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。     

Direct perception of action-scaled affordances: the shrinking gap problem

The aim of this study was to investigate the perception of possibilities for action (i.e., affordances) that depend on one's movement capabilities, and more specifically, the passability of a shrinking gap between converging obstacles. We introduce a new optical invariant that specifies in intrinsic units the minimum locomotor speed needed to safely pass through a shrinking gap. Detecting this information during self-motion requires recovering the component of the obstacles' local optical expansion attributable to obstacle motion, independent of self-motion. In principle, recovering the obstacle motion component could involve either visual or non-visual self-motion information. We investigated the visual and non-visual contributions in two experiments in which subjects walked through a virtual environment and made judgments about whether it was possible to pass through a shrinking gap. On a small percentage of trials, visual and non-visual self-motion information were independently manipulated by varying the speed with which subjects moved through the virtual environment. Comparisons of judgments on such catch trials with judgments on normal trials revealed both visual and non-visual contributions to the detection of information about minimum walking speed.     

Self-motion perception influences number processing: evidence from a parity task

We investigated the role of horizontal body motion on the processing of numbers. We hypothesized that leftward self-motion leads to shifts in spatial attention and therefore facilitates the processing of small numbers, and vice versa, we expected that rightward self-motion facilitates the processing of large numbers. Participants were displaced by means of a motion platform during a parity judgment task. We found a systematic influence of self-motion direction on number processing, suggesting that the processing of numbers is intertwined with the processing of self-motion perception. The results differed from known spatial numerical compatibility effects in that self-motion exerted a differential influence on inner and outer numbers of the given interval. The results highlight the involvement of sensory body motion information in higher-order spatial cognition.     

Perceived distance and the perceived speed of self-motion: Linear vs. angular velocity?

Experiments are reported in which it was found that, with the angular speed of a visual surround held constant, the perceived speed of rotary self-motion increased linearly with increasing perceived distance of this surround. This finding was in agreement with a motion constancy equation derived from a consideration of object-referred motion perception. Since information concerning distance is necessary for the perception of linear but not angular speed, this finding supports the conclusion that visually perceived rotary self-motion perception is dependent upon perceived linear surround motion at least in the horizontal plane. The visual motion constancy mechanism which operates for object-referred motion can apparently not be switched off for the special case of self-motion perception.     

Optical flow and viewpoint change modulate the perception and memorization of complex motion

Participants observed a point-light character (PLC) performing a gymnastic movement. They either memorized the final PLC orientation from the initial viewpoint, to match it to a test posture (memory task), or judged whether the biological motion appeared continuous (perceptual task), despite a viewpoint change. The observer could be either static or virtually in motion (pan or track) while looking at the movement from the initial viewpoint. The presence of a spatial layout during virtual self-motion induced a global optical flow specifying the translational component of the PLC movement, rendering the event more predictable for the participants. A representational momentum effect was observed in the memory task, suggesting that when a visual stimulation, such as a PLC motion, is abruptly stopped, its dynamics survive. In contrast, structural and transformational invariants specifying the PLC motion were sufficient to solve the perceptual task accurately. Finally, both the remembering of the final posture and the perception of continuity degraded with an increase in viewpoint change due to tilt/slant posture orientation matching, indicating that orientation processes interfered with event perception.     

Spatial orientation from optic flow in the central visual field

Previous research has shown that stimulation of the central visual field with radial flow patterns (produced by forward motion) can induce perceived self-motion, but has failed to demonstrate effects on postural stability of either radial flow patterns or lamellar flow patterns (produced by horizontal translation) in the central visual field. The present study examined the effects of lamellar and radial flow on postural stability when stimulation was restricted to the central visual field. Displays simulating observer motion through a volume of randomly positioned points were observed binocularly through a window that limited the field of view to 15 degrees. The velocity of each display varied according to the sum of four sine functions of prime frequencies. Changes in posture were used to measure changes in perceived spatial orientation. A frequency analysis of postural sway indicated that increased sway occurred at the frequencies of motion simulated in the display for both lamellar and radial flow. These results suggest that both radial and lamellar optic flow are effective for determining spatial orientation when stimulation is limited to the central visual field.     

Voluntary head movement and allocentric perception of space

Although visual input is egocentric, at least some visual perceptions and representations are allocentric, that is, independent of the observer's vantage point or motion. Three experiments investigated the visual perception of three-dimensional object motion during voluntary and involuntary motion in human subjects. The results show that the motor command contributes to the objective perception of space: Observers are more likely to apply, consciously and unconsciously, spatial criteria relative to an allocentric frame of reference when they are executing voluntary head movements than while they are undergoing similar involuntary displacements (which lead to a more egocentric bias). Furthermore, details of the motor command are crucial to spatial vision, as allocentric bias decreases or disappears when self-motion and motor command do not match.     

Visual distance estimation in static compared to moving virtual scenes

Visual motion is used to control direction and speed of self-motion and time-to-contact with an obstacle. In earlier work, we found that human subjects can discriminate between the distances of different visually simulated self-motions in a virtual scene. Distance indication in terms of an exocentric interval adjustment task, however, revealed linear correlation between perceived and indicated distances but with a profound distance underestimation. One possible explanation for this underestimation is the perception of visual space in virtual environments. Humans perceive visual space in natural scenes as curved, and distances are increasingly underestimated with increasing distance from the observer. Such spatial compression may also exist in our virtual environment. We therefore surveyed perceived visual space in a static virtual scene. We asked observers to compare two horizontal depth intervals, similar to experiments performed in natural space. Subjects had to indicate the size of one depth interval relative to a second interval. Our observers perceived visual space in the virtual environment as compressed, similar to the perception found in natural scenes. However, the nonlinear depth function we found can not explain the observed distance underestimation of visual simulated self-motions in the same environment.     

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.     

When Walking Makes Perception Better

ABSTRACT— When we move, the visual world moves toward us. That is, self-motion normally produces visual signals (flow) that tell us about our own motion. But these signals are distorted by our motion: Visual flow actually appears slower while we are moving than it does when we are stationary and our surroundings move past us. Although for many years these kinds of distortions have been interpreted as a suppression of flow to promote the perception of a stable world, current research has shown that these shifts in perceived visual speed may have an important function in measuring our own self-motion. Specifically, by slowing down the apparent rate of visual flow during self-motion, our visual system is able to perceive differences between actual and expected flow more precisely. This is useful in the control of action.     

Perception of visual speed while moving

During self-motion, the world normally appears stationary. In part, this may be due to reductions in visual motion signals during self-motion. In 8 experiments, the authors used magnitude estimation to characterize changes in visual speed perception as a result of biomechanical self-motion alone (treadmill walking), physical translation alone (passive transport), and both biomechanical self-motion and physical translation together (walking). Their results show that each factor alone produces subtractive reductions in visual speed but that subtraction is greatest with both factors together, approximating the sum of the 2 separately. The similarity of results for biomechanical and passive self-motion support H. B. Barlow's (1990) inhibition theory of sensory correlation as a mechanism for implementing H. Wallach's (1987) compensation for self-motion.     

Perception of self-motion from visual flow

Accurate and efficient control of self-motion is an important requirement for our daily behavior. Visual feedback about self-motion is provided by optic flow. Optic flow can be used to estimate the direction of self-motion (‘heading’) rapidly and efficiently. Analysis of oculomotor behavior reveals that eye movements usually accompany self-motion. Such eye movements introduce additional retinal image motion so that the flow pattern on the retina usually consists of a combination of self-movement and eye movement components. The question of whether this ‘retinal flow’ alone allows the brain to estimate heading, or whether an additional ‘extraretinal’ eye movement signal is needed, has been controversial. This article reviews recent studies that suggest that heading can be estimated visually but extraretinal signals are used to disambiguate problematic situations. The dorsal stream of primate cortex contains motion processing areas that are selective for optic flow and self-motion. Models that link the properties of neurons in these areas to the properties of heading perception suggest possible underlying mechanisms of the visual perception of self-motion.     

Similar effects of visual perception and imagery on simple reaction time

A longstanding issue is whether perception and mental imagery share similar cognitive and neural mechanisms. To cast further light on this problem, we compared the effects of real and mentally generated visual stimuli on simple reaction time (RT). In five experiments, we tested the effects of difference in luminance, contrast, spatial frequency, motion, and orientation. With the intriguing exception of spatial frequency, in all other tasks perception and imagery showed qualitatively similar effects. An increase in luminance, contrast, and visual motion yielded a decrease in RT for both visually presented and imagined stimuli. In contrast, gratings of low spatial frequency were responded to more quickly than those of higher spatial frequency only for visually presented stimuli. Thus, the present study shows that basic dependent variables exert similar effects on visual RT either when retinally presented or when imagined. Of course, this evidence does not necessarily imply analogous mechanisms for perception and imagery, and a note of caution in such respect is suggested by the large difference in RT between the two operations. However, the present results undoubtedly provide support for some overlap between the structural representation of perception and imagery.     

Similar effects of visual perception and imagery on simple reaction time

A longstanding issue is whether perception and mental imagery share similar cognitive and neural mechanisms. To cast further light on this problem, we compared the effects of real and mentally generated visual stimuli on simple reaction time (RT). In five experiments, we tested the effects of difference in luminance, contrast, spatial frequency, motion, and orientation. With the intriguing exception of spatial frequency, in all other tasks perception and imagery showed qualitatively similar effects. An increase in luminance, contrast, and visual motion yielded a decrease in RT for both visually presented and imagined stimuli. In contrast, gratings of low spatial frequency were responded to more quickly than those of higher spatial frequency only for visually presented stimuli. Thus, the present study shows that basic dependent variables exert similar effects on visual RT either when retinally presented or when imagined. Of course, this evidence does not necessarily imply analogous mechanisms for perception and imagery, and a note of caution in such respect is suggested by the large difference in RT between the two operations. However, the present results undoubtedly provide support for some overlap between the structural representation of perception and imagery.     

State reversals of optically induced tilt and torsional eye movements

Alternations of the state of apparent self-motion during observation of a large visual display rotating about the line of sight are associated with alternations in the magnitude of induced tilt and torsional eye rotation. In one experiment, shifts in visually induced tilt during these state alternations are found to be in the opposite direction to corresponding shifts in induced ocular torsion. In a second experiment, the reversals of self-motion perception are shown to be an intravisual phenomenon, independent of competing inputs provided by the vestibular system. These results emphasize the importance of distinguishing between visual and vestibular processes in tilt perception and ocular rotation during human orientation to gravitational vertical.     

Chromaticity, spatial complexity, and self-motion perception

The effects of visual field color and spatial complexity on self-motion perception were investigated by placing observers inside a large rotating cylinder (optokinetic drum). Under optokinetic-drum conditions visually induced self-motion (vection) is typically perceived within 30 s, even though all forms of sensory input (eg vestibular, proprioceptive, auditory), except vision, indicate that the observer is stationary. It was hypothesized that vection would be hastened and vection magnitude increased by adding chromatic colors and spatial complexity to the lining of an optokinetic drum. Addition of these visual-field characteristics results in an array that shares more visual-field characteristics with our typical environment that usually serves as a stable frame of reference regarding self-motion perception. In the color experiment, participants viewed vertical stripes that were: (i) black and white, (ii) various gray shades, or (iii) chromatic. In the spatial complexity experiment, participants were presented with: (i) black-and-white vertical stripes, or (ii) a black-and-white checkerboard pattern. Drum rotation velocity was 5 rev. min(-1) (30 degrees s(-1)), and both vection onset and magnitude were measured for 60 s trials. Results indicate that chromaticity and spatial complexity hasten the onset of vection and increase its perceived magnitude. Chromaticity and spatial complexity are common characteristics of the environments in which our visual system evolved. The presence of these visual-field features in an optic flow pattern may be treated as an indicator that the scene being viewed is stationary and that the observer is moving.     

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.     

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.     

The role of local and distal landmarks in the development of object location memory

To locate objects in the environment, animals and humans use visual and nonvisual information. We were interested in children's ability to relocate an object on the basis of self-motion and local and distal color cues for orientation. Five- to 9-year-old children were tested on an object location memory task in which, between presentation and test, the availability of local and distal cues was manipulated. Additionally, participants' viewpoint could be changed. We used a Bayesian model selection approach to compare our hypotheses. We found that, to remain oriented in space, 5-year-olds benefit from visual information in general, 7-year-olds benefit from visual cues when a viewpoint change takes place, and 9-year-olds do not benefit from the availability of visual cues for orientation but rely on self-movement cues instead. Results are discussed in terms of the adaptive combination model (Newcombe & Huttenlocher, 2006).