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Drift-balanced random stimuli: a general basis for studying non-Fourier motion perception.

To some degree, all current models of visual motion-perception mechanisms depend on the power of the visual signal in various spatiotemporal-frequency bands. Here we show how to construct counterexamples: visual stimuli that are consistently perceived as obviously moving in a fixed direction yet for which Fourier-domain power analysis yields no systematic motion components in any given direction. We provide a general theoretical framework for investigating non-Fourier motion-perception mechanisms; central are the concepts of drift-balanced and microbalanced random stimuli. A random stimulus S is drift balanced if its expected power in the frequency domain is symmetric with respect to temporal frequency, that is, if the expected power in S of every drifting sinusoidal component is equal to the expected power of the sinusoid of the same spatial frequency, drifting at the same rate in the opposite direction. Additionally, S is microbalanced if the result WS of windowing S by any space-time-separable function W is drift balanced. We prove that (i) any space-time-separable random (or nonrandom) stimulus is microbalanced; (ii) any linear combination of pairwise independent microbalanced (respectively, drift-balanced) random stimuli is microbalanced and drift balanced if the expectation of each component is uniformly zero; (iii) the convolution of independent microbalanced and drift-balanced random stimuli is microbalanced and drift balanced; (iv) the product of independent microbalanced random stimuli is microbalanced; and (v) the expected response of any Reichardt detector to any microbalanced random stimulus is zero at every instant in time. Examples are provided of classes of microbalanced random stimuli that display consistent and compelling motion in one direction. All the results and examples from the domain of motion perception are transposable to the space-domain problem of detecting orientation in a texture pattern.

Fourier Analysis

Reversed rotary motion perception.

A stroboscopically presented revolving annulus composed of dots is used to elicit rotary motion perception. Observers judge the direction of rotary motion. We find sharp and gradual transitions in the probability for reversed motion perception as a function of the angle of rotation between successive frames. These transitions reveal that matches between nonsuccessive frames can dominate motion perception. The transitions are scale invariant. The strength of a match is discussed in terms of a motion strength function, which is a separable function of the angle of rotation between successive frames and the frame repetition rate. The dependence of motion strength on the frame repetition rate (time function) is computed from the transitions. The similarity of this time function for rotary motion with the time function for linear motion [Psychol. Rev. 88, 171 (1981)] suggests that mechanisms for the discrimination of rotary motion address local detectors of linear motion.

Humans

On and off pathway contributions to apparent motion perception.

We studied the separability and/or interaction of the On and Off pathways in their role as inputs to visual motion perception. Using the long-range motion perception system, we asked if the motion system can use brightness polarity information, by testing whether correspondence is preferred between elements for which brightness polarity is preserved. We found such a preference, suggesting that brightness polarity information is indeed available to the motion system. However, under certain conditions motion is perceived even though the brightness polarity of apparent motion stimulus elements is reversed, indicating that the apparent motion system does integrate information from these two pathways. The source of the preference for maintaining polarity seems not to be the different brightnesses of the dark and bright stimulus elements, but the very fact that information must be integrated from different pathways. We relate the characteristics of the dependence of the motion perception on element contrast and contrast sign to those of previously reported visual evoked potential responses to brightness increments and decrements.

Contrast Sensitivity

Object-motion detection affected by concurrent self-motion perception: applied aspects for vehicle guidance.

Thresholds and response times for object-motion detection are significantly raised during concurrent real or visually induced self-motion perception. This was demonstrated by standardized laboratory experiments in which subjects had to react to a suprathreshold visual stimulus (1 degree-light spot moving with 5 degrees/s speed) under different stimulus conditions of simultaneously perceived self-motion. Elevated response times (mean elevation factor: 3.27) were also obtained for the detection of changes in inter-vehicle distance (headway) under real road conditions with the simultaneous involvement of self- and object-motion perception compared to a corresponding (object-motion perception) simulation in the laboratory without any self-motion. With regard to vehicle guidance, existing concepts of safe stopping distances, which depend upon adequate detection of a collision course and the corresponding reaction times, have to be recalculated.

Adult

The Aubert-Fleischl phenomenon: a temporal frequency effect on perceived velocity in afferent motion perception.

Apparent velocities of moving visual stimuli are known to be different depending on whether the subject pursues the stimulus (efferently controlled motion perception) or whether the eye is stationary and the image moves across the retina (afferent motion perception). Afferent motion perception of a periodic pattern or a moving single object causes overestimation of velocity (magnitude estimations) as compared to smooth pursuit. This socalled Aubert-Fleischl phenomenon is shown to depend on local temporal frequency stimulation on the retina caused by the repetitive passage of contrast borders of the moving periodic pattern. This is evidenced by the fact that for a given stimulus speed the amount of overestimation is a function of the spatial frequency of the pattern (or the angular subtend of a single moving object) and that the Aubert-Fleischl phenomenon is not observed if a single edge moves. Background characteristics seem not to influence the apparent velocity during smooth pursuit.

Adult

Motion perception during dichoptic viewing of moving random-dot stereograms.

The relation between binocular and monocular motion perception was investigated. A random-dot stereogram (30 X 30 deg arc), containing a central figure seen in front of the background in stereoscopic vision, was viewed dichoptically by human subjects without a fixed visual frame of reference. The images seen by the right and left eye were moved laterally according to a triangular wave form, in counterphase, but with variable amplitude ratios. Under this condition only purely lateral movement and no motion in depth of the stereogram as a whole was perceived, while stereoscopic vision of the figure-background relation was maintained. The magnitude of the binocularly perceived lateral motion, signalled by manual tracking of the perceived displacement, equalled the algebraic mean of the monocular motion percepts. As a special case, when the two images forming the stereogram were moved with equal velocities but in opposite directions they were perceived as a completely stationary, fused image in stereoscopic depth. Only the addition of a stationary reference (a bar or grating seen by both eyes) resulted in the perception of motion in depth. We conclude that a visual frame of reference is essential for perception of motion in depth but not for perception of lateral movements. Moreover, it seems likely that not absolute binocular disparity (retinal locus differences) but relative binocular disparity (differences in angular distance between two or more corresponding features in the two retinal images) is a cue for perception of depth.

Convergence, Ocular

Some additional predictions and further tests of the Marr-Ullman model of motion perception.

The Marr-Ullman model for motion detection in the human visual system functions by means of the dual input of polarity-specific edge detectors and luminance change detectors. Moulden and Begg (1986) found a polarity-specific motion aftereffect which they claimed provided support for this dual input model. The logic of their experiment is examined, and it is shown that several additional predictions arise from the Marr-Ullman model, which were not supported by Moulden and Begg's study. A more powerful experiment was carried out and these additional predictions were disconfirmed, although the polarity-specific effect did emerge. A consideration of alternative explanations of this effect led to a second experiment in which an attempt was made to discover the actual determinants of the effect. This revealed that polarity-specific units are unlikely to play any part in the phenomenon. It was concluded, in the light of this and other evidence, that one of a class of alternative models is more likely to be the actual mechanism for motion perception. However, careful consideration of the Marr-Ullman model indicated that it may be untestable in principle if various differentially weighted levels of neural integration are envisaged.

Attention

[An experiment on the theory of visual motion perception].

Two classes of theories of motion perception were studied: correlation and gradient models (in the sense of D. Marr). Random-dot kinematograms with a shifted square were presented to subjects, and the 80% threshold for detection of correct direction of movement was determined. Correlation models predict scale invariance, that is, a constant shift measured in units of texture elements of the translated pattern in spite of a geometric magnification. This was refuted for all subjects. The increase of the recognizable translation with the area of the translated form was verified except for the patterns with the largest texture elements (8 minutes of arc). This prediction, however, is not very specific for competing theories. Gradient models are not yet fully specified, and they contain some free parameters. They cannot be tested strictly, but there exist reasonable numerical parameter values by which our data can be satisfactorily explained.

Adult

Object-motion detection affected by concurrent self-motion perception: psychophysics of a new phenomenon.

Thresholds for object-motion detection are significantly raised when concurrent self-motion perception is induced by either vestibular, or visual, or cervico-somatosensory stimulation. Active sinusoidal horizontal head oscillations with compensatory vestibulo-ocular reflex (VOR) and foveal or eccentrical target presentation; 'passive' head movements with fixation suppression of the VOR; pure body oscillations with the head fixed in space (cervical stimulation); optokinetically induced apparent self-motion (circularvection). This new visual phenomenon of a physiological 'inhibitory interaction' between object- and self-motion perception seems to have a somatosensory motor analogue. It may reflect the disadventageous side effect due to unspecificness of an otherwise beneficial space constancy mechanism, which provides us with the image of a stable world during locomotion.

Adolescent

Monocular motion sensing, binocular motion perception.

The two-process account of motion perception and its binocular organization were addressed in experiments on apparent movement (AM) with three types of grating: sinusoidal; random bar width; and square-wave with missing fundamental (MF). Monocular MF gratings sampled four times per cycle of drift always appeared to move backwards. Here AM was unrelated to the spatial appearance of the pattern, and followed the motion of the dominant spatial frequency component (the third harmonic). We take this reversed AM to be characteristic of "short-range" motion sensors. It did not occur dichoptically, implying that the direction-selective mechanism of motion sensors is purely monocular. AM was seen with dichoptic presentation for all three types of grating. Performance improved with the length of the stimulus sequence, as predicted by probability summation. This result reconciles previous positive and negative findings on dichoptic AM. The perceived direction of dichoptic AM was consistent with polarity-selective matching of features over time (the "long-range process"). The most telling effect supporting feature-matching in dichoptic motion was that dichoptic MF motion reversed direction with a change in the visible features of the pattern (induced by changes in contrast and pulse duration); monocular apparent motion did not. Two routes from spatial frequency channels to the perception of object motion are discussed.

Contrast Sensitivity

Absence of smooth motion perception in color vision.

We have tested the behavioral evidence for a separation of the processing of color contrast from motion in the human visual system. Two different aspects of motion perception are examined; the identification of the direction of movement of a chromatic grating and the perception of smooth motion. The results show that color vision is at no great disadvantage in the identification of direction of movement, since this can be done at color contrasts quite close to detection threshold over a wide range of spatial and temporal frequencies. However, we find that subjects can identify direction without having the genuine perception of smooth motion. Smooth motion perception is revealed to be highly impaired since it is detected only at very high color contrasts and over a narrow range of spatial temporal conditions.

Color Perception

Motion perception and aging.

The authors used a correlated motion paradigm to investigate the effects of aging and gender on motion sensitivity. In 2 experiments with a total of 50 elderly and 50 young subjects, motion thresholds were significantly higher for elderly women. The correlated motion signal, which was embedded in random motion, may have been coherent to subjects in much the same way a form is in Witkin's Embedded Figures Test (EFT). In Experiment 2, EFT scores were obtained. A significant positive relationship between motion thresholds and EFT performance was found within each age group. Although gender-related perceptual style differences may contribute to motion perception effects, the authors argue that a common neural factor contributes to performance on both the EFT and the correlated motion task.

Adult

Intact "biological motion" and "structure from motion" perception in a patient with impaired motion mechanisms: a case study.

A series of psychophysical tests examining early and later aspects of image-motion processing were conducted in a patient with bilateral lesions involving the posterior visual pathways, affecting the lateral parietal-temporal-occipital cortex and the underlying white matter (as shown by magnetic resonance imaging studies and confirmed by neuro-ophthalmological and neuropsychological examinations). Visual acuity, form discrimination, color, and contrast-sensitivity discrimination were normal whereas spatial localization, line bisection, depth, and binocular stereopsis were severely impaired. Performance on early motion tasks was very poor. These include seeing coherent motion in random noise (Newsome & Paré, 1988), speed discrimination, and seeing two-dimensional form from relative speed of motion. However, on higher-order motion tasks the patient was able to identify actions from the evolving pattern of dots placed at the joints of a human actor (Johansson, 1973) as well as discriminating three-dimensional structure of a cylinder from motion in a dynamic random-dot field. The pattern of these results is at odds with the hypothesis that precise metrical comparison of early motion measurements is necessary for higher-order "structure from motion" tasks.

Cerebral Cortex

Does primate motion perception depend on the magnocellular pathway?

This study examined the importance of the primate magnocellular retinocortical pathway in the perception of moving stimuli. A portion of the magnocellular pathway was permanently and selectively interrupted by ibotenic acid injections in the LGN of macaque monkeys. We then tested contrast sensitivity for detecting moving stimuli, as well as two indices of motion perception, contrast sensitivity for opposite direction discrimination and speed difference thresholds, in the affected portion of the visual field. Magnocellular lesions greatly reduced detection contrast sensitivity at high temporal and low spatial frequencies and had a similar effect on contrast sensitivity for opposite direction discrimination under these same stimulus conditions. Consequently, opposite direction discriminations could be made at contrast threshold, suggesting that magnocellular lesions reduced the visibility of stimuli used to test direction perception, but did not act directly on direction perception. Magnocellular lesions also elevated speed difference thresholds under some stimulus conditions. However, this deficit was reduced or eliminated by raising the contrast of the test stimulus. Together, these findings suggest that magnocellular lesions reduce the visibility of stimuli used to test motion perception but that they do not appear to alter motion perception otherwise.

Animals

Assimilation and contrast in motion perception: explorations in cooperativity.

Motions within one region of the field influence motion seen elsewhere. To explore this phenomenon we used cinematograms comprised of alternating strips within which dots (i) tended to move in one direction, or (ii) moved in random directions (dynamic noise). When alternating strips were narrow, motion in one direction induced a similar direction of illusory motion in the adjoining dynamic noise (assimilation); when alternating strips were wide, motion tended to induce an illusory opposed motion in the dynamic noise (contrast). Since this illusory motion exhibits hysteresis, it probably results from spatially distributed, cooperative processes. The shift from assimilation to contrast, as the cinematogram's strips increase in size, suggests that facilitatory and inhibitory influences of the network extend over different distances.

Contrast Sensitivity

Influence of head or trunk oscillations on visually induced self-motion perception in humans.

The influence of concurrent head or trunk movements on optokinetically induced self-motion perception has been investigated by measuring circularvection (CV) latencies. Active head as well as passive trunk oscillation cause an increase in circularvection latency of about 50% as compared to latencies with the head and trunk stationary during full field optokinetic stimulation. The results point towards an intersensory inhibition of self-motion perception.

Adolescent

Consequences of spatial sampling for human motion perception.

This paper describes evidence for spatial aliasing in human motion perception. For a certain range of spatial frequencies, interference fringes drifting across the extrafoveal retina resemble two-dimensional spatial noise drifting in the opposite direction. For retinal locations within 10 deg of the fovea, the perceived direction of motion is veridical up to spatial frequencies near the cone Nyquist frequency, reverses between one and two times the cone Nyquist frequency, and sometimes reverses back to the correct direction at still higher frequencies. Thus two "motion nulls", or spatial frequencies at which the direction of motion is ambiguous, are typically observed at each retinal eccentricity. A computational model is described in which sinusoidal gratings are sampled by a cone mosaic and the direction of motion of the filtered output is computed. The model predicts that the second motion null, but not the first, should be relatively immune to postreceptoral processing and should roughly equal twice the cone Nyquist frequency. This prediction is confirmed by psychophysical experiments, providing a new technique to estimate cone spacing in the living human eye.

Discrimination, Psychological