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At least 163 records · Page 9

Adaptation of the phase of the human linear vestibulo-ocular reflex (LVOR) and effects on the oculomotor neural integrator

The phase of the translational linear VOR (LVOR) can be adaptively modified by exposure to a visual-vestibular mismatch. We extend here our earlier work on LVOR phase adaptation, and discuss the role of the oculomotor neural integrator. Ten subjects were oscillated laterally at 0.5 Hz, 0.3 g peak acceleration, while sitting upright on a linear sled. LVOR was assessed before and after adaptation with subjects tracking the remembered location of a target at 1 m in the dark. Phase and gain were measured by fitting sine waves to the desaccaded eye movements, and comparing sled and eye position. To adapt LVOR phase, the subject viewed a computer-generated stereoscopic visual display, at a virtual distance of 1 m, that moved so as to require either a phase lead or a phase lag of 53 deg. Adaptation lasted 20 min, during which subjects were oscillated at 0.5 Hz/0.3 g. Four of five subjects produced an adaptive change in the lag condition (range 4-45 deg), and each of five produced a change in the lead condition (range 19-56 deg), as requested. Changes in drift on eccentric gaze suggest that the oculomotor velocity-to-position integrator may be involved in the phase changes.

Non-NASA Center↗

Eye Movement Latencies to Direction Change for Different Classes of Motion

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

Mulligan, Jeffrey B.↗

Validation of a Manually Oscillating Chair for In-The-Field Assessment of Dynamic Visual Acuity on Crewmembers Within Hours of Returning From Long-Duration Spaceflight

Long-duration spaceflight results in sensorimotor adaptations, which cause functional deficits during gravitational transitions, such as landing on a planetary surface after long-duration microgravity exposure. Both the vestibular system and the central nervous system are affected by gravitational transitions. These systems are responsible for coordinating head and eye movements via the vestibulo-ocular reflex (VOR) and go through an adaptation period upon exposure to microgravity. Consequently, they must also re-adapt to Earth's gravitational environment upon landing. This re-adaptation causes decrements in gaze control and dynamic visual acuity, with crewmembers reporting oscillopsia and blurred vision caused by retinal slip, or the inability to keep an image focused on their retina. This is thought to drive motion sickness symptoms experienced by most crewmembers following landing. Retinal slip can be estimated by dynamic visual acuity (DVA); visual acuity while in motion. Previously, DVA has been assessed in the laboratory where subjects walked at 6.4 km/hr on a motorized treadmill. Using this method, Peters et al. (2011) found that DVA is worsened in astronauts by an average of 0.75 eye-chart lines one day after landing. However, it is believed that re-adaptation occurs quickly and that DVA might be worse immediately upon re-exposure to a gravitational environment. Since many crewmembers are unable to walk safely upon landing, it was necessary to develop a method for replicating the vertical head movements associated with walking. In addition, the use of a chair to imitate the head displacement caused by walking isolates eye-head interactions without allowing for trunk and lower-body compensation, as seen with treadmill walking (Mulavara & Bloomberg 2003). Therefore, a modality for assessing DVA in the field within a few hours of landing was developed. In this study, we validated the ability of a manually operated oscillating chair to reproduce the oscillatory frequency of walking on a treadmill. Healthy non-astronaut subjects (n=14) participated in one test session and completed three static (seated) and three dynamic (walking/oscillated) visual acuity tests. DVA was assessed using a motorized treadmill, an automated oscillating chair, and a manually operated chair, both developed in the Neuroscience Laboratory at JSC. The automated chair was motor-driven and set to oscillate vertically at 2 Hz with a vertical displacement of +/- 5 cm to simulate vertical translation while walking. The manually operated chair was oscillated vertically by a test operator to the beat of a metronome at 120 beats/min (2 Hz) and a vertical displacement of approximately +/- 5 cm. As the subject was oscillated, they were asked to discern the direction gap of Landolt-C optotypes of varying sizes and verbally reported the direction while an operator recorded their response using a gamepad. Subjects were outfitted with accelerometers (sampling rate = 128 Hz) on their head, trunk and lumbar spine. A fast Fourier transform was performed on the vertical trunk acceleration to compare the peak and spread of the distribution of oscillation frequencies for each oscillating condition. The spread of the frequency distribution for the manual chair was not significantly different from either the treadmill or the automated chair. However, all three conditions had similar non-zero standard error values, suggesting a variance in head movement frequency which may affect DVA. The average oscillation frequency of the manual chair (1.85 Hz) was significantly different (α=0.05) from that of treadmill walking (2.24 Hz), but not significantly different from that of the automated chair (1.85 Hz) and all three conditions had small standard errors (SEM = 0.04, 0.06, and 0.08 Hz for manual, treadmill, and automated respectively). This implies that both chairs oscillate at a frequency below that of treadmill walking, but are comparable to each other and reproducible across sessions. Additionally, DVA scores did not vary significantly across conditions. The smaller spread values of the oscillating chairs' frequencies indicated mitigation of variation induced by locomotor strategies, which enables better examination of the issue of VOR adaptation. Furthermore, due to the deconditioned state of crewmembers in the initial hours after landing, it is easier to transport a manual bouncing chair into the field and safer to perform a vision test while seated in a chair versus walking on a treadmill. Therefore, the manually oscillating chair has been deemed to meet and exceed the DVA testing capabilities previously obtained by treadmill walking.

Kreutzberg, G. A.↗

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↗

The vestibulo-ocular reflex of the squirrel monkey during eccentric rotation and roll tilt

The vestibulo-ocular reflexes (VOR) are determined not only by angular acceleration, but also by the presence of gravity and linear acceleration. This phenomenon was studied by measuring three-dimensional nystagmic eye movements, with implanted search coils, in six male squirrel monkeys during eccentric rotation. Monkeys were rotated in the dark at a constant velocity of 200 degrees/s (centrally or 79 cm off axis) with the axis of rotation always aligned with gravity and the spinal axis of the upright monkeys. The monkey's orientation (facing-motion or back-to-motion) had a dramatic influence on the VOR. These experiments show that: (a) the axis of eye rotation always shifted toward alignment with gravito-inertial force; (b) the peak value of horizontal slow phase eye velocity was greater with the monkey facing-motion than with back-to-motion; and (c) the time constant of horizontal eye movement decay was smaller with the monkey facing-motion than with back-to-motion. All of these findings were statistically significant and consistent across monkeys. In another set of tests, the same monkeys were rapidly tilted about their naso-occipital (roll) axis. Tilted orientations of 45 degrees and 90 degrees were maintained for 1 min. Other than a compensatory angular VOR during the angular rotation, no consistent eye velocity response was observed during or following the tilt for any of the six monkeys. The absence of any eye movement response following tilt weighs against the possibility that translational linear VOR responses are due to simple high-pass filtering of the otolith signals. The VOR response during eccentric rotation was divided into the more familiar angular VOR and linear VOR components. The angular component is known to depend upon semicircular canal dynamics and central influences. The linear component of the response decays rapidly with a mean duration of only 6.6 s, while the axis of eye rotation rapidly aligns (< 10 s) with gravito-inertial force. These results are consistent with the hypothesis that the measurement of gravito-inertial force by the otolith organs is resolved into central estimates of linear acceleration and gravity, such that the central estimate of gravitational force minus the central estimate of linear acceleration approximately equals the otolith measurement of gravito-inertial force.

NASA Discipline Neuroscience↗

Resolution of sensory ambiguities for gaze stabilization requires a second neural integrator

The ability to simultaneously move in the world and maintain stable visual perception depends critically on the contribution of vestibulo-ocular reflexes (VORs) to gaze stabilization. It is traditionally believed that semicircular canal signals drive compensatory responses to rotational head disturbances (rotational VOR), whereas otolith signals compensate for translational movements [translational VOR (TVOR)]. However, a sensory ambiguity exists because otolith afferents are activated similarly during head translations and reorientations relative to gravity (i.e., tilts). Extra-otolith cues are, therefore, necessary to ensure that dynamic head tilts do not elicit a TVOR. To investigate how extra-otolith signals contribute, we characterized the temporal and viewing distance-dependent properties of a TVOR elicited in the absence of a lateral acceleration stimulus to the otoliths during combined translational/rotational motion. We show that, in addition to otolith signals, angular head position signals derived by integrating sensory canal information drive the TVOR. A physiological basis for these results is proposed in a model with two distinct integration steps. Upstream of the well known oculomotor velocity-to-position neural integrator, the model incorporates a separate integration element that could represent the "velocity storage integrator," whose functional role in the oculomotor system has so far remained controversial. We propose that a key functional purpose of the velocity storage network is to temporally integrate semicircular canal signals, so that they may be used to extract translation information from ambiguous otolith afferent signals in the natural and functionally relevant bandwidth of head movements.

Non-NASA Center↗

Recovery trajectories of vestibulopathic subjects after perturbations during locomotion

We compared the mediolateral (M/L) responses to perturbations during locomotion of vestibulopathic (VP) subjects to those of controls. Eight subjects with unilateral vestibular loss (100% Reduced Vestibular Response from the caloric test) resulting from surgery for vestibular schwannoma and 11 controls were selected for this study. Despite their known vestibulopathy, all VP subjects scored within the normal range on computerized dynamic posturography Sensory Organization Tests. During gait, subjects were given surface perturbations of the right support-phase foot in two possible directions (forward-right and backward-left) at two possible magnitudes (5 and 10 cm) that were randomly mixed with trials having no perturbations. M/L stability was quantified by estimating the length of the M/L moment arm between the support foot and the trunk, and the M/L accelerations of the sternum and the head. The VP group had greater changes (p < 0.05) in their moment arm responses compared to controls. The number of steps that it took for the moment arm oscillations to return to normal and the variability in the moment arms were greater for the VP group. Differences in the sternum and head accelerations between VP and control groups were not as consistent, but there was a trend toward greater response deviations in the VP group for all 4 perturbation types. Increased response magnitude and variability of the VP group is consistent with an increase in their sensory noise of vestibular inputs due to the surgical lesion. Another possibility is a reduced sensitivity to motion inputs. This perturbation approach may prove useful for characterizing subtle vestibulopathies and similar changes in the human orientation mechanism after exposure to microgravity.

NASA Discipline Neuroscience↗

Physiologic adaptation of man in space; Proceedings of the Seventh International Man in Space Symposium, Houston, TX, Feb. 10-13, 1986

Topics discussed in this volume include space motion sickness, cardiovascular adaptation, fluid shifts, extravehicular activity, general physiology, perception, vestibular response modifications, vestibular physiology, and pharmacology. Papers are presented on the clinical characterization and etiology of space motion sickness, ultrasound techniques in space medicine, fluid shifts in weightlessness, Space Shuttle inflight and postflight fluid shifts measured by leg volume changes, and the probability of oxygen toxicity in an 8-psi space suit. Consideration is also given to the metabolic and hormonal status of crewmembers in short-term space flights, adaptive changes in perception of body orientation and mental image rotation in microgravity, the effects of a visual-vestibular stimulus on the vestibulo-ocular reflex, rotation tests in the weightless phase of parabolic flight, and the mechanisms of antimotion sickness drugs.

Holland, Albert W.↗

Dynamic visual acuity using "far" and "near" targets

CONCLUSIONS: DVA may be useful for assessing the functional consequences of an impaired gaze stabilization mechanism or for testing the effectiveness of a rehabilitation paradigm. Because target distance influences the relative contributions of canal and otolith inputs, the ability to measure DVA at near and far viewing distances may also lead to tests that will independently assess canal and otolith function. OBJECTIVE: To present and test a methodology that uses dynamic visual acuity (DVA) to assess the efficacy of compensatory gaze mechanisms during a functionally relevant activity that differentially measures canal and otolith function. MATERIAL AND METHODS: The effect of treadmill walking at a velocity of 1.79 m/s on subjects' visual acuity was assessed at each of two viewing distances. A custom-written threshold determination program was used to display Landolt C optotypes on a laptop computer screen during a "far" (4 m) target condition and on a micro-display for a "near" (50 cm) target condition. The walking acuity scores for each target distance were normalized by subtracting a corresponding acuity measure obtained while standing still on the treadmill belt. RESULTS: As predicted by subjective reports of relative target motion, the decrease in visual acuity was significantly greater (p < 0.00001) for the near compared to the far condition.

Pattern Recognition, Visual/physiology↗

The guidance of saccadic eye movements to perceptually mislocalized visual and non-visual targets

The present experiment examined whether saccadic eye movements to visual targets are dependent on the perceived directions of the targets or on their retinally specified directions. Perceptual mislocations of visual targets were induced by having the target light attached to a subject's stationary hand while his biceps or triceps muscle was vibrated. Such vibration leads to apparent extension or flexion of the subject's restrained forearm and perceived visual motion of the stationary target light. Subjects always made accurate saccadic eye movements to a visual target, even when the target was perceptually mislocalized by as much as 20 deg. By contrast, when subjects made saccadic eye movements to a nonvisual target, the location of their hand in the dark, they always looked to the perceived direction of the target even though it did not necessarily correspond to the true direction. These findings indicate that a distinction is maintained between 'reflexive aspects' of oculomotor control related to foveation and the computation of perceived visual direction.

Lackner, J. R.↗

Inactivation of Semicircular Canals Causes Adaptive Increases in Otolith-driven Tilt Responses

Growing experimental and theoretical evidence suggests a functional synergy in the processing of otolith and semicircular canal signals for the generation of the vestibulo-ocular reflexes (VORs). In this study we have further tested this functional interaction by quantifying the adaptive changes in the otolith-ocular system during both rotational and translational movements after surgical inactivation of the semicircular canals. For 0.1- 0.5 Hz (stimuli for which there is no recovery of responses from the plugged canals), pitch and roll VOR gains recovered during earth- horizontal (but not earth-vertical) axis rotations. Corresponding changes were also observed in eye movements elicited by translational motion (0.1 - 5 Hz). Specifically, torsional eye movements increased during lateral motion, whereas vertical eye movements increased during fore-aft motion. The findings indicate that otolith signals can be adapted according to compromised strategy that leads to improved gaze stabilization during motion. Because canal-plugged animals permanently lose the ability to discriminate gravitoinertial accelerations, adapted animals can use the presence of gravity through otolith-driven tilt responses to assist gaze stabilization during earth-horizontal axis rotations.

Angelaki, Dora E.↗

Reflexive obstacle avoidance for kinematically-redundant manipulators

Dexterous telerobots incorporating 17 or more degrees of freedom operating under coordinated, sensor-driven computer control will play important roles in future space operations. They will also be used on Earth in assignments like fire fighting, construction and battlefield support. A real time, reflexive obstacle avoidance system, seen as a functional requirement for such massively redundant manipulators, was developed using arm-mounted proximity sensors to control manipulator pose. The project involved a review and analysis of alternative proximity sensor technologies for space applications, the development of a general-purpose algorithm for synthesizing sensor inputs, and the implementation of a prototypical system for demonstration and testing. A 7 degree of freedom Robotics Research K-2107HR manipulator was outfitted with ultrasonic proximity sensors as a testbed, and Robotics Research's standard redundant motion control algorithm was modified such that an object detected by sensor arrays located at the elbow effectively applies a force to the manipulator elbow, normal to the axis. The arm is repelled by objects detected by the sensors, causing the robot to steer around objects in the workspace automatically while continuing to move its tool along the commanded path without interruption. The mathematical approach formulated for synthesizing sensor inputs can be employed for redundant robots of any kinematic configuration.

Karlen, James P.↗

Lightweight device to stimulate and monitor human vestibulo-ocular reflex

A helmet formed of a rigid shell is disclosed. The shell is lined with several air filled bladders to contact firmly the head of a user. The shell has a rigid chin bar supporting a bite bar connected fixedly to a mouthpiece bearing against the teeth and hard palate to firmly anchor the helmet without movement. The outer shell surface supports various air pumping bulbs and accelerometers. Separate left and right visor pivot on the side guided in a central tongue and groove track to move optical lens mounts into the user's field of vision. The chin bar is connected to the shell by a pair of releasable clasps. A safety lanyard connects to the clasps to quickly pull pins from the clasps to enable quick release in case of motion sickness.

McStravick, M. Catherine↗

Sensory processing in the vestibular nuclei during active head movements

Many secondary vestibular neurons are sensitive to head on trunk rotation during reflex-induced and voluntary head movements. During passive whole body rotation the interaction of head on trunk signals related to the vestibulo-collic reflex with vestibular signals increases the rotational gain of many secondary vestibular neurons, including many that project to the spinal cord. In some units, the sensitivity to head on trunk and vestibular input is matched and the resulting interaction produces an output that is related to the trunk velocity in space. In other units the head on trunk inputs are stronger and the resulting interaction produces an output that is larger during the reflex. During voluntary head movements, inputs related to head on trunk movement combine destructively with vestibular signals, and often cancel the sensory reafferent consequences of self-generated movements. Cancellation of sensory vestibular signals was observed in all of the antidromically identified secondary vestibulospinal units, even though many of these units were not significantly affected by reflexive head on trunk movements. The results imply that the inputs to vestibular neurons related to head on trunk rotation during reflexive and voluntary movements arise from different sources. We suggest that the relative strength of reflexive head on trunk input to different vestibular neurons might reflect the different functional roles they have in controlling the posture of the neck and body.

Non-NASA Center↗

Stimulus Characteristics for Vestibular Stochastic Resonance to Improve Balance Function

Stochastic resonance (SR) is a mechanism by which noise can enhance the response of neural systems to relevant sensory signals. Studies have shown that imperceptible stochastic vestibular electrical stimulation, when applied to normal young and elderly subjects, significantly improved their ocular stabilization reflexes in response to whole-body tilt as well as balance performance during postural disturbances. The goal of this study was to optimize the amplitude characteristics of the stochastic vestibular signals for balance performance during standing on an unstable surface. Subjects performed a standard balance task of standing on a block of foam with their eyes closed. Bipolar stochastic electrical stimulation was applied to the vestibular system using constant current stimulation through electrodes placed over the mastoid process behind the ears. Amplitude of the signals varied in the range of 0-700 microamperes. Balance performance was measured using a force plate under the foam block, and inertial motion sensors were placed on the torso and head. Balance performance with stimulation was significantly greater (10%-25%) than with no stimulation. The signal amplitude at which performance was maximized was in the range of 100-300 microamperes. Optimization of the amplitude of the stochastic signals for maximizing balance performance will have a significant impact on development of vestibular SR as a unique system to aid recovery of function in astronauts after long-duration space flight or in patients with balance disorders.

Mulavara, Ajitkumar↗

Modification of the Passive Vestibulo-Ocular Reflex During and After Short-Duration Spaceflight

The vestibulo-ocular reflex (VOR) is mediated by integration of canal and otolith inputs to generate compensatory eye movements during head movements. We hypothesized that adaptive change in vestibular processing of gravitoinertial cues would be reflected by plane specific modification of the VOR during passive whole-body rotation during and after spaceflight. Using a repeated measures design, the VOR was assessed in four payload crewmembers in yaw, pitch and roll planes during multiple sessions before, during and after an 8 day orbital mission (STS-42). Rotation was about an earth-vertical axis during ground tests, with the head located off-axis by up to 45cm during pitch and roll rotation (peak acceleration less than 0.2g). The motion profiles included sum-of-sinusoids between 0.02 - 1.39 Hz (yaw), single sinusoids between 0.05-1.25 Hz (yaw and pitch) and velocity steps (yaw, pitch and roll). Eye movements were recorded with both video and electro-oculographic techniques. As expected, VOR gain changes were greater in pitch than in yaw. During pitch rotation, there was a progressive shift in the axis of eye movements during the flight, which was also present during the early post-flight period. This increased horizontal component during pitch, most prevalent at 0.2 Hz, was interpreted as an increase in a translational vergence response elicited during eccentric rotation as subjects imagined a wall fixed target. There was also an increased horizontal component during the eccentric roll step runs performed on flight day 7. These results are consistent with a frequency-dependent increase in otolith-mediated translational VOR responses following adaptation to microgravity. We conclude that the adaptive changes in the VOR are likely to be greatest in the frequency range where there is a cross-over of otolith-mediated tilt and translation responses.

Wood, S. J.↗

Effect of Target Location on Dynamic Visual Acuity During Passive Horizontal Rotation

The vestibulo-ocular reflex (VOR) generates eye rotation to compensate for potential retinal slip in the specific plane of head movement. Dynamic visual acuity (DVA) has been utilized as a functional measure of the VOR. The purpose of this study was to examine changes in accuracy and reaction time when performing a DVA task with targets offset from the plane of rotation, e.g. offset vertically during horizontal rotation. Visual acuity was measured in 12 healthy subjects as they moved a hand-held joystick to indicate the orientation of a computer-generated Landolt C "as quickly and accurately as possible." Acuity thresholds were established with optotypes presented centrally on a wall-mounted LCD screen at 1.3 m distance, first without motion (static condition) and then while oscillating at 0.8 Hz (DVA, peak velocity 60 deg/s). The effect of target location was then measured during horizontal rotation with the optotypes randomly presented in one of nine different locations on the screen (offset up to 10 deg). The optotype size (logMar 0, 0.2 or 0.4, corresponding to Snellen range 20/20 to 20/50) and presentation duration (150, 300 and 450 ms) were counter-balanced across five trials, each utilizing horizontal rotation at 0.8 Hz. Dynamic acuity was reduced relative to static acuity in 7 of 12 subjects by one step size. During the random target trials, both accuracy and reaction time improved proportional to optotype size. Accuracy and reaction time also improved between 150 ms and 300 ms presentation durations. The main finding was that both accuracy and reaction time varied as a function of target location, with greater performance decrements when acquiring vertical targets. We conclude that dynamic visual acuity varies with target location, with acuity optimized for targets in the plane of motion. Both reaction time and accuracy are functionally relevant DVA parameters of VOR function.

Appelbaum, Meghan↗

Experimental tests of a superposition hypothesis to explain the relationship between the vestibuloocular reflex and smooth pursuit during horizontal combined eye-head tracking in humans

1. We used a modeling approach to test the hypothesis that, in humans, the smooth pursuit (SP) system provides the primary signal for cancelling the vestibuloocular reflex (VOR) during combined eye-head tracking (CEHT) of a target moving smoothly in the horizontal plane. Separate models for SP and the VOR were developed. The optimal values of parameters of the two models were calculated using measured responses of four subjects to trials of SP and the visually enhanced VOR. After optimal parameter values were specified, each model generated waveforms that accurately reflected the subjects' responses to SP and vestibular stimuli. The models were then combined into a CEHT model wherein the final eye movement command signal was generated as the linear summation of the signals from the SP and VOR pathways. 2. The SP-VOR superposition hypothesis was tested using two types of CEHT stimuli, both of which involved passive rotation of subjects in a vestibular chair. The first stimulus consisted of a "chair brake" or sudden stop of the subject's head during CEHT; the visual target continued to move. The second stimulus consisted of a sudden change from the visually enhanced VOR to CEHT ("delayed target onset" paradigm); as the vestibular chair rotated past the angular position of the stationary visual stimulus, the latter started to move in synchrony with the chair. Data collected during experiments that employed these stimuli were compared quantitatively with predictions made by the CEHT model. 3. During CEHT, when the chair was suddenly and unexpectedly stopped, the eye promptly began to move in the orbit to track the moving target. Initially, gaze velocity did not completely match target velocity, however; this finally occurred approximately 100 ms after the brake onset. The model did predict the prompt onset of eye-in-orbit motion after the brake, but it did not predict that gaze velocity would initially be only approximately 70% of target velocity. One possible explanation for this discrepancy is that VOR gain can be dynamically modulated and, during sustained CEHT, it may assume a lower value. Consequently, during CEHT, a smaller-amplitude SP signal would be needed to cancel the lower-gain VOR. This reduction of the SP signal could account for the attenuated tracking response observed immediately after the brake. We found evidence for the dynamic modulation of VOR gain by noting differences in responses to the onset and offset of head rotation in trials of the visually enhanced VOR.(ABSTRACT TRUNCATED AT 400 WORDS).

Non-NASA Center↗