Hand trajectories of vertical arm movements in one-G and zero-G environments Evidence for a central representation of gravitational force

The purpose of the present experiment was to study the way in which the central nervous system (CNS), represents gravitational force during vertical arm pointing movements. Movements in upward (against gravity) and downward (with gravity) directions, with two different mass loads (hand empty and with a hand-held 0.5-kg weight) were executed by eight subjects in a normal gravitational environment. Movements by two cosmonauts, in the two directions, were also tested in a state of weightlessness. Analyses focused upon finger trajectories in the saggital plane. Subjects in a normal gravitational environment showed curved paths for both directions and weight conditions. In addition, downward movements showed significantly smaller curvatures than upward movements. Movement times were approximately the same for all the experimental conditions. Curvature differences between upward and downward movements persisted during space flight and immediately postflight. Movement times from both cosmonauts increased slightly during flight, but returned to normal immediately on reentry in a one-G environment. Results from the present study provide evidence that gravity is centrally represented in an anticipatory fashion as a driving force during vertical arm movement planning.     

The curvature of human arm movements in the absence of visual experience

It has been suggested that the spatial path of the hand is an important controlled feature of normal human arm movements and that the desired path is a straight line through external space. Recent experiments have suggested that distortions in visual perception of external space may lead to errors in its representation and thus influence the curvature of movements. The movements of blind and normal blind-folded subjects were therefore compared in a task requiring point-to-point hand movements in six directions across a horizontal worktop. Movement curvature varied with direction in both groups but was significantly higher for the blindfolded control subjects. Thus, the normals' distorted visual experience of straight lines in some orientations may lead them to make curved movement paths. The perception of curvature was also tested in the two groups in a task in which they traced the curved edge of a ruler. The blind group were slightly better at this task, although the difference was not significant. We conclude that visual experience influences point-to-point hand movements, leading to higher curvature for movements made in the fronto-parallel plane by sighted subjects due to visual distortions. These data therefore support the hypothesis that the spatial path followed by the hand is influenced by sensory inputs and is a controlled feature of human reaching movements. The data argue against the hypothesis that movement curvature is a result of optimising only the dynamics of the limb control.     

Egocentric references and human spatial orientation in microgravity

This article describes the results of the ellipses experiment conducted during the second French-Soviet spaceflight (project Aragatz). The realization of oriented motor tasks, on the basis of internal body representation and without visual feedback, was chosen as a paradigm for studying the determinants of spatial orientation under weightlessness. The process of drawing ellipses in the air, using arm movements with axes parallel or perpendicular to the longitudinal body axis, was studied under normal gravity and in weightlessness, and recorded using a video computer motion-analyzing system (Kinesigraph). On Earth, the experiments were performed in standing and lying positions, and in flight, in the erect position with the feet fixed to the floor. In general, performance of the task in microgravity was not disturbed. Under conditions of spaceflight, the longitudinal ellipse was inclined forward in accordance with the inclination of the whole body relative to the fixed feet. On Earth, the angle between the long axes of longitudinal and transverse ellipses deviated from 90° by 20–30°. The same deviation persisted under microgravity conditions. The distinctive features of ellipses traced by individual subjects were also preserved. It is concluded that an egocentric reference system ensures normal performance of sensorimotor tasks in the absence of a gravitational reference.     

Maximizing information from space data resources: a case for expanding integration across research disciplines

Regulatory systems are affected in space by exposure to weightlessness, high-energy radiation or other spaceflight-induced changes. The impact of spaceflight occurs across multiple scales and systems. Exploring such interactions and interdependencies via an integrative approach provides new opportunities for elucidating these complex responses. This paper argues the case for increased emphasis on integration, systematically archiving, and the coordination of past, present and future space and ground-based analogue experiments. We also discuss possible mechanisms for such integration across disciplines and missions. This article then introduces several discipline-specific reviews that show how such integration can be implemented. Areas explored include: adaptation of the central nervous system to space; cerebral autoregulation and weightlessness; modelling of the cardiovascular system in space exploration; human metabolic response to spaceflight; and exercise, artificial gravity, and physiologic countermeasures for spaceflight. In summary, spaceflight physiology research needs a conceptual framework that extends problem solving beyond disciplinary barriers. Administrative commitment and a high degree of cooperation among investigators are needed to further such a process. Well-designed interdisciplinary research can expand opportunities for broad interpretation of results across multiple physiological systems, which may have applications on Earth.     

Perceptual distortion contributes to the curvature of human reaching movements

Unconstrained point-to-point human arm movements are generally gently curved, a fact which has been used to assess the validity of models of trajectory formation. In this study we examined the relationship between curvature perception and movement curvature for planar sagittal and transverse arm movements. We found a significant correlation (P<0.0001, n=16) between the curvature perceived as straight and the curvature of actual arm movements. We suggest that subjects try to make straight-line movements, but that actual movements are curved because visual perceptual distortion makes the movements appear to be straighter than they really are. We conclude that perceptual distortion of curvature contributes to the curvature seen in human point-to-point arm movements and that this must be taken into account in the assessment of models of trajectory formation.     

On the nature of the vestibular control of arm-reaching movements during whole-body rotations

Recent studies report efficient vestibular control of goal-directed arm movements during body motion. This contribution tested whether this control relies (a) on an updating process in which vestibular signals are used to update the perceived egocentric position of surrounding objects when body orientation changes, or (b) on a sensorimotor process, i.e. a transfer function between vestibular input and the arm motor output that preserves hand trajectory in space despite body rotation. Both processes were separately and specifically adapted. We then compared the respective influences of the adapted processes on the vestibular control of arm-reaching movements. The rationale was that if a given process underlies a given behavior, any adaptive modification of this process should give rise to observable modification of the behavior. The updating adaptation adapted the matching between vestibular input and perceived body displacement in the surrounding world. The sensorimotor adaptation adapted the matching between vestibular input and the arm motor output necessary to keep the hand fixed in space during body rotation. Only the sensorimotor adaptation significantly altered the vestibular control of arm-reaching movements. Our results therefore suggest that during passive self-motion, the vestibular control of arm-reaching movements essentially derives from a sensorimotor process by which arm motor output is modified on-line to preserve hand trajectory in space despite body displacement. In contrast, the updating process maintaining up-to-date the egocentric representation of visual space seems to contribute little to generating the required arm compensation during body rotations.     

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. Received: 21 May 1999 / Accepted: 25 May 1999     

Lung function is unchanged in the 1 G environment following 6-months exposure to microgravity

Many organ systems adapt in response to the removal of gravity, such as that occurring during spaceflight. Such adaptation occurs over varying time periods depending on the organ system being considered, but the effect is that upon a return to the normal 1 G environment, the organ system is ill-adapted to that environment. As a consequence, either countermeasures to the adaptive process in flight, or rehabilitation upon return to 1 G is required. To determine whether the lung changed in response to a long period without gravity, we studied numerous aspects of lung function on ten subjects (one female) before and after they were exposed to 4-6 months of microgravity (microG, weightlessness) in the normobaric normoxic environment of the International Space Station. With the exception of small (and likely physiologically inconsequential) changes in expiratory reserve volume, one index of peripheral gas mixing in the periphery of the lung, and a possible slight reduction in D(L)CO in the early postflight period despite an unchanged cardiac output, lung function was unaltered by 4-6 months in microG. These results suggest that unlike many other organ systems in the human body, lung function returns to normal after long term exposure to the removal of gravity. We conclude that that in a normoxic, normobaric environment, lung function is not a concern following long-duration future spaceflight exploration missions of up to 6 months.     

The central nervous system does not minimize energy cost in arm movements

It has been widely suggested that the many degrees of freedom of the musculoskeletal system may be exploited by the CNS to minimize energy cost. We tested this idea by having subjects making point-to-point movements while grasping a robotic manipulandum. The robot created a force field chosen such that the minimal energy hand path for reaching movements differed substantially from those observed in a null field. The results show that after extended exposure to the force field, subjects continued to move exactly as they did in the null field and thus used substantially more energy than needed. Even after practicing to move along the minimal energy path, subjects did not adapt their freely chosen hand paths to reduce energy expenditure. The results of this study indicate that for point-to-point arm movements minimization of energy cost is not a dominant factor that influences how the CNS arrives at kinematics and associated muscle activation patterns.     

Multisensory adaptation of spatial-to-motor transformations in children with developmental coordination disorder

Recent research has demonstrated that adaptation to a visuomotor distortion systematically influenced movements to auditory targets in adults and typically developing (TD) children, suggesting that the adaptation of spatial-to-motor transformations for reaching movements is multisensory (i.e., generalizable across sensory modalities). The multisensory characteristics of these transformations in children with developmental coordination disorder (DCD) have not been examined. Given that previous research has demonstrated that children with DCD have deficits in sensorimotor integration, these children may also have impairments in the formation of multisensory spatial-to-motor transformations for target-directed arm movements. To investigate this hypothesis, children with and without DCD executed discrete arm movements to visual and acoustic targets prior to and following exposure to an abrupt visual feedback rotation. Results demonstrated that the magnitudes of the visual aftereffects were equivalent in the TD children and the children with DCD, indicating that both groups of children adapted similarly to the visuomotor perturbation. Moreover, the influence of visuomotor adaptation on auditory-motor performance was similar in the two groups of children. This suggests that the multisensory processes underlying adaptation of spatial-to-motor transformations are similar in children with DCD and TD children.     

Effects of spaceflight on ocular counterrolling and the spatial orientation of the vestibular system

We recorded the horizontal (yaw), vertical (pitch), and torsional (roll) eye movements of two rhesus monkeys with scierai search coils before and after the COSMOS Biosatellite 2229 Flight. The aim was to determine effects of adaptation to microgravity on the vestibulo-ocular reflex (VOR). The animals flew for 11 days. The first postflight tests were 22 h and 55 h after landing, and testing extended for 11 days after reentry. There were four significant effects of spaceflight on functions related to spatial orientation: (1) Compensatory ocular counterrolling (OCR) was reduced by about 70% for static and dynamic head tilts with regard to gravity. The reduction in OCR persisted in the two animals throughout postflight testing. (2) The gain of the torsional component of the angular VOR (roll VOR) was decreased by 15% and 50% in the two animals over the same period. (3) An up-down asymmetry of nystagmus, present in the two monkeys before flight was reduced after exposure to microgravity. (4) The spatial orientation of velocity storage was shifted in the one monkey that could be tested soon after flight. Before flight, the yaw axis eigenvector of optokinetic afternystagmus was close to gravity when the animal was upright or tilted. After flight, the yaw orientation vector was shifted toward the body yaw axis. By 7 days after recovery, it had reverted to a gravitational orientation. We postulate that spaceflight causes changes in the vestibular system which reflect adaptation of spatial orientation from a gravitational to a body frame of reference. These changes are likely to play a role in the postural, locomotor, and gaze instability demonstrated on reentry after spaceflight.     

Slowing of human arm movements during weightlessness: the role of vision

The human motor system responds to weightlessness by the slowing of movement. It has been suggested that deficits in visuo-motor co-ordination cause this effect. We studied the mechanisms of the slowing of movement in three long-term missions to the Russian space station Mir. In particular, the role of vision in the control of movement in microgravity has been studied in these experiments on seven cosmonauts, pre-, in-, and post-flight. The cosmonauts made arm movements to visual targets under the following conditions of visual control: no visual control, interrupted visual control, and undisturbed visual control. The results showed that the slowing of movement during weightlessness was manifested by decreases of peak velocity and peak acceleration, was not associated with a prolongation of the movement phase of deceleration, and was not affected by manipulation of the conditions of visual control. The slowing of movement tended to subside after the months of the flight and completely disappeared within days after the landing. Accuracy of the movements strictly depended on the constraints imposed on the vision and remained unaffected in-flight. The data presented demonstrate that the slowing of movement in microgravity is not directly related to deficits in sensori-motor co-ordination and is not associated with a reduction of the accuracy of movement. The strategy for motor control in microgravity seems to be directed towards the generation of smooth movements and the maintenance of their accuracy. Electronic Publication     

Are arm trajectories planned in kinematic or dynamic coordinates? An adaptation study

There are several invariant features of pointto-point human arm movements: trajectories tend to be straight, smooth, and have bell-shaped velocity profiles. One approach to accounting for these data is via optimization theory; a movement is specified implicitly as the optimum of a cost function, e.g., integrated jerk or torque change. Optimization models of trajectory planning, as well as models not phrased in the optimization framework, generally fall into two main groups-those specified in kinematic coordinates and those specified in dynamic coordinates. To distinguish between these two possibilities we have studied the effects of artificial visual feedback on planar two-joint arm movements. During self-paced point-to-point arm movements the visual feedback of hand position was altered so as to increase the perceived curvature of the movement. The perturbation was zero at both ends of the movement and reached a maximum at the midpoint of the movement. Cost functions specified by hand coordinate kinematics predict adaptation to increased curvature so as to reduce the visual curvature, while dynamically specified cost functions predict no adaptation in the underlying trajectory planner, provided the final goal of the movement can still be achieved. We also studied the effects of reducing the perceived curvature in transverse movements, which are normally slightly curved. Adaptation should be seen in this condition only if the desired trajectory is both specified in kinematic coordinates and actually curved. Increasing the perceived curvature of normally straight sagittal movements led to significant (P<0.001) corrective adaptation in the curvature of the actual hand movement; the hand movement became curved, thereby reducing the visually perceived curvature. Increasing the curvature of the normally curved transverse movements produced a significant (P<0.01) corrective adaptation; the hand movement became straighter, thereby again reducing the visually perceived curvature. When the curvature of naturally curved transverse movements was reduced, there was no significant adaptation (P>0.05). The results of the curvature-increasing study suggest that trajectories are planned in visually based kinematic coordinates. The results of the curvature-reducing study suggest that the desired trajectory is straight in visual space. These results are incompatible with purely dynamicbased models such as the minimum torque change model. We suggest that spatial perception-as mediated by vision-plays a fundamental role in trajectory planning.     

The effect of visuomotor displacements on arm movement paths

The gently curved paths evident in point-to-point arm movements have been attributed to both an imperfect execution of a planned straight-hand path or as an emergent property of a control strategy in which an intrinsic cost, dependent on arm dynamics, is minimised. We used a virtual visual feedback system to test whether path curvature was mainly determined by the visually perceived or actual location of the moving limb. Hand paths were measured for movements between three pairs of targets under both veridical and uniformly translated visual feedback. This allowed us to decouple the actual and perceived hand location during movement. Under different conditions of visual feedback the curvature of the hand paths did not correlate with either the visually perceived location of the limb or the actual location but rather with the relative displacement between the actual and visually perceived limb locations. The results are consistent with the hypothesis that in planning a movement the internal estimate of intrinsic coordinates, such as joint angles, is at least partially derived from visual information. Received: 31 August 1998 / Accepted: 8 March 1999     

Adaptation of postural control to weightlessness

Summary Adaptation of motor control to weightlessness was studied during a 7-day spaceflight. The maintenance of control of upright posture was examined during a voluntary raising movement of the arm and during the voluntary raising on tiptoe. In order to evaluate the contribution of visual cues, three types of visual situations were examined: normal vision, central vision, and without vision. On the basis of cinematographic and mechanographic data, the postural perturbations consecutive to the movement of a body part in conditions of weightlessness were found to be similar to those observed on earth. However, in weightlessness, in contrast to the ground-based situation, erectness of posture was maintained primarily due to the predominant contraction of the ankle flexor muscles. The sequences of postural leg muscle activity associated with the arm or foot movement were well structured and varied slightly in the course of the flight. In addition, the initial posture, that is the erect posture before the movement was executed, changed throughout the flight from an exaggerated oblique position to a terrestrial standing position. Visual information was preponderant at the beginning of the space mission for the recalibration of other sensory cues affected by weightlessness. The findings are indicative of two types of adaptation of the central program of posture regulation to weightlessness: fast, short-term adaptation, characterized by a quasi-instantaneous redistribution of motor commands between ankle flexors and extensors (an operative process) and slow, long-term adaptation, exemplified by the loss of anticipatory activation of certain muscles by the end of the flight (a conservative process).     

The Role of Gravitation-Dependent Systems in Visual Tracking

The effects of prolonged microgravity conditions on the performance of visual tracking functions such as fixational rotations of the eyes (saccades), smooth tracking of linear and curved movements of a foveal point stimulus, and following a vertical pendulum-like movement of foveoretinal optokinetic stimuli were studied. Experiments were performed on 31 cosmonauts in freefall conditions, in ten cases followed by additional studies after a cycle of head movements and in 14 after resting. These experiments showed that while intrinsic visual functions were retained in microgravity conditions, there were decreases in the precision and speed measures of all types of visual tracking (fixational rotations of the eyes, smooth tracking) and, in some cases, complete degradation of the smooth tracking reflex, an increase in the time taken to fix the gaze on a target (by factors of 2 or more), and decreases in the frequency of stimulus tracking. During the initial period of adaptation to the altered gravitational conditions and periodically during prolonged flight, the system of smooth visual tracking was found to undergo a transition to a strategy of saccadic approximation, in which gaze tracks the movement of the target using a set of macro- or microsaccadic movements. These impairments, seen in virtually all the cosmonauts, resulted from vestibular deprivation (functional deafferentation of the otolith input) in conditions of weightlessness, while in cosmonauts conceptualizing space on the basis of perceiving the positions of the feet and head additionally showed support-tactile deprivation.     

Effects of cosmonaut vestibular training on vestibular function prior to spaceflight

The purpose of this study was to quantify the effects of repetitive Coriolis and cross-coupled stimulations, similar to the vestibular training the cosmonauts are exposed to prior to their spaceflight, on vestibular function in control subjects on Earth. Ten volunteers were passively rotated in yaw on a rotating chair while executing standardized pitch head-and-trunk movements. The chair stopped to change direction after 12 head-and-trunk movements were made. The runs were grouped in sessions of ten,which were repeated daily for 10 days. The severity of motion sickness was assessed by subjective judgment and measurements of skin pallor and salivary total protein concentration, and nystagmus was recorded. The severity of motion sickness and nystagmus decreased during cosmonaut vestibular training (CVT). One month after the end of CVT, nystagmus responses were still about 20–30% lower than control values. These results indicate that CVT induces a habituation of vestibular responses. One important implication of this experiment concerns space studies on cosmonauts who are exposed to such vestibular training prior to their spaceflight. Electronic Publication     

Visual-shift adaptation is composed of separable sensory and task-dependent effects

Visuomotor coordination requires both the accurate alignment of spatial information from different sensory streams and the ability to convert these sensory signals into accurate motor commands. Both of these processes are highly plastic, as illustrated by the rapid adaptation of goal-directed movements following exposure to shifted visual feedback. Although visual-shift adaptation is a widely used model of sensorimotor learning, the multifaceted adaptive response is typically poorly quantified. We present an approach to quantitatively characterizing both sensory and task-dependent components of adaptation. Sensory aftereffects are quantified with "alignment tests" that provide a localized, two-dimensional measure of sensory recalibration. These sensory effects obey a precise form of "additivity," in which the shift in sensory alignment between vision and the right hand is equal to the vector sum of the shifts between vision and the left hand and between the right and left hands. This additivity holds at the exposure location and at a second generalization location. These results support a component transformation model of sensory coordination, in which eye-hand and hand-hand alignment relies on a sequence of shared sensory transformations. We also ask how these sensory effects compare with the aftereffects measured in target reaching and tracking tasks. We find that the aftereffect depends on both the task performed during feedback-shift exposure and on the testing task. The results suggest the presence of both a general sensory recalibration and task-dependent sensorimotor effect. The task-dependent effect is observed in highly stereotyped reaching movements, but not in the more variable tracking task.     

Biomechanical analysis of running in weightlessness on a treadmill equipped with a subject loading system

One countermeasure used during long-duration spaceflight to maintain bone and muscle mass is a treadmill equipped with a subject loading system (SLS) that simulates gravity. To date, little is known about the biomechanics of running in weightlessness on such a treadmill-SLS system. We have designed an instrumented treadmill/force plate to compare the biomechanics of running in weightlessness to running on Earth. Gravity was simulated by two pneumatic pistons pulling downward on a subject’s harness, with a force approximately equal to body weight on Earth. Four transducers, mounted under the treadmill, measured the three components of the reaction force exerted by the tread belt under the foot. A high-speed video camera recorded the movements of limb segments while the electromyography of the four lower limb muscles was registered. Experiments in weightlessness were conducted during the European Space Agency parabolic flight campaigns. Control experiments were performed on the same subjects on Earth. When running on the treadmill with an SLS, the bouncing mechanism of running is preserved. Depending on the speed of progression, the ground reaction forces, contact and aerial times, muscular work and bone stress differed by a maximum of ±5–15% during running on the treadmill with an SLS, as compared to that on Earth. The movements of the lower limb segments and the EMG patterns of the lower limb muscles were also comparable. Thus, the biomechanics of running on Earth can reasonably be duplicated in weightlessness using a treadmill with an SLS that generates a pull-down force close to body weight on Earth.