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.     

Kinetic analysis of arm reaching movements during voluntary and passive rotation of the torso

Reaching movements made to targets during exposure to passive constant velocity rotation show significant endpoint errors. By contrast, reaching movements made during voluntary rotation of the torso are accurate. In both cases, as a consequence of the simultaneous motion of the arm and the torso, Coriolis forces are generated on the arm tending to deflect its path. Our goal in the present paper was to determine whether during voluntary torso rotations arm movement accuracy is preserved by feed forward compensations for self-generated Coriolis forces. To test this hypothesis we analyzed and quantified the contribution of torso rotation and translation to arm dynamics and compared the kinematics and kinetics of pointing movements during voluntary and passive torso rotation. Coriolis torques at the shoulder increase nearly sixfold in voluntary turn and reach movements relative to reaches made without torso rotation, yet are equally accurate. Coriolis torques during voluntary turn and reach movements are more than double those produced by reaching movements during passive body rotation at 60°/s. Nevertheless, the endpoints of the reaches made during voluntary rotation are not deviated, but those of reaches made during passive rotation are deviated in the direction of the Coriolis forces generated during the movements. We conclude that there is anticipatory pre-programmed compensation for self-generated Coriolis forces during voluntary torso rotation contingent on intended torso motion and arm trajectory.     

Influence of interaction force levels on degree of motor adaptation in a stable dynamic force field

Studies have shown that the point-to-point reaching movements of subjects seated in a dark, rotating room demonstrate errors in movement trajectories and endpoints, consistent with the direction of the Coriolis force perturbations created by room rotation. Adaptation of successive reaches and the presence of postrotation aftereffects have indicated that subjects form internal models of the Coriolis field dynamics in order to make appropriate movement corrections. It has been argued that these findings are inconsistent with predictions of peripheral stabilization assumed in equilibrium-point models of motor control. A possibility that has been raised, however, is that the Coriolis field findings may in fact stem from changes in control commands elicited due to the magnitude and destabilizing nature of the Coriolis perturbations. That is, it has been suggested that a perturbation threshold exists, below which central reactions are not necessary in order to maintain movement stability. We tested the existence of a perturbation threshold in normal-speed reaching movements. Twelve normal human subjects performed non-visually guided reaching movements while grasping a robotic manipulandum. The endpoints and trajectory deviations of their movements were measured before, during, and after a position-dependent force field (similar to a Coriolis field in terms of the time history of applied forces) was applied to their movements. We examined the responses to a range of perturbation field strengths from small to considerable. Our experimental results demonstrated a substantial adaptation response over the entire range of perturbation field magnitudes examined. Neither the amount of adaptation after 5 trials nor after 25 trials was found to change as disturbance magnitudes decreased. These findings indicate that there is an adaptive response even for small perturbations; i.e., threshold behavior was not found. This result contradicts the assertion that peripheral stabilization mechanisms enable the central controller to ignore small details of peripheral or environmental dynamics. Our findings instead point to a central dynamic modeler that is both highly sensitive and continually active. The results of our study also showed that subjects were able to maintain baseline pointing accuracies despite exposure to perturbation forces of sizeable magnitude (more than 7 N).     

Coordinated turn-and-reach movements. I. Anticipatory compensation for self-generated coriolis and interaction torques

When reaching movements involve simultaneous trunk rotation, additional interaction torques are generated on the arm that are absent when the trunk is stable. To explore whether the CNS compensates for such self-generated interaction torques, we recorded hand trajectories in reaching tasks involving various amplitudes and velocities of arm extension and trunk rotation. Subjects pointed to three targets on a surface slightly above waist level. Two of the target locations were chosen so that a similar arm configuration relative to the trunk would be required for reaching to them, one of these targets requiring substantial trunk rotation, the other very little. Significant trunk rotation was necessary to reach the third target, but the arm's radial distance to the body remained virtually unchanged. Subjects reached at two speeds-a natural pace (slow) and rapidly (fast)-under normal lighting and in total darkness. Trunk angular velocity and finger velocity relative to the trunk were higher in the fast conditions but were not affected by the presence or absence of vision. Peak trunk velocity increased with increasing trunk rotation up to a maximum of 200 degrees /s. In slow movements, peak finger velocity relative to the trunk was smaller when trunk rotation was necessary to reach the targets. In fast movements, peak finger velocity was approximately 1.7 m/s for all targets. Finger trajectories were more curved when reaching movements involved substantial trunk rotation; however, the terminal errors and the maximal deviation of the trajectory from a straight line were comparable in slow and fast movements. This pattern indicates that the larger Coriolis, centripetal, and inertial interaction torques generated during rapid reaches were compensated by additional joint torques. Trajectory characteristics did not vary with the presence or absence of vision, indicating that visual feedback was unnecessary for anticipatory compensations. In all reaches involving trunk rotation, the finger movement generally occurred entirely during the trunk movement, indicating that the CNS did not minimize Coriolis forces incumbent on trunk rotation by sequencing the arm and trunk motions into a turn followed by a reach. A simplified model of the arm/trunk system revealed that additional interaction torques generated on the arm during voluntary turning and reaching were equivalent to < or =1.8 g (1 g = 9.81 m/s(2)) of external force at the elbow but did not degrade performance. In slow-rotation room studies involving reaching movements during passive rotation, Coriolis forces as small as 0.2 g greatly deflect movement trajectories and endpoints. We conclude that compensatory motor innervations are engaged in a predictive fashion to counteract impending self-generated interaction torques during voluntary reaching movements.     

Balance in a rotating artificial gravity environment

When subjects stand at the center of a fully enclosed room that is rotating at constant velocity, their natural postural sway generates Coriolis forces that destabilize their center of mass and head. We quantitatively assessed how exposure to constant velocity rotation at 10 rpm affected postural control. Twelve subjects stood in a heel-to-toe stance in the rotating room. Each test session involved three phases: (1) pre-rotation, (2) per-rotation, and (3) post-rotation. In each phase, subjects were tested in both eyes open and eyes closed conditions. Four measures were used to characterize center of mass movement and head movement: mean sway amplitude, total power, mean power frequency, and frequency of maximum power. Each measure was computed for anterior-posterior and medial-lateral sway. Both anterior-posterior and medial-lateral head and center of mass sway during rotation had significantly greater mean sway amplitude and total power compared with pre- and post-rotation values. Mean power frequency and frequency of maximum power were little affected. Eyes open conditions were significantly more stable in all test phases than eyes-closed, but vision did not completely suppress the effects of rotation. The greatest effect of rotation was in the eyes-closed condition with mean sway amplitude and total power increasing more than twofold. Inverted pendulum sway was maintained in all phases of both test conditions. No aftereffects of rotation were present after the four 25-s exposures each subject received. We expect that with longer exposure periods and with active generation of body sway subjects would both adapt to rotation and exhibit post-rotary aftereffects. Electronic Publication     

Moving objects in a rotating environment: rapid prediction of Coriolis and centrifugal force perturbations

Grip force adaptation to Coriolis and centrifugal force perturbations was tested in healthy subjects. Eight subjects were seated in a rotating chamber in a rotating axis position. They each grasped an instrumented object resting on the thumb, which was stabilized by the other fingers from above. Subjects performed horizontal point-to-point movements with the grasped object away and towards the trunk. These movements were directed in a nonparallel fashion towards the axis of rotation prior (40 pre-rotational movements), during (80 per-rotational movements) and following (40 post-rotational movements) clockwise body rotation. During pre- and post-rotational movements two load force peaks of similar magnitude occurred during the acceleratory and deceleratory phases of the movements. Accordingly, a Coriolis force, which was orthogonal and proportional to the linear velocity of the moving arm, as well as a centrifugal force proportional to the systems squared angular velocity and movement amplitude developed during per-rotational movements. The load perturbations altered the load force profile in a characteristic way. The first 10 per-rotational movement sequence revealed that there was a less precise coupling between grip and load force magnitudes and a reduced temporo-spatial co-ordination between grip and load force profiles. With increasing number of per-rotational movements, there was significant improvement in the temporo-spatial co-ordination and in the coupling in force magnitude between grip and load force profiles, indicating an ongoing adaptation process. The coupling between grip and load forces proved to be similarly precise for the last 10 per-rotational movements and for pre-rotational movements, suggesting complete adaptation. Significant effects were observed for the first post rotational movements following adaptation to the per-rotational load characteristics both for the temporal co-ordination between grip and load forces and for the coupling in force magnitudes. However, the last 10 post-rotational movements proved to be similarly precise with comparison to pre-rotational performance in terms of grip force regulation with movement-induced loads. The results are discussed within the context of the CNS ability to use internal models when planning and processing anticipatory grip force adjustments during manipulative tasks.     

Sensorimotor adaptation to inertial forces in a multi-force environment does not depend on the number of targets: indirect validation of the altered-proprioception hypothesis

The ability of our sensorimotor system to adapt to changing and complex environmental demands has been under experimental scrutiny for more than a century. Previous works have shown that aimed arm movements adapt quickly and completely to Coriolis force, but incompletely to the combination of Coriolis and centrifugal forces without visual cues. Two hypotheses may be advanced to explain this discrepancy: the workspace-exploration hypothesis, and the degraded-proprioception hypothesis. The aim of this study was to distinguish between the above two alternatives by comparing adaptive improvement during off-axis rotation in subjects pointing at one, three or seven different targets in complete darkness. Two main results emerge: (a) off-axis rotation led initially to errors in the direction of Coriolis force and in the opposite direction of the centrifugal force; (b) the size of the visited workspace has no effect on the way the subjects adapt to a multi-force environment. The lack of a target-number effect and the persistence of lateral errors in the pointing movements performed during rotation of the platform, support the degraded-proprioception rather than the workspace-exploration hypothesis of adaptation to a multi-force environment.     

Features of motor performance that drive adaptation in rapid hand movements

In order to explore how subjects correct for errors in movement and adapt their motor programs, we studied rapid hand movements. Subjects grasped a grooved knob and made brisk turning movements to various targets, similar to tuning a radio dial. A motor attached to the knob shaft was configured to apply a destabilizing negative viscous perturbation. Following a period of practice with no perturbations, the negative viscosity was engaged, which caused a large change in overall kinematics: the peak velocity increased, the movement amplitude was too large, and discrete corrective submovements were generated to bring the pointer back onto the target. After about an hour and nearly 1000 trials, subjects learned to move accurately in the new dynamic environment, returning their overall kinematics near to previous levels. Measures of performance included the endpoint error of the primary movement (the initial movement segment), the frequency and amplitude of corrective submovements, task success rate, mean squared jerk, and deviation from a "normal" angular velocity temporal profile. Both the amplitude and frequency of corrective submovements decreased progressively during adaptation as the subjects made fewer target overshoot errors. These results are consistent with motor learning schemes in which adaptation of the motor controller is driven by an attempt to reduce the endpoint error of the primary movement. While there have been many theories regarding what is being optimized in motor control, in general, biologically plausible mechanisms for implementing these schemes have not been described. A biologically plausible optimization criterion is the minimization of the occurrence and amplitude of corrective submovements, since the latter have been proposed as realistic climbing fiber training signals for adaptive changes in the cerebellum. We postulate that the other criteria that have been proposed are instead secondary to an increased accuracy of the primary movement and a corresponding decrease in the occurrence and amplitude of corrective submovements. Electronic Publication     

Endpoints of arm movements to visual targets

Reaching out for objects with an unseen arm involves using both visual and kinesthetic information. Neither visual nor kinesthetic information is perfect. Each is subject to both constant and variable errors. To evaluate how such errors influence performance in natural goal-directed movements, we asked subjects to align a real 5-cm cube, which they held in their hand but could not see, with a three-dimensional visual simulation of such a cube. The simulated cube was presented at one of four target locations at the corners of an imaginary tetraeder. Subjects made successive, self-paced movements between these target locations. They could not see anything except the simulated cube throughout the experiment. Initial analysis of the spatial dispersion of movement endpoints demonstrated that the major source of errors under these conditions was visual. Further analysis of the relationship between variability of the starting positions and endpoints showed that the errors were primarily in judging the endpoint, rather than the direction or amplitude of the required movement vector. The findings support endpoint control of human goal-directed movements.     

Labyrinthine lesions and motion sickness susceptibility

The angular vestibulo-ocular reflex (aVOR) has a fast pathway, which mediates compensatory eye movements, and a slow (velocity storage) pathway, which determines its low frequency characteristics and orients eye velocity toward gravity. We have proposed that motion sickness is generated through velocity storage, when its orientation vector, which lies close to the gravitational vertical, is misaligned with eye velocity during head motion. The duration of the misalignment, determined by the dominant time constant of velocity storage, causes the buildup of motion sickness. To test this hypothesis, we studied bilateral labyrinthine-defective subjects with short vestibular time constants but normal aVOR gains for their motion sickness susceptibility. Time constants and gains were taken from rotational responses. Motion sickness was generated by rolling the head while rotating, and susceptibility was assessed by the number of head movements made before reaching intolerable levels of nausea. More head movements signified lower motion sickness susceptibility. Labyrinthine-defective subjects made more head movements on their first exposure to roll while rotating than normals (39.8 ± 7.2 vs 13.7 ± 5.5; P < 0.0001). Normals were tested eight times, which habituated their time constants and reduced their motion sickness susceptibility. Combining data from all subjects, there was a strong inverse relationship between time constants and number of head movements (r = 0.94), but none between motion sickness susceptibility and aVOR gains. This provides further evidence that motion sickness is generated through velocity storage, not the direct pathway, and suggests that motion sickness susceptibility can be reduced by reducing the aVOR time constant.     

Prism adaptation during walking generalizes to reaching and requires the cerebellum

Adaptation of arm movements to laterally displacing prism glasses is usually highly specific to body part and movement type and is known to require the cerebellum. Here, we show that prism adaptation of walking trajectory generalizes to reaching (a different behavior involving a different body part) and that this adaptation requires the cerebellum. In experiment 1, healthy control subjects adapted to prisms during either reaching or walking and were tested for generalization to the other movement type. We recorded lateral deviations in finger endpoint position and walking direction to measure negative aftereffects and generalization. Results showed that generalization of prism adaptation is asymmetric: walking generalizes extensively to reaching, but reaching does not generalize to walking. In experiment 2, we compared the performance of cerebellar subjects versus healthy controls during the prism walking adaptation. We measured rates of adaptation, aftereffects, and generalization. Cerebellar subjects had reduced adaptation magnitudes, slowed adaptation rates, decreased negative aftereffects, and poor generalization. Based on these experiments, we propose that prism adaptation during whole body movements through space invokes a more general system for visuomotor remapping, involving recalibration of higher-order, effector-independent brain regions. In contrast, prism adaptation during isolated movements of the limbs is probably recalibrated by effector-specific mechanisms. The cerebellum is an essential component in the network for both types of prism adaptation.     

Coordinated turn-and-reach movements. II. Planning in an external frame of reference

The preceding study demonstrated that normal subjects compensate for the additional interaction torques generated when a reaching movement is made during voluntary trunk rotation. The present paper assesses the influence of trunk rotation on finger trajectories and on interjoint coordination and determines whether simultaneous turn-and-reach movements are most simply described relative to a trunk-based or an external reference frame. Subjects reached to targets requiring different extents of arm joint and trunk rotation at a natural pace and quickly in normal lighting and in total darkness. We first examined whether the larger interaction torques generated during rapid turn-and-reach movements perturb finger trajectories and interjoint coordination and whether visual feedback plays a role in compensating for these torques. These issues were addressed using generalized Procrustes analysis (GPA), which attempts to overlap a group of configurations (e.g., joint trajectories) through translations and rotations in multi-dimensional space. We first used GPA to identify the mean intrinsic patterns of finger and joint trajectories (i.e., their average shape irrespective of location and orientation variability in the external and joint workspaces) from turn-and-reach movements performed in each experimental condition and then calculated their curvatures. We then quantified the discrepancy between each finger or joint trajectory and the intrinsic pattern both after GPA was applied individually to trajectories from a pair of experimental conditions and after GPA was applied to the same trajectories pooled together. For several subjects, joint trajectories but not finger trajectories were more curved in fast than slow movements. The curvature of both joint and finger trajectories of turn-and-reach movements was relatively unaffected by the vision conditions. Pooling across speed conditions significantly increased the discrepancy between joint but not finger trajectories for most subjects, indicating that subjects used different patterns of interjoint coordination in slow and fast movements while nevertheless preserving the shape of their finger trajectory. Higher movement speeds did not disrupt the arm joint rotations despite the larger interaction torques generated. Rather, subjects used the redundant degrees of freedom of the arm/trunk system to achieve similar finger trajectories with differing joint configurations. We examined finger movement patterns and velocity profiles to determine the frame of reference in which turn-and-reach movements could be most simply described. Finger trajectories of turn-and-reach movements had much larger curvatures and their velocity profiles were less smooth and less bell-like in trunk-based coordinates than in external coordinates. Taken together, these results support the conclusion that turn-and-reach movements are controlled in an external frame of reference.     

Visual gravity influences arm movement planning

When submitted to a visuomotor rotation, subjects show rapid adaptation of visually guided arm reaching movements, indicated by a progressive reduction in reaching errors. In this study, we wanted to make a step forward by investigating to what extent this adaptation also implies changes into the motor plan. Up to now, classical visuomotor rotation paradigms have been performed on the horizontal plane, where the reaching motor plan in general requires the same kinematics (i.e., straight path and symmetric velocity profile). To overcome this limitation, we considered vertical and horizontal movement directions requiring specific velocity profiles. This way, a change in the motor plan due to the visuomotor conflict would be measurable in terms of a modification in the velocity profile of the reaching movement. Ten subjects performed horizontal and vertical reaching movements while observing a rotated visual feedback of their motion. We found that adaptation to a visuomotor rotation produces a significant change in the motor plan, i.e., changes to the symmetry of velocity profiles. This suggests that the central nervous system takes into account the visual information to plan a future motion, even if this causes the adoption of nonoptimal motor plans in terms of energy consumption. However, the influence of vision on arm movement planning is not fixed, but rather changes as a function of the visual orientation of the movement. Indeed, a clear influence on motion planning can be observed only when the movement is visually presented as oriented along the vertical direction. Thus vision contributes differently to the planning of arm pointing movements depending on motion orientation in space.     

Comparison of variability of initial kinematics and endpoints of reaching movements

 The accuracy of reaching movements to memorized visual target locations is presumed to be determined largely by central planning processes before movement onset. If so, then the initial kinematics of a pointing movement should predict its endpoint. Our study examined this hypothesis by testing the correlation between peak acceleration, peak velocity, and movement amplitude and the correspondence between the respective spatial positions of these kinematic landmarks. Subjects made planar horizontal reaching movements to targets located at five different distances and along five radially arrayed directions without visual feedback during the movements.The spatial dispersion of the positions of peak acceleration, peak velocity, and endpoint all tended to form ellipses oriented along the movement trajectory. However, whereas the peaks of acceleration and velocity scaled strongly with movement amplitude for all of the movements made at the five target distances in any one direction, the correlations with movement amplitude were more modest for trajectories aimed at each target separately. Furthermore, the spatial variability in direction and extent of the distribution of positions of peak acceleration and peak velocity did not scale differently with target distance, whereas they did for endpoint distributions. Therefore, certain features of the final kinematics are evident in the early kinematics of the movements as predicted by the hypothesis that they reflect planning processes. However, endpoint distributions were not completely predetermined by the Initial kinematics. In contrast, multivariate analysis suggests that adjustments to movement duration help compensate for the variability of the initial kinematics to achieve desired movement amplitude. These compensatory adjustments do not contradict the general conclusion that the systematic patterns in the spatial variability observed in this study reflect planning processes. On the contrary, and consistent with that conclusion, our results provide further evidence that direction and extent of reaching movements are planned and determined in parallel over time. Received: 23 March 1998 / Accepted: 2 September 1998     

Optimal impedance control for task achievement in the presence of signal-dependent noise

There is an infinity of impedance parameter values, and thus different co-contraction levels, that can produce similar movement kinematics from which the CNS must select one. Although signal-dependent noise (SDN) predicts larger motor-command variability during higher co-contraction, the relationship between impedance and task performance is not theoretically obvious and thus was examined here. Subjects made goal-directed, single-joint elbow movements to either move naturally to different target sizes or voluntarily co-contract at different levels. Stiffness was estimated as the weighted summation of rectified EMG signals through the index of muscle co-contraction around the joint (IMCJ) proposed previously. When subjects made movements to targets of different sizes, IMCJ increased with the accuracy requirements, leading to reduced endpoint deviations. Therefore without the need for great accuracy, subjects accepted worse performance with lower co-contraction. When subjects were asked to increase co-contraction, the variability of EMG and torque both increased, suggesting that noise in the neuromotor command increased with muscle activation. In contrast, the final positional error was smallest for the highest IMCJ level. Although co-contraction increases the motor-command noise, the effect of this noise on the task performance is reduced. Subjects were able to regulate their impedance and control endpoint variance as the task requirements changed, and they did not voluntarily select the high impedance that generated the minimum endpoint error. These data contradict predictions of the SDN-based theory, which postulates minimization of only endpoint variance and thus require its revision.     

Postural configuration does not alter unperturbed or perturbed reach movement kinematics

This study investigated whether postural configuration has a significant effect upon the kinematics of arm movements when humans performed unconstrained reach movements to visual targets. Eight subjects were required to reach to static visual targets (unperturbed REACH movements) or correct reach movements when the position of a target unexpectedly changed during the execution of a planned movement (perturbed reaches, or online corrections, OC). Subjects were required to execute REACH and OC movements in sitting and standing (STAND) positions. The height of the targets, distance from the right shoulder (acromion) and eccentricity in terms of the body midline were standardized between the two postural conditions before movements begun. Unperturbed REACH movements were executed to a central target placed at 130 % of outstretched arm length, along the midline (0°). Perturbed (OC) movements involved subjects initiating an arm movement to the 0° target upon its illumination. Two hundred milliseconds after the onset of the hand movement, the 0° target was extinguished and the target at 60° to the right of the midline (still at 130 % outstretched arm distance) illuminated. Subjects had to correct their reach movements online to the new target. Results demonstrated that, despite evident differences in postural kinematics between the four experimental conditions (e.g. pelvis obliquity and trunk/pelvis rotation), postural configuration had little or no effect upon the endpoint kinematics of the finger. Most importantly, the STAND position, with its greater postural constraints, did not affect the time taken to initiate an OC, nor did it lengthen the time taken to complete the REACH or OC movements. Our results suggest, therefore, that postural constraints are accounted for by the central nervous system when executing complex arm reaching movements.     

Detection of cerebral oxyhaemoglobin changes during vestibular Coriolis cross-coupling stimulation using near infrared spectroscopy

Near infrared spectroscopy (NIRS) has been successful in monitoring cerebral haemodynamics when the subject is immobilized during surgery, and when there is a drastic depletion of blood from the cerebral cortex during positive acceleration. In this study, we monitored subtle changes of cerebral oxygen level using NIRS during vestibular stimulation. For the control conditions, cerebral oxygen status was monitored in six stationary subjects sitting upright, and while they executed head movements in the pitch axis with eyes opened and eyes closed. The experimental conditions involved the subjects making a head movement which required a 45 degrees pitch-down followed by a return to upright head movements 12 s later during yaw rotation (Coriolis cross coupling) at 10 and 20 rotations per minute (rpm) in a random order. Oxyhaemoglobin (O(2)Hb), deoxyhaemoglobin (HHb) and total haemoglobin levels were recorded every 0.5 s from both the parietal and the occipital lobe simultaneously. A significant rotation effect was observed in total Hb level changes from baseline in both regions. Occipital O(2)Hb increased significantly after the head movement with eyes opened at 20 rpm. Our findings appear to be consistent with previous vestibular studies that significant changes in brain blood flow occur during caloric stimulation. NIRS can be used to monitor discrete cortical blood flow changes resulting from vestibular and other forms of stimulation.     

Functional significance of stiffness in adaptation of multijoint arm movements to stable and unstable dynamics

This study compared the mechanisms of adaptation to stable and unstable dynamics from the perspective of changes in joint mechanics. Subjects were instructed to make point to point movements in force fields generated by a robotic manipulandum which interacted with the arm in either a stable or an unstable manner. After subjects adjusted to the initial disturbing effects of the force fields they were able to produce normal straight movements to the target. In the case of the stable interaction, subjects modified the joint torques in order to appropriately compensate for the force field. No change in joint torque or endpoint force was required or observed in the case of the unstable interaction. After adaptation, the endpoint stiffness of the arm was measured by applying displacements to the hand in eight different directions midway through the movements. This was compared to the stiffness measured similarly during movements in a null force field. After adaptation, the endpoint stiffness under both the stable and unstable dynamics was modified relative to the null field. Adaptation to unstable dynamics was achieved by selective modification of endpoint stiffness in the direction of the instability. To investigate whether the change in endpoint stiffness could be accounted for by change in joint torque or endpoint force, we estimated the change in stiffness on each trial based on the change in joint torque relative to the null field. For stable dynamics the change in endpoint stiffness was accurately predicted. However, for unstable dynamics the change in endpoint stiffness could not be reproduced. In fact, the predicted endpoint stiffness was similar to that in the null force field. Thus, the change in endpoint stiffness seen after adaptation to stable dynamics was directly related to changes in net joint torque necessary to compensate for the dynamics in contrast to adaptation to unstable dynamics, where a selective change in endpoint stiffness occurred without any modification of net joint torque.     

Rapid adaptation of torso pointing movements to perturbations of the base of support

We investigated whether pointing movements made with the torso would adapt to movement-contingent augmentation or attenuation of their spatial amplitude. The pointing task required subjects standing on a platform in the dark to orient the mid-sagittal plane of their torso to the remembered locations of just extinguished platform-fixed visual targets without moving their feet. Subjects alternated pointing at two chest-high targets, 60° apart, (1) in a baseline period with the stance platform stationary, (2) during exposure to concomitant contra or ipsiversive platform rotations that grew incrementally to 50% of the velocity of torso rotation, and (3) after return in one step to stationary platform conditions. The velocity and amplitude of torso movements relative to space decreased 25–50% during exposure to contraversive platform rotations and increased 20–50% during ipsiversive rotations. Torso rotation kinematics relative to the platform (as well as the platform-fixed targets and feet) remained virtually constant throughout the incremental exposure period. Subjects were unaware of the altered motion of their body in space imposed by the platform and did not perceive their motor adjustments. Upon return to stationary conditions, torso rotation movements were smaller and slower following adaptation to contraversive rotations and larger and faster after ipsiversive platform rotations. These results indicate a rapid sensory-motor recalibration to the altered relationship between spatial (inertial) torso motion and intended torso motion relative to the feet, and rapid re-adaptation to normal conditions. The adaptive system producing such robust torso regulation provides a critical basis for control of arm, head, and eye movements.