion of the (1-D) components, and then calculating a speed and direction which is consistent with those 1-D motions. We find that the direction of motion of a plaid (components 1.6 c/deg orientated +60 degrees and -60 degrees) can be discriminated at velocities so low that the direction of motion of its components is not discriminable. This finding is not consistent with the "two-stage" hypothesis in the form that it is usually expressed. We suggest that mechanisms sensitive to the motion of local elements in the pattern, such as edges, could also contribute to the first stage of the analysis of plaid motion.     

The oblique plaid effect

Plaids are ambiguous stimuli that can be perceived either as a coherent pattern moving rigidly or as two gratings sliding over each other. Here we report a new factor that affects the relative strength of coherency versus transparency: the global direction of motion of the plaid. Plaids moving in oblique directions are perceived as sliding more frequently than plaids moving in cardinal directions. We term this the oblique plaid effect. There is also a difference between the two cardinal directions: for most observers, plaids moving in horizontal directions cohere more than plaids moving in vertical directions. Two measures were used to quantify the relative strength of coherency vs. transparency: C/[C+T] and RTtransp. Those measures were derived from dynamics data obtained in long-duration trials (>1 min) where observers continually indicated their percept. The perception of plaids is bi-stable: over time it alternates between coherency and transparency, and the dynamics data reveal the relative strength of the two interpretations [Vision Research 43 (2003) 531]. C/[C+T] is the relative cumulative time spent perceiving coherency; RTtransp is the time between stimulus onset and the first report of transparency. The dynamics-based measures quantify the relative strength of coherency over a wider range of parameters than brief-presentation 2AFC methods, and exposed an oblique plaid effect in the entire range tested. There was no interaction between the effect of the global direction of motion and the effect of gratings' orientations. Thus, the oblique plaid effect is due to anisotropies inherent to motion mechanisms, not a bi-product of orientation anisotropies. The strong effect of a plaid's global direction on its tendency to cohere imposes new and important constraints on models of motion integration and transparency. Models that rely solely on relative differences in directions and/or orientations in the stimulus cannot predict our results. Instead, models should take into account anisotropies in the neuronal populations that represent the coherent percept (integrated motion) and those that represent the transparent percept (segmented motion). Furthermore, the oblique plaid effect could be used to test whether neuronal populations supposed to be involved in plaid perception display tuning biases in favor of cardinal directions.     

Multiple gain control processes in contrast-contrast phenomena

Spatial interactions among orientation-tuned gain control processes are presumed to mediate center-surround contrast-contrast phenomena. In this paper, we assess contributions of gain control processes that pool over orientation. We measured the apparent contrast of a luminance-modulated center disk embedded in various modulated surrounds. In all conditions, observers compared the apparent contrast of the test center to an identically modulated disk with no surround. When center and surround are simple, vertical sinusoids and presented in phase, suppression depends upon surround contrast and is marked at high contrasts. When components are presented 180 degrees out of phase, no suppression occurs at any contrast. When a horizontal component is added to the surround, much less suppression occurs. However, strong suppression is reinstated when both center and surround are plaids. Neither of the latter two effects are phase dependent. We suggest that two different sources of gain control are revealed by the simple sinusoidal and the plaid stimuli. One is orientation tuned and phase-dependent. The other pools over all orientations and includes neurons tuned to multiple phases.     

Lower threshold of motion for one and two dimensional patterns in central and peripheral vision.

Lower motion thresholds for discriminating opposing motion directions were compared for one dimensional (grating) and two dimensional (plaid) stimuli in central and peripheral vision. The results were consistent with a two-stage model of motion sensitivity in which threshold-limiting noise occurs at both stages, and the speed as well as the direction of the resultant motion is determined by intersection-of-constraints (IOC) from the component motions. The results do not support a purely geometric interpretation of the IOC model, in which thresholds for plaid stimuli are related to thresholds of component gratings by a geometric factor. Neither do the data favour explanations in which local luminance features (i.e. blobs) are detected and their velocity determined. Monte-Carlo simulations of the two-stage process predict thresholds across variations in component direction, contrast, and visual field eccentricity. Lower motion thresholds for gratings and plaids both follow a saturating function of contrast; the fit between grating and plaid data is improved when the plaid contrast is expressed in terms of the contrast of its components. Although less contrast saturation was found in the periphery, in relative terms, plaid and grating motion thresholds were similar in central and peripheral vision, implying cortical magnifications are similar for mechanisms which process grating and plaid motion.     

Center-surround inhibition deepens binocular rivalry suppression

When dissimilar stimuli are presented to each eye, perception alternates between both images--a phenomenon known as binocular rivalry. It has been shown that stimuli presented in proximity of rival targets modulate the time each target is perceptually dominant. For example, presenting motion to the region surrounding the rival targets decreases the predominance of the same-direction target. Here, using a stationary concentric grating rivaling with a drifting grating, we show that a drifting surround grating also increases the depth of binocular rivalry suppression, as measured by sensitivity to a speed discrimination probe on the rival grating. This was especially so when the surround moved in the same direction as the grating, and was slightly weaker for opposed directions. Suppression in both cases was deeper than a no-surround control condition. We hypothesize that surround suppression often observed in area MT (V5)-a visual area implicated in visual motion perception-is responsible for this increase in suppression. In support of this hypothesis, monocular and binocular surrounds were both effective in increasing suppression depth, as were surrounds contralateral to the probed eye. Static and orthogonal motion surrounds failed to add to the depth of rivalry suppression. These results implicate a higher-level, fully binocular area whose surround inhibition provides an additional source of suppression which sums with rivalry suppression to effectively deepen suppression of an unseen rival target.     

Color and luminance in the perception of 1- and 2-dimensional motion.

An isoluminant color grating usually appears to move more slowly than a luminance grating that has the same physical speed. Yet a grating defined by both color and luminance is seen as perceptually unified and moving at a single intermediate speed. In experiments measuring perceived speed and direction, it was found that color- and luminance-based motion signals are combined differently in the perception of 1-D motion than they are in the perception of 2-D motion. Adding color to a moving 1-D luminance pattern, a grating, slows its perceived speed. Adding color to a moving 2-D luminance pattern, a plaid made of orthogonal gratings, leaves its perceived speed unchanged. Analogous results occur for the perception of the direction of 2-D motion. The visual system appears to discount color when analyzing the motion of luminance-bearing 2-D patterns. This strategy has adaptive advantages, making the sensing of object motion more veridical without sacrificing the ability to see motion at isoluminance.     

Effect of contrast on the perceived direction of a moving plaid

We performed a series of experiments examining the effect of contrast on the perception of moving plaids. This was done to test the hypothesis put forth by Adelson and Movshon (1982) that the human visual system determines the direction of a moving plaid in a two-staged process: decomposition into component motion followed by application of the intersection of constraints rule. Although there is recent evidence that the first tenet of their hypothesis is correct, i.e. that plaid motion is initially decomposed into the motion of the individual grating components (Movshon, Adelson, Gizzi & Newsome, 1986; Welch, 1989), the nature of the second-stage combination rule has not as yet been established. We found that when the gratings within the plaid are of different contrast, the perceived direction is not predicted by the intersection of constraints rule. There is a strong (up to 20 deg) bias in the direction of the higher-contrast grating. A revised model, which incorporates a contrast-dependent weighting of perceived grating speed as observed for 1-D patterns (Thompson, 1982), can quantitatively predict most of our results. We discuss our results in the context of various models of human visual motion processing and of physiological responses of neurons in the primate visual system.     

Evidence for a Feature Tracking Explanation of Why Type II Plaids Move in the Vector Sum Direction at Short Durations

When two moving sinusoidal gratings, with similar spatial frequency, contrast, phase, but different orientation are combined to form a plaid, their perceived direction of motion has been predicted by the intersection of constraints rule (IOC) (Adelson & Movshon, Nature, 300, 523–525, 1982). However, at short durations (60 msec) the direction of perceived motion has been predicted by the vector sum direction for “Type II” plaids (Yo & Wilson, Vision Research, 32, 1, 1992). Type II plaids are the set of plaids where the components are both located on one side of the resultant computed using the IOC rule. Yo and Wilson suggest that the vector sum direction is observed for Type II plaids at short durations because non-Fourier information is not available and direction is computed from Fourier information only. The first experiment in this study replicates the original Yo and Wilson result using similar stimuli but a simpler task; perceived direction was measured using a direction discrimination task instead of the method of adjustment used by Yo and Wilson. The second experiment provides evidence against generalizing the result to all Type II plaids. A systematic set of type II plaids that varied only in terms of the orientation of the second component provided an ideal set because their predicted motion direction followed very different patterns when predicted by the IOC and vector sum computations. The results obtained were predicted more accurately by the IOC than the vector sum. Experiment 3 provides further evidence that movement in the vector sum direction is not a general property of type II plaids. A small change to the velocity of one of the components of a plaid previously perceived in the vector sum direction had the effect of shifting the perceived motion in the IOC direction, despite increasing the difference between the IOC and VS predictions. This result is not consistent with Yo and Wilson's hypothesis that Type II plaids move in the vector sum direction because of a temporal delay between Fourier and non-Fourier information. Computational analysis of the stimuli used in both the current and original experiments revealed a possible explanation of the results in terms of a contribution from local feature tracking rather than a vector sum operation. Copyright © 1996 Elsevier Science Ltd.     

The role of motion streaks in the perception of the kinetic Zollner illusion

In classic geometric illusions such as the Zollner illusion, vertical lines superimposed on oriented background lines appear tilted in the direction opposite to the background. In kinetic forms of this illusion, an object moving over oriented background lines appears to follow a titled path, again in the direction opposite to the background. Existing literature does not proffer a complete explanation of the effect. Here, it is suggested that motion streaks underpin the illusion; that the effect is a consequence of interactions between detectors tuned to the orientation of background lines and those sensing the motion streaks that arise from fast object motion. This account was examined in the present study by measuring motion-tilt induction under different conditions in which the strength or salience of motion streaks was attenuated: by varying object speed (Experiment 1), contrast (Experiment 2), and trajectory/length by changing the element life-time within the stimulus (Experiment 3). It was predicted that, as motion streaks become less available, background lines would less affect the perceived direction of motion. Consistent with this prediction, the results indicated that, with a reduction in object speed below that required to generate motion streaks (< 1.12°/s), Weber contrast (< 0.125) and motion streak length (two frames) reduced or extinguished the motion-tilt-induction effect. The findings of the present study are consistent with previous reports and computational models that directly combine form and motion information to provide an effective determinant of motion direction.     

Induced steady color shifts from temporally varying surrounds

The color appearance of a physically steady central region can appear to vary over time if a surrounding chromatic light varies in time. The induced temporal variation, however, is strongly attenuated at surround temporal frequencies above approximately 3 Hz. At these higher temporal frequencies, the central region appears steady (De Valois et al., 1986). The posited explanation is a cortical low-pass temporal filter. Here, we investigate whether higher temporal-frequency surrounds induce color shifts in the steady appearance of the central test. Surrounds modulated in time along the l or s chromatic direction of MacLeod-Boynton color space were symmetric around equal-energy white (EEW). The temporal frequency of the surround was varied. If observers perceived the central test to be temporally modulating between two points in time, they set two separate matches to the extreme points of this modulation. If the central test appeared steady in time, then color matches were made to this steady appearance. Corroborating previous reports, measurements showed that surround temporal frequencies below approximately 3 Hz induced temporal modulation. At higher temporal frequencies, however, the surround induced steady color shifts, compared to a steady surround at its time average (EEW). The measurements imply that a nonlinear neural process affects chromatic induction from time-varying context.     

Motion Coherence Across Different Chromatic Axes

It has been reported that equiluminant plaid patterns constructed from component gratings modulated along different axes of a cardinal colour space fail to create a coherent impression of two-dimensional motion Krauskopf and Farell (1990). Nature, 348, 328–331. In this paper we assess whether this lack of interaction between cardinal axes is a general finding or is instead dependent upon specific stimulus parameters. Type I and Type II plaids were made from sinusoidal components (1 cpd) each modulated along axes in a cardinal colour space and presented at equivalent perceived contrasts. The spatial angular difference between the two components was varied from 5 to 90 deg whilst keeping the Intersection of Constraints (I.O.C.) solution of the pattern constant. Observers were required to indicate the perceived direction of motion of the pattern in a single interval direction-identification task. We find that: (i) When plaids were made from components modulated along the same cardinal axis, coherent “pattern” motion was perceived at all angular differences. As the angular difference between the components decreased in a Type II plaid, the perceived direction of motion moved closer to the I.O.C. solution and away from that predicted by the vector sum. (ii) A plaid made from components modulated along red-green and blue-yellow cardinal axes (cross-cardinal axis) did not cohere at high angular differences (>30 deg) but had a perceived direction of the fastest moving component. At lower angular differences, however, pattern motion was detected and approached the I.O.C. solution in much the same way as a same-cardinal axis Type II plaid. (iii) A plaid made from a luminance grating and a cardinal chromatic grating (red-green or blue-yellow) failed to cohere under all conditions, demonstrating that there is no interaction between luminance and chromatic cardinal axes. These results indicate that there are conditions under which red-green and blue-yellow cardinal components interact for the purposes of motion detection. Copyright © 1996 Elsevier Science Ltd.     

Cross-domain adaptation reveals that a common mechanism computes stereoscopic (cyclopean) and luminance plaid motion

Across three experiments, this study investigated the visual processing of moving stereoscopic plaid patterns (plaids created with cyclopean components defined by moving binocular disparity embedded in a dynamic random-dot stereogram). Results showed that adaptation to a moving stereoscopic plaid or its components affected the perceived coherence of a luminance test plaid, and vice versa. Cross-domain adaptation suggests that stereoscopic and luminance motion signals feed into a common pattern-motion mechanism, consistent with the idea that stereoscopic motion signals are computed early in the motion processing stream.     

Bivectorial transparent stimuli simultaneously adapt mechanisms at different levels of the motion pathway

The motion aftereffect (MAE) to drifting bivectorial stimuli, such as plaids, is usually univectorial and in a direction opposite to the pattern direction of the plaid. This is true for plaids that are perceived as coherent, but also for other plaids which are seen as transparent for most or all of the adaptation period. The underlying mechanisms of this MAE are still not well understood. In order to assess these mechanisms further, we measured static and dynamic MAEs and their interocular transfer (IOT). Adaptation stimuli were plaids with small (coherent) and large (transparent) angles between the directions of the component gratings and a horizontal grating, which were adjusted in spatial frequency and drift velocity so that the pattern speed and vertical periodicity remained constant. Test stimuli were horizontal static or counterphasing gratings with the same periodicity as the adaptation stimuli. MAE duration was measured for monocular, binocular and IOT conditions. All static MAEs were smallest for the transparent plaid and largest for the grating, while all dynamic MAEs were constant across adaptation stimuli. IOT was twice as big for dynamic MAEs as for static MAEs, and did not vary with the adaptation stimuli. Other adaptation stimuli were plaids that differed in intersection luminance, contrast or spatial frequency, resulting in different amounts of perceived coherence. MAEs and IOT did not vary with perceived coherence. The results suggest that the MAE for bivectorial stimuli consists of low-level adaptation (dependent on local component properties, small IOT), as well as high-level adaptation (dependent on global integrated pattern properties, large IOT), which can be measured independently with static and dynamic test stimuli.