Heading detection using motion templates and eye velocity gain fields

Eye or head rotation would influence perceived heading direction if it were coded by cells tuned only to retinal flow patterns that correspond to linear self-movement. We propose a model for heading detection based on motion templates that are also Gaussian-tuned to the amount of rotational flow. Such retinal flow templates allow explicit use of extra-retinal signals to create templates tuned to head-centric flow as seen by the stationary eye. Our model predicts an intermediate layer of 'eye velocity gain fields' in which 'rate-coded' eye velocity is multiplied with responses of templates sensitive to specific retinal flow patterns. By combination of the activities of one retinal flow template and many units with an eye velocity gain field, a new type of unit appears: its preferred retinal flow changes dynamically in accordance with the eye rotation velocity. This unit's activity becomes thereby approximately invariant to the amount of eye rotation. The units with eye velocity gain fields from the motion-analogue of the units with eye position gain fields found in area 7a, which according to our general approach, are needed to transform position from retino-centric to head-centric coordinates. The rotation-tuned templates can also provide rate-coded visual estimates of eye rotation to allow a pure visual compensation for rotational flow. Our model is consistent with psychophysical data that indicate a role for extra-retinal as well as visual rotation signals in the correct perception of heading.     

Nitric oxide is involved in the inhibitory neurotransmission and endothelium-dependent relaxations of human small penile arteries

When rotating stripes or other periodic stimuli cross the retina at a critical rate, a reversal in the direction of motion of the stimuli is often seen. This illusion of motion perception was used to explore the roles of retinal and perceived motion in the generation of optokinetic nystagmus. Here we show that optokinetic nystagmus is disrupted during the perception of this illusion. Thus, when perceived and actual motion are in conflict, subjects fail to track the veridical movement. This observation suggests that the perception of motion can directly influence optokinetic nystagmus, even in the presence of a moving retinal image. A conflict in the neural representation of motion in different brain areas may explain these findings.     

A note on formal learning theory

Previous investigations have challenged the generality of the claim that perceived motion in an effective stimulus for smooth pursuit eye movements. The experiments extend the scope of these investigations. Three experiments test the hypothesis that perceived motion can serve as the stimulus for pursuit when the eye movement does not generate constraining retinal error information. Observers viewed retinally stabilized displays that elicited the perception that a stationary target was moving or that a moving target was moving faster than it was actually moving. The results failed to confirm the hypothesis. Relevant literature is reviewed. We conclude that perceived movement can act as a stimulus for pursuit only when the "perceptual target" has no retinal counterpart.     

Guidance of locomotion on foot uses perceived target location rather than optic flow

What visual information do we use to guide movement through our environment? Self-movement produces a pattern of motion on the retina, called optic flow. During translation, the direction of movement (locomotor direction) is specified by the point in the flow field from which the motion vectors radiate - the focus of expansion (FoE) [1-3]. If an eye movement is made, however, the FoE no longer specifies locomotor direction [4], but the 'heading' direction can still be judged accurately [5]. Models have been proposed that remove confounding rotational motion due to eye movements by decomposing the retinal flow into its separable translational and rotational components ([6-7] are early examples). An alternative theory is based upon the use of invariants in the retinal flow field [8]. The assumption underpinning all these models (see also [9-11]), and associated psychophysical [5,12,13] and neurophysiological studies [14-16], is that locomotive heading is guided by optic flow. In this paper we challenge that assumption for the control of direction of locomotion on foot. Here we have explored the role of perceived location by recording the walking trajectories of people wearing displacing prism glasses. The results suggest that perceived location, rather than optic or retinal flow, is the predominant cue that guides locomotion on foot.     

Biological movements look uniform: Evidence of motor-perceptual interactions.

Demonstrated a visual dynamic illusion in 6 experiments. Previous work has shown that in 2-dimensional (2-D) drawing movements, tangential velocity and radius of curvature covary in a constrained manner. The velocity of point stimuli is perceived as uniform if and only if this biological constraint is satisfied. The illusion is conspicuous: The variations of velocity in the stimuli exceed 200%. Yet movements are perceived as uniform. Conversely, 2-D stimuli moving at constant velocity are perceived as strongly nonuniform. The illusion is robust: Exposure to true constant velocity fails to suppress it. Results cannot be explained entirely by the kinetic depth effect. The illusion is evidence of a coupling between motor and perceptual processes: Even in the absence of any intention to perform a movement, certain properties of the motor system implicitly influence perceptual interpretation of the visual stimulus. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Estimating heading during real and simulated eye movements

The ability to judge heading during tracking eye movements has recently been examined by several investigators. To assess the use of retinal-image and extra-retinal information in this task, the previous work has compared heading judgments with executed as opposed to simulated eye movements. For eye movement velocities greater than 1 deg/sec, observers seem to require the eye-velocity information provided by extra-retinal signals that accompany tracking eye movements. When those signals are not provided, such as with simulated eye movements, observers perceive their self-motion as curvilinear translation rather than the linear translation plus eye rotation being presented. The interpretation of the previous results is complicated, however, by the fact that the simulated eye movement condition may have created a conflict between two possible estimates of the heading: one based on extra-retinal solutions and the other based on retina-image solutions. In four experiments, we minimized this potential conflict by having observers judge heading in the presence of rotations consisting of mixtures of executed and simulated eye movements. The results showed that the heading is estimated more accurately when rotational flow is created by executed eye movements alone. In addition, the magnitude of errors in heading estimates is essentially proportional to the amount of rotational flow created by a simulated eye rotation (independent of the total magnitude of the rotational flow). The fact that error magnitude is proportional to the amount of simulated rotation suggests that the visual system attributes rotational flow unaccompanied by an eye movement to a displacement of the direction of translation in the direction of the simulated eye rotation.     

Depth perception in motion parallax and stereokinesis.

Perceived depth in the stereokinetic effect (SKE) illusion and in the monocular derivation of depth from motion parallax were compared. Motion parallax gradients of velocity can be decomposed into 2 components: object- and observer-relative transformations. SKE displays present only the object-relative component. Observers were asked to estimate the magnitude and near–far order of depth in motion parallax and SKE displays. Monocular derivation of depth magnitude from motion parallax is fully accounted for by the perceptual response to the SKE, and observer-relative transformations absent in the SKE are of perceptual utility only as determinants of the near–far signing of perceived sequential depth. The amount of depth and rigidity perceived in motion parallax and SKE displays covaries with the projective size of the stimuli. The monocular derivation of depth from motion is mediated by a perceptual heuristic of which the SKE is symptomatic. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Temporal properties of visual motion signals for the initiation of smooth pursuit eye movements in monkeys

1. Our goal was to assess whether visual motion signals related to changes in image velocity contribute to pursuit eye movements. We recorded the smooth eye movements evoked by ramp target motion at constant speed. In two different kinds of stimuli, the onset of target motion provided either an abrupt, step change in target velocity or a smooth target acceleration that lasted 125 ms followed by prolonged target motion at constant velocity. We measured the eye acceleration in the first 100 ms of pursuit. Because of the 100-ms latency from the onset of visual stimuli to the onset of smooth eye movement, the eye acceleration in this 100-ms interval provides an estimate of the open-loop response of the visuomotor pathways that drive pursuit. 2. For steps of target velocity, eye acceleration in the first 100 ms of pursuit depended on the "motion onset delay," defined as the interval between the appearance of the target and the onset of motion. If the motion onset delay was > 100 ms, then the initial eye movement consisted of separable early and late phases of eye acceleration. The early phase dominated eye acceleration in the interval from 0 to 40 ms after pursuit onset and was relatively insensitive to image speed. The late phase dominated eye acceleration in the interval 40-100 ms after the onset of pursuit and had an amplitude that was proportional to image speed. If there was no delay between the appearance of the target and the onset of its motion, then the early component was not seen, and eye acceleration was related to target speed throughout the first 100 ms of pursuit. 3. For step changes of target velocity, the relationship between eye acceleration in the first 40 ms of pursuit and target velocity saturated at target speeds > 10 degrees /s. In contrast, the relationship was nearly linear when eye acceleration was measured in the interval 40-100 ms after the onset of pursuit. We suggest that the first 40 ms of pursuit are driven by a transient visual motion input that is related to the onset of target motion (motion onset transient component) and that the next 60 ms are driven by a sustained visual motion input (image velocity component). 4. When the target accelerated smoothly for 125 ms before moving at constant speed, the initiation of pursuit resembled that evoked by steps of target velocity. However, the latency of pursuit was consistently longer for smooth target accelerations than for steps of target velocity.(ABSTRACT TRUNCATED AT 400 WORDS)     

Global induced motion and visual stability in an optic flow illusion

When an expansion flow field of moving dots is overlapped by planar motion, observers perceive an illusory displacement of the focus of expansion (FOE) in the direction of the planar motion (Duffy and Wurtz, Vision Research, 1993;33:1481-1490). The illusion may be a consequence of induced motion, wherein an induced component of motion relative to planar dots is added to the motions of expansion dots to produce the FOE shift. While such a process could be mediated by local 'center-surround' receptive fields, the effect could also be due to a higher level process which detects and subtracts large-field planar motion from the flow field. We probed the mechanisms underlying this illusion by adding varying amounts of rotation to the expansion stimulus, and by varying the speed and size of the planar motion field. The introduction of rotation into the stimulus produces an illusory shift in a direction perpendicular to the planar motion. Larger FOE shifts were perceived for greater speeds and sizes of planar motion fields, although the speed effect saturated at high speeds. While the illusion appears to share a common mechanism with center-surround induced motion, our results also point to involvement of a more global mechanism that subtracts coherent planar motion from the flow field. Such a process might help to maintain visual stability during eye movements.     

Estimating heading during eye movements

In eight experiments, we examined the ability to judge heading during tracking eye movements. To assess the use of retinal-image and extra-retinal information in this task, we compared heading judgments with executed as opposed to simulated eye movements. In general, judgments were much more accurate during executed eye movements. Observers in the simulated eye movement condition misperceived their self-motion as curvilinear translation rather than the linear translation plus eye rotation that was simulated. There were some experimental conditions in which observers could judge heading reasonably accurately during simulated eye movements; these included conditions in which eye movement velocities were 1 deg/sec or less and conditions which made available a horizon cue that exists for locomotion parallel to a ground plane with a visible horizon. Overall, our results imply that extra-retinal, eye-velocity signals are used in determining heading under many, perhaps most, viewing conditions.     

Reversed visual motion and self-sustaining eye oscillations

A random-dot field undergoing counterphase flicker paradoxically appears to move in the same direction as head and eye movements, i.e. opposite to the optic-flow field. The effect is robust and occurs over a wide range of flicker rates and pixel sizes. The phenomenon can be explained by reversed phi motion caused by apparent pixel movement between successive retinal images. The reversed motion provides a positive feedback control of the display, whereas under normal conditions retinal signals provide a negative feedback. This altered polarity invokes self-sustaining eye movements akin to involuntary optokinetic nystagmus.     

Self-motion path discrimination: effects of image stabilization

Path-deviation thresholds were measured as the effects of eye movements in the retinal flow were minimized through image stabilization. Thresholds obtained with image stabilization were compared to those obtained with unstabilized viewing to determine whether the elimination of eye movements from the retinal flow improves self-motion judgments. The results showed that, at slow forward speeds, eliminating the retinal effects of eye movements did not improve path-discrimination performance; subjects required more of an angular deviation to discriminate a circular from a straight motion path with image stabilization than with unstabilized viewing. In an effort to understand the results, eye movements were measured in unstabilized viewing conditions, and the measured eye velocities were used to estimate the retinal-image motion. The results showed that, for slow forward speeds, eye movements increased the average retinal speed, independent of the circular flow direction. At fast forward speeds, there was no significant increase in the average retinal-image speed due to eye movements. A parsimonious explanation for the decreased performance with image stabilization at the slow forward speed is that retinal-image motion was too slow to optimally stimulate the visual motion sensors.     

Perception of timing of kinesthetic stimuli

A psychophysical method was used to estimate the timing of perception of kinesthetic stimuli with different velocities in normal volunteers. A 1 ms auditory click occurred randomly before or after an imposed flexion movement at either 20, 40 or 60 deg/s of the metacarpophalangeal joint. Subjects reported whether the click was perceived before or after the movement onset (experiment 1) or perception of movement velocity (experiment 2). The time at which there was a 50% chance that subjects reported movement or velocity perception after the click was taken as an estimate of the time subjects perceived the stimuli. The difference in time of perceived movement velocity discrimination and movement onset was only significant when the velocity was 20 deg/s (52 ms). This suggests that movement onset and identification of the velocity of the faster movements are perceived nearly simultaneously.     

Visual perception modifies goal-directed movement control: supporting evidence from a visual perturbation paradigm

It is well known that dynamic visual information influences movement control, whereas the role played by background visual information is still largely unknown. Evidence coming mainly from eye movement and manual tracking studies indicates that background visual information modifies motion perception and might influence movement control. The goal of the present study was to test this hypothesis. Subjects had to apply pressure on a strain gauge to displace in a single action a cursor shown on a video display and to immobilize it on a target shown on the same display. In some instances, the visual background against which the cursor moved was unexpectedly perturbed in a direction opposite to (Experiment 1), or in the same direction as (Experiment 2) the cursor controlled by the subject. The results of both experiments indicated that the introduction of a visual perturbation significantly affected aiming accuracy. These results suggest that background visual information is used to evaluate the velocity of the aiming cursor, and that this perceived velocity is fed back to the control system, which uses it for on-line corrections.     

Pursuit eye movements and the depth-movement phenomenon in dynamic visual noise

The ability to generate voluntary pursuit eye movements in the absence of retinal-contour motion cues was assessed on the basis of observers' perceptions of depth and motion when they viewed dynamic visual noise with a filter over one eye. The results indicated that the depth-movement phenomenon yielded robust pursuit with the velocity an inverse function of filter density. These data suggest that retinal-contour motion cues are not necessary and that perceived motion is sufficient to drive pursuit.     

Postsaccadic enhancement of initiation of smooth pursuit eye movements in monkeys

Step-ramp target motion evokes a characteristic sequence of presaccadic smooth eye movement in the direction of the target ramp, catch-up targets to bring eye position close to the position of the moving target, and postsaccadic eye velocities that nearly match target velocity. I have analyzed this sequence of eye movements in monkeys to reveal a strong postsaccadic enhancement of pursuit eye velocity and to document the conditions that lead to that enhancement. Smooth eye velocity was measured in the last 10 ms before and the first 10 ms after the first saccade evoked by step-ramp target motion. Plots of eye velocity as a function of time after the onset of the target ramp revealed that eye velocity at a given time was much higher if measured after versus before the saccade. Postsaccadic enhancement of pursuit was recorded consistently when the target stepped 3 degrees eccentric on the horizontal axis and moved upward, downward, or away from the position of fixation. To determine whether postsaccadic enhancement of pursuit was invoked by smear of the visual scene during a saccade, I recorded the effect of simulated saccades on the presaccadic eye velocity for step-ramp target motion. The 3 degrees simulated saccade, which consisted of motion of a textured background at 150 degrees/s for 20 ms, failed to cause any enhancement of presaccadic eye velocity. By using a strategically selected set of oblique target steps with horizontal ramp target motion, I found clear enhancement for saccades in all directions, even those that were orthogonal to target motion. When the size of the target step was varied by up to 15 degrees along the horizontal meridian, postsaccadic eye velocity did not depend strongly either on the initial target position or on whether the target moved toward or away from the position of fixation. In contrast, earlier studies and data in this paper show that presaccadic eye velocity is much stronger when the target is close to the center of the visual field and when the target moves toward versus away from the position of fixation. I suggest that postsaccadic enhancement of pursuit reflects activation, by saccades, of a switch that regulates the strength of transmission through the visual-motor pathways for pursuit. Targets can cause strong visual motion signals but still evoke low presaccadic eye velocities if they are ineffective at activating the pursuit system.     

Directional organization of eye movement and visual signals in the floccular lobe of the monkey cerebellum

The floccular lobe of the monkey is critical for the generation of visually-guided smooth eye movements. The present experiments reveal physiological correlates of the directional organization in the primate floccular lobe by examining the selectivity for direction of eye motion and visual stimulation in the firing of individual Purkinje cells (PCs) and mossy fibers. During tracking of sinusoidal target motion along different axes in the frontoparallel plane, PCs fell into two classes based on the axis that caused the largest modulation of simple-spike firing rate. For "horizontal" PCs, the response was maximal during horizontal eye movements, with increases in firing rate during pursuit toward the side of recording (ipsiversive). For "vertical" PCs, the response was maximal during eye movement along an axis just off pure vertical, with increases in firing rate during pursuit directed downward and slightly contraversive. During pursuit of target motion at constant velocity, PCs again fell into horizontal and vertical classes that matched the results from sinusoidal tracking. In addition, the directional tuning of the sustained "eye velocity" and transient "visual" components of the neural responses obtained during constant velocity tracking were very similar. PCs displayed very broad tuning approximating a cosine tuning curve; the mean half-maximum bandwidth of their tuning curves was 170-180 degrees. Other cerebellar elements, related purely to eye movement and presumed to be mossy fibers, exhibited tuning approximately 40 degrees narrower than PCs and had best directions that clustered around the four cardinal directions. Our data indicate that the motion signals encoded by PCs in the monkey floccular lobe are segregated into channels that are consistent with a coordinate system defined by the vestibular apparatus and eye muscles. The differences between the tuning properties exhibited by PCs compared with mossy fibers indicate that a spatial transformation occurs within the floccular lobe.     

A jitter after-effect reveals motion-based stabilization of vision

A shaky hand holding a video camera invariably turns a treasured moment into an annoying, jittery momento. More recent consumer cameras thoughtfully offer stabilization mechanisms to compensate for our unsteady grip. Our eyes face a similar challenge in that they are constantly making small movements even when we try to maintain a fixed gaze. What should be substantial, distracting jitter passes completely unseen. Position changes from large eye movements (saccades) seem to be corrected on the basis of extraretinal signals such as the motor commands sent to the eye muscle, and the resulting motion responses seem to be simply switched off. But this approach is impracticable for incessant, small displacements, and here we describe a novel visual illusion that reveals a compensation mechanism based on visual motion signals. Observers were adapted to a patch of dynamic random noise and then viewed a larger pattern of static random noise. The static noise in the unadapted regions then appeared to 'jitter' coherently in random directions. Several observations indicate that this visual jitter directly reflects fixational eye movements. We propose a model that accounts for this illusion as well as the stability of the visual world during small and/or slow eye movements such as fixational drift, smooth pursuit and low-amplitude mechanical vibrations of the eyes.     

Taking a long look at action and time perception.

Everyone has probably experienced chronostasis, an illusion of time that can cause a clock's second hand to appear to stand still during an eye movement. Though the illusion was initially thought to reflect a mechanism for preserving perceptual continuity during eye movements, an alternative hypothesis has been advanced that overestimation of time might be a general effect of any action. Contrary to both of these hypotheses, the experiments reported here suggest that distortions of time perception related to an eye movement are not distinct from temporal distortions for other kinds of responses. Moreover, voluntary action is neither necessary nor sufficient for overestimation effects. These results lead to a new interpretation of chronostasis based on the role of attention and memory in time estimation. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

A model for the fluid motion of vitreous humour of the human eye during saccadic movement

During saccadic motion the eyewall moves in a manner similar to a sinusoid or at least can be represented by a sine Fourier series. Motion of the vitreous is induced by the saccade and the vitreo-retinal interface is subjected to a time-dependent shear. This force may be a significant factor for retinal tearing in the neighbourhood of small retinal holes or tears. An analytical viscoelastic model and a numerical, Newtonian model of the motion of the vitreous are presented and compared. Under sinusoidal boundary motion the analytical model shows that a viscous wave propagates inward toward the axis of rotation and the characteristic length of this wave is a function of the Womersley number. The numerical solution indicates that the vitreous moves similarly to the analytical result with small secondary motion; however, this motion allows complete recirculation of the vitreous over large timescales. Excellent agreement is found between the analytical and numerical models. The time-dependent fluid shear is evaluated and from the analytical solution the maximum value of this is found to be proportional to R0 square root of v(omega)3, where R0 is the eye radius, v the modified complex viscosity and omega the sinusoidal frequency. This indicates that myopes have a larger shear force exerted on them by virtue of the larger eye size. Further work is directed toward a model which links the stress found in the sclera to that exerted on the vitreo-retinal interface by the vitreous fluid motion.