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.     

Retinal flow is sufficient for steering during observer rotation

How do people control locomotion while their eyes are simultaneously rotating? A previous study found that during simulated rotation, they can perceive a straight path of self-motion from the retinal flow pattern, despite conflicting extraretinal information, on the basis of dense motion parallax and reference objects. Here we report that the same information is sufficient for active control ofjoystick steering. Participants steered toward a target in displays that simulated a pursuit eye movement. Steering was highly inaccurate with a textured ground plane (motion parallax alone), but quite accurate when an array of posts was added (motion parallax plus reference objects). This result is consistent with the theory that instantaneous heading is determined from motion parallax, and the path of self-motion is determined by updating heading relative to environmental objects. Retinal flow is thus sufficient for both perceiving self-motion and controlling self-motion with a joystick; extraretinal and positional information can also contribute, but are not necessary.     

自身运动认知的神经机制

通过光流信息来指导个体在环境中有效移动是我们视觉神经系统的一项核心任务。在灵长类的大脑皮层, 视觉运动的信息加工是由位于背侧通路的一系列脑区来完成的, 这一信息通路主要参与运动和空间动作的分析。在高级视皮层, 视觉系统很可能利用非视觉信息来补偿因眼动造成的光流模式扭曲, 以重建对自身运动方向的正确表征。根据目前研究进展, MST和VIP这两个位于顶叶的脑区都参与了自身运动认知过程, 并且对精确的自身运动方向判断是不可或缺的。本文系统介绍了近些年来在自身运动认知神经机制研究领域的进展, 尤其是神经生理学家们利用非人灵长类动物模型在自身运动认知皮层处理机制方面的成果。同时也提出了一些深入研究急需解决的关键问题。     

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.     

Pursuit compensation during self-motion

The pattern of motion in the retinal image during self-motion contains information about the person's movement. Pursuit eye movements perturb the pattern of retinal-image motion, complicating the problem of self-motion perception. A question of considerable current interest is the relative importance of retinal and extra-retinal signals in compensating for these effects of pursuit on the retinal image. We addressed this question by examining the effect of prior motion stimuli on self-motion judgments during pursuit. Observers viewed 300 ms random-dot displays simulating forward self-motion during pursuit to the right or to the left; at the end of each display a probe appeared and observers judged whether they would pass left or right of it. The display was preceded by a 300 ms dot pattern that was either stationary or moved in the same direction as, or opposite to, the eye movement. This prior motion stimulus had a large effect on self-motion judgments when the simulated scene was a frontoparallel wall (experiment 1), but not when it was a three-dimensional (3-D) scene (experiment 2). Corresponding simulated-pursuit conditions controlled for purely retinal motion aftereffects, implying that the effect in experiment 1 is mediated by an interaction between retinal and extra-retinal signals. In experiment 3, we examined self-motion judgments with respect to a 3-D scene with mixtures of real and simulated pursuit. When real and simulated pursuits were in opposite directions, performance was determined by the total amount of pursuit-related retinal motion, consistent with an extra-retinal 'trigger' signal that facilitates the action of a retinally based pursuit-compensation mechanism. However, results of experiment 1 without a prior motion stimulus imply that extra-retinal signals are more informative when retinal information is lacking. We conclude that the relative importance of retinal and extra-retinal signals for pursuit compensation varies with the informativeness of the retinal motion pattern, at least for short durations. Our results provide partial explanations for a number of findings in the literature on perception of self-motion and motion in the frontal plane.     

Perceiving self-motion in depth: The role of stereoscopic motion and changing-size cues

During self-motions, different patterns of optic flow are presented to the left and right eyes. Previous research has, however, focused mainly on the self-motion information contained in a single pattern of optic flow. The present experiments investigated the role that binocular disparity plays in the visual perception of self-motion, showing that the addition of stereoscopic cues to optic flow significantly improves forward linear vection in central vision. Improvements were also achieved by adding changingsize cues to sparse (but not dense) flow patterns. These findings showed that assumptions in the heading literature that stereoscopic cues facilitate self-motion only when the optic flow has ambiguous depth ordering do not apply to vection. Rather, it was concluded that both stereoscopic and changingsize cues provide additional motion-in-depth information that is used in perceiving self-motion.     

Perception of heading without retinal optic flow

How do we determine where we are heading during visually controlled locomotion? Psychophysical research has shown that humans are quite good at judging their travel direction, or heading, from retinal optic flow. Here we show that retinal optic flow is sufficient, but not necessary, for determining heading. By using a purely cyclopean stimulus (random dot cinematogram), we demonstrate heading perception without retinal optic flow. We also show that heading judgments are equally accurate for the cyclopean stimulus and a conventional optic flow stimulus, when the two are matched for motion visibility. The human visual system thus demonstrates flexible, robust use of available visual cues for perceiving heading direction.     

Contribution of extraretinal signals to the scaling of object distance during self-motion

We investigated the role of extraretinal information in the perception of absolute distance. In a computer-simulated environment, monocular observers judged the distance of objects positioned at different locations in depth while performing frontoparallel movements of the head. The objects were spheres covered with random dots subtending three different visual angles. Observers viewed the objects ateye level, either in isolation or superimposed on a ground floor. The distance and size of the spheres were covaried to suppress relative size information. Hence, the main cues to distance were the motion parallax and the extraretinal signals. In three experiments, we found evidence that (1) perceived distance is correlated with simulated distance in terms of precision and accuracy, (2) the accuracy in the distance estimate is slightly improved by the presence of a ground-floor surface, (3) the perceived distance is not altered significantly when the visual field size increases, and (4) the absolute distance is estimated correctly during self-motion. Conversely, stationary subjects failed to report absolute distance when they passively observed a moving object producing the same retinal stimulation, unless they could rely on knowledge of the three-dimensional movements.     

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.     

Simulated angular head oscillation enhances vection in depth

Research has shown that adding simulated linear head oscillation to radial optic flow displays enhances the illusion of self-motion in depth (ie linear vection). We examined whether this oscillation advantage for vection was due to either the added motion parallax or retinal slip generated by insufficient compensatory eye movement during display oscillation. We constructed radial flow displays which simulated 1 Hz horizontal linear head oscillation (generates motion parallax) or angular head oscillation in yaw (generates no motion parallax). We found that adding simulated angular or linear head oscillation to radial flow increased the strength of linear vection in depth. Neither type of simulated head oscillation significantly reduced vection onset latencies relative to pure radial flow. Simultaneous eye-movement recordings showed that slow-phase ocular following responses (OFRs) were induced in both linear and angular viewpoint oscillation conditions. Vection strength was significantly reduced by active central fixation when viewing displays which simulated angular, but not linear, head oscillation. When these displays with angular oscillation were viewed without stable fixation, vection strength was found to increase with the velocity and regularity of the OFR. We conclude that vection improvements observed during central viewing of displays with angular viewpoint oscillation depend on the generation of eye movements.     

The role of central and peripheral vision in perceiving the direction of self-motion.

Three experiments were performed to examine the role that central and peripheral vision play in the perception of the direction of translational self-motion, or heading, from optical flow. When the focus of radial outflow was in central vision, heading accuracy was slightly higher with central circular displays (10 degrees-25 degrees diameter) than with peripheral annular displays (40 degrees diameter), indicating that central vision is somewhat more sensitive to this information. Performance dropped rapidly as the eccentricity of the focus of outflow increased, indicating that the periphery does not accurately extract radial flow patterns. Together with recent research on vection and postural adjustments, these results contradict the peripheral dominance hypothesis that peripheral vision is specialized for perception of self-motion. We propose a functional sensitivity hypothesis--that self-motion is perceived on the basis of optical information rather than the retinal locus of stimulation, but that central and peripheral vision are differentially sensitive to the information characteristic of each retinal region.     

The role of central and peripheral vision in perceiving the direction of self-motion

Three experiments were performed to examine the role that central and peripheral vision play in the perception of the direction of translational self-motion, or heading, from optical flow. When the focus of radial outflow was in central vision, heading accuracy was slightly higher with central circular displays (10°–25° diameter) than with peripheral annular displays (40° diameter), indicating that central vision is somewhat more sensitive to this information. Performance dropped rapidly as the eccentricity of the focus of outflow increased, indicating that the periphery does not accurately extract radial flow patterns. Together with recent research on vection and postural adjustments, these results contradict theperipheral dominance hypothesis that peripheral vision is specialized for perception of self-motion. We propose afunctional sensitivity hypothesis—that. self-motion is perceived on the basis of optical information rather than the retinal locus of stimulation, but that central and peripheral vision are differentially sensitive to the information characteristic of each retinal region.     

Auditory clicks distort perceived velocity but only when the system has to rely on extraretinal signals

Previous work has found that repetitive auditory stimulation (click trains) increases the subjective velocity of subsequently presented moving stimuli. We ask whether the effect of click trains is stronger for retinal velocity signals (produced when the target moves across the retina) or for extraretinal velocity signals (produced during smooth pursuit eye movements, when target motion across the retina is limited). In Experiment 1, participants viewed leftward or rightward moving single dot targets, travelling at speeds from 7.5 to 17.5 deg/s. They estimated velocity at the end of each trial. Prior presentation of auditory click trains increased estimated velocity, but only in the pursuit condition, where estimates were based on extraretinal velocity signals. Experiment 2 generalized this result to vertical motion. Experiment 3 found that the effect of clicks during pursuit disappeared when participants tracked across a visually textured background that provided strong local motion cues. Together these results suggest that auditory click trains selectively affect extraretinal velocity signals. This novel finding suggests that the cross-modal integration required for auditory click trains to influence subjective velocity operates at later stages of processing.     

The discrimination of heading from optic flow is not retinally invariant.

Three experiments were conducted to determine whether the discrimination of heading from optic flow is retinally invariant and to determine the importance of acuity in accounting for heading eccentricity effects. In the first experiment, observers were presented with radial flow fields simulating forward translation through a three-dimensional volume of dots. The flow fields subtended 10 degrees of visual angle and were presented at 0 degree, 10 degrees, 20 degrees, and 40 degrees of retinal eccentricity. The observers were asked to indicate whether the simulated movement was to the right or the left of a target that appeared at the end of the display sequence. Eye movements were monitored with an electrooculogram apparatus. In a second experiment, static acuity thresholds were derived for each of the observers at the same retinal eccentricities. There was a significant increase in heading detection thresholds with retinal eccentricity (from 0.92 degree at 0 degree retinal eccentricity to 3.47 degrees at 40 degrees). An analysis of covariance indicated that the variation in sensitivity to radial flow, as a function of retinal eccentricity, is independent of acuity. Similar results were obtained when the Vernier acuity of observers was measured. These results suggest that the discrimination of heading from radial flow is not retinally invariant.     

Stimulus eccentricity and spatial frequency interact to determine circular vection.

While early research suggested that peripheral vision dominates the perception of self-motion, subsequent studies found little or no effect of stimulus eccentricity. In contradiction to these broad notions of 'peripheral dominance' and 'eccentricity independence', the present experiments showed that the spatial frequency of optic flow interacts with its eccentricity to determine circular vection magnitude--central stimulation producing the most compelling vection for high-spatial-frequency stimuli and peripheral stimulation producing the most compelling vection for lower-spatial-frequency stimuli. This interaction appeared to be due, in part at least, to the effect that the higher-spatial-frequency moving pattern had on subjects' ability to organise optic flow into related motion about a single axis. For example, far-peripheral exposure to this high-spatial-frequency pattern caused many subjects to organise the optic flow into independent local regions of motion (a situation which clearly favoured the perception of object motion not self-motion). It is concluded that both high-spatial-frequency and low-spatial-frequency mechanisms are involved in the visual perception of self-motion--with their activities depending on the nature and eccentricity of the motion stimulation.     

The discrimination of heading from optic flow is not retinally invariant

Three experiments were conducted to determine whether the discrimination of heading from optic flow is retinally invariant and to determine the importance of acuity in accounting for heading eccentricity effects. In the first experiment, observers were presented with radial flow fields simulating forward translation through a three-dimensional volume of dots. The flow fields subtended 10° of visual angle and were presented at 0°, 10°, 20°, and 40° of retinal eccentricity. The observers were asked to indicate whether the simulated movement was to the right or the left of a target that appeared at the end of the display sequence. Eye movements were monitored with an electrooculogram apparatus. In a second experiment, static acuity thresholds were derived for each of the observers at the same retinal eccentricities. There was a significant increase in heading detection thresholds with retinal eccentricity (from 0.92° at 0° retinal eccentricity to 3.47° at 40°). An analysis of covariance indicated that the variation in sensitivity to radial flow, as a function of retinal eccentricity, is independent of acuity. Similar results were obtained when the Vernier acuity of observers was measured. These results suggest that the discrimination of heading from radial flow is not retinally invariant.     

Stability in young infants' discrimination of optic flow

Although considerable progress has been made in understanding how adults perceive their direction of self-motion, or heading, from optic flow, little is known about how these perceptual processes develop in infants. In 3 experiments, the authors explored how well 3- to 6-month-old infants could discriminate between optic flow patterns that simulated changes in heading direction. The results suggest that (a) prior to the onset of locomotion, the majority of infants discriminate between optic flow displays that simulate only large (> 22 deg.) changes in heading, (b) there is minimal development in sensitivity between 3 and 6 months, and (c) optic flow alone is sufficient for infants to discriminate heading. These data suggest that spatial abilities associated with the dorsal visual stream undergo prolonged postnatal development and may depend on locomotor experience.     

Coherent modulation of stimulus colour can affect visually induced self-motion perception

The effects of dynamic colour modulation on vection were investigated to examine whether perceived variation of illumination affects self-motion perception. Participants observed expanding optic flow which simulated their forward self-motion. Onset latency, accumulated duration, and estimated magnitude of the self-motion were measured as indices of vection strength. Colour of the dots in the visual stimulus was modulated between white and red (experiment 1), white and grey (experiment 2), and grey and red (experiment 3). The results indicated that coherent colour oscillation in the visual stimulus significantly suppressed the strength of vection, whereas incoherent or static colour modulation did not affect vection. There was no effect of the types of the colour modulation; both achromatic and chromatic modulations turned out to be effective in inhibiting self-motion perception. Moreover, in a situation where the simulated direction of a spotlight was manipulated dynamically, vection strength was also suppressed (experiment 4). These results suggest that observer's perception of illumination is critical for self-motion perception, and rapid variation of perceived illumination would impair the reliabilities of visual information in determining self-motion.     

Optical motions as information for unsigned depth

Optical motions and gradients of retinal flow have been assumed to be an important source of information for the perception of spatial layout. In the case of lateral parallax, however, the complicating effects of smooth eye movements on retinal flow fields and the known insensitivity of the visual system to absolute motion suggest that optical motions alone cannot provide the basis for accurate perception of the direction (sign) of depth relations. At most they can provide information for "unsigned" depth. Results of two experiments support the view that differential optical motions result in a strong impression of separation of objects in depth, but that the determination of near/far relations normally depends on other sources of information.