ALBERT

Description: 1. Reaching movements made in a rotating room generate Coriolis forces that are directly proportional to the cross product of the room's angular velocity and the arm's linear velocity. Such Coriolis forces are inertial forces not involving mechanical contact with the arm. 2. We measured the trajectories of arm movements made in darkness to a visual target that was extinguished at the onset of each reach. Prerotation subjects pointed with both the right and left arms in alternating sets of eight movements. During rotation at 10 rpm, the subjects reached only with the right arm. Postrotation, the subjects pointed with the left and right arms, starting with the left, in alternating sets of eight movements. 3. The initial perrotary reaching movements of the right arm were highly deviated both in movement path and endpoint relative to the prerotation reaches of the right arm. With additional movements, subjects rapidly regained straight movement paths and accurate endpoints despite the absence of visual or tactile feedback about reaching accuracy. The initial postrotation reaches of the left arm followed straight paths to the wrong endpoint. The initial postrotation reaches of the right arm had paths with mirror image curvature to the initial perrotation reaches of the right arm but went to the correct endpoint. 4. These observations are inconsistent with current equilibrium point models of movement control. Such theories predict accurate reaches under our experimental conditions. Our observations further show independent implementation of movement and posture, as evidenced by transfer of endpoint adaptation to the nonexposed arm without transfer of path adaptation. Endpoint control may occur at a relatively central stage that represents general constraints such as gravitoinertial force background or egocentric direction relative to both arms, and control of path may occur at a more peripheral stage that represents moments of inertia and muscle dynamics unique to each limb. 5. Endpoint and path adaptation occur despite the absence both of mechanical contact cues about the perturbing force and visual or tactile cues about movement accuracy. These findings point to the importance of muscle spindle signals, monitoring of motor commands, and possibly joint and tendon receptors in a detailed trajectory monitoring process. Muscle spindle primary and secondary afferent signals may differentially influence adaptation of movement shape and endpoint, respectively.

Description: 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.

Description: 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.

Description: 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. Copyright 2000 Wiley-Liss, Inc.

Description: 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.

Description: In an earlier study, the authors found that severely motion sick individuals could be greatly relieved of their symptoms by intramuscular injections of promethazine (50 mg) or scopolamine (.5 mg). Comparable 50-mg injections of promethazine also have been found effective in alleviating symptoms of space motion sickness. The concern has risen, however, that such drugs may delay or retard the acquisition of adaptation to stressful environments. In the current study, we controlled arousal using a mental arithmetic task and precisely equated the exposure history (number of head movements during rotation) of a placebo, control group and an experimental group who had received promethazine. No differences in total adaptation or in rates of adaptation were present between the two groups. Another experimental group also received promethazine and was allowed to make as many head movements as they could, before reaching nausea, up to 800. This group showed a greater level of adaptation than the placebo group. These results suggest a strategy for dealing with space motion sickness that is described.

Description: Microgravity provides unique, though experimentally challenging, opportunities to study motor control. A traditional research focus has been the effects of linear acceleration on vestibular responses to angular acceleration. Evidence is accumulating that the high-frequency vestibulo-ocular reflex (VOR) is not affected by transitions from a 1 g linear force field to microgravity (〈1 g); however, it appears that the three-dimensional organization of the VOR is dependent on gravitoinertial force levels. Some of the observed effects of microgravity on head and arm movement control appear to depend on the previously undetected inputs of cervical and brachial proprioception, which change almost immediately in response to alterations in background force levels. Recent studies of post-flight disturbances of posture and locomotion are revealing sensorimotor mechanisms that adjust over periods ranging from hours to weeks.

Description: 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.

Description: 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.

Description: 1. Forward reaching movements made during body rotation generate tangential Coriolis forces that are proportional to the cross product of the angular velocity of rotation and the linear velocity of the arm. Coriolis forces are inertial forces that do not involve mechanical contact. Virtually no constant centrifugal forces will be present in the background when motion of the arm generates transient Coriolis forces if the radius of body rotation is small. 2. We measured the trajectories of arm movements made in darkness to a visual target that was extinguished as movement began. The reaching movements were made prerotation, during rotation at 10 rpm in a fully enclosed rotating room, and postrotation. During testing the subject was seated at the center of the room and pointed radially. Neither visual nor tactile feedback about movement accuracy was present. 3. In experiment 1, subjects reached at a fast or slow rate and their hands made contact with a horizontal surface at the end of the reach. Their initial perrotary movements were highly significantly deviated relative to prerotation in both trajectories and end-points in the direction of the transient Coriolis forces that had been generated during the reaches. Despite the absence of visual and tactile feedback about reaching accuracy, all subjects rapidly regained straight movement trajectories and accurate endpoints. Postrotation, transient errors of opposite sign were present for both trajectories and endpoints. 4. In a second experiment the conditions were identical except that subjects pointed just above the location of the extinguished target so that no surface contact was involved. All subjects showed significant initial perrotation deviations of trajectories and endpoints in the direction of the transient Coriolis forces. With repeated reaches the trajectories, as viewed from above, again became straight, but there was only partial restoration of endpoint accuracy, so that subjects reached in a straight line to the wrong place. Aftereffects of opposite sign were transiently present in the postrotary movements. 5. These observations fail to support current equilibrium point models, both alpha and lambda, of movement control. Such theories would not predict endpoint errors under our experimental conditions, in which the Coriolis force is absent at the beginning and end of a movement. Our results indicate that detailed aspects of movement trajectory are being continuously monitored on the basis of proprioceptive feedback in relation to motor commands. Adaptive compensations can be initiated after one perturbation despite the absence of either visual or tactile feedback about movement trajectory and endpoint error. Moreover, movement trajectory and end-point can be remapped independently.(ABSTRACT TRUNCATED AT 400 WORDS).