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At least 361 records · Page 20

Yaw and pitch visual-vestibular interaction in weightlessness

Both yaw and pitch visual-vestibular interactions at two separate frequencies of chair rotation (0.2 and 0.8 Hz) in combination with a single velocity of optokinetic stimulus (36 degrees/s) were used to investigate the effects of sustained weightlessness on neural strategies adopted by astronaut subjects to cope with the stimulus rearrangement of spaceflight. Pitch and yaw oscillation in darkness at 0.2 and 0.8 Hz without optokinetic stimulation, and constant velocity linear optokinetic stimulation at 18, 36, and 54 degrees/s presented relative to the head with the subject stationary, were used as controls for the visual-vestibular interactions. The results following 8 days of space flight showed no significant changes in: (1) either the horizontal and vertical vestibulo-ocular reflex (VOR) gain, phase, or bias; (2) the yaw visual-vestibular response (VVR); or (3) the horizontal or vertical optokinetic (OKN) slow phase velocity (SPV). However, significant changes were observed: (1) when during pitch VVR at 0.2 Hz late inflight, the contribution of the optokinetic input to the combined oculomotor response was smaller than during the stationary OKN SPV measurements, followed by an increased contribution during the immediate postflight testing; and (2) when during pitch VVR at 0.8 Hz, the component of the combined oculomotor response due to the underlying vertical VOR was more efficiently suppressed early inflight and less suppressed immediately postflight compared with preflight observations. The larger OKN response during pitch VVR at 0.2 Hz and the better suppression of VOR during pitch VVR at 0.8 Hz postflight are presumably due to the increased role of vision early inflight and immediately after spaceflight, as previously observed in various studies. These results suggest that the subjects adopted a neural strategy to structure their spatial orientation in weightlessness by reweighting visual, otolith, and perhaps tactile/somatic signals.

NASA Center JSC↗

Artificial gravity considerations for a mars exploration mission

Artificial gravity (AG), as a means of preventing physiological deconditioning of astronauts during long-duration space flights, presents certain special challenges to the otolith organs and the adaptive capabilities of the CNS. The key issues regarding the choice of AG acceleration, radius, and rotation rate are reviewed from the viewpoints of physiological requirements and human factors disturbances. Head movements and resultant Coriolis forces on the rotating platform may limit the usefulness of economical short centrifuges for other than brief periods of intermittent stimulation.

Review↗

Functional organization of primate translational vestibulo-ocular reflexes and effects of unilateral labyrinthectomy

Translational vestibulo-ocular reflexes (trVORs) are characterized by distinct spatio-temporal properties and sensitivities that are proportional to the inverse of viewing distance. Anodal (inhibitory) labyrinthine stimulation (100 microA, < 2 s) during motion decreased the high-pass filtered dynamics, as well as horizontal trVOR sensitivity and its dependence on viewing distance. Cathodal (excitatory) currents had opposite effects. Translational VORs were also affected after unilateral labyrinthectomy. Animals lost their ability to modulate trVOR sensitivity as a function of viewing distance acutely after the lesion. These deficits partially recovered over time, albeit a significant reduction in trVOR sensitivity as a function of viewing distance remained in compensated animals. During fore-aft motion, the effects of unilateral labyrinthectomy were more dramatic. Both acute and compensated animals permanently lost their ability to modulate fore-aft trVOR responses as a function of target eccentricity. These results suggest that (1) the dynamics and viewing distance-dependent properties of the trVORs are very sensitive to changes in the resting firing rate of vestibular afferents and, consequently, vestibular nuclei neurons; (2) the most irregularly firing primary otolith afferents that are most sensitive to labyrinthine electrical stimulation might contribute to reflex dynamics and sensitivity; (3) inputs from both labyrinths are necessary for the generation of the translational VORs.

Non-NASA Center↗

Signal processing related to the vestibulo-ocular reflex during combined angular rotation and linear translation of the head

The contributions of vestibular nerve afferents and central vestibular pathways to the angular (AVOR) and linear (LVOR) vestibulo-ocular reflex were studied in squirrel monkeys during fixation of near and far targets. Irregular vestibular afferents did not appear to be necessary for the LVOR, since when they were selectively silenced with galvanic currents the LVOR was essentially unaffected during both far- and near-target viewing. The linear translation signals generated by secondary AVOR neurons in the vestibular nuclei were, on average, in phase with head velocity, inversely related to viewing distance, and were nearly as strong as AVOR-related signals. We suggest that spatial-temporal transformation of linear head translation signals to angular eye velocity commands is accomplished primarily by the addition of viewing distance multiplied, centrally integrated, otolith regular afferent signals to angular VOR pathways.

NASA Discipline Neuroscience↗

Effects of static orientation upon human optokinetic afternystagmus

"Normal" human subjects were placed in a series of 5 static orientations with respect to gravity and were asked to view an optokinetic display moving at a constant angular velocity. The axis of rotation coincided with the subject's rostro-caudal axis and produced horizontal optokinetic nystagmus and afternystagmus. Wall (1) previously reported that these optokinetic afternystagmus responses were not well characterized by parametric fits to slow component velocity. The response for nose-up, however, was larger than for nose-down. This suggested that the horizontal eye movements measured during optokinetic stimulation might include an induced linear VOR component as presented in the body of this paper. To investigate this hypothesis, another analysis of these data has been made using cumulative slow component eye position. Some subjects' responses had reversals in afternystagmus direction. These reversals were "filled in" by a zero slow component velocity. This method of analysis gives a much more consistent result across subjects and shows that, on average, responses from the nose-down horizontal (prone) orientation are greatly reduced (p < 0.05) compared to other horizontal and vertical orientations. Average responses are compared to responses predicted by a model previously used to predict successfully the responses to post-rotatory nystagmus after earth horizontal axis rotation. Ten of 11 subjects had larger responses in their supine than their prone orientation. Application of horizontal axis optokinetic afternystagmus for clinical otolith function testing, and implications for altered gravity experiments are discussed.

Non-NASA Center↗

Vestibulospinal adaptation to microgravity

Human balance control is known to be transiently disrupted after spaceflight; however, the mechanisms responsible for postflight postural ataxia are still under investigation. In this report, we propose a conceptual model of vestibulospinal adaptation based on theoretical adaptive control concepts and supported by the results from a comprehensive study of balance control recovery after spaceflight. The conceptual model predicts that immediately after spaceflight the balance control system of a returning astronaut does not expect to receive gravity-induced afferent inputs and that descending vestibulospinal control of balance is disrupted until the central nervous system is able to cope with the newly available vestibular otolith information. Predictions of the model are tested using data from a study of the neurosensory control of balance in astronauts immediately after landing. In that study, the mechanisms of sensorimotor balance control were assessed under normal, reduced, and/or altered (sway-referenced) visual and somatosensory input conditions. We conclude that the adaptive control model accurately describes the neurobehavioral responses to spaceflight and that similar models of altered sensory, motor, or environmental constraints are needed clinically to predict responses that patients with sensorimotor pathologies may have to various visual-vestibular or changing stimulus environments.

manned↗

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↗

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

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

NASA Discipline Neuroscience↗

Differential central projections of vestibular afferents in pigeons

The question of whether a differential distribution of vestibular afferent information to central nuclear neurons is present in pigeons was studied using neural tracer compounds. Discrete tracing of afferent fibers innervating the individual semicircular canal and otolith organs was produced by sectioning individual branches of the vestibular nerve that innervate the different receptor organs and applying crystals of horseradish peroxidase, or a horseradish peroxidase/cholera toxin mixture, or a biocytin compound for neuronal uptake and transport. Afferent fibers and their terminal distributions within the brainstem and cerebellum were visualized subsequently. Discrete areas in the pigeon central nervous system that receive primary vestibular input include the superior, dorsal lateral, ventral lateral, medial, descending, and tangential vestibular nuclei; the A and B groups; the intermediate, medial, and lateral cerebellar nuclei; and the nodulus, the uvula, and the paraflocculus. Generally, the vertical canal afferents projected heavily to medial regions in the superior and descending vestibular nuclei as well as the A group. Vertical canal projections to the medial and lateral vestibular nuclei were observed but were less prominent. Horizontal canal projections to the superior and descending vestibular nuclei were much more centrally located than those of the vertical canals. A more substantial projection to the medial and lateral vestibular nuclei was seen with horizontal canal afferents compared to vertical canal fibers. Afferents innervating the utricle and saccule terminated generally in the lateral regions of all vestibular nuclei in areas that were separate from the projections of the semicircular canals. In addition, utricular fibers projected to regions in the vestibular nuclei that overlapped with the horizontal semicircular canal terminal fields, whereas saccular afferents projected to regions that received vertical canal fiber terminations. Lagenar afferents projected throughout the cochlear nuclei, to the dorsolateral regions of the cerebellar nuclei, and to lateral regions of the superior, lateral, medial, and descending vestibular nuclei.

NASA Discipline Neuroscience↗

Reduction of the elevator illusion from continued hypergravity exposure and visual error-corrective feedback

Ten subjects served as their own controls in two conditions of continuous, centrifugally produced hypergravity (+2 Gz) and a 1-G control condition. Before and after exposure, open-loop measures were obtained of (1) motor control, (2) visual localization, and (3) hand-eye coordination. During exposure in the visual feedback/hypergravity condition, subjects received terminal visual error-corrective feedback from their target pointing, and in the no-visual feedback/hypergravity condition they pointed open loop. As expected, the motor control measures for both experimental conditions revealed very short lived underreaching (the muscle-loading effect) at the outset of hypergravity and an equally transient negative aftereffect on returning to 1 G. The substantial (approximately 17 degrees) initial elevator illusion experienced in both hypergravity conditions declined over the course of the exposure period, whether or not visual feedback was provided. This effect was tentatively attributed to habituation of the otoliths. Visual feedback produced a smaller additional decrement and a postexposure negative after-effect, possible evidence for visual recalibration. Surprisingly, the target-pointing error made during hypergravity in the no-visual-feedback condition was substantially less than that predicted by subjects' elevator illusion. This finding calls into question the neural outflow model as a complete explanation of this illusion.

NASA Program Space Physiology and Countermeasures↗

Space flight and neurovestibular adaptation

Space flight represents a form of sensory stimulus rearrangement requiring modification of established terrestrial response patterns through central reinterpretation. Evidence of sensory reinterpretation is manifested as postflight modifications of eye/head coordination, locomotor patterns, postural control strategies, and illusory perceptions of self or surround motion in conjunction with head movements. Under normal preflight conditions, the head is stabilized during locomotion, but immediately postflight reduced head stability, coupled with inappropriate eye/head coordination, results in modifications of gait. Postflight postural control exhibits increased dependence on vision which compensates for inappropriate interpretation of otolith and proprioceptive inputs. Eye movements compensatory for perceived self motion, rather than actual head movements have been observed postflight. Overall, the in-flight adaptive modification of head stabilization strategies, changes in head/eye coordination, illusionary motion, and postural control are maladaptive for a return to the terrestrial environment.

Review, Tutorial↗

Recovery of postural equilibrium control following space flight

DSO 605 represents the first large study of balance control following spaceflight. Data collected during DSO 605 confirm the theory that postural ataxia following short duration spaceflight is of vestibular origin. We used the computerized dynamic posturography technique developed by Nashner et al. to study the role of the vestibular system in balance control in astronauts during quiet stance before and after spaceflight. Our results demonstrate unequivocally that balance control is disrupted in all astronauts immediately after return from space. The most severely affected returning crew members performed in the same way as vestibular deficient patients exposed to this test battery. We conclude that otolith mediated spatial reference provided by the terrestrial gravitational force vector is not used by the astronauts balance control systems immediately after spaceflight. Because the postflight ataxia appears to be mediated primarily by CNS adaptation to the altered vestibular inputs caused by loss of gravitational stimulation, we believe that intermittent periods of exposure to artificial gravity may provide an effective in-flight countermeasure. Specifically, we propose that in-flight centrifugation will allow crew members to retain their terrestrial sensory-motor adapted states while simultaneously developing microgravity adapted states. The dual-adapted astronaut should be able to make the transition from microgravity to unit gravity with minimal sensory-motor effects. We have begun a ground based program aimed at developing short arm centrifuge prescriptions designed to optimize adaptation to altered gravitational environments. Results from these experiments are expected to lead directly to in-flight evaluation of the proposed centrifuge countermeasure. Because our computerized dynamic posturography system was able to (1) quantify the postflight postural ataxia reported by crew members and observed by flight surgeons and scientists, (2) track the recovery of normal (preflight) balance control, (3) differentiate between rookie and veteran subjects, and (4) provide normative and clinical databases for comparison, and because our study successfully characterized postflight balance control recovery in a large cross-section of Shuttle crew members, we recommend that this system and protocol be adopted as a standard dependent measure for evaluating the efficacy of countermeasures and/or evaluating the postflight effects of changing mission durations or activities.

Paloski, William H.↗

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↗

Encoding of head acceleration in vestibular neurons. I. Spatiotemporal response properties to linear acceleration

1. Extracellular recordings were made in and around the medial vestibular nuclei in decerebrated rats. Neurons were functionally identified according to their semicircular canal input on the basis of their responses to angular head rotations around the yaw, pitch, and roll head axes. Those cells responding to angular acceleration were classified as either horizontal semicircular canal-related (HC) or vertical semicircular canal-related (VC) neurons. The HC neurons were further characterized as either type I or type II, depending on the direction of rotation producing excitation. Cells that lacked a response to angular head acceleration, but exhibited sensitivity to a change in head position, were classified as purely otolith organ-related (OTO) neurons. All vestibular neurons were then tested for their response to sinusoidal linear translation in the horizontal head plane. 2. Convergence of macular and canal inputs onto central vestibular nuclei neurons occurred in 73% of the type I HC, 79% of the type II HC, and 86% of the VC neurons. Out of the 223 neurons identified as receiving macular input, 94 neurons were further studied, and their spatiotemporal response properties to sinusoidal stimulation with pure linear acceleration were quantified. Data were obtained from 33 type I HC, 22 type II HC, 22 VC, and 17 OTO neurons. 3. For each neuron the angle of the translational stimulus vector was varied by 15, 30, or 45 degrees increments in the horizontal head plane. In all tested neurons, a direction of maximum sensitivity was identified. An interesting difference among neurons was their response to translation along the direction perpendicular to that that produced the maximum response ("null" direction). For the majority of neurons tested, it was possible to evoke a nonzero response during stimulation along the null direction always had response phases that varied as a function of stimulus direction. 4. These spatiotemporal response properties were quantified in two independent ways. First, the data were evaluated on the basis of the traditional one-dimensional principle governed by the "cosine gain rule" and constant response phase at different stimulus orientations. Second, the response gain and phase values that were empirically determined for each orientation of the applied linear stimulus vector were fitted on the basis of a newly developed formalism that treats neuronal responses as exhibiting two-dimensional spatial sensitivity. Thus two response vectors were determined for each neuron on the basis of its response gain and phase at different stimulus directions in the horizontal head plane.(ABSTRACT TRUNCATED AT 400 WORDS).

NASA Discipline Neuroscience↗

An analysis of ocular counterrolling in response to body positions in three-dimensional space

Four normal subjects underwent ocular counterrolling testing in a tiltable chair. Measurements were taken in 62 different body positions in steps of 30 degrees varied rolls and pitches. In each body position the eyes were recorded on video and their roll angle was determined automatically by computer analysis. The ocular counterrolling profile showed a periodic characteristic with maximal amplitude at roll tilts of 60 degrees. In this study we can clearly show that the eyes' rolling response is not systematically affected when lateral body tilts are combined with any tilts in the pitch direction. This undoubtedly implies that the ocular counterrolling was mainly stimulated by the subject's roll angle. As an empirical contribution, this study provides new data specially to be used in modelling and simulating the function of otolith organs.

Posture/physiology↗

Modeling the spatiotemporal organization of velocity storage in the vestibuloocular reflex by optokinetic studies

1. A generalized three-dimensional state space model of visual vestibular interaction was developed. Matrix and dynamical system operators associated with inputs from the semicircular canals, otolith velocity estimator, and the visual system have been incorporated into the model, which focus on their relationship to the velocity storage integrator. 2. A relationship was postulated between the eigenvalues and the direction of the eigenvectors of the system matrix and the orientation of the spatial vertical. It was assumed that the system matrix for a tilted position was a composition of two linear transformations of the system matrix for the upright position. One transformation modifies the eigenvalues of the system matrix, whereas another rotates the eigenvectors of the system matrix. The pitch and roll eigenvectors rotate with the head, whereas the yaw axis eigenvector remains approximately spatially invariant. 3. Based on the three-dimensional model, a computational procedure was formulated to identify the eigenvalues and eigenvectors of the system matrix with the use of a modification of the marquardt algorithm. With the use of data obtained from a monkey, it was shown that the three-dimensional behavior of velocity storage cannot be predicted solely in terms of its time constants, i.e., the inverse of its eigenvalues. With the use of the same eigenvalues the data could either be fit or not fit, depending on the eigenvector directions. Therefore, it is necessary to specify eigenvector directions when characterizing velocity storage in three dimensions. 4. Parameters found with the use of the Marquardt algorithm were incorporated into the model. Diagonal matrices in a head coordinate frame were introduced for coupling the visual system to the integrator and to the direct optokinetic pathway. Simulations of optokinetic nystagmus (OKN) and optokinetic after-nystagmus (OKAN) were run. The model predicted the behavior of yaw and pitch OKN and OKAN when the animal is upright. It also predicted the cross-coupling in the side down position. The trajectories in velocity space were also accurately simulated. 5. One of the predictions of the model is that when the stimulus direction is along an eigenvector, the trajectory in velocity space is a straight line. Using the "spectral width" of the residuals from a straight line sequence during OKAN, we developed a methodology to estimate how close the OKAN decay was to an eigenvector trajectory. 6. Thus we have developed a model-based approach for studying and interpreting the response characteristics of velocity storage in three dimensions.(ABSTRACT TRUNCATED AT 400 WORDS).

Non-NASA Center↗

M.I.T./Canadian vestibular experiments on the Spacelab-1 mission: 2. Visual vestibular tilt interaction in weightlessness

Adaptation to weightlessness includes the substitution of other sensory signals for the no longer appropriate graviceptor information concerning static spatial orientation. Visual-vestibular interaction producing roll circularvection was studied in weightlessness to assess the influence of otolith cues on spatial orientation. Preliminary results from four subjects tested on Spacelab-1 indicate that visual orientation effects were stronger in weightlessness than pre-flight. The rod and frame test of visual field dependence showed a weak post-flight increase in visual influence. Localized tactile cues applied to the feet in space reduced subjective vection strength.

Non-NASA Center↗

Motion Cueing Algorithm Development: Human-Centered Linear and Nonlinear Approaches

While the performance of flight simulator motion system hardware has advanced substantially, the development of the motion cueing algorithm, the software that transforms simulated aircraft dynamics into realizable motion commands, has not kept pace. Prior research identified viable features from two algorithms: the nonlinear "adaptive algorithm", and the "optimal algorithm" that incorporates human vestibular models. A novel approach to motion cueing, the "nonlinear algorithm" is introduced that combines features from both approaches. This algorithm is formulated by optimal control, and incorporates a new integrated perception model that includes both visual and vestibular sensation and the interaction between the stimuli. Using a time-varying control law, the matrix Riccati equation is updated in real time by a neurocomputing approach. Preliminary pilot testing resulted in the optimal algorithm incorporating a new otolith model, producing improved motion cues. The nonlinear algorithm vertical mode produced a motion cue with a time-varying washout, sustaining small cues for longer durations and washing out large cues more quickly compared to the optimal algorithm. The inclusion of the integrated perception model improved the responses to longitudinal and lateral cues. False cues observed with the NASA adaptive algorithm were absent. The neurocomputing approach was crucial in that the number of presentations of an input vector could be reduced to meet the real time requirement without degrading the quality of the motion cues.

Houck, Jacob A.↗