Virtual reality as a countermeasure for astronaut motion sickness during simulated post-flight water landings.

Entry motion sickness (EMS) affects crewmembers upon return to Earth following extended adaptation to microgravity. Anticholinergic pharmaceuticals (e.g., Meclizine) are often taken prior to landing; however, they have operationally adverse side effects (e.g., drowsiness). There is a need to develop non-pharmaceutical countermeasures to EMS. We assessed the efficacy of a technological countermeasure providing external visual cues following splashdown, where otherwise only nauseogenic internal cabin visual references are available. Our countermeasure provided motion-congruent visual cues of an Earth-fixed scene in virtual reality, which was compared to a control condition with a head-fixed fixation point in virtual reality in a between-subject design with 15 subjects in each group. We tested the countermeasure's effectiveness at mitigating motion sickness symptoms at the end of a ground-based reentry analog: approximately 1 h of 2Gx centrifugation followed by up to 1 h of wave-like motion. Secondarily, we explored differences in vestibular-mediated balance performance between the two conditions. While Motion Sickness Questionnaire outcomes did not differ detectably between groups, we found significantly better survival rates (with dropout dictated by reporting moderate nausea consecutively over 2 min) in the visual countermeasure group than the control group (79% survival vs. 33%, t(14) = 2.50, p = 0.027). Following the reentry analogs, subjects demonstrated significantly higher sway prior to recovery (p = 0.0004), which did not differ between control and countermeasure groups. These results imply that providing motion-congruent visual cues may be an effective mean for curbing the development of moderate nausea and increasing comfort following future space missions.

The oculogyral illusion: retinal and oculomotor factors.

Subjects in a dark chamber exposed to angular acceleration while viewing a head-fixed target experience motion and displacement of the target relative to their body. Competing explanations of this phenomenon, known as the oculogyral illusion, have attributed it to the suppression of the vestibulo-ocular reflex (VOR) or to retinal slip. In the dark, the VOR evokes compensatory eye movements in the direction opposite to body acceleration. A head-fixed visual target will tend to suppress these eye movements. The VOR suppression hypothesis attributes the oculogyral illusion to the signals that prevent reflexive deviation of the eyes from the target thus resulting in apparent target displacement in the direction of acceleration. The retinal slip hypothesis attributes the illusion to inadequate fixation of the target with the eyes being involuntarily deviated in the direction opposite acceleration, the retinal slip being interpreted as target displacement in the direction of acceleration. Another possibility is that the illusion could arise from a change in the representation of the perceived head midline. To evaluate these three alternative hypotheses, we tested 8 subjects at 4 acceleration rates (2, 10, 20, 30°/s²) in each of three conditions: (a) fixate and point to a target light; (b) fixate to the target light and point to the head midline; (c) look straight ahead in the dark. The displacement magnitude of the oculogyral illusion was least at 2°/s² ≈ 2° and was ≈10° at the other acceleration rates. The presence of the target light significantly attenuated eye movements relative to the dark condition, but eye movements were still present at the 10, 20, and 30°/s² accelerations. The eye velocity profiles in the dark at different acceleration rates did not show a one-to-one inverse mapping to the magnitude of the oculogyral illusion at those rates. The perceived head midline was not significantly displaced at any of the acceleration rates. The oculogyral illusion thus has at least two contributing factors: the suppression of nystagmus at low acceleration rates and at higher acceleration rates, a partial suppression coupled with an integration of the drift of the eyes with respect to the fixation target.

Vertical frames of reference and control of body orientation.

The present paper aims at critically reviewing the most outstanding and recent studies regarding the control of body orientation in the vertical space. A first part defines the general concepts used throughout this manuscript. The second part investigates the vertical perception and the main factors which affect it, while trying to overcome the five areas of theoretical and experimental controversies we have identified in the literature. The third part of this review presents the different theoretical models of the vertical perception and body orientation in space. Finally, the last part focuses on the functional coupling between perception of the vertical and orientation of the body in space. It considers more particularly how these two dimensions interact for explaining the observed behaviors.

Influence of gravitoinertial force level on the subjective vertical during recumbent yaw axis body tilt.

We tilted recumbent subjects at various angles about their yaw (foot to head) axis and had them indicate the direction of their subjective vertical and apparent head midline about the same axis. One set of tests was conducted during parabolic flight maneuvers where the background gravitoinertial acceleration varied from 0 to 1.8g. The blindfolded subjects (n = 6) were tested supine and at tilts of 60 degrees and 30 degrees left and right about their horizontal long body axis. They used a gravity neutral joystick to indicate their subjective vertical or their head midline continuously from the high force through the 0g portions of parabolas. In 0g, all subjects felt supine and oriented the joystick perpendicular to their body when indicating the subjective vertical. This points to strong influences of the symmetric somatic touch and pressure cues from the apparatus on orientation when the otolith organs are unloaded. In contrast to the settings in 0g, settings of the subjective vertical in 1g and 1.8g varied as a function of body orientation. However, the settings did not differ between 1g and 1.8g test conditions. Subjective vertical judgments were also made by subjects (n = 11) in the Brandeis slow rotation room, with the room stationary and rotating at a speed that produced a 2g resultant of gravitational and centrifugal acceleration. There were no differences between settings of the subjective vertical made in 1g and 2g. The similarity of 1g and hyper-g settings during recumbent yaw tilts, both in parabolic flight and in the rotating room, contrasts with the previously observed, strong influence which force levels above 1g have on settings of the subjective vertical during tilt of the body in pitch or roll. The findings for all three axes are consistent with a recently developed model of static spatial orientation.

Mechanisms of human static spatial orientation.

We have developed a tri-axial model of spatial orientation applicable to static 1g and non-1g environments. The model attempts to capture the mechanics of otolith organ transduction of static linear forces and the perceptual computations performed on these sensor signals to yield subjective orientation of the vertical direction relative to the head. Our model differs from other treatments that involve computing the gravitoinertial force (GIF) vector in three independent dimensions. The perceptual component of our model embodies the idea that the central nervous system processes utricular and saccular stimuli as if they were produced by a GIF vector equal to 1g, even when it differs in magnitude, because in the course of evolution living creatures have always experienced gravity as a constant. We determine just two independent angles of head orientation relative to the vertical that are GIF dependent, the third angle being derived from the first two and being GIF independent. Somatosensory stimulation is used to resolve our vestibular model's ambiguity of the up-down directions. Our otolith mechanical model takes into account recently established non-linear behavior of the force-displacement relationship of the otoconia, and possible otoconial deflections that are not co-linear with the direction of the input force (cross-talk). The free parameters of our model relate entirely to the mechanical otolith model. They were determined by fitting the integrated mechanical/perceptual model to subjective indications of the vertical obtained during pitch and roll body tilts in 1g and 2g force backgrounds and during recumbent yaw tilts in 1g. The complete data set was fit with very little residual error. A novel prediction of the model is that background force magnitude either lower or higher than 1g will not affect subjective vertical judgments during recumbent yaw tilt. These predictions have been confirmed in recent parabolic flight experiments.

Vertical linear self-motion perception during visual and inertial motion: more than weighted summation of sensory inputs.

We evaluated visual and vestibular contributions to vertical self motion perception by exposing subjects to various combinations of 0.2 Hz vertical linear oscillation and visual scene motion. The visual stimuli presented via a head-mounted display consisted of video recordings of the test chamber from the perspective of the subject seated in the oscillator. In the dark, subjects accurately reported the amplitude of vertical linear oscillation with only a slight tendency to underestimate it. In the absence of inertial motion, even low amplitude oscillatory visual motion induced the perception of vertical self-oscillation. When visual and vestibular stimulation were combined, self-motion perception persisted in the presence of large visual-vestibular discordances. A dynamic visual input with magnitude discrepancies tended to dominate the resulting apparent self-motion, but vestibular effects were also evident. With visual and vestibular stimulation either spatially or temporally out-of-phase with one another, the input that dominated depended on their amplitudes. High amplitude visual scene motion was almost completely dominant for the levels tested. These findings are inconsistent with self-motion perception being determined by simple weighted summation of visual and vestibular inputs and constitute evidence against sensory conflict models. They indicate that when the presented visual scene is an accurate representation of the physical test environment, it dominates over vestibular inputs in determining apparent spatial position relative to external space.

Analysis of human postural responses to recoverable falls.

We studied the kinematics and kinetics of human postural responses to "recoverable falls." To induce brief falling we used a Hold and Release (H&R) paradigm. Standing subjects actively resisted a force applied to their sternum. When this force was quickly released they were suddenly off balance. For a brief period, approximately 125 ms, until restoring forces were generated to shift the center of foot pressure in front of the center of mass, the body was in a forward fall acted on by gravity and ground support forces. We were able to describe the whole-body postural behavior following release using a multilink inverted pendulum model in a regime of "small oscillations." A three-segment model incorporating upper body, upper leg, and lower leg, with active stiffness and damping at the joints was fully adequate to fit the kinematic data from all conditions. The significance of our findings is that in situations involving recoverable falls or loss of balance the earliest responses are likely dependent on actively-tuned, reflexive mechanisms yielding stiffness and damping modulation of the joints. We demonstrate that haptic cues from index fingertip contact with a stationary surface lead to a significantly smaller angular displacement of the torso and a more rapid recovery of balance. Our H&R paradigm and associated model provide a quantifiable approach to studying recovery from potential falling in normal and clinical subjects.

Stabilization of posture by precision touch of the index finger with rigid and flexible filaments.

Light touch of the index finger with a stationary surface at non-mechanically supportive force levels (<100 g) greatly attenuates the body sway of standing subjects. In three experiments, we evaluated the properties of finger contact and of the contacted object necessary to produce postural stabilization in subjects standing heel-to-toe with eyes closed, as well as how accurately hand position can be controlled. Experiment 1 involved finger contact with flexible filaments of different bending strengths, a flat surface, and an imagined spatial position. Contact with the flat surface was most effective in attenuating sway; the flexible filaments were much less effective but still significantly better than imagined contact. Experiment 2 compared the effectiveness of finger contact with a flexible filament, a rigid filament of the same diameter, a flat surface, and an imagined spatial position. The rigid filament and flat surface conditions were equally effective in attenuating body sway and were greatly superior to contact with the flexible filament, which was superior to imagined contact. Experiment 3 included five conditions: arms by sides; finger "contact" with an imagined spatial position; finger contact with a flat surface; finger contact with a flexible filament attempting to maintain it bent; and contact with the flexible filament attempting not to bend it. The arms by sides and finger "contact" with an imagined position conditions did not differ significantly; all three conditions involving actual finger contact showed significantly less center of pressure and hand sway, but contact with the flat surface was most effective in attenuating both postural and hand displacement. In all three experiments, the level of force applied in fingertip contact conditions was far below that necessary to provide mechanical stabilization. Our findings indicate that: (1) stimulation of a small number of receptors in the fingertip is adequate to allow stabilization of sway, (2) fingertip force levels as low as 5-10 g provide some stabilization, (3) contact with a stationary spatial referent is most effective, and (4) independent control of arm and torso occurs when finger contact is allowed.

Gravitoinertial force magnitude and direction influence head-centric auditory localization.

We measured the influence of gravitoinertial force (GIF) magnitude and direction on head-centric auditory localization to determine whether a true audiogravic illusion exists. In experiment 1, supine subjects adjusted computer-generated dichotic stimuli until they heard a fused sound straight ahead in the midsagittal plane of the head under a variety of GIF conditions generated in a slow-rotation room. The dichotic stimuli were constructed by convolving broadband noise with head-related transfer function pairs that model the acoustic filtering at the listener's ears. These stimuli give rise to the perception of externally localized sounds. When the GIF was increased from 1 to 2 g and rotated 60 degrees rightward relative to the head and body, subjects on average set an acoustic stimulus 7.3 degrees right of their head's median plane to hear it as straight ahead. When the GIF was doubled and rotated 60 degrees leftward, subjects set the sound 6.8 degrees leftward of baseline values to hear it as centered. In experiment 2, increasing the GIF in the median plane of the supine body to 2 g did not influence auditory localization. In experiment 3, tilts up to 75 degrees of the supine body relative to the normal 1 g GIF led to small shifts, 1--2 degrees, of auditory setting toward the up ear to maintain a head-centered sound localization. These results show that head-centric auditory localization is affected by azimuthal rotation and increase in magnitude of the GIF and demonstrate that an audiogravic illusion exists. Sound localization is shifted in the direction opposite GIF rotation by an amount related to the magnitude of the GIF and its angular deviation relative to the median plane.

Coriolis-force-induced trajectory and endpoint deviations in the reaching movements of labyrinthine-defective subjects.

When reaching movements are made during passive constant velocity body rotation, inertial Coriolis accelerations are generated that displace both movement paths and endpoints in their direction. These findings directly contradict equilibrium point theories of movement control. However, it has been argued that these movement errors relate to subjects sensing their body rotation through continuing vestibular activity and making corrective movements. In the present study, we evaluated the reaching movements of five labyrinthine-defective subjects (lacking both semicircular canal and otolith function) who cannot sense passive body rotation in the dark and five age-matched, normal control subjects. Each pointed 40 times in complete darkness to the location of a just extinguished visual target before, during, and after constant velocity rotation at 10 rpm in the center of a fully enclosed slow rotation room. All subjects, including the normal controls, always felt completely stationary when making their movements. During rotation, both groups initially showed large deviations of their movement paths and endpoints in the direction of the transient Coriolis forces generated by their movements. With additional per-rotation movements, both groups showed complete adaptation of movement curvature (restoration of straight-line reaches) during rotation. The labyrinthine-defective subjects, however, failed to regain fully accurate movement endpoints after 40 reaches, unlike the control subjects who did so within 11 reaches. Postrotation, both groups' movements initially had mirror image curvatures to their initial per-rotation reaches; the endpoint aftereffects were significantly different from prerotation baseline for the control subjects but not for the labyrinthine-defective subjects reflecting the smaller amount of endpoint adaptation they achieved during rotation. The labyrinthine-defective subjects' movements had significantly lower peak velocity, higher peak elevation, lower terminal velocity, and a more vertical touchdown than those of the control subjects. Thus the way their reaches terminated denied them the somatosensory contact cues necessary for full endpoint adaptation. These findings fully contradict equilibrium point theories of movement control. They emphasize the importance of contact cues in adaptive movement control and indicate that movement errors generated by Coriolis perturbations of limb movements reveal characteristics of motor planning and adaptation in both healthy and clinical populations.

Fingertip contact suppresses the destabilizing influence of leg muscle vibration.

Touch of the hand with a stationary surface at nonmechanically supportive force levels (<1 N) greatly attenuates postural sway during quiet stance. We predicted such haptic contact would also suppress the postural destabilization caused by vibrating the right peroneus brevis and longus muscles of subjects standing heel-to-toe with eyes closed. In experiment 1, ten subjects were tested under four conditions: no-vibration, no-touch; no-vibration, touch; vibration, no-touch; and vibration, touch. A hand-held physiotherapy vibrator (120 Hz) was applied approximately 5 cm above the malleolous to stimulate the peroneus longus and brevis tendons. Touch conditions involved contact of the right index finger with a laterally positioned surface (<1 N of force) at waist height. Vibration in the absence of finger contact greatly increased the mean sway amplitude of the center of pressure and of the head relative to the no-vibration, no-touch control condition (P < 0.001). The touch, no-vibration and touch-vibration conditions were not significantly different (P > 0.05) from each other and both had significantly less mean sway amplitude of head and of center of pressure than the other conditions (P < 0.01). In experiment 2, eight subjects stood heel-to-toe under touch and no-touch conditions involving 40-s duration trials of peroneus tendon vibration at different duty cycles: 1-, 2-, 3-, and 4-s ON and OFF periods. The vibrator was attached to the subject's leg and remotely activated. In the no-touch conditions, subjects showed periodic postural disruptions contingent on the duty cycle and mirror image rebounds with the offset of vibration. In the touch conditions, subjects were much less disrupted and showed compensations occurring within 500 ms of vibration onset and mirror image rebounds with vibration offset. Subjects were able to suppress almost completely the destabilizing influence of the vibration in the 3- and 4-s duty cycle trials. These experiments show that haptic contact of the hand with a stable surface can suppress abnormal proprioceptive and motor signals in leg muscles.

Artificial gravity as a countermeasure in long-duration space flight.

Long-duration exposure to weightlessness results in bone demineralization, muscle atrophy, cardiovascular deconditioning, altered sensory-motor control, and central nervous system reorganizations. Exercise countermeasures and body loading methods so far employed have failed to prevent these changes. A human mission to Mars might last 2 or 3 years and without effective countermeasures could result in dangerous levels of bone and muscle loss. Artificial gravity generated by rotation of an entire space vehicle or of an inner chamber could be used to prevent structural changes. Some of the physical characteristics of rotating environments are outlined along with their implications for human performance. Artificial gravity is the centripetal force generated in a rotating vehicle and is proportional to the product of the square of angular velocity and the radius of rotation. Thus, for a particular g-level, there is a tradeoff between velocity of rotation and radius. Increased radius is vastly more expensive to achieve than velocity, so it is important to know the highest rotation rates to which humans can adapt. Early studies suggested that 3 rpm might be the upper limit because movement control and orientation were disrupted at higher velocities and motion sickness and chronic fatigue were persistent problems. Recent studies, however, are showing that, if the terminal velocity is achieved over a series of gradual steps and many body movements are made at each dwell velocity, then full adaptation of head, arm, and leg movements is possible. Rotation rates as high as 7.5-10 rpm are likely feasible. An important feature of the new studies is that they provide compelling evidence that equilibrium point theories of movement control are inadequate. The central principles of equilibrium point theories lead to the equifinality prediction, which is violated by movements made in rotating reference frames.

Congenitally blind individuals rapidly adapt to coriolis force perturbations of their reaching movements.

Reaching movements made to visual targets in a rotating room are initially deviated in path and endpoint in the direction of transient Coriolis forces generated by the motion of the arm relative to the rotating environment. With additional reaches, movements become progressively straighter and more accurate. Such adaptation can occur even in the absence of visual feedback about movement progression or terminus. Here we examined whether congenitally blind and sighted subjects without visual feedback would demonstrate adaptation to Coriolis forces when they pointed to a haptically specified target location. Subjects were tested pre-, per-, and postrotation at 10 rpm counterclockwise. Reaching to straight ahead targets prerotation, both groups exhibited slightly curved paths. Per-rotation, both groups showed large initial deviations of movement path and curvature but within 12 reaches on average had returned to prerotation curvature levels and endpoints. Postrotation, both groups showed mirror image patterns of curvature and endpoint to the per-rotation pattern. The groups did not differ significantly on any of the performance measures. These results provide compelling evidence that motor adaptation to Coriolis perturbations can be achieved on the basis of proprioceptive, somatosensory, and motor information in the complete absence of visual experience.

Reaching during virtual rotation: context specific compensations for expected coriolis forces.

Subjects who are in an enclosed chamber rotating at constant velocity feel physically stationary but make errors when pointing to targets. Reaching paths and endpoints are deviated in the direction of the transient inertial Coriolis forces generated by their arm movements. By contrast, reaching movements made during natural, voluntary torso rotation seem to be accurate, and subjects are unaware of the Coriolis forces generated by their movements. This pattern suggests that the motor plan for reaching movements uses a representation of body motion to prepare compensations for impending self-generated accelerative loads on the arm. If so, stationary subjects who are experiencing illusory self-rotation should make reaching errors when pointing to a target. These errors should be in the direction opposite the Coriolis accelerations their arm movements would generate if they were actually rotating. To determine whether such compensations exist, we had subjects in four experiments make visually open-loop reaches to targets while they were experiencing compelling illusory self-rotation and displacement induced by rotation of a complex, natural visual scene. The paths and endpoints of their initial reaching movements were significantly displaced leftward during counterclockwise illusory rotary displacement and rightward during clockwise illusory self-displacement. Subjects reached in a curvilinear path to the wrong place. These reaching errors were opposite in direction to the Coriolis forces that would have been generated by their arm movements during actual torso rotation. The magnitude of path curvature and endpoint errors increased as the speed of illusory self-rotation increased. In successive reaches, movement paths became straighter and endpoints more accurate despite the absence of visual error feedback or tactile feedback about target location. When subjects were again presented a stationary scene, their initial reaches were indistinguishable from pre-exposure baseline, indicating a total absence of aftereffects. These experiments demonstrate that the nervous system automatically compensates in a context-specific fashion for the Coriolis forces associated with reaching movements.

Aspects of body self-calibration.

The representation of body orientation and configuration is dependent on multiple sources of afferent and efferent information about ongoing and intended patterns of movement and posture. Under normal terrestrial conditions, we feel virtually weightless and we do not perceive the actual forces associated with movement and support of our body. It is during exposure to unusual forces and patterns of sensory feedback during locomotion that computations and mechanisms underlying the ongoing calibration of our body dimensions and movements are revealed. This review discusses the normal mechanisms of our position sense and calibration of our kinaesthetic, visual and auditory sensory systems, and then explores the adaptations that take place to transient Coriolis forces generated during passive body rotation. The latter are very rapid adaptations that allow body movements to become accurate again, even in the absence of visual feedback. Muscle spindle activity interpreted in relation to motor commands and internally modeled reafference is an important component in permitting this adaptation. During voluntary rotary movements of the body, the central nervous system automatically compensates for the Coriolis forces generated by limb movements. This allows accurate control to be maintained without our perceiving the forces generated.

Human orientation and movement control in weightless and artificial gravity environments.

Our goal is to summarize what has been learned from studies of human movement and orientation control in weightless conditions. An understanding of the physics of weightlessness is essential to appreciate the dramatic consequences of the absence of continuous contact forces on orientation and posture. Eye, head, arm, leg, and whole body movements are discussed, but only experiments whose results seem relatively incontrovertible are included. Emphasis is placed on distinguishing between virtually immediate adaptive compensations to weightlessness and those with longer time courses. The limitations and difficulties of performing experiments in weightless conditions are highlighted. We stress that when astronauts and cosmonauts return from extended space flight they do so with both physical "plant" and neural "controller" structurally and functionally altered. Recent developments in adapting humans to artificial gravity conditions are discussed as a way of maintaining sensory-motor and structural integrity in extended missions involving transitions between different force environments.

Haptic stabilization of posture: changes in arm proprioception and cutaneous feedback for different arm orientations.

Postural sway during quiet stance is attenuated by actively maintained contact of the index finger with a stationary surface, even if the level of applied force (<1 N) cannot provide mechanical stabilization. In this situation, changes in force level at the fingertip lead changes in center of foot pressure by approximately 250 ms. These and related findings indicate that stimulation of the fingertip combined with proprioceptive information about the hand and arm can serve as an active sensor of body position relative to the point of contact. A geometric analysis of the relationship between hand and torso displacement during body sway led to the prediction that arm and hand proprioceptive and finger somatosensory information about body sway would be maximized with finger contact in the plane of body sway. Therefore, the most postural stabilization should be possible with such contact. To test this analysis, subjects touched a laterally versus anteriorly placed surface while in each of two stances: the heel-to-toe tandem Romberg stance that reduces medial-lateral stability and the heel-to-heel, toes-outward, knees-bent, "duck stance" that reduces fore-aft stability. Postural sway was always least with finger contact in the unstable plane: for the tandem stance, lateral fingertip contact was significantly more effective than frontal contact, and, for the duck stance, frontal contact was more effective than lateral fingertip contact. Force changes at the fingertip led changes in center of pressure of the feet by approximately 250 ms for both fingertip contact locations for both test stances. These results support the geometric analysis, which showed that 1) arm joint angles change by the largest amount when fingertip contact is maintained in the plane of greatest sway, and 2) the somatosensory cues at the fingertip provide both direction and amplitude information about sway when the finger is contacting a surface in the unstable plane.

Precision contact of the fingertip reduces postural sway of individuals with bilateral vestibular loss.

Contact of the hand with a stationary surface attenuates postural sway in normal individuals even when the level of force applied is mechanically inadequate to dampen body motion. We studied whether subjects without vestibular function would be able to substitute contact cues from the hand for their lost labyrinthine function and be able to balance as well as normal subjects in the dark without finger contact. We also studied the relative contribution of sight of the test chamber to the two groups. Subjects attempted to maintain a tandem Romberg stance for 25 s under three levels of fingertip contact: no contact; light-touch contact, up to 1 N (approximately 100 g) force; and unrestricted contact force. Both eyes open and eyes closed conditions were evaluated. Without contact, none of the vestibular loss subjects could stand for more than a few seconds in the dark without falling; all the normals could. The vestibular loss subjects were significantly more stable in the dark with light touch of the index finger than the normal subjects in the dark without touch. They also swayed less in the dark with light touch than when permitted sight of the test chamber without touch, and less with sight and touch than just sight. The normal subjects swayed less in the dark with touch than without, and less with sight and touch than sight alone. These findings show that during quiet stance light touch of the index finger with a stationary surface can be as effective or even more so than vestibular function for minimizing postural sway.

Adaptation in a rotating artificial gravity environment.

The centripetal force generated by a rotating space vehicle is a potential source of artificial gravity. Minimizing the cost of such a vehicle dictates using the smallest radius and highest rotation rate possible, but head movements made at high rotation rates generate disorienting, nauseogenic cross-coupled semicircular canal stimulation. Early studies suggested 3 or 4 rpm as the highest rate at which humans could adapt to this vestibular stimulus. These studies neglected the concomitant Coriolis force actions on the head/neck system. We assessed non-vestibular Coriolis effects by measuring arm and leg movements made in the center of a rotating room turning at 10 rpm and found that movement endpoints and trajectories are initially deviated; however, subjects readily adapt with 10-20 additional movements, even without seeing their errors. Equilibrium point theories of motor control errantly predict that Coriolis forces will not cause movement endpoint errors so that subjects will not have to adapt their reaching movements during rotation. Adaptation of movement trajectory acquired during Coriolis force perturbations of one arm transfers to the unexposed arm but there is no intermanual transfer of endpoint adaptation indicating that neuromotor representations of movement endpoint and trajectory are separable and can adapt independently, also contradictory to equilibrium point theories. Touching a surface at the end of reaching movements is required for complete endpoint adaptation in darkness but trajectory adapts completely with or without terminal contact. We have also made the first kinematic measurements of unconstrained head movements during rotation, these movements show rapid adaptation to Coriolis force perturbations. Our results point to methods for achieving full compensation for rotation up to 10 rpm.

Gravitoinertial force background level affects adaptation to coriolis force perturbations of reaching movements.

We evaluated the combined effects on reaching movements of the transient, movement-dependent Coriolis forces and the static centrifugal forces generated in a rotating environment. Specifically, we assessed the effects of comparable Coriolis force perturbations in different static force backgrounds. Two groups of subjects made reaching movements toward a just-extinguished visual target before rotation began, during 10 rpm counterclockwise rotation, and after rotation ceased. One group was seated on the axis of rotation, the other 2.23 m away. The resultant of gravity and centrifugal force on the hand was 1.0 g for the on-center group during 10 rpm rotation, and 1.031 g for the off-center group because of the 0.25 g centrifugal force present. For both groups, rightward Coriolis forces, approximately 0.2 g peak, were generated during voluntary arm movements. The endpoints and paths of the initial per-rotation movements were deviated rightward for both groups by comparable amounts. Within 10 subsequent reaches, the on-center group regained baseline accuracy and straight-line paths; however, even after 40 movements the off-center group had not resumed baseline endpoint accuracy. Mirror-image aftereffects occurred when rotation stopped. These findings demonstrate that manual control is disrupted by transient Coriolis force perturbations and that adaptation can occur even in the absence of visual feedback. An increase, even a small one, in background force level above normal gravity does not affect the size of the reaching errors induced by Coriolis forces nor does it affect the rate of reacquiring straight reaching paths; however, it does hinder restoration of reaching accuracy.