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At least 19 records
All sourcesNASA NTRSDOE OSTI
NASA NTRS2003

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↗
NASA NTRS1995

Geometric adjustments to account for eye eccentricity in processing horizontal and vertical eye and head movement data

Neglecting the eccentric position of the eyes in the head can lead to erroneous interpretation of ocular motor data, particularly for near targets. We discuss the geometric effects that eye eccentricity has on the processing of target-directed eye and head movement data, and we highlight two approaches to processing and interpreting such data. The first approach involves determining the true position of the target with respect to the location of the eyes in space for evaluating the efficacy of gaze, and it allows calculation of retinal error directly from measured eye, head, and target data. The second approach effectively eliminates eye eccentricity effects by adjusting measured eye movement data to yield equivalent responses relative to a specified reference location (such as the center of head rotation). This latter technique can be used to standardize measured eye movement signals, enabling waveforms collected under different experimental conditions to be directly compared, both with the measured target signals and with each other. Mathematical relationships describing these approaches are presented for horizontal and vertical rotations, for both tangential and circumferential display screens, and efforts are made to describe the sensitivity of parameter variations on the calculated results.

NASA Discipline Neuroscience↗
NASA NTRS1996

Measuring eye movements during locomotion: filtering techniques for obtaining velocity signals from a video-based eye monitor

Video-based eye-tracking systems are especially suited to studying eye movements during naturally occurring activities such as locomotion, but eye velocity records suffer from broad band noise that is not amenable to conventional filtering methods. We evaluated the effectiveness of combined median and moving-average filters by comparing prefiltered and postfiltered records made synchronously with a video eye-tracker and the magnetic search coil technique, which is relatively noise free. Root-mean-square noise was reduced by half, without distorting the eye velocity signal. To illustrate the practical use of this technique, we studied normal subjects and patients with deficient labyrinthine function and compared their ability to hold gaze on a visual target that moved with their heads (cancellation of the vestibulo-ocular reflex). Patients and normal subjects performed similarly during active head rotation but, during locomotion, patients held their eyes more steadily on the visual target than did subjects.

NASA Discipline Neuroscience↗
NASA NTRS1993

An Investigation of Horizontal Combined Eye-Head Tracking in Patients with Abnormal Vestibular and Smooth Pursuit Eye Movements

We investigated the interaction of smooth ocular pursuit (SP) and the vestibulo-ocular reflex (VOR) during horizontal, combined eye-head tracking (CEHT) in patients with abnormalities of either the VOR or SP movements. Our strategy was to apply transient stimuli that capitalized on the different latencies to onset of SP and the VOR. During CEHT of a target moving at 15 deg/sec, normal subjects and patients with VOR deficits all tracked the target with a gain close to 1.O. When the heads of normal subjects were suddenly and unexpectedly braked to a halt during CEHT, the eye promptly began to move in the orbit to track the target, but eye-in-orbit velocity transiently fell to about 60-70% of target velocity. In patients with deficient labyrinthine function, following the onset of the head brake, eye movements to track the target were absent, and SP movements were not generated until about 100 msec later. In patients with deficient SP, CEHT was superior to SP tracking with the head stationary; after the onset of the head brake, tracking eye movements were initiated promptly, but eye velocity was less than 50% of target velocity and increased only slightly thereafter. These results indicate that at least two mechanisms operate to overcome the VOR and allow gaze to track the target during CEHT: (1) the SP system provides a signal to cancel a normally-operating VOR (this cancellation signal is not needed by labyrinthine-deficient patients who have no VOR to cancel), and (2) a reduction of the gain of the VOR is achieved, an ability that is preserved even in patients with cerebral lesions that impair SP.

Huebner, William P.↗
NASA NTRSDate not supplied

How Eye Drift Adapts to the History of Eye Positions

Oculomotor behavior is a powerful tool to probe brain function and disease. Previous studies have shown how fixational drift velocities depend on eccentricity (Bertolini et al., 2013; Romano et al., 2017) and how fixational drift can adapt after prolonged eccentric fixation (Otero-Millan et al., 2019). Here, eighteen subjects completed two experiments where we measured drift velocity while fixating a briefly flashed target, to remove retinal slip cues. In the first experiment, we measured their baseline relationship between fixational drift velocity and eye position. In the second experiment, we investigated the adaptation of that relationship after prolonged fixation at a very eccentric position. This adaptation usually manifests by the presence of rebound nystagmus (RN) upon returning to central gaze. Our results replicate the finding that drift increases at central fixation in the direction consistent with rebound nystagmus, that is, towards the previously held eccentric position, highlighting that drift velocity depends on the history of eye positions. When examining the entire relationship between drift velocity and eye position we find a mostly linear relationship but with increased slope relative to the baseline condition. However, the adaptation of the integrator cannot be modeled by a simple change in leakiness, as that would not result in increased drift at central gaze. Other mechanisms of adaptation such as shifts in the position or velocity set-points for gaze holding could be present but are indistinguishable with the current data.

eye movement↗
NASA NTRS1989

Eyes open versus eyes closed - Effect on human rotational responses

The effect of eyelid closure on the response to rotational vestibular stimulation was assessed by evaluating 16 normal human subjects with both earth vertical axis (EVA) and earth horizontal axis (EHA) yaw rotations with either eyes closed (EC) or eyes open in the dark (EOD). Results indicated that for EVA rotation, the subjects' responses were of larger magnitude and less variable with EOD than with EC. However, for EHA rotation, responses were of larger magnitude and equally variable with EC as compared to EOD. Data also indicated that the quality of the EHA response with EC was altered because eyelid closure influenced the amount of periodic gaze. It is concluded that eyelid closure has an effect upon both canalocular and otolithocular reflexes and it is suggested that both EVA and EHA rotational testing be performed with EOD rather than with EC.

Wall, Conrad, III↗
NASA NTRS1999

Effect of viewing distance on the generation of vertical eye movements during locomotion

Vertical head and eye coordination was studied as a function of viewing distance during locomotion. Vertical head translation and pitch movements were measured using a video motion analysis system (Optotrak 3020). Vertical eye movements were recorded using a video-based pupil tracker (Iscan). Subjects (five) walked on a linear treadmill at a speed of 1.67 m/s (6 km/h) while viewing a target screen placed at distances ranging from 0.25 to 2.0 m at 0. 25-m intervals. The predominant frequency of vertical head movement was 2 Hz. In accordance with previous studies, there was a small head pitch rotation, which was compensatory for vertical head translation. The magnitude of the vertical head movements and the phase relationship between head translation and pitch were little affected by viewing distance, and tended to orient the naso-occipital axis of the head at a point approximately 1 m in front of the subject (the head fixation distance or HFD). In contrast, eye velocity was significantly affected by viewing distance. When viewing a far (2-m) target, vertical eye velocity was 180 degrees out of phase with head pitch velocity, with a gain of 0. 8. This indicated that the angular vestibulo-ocular reflex (aVOR) was generating the eye movement response. The major finding was that, at a close viewing distance (0.25 m), eye velocity was in phase with head pitch and compensatory for vertical head translation, suggesting that activation of the linear vestibulo-ocular reflex (lVOR) was contributing to the eye movement response. There was also a threefold increase in the magnitude of eye velocity when viewing near targets, which was consistent with the goal of maintaining gaze on target. The required vertical lVOR sensitivity to cancel an unmodified aVOR response and generate the observed eye velocity magnitude for near targets was almost 3 times that previously measured. Supplementary experiments were performed utilizing body-fixed active head pitch rotations at 1 and 2 Hz while viewing a head-fixed target. Results indicated that the interaction of smooth pursuit and the aVOR during visual suppression could modify both the gain and phase characteristics of the aVOR at frequencies encountered during locomotion. When walking, targets located closer than the HFD (1.0 m) would appear to move in the same direction as the head pitch, resulting in suppression of the aVOR. The results of the head-fixed target experiment suggest that phase modification of the aVOR during visual suppression could play a role in generating eye movements consistent with the goal of maintaining gaze on targets closer than the HFD, which would augment the lVOR response.

NASA Discipline Neuroscience↗
NASA NTRS1996

Quantifying the Correlation Between Eye-Movement and Perceptual Responses to Moving Plaids

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.

Beutter, Brent R.↗
NASA NTRS1997

Kinematic principles of primate rotational vestibulo-ocular reflex. II. Gravity-dependent modulation of primary eye position

The kinematic constraints of three-dimensional eye positions were investigated in rhesus monkeys during passive head and body rotations relative to gravity. We studied fast and slow phase components of the vestibulo-ocular reflex (VOR) elicited by constant-velocity yaw rotations and sinusoidal oscillations about an earth-horizontal axis. We found that the spatial orientation of both fast and slow phase eye positions could be described locally by a planar surface with torsional variation of <2.0 +/- 0.4 degrees (displacement planes) that systematically rotated and/or shifted relative to Listing's plane. In supine/prone positions, displacement planes pitched forward/backward; in left/right ear-down positions, displacement planes were parallel shifted along the positive/negative torsional axis. Dynamically changing primary eye positions were computed from displacement planes. Torsional and vertical components of primary eye position modulated as a sinusoidal function of head orientation in space. The torsional component was maximal in ear-down positions and approximately zero in supine/prone orientations. The opposite was observed for the vertical component. Modulation of the horizontal component of primary eye position exhibited a more complex dependence. In contrast to the torsional component, which was relatively independent of rotational speed, modulation of the vertical and horizontal components of primary position depended strongly on the speed of head rotation (i.e., on the frequency of oscillation of the gravity vector component): the faster the head rotated relative to gravity, the larger was the modulation. Corresponding results were obtained when a model based on a sinusoidal dependence of instantaneous displacement planes (and primary eye position) on head orientation relative to gravity was fitted to VOR fast phase positions. When VOR fast phase positions were expressed relative to primary eye position estimated from the model fits, they were confined approximately to a single plane with a small torsional standard deviation ( approximately 1.4-2.6 degrees). This reduced torsional variation was in contrast to the large torsional spread (well >10-15 degrees ) of fast phase positions when expressed relative to Listing's plane. We conclude that primary eye position depends dynamically on head orientation relative to space rather than being fixed to the head. It defines a gravity-dependent coordinate system relative to which the torsional variability of eye positions is minimized even when the head is moved passively and vestibulo-ocular reflexes are evoked. In this general sense, Listing's law is preserved with respect to an otolith-controlled reference system that is defined dynamically by gravity.

NASA Discipline Neuroscience↗
NASA NTRS1992

Stabilization of gaze during circular locomotion in darkness. II. Contribution of velocity storage to compensatory eye and head nystagmus in the running monkey

1. Yaw eye in head (Eh) and head on body velocities (Hb) were measured in two monkeys that ran around the perimeter of a circular platform in darkness. The platform was stationary or could be counterrotated to reduce body velocity in space (Bs) while increasing gait velocity on the platform (Bp). The animals were also rotated while seated in a primate chair at eccentric locations to provide linear and angular accelerations similar to those experienced while running. 2. Both animals had head and eye nystagmus while running in darkness during which slow phase gaze velocity on the body (Gb) partially compensated for body velocity in space (Bs). The eyes, driven by the vestibuloocular reflex (VOR), supplied high-frequency characteristics, bringing Gb up to compensatory levels at the beginning and end of the slow phases. The head provided substantial gaze compensation during the slow phases, probably through the vestibulocollic reflex (VCR). Synchronous eye and head quick phases moved gaze in the direction of running. Head movements occurred consistently only when animals were running. This indicates that active body and limb motion may be essential for inducing the head-eye gaze synergy. 3. Gaze compensation was good when running in both directions in one animal and in one direction in the other animal. The animals had long VOR time constants in these directions. The VOR time constant was short to one side in one animal, and it had poor gaze compensation in this direction. Postlocomotory nystagmus was weaker after running in directions with a long VOR time constant than when the animals were passively rotated in darkness. We infer that velocity storage in the vestibular system had been activated to produce continuous Eh and Hb during running and to counteract postrotatory afterresponses. 4. Continuous compensatory gaze nystagmus was not produced by passive eccentric rotation with the head stabilized or free. This indicates that an aspect of active locomotion, most likely somatosensory feedback, was responsible for activating velocity storage. 5. Nystagmus was compared when an animal ran in darkness and in light. the beat frequency of eye and head nystagmus was lower, and the quick phases were larger in darkness. The duration of head and eye quick phases covaried. Eye quick phases were larger when animals ran in darkness than when they were passively rotated. The maximum velocity and duration of eye quick phases were the same in both conditions. 6. The platform was counterrotated under one monkey in darkness while it ran in the direction of its long vestibular time constant.(ABSTRACT TRUNCATED AT 400 WORDS).

NASA Discipline Neuroscience↗
NASA NTRS1992

Stabilization of gaze during circular locomotion in light. I. Compensatory head and eye nystagmus in the running monkey

1. A rhesus and cynomolgus monkey were trained to run around the perimeter of a circular platform in light. We call this "circular locomotion" because forward motion had an angular component. Head and body velocity in space were recorded with angular rate sensors and eye movements with electrooculography (EOG). From these measurements we derived signals related to the angular velocity of the eyes in the head (Eh), of the head on the body (Hb), of gaze on the body (Gb), of the body in space (Bs), of gaze in space (Gs), and of the gain of gaze (Gb/Bs). 2. The monkeys had continuous compensatory nystagmus of the head and eyes while running, which stabilized Gs during the slow phases. The eyes established and maintained compensatory gaze velocities at the beginning and end of the slow phases. The head contributed to gaze velocity during the middle of the slow phases. Slow phase Gb was as high as 250 degrees/s, and targets were fixed for gaze angles as large as 90-140 degrees. 3. Properties of the visual surround affected both the gain and strategy of gaze compensation in the one monkey tested. Gains of Eh ranged from 0.3 to 1.1 during compensatory gaze nystagmus. Gains of Hb varied around 0.3 (0.2-0.7), building to a maximum as Eh dropped while running past sectors of interest. Consistent with predictions, gaze gains varied from below to above unity, when translational and angular body movements with regard to the target were in opposite or the same directions, respectively. 4. Gaze moved in saccadic shifts in the direction of running during quick phases. Most head quick phases were small, and at times the head only paused during an eye quick phase. Eye quick phases were larger, ranging up to 60 degrees. This is larger than quick phases during passive rotation or saccades made with the head fixed. 5. These data indicate that head and eye nystagmus are natural phenomena that support gaze compensation during locomotion. Despite differential utilization of the head and eyes in various conditions, Gb compensated for Bs. There are various frames of reference in which an estimate of angular velocity that drives the head and eyes could be based. We infer that body in space velocity (Bs) is likely to be represented centrally to provide this signal.

Non-NASA Center↗
NASA NTRS1992

Comparison of Predictable Smooth Ocular and Combined Eye-Head Tracking Behaviour in Patients with Lesions Affecting the Brainstem and Cerebellum

We compared the ability of eight normal subjects and 15 patients with brainstem or cerebellar disease to follow a moving visual stimulus smoothly with either the eyes alone or with combined eye-head tracking. The visual stimulus was either a laser spot (horizontal and vertical planes) or a large rotating disc (torsional plane), which moved at one sinusoidal frequency for each subject. The visually enhanced Vestibulo-Ocular Reflex (VOR) was also measured in each plane. In the horizontal and vertical planes, we found that if tracking gain (gaze velocity/target velocity) for smooth pursuit was close to 1, the gain of combined eye-hand tracking was similar. If the tracking gain during smooth pursuit was less than about 0.7, combined eye-head tracking was usually superior. Most patients, irrespective of diagnosis, showed combined eye-head tracking that was superior to smooth pursuit; only two patients showed the converse. In the torsional plane, in which optokinetic responses were weak, combined eye-head tracking was much superior, and this was the case in both subjects and patients. We found that a linear model, in which an internal ocular tracking signal cancelled the VOR, could account for our findings in most normal subjects in the horizontal and vertical planes, but not in the torsional plane. The model failed to account for tracking behaviour in most patients in any plane, and suggested that the brain may use additional mechanisms to reduce the internal gain of the VOR during combined eye-head tracking. Our results confirm that certain patients who show impairment of smooth-pursuit eye movements preserve their ability to smoothly track a moving target with combined eye-head tracking.

Grant, Michael P.↗
NASA NTRS1999

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↗
NASA NTRS1989

Optical, gravitational, and kinesthetic determinants of judged eye level

Subjects judged eye level, defined in three distinct ways relative to three distinct reference planes: a gravitational horizontal, giving the gravitationally referenced eye level (GREL); a visible surface, giving the surface-referenced eye level (SREL); and a plane fixed with respect to the head, giving the head-referenced eye level (HREL). The information available for these judgements was varied by having the subjects view an illuminated target that could be placed in a box which: (1) was pitched at various angles, (2) was illuminated or kept in darkness, (3) was moved to different positions along the subject's head-to-foot body axis, and (4) was viewed with the subjects upright or reclining. The results showed: (1) judgements of GREL made in the dark were 2.5 deg lower than in the light, with a significantly greater variability; (2) judged GREL was shifted approximately half of the way toward SREL when these two eye levels did not coincide; (3) judged SREL was shifted about 12 percent of the way toward HREL when these two eye levels did not coincide, (4) judged HREL was shifted about half way toward SREL when these two eye level did not coincide and when the subject was upright (when the subject was reclining, HREL was shifted approx. 90 percent toward SREL); (5) the variability of the judged HREL in the dark was nearly twice as great with the subject reclining than with the subject upright. These results indicate that gravity is an important source of information for judgement of eye level. In the absence of information concerning the direction of gravity, the ability to judge HREL is extremely poor. A visible environment does not seem to afford precise information as to judgements of direction, but it probably does afford significant information as to the stability of these judgements.

Stoper, Arnold E.↗
NASA NTRS1990

NASA supporting studies for microgravity research on eye movements

The purpose of the work on this project was to provide support for ground-based studies on the effects of gravity on eye movements. The effects of microgravity on the optokinetic eye movements of humans are investigated. OKN was induced by having subjects watch 3.3 deg stripes moving at 35 deg/s for 45 s in a binocular, head-fixed apparatus. The field (hor., 88 deg; vert., 72 deg), was rotated about axes that were upright or tilted 45 deg or 90 deg. The head was upright or tilted 45 deg on the body. Head-horizontal (yaw axis) and head-vertical (pitch axis) components of OKN were recorded with electro-oculography (EOG). Slow phase velocity vectors were determined relative to gravity. With the head upright, the axis of eye rotation during yaw axis OKN was coincident with the stimulus axis and the spatial vertical. With the head tilted 45 deg on the body, a persistent vertical component of eye velocity developed during yaw axis stimulation, and there was an average shift of the axis of eye rotation toward the spatial vertical of approximately 18 deg in six subjects. During oblique optokinetic stimulation with the head upright, the axis of eye rotation shifted 12 deg toward the spatial vertical. When the head was tilted, the axis of eye rotation rotated to the other side of the spatial vertical by 5.4 deg during the same oblique stimulation. This counter-rotation of the axis of eye rotation is similar to the 'Muller (E) effect', in which the perception of the upright counter-rotates to the opposite side of the spatial vertical when subjects are tilted in darkness. The data were simulated by a model of OKN. Despite the short OKAN time constants, strong horizontal to vertical cross-coupling was produced if the horizontal and vertical time constants were in proper ratio, and there was no suppression of nystagmus orthogonal to the stimulus direction. This shows that the spatial orientation of OKN can be due to a restructuring of the system matrix of velocity storage as a function of gravity. It is concluded that although human OKAN is weak, velocity storage orients the slow phase velocity of OKN towards the spatial vertical.

Cohen, Bernard↗
NASA NTRS2000

Role of muscle pulleys in producing eye position-dependence in the angular vestibuloocular reflex: a model-based study

It is well established that the head and eye velocity axes do not always align during compensatory vestibular slow phases. It has been shown that the eye velocity axis systematically tilts away from the head velocity axis in a manner that is dependent on eye-in-head position. The mechanisms responsible for producing these axis tilts are unclear. In this model-based study, we aimed to determine whether muscle pulleys could be involved in bringing about these phenomena. The model presented incorporates semicircular canals, central vestibular pathways, and an ocular motor plant with pulleys. The pulleys were modeled so that they brought about a rotation of the torque axes of the extraocular muscles that was a fraction of the angle of eye deviation from primary position. The degree to which the pulleys rotated the torque axes was altered by means of a pulley coefficient. Model input was head velocity and initial eye position data from passive and active yaw head impulses with fixation at 0 degrees, 20 degrees up and 20 degrees down, obtained from a previous experiment. The optimal pulley coefficient required to fit the data was determined by calculating the mean square error between data and model predictions of torsional eye velocity. For active head impulses, the optimal pulley coefficient varied considerably between subjects. The median optimal pulley coefficient was found to be 0.5, the pulley coefficient required for producing saccades that perfectly obey Listing's law when using a two-dimensional saccadic pulse signal. The model predicted the direction of the axis tilts observed in response to passive head impulses from 50 ms after onset. During passive head impulses, the median optimal pulley coefficient was found to be 0.21, when roll gain was fixed at 0.7. The model did not accurately predict the alignment of the eye and head velocity axes that was observed early in the response to passive head impulses. We found that this alignment could be well predicted if the roll gain of the angular vestibuloocular reflex was modified during the initial period of the response, while pulley coefficient was maintained at 0.5. Hence a roll gain modification allows stabilization of the retinal image without requiring a change in the pulley effect. Our results therefore indicate that the eye position-dependent velocity axis tilts could arise due to the effects of the pulleys and that a roll gain modification in the central vestibular structures may be responsible for countering the pulley effect.

NASA Discipline Neuroscience↗
NASA NTRS2006

Modification of Eye Movements and Motion Perception during Off-Vertical Axis Rotation

Constant velocity Off-Vertical Axis Rotation (OVAR) imposes a continuously varying orientation of the head and body relative to gravity. The ensuing ocular reflexes include modulation of both torsional and horizontal eye movements as a function of the varying linear acceleration along the lateral plane, and modulation of vertical and vergence eye movements as a function of the varying linear acceleration along the sagittal plane. Previous studies have demonstrated that tilt and translation otolith-ocular responses, as well as motion perception, vary as a function of stimulus frequency during OVAR. The purpose of this study is to examine normative OVAR responses in healthy human subjects, and examine adaptive changes in astronauts following short duration space flight at low (0.125 Hz) and high (0.5 Hz) frequencies. Data was obtained on 24 normative subjects (14 M, 10 F) and 14 (13 M, 1F) astronaut subjects. To date, astronauts have participated in 3 preflight sessions (n=14) and on R+0/1 (n=7), R+2 (n= 13) and R+4 (n= 13) days after landing. Subjects were rotated in darkness about their longitudinal axis 20 deg off-vertical at constant rates of 45 and 180 deg/s, corresponding to 0.125 and 0.5 Hz. Binocular responses were obtained with video-oculography. Perceived motion was evaluated using verbal reports and a two-axis joystick (pitch and roll tilt) mounted on top of a two-axis linear stage (anterior-posterior and medial-lateral translation). Eye responses were obtained in ten of the normative subjects with the head and trunk aligned, and then with the head turned relative to the trunk 40 deg to the right or left of center. Sinusoidal curve fits were used to derive amplitude, phase and bias of the responses over several cycles at each stimulus frequency. Eye responses during 0.125 Hz OVAR were dominated by modulation of torsional and vertical eye position, compensatory for tilt relative to gravity. While there is a bias horizontal slow phase velocity (SPV), the modulation of horizontal and vergence SPV is negligible at this lower stimulus frequency. Eye responses during 0.5 Hz OVAR; however, are characterized by modulation of horizontal and vergence SPV, compensatory for translation in the lateral and sagittal planes, respectively. Neither amplitude nor bias velocities were significantly altered by head-on-trunk position. The phases of the ocular reflexes, on the other hand, shifted towards alignment with the head. During the lower frequency OVAR, subjects reported the perception of progressing along the edge of a cone. During higher frequency OVAR, subjects reported the perception of progressing along the edge of an upright cylinder. In contrast to the eye movements, the phase of both perceived tilt and translation motion is not altered by stimulus frequency. Preliminary results from astronaut data suggest that the ocular responses are not substantially altered by short-duration spaceflight. However, compared to preflight averages, astronauts reported greater amplitude of both perceived tilt and translation at low and high frequency, respectively, during early post-flight testing. We conclude that the neural processing to distinguish tilt and translation linear acceleration stimuli differs between eye movements and motion perception. The results from modifying head-on-trunk position are consistent with the modulation of ocular reflexes during OVAR being primarily mediated by the otoliths in response to the sinusoidally varying linear acceleration along the interaural and naso-occipital head axis. While the tilt and translation ocular reflexes appear to operate in an independent fashion, the timing of perceived tilt and translation influence each other. We conclude that the perceived motion path during linear acceleration in darkness results from a composite representation of tilt and translation inputs from both vestibular and somatosensory systems.

Wood, S. J.↗
NASA NTRS2007

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.↗