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Human eye tracking of a foveal afterimage during angular head oscillation in the dark produced smooth eye movements exceeding those for normal vestibular nystagmus, and a reduction in the frequency of the fast phase component of nystagmus eye movements. These results may support a closed loop extension of the corollary discharge theory, with oculomotor commands based on perceived object velocity.
Pilot eye movement data by recording eye fixations on flight instruments
Accurate motor control requires adaptive processes that correct for gradual and rapid perturbations in the properties of the controlled object. The ability to quickly switch between different movement synergies using sensory cues, referred to as context-dependent adaptation, is a subject of considerable interest at present. The potential function of the cerebellum in context-dependent adaptation remains uncertain, but the data reviewed below suggest that it may play a fundamental role in this process.
In the present paper, we explore effects on the human of exposure to a visual virtual environment which has been enslaved to simulate the human user's head movements or eye movements. Specifically, we have studied the capacity of our experimental subjects to maintain stable spatial orientation in the context of moving their entire visible surroundings by using the parameters of the subjects' natural movements. Our index of the subjects' spatial orientation was the extent of involuntary sways of the body while attempting to stand still, as measured by translations and rotations of the head. We also observed, informally, their symptoms of motion sickness.
Visual suppression during passive eye movement compared to active eye movement to determine origin as retinal or central
To determine the extent to which the pursuit eye-movement and perceptual systems share a common neural motion-processing pathway, we simultaneously measure the psychophysical and eye-movement responses to drifting plaids in a direction-discrimination task. Correlation between the noise in the eye movements and percepts would suggest a shared neural substrate. Three observers were asked to track a plaid (Type I 90 deg; TF = 4 Hz, SF = 0.6 c/d; direction = -2, 0, or 2 deg), and to respond whether the motion was to the right or left of vertical. The eye-movement direction is computed from the initial 300 ms of saccade-free tracking (near open-loop). On each trial, we use the eye-movement direction to predict the perceptual response. We then compute the probability of successfully predicting the observer's response. We also use SOC analysis as an alternate measure of the correlation. We compare the data from both these analyses to the predictions of two models in which the eye movements and the percept are driven by either a common noisy signal (correlated model) or by two separate noisy signals (uncorrelated model). We also allow for the fact that our measured eye movements are degraded by eye-tracker noise. This noise causes our measured correlation to be lower than the actual biological correlation. The correlations in our data are higher than those predicted by the uncorrelated model, and are similar to the predictions of the correlated model. The proportion of perceptual responses correctly predicted by the eye movements were 0.66, 0.60, 0.73 for the -2, 0, 2 deg stimuli respectively (correlated model 0.65, 0.60, 0.70; uncorrelated model 0.58, 0.51, 0.60).The SOC proportions were 0.71, 0.64, 0.72 (correlated model 0.66, 0.64, 0.69; uncorrelated model 0.5, 0.5, 0.5). These results show that the oculomotor and perceptual system share a performance-limiting noise source and provide strong evidence that a common neural mechanism (perhaps MT or MST) drives both perception and smooth eye movements.
The firing behavior of 51 non-eye movement related central vestibular neurons that were sensitive to passive head rotation in the plane of the horizontal semicircular canal was studied in three squirrel monkeys whose heads were free to move in the horizontal plane. Unit sensitivity to active head movements during spontaneous gaze saccades was compared with sensitivity to passive head rotation. Most units (29/35 tested) were activated at monosynaptic latencies following electrical stimulation of the ipsilateral vestibular nerve. Nine were vestibulo-spinal units that were antidromically activated following electrical stimulation of the ventromedial funiculi of the spinal cord at C1. All of the units were less sensitive to active head movements than to passive whole body rotation. In the majority of cells (37/51, 73%), including all nine identified vestibulo-spinal units, the vestibular signals related to active head movements were canceled. The remaining units (n = 14, 27%) were sensitive to active head movements, but their responses were attenuated by 20-75%. Most units were nearly as sensitive to passive head-on-trunk rotation as they were to whole body rotation; this suggests that vestibular signals related to active head movements were cancelled primarily by subtraction of a head movement efference copy signal. The sensitivity of most units to passive whole body rotation was unchanged during gaze saccades. A fundamental feature of sensory processing is the ability to distinguish between self-generated and externally induced sensory events. Our observations suggest that the distinction is made at an early stage of processing in the vestibular system.
Rates of waking eye movement to rapid eye movement in sleep conditioned monkeys using electrooculogram
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).
A problem often encountered when eye-movement measurement is conducted is the choice of 'indices' or 'statistics' available to present such information. The present study reports the use of the Time-Locked Time-History as a technique of value in the examination and presentation of eye-movement data. Plots created using this technique are labeled time-locked time-histories as they illustrate subject eye lookpoint during a period of time before and after a certain time-locking event. Events that occur with some degree of repetition, such as the onset or termination of control activities, warning signals, or changes in indicator positions may be utilized as time-locking events. The present study reports the use of this technique in an eye-movement study using a secondary task in which the subject must discriminate specific types of information in the display.
Saccadic eye movements were studied in six subjects during two Space Shuttle missions. Reaction time, peak velocity and accuracy of horizontal, visually-guided saccades were examined preflight, inflight and postflight. Conventional electro-oculography was used to record eye position, with the subjects responding to pseudo-randomly illuminated targets at 0 deg and + or - 10 deg and 20 deg visual angles. In all subjects, preflight measurements were within normal limits. Reaction time was significantly increased inflight, while peak velocity was significantly decreased. A tendency toward a greater proportion of hypometric saccades inflight was also noted. Possible explanations for these changes and possible correlations with space motion sickness are discussed.
Eye movement data and other parameters including instrument readings, aircraft state and position variables, and control maneuvers were recorded while pilots flew ILS simulations in a B 737. The experiment itself employed seven airline pilots, each of whom flew approximately 40 approach/landing sequences. The simulator was equipped with a night visual scene but the scene was fogged out down to approximately 60 meters (200 ft). The instrument scanning appeared to follow aircraft parameters not physical position of instruments. One important implication of the results is: pilots look for categories or packets of information. Control inputs were tabulated according to throttle, wheel position, column, and pitch trim changes. Three seconds of eye movements before and after the control input were then obtained. Analysis of the eye movement data for the controlling periods showed clear patterns. The results suggest a set of miniscan patterns which are used according to the specific details of the situation. A model is developed which integrates scanning and controlling. Differentiations are made between monitoring and controlling scans.
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.
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 .
Eye movements made by various subjects in the performance of a precognitive tracking task are studied. The tracking task persented by an antiaircraft artillery (AAA) simulator has an input forcing function represented by a deterministic aircraft fly-by. The performance of subjects is ranked by two metrics. Good, mediocre, and poor trackers are selected for analysis based on performance during the difficult segment of the tracking task and over replications. Using phase planes to characterize both the eye movement patterns and the displayed error signal, a simple metric is developed to study these patterns. Two characterizations of eye movement strategies are defined and quantified. Using these two types of eye strategies, two conclusions are obtained about good, mediocre, and poor trackers. First, the eye tracker who used a fixed strategy will consistently perform better. Secondly, the best fixed strategy is defined as a Crosshair Fixator.
Warren and Hannon (1988, 1990), while studying the perception of heading during eye movements, concluded that people do not require extraretinal information to judge heading with eye/head movements present. Here, heading judgments are examined at higher, more typical eye movement velocities than the extremely slow tracking eye movements used by Warren and Hannon. It is found that people require extraretinal information about eye position to perceive heading accurately under many viewing conditions.
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.