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.     

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.     

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.     

Contribution of execution noise to arm movement variability in three-dimensional space

Reaching movements are subject to noise associated with planning and execution, but precisely how these noise sources interact to determine patterns of endpoint variability in three-dimensional space is not well understood. For frontal plane movements, variability is largest along the depth axis (the axis along which visual planning noise is greatest), with execution noise contributing to this variability along the movement direction. Here we tested whether these noise sources interact in a similar way for movements directed in depth. Subjects performed sequences of two movements from a single starting position to targets that were either both contained within a frontal plane ("frontal sequences") or where the first was within the frontal plane and the second was directed in depth ("depth sequences"). For both sequence types, movements were performed with or without visual feedback of the hand. When visual feedback was available, endpoint distributions for frontal and depth sequences were generally anisotropic, with the principal axes of variability being strongly aligned with the depth axis. Without visual feedback, endpoint distributions for frontal sequences were relatively isotropic and movement direction dependent, while those for depth sequences were similar to those with visual feedback. Overall, the results suggest that in the presence of visual feedback, endpoint variability is dominated by uncertainty associated with planning and updating visually guided movements. In addition, the results suggest that without visual feedback, increased uncertainty in hand position estimation effectively unmasks the effect of execution-related noise, resulting in patterns of endpoint variability that are highly movement direction dependent.     

The role of the cross-sensory error signal in visuomotor adaptation

Reaching to targets with misaligned visual feedback of the hand leads to changes in proprioceptive estimates of hand position and reach aftereffects. In such tasks, subjects are able to make use of two error signals: the discrepancy between the desired and actual movement, known as the sensorimotor error signal, and the discrepancy between visual and proprioceptive estimates of hand position, which we refer to as the cross-sensory error signal. We have recently shown that mere exposure to a sensory discrepancy in the absence of goal-directed movement (i.e. no sensorimotor error signal) is sufficient to produce similar changes in felt hand position and reach aftereffects. Here, we sought to determine the extent that this cross-sensory error signal can contribute to proprioceptive recalibration and movement aftereffects by manipulating the magnitude of this signal in the absence of volitional aiming movements. Subjects pushed their hand out along a robot-generated linear path that was gradually rotated clockwise relative to the path of a cursor. On all trials, subjects viewed a cursor that headed directly towards a remembered target while their hand moved out synchronously. After exposure to a 30° rotated hand-cursor distortion, subjects recalibrated their sense of felt hand position and adapted their reaches. However, no additional increases in recalibration or aftereffects were observed following further increases in the cross-sensory error signal (e.g. up to 70°). This is in contrast to our previous study where subjects freely reached to targets with misaligned visual hand position feedback, hence experiencing both sensorimotor and cross-sensory errors, and the distortion magnitude systematically predicted increases in proprioceptive recalibration and reach aftereffects. Given these findings, we suggest that the cross-sensory error signal results in changes to felt hand position which drive partial reach aftereffects, while larger aftereffects that are produced after visuomotor adaptation (and that vary with the size of distortion) are related to the sensorimotor error signal.     

Evidence for distinct, differentially adaptable sensorimotor transformations for reaches to visual and proprioceptive targets

Recent evidence suggests that planning a reaching movement entails similar stages and common networks irrespective of whether the target location is defined through visual or proprioceptive cues. Here we test whether the transformations that convert the sensory information regarding target location into the required motor output are common for both types of reaches. To do so, we adaptively modified these sensorimotor transformations through exposure to displacing prisms and hypothesized that if they are common to both types of reaches, the aftereffects observed for reaches to visual targets would generalize to reaches to a proprioceptive target. Subjects (n = 16) were divided into two groups that differed with respect to the sensory modality of the targets (visual or proprioceptive) used in the pre- and posttests. The adaptation phase was identical for both groups and consisted of movements toward visual targets while wearing 10.5 degrees horizontally displacing prisms. We observed large aftereffects consistent with the magnitude of the prism-induced shift when reaching toward visual targets in the posttest, but no significant aftereffects for movements toward the proprioceptive target. These results provide evidence that distinct, differentially adaptable sensorimotor transformations underlie the planning of reaches to visual and proprioceptive targets.     

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.     

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).     

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     

Real-time error detection but not error correction drives automatic visuomotor adaptation

We investigated the role of visual feedback of task performance in visuomotor adaptation. Participants produced novel two degrees of freedom movements (elbow flexion–extension, forearm pronation–supination) to move a cursor towards visual targets. Following trials with no rotation, participants were exposed to a 60° visuomotor rotation, before returning to the non-rotated condition. A colour cue on each trial permitted identification of the rotated/non-rotated contexts. Participants could not see their arm but received continuous and concurrent visual feedback (CF) of a cursor representing limb position or post-trial visual feedback (PF) representing the movement trajectory. Separate groups of participants who received CF were instructed that online modifications of their movements either were, or were not, permissible as a means of improving performance. Feedforward-mediated performance improvements occurred for both CF and PF groups in the rotated environment. Furthermore, for CF participants this adaptation occurred regardless of whether feedback modifications of motor commands were permissible. Upon re-exposure to the non-rotated environment participants in the CF, but not PF, groups exhibited post-training aftereffects, manifested as greater angular deviations from a straight initial trajectory, with respect to the pre-rotation trials. Accordingly, the nature of the performance improvements that occurred was dependent upon the timing of the visual feedback of task performance. Continuous visual feedback of task performance during task execution appears critical in realising automatic visuomotor adaptation through a recalibration of the visuomotor mapping that transforms visual inputs into appropriate motor commands.     

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.     

Using arm configuration to learn the effects of gyroscopes and other devices

Previous studies have perturbed the association between motor commands and arm movements by applying forces to the arm during two-dimensional movements. These studies have revealed that, when the normal hand path is perturbed, subjects gradually adapt their motor commands to return to this path. The present study used the spin of a gyroscope to create a complex perturbation, as subjects reached to targets presented in three dimensions. Hand path did not change, but the whole-arm geometry ("arm configuration" in four dimensions) was altered. Over a series of several hundred reaches to various targets, subjects gradually returned the arm movement to its normal configuration. Furthermore, during the course of this learning, subjects used a strategy that involved manipulating arm posture. A similar strategy was observed when subjects made reaching movements with a rod attached to the upper arm to change its inertial characteristics. In both cases, the gradual return to the normal arm movement was accomplished without an increase in kinetic energy, suggesting that arm postures and movements (kinematics) and muscular forces (kinetics) may be mutually optimized. In contrast to previous studies, the present results highlight the role of arm configuration (rather than hand path) in learning and control.     

The effect of visuomotor adaptation on proprioceptive localization: the contributions of perceptual and motor changes

Reaching movements are rapidly adapted following training with rotated visual feedback of the hand (motor recalibration). Our laboratory has also found that visuomotor adaptation results in changes in estimates of felt hand position (proprioceptive recalibration) in the direction of the visuomotor distortion (Cressman and Henriques 2009, 2010; Cressman et al. 2010). In the present study, we included an additional method for measuring hand proprioception [specifically, proprioceptive-guided reaches of the unadapted (left) hand to the robot-guided adapted (right) hand-target] and compared this with our original perceptual task (estimating the felt hand position of the adapted hand relative to visual reference markers/the body midline), as well as to no-cursor reaches with the adapted hand (reaching to visual and midline-targets), to better identify whether changes in reaching following adaptation to a 50° rightward-rotated cursor reflect sensory or motor processes. Results for the proprioceptive estimation task were consistent with previous findings; subjects felt their hand to be aligned with a reference marker when it was shifted approximately 4° more in the direction of the visuomotor distortion following adaptation compared with baseline conditions. Moreover, we found similar changes in the proprioceptive-guided reaching task such that subjects misreached 5° in the direction of the cursor rotation. However, these results were true only for proprioceptive-guided reaches to the adapted hand, as reaches to the body midline were not affected by adaptation. This suggests that proprioceptive recalibration is restricted to the adapted hand and does not generalize to the rest of the body; this truly reflects a change in the sensory representation of the hand rather than changes in the motor program. This is in contrast to no-cursor reaches made with the adapted hand, which show reach after-effects for both visual targets and the midline, suggesting that reaches with the adapted hand reflect more of a change in the motor system. Our results also shed light on previous studies that may have misattributed these sensory and motor changes.     

Vestibular contribution to the planning of reach trajectories

Reaching for an object while simultaneously rotating induces Coriolis and centrifugal inertial forces on the arm that require compensatory actions to maintain accuracy. We investigated whether the nervous system uses vestibular signals of head rotation to predict inertial forces. Human subjects reached in darkness to a remembered target 33 cm distant. Subjects were stationary, but experienced a strong vestibular rotation signal. We achieved this by rotating subjects at 360°/s in yaw for 2 min and then stopping, and subjects reached during the ‘post-rotary’ period when the deceleration is interpreted by the vestibular system as a rotation in the opposite direction. Arm trajectories were straight in control trials without a rotary stimulus. With vestibular stimulation, trajectory curvature increased an average of 3 cm in the direction of the vestibular stimulation (e.g., to the right for a rightward yaw stimulus). Vestibular-induced curvature returned rapidly to normal, with an average time constant of 6 s. Movements also became longer as the vestibular stimulus diminished, and returned towards normal length with an average time constant of 5.6 s. In a second experiment we compared reaching with preferred and non-preferred hands, and found that they were similarly affected by vestibular stimulation. The reach curvatures were in the expected direction if the nervous system anticipated and attempted to counteract the presence of Coriolis forces based on the vestibular signals. Similarly, the shorter reaches may have occurred because the nervous system was attempting to compensate for an expected centrifugal force. Since vestibular stimulation also alters the perceived location of targets, vestibular signals probably influence all stages of the sensorimotor pathway transforming the desired goal of a reach into specific motor-unit innervation.     

Visuomotor adaptation and generalization with repeated and varied training

Many studies have shown that reaching movements to visual targets can rapidly adapt to altered visual feedback of hand motion (i.e., visuomotor rotation) and generalize to new target directions. This generalization is thought to reflect the acquisition of a neural representation of the novel visuomotor environment that is localized to the particular trained direction. In these studies, participants perform movements to a small number of target locations repeatedly. However, it is unclear whether adaptation and generalization are comparable when target locations are constantly varied and participants reach to visual targets one time only. Here, we compared performance for reaches to a 30° counter-clockwise visuomotor rotation to four targets, spaced 90° apart across four areas of workspace 18 times each (repeated practice (RP)) with one time only reaching movements to 72 targets, spaced 5° apart (varied practice (VP)). For both training groups, participants performed 18 reaches to radial targets (either at the repeated or varied location) in a specific area of the workspace (i.e., one of four quadrants) before reaching in the adjacent workspace. We found that the RP group adapted more completely compared to the VP group. Conversely, the VP group generalized to new target directions more completely when reaching without cursor feedback compared to the RP group. This suggests that RP and VP follow a mainly common pattern of adaptation and generalization represented in the brain, with benefits of faster adaptation with RP and more complete generalization with VP.     

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.     

Hand and joint paths during reaching movements with and without vision

 This study examines whether the kinematics of pointing movements are altered by the sensory systems used to select spatial targets and to guide movement. Hand and joint paths of visually guided reaching movements of human subjects were compared with two non-visual conditions where only proprioception was available: (1) movements of the same subjects with blindfolds, and (2) movements by congenitally blind subjects. While hand-path curvatures were overall quite small, sighted subjects wearing a blindfold showed a statistical increase in hand-path curvature compared with their visually guided movements. Blindfolded subjects also showed greater hand-path curvature than blind subjects. These increases in hand-path curvature for blindfolded subjects did not always lead to a decrease in joint-path curvature. While there were differences between blind subjects and sighted subjects using vision for some movement directions, there was no systematic difference between these two groups. The magnitude of joint-path curvature showed much greater variation than hand-path curvature across the movement directions. We found variation in joint-path curvature to be correlated to two factors, one spatial and one geometrical. For all subject groups, joint-path curvature tended to be smaller for sagittal-plane movements than for transverse or diagonal movements. As well, we found that the magnitude of joint-path curvature was also related to the relative motion at each joint. Joint-path curvature tended to increase when movements predominantly involved changes in shoulder angle and was minimal when movements predominantly involved elbow motion. The consistently small curvatures of hand trajectory across blind and sighted subjects emphasize the powerful tendency of the motor system to generate goal-directed reaching movements with relatively straight hand trajectories, even when deprived of visual feedback from very early in life. Received: 16 July 1997 / Accepted: 20 May 1998     

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.     

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.     

Reaching to ipsilateral or contralateral targets: within-hemisphere visuomotor processing cannot explain hemispatial differences in motor control

Aiming movements made to visual targets on the same side of the body as the reaching hand typically show advantages as compared to aiming movements made to targets on the opposite side of the body midline in the contralateral visual field. These advantages for ipsilateral reaches include shorter reaction time, higher peak velocity, shorter duration and greater endpoint accuracy. It is commonly hypothesized that such advantages are related to the efficiency of intrahemispheric processing, since, for example, a left-sided target would be initially processed in the visual cortex of the right hemisphere and that same hemisphere controls the motor output to the left hand. We tested this hypothesis by examining the kinematics of aiming movements made by 26 right-handed subjects to visual targets briefly presented in either the left or the right visual field. In one block of trials, the subjects aimed their finger directly towards the target; in the other block, subjects were required to aim their movement to the mirror symmetrical position on the opposite side of the fixation light from the target. For the three kinematic measures in which hemispatial differences were obtained (peak velocity, duration and percentage of movement time spent in deceleration), the advantages were related to the side to which the motor response was directed and not to the side where the target was presented. In addition, these effects tended to be larger in the right hand than in the left, particularly for the percentage of the movement time spent in deceleration. The results are interpreted in terms of models of biomechanical constraints on contralateral movements, which are independent of the hemispace of target presentation.