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Isolation, sensory deprivation, and sensory rearrangement effects on visual, auditory, and somesthetic sensation, perception, and spatial orientation
When head motion includes a linear velocity component, eye velocity required to track an earth-fixed target depends upon: a) angular and linear head velocity, b) target distance, and c) direction of gaze relative to the motion trajectory. Recent research indicates that eye movements (LVOR), presumably otolith-mediated, partially compensate for linear velocity in small head excursions on small devices. Canal-mediated eye velocity (AVOR), otolith-mediated eye velocity (LVOR), and Ocular Torsion (OT) can be measured, one by one, on small devices. However, response dynamics that depend upon the ratio of linear to angular velocity in the motion trajectory and on subject orientation relative to the trajectory are present in a centrifuge paradigm. With this paradigm, two 3-min runs yields measures of: LVOR differentially modulated by different subject orientations in the two runs; OT dynamics in four conditions; two directions of "steady-state" OT, and two directions of AVOR. Efficient assessment of the dynamics (and of the underlying central integrative processes) may require a centrifuge radius of 1.0 meters or more. Clinical assessment of the spatial orientation system should include evaluation of central integrative processes that determine the dynamics of these responses.
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.
This article summarizes a variety of newly published findings obtained by the Neuroscience Laboratory, Johnson Space Center, and attempts to place this work within a historical framework of previous results on posture, locomotion, motion sickness, and perceptual responses that have been observed in conjunction with space flight. In this context, we have taken the view that correct transduction and integration of signals from all sensory systems is essential to maintaining stable vision, postural and locomotor control, and eye-hand coordination as components of spatial orientation. The plasticity of the human central nervous system allows individuals to adapt to altered stimulus conditions encountered in a microgravity environment. However, until some level of adaptation is achieved, astronauts and cosmonauts often experience space motion sickness, disturbances in motion control and eye-hand coordination, unstable vision, and illusory motion of the self, the visual scene, or both. Many of the same types of disturbances encountered in space flight reappear immediately after crew members return to earth. The magnitude of these neurosensory, sensory-motor and perceptual disturbances, and the time needed to recover from them, tend to vary as a function of mission duration and the space travelers prior experience with the stimulus rearrangement of space flight. To adequately chart the development of neurosensory changes associated with space flight, we recommend development of enhanced eye movement systems and body position measurement. We also advocate the use of a human small radius centrifuge as both a research tool and as a means of providing on-orbit countermeasures that will lessen the impact of living for long periods of time with out exposure to altering gravito-inertial forces. Copyright 1998 Elsevier Science B.V.
Reason and Brand (1975) noted that motion sickness occurs in many situations involving either passive body motion or active interaction with the world via indirect sensorimotor interfaces (e.g., prism spectacles). As might be expected, motion sickness is being reported in VEs that involve apparent self-motion through space, the best known examples being flight simulators (Kennedy et al., 1990). The goals of this paper are to introduce the motion-sickness symptomatology; to outline some concepts that are central to theories of motion sickness, spatial orientation, and adaptation; and to discuss the implications of some trends in VE research and development.
Adaptation to research paradigms such as rotating rooms and optical alteration of visual feedback during movement results in development of perceptual-motor programs that provide the reflexive assistance that is necessary to skilled control of movement and balance. The discomfort and stomach awareness that occur during the adaptation process has been attributed to conflicting sensory information about the state of motion. Vestibular signals depend on the kinematics of head movements irrespective of the presence or absence of signals from other senses. We propose that sensory conflict when vestibular signals are at least one component of the conflict are innately disturbing and unpleasant. This innate reaction is part of a continuum that operates early in life to prevent development of inefficient perceptual-motor programs. This reaction operates irrespective of and in addition to reward and punishment from parental guidance or goal attainment to yield efficient control of whole body movement in the operating environment of the individual. The same mechanism is involved in adapting the spatial orientation system to strange environments. This conceptual model "explains" why motion sickness is associated with adaptation to novel environments and is in general consistent with motion sickness literature.
The purpose is to discuss the role of geometry in determining the perception of spatial layout and perceived orientation in pictures viewed at an angle. This discussion derives from Cutting's (1988) suggestion, based on his analysis of some of the author's data (Goldstein, 1987), that the changes in perceived orientation that occur when pictures are viewed at an angle can be explained in terms of geometrically produced changes in the picture's virtual space.
A previously formulated mathematical model, describing how observers reconstruct three-dimensional spatial layouts from perspective projections, has been extended to more complex situations. The model assumes that the observer has a priori knowledge of certain characteristics of the viewed objects, like size, shape, or parallelism or perpendicularity of lines or planes. These assumptions are used in a three-dimensional process to reconstruct a spatial layout that `best matches' the perceived lines of sight to the object coordinates. Sources of errors and biases in this process are specified and their effects on model outputs are discussed. An experiment, in which eight subjects judged the relative direction of one object with respect to another, has been conducted to validate the model. The model has been found to generally reproduce the systematic trends of the experimental results and also has provided an analytical explanation for them. The mathematical model is expected to be a useful tool in analyzing and developing pictorial perspective flight displays.
To better understand the mechanisms of human adaptation to rotating environments, we exposed 19 healthy subjects and 8 vestibular-deficient subjects ("abnormal"; four bilateral and four unilateral lesions) to an interaural centripetal acceleration of 1 g (resultant 45 degrees roll-tilt of 1.4 g) on a 0.8-m-radius centrifuge for periods of 90 min. The subjects sat upright (body z-axis parallel to centrifuge rotation axis) in the dark with head stationary, except during 4 min of every 10 min, when they performed head saccades toward visual targets switched on at 3- to 5-s intervals at random locations (within +/- 30 degrees) in the earth-horizontal plane. Eight of the normal subjects also performed the head saccade protocol in a stationary chair adjusted to a static roll-tilt angle of 45 degrees for 90 min (reproducing the change in orientation but not the magnitude of the gravitoinertial force on the centrifuge). Eye movements, including voluntary saccades directed along perceived earth- and head-referenced planes, were recorded before, during, and immediately after centrifugation. Postural center of pressure (COP) and multisegment body kinematics were also gathered before and within 10 min after centrifugation. Normal subjects overestimated roll-tilt during centrifugation and revealed errors in perception of head-vertical provided by directed saccades. Errors in this perceptual response tended to increase with time and became significant after approximately 30 min. Motion-sickness symptoms caused approximately 25% of normal subjects to limit their head movements during centrifugation and led three normal subjects to stop the test early. Immediately after centrifugation, subjects reported feeling tilted 10 degrees in the opposite direction, which was in agreement with the direction of their earth-referenced directed saccades. Postural COP, segmental body motion amplitude, and hip-sway frequency increased significantly after centrifugation. These postural effects were short-lived, however, with a recovery time of several postural test trials (minutes). There were also asymmetries in the direction of postcentrifugation COP and head tilt which depended on the subject's orientation during the centrifugation adaptation period (left ear or right ear out). The amount of total head movements during centrifugation correlated poorly or inversely with postcentrifugation postural stability, and the most unstable subject made no head movements. There was no decrease in postural stability after static tilt, although these subjects also reported a perceived tilt briefly after return to upright, and they also had COP asymmetries. Abnormal subjects underestimated roll-tilt during centrifugation, and their directed saccades revealed permanent spatial distortions. Bilateral abnormal subjects started out with poor postural control, but showed no postural decrements after centrifugation, while unilateral abnormal subjects had varying degrees of postural decrement, both in their everyday function and as a result of experiencing the centrifugation. In addition, three unilateral, abnormal subjects, who rode twice in opposite orientations, revealed a consistent orthogonal pattern of COP offsets after centrifugation. These results suggest that both orientation and magnitude of the gravitoinertial vector are used by the central nervous system for calibration of multiple orientation systems. A change in the background gravitoinertial force (otolith input) can rapidly initiate postural and perceptual adaptation in several sensorimotor systems, independent of a structured visual surround.
1. A simplified three-dimensional state space model of visual vestibular interaction was formulated. Matrix and dynamical system operators representing coupling from the semicircular canals and the visual system to the velocity storage integrator were incorporated into the model. 2. It was postulated 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 while another rotates the pitch and roll eigenvectors with the head, while maintaining the yaw axis eigenvector approximately spatially invariant. Using this representation, the response characteristics of the pitch, roll, and yaw eye velocity were obtained in terms of the eigenvalues and associated eigenvectors. 3. Using OKAN data obtained from monkeys and comparing to the model predictions, the eigenvalues and eigenvectors of the system matrix were identified as a function of tilt to the side or of tilt to the prone positions, using a modification of the Marquardt algorithm. The yaw eigenvector for right-side-down tilt and for downward pitch cross-coupling was approximately 30 degrees from the spatial vertical. For the prone position, the eigenvector was computed to be approximately 20 degrees relative to the spatial vertical. For both side-down and prone positions, oblique OKN induced along eigenvector directions generated OKAN which decayed to zero along a straight line with approximately a single time constant. This was verified by a spectral analysis of the residual sequence about the straight line fit to the decaying data. The residual sequence was associated with a narrow autocorrelation function and a wide power spectrum. 4. Parameters found using the Marquardt algorithm were incorporated into the model. Diagonal matrices in a head coordinate frame were introduced to represent the direct pathway and the coupling of the visual system to the integrator. Model simulations predicted the behavior of yaw and pitch OKN and OKAN when the animal was upright, as well as the cross-coupling in the tilted position. The trajectories in velocity space were also accurately simulated. 5. There were similarities between the monkey eigenvectors and human perception of the spatial vertical. For side-down tilts and downward eye velocity cross-coupling, there was only an Aubert (A) effect. For upward eye velocity cross-coupling there were both Muller (E) and Aubert (A) effects. The mean of the eigenvectors for upward and downward eye velocities overlay human 1 x g perceptual data.(ABSTRACT TRUNCATED AT 400 WORDS).
Error and control efficiency analysis of pilots exposed to simulated pitch, roll, yaw, and altitude variations
Two experiments were conducted. In Experiment I, each of eight Ss attempted to place himself at 13 different goal orientations between prone and supine. Deviations of achieved body pitch angles from goal orientations were determined. In Experiment II, each of eight Ss attempted to align a visual target with his morphological horizon while he was placed at each of the 13 goal orientations. Changes in settings of the target were examined. Results indicate that Ss underestimate body pitch when they are tilted less than 60 deg backward or forward from the vertical, overestimate body pitch when they are nearly prone, and accurately estimate body pitch when they are nearly supine. In contrast, Ss set the visual target maximally above the morphological horizon when they are tilted 30 deg forward from the vertical. The findings are discussed in terms of common and different physiological mechanism that may underlie judgments of these types.
A visual display rotating in a frontal plane induces effects equivalent to a change in the apparent direction of gravity. Magnitude of visual tilt was measured as a function of time from onset of rotation, velocity of rotation, and area and retinal location of the stimulating field. The mejor part of the tilt occurs within 30 sec from onset of stimulation. It increases with angular velocity, but independently of area and location of field, up to about 30 to 40 deg of rotation per sec and then levels off. Tilt increases with field size but the effect of thin ring-fields increases with retinal eccentricity. The interaction of visual and nonvisual determinants of the induced effects is discussed.
The compensation signals from somatosensory and vestibular receptors, which act to compensate for disturbances produced by the displacement of our eyes relative to other parts of our bodies and for visual disturbances such as tilted frames or moving visual fields, are investigated. Disturbances were evoked by tilting the head and by rotating a large visual display, while compensation signals related to gravity were altered by placing the subjects horizontally on a board or seating them vertically. The first experiment studied the effects of visual disturbance on the ability of supine observers to set a line to the longitudinal body axis while the head was tilted toward one shoulder or while the head was straight. Results showed that the effects of the visual disturbance were greater when the head was tilted than when it was straight, which indicates that the effects of visual disturbance were greater for a task that required compensation. The second experiment compared the performance of supine and erect observers. No differences were found between the performance of observers on a task requiring the use of compensation signals under these two conditions, which suggests that the enrichment of compensatory signals did not reduce the effects of visual disturbancs.
From a sensory point of view, space flight represents a form of 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. Appropriate countermeasures for long-duration flights will rely on preflight adaptation and in-flight training.
Artificial representations of virtual worlds are becoming more common due to advances in the technology of image generation and display systems. Application areas include flight simulation, mission rehearsal, teleoperator systems, and virtual reality systems. System developers should be forewarned that some proportion of users will experience perceptual anomalies and symptoms of motion sickness as a result of travel through virtual space.
Two part-task preflight adaptation trainers (PATs) are being developed at the NASA Johnson Space Center to preadapt astronauts to novel sensory stimulus conditions similar to those present in microgravity to facilitate adaptation to microgravity and readaptation to Earth. This activity is a major component of a general effort to develop countermeasures aimed at minimizing sensory and sensorimotor disturbances and Space Motion Sickness (SMS) associated with adaptation to microgravity and readaptation to Earth. Design principles for the development of the two trainers are discussed, along with a detailed description of both devices. In addition, a summary of four ground-based investigations using one of the trainers to determine the extent to which various novel sensory stimulus conditions produce changes in compensatory eye movement responses, postural equilibrium, motion sickness symptoms, and electrogastric responses are presented. Finally, a brief description of the general concept of dual-adopted states that underly the development of the PATs, and ongoing and future operational and basic research activities are presented.
Eight chronic vestibular deficient (VD) patients (bilateral N = 4, unilateral N = 4, ages 18-67 were exposed to an interaural centripetal acceleration of 1 G (resultant 45 degree roll tilt of 1.4 G) on a 0.8 meter radius centrifuge for up to 90 minutes in the dark. The patients sat with head fixed upright, except every 4 of 10 minutes when instructed to point their nose and eyes towards a visual target (switched on every 3 to 5 seconds at random places within plus or minus 30 deg) in the Earth horizontal plane. Eye movements, including directed saccades for subjective Earth-and head-referenced planes, were recorded before, during, and after centrifugation using electro-oculography. Postural sway was measured before and within ten minutes after centrifugation using a sway-referenced or earth-fixed support surface, and with or without a head movement sequence. The protocol was selected for each patient based on the most challenging condition in which the patient was able to maintain balance with eyes closed.