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At least 55 records · Page 3

Effects of voluntary eye movement and convergence on the binocular appreciation of depth.

Scaling techniques were employed to establish the relation between perceived distance ratio and physical distance ratio. Measurements were made both with and without free eye movement and under two states of convergence. The results were confirmed using a matching technique. With free eye movement, the perceived ratio is a monotonic increasing function of the physical ratio. Without eye movement, the perceived ratio generally increases, then decreases, as the physical ratio increases. For a given physical ratio, perceived distance ratio is less in the absence of voluntary eye movements. Convergence produces depth micropsia when eye movements are permitted, but not in their absence.

Foley, J. M.↗

Perceiving the coherent movements of spatially-separated features

When a partially-occluded object is represented in an image, it is defined by a set of spatially-separated features that may be registered at different spatial scales. To understand the image, human vision must organize these fragmented optical features into common and distinct object surfaces. Although the common fate of moving features often is considered a primary source of reliable information for image segmentation, little is known of the visual system's capacity to discriminate the coherent relative movements of spatially-separated features. In a series of experiments, observers viewed elements whose movements were correlated (direction and magnitude) or were uncorrelated. Our results indicate that observers can discriminate the two types of movement about as well as they can detect any movement at all. Moreover, the ability to perceive coherent movements is maintained under a variety of conditions including differences in the elements' spatial frequency content, spatial position and contrast, and temporal phase shifts between the spatially-correlated displacements. These results suggest that coherent relative motion may be a fundamental source of information exploited by vision, despite considerable variability in the spatial and temporal characteristics of the individual features.

Mowafy, Lyn↗

Torsional Eye Movements Evoked by Unilateral Labyrinthine Galvanic Polarizations in the Squirrel Monkey

Electrical stimulation of vestibular-nerve afferents innervating the semicircular canals has been used to identify the extraocular muscles receiving activation or inhibition by individual ampullary nerves. This technique was originally developed by Szentagothai (1950) and led to the description of three neuron reflex arcs that connect each semicircular canal through an interneuron traversing in the region of the medial longitudinal fasciculus to one ipsilateral and one contralateral eye muscle. Selective ampullary nerve stimulation was subsequently used by Cohen and colleagues (Cohen and Suzuki, 1963; Cohen et al., 1964; Suzuki et al., 1964; Cohen et al., 1966) to study movements of the eyes and activation of individual extraocular muscles in response to stimulation of combinations of ampullary nerves. This work led to a description of the now familiar relationships between activation of a semicircular canal ampullary nerves and the anticipated movement in each eye. Disconjugacy of eye movements induced by individual vertical canal stimulation and dependence of the pulling direction of vertical recti and oblique muscles on eye position were also defined in these experiments. Subsequent studies have defined the mechanisms by which externally applied galvanic currents result in a change in vestibular-nerve afferent discharge. The currents appear to act at the spike trigger site. Perilymphatic cathodal currents depolarize the trigger site and lead to excitation whereas anodal currents hyperpolarize and result in inhibition. Afferents innervating all five vestibular endorgans appear to be affected equally by the currents (Goldberg et al., 1984). Irregularly discharging afferents are about 5-10 times more sensitive than regularly discharging ones because of the steeper slope of the former's faster postspike recovery of excitability in encoder sensitivity (Smith and Goldberg, 1986). Response adaptation similar to that noted during acceleration steps is apparent for longer periods of current administration. This adaptation is manifested as a perstimulus return toward resting discharge and poststimulus after-response in the opposite direction (Goldberg et al., 1984; Minor and Goldberg, l991). Cathodal currents (with respect to the perilymphatic space of the vestibule) are excitatory whereas anodal currents are inhibitory. Horizontal eye movements evoked by unilateral galvanic polarizations administered through chronically implanted labyrinthine stimulating electrodes have been studied in alert squirrel monkeys (Minor and Goldberg, 1991). We sought to extend this analysis by recording three-dimensional eye movements during galvanic stimulation. As predicted based upon roughly equal stimulation of ampullary nerves innervating the vertical canals, a substantial torsional component to the nystagmus is noted. The trajectory of torsional slow phases and nystagmus profile after the polarization provide insight into the central mechanisms that influence these responses.

Minor, Lloyd B.↗

Head and Trunk Movement Control During Locomotion After Long-Duration Spaceflight

Exposure to the microgravity environment encountered during spaceflight induces adaptive alteration in sensorimotor function that leads to postflight disturbances in locomotor control. Head and trunk movement control plays a central role in maintaining gaze stability and in providing a stable reference system to permit spatial navigation in a complex and constantly varying environment. The goal of the present study was to investigate the effects of long-duration spaceflight (3-6 months) on head and trunk movement control during postflight terrestrial locomotion. Before and after spaceflight, subjects walked on a motorized treadmill while performing a challenging gaze stabilization task requiring number recognition. Head and trunk kinematic data were collected with a video-based motion analysis system. Analysis of roll, pitch and yaw head and trunk movements during treadmill walking revealed postflight alterations in head and trunk movement control in all three planes of motion. Subjects also experienced oscillopsia during postflight walking which led to impairment in performance of the number recognition task. These data indicate that exposure to long-duration space flight causes alteration in head and trunk movement control during postflight locomotion. These changes have implications for the control of gaze and maintenance of dynamic stability during walking after long-duration spaceflight.

Bloomberg, Jacob J.↗

Vertex Movement for Mission Status Graphics: A Polar-Star Display

Humans are traditionally bad monitors, especially over long periods of time on reliable systems, and they are being called upon to do this more and more as systems become further automated. Because of this, there is a need to find a way to display the monitoring information to the human operator in such a way that he can notice pertinent deviations in a timely manner. One possible solution is to use polar-star displays that will show deviations from normal in a more salient manner. A polar-star display uses a polygon's vertices to report values. An important question arises, though, of how the vertices should move. This experiment investigated two particular issues of how the vertices should move: (1) whether the movement of the vertices should be continuous or discrete and (2) whether the parameters that made up each vertex should always move in one direction regardless of parameter sign or move in both directions indicating parameter sign. The results indicate that relative movement direction is best. Subjects performed better with this movement type and they subjectively preferred it to the absolute movement direction. As for movement type, no strong preferences were shown.

Trujillo, Anna↗

Analysis of the Trial-By-Trial Correlation Between Eye Movement and Perceptual Responses to Moving Plaids

We have previously shown that windowing a drifting plaid with an asymmetric spatial Gaussian produces systematic biases of similar magnitude in both the perceived direction of motion and the direction of the eye movement response. To further investigate this, we simultaneously measured the pyschophysical and eye-movement responses (with an ISCAN RK426 IR tracker)to drifting plaids in a direction discrimination task. Three observers were instructed to track a plaid (Type I 90 deg.; TF = 4 Hz, SF = 0.6 c/d; windowed by a circular spatial gaussian) and to respond whether the motion was to the right or left of pure vertical. Three plaid directions were presented (-2, 0, 2 deg. with respect to straight down), so there was uncertainty in the perceptual judgements. This enabled us to examine the trial-by-trial relationship between the eye movements and psychophysical responses. The eye-movement direction was computed to be the slope of the best fitting line to the initial 300 ms of saccade-free tracking (near open-loop). Again, we found the mean psychophysical and oculomotor responses were correlated. The mean percentages of rightward responses were 6, 27, 87%, and the mean normalized eye-movement directions were -1.5, 0, and 1.0 deg. for the -2, 0, and 2 deg. stimuli respectively. We also computed the trial-by-trial correlation by performing an SOC analysis. The mean correlations were 0.52, 0.66, and 0.67 for our three observers. Because of eye-tracker measurement noise, the above results are lower limits on the actual correlation. These results support the view that the oculomotor and perceptual system share common motion inputs.

Beutter, Brent R.↗

Eye Movements Reveal Hierarchical Motion Processing

Purpose: In the analysis of visual motion, local features such as orientation are analyzed early in the cortical processing stream (V1), while integration across orientation and space is thought to occur in higher cortical areas such as MT, MST, etc. If all areas provide inputs to eye movement control centers, we would expect that local properties would drive eye movements with relatively short latencies, while global properties would require longer latencies. When such latencies are observed, they can provide information about when (and where?) various stimulus properties are analyzed. Methods: The stimulus employed was an elliptical Gabor patch with a drifting carrier, in which the orientations of the carrier grating and the contrast window were varied independently. We have previously demonstrated that the directional percepts evoked by this stimulus vary between the "grating direction" (the normal to the grating's orientation) and the "window direction", and that similar effects can be observed in reflexive eye movements. Subjects viewed such a stimulus while attempting to maintain steady fixation on the center of the pattern, and the small reflexive eye movements ("stare OKN") were recorded. In the middle of the trial, the orientation of either the grating or the window was rotated smoothly by 30 degrees. Results: Responses to the shift of both grating orientation and window orientation are seen in the average OKN slow phase velocity. Grating rotations produce a rapid OKN rotation to the grating direction (100 ms latency, 300 ms time constant), followed by a slower rebound to the steady state perceived direction midway between the grating and window directions. Window rotations, on the other hand, evoke a slower response (200 ms latency, 500 ms time constant). Conclusions: The results demonstrate multiple cortical inputs to eye movement control: a fast, early input driven by orientation, and a slower input from higher areas sensitive to global stimulus properties.

Mulligan, Jeffrey B.↗

Eye Movement Latencies to Direction Change for Different Classes of Motion

In the analysis of visual motion, local features such as orientation are analyzed early in the cortical processing stream (V1), while integration across orientation and space is thought to occur in higher cortical areas such as MT, MST, etc. If all areas provide inputs to eye movement control centers, we would expect that local properties would drive eye movements with relatively short latencies, while global properties would require longer latencies. When such latencies are observed, they can provide information about when (and where?) various stimulus properties are analyzed. To this end, a stimulus was employed in which local and global properties determining perceived direction-of-motion could be manipulated independently: an elliptical Gabor patch with a drifting carrier, with variable orientation of the carrier grating and the contrast window. We have previously demonstrated that the directional percepts evoked by this stimulus vary between the "grating direction" (the normal to the grating's orientation) and the "window direction" (ARVO 91, 94), and that similar effects can be observed in reflexive eye movements (ARVO 95). Subjects viewed such a stimulus while attempting to maintain steady fixation on the center of the pattern, and the small reflexive eye movements ("stare OKN") were recorded. In the middle of the trial, the orientation of either the grating or the window was rotated smoothly by 30 degrees. Responses to the shift of both grating orientation and window orientation are seen in the average OKN slow phase velocity. Grating rotations produce a rapid OKN rotation to the grating direction (100 ms latency, 300 ms time constant), followed by a slower rebound to the steady state perceived direction midway between the grating and window directions. Window rotations, on the other hand, evoke a slower response (200 ms latency, 500 ms time constant). The results demonstrate multiple cortical inputs to eye movement control: a taste early input driven by orientation, and a slower input from higher areas sensitive to global stimulus properties.

Mulligan, Jeffrey B.↗

Pursuit Eye Movements

When viewing objects, primates use a combination of saccadic and pursuit eye movements to stabilize the retinal image of the object of regard within the high-acuity region near the fovea. Although these movements involve widespread regions of the nervous system, they mix seamlessly in normal behavior. Saccades are discrete movements that quickly direct the eyes toward a visual target, thereby translating the image of the target from an eccentric retinal location to the fovea. In contrast, pursuit is a continuous movement that slowly rotates the eyes to compensate for the motion of the visual target, minimizing the blur that can compromise visual acuity. While other mammalian species can generate smooth optokinetic eye movements - which track the motion of the entire visual surround - only primates can smoothly pursue a single small element within a complex visual scene, regardless of the motion elsewhere on the retina. This ability likely reflects the greater ability of primates to segment the visual scene, to identify individual visual objects, and to select a target of interest.

Krauzlis, Rich↗

Premotor neurons encode torsional eye velocity during smooth-pursuit eye movements

Responses to horizontal and vertical ocular pursuit and head and body rotation in multiple planes were recorded in eye movement-sensitive neurons in the rostral vestibular nuclei (VN) of two rhesus monkeys. When tested during pursuit through primary eye position, the majority of the cells preferred either horizontal or vertical target motion. During pursuit of targets that moved horizontally at different vertical eccentricities or vertically at different horizontal eccentricities, eye angular velocity has been shown to include a torsional component the amplitude of which is proportional to half the gaze angle ("half-angle rule" of Listing's law). Approximately half of the neurons, the majority of which were characterized as "vertical" during pursuit through primary position, exhibited significant changes in their response gain and/or phase as a function of gaze eccentricity during pursuit, as if they were also sensitive to torsional eye velocity. Multiple linear regression analysis revealed a significant contribution of torsional eye movement sensitivity to the responsiveness of the cells. These findings suggest that many VN neurons encode three-dimensional angular velocity, rather than the two-dimensional derivative of eye position, during smooth-pursuit eye movements. Although no clear clustering of pursuit preferred-direction vectors along the semicircular canal axes was observed, the sensitivity of VN neurons to torsional eye movements might reflect a preservation of similar premotor coding of visual and vestibular-driven slow eye movements for both lateral-eyed and foveate species.

Non-NASA Center↗

Inhibition of polar calcium movement and gravitropism in roots treated with auxin-transport inhibitors

Primary roots of maize (Zea mays L.) and pea (Pisum sativum L.) exhibit strong positive gravitropism. In both species, gravistimulation induces polar movement of calcium across the root tip from the upper side to the lower side. Roots of onion (Allium cepa L.) are not responsive to gravity and gravistimulation induces little or no polar movement of calcium across the root tip. Treatment of maize or pea roots with inhibitors of auxin transport (morphactin, naphthylphthalamic acid, 2,3,5-triiodobenzoic acid) prevents both gravitropism and gravity-induced polar movement of calcium across the root tip. The results indicate that calcium movement and auxin movement are closely linked in roots and that gravity-induced redistribution of calcium across the root cap may play an important role in the development of gravitropic curvature.

Indoleacetic Acids/antagonists & inhibitors/metabo↗

Three-dimensional organization of vestibular-related eye movements to off-vertical axis rotation and linear translation in pigeons

During linear accelerations, compensatory reflexes should continually occur in order to maintain objects of visual interest as stable images on the retina. In the present study, the three-dimensional organization of the vestibulo-ocular reflex in pigeons was quantitatively examined during linear accelerations produced by constant velocity off-vertical axis yaw rotations and translational motion in darkness. With off-vertical axis rotations, sinusoidally modulated eye-position and velocity responses were observed in all three components, with the vertical and torsional eye movements predominating the response. Peak torsional and vertical eye positions occurred when the head was oriented with the lateral visual axis of the right eye directed orthogonal to or aligned with the gravity vector, respectively. No steady-state horizontal nystagmus was obtained with any of the rotational velocities (8-58 degrees /s) tested. During translational motion, delivered along or perpendicular to the lateral visual axis, vertical and torsional eye movements were elicited. No significant horizontal eye movements were observed during lateral translation at frequencies up to 3 Hz. These responses suggest that, in pigeons, all linear accelerations generate eye movements that are compensatory to the direction of actual or perceived tilt of the head relative to gravity. In contrast, no translational horizontal eye movements, which are known to be compensatory to lateral translational motion in primates, were observed under the present experimental conditions.

NASA Discipline Neuroscience↗

Properties of cerebellar fastigial neurons during translation, rotation, and eye movements

The most medial of the deep cerebellar nuclei, the fastigial nucleus (FN), receives sensory vestibular information and direct inhibition from the cerebellar vermis. We investigated the signal processing in the primate FN by recording single-unit activities during translational motion, rotational motion, and eye movements. Firing rate modulation during horizontal plane translation in the absence of eye movements was observed in all non-eye-movement-sensitive cells and 26% of the pursuit eye-movement-sensitive neurons in the caudal FN. Many non-eye-movement-sensitive cells recorded in the rostral FN of three fascicularis monkeys exhibited convergence of signals from both the otolith organs and the semicircular canals. At low frequencies of translation, the majority of these rostral FN cells changed their firing rates in phase with head velocity rather than linear acceleration. As frequency increased, FN vestibular neurons exhibited a wide range of response dynamics with most cells being characterized by increasing phase leads as a function of frequency. Unlike cells in the vestibular nuclei, none of the rostral FN cells responded to rotational motion alone, without simultaneously exhibiting sensitivity to translational motion. Modulation during earth-horizontal axis rotation was observed in more than half (77%) of the neurons, although with smaller gains than during translation. In contrast, only 47% of the cells changed their firing rates during earth-vertical axis rotations in the absence of a dynamic linear acceleration stimulus. These response properties suggest that the rostral FN represents a main processing center of otolith-driven information for inertial motion detection and spatial orientation.

Non-NASA Center↗

Vestibular and Non-vestibular Contributions to Eye Movements that Compensate for Head Rotations during Viewing of Near Targets

We studied horizontal eye movements induced by en-bloc yaw rotation, over a frequency range 0.2 - 2.8 Hz, in 10 normal human subjects as they monocularly viewed a target located at their near point of focus. We measured gain and phase relationships between eye-in-head velocity and head velocity when the near target was either earth-fixed or head-fixed. During viewing of the earth-fixed near target, median gain was 1.49 (range 1.24 - 1.87) at 0.2 Hz for the group of subjects, but declined at higher frequencies, so that at 2.8 Hz median gain was 1.08 (range 0.68 - 1.67). During viewing of the head-fixed near target , median gain was 0.03 (range 0.01 - 0.10) at 0.2 Hz for the group of subjects, but increased at higher frequencies, so that at 2.8 Hz median gain was 0.71 (range 0.28 - 0.94). We estimated the vestibular contribution to these responses vestibulo-ocular reflex gain (Gvor) by applying transient head perturbations (peak acceleration> 1,000 deg/s(exp 2)) during sinusoidal rotation under the two viewing conditions. Median Gvor, estimated < 70ms after the onset of head perturbation, was 0.98 (range 0.39 - 1.42) while viewing the earth-fixed near target, and 0.97 (range 0.37 - 1.33) while viewing the head-fixed near target. For the group of subjects, 9 out of 10 subjects showed no significant difference of Gvor between the two viewing conditions ( p > 0.053 ) at all test frequencies. Since Gvor accounted for only -73% of the overall response gain during viewing of the earth-fixed target, we investigated the relative contributions of non-vestibular factors. When subjects viewed the earth-fixed target under strobe illumination, to eliminate retinal image slip information, the gain of compensatory eye movements declined compared with viewing in ambient room light. During sum-of-sine head rotations, while viewing the earth-fixed target, to Han et al./VOR during near-viewing minimize contributions from predictive mechanisms, gain also declined Nonetheless, simple superposition of smooth-pursuit tracking of sinusoidal target motion could not fully account for the overall response at higher frequencies, suggesting other nonvestibular contributions. During binocular viewing conditions when vergence angle was significantly greater than monocular viewing (p < 0.00l), the gain of compensatory eye movements did not show proportional change; indeed, gain could not be correlated with vergence angle during monocular or binocular viewing. We conclude that several separate factors contribute to generate eye rotations during sinusoidal yaw head rotations while viewing a near target; these include the VOR, visual-tracking eye movements that utilize retinal image motion, predictive eye movements and, possibly, other unidentified non-vestibular factors. For these experiments, vergence was not an important determinant of response gam.

Han, Yanning H.↗

Vestibular and Non-vestibular Contributions to Eye Movements that Compensate for Head Rotations during Viewing of Near Targets

We studied horizontal eye movements induced by en-bloc yaw rotation, over a frequency range 0.2 - 2.8 Hz, in 10 normal human subjects as th ey monocularly viewed a target located at their near point of focus. We measured gain and phase relationships between eye-in-head velocity and head velocity when the near target was either earth-fixed or head-fixed. During viewing of the earth-fixed near target,median gain was 1.49 (range 1.24 - 1.87) at 0.2 Hz for the group of subjects, but decl ined at higher frequencies, so that at 2.8 Hz median gain was 1.08 (r ange 0.68 - 1.67). During viewing of the head-fixed near target, median gain was 0.03 (range 0.01 - 0.10) at 0.2 Hz for the group of subjec ts, but increased at higher frequencies, so that at 2.8 Hz median gai n was 0.71 (range 0.28 - 0.94). We estimated the vestibular contribution to these responses (vestibulo-ocular reflex gain, Gvor) by applyin g transient head perturbations (peak acceleration> 1,000 deg's(exp 2) ) during sinusoidal rotation under the two viewing conditions. Median Gvor, estimated < 70m after the onset of head perturbation, was 0.98 (range 0.39 - 1.42) while viewing the earth-fixed near target, and 0. 97 (range 0.37 - 1.33) while viewing the head-fixed near target. For the group of subjects, 9 out of 10 subjects showed no sigificant diff erence of Gvor between the two viewing conditions ( p > 0.053 ) at all test frequencies. Since Gvor accounted for only approximately 73% of the overall response gain during viewing of the earth-fixed target, we investigated the relative contributions of non-vestibular factors. When subjects viewed the earth-fixed target under strobe illumination , to eliminate retinal image slip information, the gain of compensato ry eye movements declined compared with viewing in ambient room light . During sum-of-sine head rotations, while viewing the earth-fixed target, to minimize contributions from predictive mechanisms, gain also declined Nonetheless, simple superposition of smooth-pursuit tracking of sinusoidal target motion could not fully account for the overall r esponse at higher frequencies, suggesting other non-vestibular contributions. During binocular viewing conditions when vergence angle was s ignificantly greater than monocular viewing (p < 0.001), this gain of compensatory eye movements did not show proportional change; indeed, gain could not be correlated with vergence angle during monocular or binocular viewing. We conclude that several separate factors contribute to generate eye rotations during sinusoidal yaw head rotations whi le viewing a near target; these include the VOR, visual-tracking eye movements that utilize retinal image motion, predictive eye movements and, possibly, other unidentified nonvestibular factors. For these experiments, vergence was not an important determinant of response gain .

Han, Yanning H.↗

Vestibulo-Cervico-Ocular Responses and Tracking Eye Movements after Prolonged Exposure to Microgravity

The vestibular function and tracking eye movements were investigated in 12 Russian crew members of ISS missions on days 1(2), 4(5-6), and 8(9-10) after prolonged exposure to microgravity (126 to 195 days). The spontaneous oculomotor activity, static torsional otolith-cervico-ocular reflex, dynamic vestibulo-cervico-ocular responses, vestibular reactivity, tracking eye movements, and gaze-holding were studied using videooculography (VOG) and electrooculography (EOG) for parallel eye movement recording. On post-flight days 1-2 (R+1-2) some cosmonauts demonstrated: - an increased spontaneous oculomotor activity (floating eye movements, spontaneous nystagmus of the typical and atypical form, square wave jerks, gaze nystagmus) with the head held in the vertical position; - suppressed otolith function (absent or reduced by one half amplitude of torsional compensatory eye counter-rolling) with the head inclined statically right- or leftward by 300; - increased vestibular reactivity (lowered threshold and increased intensity of the vestibular nystagmus) during head turns around the longitudinal body axis at 0.125 Hz; - a significant change in the accuracy, velocity, and temporal characteristics of the eye tracking. The pattern, depth, dynamics, and velocity of the vestibular function and tracking eye movements recovery varied with individual participants in the investigation. However, there were also regular responses during readaptation to the normal gravity: - suppression of the otolith function was typically accompanied by an exaggerated vestibular reactivity; - the structure of visual tracking (the accuracy of fixational eye rotations, smooth tracking, and gaze-holding) was disturbed (the appearance of correcting saccades, the transition of smooth tracking to saccadic tracking) only in those cosmonauts who, in parallel to an increased reactivity of the vestibular input, also had central changes in the oculomotor system (spontaneous nystagmus, gaze nystagmus).

Kornilova, L. N.↗

Adjustment of saccade characteristics during head movements.

Saccade characteristics have been studied during coordinated eye-head movements in monkeys. Amplitude, duration, and peak velocity of saccades with head turning were compared with saccades executed while the head was artificially restrained. The results indicate that the saccade characteristics are modulated as a function of head movement, hence the gaze movement (eye+head) exactly matches saccades with head fixed. Saccade modulation is achieved by way of negative vestibulo-ocular feedback. The neck proprioceptors, because of their longer latency, are effective only if the head starts moving prior to the onset of saccade. It is concluded that saccades make with head turning are not 'ballistic' movements because their trajectory is not entirely predetermined by a central command.

Morasso, P.↗

Eye movements during afterimage tracking under sinusoidal and random vestibular stimulation

The smooth portion of the horizontal vestibulo-ocular reflex was analyzed in terms of the frequency response, relating slow-phase eye velocity to angular velocity of a rotating chair under four different cases: sinusoidal rotation about a vertical axis in total darkness, sinusoidal rotation during afterimage tracking, pseudorandom head rotation in total darkness, and pseudorandom head rotation during afterimage tracking. Eye movements were recorded using a photoelectric limbus tracking method. The observation that the presence of an afterimage during vestibular stimulation increases the velocity of slow-phase eye movements is in support of the theory that such slow-phase movements are generated, at least in part, by the perceived velocity of the target. Since the target is immobilized on the retina, this perceived velocity is clearly not generated by retinal slip, but rather by a mechanism related to the eye movement such as corollary discharge.

Yasui, S.↗