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NASA NTRS2010

Direct Relationship Between Perceptual and Motor Variability

The time that elapses between stimulus onset and the onset of a saccadic eye movement is longer and more variable than can be explained by neural transmission times and synaptic delays (Carpenter, 1981, in: Eye Movements: Cognition & Visual Perception, Earlbaum). In theory, noise underlying response-time (RT) variability could arise at any point along the sensorimotor cascade, from sensory noise arising Vvithin the early visual processing shared Vvith perception to noise in the motor criterion or commands necessary to trigger movements. These two loci for internal noise can be distinguished empirically; sensory internal noise predicts that response time Vvill correlate Vvith perceived stimulus magnitude whereas motor internal noise predicts no such correlation. Methods. We used the data described by Liston and Stone (2008, JNS 28:13866-13875), in which subjects performed a 2AFC saccadic brightness discrimination task and the perceived brightness of the chosen stimulus was then quantified in a second 21FC perceptual task. Results. We binned each subject's data into quartiles for both signal strength (from dimmest to brightest) and RT (from slowest to fastest) and analyzed the trends in perceived brightness. We found significant effects of both signal strength (as expected) and RT on normalized perceived brightness (both p less than 0.0001, 2-way ANOVA), without significant interaction (p = 0.95, 2-way ANOVA). A plot of normalized perceived brightness versus normalized RT show's that more than half of the variance was shared (r2 = 0.56, P less than 0.0001). To rule out any possibility that some signal-strength related artifact was generating this effect, we ran a control analysis on pairs of trials with repeated presentations of identical stimuli and found that stimuli are perceived to be brighter on trials with faster saccades (p less than 0.001, paired t-test across subjects). Conclusion. These data show that shared early visual internal noise jitters perceived brightness and the saccadic motor output in parallel. While the present correlation could theoretically result, either directly or indirectly, from some low-level brainstem or retinal mechanism (e.g., arousal, pupil size, photoreceptor noise) that influences both visual and oculomotor circuits, this is unlikely given the earlier fin ding that the variability in perceived motion direction and smooth-pursuit motor output is highly correlated (Stone and Krauzlis, 2003, JOV 3:725-736), suggesting that cortical circuits contribute to the shared internal noise.

Liston, Dorion B.↗
NASA NTRS2006

Modification of Eccentric Gaze-Holding

Clear vision and accurate localization of objects in the environment are prerequisites for reliable performance of motor tasks. Space flight confronts the crewmember with a stimulus rearrangement that requires adaptation to function effectively with the new requirements of altered spatial orientation and motor coordination. Adaptation and motor learning driven by the effects of cerebellar disorders may share some of the same demands that face our astronauts. One measure of spatial localization shared by the astronauts and those suffering from cerebellar disorders that is easily quantified, and for which a neurobiological substrate has been identified, is the control of the angle of gaze (the "line of sight"). The disturbances of gaze control that have been documented to occur in astronauts and cosmonauts, both in-flight and postflight, can be directly related to changes in the extrinsic gravitational environment and intrinsic proprioceptive mechanisms thus, lending themselves to description by simple non-linear statistical models. Because of the necessity of developing robust normal response populations and normative populations against which abnormal responses can be evaluated, the basic models can be formulated using normal, non-astronaut test subjects and subsequently extended using centrifugation techniques to alter the gravitational and proprioceptive environment of these subjects. Further tests and extensions of the models can be made by studying abnormalities of gaze control in patients with cerebellar disease. A series of investigations were conducted in which a total of 62 subjects were tested to: (1) Define eccentric gaze-holding parameters in a normative population, and (2) explore the effects of linear acceleration on gaze-holding parameters. For these studies gaze-holding was evaluated with the subjects seated upright (the normative values), rolled 45 degrees to both the left and right, or pitched back 30 and 90 degrees. In a separate study the further effects of acceleration on gaze stability was examined during centrifugation (+2 G (sub x) and +2 G (sub z) using a total of 23 subjects. In all of our investigations eccentric gaze-holding was established by having the subjects acquire an eccentric target (+/-30 degrees horizontal, +/- 15 degrees vertical) that was flashed for 750 msec in an otherwise dark room. Subjects were instructed to hold gaze on the remembered position of the flashed target for 20 sec. Immediately following the 20 sec period, subjects were cued to return to the remembered center position and to hold gaze there for an additional 20 sec. Following this 20 sec period the center target was briefly flashed and the subject made any corrective eye movement back to the true center position. Conventionally, the ability to hold eccentric gaze is estimated by fitting the natural log of centripetal eye drifts by linear regression and calculating the time constant (G) of these slow phases of "gaze-evoked nystagmus". However, because our normative subjects sometimes showed essentially no drift (tau (sub c) = m), statistical estimation and inference on the effect of target direction was performed on values of the decay constant theta = 1/(tau (sub c)) which we found was well modeled by a gamma distribution. Subjects showed substantial variance of their eye drifts, which were centrifugal in approximately 20 % of cases, and > 40% for down gaze. Using the ensuing estimated gamma distributions, we were able to conclude that rightward and leftward gaze holding were not significantly different, but that upward gaze holding was significantly worse than downward (p<0.05). We also concluded that vertical gaze holding was significantly worse than horizontal (p<0.05). In the case of left and right roll, we found that both had a similar improvement to horizontal gaze holding (p<0.05), but didn't have a significant effect on vertical gaze holding. For pitch tilts, both tilt angles significantly decreased gaze-holding ility in all directions (p<0.05). Finally, we found that hyper-g centrifugation significantly decreased gaze holding ability in the vertical plane. The main findings of this study are as follows: (1) vertical gaze-holding is less stable than horizontal, (2) gaze-holding to upward targets is less stable than to downward targets, (3) tilt affects gaze holding, and (4) hyper-g affects gaze holding. This difference between horizontal and vertical gaze-holding may be ascribed to separate components of the velocity-to-position neural integrator for eye movements, and to differences in orbital mechanics. The differences between upward and downward gaze-holding may be ascribed to an inherent vertical imbalance in the vestibular system. Because whole body tilt and hyper-g affects gaze-holding, it is implied that the otolith organs have direct connections to the neural integrator and further studies of astronaut gaze-holding are warranted. Our statistical method for representing the range of normal eccentric gaze stability can be readily applied to normals who maybe exposed to environments which may modify the central integrator and require monitoring, and to evaluate patients with gaze-evoked nystagmus by comparing to the above established normative criteria.

Reschke, M. F.↗
NASA NTRS1992

A Reevaluation of the Vestibulo-Ocular Reflex: New Ideas of its Purpose, Properties, Neural Substrate, and Disorders

Conventional views of the Vestibulo-Ocular Reflex (VOR) have emphasized testing with caloric stimuli and by passively rotating patients at low frequencies in a chair. The properties of the VOR tested under these conditions differ from the performance of this reflex during the natural function for which it evolved-locomotion. Only the VOR (and not visually mediated eye movements) can cope with the high-frequency angular and linear perturbations of the head that occur during locomotion; this is achieved by generating eye movements at short latency (less than 16 msec). Interpretation of vestibular testing is enhanced by the realization that, although the di- and trisynaptic components of the VOR are essential for this short-latency response, the overall accuracy and plasticity of the VOR depend upon a distributed, parallel network of neurons involving the vestibular nuclei. Neurons in this network variously encode inputs from the labyrinthine semicircular canals and otoliths, as well as from the visual and somatosensory systems. The central vestibular pathways branch to contact vestibular cortex (for perception) and the spinal cord (for control of posture). Thus, the vestibular nuclei basically coordinate the stabilization of gaze and posture, and contribute to the perception of verticality and self-motion. Consequently, brainstem disorders that disrupt the VOR cause not just only nystagmus, but also instability of posture (eg, increased fore-aft sway in patients with downbeat nystagmus) and disturbance of spatial orientation (eg, tilt of the subjective visual vertical in Wallenberg's syndrome).

Leigh, R. John↗
NASA NTRS1995

Dynamic properties of the human vestibulo-ocular reflex during head rotations in roll

We investigated the dynamic properties of the human vestibulo-ocular reflex (VOR) during roll head rotations in three human subjects using the magnetic search coil technique. In the first of two experiments, we quantify the behavior of the ocular motor plant in the torsional plane. The subject's eye was mechanically displaced into intorsion, extorsion or abduction, and the dynamic course of return of the eye to its resting position was measured. The mean predominant time constants of return were 210 msec from intorsion, 83 msec from extorsion, and 217 msec from abduction, although there was considerable variability of results from different trials and subjects. In the second experiment, we quantify the efficacy of velocity-to-position integration of the vestibular signal. Position-step stimuli were used to test the torsional or horizontal VOR, being applied with subjects heads erect or supine. After a torsional position-step, the eye drifted back to its resting position, but after a horizontal position-step the eye held its new horizontal position. To interpret these responses we used a simple model of the VOR with parameters of the ocular motor plant set to values determined during Exp 1. The time constant of the velocity-to-position neural integrator was smaller (typically 2 sec) in the torsional plane than in the horizontal plane (> 20 sec). No disconjugacy of torsional eye movements was observed. Thus, the dynamic properties of the VOR in roll differ significantly from those of the VOR in yaw, reflecting different visual demands placed on this reflex in these two planes.

NASA Discipline Neuroscience↗
NASA NTRS1993

A reevaluation of the vestibulo-ocular reflex: new ideas of its purpose, properties, neural substrate, and disorders

Conventional views of the vestibulo-ocular reflex (VOR) have emphasized testing with caloric stimuli and by passively rotating patients at low frequencies in a chair. The properties of the VOR tested under these conditions differ from the performance of this reflex during the natural function for which it evolved--locomotion. Only the VOR (and not visually mediated eye movements) can cope with the high-frequency angular and linear perturbations of the head that occur during locomotion; this is achieved by generating eye movements at short latency (< 16 msec). Interpretation of vestibular testing is enhanced by the realization that, although the di- and trisynaptic components of the VOR are essential for this short-latency response, the overall accuracy and plasticity of the VOR depend upon a distributed, parallel network of neurons involving the vestibular nuclei. Neurons in this network variously upon a distributed, parallel network of neurons involving the vestibular nuclei. Neurons in this network variously encode inputs from the labyrinthine semicircular canals and otoliths, as well as from the visual and somatosensory systems. The central vestibular pathways branch to contact vestibular cortex (for perception) and the spinal cord (for control of posture). Thus, the vestibular nuclei basically coordinate the stabilization of gaze and posture, and contribute to the perception of verticality and self-motion. Consequently, brainstem disorders that disrupt the VOR cause not just only nystagmus, but also instability of posture (eg, increased fore-aft sway in patients with downbeat nystagmus) and disturbance of spatial orientation (eg, tilt of the subjective visual vertical in Wallenberg's syndrome).

Review↗
NASA NTRS1996

Saccades to remembered targets: the effects of smooth pursuit and illusory stimulus motion

1. Measurements were made in four normal human subjects of the accuracy of saccades to remembered locations of targets that were flashed on a 20 x 30 deg random dot display that was either stationary or moving horizontally and sinusoidally at +/-9 deg at 0.3 Hz. During the interval between the target flash and the memory-guided saccade, the "memory period" (1.4 s), subjects either fixated a stationary spot or pursued a spot moving vertically sinusoidally at +/-9 deg at 0.3 Hz. 2. When saccades were made toward the location of targets previously flashed on a stationary background as subjects fixated the stationary spot, median saccadic error was 0.93 deg horizontally and 1.1 deg vertically. These errors were greater than for saccades to visible targets, which had median values of 0.59 deg horizontally and 0.60 deg vertically. 3. When targets were flashed as subjects smoothly pursued a spot that moved vertically across the stationary background, median saccadic error was 1.1 deg horizontally and 1.2 deg vertically, thus being of similar accuracy to when targets were flashed during fixation. In addition, the vertical component of the memory-guided saccade was much more closely correlated with the "spatial error" than with the "retinal error"; this indicated that, when programming the saccade, the brain had taken into account eye movements that occurred during the memory period. 4. When saccades were made to targets flashed during attempted fixation of a stationary spot on a horizontally moving background, a condition that produces a weak Duncker-type illusion of horizontal movement of the primary target, median saccadic error increased horizontally to 3.2 deg but was 1.1 deg vertically. 5. When targets were flashed as subjects smoothly pursued a spot that moved vertically on the horizontally moving background, a condition that induces a strong illusion of diagonal target motion, median saccadic error was 4.0 deg horizontally and 1.5 deg vertically; thus the horizontal error was greater than under any other experimental condition. 6. In most trials, the initial saccade to the remembered target was followed by additional saccades while the subject was still in darkness. These secondary saccades, which were executed in the absence of visual feedback, brought the eye closer to the target location. During paradigms involving horizontal background movement, these corrections were more prominent horizontally than vertically. 7. Further measurements were made in two subjects to determine whether inaccuracy of memory-guided saccades, in the horizontal plane, was due to mislocalization at the time that the target flashed, misrepresentation of the trajectory of the pursuit eye movement during the memory period, or both. 8. The magnitude of the saccadic error, both with and without corrections made in darkness, was mislocalized by approximately 30% of the displacement of the background at the time that the target flashed. The magnitude of the saccadic error also was influenced by net movement of the background during the memory period, corresponding to approximately 25% of net background movement for the initial saccade and approximately 13% for the final eye position achieved in darkness. 9. We formulated simple linear models to test specific hypotheses about which combinations of signals best describe the observed saccadic amplitudes. We tested the possibilities that the brain made an accurate memory of target location and a reliable representation of the eye movement during the memory period, or that one or both of these was corrupted by the illusory visual stimulus. Our data were best accounted for by a model in which both the working memory of target location and the internal representation of the horizontal eye movements were corrupted by the illusory visual stimulus. We conclude that extraretinal signals played only a minor role, in comparison with visual estimates of the direction of gaze, in planning eye movements to remembered targ.

Clinical Trial↗
NASA NTRS1997

Models of Speed Discrimination

The prime purpose of this project was to investigate various theoretical issues concerning the integration of information across visual space. To date, most of the research efforts in the study of the visual system seem to have been focused in two almost non-overlaping directions. One research focus has been the low level perception as studied by psychophysics. The other focus has been the study of high level vision exemplified by the study of object perception. Most of the effort in psychophysics has been devoted to the search for the fundamental "features" of perception. The general idea is that the most peripheral processes of the visual system decompose the input into features that are then used for classification and recognition. The experimental and theoretical focus has been on finding and describing these analyzers that decompose images into useful components. Various models are then compared to the physiological measurements performed on neurons in the sensory systems. In the study of higher level perception, the work has been focused on the representation of objects and on the connections between various physical effects and object perception. In this category we find the perception of 3D from a variety of physical measurements including motion, shading and other physical phenomena. With few exceptions, there seem to be very limited development of theories describing how the visual system might combine the output of the analyzers to form the representation of visual objects. Therefore, the processes underlying the integration of information over space represent critical aspects of vision system. The understanding of these processes will have implications on our expectations for the underlying physiological mechanisms, as well as for our models of the internal representation for visual percepts. In this project, we explored several mechanisms related to spatial summation, attention, and eye movements. The project comprised three components: 1. Modeling visual search for the detection of speed deviation. 2. Perception of moving objects. 3. Exploring the role of eye movements in various visual tasks.

Source record↗
NASA NTRS1973

Automated nystagmus analysis

Several methods have recently been used for on-line analysis of nystagmus: A digital computer program has been developed to accept sampled records of eye position, detect fast phase components, and output cumulative slow phase position, continuous slow phase velocity, instantaneous fast phase frequency, and other parameters. The slow phase velocity is obtained by differentiation of the calculated cumulative position rather than the original eye movement record. Also, a prototype analog device has been devised which calculates the velocity of the slow phase component during caloric testing. Examples of clinical and research eye movement records analyzed with these devices are shown.

Oman, C. M.↗
NASA NTRS1998

STS-90 Day 10 Highlights

On this tenth day of the STS-90 mission, the flight crew, Cmdr. Richard A. Searfoss, Pilot Scott D. Altman, and Mission Specialists Richard M. Linnehan, Dafydd Rhys Williams and Kathryn P. Hire, and Payload Specialists Jay C. Buckey and James A. Pawelczyk have a relatively light day of scientific activity on board Columbia. The science crew of Mission Specialists Rick Linnehan and Dave Williams, along with Payload Specialists Jay Buckey and Jim Pawelczyk, continue investigations into how the human nervous system adapts to the weightlessness of space. All four serve as subjects in a vestibular experiment that uses an on-board rotating chair. The Visual and Vestibular Integration System (VVIS) correlates eye movements with balance. Developed by the European Space Agency, the chair stimulates the human balance system with both spinning and tilting sensations. Infrared video cameras observe and capture the eye movements that accompany the exercise.

Source record↗
NASA NTRS2006

"Far" and "Near" Visual Acuity While Walking and the Collective Contributions of Non-Ocular Mechanisms to Gaze Stabilization

Gaze stabilization was quantified in subjects (n=11) as they walked on a motorized treadmill (1.8 m/s) and viewed visual targets at two viewing distances. A "far" target was positioned at 4 m (FAR) in front of the subject and the "near" target was placed at a distance of 0.5 m (NEAR). A direct measure of visual acuity was used to assess the overall effectiveness of the gaze stabilization system. The contributions of nonocular mechanisms to the gaze goal were also quantified using a measure of the distance between the subject and point in space where fixation of the visual target would require the least eye movement amplitude (i.e. the head fixation distance (HFD)). Kinematic variables mirrored those of previous investigations with the vertical trunk translation and head pitch signals, and the lateral translation and head yaw signals maintaining what appear as antiphase relationships. However, an investigation of the temporal relationships between the maxima and minima of the vertical translation and head pitch signals show that while the maximum in vertical translation occurs at the point of the minimum head pitch signal, the inverse is not true. The maximum in the head pitch signal lags the vertical translation minimum by an average of greater than 12 percent of the step cycle time. Three HFD measures, one each for data in the sagittal and transverse planes, and one that combined the movements from both planes, all revealed changes between the FAR and NEAR target viewing conditions. This reorganization of the nonocular degrees of freedom while walking was consistent with a strategy to reduce the magnitude of the eye movements required when viewing the NEAR target. Despite this reorganization, acuity measures show that image stabilization is not occurring while walking and viewing the NEAR target. Group means indicate that visual acuity is not affected while walking in the FAR condition, but a decrement of 0.15 logMAR (i.e. 1.5 eye chart lines) exists between the standing and walking acuity measures when viewing the NEAR target.

Peters, Brian T.↗
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 NTRS2003

Investigation of Neural Strategies of Visual Search

The goal of this project was to measure how neurons in the superior colliculus (SC) change their activity during a visual search task. Specifically, we proposed to measure how the activity of these neurons was altered by the discriminability of visual targets and to test how these changes might predict the changes in the subjects performance. The primary rationale for this study was that understanding how the information encoded by these neurons constrains overall search performance would foster the development of better models of human performance. Work performed during the period supported by this grant has achieved these aims. First, we have recorded from neurons in the superior colliculus (SC) during a visual search task in which the difficulty of the task and the performance of the subject was systematically varied. The results from these single-neuron physiology experiments shows that prior to eye movement onset, the difference in activity across the ensemble of neurons reaches a fixed threshold value, reflecting the operation of a winner-take-all mechanism. Second, we have developed a model of eye movement decisions based on the principle of winner-take-all . The model incorporates the idea that the overt saccade choice reflects only one of the multiple saccades prepared during visual discrimination, consistent with our physiological data. The value of the model is that, unlike previous models, it is able to account for both the latency and the percent correct of saccade choices.

Krauzlis, Richard J.↗
NASA NTRS1981

Effects of rectilinear acceleration, optokinetic and caloric stimuli in space

The set of experiments comprising the Spacelab 1ES201 package designed to investigate the human vestibular system and equilibratory function in weightlessness are described. The specific objectives of the experiments include: (1) the determination of the threshold of perception of linear oscillatory motion; (2) measurement of physiological and subjective responses to supra threshold, linear and angular motion stimuli; (3) study of the postural adjustments, eye movements, and illusions of attitude and motion evoked by optokinetic stimuli, (i.e., moving visual patterns) in order to assess visual/vestibular interactions; (4) examination of the effect of thermal stimulations of the vestibular apparatus to determine if the eye movements elicited by the 'caloric test' are used by a density gradient in the semicircular canal; and (5) investigation of the pathogenesis of space motion sickness by recording signs and symptoms during the course of vestibular stimulation and, specifically, when the test subject is exposed to sustained, linear oscillatory motion.

Vonbaumgarten, R.↗
NASA NTRS1981

Physiological mechanisms of the nystagmus produced by rotations about an earth-horizontal axis

The physiological basis of the nystagmus produced by rotation about an earth-horizontal axis is investigated with particular emphasis on the unidirectional nystagmus attributed to a bias component. Eye movement recordings were made with dc electro-oculography in alert squirrel monkeys and afferent responses were recorded from semicircular canals and otolith receptors of anesthetized animals upon rotations in the pitch and yaw planes. The eye-movement recordings show the rotation responses in the squirrel monkey to resemble those of other species, including persistent horizontal and vertical nystagmuses during yaw and pitch rotations, respectively, a unidirectional nystagmus at low rotation speeds, and postrotatory responses of relatively small amplitude and duration. The vestibular nerve recordings do not show a directionally specific dc response that can account for the bias component, but instead exhibit sinusoidal responses of peak amplitudes 0-15 and 25-75 spikes/sec for the canals and otolith, respectively. Results thus indicate that the dc signal to the oculomotor centers responsible for the nystagmus is of central origin, most likely based on some transformation of the otolith signals.

Goldberg, J. M.↗
NASA NTRS2000

Primate translational vestibuloocular reflexes. I. High-frequency dynamics and three-dimensional properties during lateral motion

The dynamics and three-dimensional (3-D) properties of the primate translational vestibuloocular reflex (trVOR) for high-frequency (4-12 Hz, +/-0.3-0.4 g) lateral motion were investigated during near-target viewing at center and eccentric targets. Horizontal response gains increased with frequency and depended on target eccentricity. The larger the horizontal and vertical target eccentricity, the steeper the dependence of horizontal response gain on frequency. In addition to horizontal eye movements, robust torsional response components also were present at all frequencies. During center-target fixation, torsional response phase was opposite (anticompensatory) to that expected for an "apparent" tilt response. Instead torsional response components depended systematically on vertical-target eccentricity, increasing in amplitude when looking down and reversing phase when looking up. As a result the trVOR eye velocity vector systematically tilted away from a purely horizontal direction, through an angle that increased with vertical eccentricity with a slope of approximately 0.7. This systematic dependence of torsional eye velocity tilt on vertical eye position suggests that the trVOR might follow the 3-D kinematic requirements that have been shown to govern visually guided eye movements and near-target fixation.

NASA Discipline Neuroscience↗
NASA NTRS2005

Descending projections of the hamster intergeniculate leaflet: relationship to the sleep/arousal and visuomotor systems

The intergeniculate leaflet (IGL), homolog of the primate pregeniculate nucleus, modulates circadian rhythms. However, its extensive anatomical connections suggest that it may regulate other systems, particularly those for visuomotor function and sleep/arousal. Here, descending IGL-efferent pathways are identified with the anterograde tracer, Phaseolus vulgaris leucoagglutinin, with projections to over 50 brain stem nuclei. Projections of the ventral lateral geniculate are similar, but more limited. Many of the nuclei with IGL afferents contribute to circuitry governing visuomotor function. These include the oculomotor, trochlear, anterior pretectal, Edinger-Westphal, and the terminal nuclei; all layers of the superior colliculus, interstitial nucleus of the medial longitudinal fasciculus, supraoculomotor periaqueductal gray, nucleus of the optic tract, the inferior olive, and raphe interpositus. Other target nuclei are known to be involved in the regulation of sleep, including the lateral dorsal and pedunculopontine tegmentum. The dorsal raphe also receives projections from the IGL and may contribute to both sleep/arousal and visuomotor function. However, the locus coeruleus and medial vestibular nucleus, which contribute to sleep and eye movement regulation and which send projections to the IGL, do not receive reciprocal projections from it. The potential involvement of the IGL with the sleep/arousal system is further buttressed by existing evidence showing IGL-efferent projections to the ventrolateral preoptic area, dorsomedial, and medial tuberal hypothalamus. In addition, the great majority of all regions receiving IGL projections also receive input from the orexin/hypocretin system, suggesting that this system contributes not only to the regulation of sleep, but to eye movement control as well.

Non-NASA Center↗
NASA NTRS1987

Comparison of smooth pursuit and combined eye-head tracking in human subjects with deficient labyrinthine function

The effects of deficient labyrinthine function on smooth visual tracking with the eyes and head were investigated, using ten patients with bilateral peripheral vestibular disease and ten normal controls. Active, combined eye-head tracking (EHT) was significantly better in patients than smooth pursuit with the eyes alone, whereas normal subjects pursued equally well in both cases. Compensatory eye movements during active head rotation in darkness were always less in patients than in normal subjects. These data were used to examine current hypotheses that postulate central cancellation of the vestibulo-ocular reflex (VOR) during EHT. A model that proposes summation of an integral smooth pursuit command and VOR/compensatory eye movements is consistent with the findings. Observation of passive EHT (visual fixation of a head-fixed target during en bloc rotation) appears to indicate that in this mode parametric gain changes contribute to modulation of the VOR.

Leigh, R. J.↗
NASA NTRS2009

Translational Vestibulo-Ocular Reflexes During Off-Vertical Axis Rotation

The translational vestibulo-ocular reflex (tVOR) is an otolith-mediated response that stabilizes near vision during linear acceleration at higher frequencies where visually mediated reflexes are not adequate. The modulation of horizontal and vergence eye movements during Off-Vertical Axis Rotation (OVAR) are presumed to reflect the tVOR in response to the continuously varying linear acceleration in the interaural and nasooccipital axes, respectively. The purpose of this study was to examine the effect of frequency and fixation distance on the modulation of slow phase eye velocity (SPV) as further evidence that the tVOR is elicited during OVAR. Eighteen subjects were rotated about their longitudinal axis tilted by 30 deg off-vertical. Rotational velocities varied between 18 and 288 deg/sec corresponding to a frequency range of 0.05 to 0.8 Hz. Fixation distance was altered by asking subjects to imagine stationary targets that were briefly presented at 0.5, 1 and 2 m during some rotation cycles. The target flash was 40 msec in the nose-up position at eye level. Oculomotor responses were recorded in the dark using infrared binocular videography. Sinusoidal curve fits were used to derive amplitude, phase and bias velocity of the eye movements across multiple rotation cycles. Consistent with previous studies, the modulation of both horizontal and vergence SPV increased with stimulus frequency. The effect of fixation distance was negligible at lower frequencies. The modulation of horizontal and vergence SPV was; however, proportional to fixation distance during OVAR at 0.8 Hz. This increasing sensitivity and dependence on fixation distance of horizontal and vergence SPV during OVAR is consistent with tVOR characteristics measured during other types of linear motion. We conclude that the modulation of horizontal and vergence SPV will be diagnostically more useful at higher stimulus frequencies where the tVOR is more robust.

Wood, Scott J.↗