Independent aftereffects of attention and motion.

In the motion aftereffect (MAE), a stationary pattern appears to move in the opposite direction to previously viewed motion. Here we report an MAE that is observed for a putatively high level of visual analysis-attentive tracking. These high-level MAEs, visible on dynamic (but not static) tests, suggest that attentive tracking does not simply enhance low-level motion signals but, rather, acts at a subsequent stage. MAEs from tracking (1) can overrule competing MAEs from adaptation to low-level motion, (2) can be established opposite to low-level MAEs seen on static tests at the same location, and (3), most striking, are specific to the overall direction of object motion, even at nonadapted locations. These distinctive properties suggest MAEs from attentive tracking can serve as valuable probes for understanding the mechanisms of high-level vision and attention.

The motion aftereffect.

The motion aftereffect is a powerful illusion of motion in the visual image caused by prior exposure to motion in the opposite direction. For example, when one looks at the rocks beside a waterfall they may appear to drift upwards after one has viewed the flowing water for a short period-perhaps 60 seconds. The illusion almost certainly originates in the visual cortex, and arises from selective adaptation in cells tuned to respond to movement direction. Cells responding to the movement of the water suffer a reduction in responsiveness, so that during competitive interactions between detector outputs, false motion signals arise. The result is the appearance of motion in the opposite direction when one later gazes at the rocks. The adaptation is not confined to just one population of cells, but probably occurs at several cortical sites, reflecting the multiple levels of processing involved in visual motion analysis. The effect is unlikely to be caused by neural fatigue; more likely, the MAE and similar adaptation effects provide a form of error-correction or coding optimization, or both.

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.

Attentional modulation of adaptation to two-component transparent motion.

We have studied the effects of voluntary attention on the induction of motion aftereffects (MAEs). While adapting, observers paid attention to one of two transparently displayed random dot patterns, moving concurrently in opposite directions. Selective attention was found to modulate the susceptibility to motion adaptation very substantially. To measure the strength of the induced MAEs we modulated the signal-to-noise ratio of a real motion signal in a random dot pattern that was used to balance the aftereffect. Results obtained for adapting to single motion vectors show that the MAE can be represented as a shift of the psychometric function for motion direction discrimination. Selective attention to the different components of transparent motion altered the susceptibility to adaptation. Shifting attention from one component to the other caused a large shift of the psychometric curves, about 70-75% of the shift measured for the separate components of the transparent adapting stimulus. We conclude that attention can differentiate between spatially superimposed motion vectors and that attention modulates the activity of motion mechanisms before or at the level where adaptation gives rise to MAEs. The results are discussed in light of the role of attention in visual perception and the physiological site for attentional modulation of MAEs.

Temporal and spatial frequency tuning of the flicker motion aftereffect.

The motion aftereffect (MAE) was used to study the temporal and spatial frequency selectivity of the visual system at supra-threshold contrasts. Observers adapted to drifting sine-wave gratings of a range of spatial and temporal frequencies. The magnitude of the MAE induced by the adaptation was measured with counterphasing test gratings of a variety of spatial and temporal frequencies. Independently of the spatial or temporal frequency of the adapting grating, the largest MAE was found with slowly counterphasing test gratings (at approximately 0.125-0.25 Hz). The largest MAEs were also found when the test grating was of similar spatial frequency to that of the adapting grating, even at very low spatial frequencies (0.125 c/deg). These data suggest that MAEs are dominated by a single, low-pass temporal frequency mechanism and by a series of band-pass spatial frequency mechanisms. The band-pass spatial frequency tuning even at low spatial frequencies suggests that the "lowest adaptable channel" concept [Cameron et al. (1992). Vision Research, 32, 561-568] may be an artifact of disadvantaged low spatial frequencies using static test patterns.

Limits of attentive tracking reveal temporal properties of attention.

The maximum speed for attentive tracking of targets was measured in three types of (radial) motion displays: ambiguous motion where only attentive tracking produced an impression of direction, apparent motion, and continuous motion. The upper limit for tracking (about 50 deg s-1) was an order of magnitude lower than the maximum speed at which motion can be perceived for some of these stimuli. In all cases but one, the ultimate limit appeared to be one of temporal frequency, 4-8 Hz, not retinal speed or rotation rate. It was argued that this rate reflects the temporal resolution of attention, the maximum rate at which events can be individuated from those that precede or follow them. In one condition, evidence was also found for a speed limit to attentive tracking, a maximum rate at which attention could follow a path around the display.

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.

Temporal integration of random dot apparent motion information in human central vision.

Human motion perception is assumed to be functionally described by an array of bi-local detectors feeding later, higher order computational stages. Using this model as a guide, improvement of spatio-temporal displacement sensitivity by temporal integration (summation) was measured in human central vision using random dot pattern apparent-motion stimuli. Our results agree with previous experiments with regard to improvement of maximum perceivable spatial displacement but show that contrary to previous results the minimum perceivable spatial displacement can be improved in a similar manner. Furthermore, stimulus duration is a more accurate predictor of sensitivity than the number of frames in the stimulus over a wide range of stimulus parameter values. Finally, our results indicate that temporal tuning of motion detectors is inversely related to the size of the spatial pattern displacement.

An analysis of the temporal integration mechanism in human motion perception.

We present a model for the temporal integration of apparent motion information. The model is constructed by considering psychophysical and neurophysiological data, and consists of the leaky integration of pulsatile motion detector responses to apparent motion stimuli. Each pulse represents a motion detector populational response to a discrete spatial displacement of the spatial pattern. Temporal contrast sensitivity determines the shape of constant-stimulus-duration threshold curves for image frame exposure durations less than about 133 msec. The shape of the threshold curve for image frame exposure durations greater than about 133 msec is determined by the leaky integrator time constant and the shape of the pulses emitted by the motion detectors. The leaky integrator model exhibits threshold saturation behaviour (the reaching of a maximum sensitivity or minimum threshold) seen in psychophysical data as well as dependence of saturation time on the frame rate of the apparent motion stimulus. A low frame rate results in a longer time-to-saturation because the leaky integrator discharges more between detector output pulses. When the motion detector output pulses are far enough apart there is effectively no temporal integration and therefore no threshold improvement over time. Finally, the behaviour of the psychophysical threshold curves across spatial displacement sizes is consistent with a populational-response threshold mechanism combined with spatial summation over a non-uniform distribution of detector types across the visual field.

Spatial summation and its interaction with the temporal integration mechanism in human motion perception.

The combination of visual motion information over visual space (spatial summation) and stimulus duration (temporal integration) was investigated using a random-pixel array (spatiotemporally broad-band) apparent motion stimulus designed to isolate specific populations of visual motion detectors. The results indicate that, in agreement with results from spatiotemporally narrow-band stimuli, spatial summation follows the form of linear probabilistic summation rather than non-linear probabilistic summation. Linear probabilistic summation holds for a wide range of stimulus parameters and when changing either motion stimulus height or width. Linear probabilistic summation breaks down when the motion display region approaches a height and/or width that is related to the spatial displacement size, not the speed, of the random-pixel array. This height and width (termed the critical height and width, or critical dimension), increases with spatial displacement size and can be interpreted as a measure of the basic dimensions of the selected motion detector population's receptive field. The critical height is smaller than the critical width, a result that is consistent with a motion detector receptive field that is elongated in the direction of motion. Perhaps most importantly, the mechanisms of temporal integration and spatial summation can work independently under a wide range of conditions. Finally, the results provide evidence for a short-term inhibitory phenomenon from the edges of the useful display area that affects the visibility of the motion.

Movement aftereffect of bi-vectorial transparent motion.

Two moving random-pixel arrays (RPAs) were presented simultaneously in the same target field. These RPAs are perceived as two superimposed transparent moving sheets. Although two directions are perceived simultaneously during stimulus presentation, the movement aftereffect (MAE) is unidirectional. The visual system averages both motion signals in the MAE. For motion vectors of equal magnitude and perpendicular direction the MAE direction is the inverse of the sum of both vectors. In the first experiment we measured perceived direction of the MAE of transparent motion for a range of speed combinations. Results indicate that vector summation only predicts the correct MAE direction for combinations of equal speeds. It is suggested that the direction of the MAE of transparent motion is a resultant of the weighted summation of the component inducing vectors. The question then arises what determines the weighting factors. Directional sensitivity and MAE duration of the individual vectors under transparent conditions were measured and used to weigh the vectors and predict the MAE direction of transparent motion. Statistical analyses showed that MAE duration is a better basis to determine the weighting factors predicting the direction of the MAE of transparent motion than component sensitivity. The direction of the MAE of transparent motion thus seems to be determined by the amount of adaptation to the component vectors as reflected by MAE duration. The results suggest that this gain control cannot be located in the individual motion detectors and must be situated at or after some subsequent cooperation stage of the human motion analysis system.

Spatio-temporal characteristics of human motion perception.

A bi-local detector array model was assumed to describe the functional performance of monocular motion perception. Distributions of model parameters were measured in human vision at several positions in the visual field. The stimulus paradigm was designed to measure directional motion perception thresholds for individual combinations of spatial displacement and temporal delay in random dot apparent motion stimuli. The resulting data support previous results on perceivable spatial displacement limits in human vision but also indicate that both minimum and maximum perceivable spatial displacement thresholds in human observers have a similar dependence on temporal delay. This dependence changes with eccentricity in the visual field in a qualitatively similar manner but by quantitatively different factors. A description of possible biological properties of the bi-local detector population is presented that may explain how detection of spatio-temporal pattern displacements can be performed by a single system. Such a system also predicts that minimum and maximum perceivable spatial displacement thresholds should scale with visual field eccentricity in a manner consistent with our results.

Linking lower and higher stages of motion processing?

The spatial frequency selectivity of motion detection mechanisms can be measured by comparing the magnitude of motion aftereffects (MAEs) as a function of the spatial frequency of the adapting and test gratings. For static test gratings, narrow spatial frequency tuning has been reported in a number of studies. However, for dynamic test patterns, reports have been conflicting. Ashida & Osaka [(1994). Perception, 23, 1313-1320] found no tuning whereas Bex et al. [(1996) Vision Research, 36, 2721-2727] reported a narrow tuning. The main difference between the two studies was the temporal frequency of the test pattern. In this study we measured the spatial frequency tuning of the MAE using test patterns for a range of temporal frequencies. The results confirmed that there was narrow spatial frequency tuning when the test pattern was counterphasing at a low temporal frequency. However, the spatial frequency selectivity broadened as the temporal frequency of the test pattern was increased.

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.

Recovery from motion adaptation is delayed by successively presented orthogonal motion.

Following a period of adaptation to a pattern moving in a particular direction, a subsequently viewed stationary pattern appears to move in the opposite direction for some time: the movement after effect (MAE). The MAE lasts longer when the test pattern is not immediately or not continuously presented after adaptation. This phenomenon is called storage. So far research indicates that storage only occurs when textured visual stimulation is absent during part of the test phase or if the processing of a stationary test stimulus is prevented (e.g. by binocular rivalry). We present evidence that storage-like phenomena can occur even while a textured and moving visual stimulus is phenomenally present. We adapted binocularly to uni-directional motion of a random-pixel array M1 for 60 sec. This stimulus was immediately followed by another moving pattern M2. Its motion direction was orthogonal to that of M1. The presentation time of M2 was the independent variable. A stationary pattern was presented immediately after presentation of M2. The direction of the resulting integrated uni-directional MAE was measured. For short presentation times of M2 there is an integrated uni-directional MAE, which shows an interaction of the output of units stimulated by both moving patterns. However, it appeared that the effect of M1 on the direction of this combined uni-directional MAE is much longer present than would be expected from the MAE duration of M1, when tested in isolation.

Systematic eye movements do not account for the perception of motion during attentive tracking.

It has been suggested that attention can disambiguate stimuli that have equal motion energy in opposite directions (e.g. a counterphasing grating), such that a clear motion direction is perceived. The direction of this movement is determined by the observer and can be changed at will. Assuming that the responses of front-end motion detectors are equal for the two opponent directions, it has been proposed that the unambiguous motion perceived with attentive tracking arises from an independent mechanism that monitors the shifts of attention directed to the moving feature of interest. However, while perceiving motion under attentive tracking conditions, observers often report a strong impression that they are making eye movements. In this study, we investigated whether systematic eye movements are present during attentive tracking and, as a result, could be responsible for the subjective experience of movement. We had observers track an object in smooth motion, apparent motion and ambiguous motion, either with eye movements or with attention. The results show that there are negligible eye movements during attentive tracking, which are neither systematic nor correlated with the stimulus. Given that neither eye movements nor retinal image motion can account for subjectively perceived motion, as well as the absence of any other plausible explanation, we find it tempting evidence for an earlier suggestion that the percept of movement must arise from a specialized mechanism.

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.

Responses of complex cells in area 17 of the cat to bi-vectorial transparent motion.

We examined the responses to transparent motion of complex cells in cat area 17 which show directional selectivity to moving random pixel arrays (RPAs). The response to an RPA moving in the cell's preferred direction is inhibited when a second RPA is transparently moving in another direction. The inhibition by the second pattern is quantified as a function of its direction. The response to a pattern moving in the preferred direction is never completely suppressed, not even when a second pattern is moving transparently in the opposite direction. To the extent that supra-spontaneous firing rates signal the presence of the optimal velocity vector, these cells therefore still signal the presence of this line-label stimulus despite additional opposing, or otherwise directed, motion components. The results confirm previous suggestions that, for the computation of motion energy in cat area 17 complex cells, a full opponent stage is not plausible. Furthermore, we show that the response to a combination of two motion vectors can be predicted by the average of the responses to the individual components.

Directional motion sensitivity under transparent motion conditions.

We measured directional sensitivity to a foreground pattern while an orthogonally directed background pattern was present under transparent motion conditions. For both foreground and background pattern, the speed was varied between 0.5 and 28 deg sec-1. A multi-step paradigm was employed which results in a better estimation of the suppressive or facilitatory effects than previously applied single-step methods (e.g. measuring Dmax or Dmin). Moreover, our method gives insight into the interactions for a wide range of speed and not just the extreme motion thresholds (the D-values). We found that high background speeds have an inhibitory effect on the detection of a range of high foreground speeds and low background speeds have an inhibitory effect on a range of low foreground speeds. Intermediate background pattern speeds inhibit the detection of both low and high foreground pattern speeds and do so in a systemic manner.

Recovery from adaptation for dynamic and static motion aftereffects: evidence for two mechanisms.

The motion aftereffect (MAE) is an illusory drift of a physically stationary pattern induced by prolonged viewing of a moving pattern. Depending on the nature of the test pattern the MAE can be phenomenally different. This difference in appearance has led to the suggestion that different underlying mechanisms may be responsible and several reports show that this might be the case. Here, we tested whether differences in MAE duration obtained with stationary test patterns and dynamic test patterns can be explained by a single underlying mechanism. We find the results support the existence of (at least) two mechanisms. The two mechanisms show different characteristics: the static MAE (i.e. the MAE tested with a static test pattern) is almost completely stored when the static test is preceded by a dynamic test; in contradistinction, the dynamic MAE is not stored when dynamic testing is preceded by a static test pattern.