Additivity of retinal and pursuit velocity in the perceptions of depth and rigidity from object-produced motion parallax

Two psychophysical experiments were conducted to investigate the mechanism that generates stable depth structure from retinal motion combined with extraretinal signals from pursuit eye movements. Stimuli consisted of random dots that moved horizontally in one direction (ie stimuli had common motion on the retina), but at different speeds between adjacent rows. The stimuli were presented with different speeds of pursuit eye movements whose direction was opposite to that of the common retinal motion. Experiment 1 showed that the rows moving faster on the retina appeared closer when viewed without eye movements; however, they appeared farther when pursuit speed exceeded the speed of common retinal motion. The 'transition' speed of the pursuit eye movement was slightly, but consistently, larger than the speed of common retinal motion. Experiment 2 showed that parallax thresholds for perceiving relative motion between adjacent rows were minimum at the transition speed found in experiment 1. These results suggest that the visual system calculates head-centric velocity, by adding retinal velocity and pursuit velocity, to obtain a stable depth structure.     

The interaction between duration, velocity and repetitive auditory stimulation

Repetitive auditory stimulation (with click trains) and visual velocity signals both have intriguing effects on the subjective passage of time. Previous studies have established that prior presentation of auditory clicks increases the subjective duration of subsequent sensory input, and that faster moving stimuli are also judged to have been presented for longer (the time dilation effect). However, the effect of clicks on velocity estimation is unknown, and the nature of the time dilation effect remains ambiguous. Here were present a series of five experiments to explore these phenomena in more detail. Participants viewed a rightward moving grating which traveled at velocities ranging from 5 to 15°/s and which lasted for durations of 500 to 1500 ms. Gratings were preceded by clicks, silence or white noise. It was found that both clicks and higher velocities increased subjective duration. It was also found that the time dilation effect was a constant proportion of stimulus duration. This implies that faster velocity increases the rate of the pacemaker component of the internal clock. Conversely, clicks increased subjective velocity, but the magnitude of this effect was not proportional to actual velocity. Through considerations of these results, we conclude that clicks independently affect velocity and duration representations.     

Visual localization and estimation of extent of target motion during ocular pursuit: a common mechanism?

The ability to localize a visual target and to estimate the distance through which it moves was studied during ocular pursuit. In the first experiment observers had to localize the position of a visually tracked moving target when they heard an acoustic signal. The signal was sounded near the beginning or near the end of the motion. The distance between the perceived positions was shorter than the distance between the corresponding physical positions of the target. The 'shortening' became more pronounced with higher tracking velocity. In another condition the observers estimated the length of the motion path between two successive sound signals, one presented near the beginning and one near the end of the motion. The length of path travelled was underestimated, the effect being stronger with higher tracking velocity. In the second experiment this effect of velocity on the underestimation of distance was shown to exist only during ocular pursuit and not during steady fixation. The hypothesis that localization and estimation of distance during ocular pursuit share a common mechanism is discussed.     

Retinal and extraretinal information in movement perception: how to invert the Filehne illusion

During a pursuit eye movement made in darkness across a small stationary stimulus, the stimulus is perceived as moving in the opposite direction to the eyes. This so-called Filehne illusion is usually explained by assuming that during pursuit eye movements the extraretinal signal (which informs the visual system about eye velocity so that retinal image motion can be interpreted) falls short. A study is reported in which the concept of an extraretinal signal is replaced by the concept of a reference signal, which serves to inform the visual system about the velocity of the retinae in space. Reference signals are evoked in response to eye movements, but also in response to any stimulation that may yield a sensation of self-motion, because during self-motion the retinae also move in space. Optokinetic stimulation should therefore affect reference signal size. To test this prediction the Filehne illusion was investigated with stimuli of different optokinetic potentials. As predicted, with briefly presented stimuli (no optokinetic potential) the usual illusion always occurred. With longer stimulus presentation times the magnitude of the illusion was reduced when the spatial frequency of the stimulus was reduced (increased optokinetic potential). At very low spatial frequencies (strongest optokinetic potential) the illusion was inverted. The significance of the conclusion, that reference signal size increases with increasing optokinetic stimulus potential, is discussed. It appears to explain many visual illusions, such as the movement aftereffect and center-surround induced motion, and it may bridge the gap between direct Gibsonian and indirect inferential theories of motion perception.     

Judgments of synchrony between auditory and moving or still visual stimuli.

The flash-lag effect is a visual illusion wherein intermittently flashed, stationary stimuli seem to trail after a moving visual stimulus despite being flashed synchronously. We tested hypotheses that the flash-lag effect is due to spatial extrapolation, shortened perceptual lags, or accelerated acquisition of moving stimuli, all of which call for an earlier awareness of moving visual stimuli over stationary ones. Participants judged synchrony of a click either to a stationary flash of light or to a series of adjacent flashes that seemingly bounced off or bumped into the edge of the visual display. To be judged synchronous with a stationary flash, audio clicks had to be presented earlier--not later--than clicks that went with events, like a simulated bounce (Experiment 1) or crash (Experiments 2-4), of a moving visual target. Click synchrony to the initial appearance of a moving stimulus was no different than to a flash, but clicks had to be delayed by 30-40 ms to seem synchronous with the final (crash) positions (Experiment 2). The temporal difference was constant over a wide range of motion velocity (Experiment 3). Interrupting the apparent motion by omitting two illumination positions before the last one did not alter subjective synchrony, nor did their occlusion, so the shift in subjective synchrony seems not to be due to brightness contrast (Experiment 4). Click synchrony to the offset of a long duration stationary illumination was also delayed relative to its onset (Experiment 5). Visual stimuli in motion enter awareness no sooner than do stationary flashes, so motion extrapolation, latency difference, and motion acceleration cannot explain the flash-lag effect.     

Keeping balance during head-free smooth pursuit: The role of aging

Standing balance is often more unstable when visually pursuing a moving target than when fixating on a stationary one. These effects are common in both young and older adults when the head is restrained during visual task performance. The present study focused on the role of head motion on standing balance during smooth pursuit as a function of age. Three predictions were tested: a) standing balance is compromised to a greater extent in older than young adults by gaze target pursuit compared to fixation, b) older adults pursue a moving target with greater and more variable head rotation than young adults, and c) greater and more variable head rotation during the smooth pursuit task is associated with greater Center of Pressure (CoP) sway. Twenty-two (22) older (age: 71.7 ± 8.1, 12 M / 10 F) and twenty-three (23) young adults (age: 23.6 ± 2.5, 12 M / 11 F) stood on a force plate while either fixating a stationary or smoothly pursuing a horizontally moving target (31.9° peak-to-peak visual angle). CoP (Bertec Balance Plate), head kinematics (Vicon Motion Analysis) and head-unconstrained gaze (Pupil Labs Invisible) were synchronously recorded. The root means square (RMS) of CoP velocity increased during smooth pursuit compared to fixation regardless of age (p < .05), while the interquartile CoP range increased only in older and not in young participants (p < .05). We also calculated the head rotation range (peak to peak cycle amplitude) of motion and variability (SD of range of motion) across the cycles of the smooth pursuit task. Older adults pursued the moving target employing more variable (p = .022) head yaw rotation than young participants although the mean range of head rotation was similar between groups (p =. 077). The amplitude and variability of head yaw rotation did not correlate with CoP sway measures. Results suggest that head-free pursuing of a moving target decreased balance to a greater extent in old than young individuals when compared to fixation. Nevertheless, postural sway during head-free smooth pursuit was not associated with the extent or variability of head rotation.     

Extraretinal and retinal amplitude and phase errors during Filehne illusion and path perception

Pursuit eye movements give rise to retinal motion. To judge stimulus motion relative to the head, the visual system must correct for the eye movement by using an extraretinal, eye-velocity signal. Such correction is important in a variety of motion estimation tasks including judgments of object motion relative to the head and judgments of self-motion direction from optic flow. The Filehne illusion (where a stationary object appears to move opposite to the pursuit) results from a mismatch between retinal and extraretinal speed estimates. A mismatch in timing could also exist. Speed and timing errors were investigated using sinusoidal pursuit eye movements. We describe a new illusion--the slalom illusion--in which the perceived direction of self-motion oscillates left and right when the eyes move sinusoidally. A linear model is presented that determines the gain ratio and phase difference of extraretinal and retinal signals accompanying the Filehne and slalom illusions. The speed mismatch and timing differences were measured in the Filehne and self-motion situations using a motion-nulling procedure. Timing errors were very small for the Filehne and slalom illusions. However, the ratios of extraretinal to retinal gain were consistently less than 1, so both illusions are the consequence of a mismatch between estimates of retinal and extraretinal speed. The relevance of the results for recovering the direction of self-motion during pursuit eye movements is discussed.     

Adaptation to auditory motion in the horizontal plane: effect of prior exposure to motion on motion detectability.

Thresholds for auditory motion detectability were measured in a darkened anechoic chamber while subjects were adapted to horizontally moving sound sources of various velocities. All stimuli were 500-Hz lowpass noises presented at a level of 55 dBA. The threshold measure employed was the minimum audible movement angle (MAMA)--that is, the minimum angle a horizontally moving sound must traverse to be just discriminable from a stationary sound. In an adaptive, two-interval forced-choice procedure, trials occurred every 2-5 sec (Experiment 1) or every 10-12 sec (Experiment 2). Intertrial time was "filled" with exposure to the adaptor--a stimulus that repeatedly traversed the subject's front hemifield at ear level (distance: 1.7 m) at a constant velocity (-150 degrees/sec to +150 degrees/sec) during a run. Average MAMAs in the control condition, in which the adaptor was stationary (0 degrees/sec,) were 2.4 degrees (Experiment 1) and 3.0 degrees (Experiment 2). Three out of 4 subjects in each experiment showed significantly elevated MAMAs (by up to 60%), with some adaptors relative to the control condition. However, there were large intersubject differences in the shape of the MAMA versus adaptor velocity functions. This loss of sensitivity to motion that most subjects show after exposure to moving signals is probably one component underlying the auditory motion aftereffect (Grantham, 1989), in which judgments of the direction of moving sounds are biased in the direction opposite to that of a previously presented adaptor.     

Distractor interference during smooth pursuit eye movements

When 2 targets for pursuit eye movements move in different directions, the eye velocity follows the vector average (S. G. Lisberger & V. P. Ferrera, 1997). The present study investigates the mechanisms of target selection when observers are instructed to follow a predefined horizontal target and to ignore a moving distractor stimulus. Results show that at 140 ms after distractor onset, horizontal eye velocity is decreased by about 25%. Vertical eye velocity increases or decreases by 1 degrees /s in the direction opposite from the distractor. This deviation varies in size with distractor direction, velocity, and contrast. The effect was present during the initiation and steady-state tracking phase of pursuit but only when the observer had prior information about target motion. Neither vector averaging nor winner-take-all models could predict the response to a moving to-be-ignored distractor during steady-state tracking of a predefined target. The contributions of perceptual mislocalization and spatial attention to the vertical deviation in pursuit are discussed.     

Evaluation of an imputed pitch velocity model of the auditory tau effect

This article extends an imputed pitch velocity model of the auditory kappa effect proposed by Henry and McAuley (2009a) to the auditory tau effect. Two experiments were conducted using an AXB design in which listeners judged the relative pitch of a middle target tone (X) in ascending and descending three-tone sequences. In Experiment 1, sequences were isochronous, establishing constant fast, medium, and slow velocity conditions. No systematic distortions in perceived target pitch were observed, and thresholds were similar across velocity conditions. Experiment 2 introduced to-be-ignored variations in target timing. Variations in target timing that deviated from constant velocity conditions introduced systematic distortions in perceived target pitch, indicative of a robust auditory tau effect. Consistent with an auditory motion hypothesis, the magnitude of the tau effect was larger at faster velocities. In addition, the tau effect was generally stronger for descending sequences than for ascending sequences. Combined with previous work on the auditory kappa effect, the imputed velocity model and associated auditory motion hypothesis provide a unified quantitative account of both auditory tau and kappa effects. In broader terms, these findings add support to the view that pitch and time relations in auditory patterns are fundamentally interdependent.     

Visual short-term memory during smooth pursuit eye movements

Visual short-term memory (VSTM) was probed while observers performed smooth pursuit eye movements. Smooth pursuit keeps a moving object stabilized in the fovea. VSTM capacity for position was reduced during smooth pursuit compared with a condition with eye fixation. There was no difference between a condition in which the items were approximately stabilized on the retina because they moved with the pursuit target and a condition in which the items moved across the retina because they were stationary in space. The reduction of capacity for position was eliminated when miniature items were presented on the pursuit target. Similarly, VSTM capacity for color did not differ between smooth pursuit and fixation. The results suggest that visuospatial attention is tied to the target during smooth pursuit, which impairs VSTM for the position of peripheral objects. Sensory memory during smooth pursuit was only slightly impaired.     

Adaptation to auditory motion in the horizontal plane: Effect of prior exposure to motion on motion detectability

Thresholds for auditory motion detectability were measured in a darkened anechoic chamber while subjects were adapted to horizontally moving sound saurces of various-velocities. All stimuli were 500-Hz lowpass noises presented at a level of 55 dBA. The threshold measure employed was the minimum audible movement angle(MAMA)—that is, the minimum angle a horizontally moving sound must traverse to be just discriminable from a stationary sound. In an adaptive, two-interval forced-choice procedure, trials occurred every 2-5 sec (Experiment 1) or every 10–12 sec (Experiment 2). Intertrial time was “filled” with exposure to the adaptor—a stimulus that repeatedly traversed the subject’s front hemifield at ear level (distance: 1.7 m) at a constant velocity (?150°/secto + 150°/sec)during a run. Average MAMAs in the control condition, in which the adaptor was stationary (0°/sec), were 2.4° (Experiment 1) and 3.0° (Experiment 2). Three out of 4 subjects in each experiment showed significantly elevated MAMAs (by up to 60%), with some adaptors relative to the control condition. However, there were large intersubject differences in the shape of the MAMA versus adaptor velocity functions. This loss of sensitivity to motion that most subjects show after exposure to moving signals is probably one component underlying the auditory motion aftereffect (Grantham, 1989), in which judgmentsof the direction-afmoving sounds are biased in the direction opposite to that of a previously presented adaptor.     

Neurophysiology and neuroanatomy of smooth pursuit in humans

Smooth pursuit eye movements enable us to focus our eyes on moving objects by utilizing well-established mechanisms of visual motion processing, sensorimotor transformation and cognition. Novel smooth pursuit tasks and quantitative measurement techniques can help unravel the different smooth pursuit components and complex neural systems involved in its control. The maintenance of smooth pursuit is driven by a combination of the prediction of target velocity and visual feedback about performance quality, thus a combination of retinal and extraretinal information that has to be integrated in various networks. Different models of smooth pursuit with specific in- and output parameters have been developed for a better understanding of the underlying neurophysiological mechanisms and to make quantitative predictions that can be tested in experiments. Functional brain imaging and neurophysiological studies have defined motion sensitive visual area V5, frontal (FEF) and supplementary (SEF) eye fields as core cortical smooth pursuit regions. In addition, a dense neural network is involved in the adjustment of an optimal smooth pursuit response by integrating also extraretinal information. These networks facilitate interaction of the smooth pursuit system with multiple other visual and non-visual sensorimotor systems on the cortical and subcortical level. Future studies with fMRI advanced techniques (e.g., event-related fMRI) promise to provide an insight into how smooth pursuit eye movements are linked to specific brain activation. Applying this approach to neurological and also mental illness can reveal distinct disturbances within neural networks being present in these disorders and also the impact of medication on this circuitry.     

Neurophysiology and neuroanatomy of smooth pursuit in humans

Smooth pursuit eye movements enable us to focus our eyes on moving objects by utilizing well-established mechanisms of visual motion processing, sensorimotor transformation and cognition. Novel smooth pursuit tasks and quantitative measurement techniques can help unravel the different smooth pursuit components and complex neural systems involved in its control. The maintenance of smooth pursuit is driven by a combination of the prediction of target velocity and visual feedback about performance quality, thus a combination of retinal and extraretinal information that has to be integrated in various networks. Different models of smooth pursuit with specific in- and output parameters have been developed for a better understanding of the underlying neurophysiological mechanisms and to make quantitative predictions that can be tested in experiments. Functional brain imaging and neurophysiological studies have defined motion sensitive visual area V5, frontal (FEF) and supplementary (SEF) eye fields as core cortical smooth pursuit regions. In addition, a dense neural network is involved in the adjustment of an optimal smooth pursuit response by integrating also extraretinal information. These networks facilitate interaction of the smooth pursuit system with multiple other visual and non-visual sensorimotor systems on the cortical and subcortical level. Future studies with fMRI advanced techniques (e.g., event-related fMRI) promise to provide an insight into how smooth pursuit eye movements are linked to specific brain activation. Applying this approach to neurological and also mental illness can reveal distinct disturbances within neural networks being present in these disorders and also the impact of medication on this circuitry.     

Target velocity effects on manual interception kinematics

Participants generated manual interception movements toward a target cursor that moved across a computer screen. The target reached its peak velocity either during the first third, at the midpoint, or during the last third of the movement. In Experiment 1 the view of the target was available for either the first 316, 633, 950, or 1267 ms, after which it disappeared. Results showed that for all viewing conditions, the timing of the interception velocity was related to the temporal properties of the target's trajectory. In Experiment 2, when the portion of the target trajectory that was viewed was reversed (such that participants did not see the first 316, 633, 950, or 1267 ms of the trajectory, but instead saw only the later portions of the trajectory), there was no clear relationship between the target trajectory and the timing of the aiming trajectory. These results suggest that participants use visual information early in the target's trajectory to form a representation of the target motion that is used to facilitate manual interception.     

The spoke brightness illusion originates at an early motion processing stage

When a bright white disk revolves around a fixation point on a gray background, observers perceive a "spoke": a dark gray region that connects the disk with the fixation point. Our first experiment suggests that motion across the retina is both necessary and sufficient for spokes: The illusion occurs when a disk moves across the retina even though it is perceived to be stationary, but the illusion does not occur when the disk appears to move while remaining stationary on the retina. A second experiment shows that the strength of the illusion decreases with decreasing luminance contrast until subjective equiluminance, where little or no spoke is perceived. These results suggest that spokes originate at an early, predominantly luminance-based stage of motion processing, before the visual system discounts retinal motion caused by smooth pursuit.     

Congruency effects between auditory and tactile motion: Extending the phenomenon of cross-modal dynamic capture

Behavioral studies of multisensory integration in motion perception have focused on the particular case of visual and auditory signals. Here, we addressed a new case: audition and touch. In Experiment 1, we tested the effects of an apparent motion stream presented in an irrelevant modality (audition or touch) on the perception of apparent motion streams in the other modality (touch or audition, respectively). We found significant congruency effects (lower performance when the direction of motion in the irrelevant modality was incongruent with the direction of the target) for the two possible modality combinations. This congruency effect was asymmetrical, with tactile motion distractors having a stronger influence on auditory motion perception than vice versa. In Experiment 2, we used auditory motion targets and tactile motion distractors while participants adopted one of two possible postures: arms uncrossed or arms crossed. The effects of tactile motion on auditory motion judgments were replicated in the arms-uncrossed posture, but they dissipated in the arms-crossed posture. The implications of these results are discussed in light of current findings regarding the representation of tactile and auditory space.     

A comparison of visual and auditory representational momentum in spatial tasks

Similarities have been observed in the localization of the final position of moving visual and moving auditory stimuli: Perceived endpoints that are judged to be farther in the direction of motion in both modalities likely reflect extrapolation of the trajectory, mediated by predictive mechanisms at higher cognitive levels. However, actual comparisons of the magnitudes of displacement between visual tasks and auditory tasks using the same experimental setup are rare. As such, the purpose of the present free-field study was to investigate the influences of the spatial location of motion offset, stimulus velocity, and motion direction on the localization of the final positions of moving auditory stimuli (Experiment 1 and 2) and moving visual stimuli (Experiment 3). To assess whether auditory performance is affected by dynamically changing binaural cues that are used for the localization of moving auditory stimuli (interaural time differences for low-frequency sounds and interaural intensity differences for high-frequency sounds), two distinct noise bands were employed in Experiments 1 and 2. In all three experiments, less precise encoding of spatial coordinates in paralateral space resulted in larger forward displacements, but this effect was drowned out by the underestimation of target eccentricity in the extreme periphery. Furthermore, our results revealed clear differences between visual and auditory tasks. Displacements in the visual task were dependent on velocity and the spatial location of the final position, but an additional influence of motion direction was observed in the auditory tasks. Together, these findings indicate that the modality-specific processing of motion parameters affects the extrapolation of the trajectory.     

Peak velocity as a cue in audiovisual synchrony perception of rhythmic stimuli

This study investigated audiovisual synchrony perception in a rhythmic context, where the sound was not consequent upon the observed movement. Participants judged synchrony between a bouncing point-light figure and an auditory rhythm in two experiments. Two questions were of interest: (1) whether the reference in the visual movement, with which the auditory beat should coincide, relies on a position or a velocity cue; (2) whether the figure form and motion profile affect synchrony perception. Experiment 1 required synchrony judgment with regard to the same (lowest) position of the movement in four visual conditions: two figure forms (human or non-human) combined with two motion profiles (human or ball trajectory). Whereas figure form did not affect synchrony perception, the point of subjective simultaneity differed between the two motions, suggesting that participants adopted the peak velocity in each downward trajectory as their visual reference. Experiment 2 further demonstrated that, when judgment was required with regard to the highest position, the maximal synchrony response was considerably low for ball motion, which lacked a peak velocity in the upward trajectory. The finding of peak velocity as a cue parallels results of visuomotor synchronization tasks employing biological stimuli, suggesting that synchrony judgment with rhythmic motions relies on the perceived visual beat.     

An acceleration illusion caused by underestimation of stimulus velocity during pursuit eye movements: Aubert-Fleischl revisited

When the eyes pursue a fixation point that sweeps across a moving background pattern, and the fixation point is suddenly made to stop, the ongoing motion of the background pattern seems to accelerate to a higher velocity. Experiment I showed that this acceleration illusion is not caused by the sudden change in (i) the relative velocity between background and fixation point, (ii) the velocity of the retinal image of the background pattern, or (iii) the motion of the retinal image of the rims of the CRT screen on which the experiment was carried out. In experiment II the magnitude of the illusion was quantified. It is strongest when background and eyes move in the same direction. When they move in opposite directions it becomes less pronounced (and may disappear) with higher background velocities. The findings are explained in terms of a model proposed by the first author, in which the perception of object motion and velocity derives from the interaction between retinal slip velocity information and the brain's 'estimate' of eye velocity in space. They illustrate that the classic Aubert-Fleischl phenomenon (a stimulus seems to be moving slower when pursued with the eyes than when moving in front of stationary eyes) is a special case of a more general phenomenon: whenever we make a pursuit eye movement we underestimate the velocity of all stimuli in our visual field which happen to move in the same direction as our eyes, or which move slowly in the direction opposite to our eyes.