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At least 37 records · Page 2

Binocular Coordination of the Human Vestibulo-Ocular Reflex during Off-axis Pitch Rotation

Head movements in the sagittal pitch plane typically involve off-axis rotation requiring both vertical and horizontal vergence ocular reflexes to compensate for angular and translational motion relative to visual targets of interest. The purpose of this study was to compare passive pitch VOR responses during rotation about an Earth-vertical axis (canal only cues) with off-axis rotation (canal and otolith cues). Methods. Eleven human subjects were oscillated sinusoidally at 0.13, 0.3 and 0.56 Hz while lying left-side down with the interaural axis either aligned with the axis of rotation or offset by 50 cm. In a second set of measurements, twelve subjects were also tested during sinusoidally varying centrifugation over the same frequency range. The modulation of vertical and horizontal vergence ocular responses was measured with a binocular videography system. Results. Off-axis pitch rotation enhanced the vertical VOR at lower frequencies and enhanced the vergence VOR at higher frequencies. During sinusoidally varying centrifugation, the opposite trend was observed for vergence, with both vertical and vergence vestibulo-ocular reflexes being suppressed at the highest frequency. Discussion. These differential effects of off-axis rotation over the 0.13 to 0.56 Hz range are consistent with the hypothesis that otolith-ocular reflexes are segregated in part on the basis of stimulus frequency. At the lower frequencies, tilt otolith-ocular responses compensate for declining canal input. At higher frequencies, translational otolith-ocular reflexes compensate for declining visual contributions to the kinematic demands required for fixating near targets.

Wood, S. J.↗

Three-dimensional ocular kinematics during eccentric rotations: evidence for functional rather than mechanical constraints

Previous studies have reported that the translational vestibuloocular reflex (TVOR) follows a three-dimensional (3D) kinematic behavior that is more similar to visually guided eye movements, like pursuit, rather than the rotational VOR (RVOR). Accordingly, TVOR rotation axes tilted with eye position toward an eye-fixed reference frame rather than staying relatively fixed in the head like in the RVOR. This difference arises because, contrary to the RVOR where peripheral image stability is functionally important, the TVOR like pursuit and saccades cares to stabilize images on the fovea. During most natural head and body movements, both VORs are simultaneously activated. In the present study, we have investigated in rhesus monkeys the 3D kinematics of the combined VOR during yaw rotation about eccentric axes. The experiments were motivated by and quantitatively compared with the predictions of two distinct hypotheses. According to the first (fixed-rule) hypothesis, an eye-position-dependent torsion is computed downstream of a site for RVOR/TVOR convergence, and the combined VOR axis would tilt through an angle that is proportional to gaze angle and independent of the relative RVOR/TVOR contributions to the total eye movement. This hypothesis would be consistent with the recently postulated mechanical constraints imposed by extraocular muscle pulleys. According to the second (image-stabilization) hypothesis, an eye-position-dependent torsion is computed separately for the RVOR and the TVOR components, implying a processing that takes place upstream of a site for RVOR/TVOR convergence. The latter hypothesis is based on the functional requirement that the 3D kinematics of the combined VOR should be governed by the need to keep images stable on the fovea with slip on the peripheral retina being dependent on the different functional goals of the two VORs. In contrast to the fixed-rule hypothesis, the data demonstrated a variable eye-position-dependent torsion for the combined VOR that was different for synergistic versus antagonistic RVOR/TVOR interactions. Furthermore, not only were the eye-velocity tilt slopes of the combined VOR as much as 10 times larger than what would be expected based on extraocular muscle pulley location, but also eye velocity during antagonistic RVOR/TVOR combinations often tilted opposite to gaze. These results are qualitatively and quantitatively consistent with the image-stabilization hypothesis, suggesting that the eye-position-dependent torsion is computed separately for the RVOR and the TVOR and that the 3D kinematics of the combined VOR are dependent on functional rather than mechanical constraints.

Non-NASA Center↗

Effect of Vestibular Impairment on Cerebral Blood Flow Response to Dynamic Roll Tilt

Change to upright posture results in reductions in cerebral perfusion pressure due to hydrostatic pressure changes related to gravity. Since vestibular organs, specifically the otoliths, provide information on position relative to gravity, vestibular inputs may assist in adaptation to the upright posture. The goal of this study was to examine the effect of direct vestibular stimulation on cerebral blood flow (CBF). To examine the role of otolith inputs we screened 165 subjects for vestibular function and classified subjects as either normal or impaired based on ocular torsion. Ocular torsion, an indication of otolith function, was assessed during sinusoidal roll tilt of 20 degrees at 0.01 Hz (100 sec per cycle). Subjects with torsion one SD below the mean were classified as impaired while subjects one SD above the mean were considered normal. During one session subjects were placed in a chair that was sinusoidally rotated 25 degrees in the roll plane at five frequencies: 0.25 & 0.125 Hz for 80 sec, 0.0625 Hz for 160 sec and 0.03125 Hz and 0.015625 Hz for 320 sec. During testing, CBF (transcranial Doppler), blood pressure (Finapres), and end tidal CO2 (Puritan Bennet) were measured continuously. Ocular torsion was assessed from infrared images of the eyes. All rotations were done in the dark with subjects fixated on a red LED directly at the center of rotation. In the normal group, dynamic tilt resulted in significant changes in both blood pressure and cerebral blood flow velocity that was related to the frequency of stimulus. In contrast the impaired group did not show similar patterns. As expected normal subjects demonstrated significant ocular torsion that was related to stimulus frequency while impaired subjects had minimal changes. These data suggest that vestibular inputs have direct effects on cerebral blood flow regulation during dynamic tilt. Supported by NASA.

Serrador, J. M.↗

Brain-Computer Interfaces for 1-D and 2-D Cursor Control: Designs Using Volitional Control of the EEG Spectrum or Steady-State Visual Evoked Potentials

We have developed and tested two EEG-based brain-computer interfaces (BCI) for users to control a cursor on a computer display. Our system uses an adaptive algorithm, based on kernel partial least squares classification (KPLS), to associate patterns in multichannel EEG frequency spectra with cursor controls. Our first BCI, Target Practice, is a system for one-dimensional device control, in which participants use biofeedback to learn voluntary control of their EEG spectra. Target Practice uses a KF LS classifier to map power spectra of 30-electrode EEG signals to rightward or leftward position of a moving cursor on a computer display. Three subjects learned to control motion of a cursor on a video display in multiple blocks of 60 trials over periods of up to six weeks. The best subject s average skill in correct selection of the cursor direction grew from 58% to 88% after 13 training sessions. Target Practice also implements online control of two artifact sources: a) removal of ocular artifact by linear subtraction of wavelet-smoothed vertical and horizontal EOG signals, b) control of muscle artifact by inhibition of BCI training during periods of relatively high power in the 40-64 Hz band. The second BCI, Think Pointer, is a system for two-dimensional cursor control. Steady-state visual evoked potentials (SSVEP) are triggered by four flickering checkerboard stimuli located in narrow strips at each edge of the display. The user attends to one of the four beacons to initiate motion in the desired direction. The SSVEP signals are recorded from eight electrodes located over the occipital region. A KPLS classifier is individually calibrated to map multichannel frequency bands of the SSVEP signals to right-left or up-down motion of a cursor on a computer display. The display stops moving when the user attends to a central fixation point. As for Target Practice, Think Pointer also implements wavelet-based online removal of ocular artifact; however, in Think Pointer muscle artifact is controlled via adaptive normalization of the SSVEP. Training of the classifier requires about three minutes. We have tested our system in real-time operation in three human subjects. Across subjects and sessions, control accuracy ranged from 80% to 100% correct with lags of 1-5 seconds for movement initiation and turning.

Trejo, Leonard J.↗

Visual suppression of the vestibulo-ocular reflex during space flight

Visual suppression of the vestibulo-ocular reflex was studied in 16 subjects on 4 Space Shuttle missions. Eye movements were recorded by electro-oculography while subjects fixated a head mounted target during active sinusoidal head oscillation at 0.3 Hz. Adequacy of suppression was evaluated by the number of nystagmus beats, the mean amplitude of each beat, and the cumulative amplitude of nystagmus during two head oscillation cycles. Vestibulo-ocular reflex suppression was unaffected by space flight. Subjects with space motion sickness during flight had significantly more nystagmus beats than unaffected individuals. These susceptible subjects also tended to have more nystagmus beats before flight.

Uri, John J.↗

Vestibular-ocular accommodation reflex in man

Stimulation of the vestibular system by angular acceleration produces widespread sensory and motor effects. The present paper studies a motor effect which has not been reported in the literature, i.e., the influence of rotary acceleration of the body on ocular accommodation. The accommodation of 10 young men was recorded before and after a high-level deceleration to zero velocity following 30 sec of rotating. Accommodation was recorded continuously on an infrared optometer for 110 sec under two conditions: while the subjects observed a target set at the far point, and while they viewed the same target through a 0.3-mm pinhole. Stimulation by high-level rotary deceleration produced positive accommodation or a pseudomyopia under both conditions, but the positive accommodation was substantially greater and lasted much longer during fixation through the pinhole. It is hypothesized that this increase in accommodation is a result of a vestibular-ocular accommodation reflex.

Clark, B.↗

Rapid motor learning in the translational vestibulo-ocular reflex

Motor learning was induced in the translational vestibulo-ocular reflex (TVOR) when monkeys were repeatedly subjected to a brief (0.5 sec) head translation while they tried to maintain binocular fixation on a visual target for juice rewards. If the target was world-fixed, the initial eye speed of the TVOR gradually increased; if the target was head-fixed, the initial eye speed of the TVOR gradually decreased. The rate of learning acquisition was very rapid, with a time constant of approximately 100 trials, which was equivalent to <1 min of accumulated stimulation. These learned changes were consolidated over >or=1 d without any reinforcement, indicating induction of long-term synaptic plasticity. Although the learning generalized to targets with different viewing distances and to head translations with different accelerations, it was highly specific for the particular combination of head motion and evoked eye movement associated with the training. For example, it was specific to the modality of the stimulus (translation vs rotation) and the direction of the evoked eye movement in the training. Furthermore, when one eye was aligned with the heading direction so that it remained motionless during training, learning was not expressed in this eye, but only in the other nonaligned eye. These specificities show that the learning sites are neither in the sensory nor the motor limb of the reflex but in the sensory-motor transformation stage of the reflex. The dependence of the learning on both head motion and evoked eye movement suggests that Hebbian learning may be one of the underlying cellular mechanisms.

Non-NASA Center↗

Primate translational vestibuloocular reflexes. IV. Changes after unilateral labyrinthectomy

The effects of unilateral labyrinthectomy on the properties of the translational vestibuloocular reflexes (trVORs) were investigated in rhesus monkeys trained to fixate near targets. Translational motion stimuli consisted of either steady-state lateral and fore-aft sinusoidal oscillations or short-lasting transient displacements. During small-amplitude, steady-state sinusoidal lateral oscillations, a small decrease in the horizontal trVOR sensitivity and its dependence on viewing distance was observed during the first week after labyrinthectomy. These deficits gradually recovered over time. In addition, the vertical response component increased, causing a tilt of the eye velocity vector toward the lesioned side. During large, transient lateral displacements, the deficits were larger and longer lasting. Responses after labyrinthectomy were asymmetric, with eye velocity during movements toward the side of the lesion being more compromised. The most profound effect of the lesions was observed during fore-aft motion. Whereas responses were kinematically appropriate for fixation away from the side of the lesion (e.g., to the left after right labyrinthectomy), horizontal responses were anticompensatory during fixation at targets located ipsilateral to the side of the lesion (e.g., for targets to the right after right labyrinthectomy). This deficit showed little recovery during the 3-mo post-labyrinthectomy testing period. These results suggest that inputs from both labyrinths are important for the proper function of the trVORs, although the details of how bilateral signals are processed and integrated remain unknown.

NASA Discipline Neuroscience↗

Contribution of the cerebellar flocculus to gaze control during active head movements

The flocculus and ventral paraflocculus are adjacent regions of the cerebellar cortex that are essential for controlling smooth pursuit eye movements and for altering the performance of the vestibulo-ocular reflex (VOR). The question addressed in this study is whether these regions of the cerebellum are more globally involved in controlling gaze, regardless of whether eye or active head movements are used to pursue moving visual targets. Single-unit recordings were obtained from Purkinje (Pk) cells in the floccular region of squirrel monkeys that were trained to fixate and pursue small visual targets. Cell firing rate was recorded during smooth pursuit eye movements, cancellation of the VOR, combined eye-head pursuit, and spontaneous gaze shifts in the absence of targets. Pk cells were found to be much less sensitive to gaze velocity during combined eye-head pursuit than during ocular pursuit. They were not sensitive to gaze or head velocity during gaze saccades. Temporary inactivation of the floccular region by muscimol injection compromised ocular pursuit but had little effect on the ability of monkeys to pursue visual targets with head movements or to cancel the VOR during active head movements. Thus the signals produced by Pk cells in the floccular region are necessary for controlling smooth pursuit eye movements but not for coordinating gaze during active head movements. The results imply that individual functional modules in the cerebellar cortex are less involved in the global organization and coordination of movements than with parametric control of movements produced by a specific part of the body.

NASA Discipline Neuroscience↗

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.↗

Primate translational vestibuloocular reflexes. II. Version and vergence responses to fore-aft motion

To maintain binocular fixation on near targets during fore-aft translational disturbances, largely disjunctive eye movements are elicited the amplitude and direction of which should be tuned to the horizontal and vertical eccentricities of the target. The eye movements generated during this task have been investigated here as trained rhesus monkeys fixated isovergence targets at different horizontal and vertical eccentricities during 10 Hz fore-aft oscillations. The elicited eye movements complied with the geometric requirements for binocular fixation, although not ideally. First, the corresponding vergence angle for which the movement of each eye would be compensatory was consistently less than that dictated by the actual fixation parameters. Second, the eye position with zero sensitivity to translation was not straight ahead, as geometrically required, but rather exhibited a systematic dependence on viewing distance and vergence angle. Third, responses were asymmetric, with gains being larger for abducting and downward compared with adducting and upward gaze directions, respectively. As frequency was varied between 4 and 12 Hz, responses exhibited high-pass filter properties with significant differences between abduction and adduction responses. As a result of these differences, vergence sensitivity increased as a function of frequency with a steeper slope than that of version. Despite largely undercompensatory version responses, vergence sensitivity was closer to ideal. Moreover, the observed dependence of vergence sensitivity on vergence angle, which was varied between 2.5 and 10 MA, was largely linear rather than quadratic (as geometrically predicted). We conclude that the spatial tuning of eye velocity sensitivity as a function of gaze and viewing distance follows the general geometric dependencies required for the maintenance of foveal visual acuity. However, systematic deviations from ideal behavior exist that might reflect asymmetric processing of abduction/adduction responses perhaps because of different functional dependencies of version and vergence eye movement components during translation.

NASA Program Biomedical Research and Countermeasur↗

Advanced Techniques for Assessment of Postural and Locomotor Ataxia, Spatial Orientation, and Gaze Stability

In addition to adapting to microgravity, major neurovestibular problems of space flight include postflight difficulties with standing, walking, turning corners, and other activities that require stable upright posture and gaze stability. These difficulties inhibit astronauts' ability to stand or escape from their vehicle during emergencies. The long-ter7n goal of the NSBRI is the development of countermeasures to ameliorate the effects of long duration space flight. These countermeasures must be tested with valid and reliable tools. This project aims to develop quantitative, parametric approaches for assessing gaze stability and spatial orientation during normal gait and when gait is perturbed. Two of this year's most important findings concern head fixation distance and ideal trajectory analysis. During a normal cycle of walking the head moves up and down linearly. A simultaneous angular pitching motion of the head keeps it aligned toward an imaginary point in space at a distance of about one meter in front of a subject and along the line of march. This distance is called the head fixation distance. Head fixation distance provides the fundamental framework necessary for understanding the functional significance of the vestibular reflexes that couple head motion to eye motion. This framework facilitates the intelligent design of counter-measures for the effects of exposure to microgravity upon the vestibular ocular reflexes. Ideal trajectory analysis is a simple candidate countermeasure based upon quantifying body sway during repeated up and down stair stepping. It provides one number that estimates the body sway deviation from an ideal sinusoidal body sway trajectory normalized on the subject's height. This concept has been developed with NSBRI funding in less than one year. These findings are explained in more detail below. Compared to assessments of the vestibuo-ocular reflex, analysis of vestibular effects on locomotor function is relatively less well developed and quantified. We are improving this situation by applying methodologies such as nonlinear orbital stability to quantify responses and by using multivariate statistical approaches to link together the responses across separate tests. In this way we can exploit the information available and increase the ability to discriminate between normal and pathological responses. Measures of stability and orientation are compared to measures such as dynamic visual acuity and with balance function tests. The responses of normal human subjects and of patients having well documented pathophysiologies are being characterized. When these studies are completed, we should have a clearer idea about normal and abnormal patterns of eye, head, and body movements during locomotion and their stability in a wide range of environments. We plan eventually to use this information to validate the efficacy of candidate neurovestibular and neuromuscular rehabilitative techniques. Some representative studies made during this year are summarized.

Wall, Conrad., III↗

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↗

Primate translational vestibuloocular reflexes. III. Effects of bilateral labyrinthine electrical stimulation

The effects of functional, reversible ablation and potential recruitment of the most irregular otolith afferents on the dynamics and sensitivity of the translational vestibuloocular reflexes (trVORs) were investigated in rhesus monkeys trained to fixate near and far targets. Translational motion stimuli consisted of either steady-state lateral and fore-aft sinusoidal oscillations or short-lasting transient lateral head displacements. Short-duration (usually <2 s) anodal (inhibitory) and cathodal (excitatory) currents (50-100 microA) were delivered bilaterally during motion. In the presence of anodal labyrinthine stimulation, trVOR sensitivity and its dependence on viewing distance were significantly decreased. In addition, anodal currents significantly increased phase lags. During transient motion, anodal stimulation resulted in significantly lower initial eye acceleration and more sluggish responses. Cathodal currents tended to have opposite effects. The main characteristics of these results were simulated by a simple model where both regularly and irregularly discharging afferents contribute to the trVORs. Anodal labyrinthine currents also were found to decrease eye velocity during long-duration, constant velocity rotations, although results were generally more variable compared with those during translational motion.

NASA Program Biomedical Research and Countermeasur↗

Inertial vestibular coding of motion: concepts and evidence

Central processing of inertial sensory information about head attitude and motion in space is crucial for motor control. Vestibular signals are coded relative to a non-inertial system, the head, that is virtually continuously in motion. Evidence for transformation of vestibular signals from head-fixed sensory coordinates to gravity-centered coordinates have been provided by studies of the vestibulo-ocular reflex. The underlying central processing depends on otolith afferent information that needs to be resolved in terms of head translation related inertial forces and head attitude dependent pull of gravity. Theoretical solutions have been suggested, but experimental evidence is still scarce. It appears, along these lines, that gaze control systems are intimately linked to motor control of head attitude and posture.

Review↗

Human discrimination of visual direction of motion with and without smooth pursuit eye movements

It has long been known that ocular pursuit of a moving target has a major influence on its perceived speed (Aubert, 1886; Fleischl, 1882). However, little is known about the effect of smooth pursuit on the perception of target direction. Here we compare the precision of human visual-direction judgments under two oculomotor conditions (pursuit vs. fixation). We also examine the impact of stimulus duration (200 ms vs. ~800 ms) and absolute direction (cardinal vs. oblique). Our main finding is that direction discrimination thresholds in the fixation and pursuit conditions are indistinguishable. Furthermore, the two oculomotor conditions showed oblique effects of similar magnitudes. These data suggest that the neural direction signals supporting perception are the same with or without pursuit, despite remarkably different retinal stimulation. During fixation, the stimulus information is restricted to large, purely peripheral retinal motion, while during steady-state pursuit, the stimulus information consists of small, unreliable foveal retinal motion and a large efference-copy signal. A parsimonious explanation of our findings is that the signal limiting the precision of direction judgments is a neural estimate of target motion in head-centered (or world-centered) coordinates (i.e., a combined retinal and eye motion signal) as found in the medial superior temporal area (MST), and not simply an estimate of retinal motion as found in the middle temporal area (MT).

Saccades/physiology↗

Eye and head motion during head turns in spaceflight

Eye-head motion was studied pre-, in- and postflight during single voluntary head turns. A transient increase in vestibulo-ocular reflex (VOR) gain occurred early in the flight, but later trended toward normal. This increased gain was produced by a relative increase in eye counterrotation velocity. Asymmetries in gain with right and left turns also occurred, caused by asymmetries in eye counterrotation velocities. These findings were remarkably similar to those from Soviet primate studies using gaze fixation targets, except the human study trended more rapidly toward normal. These findings differ substantially from those measuring VOR gain by head oscillation, in which no significant changes were found inflight. No visual disturbances were noted in either test condition or in normal activities. These head turn studies are the only ones to date documenting any functional change in VOR in weightlessness.

Thornton, William E.↗

Short-term adaptation of the VOR: non-retinal-slip error signals and saccade substitution

We studied short-term (30 min) adaptation of the vestibulo-ocular reflex (VOR) in five normal humans using a "position error" stimulus without retinal image motion. Both before and after adaptation a velocity gain (peak slow-phase eye velocity/peak head velocity) and a position gain (total eye movement during chair rotation/amplitude of chair motion) were measured in darkness using search coils. The vestibular stimulus was a brief ( approximately 700 ms), 15 degrees chair rotation in darkness (peak velocity 43 degrees /s). To elicit adaptation, a straight-ahead fixation target disappeared during chair movement and when the chair stopped the target reappeared at a new location in front of the subject for gain-decrease (x0) adaptation, or 10 degrees opposite to chair motion for gain-increase (x1.67) adaptation. This position-error stimulus was effective at inducing VOR adaptation, though for gain-increase adaptation the primary strategy was to substitute augmenting saccades during rotation while for gain-decrease adaptation both corrective saccades and a decrease in slow-phase velocity occurred. Finally, the presence of the position-error signal alone, at the end of head rotation, without any attempt to fix upon it, was not sufficient to induce adaptation. Adaptation did occur, however, if the subject did make a saccade to the target after head rotation, or even if the subject paid attention to the new location of the target without actually looking at it.

Non-NASA Center↗