The extent of interlimb transfer following adaptation to a novel visuomotor condition does not depend on awareness of the condition

There is a controversy in the literature as to whether transfer of motor learning across the arms occurs because of an individual's cognitive awareness of the learned condition. The purpose of this study was to test whether the extent of interlimb transfer following adaptation to a novel visuomotor rotation with one arm, as well as the rate of learning acquired by the other arm, would vary depending on the subjects' awareness of the rotation condition. Awareness of the condition was varied by employing three experimental conditions. In one condition, visual rotation of the display up to 32° was gradually introduced to minimize the subjects' awareness of the rotation during targeted reaching movement. In another condition, the 32° rotation was abruptly introduced from the beginning of the adaptation session. Finally, the subjects were informed regarding the rotation prior to the adaptation session. After adaptation with the left arm under the three conditions, subjects performed reaching movement with the right arm under the same 32° rotation condition. Our results showed that the amount of initial transfer, and also the changes in performance with the right arm, did not vary significantly across the three conditions. This finding suggests that interlimb transfer of visuomotor adaptation does not occur based on an individual's awareness of the manipulation, but rather as a result of implicit generalization of the obtained visuomotor transformation across the arms.     

Evidence for a dynamic-dominance hypothesis of handedness

Handedness is a prominent behavioral phenomenon that emerges from asymmetrical neural organization of human motor systems. However, the aspects of motor performance that correspond to handedness remain largely undetermined. A recent study examining interlimb differences in coordination of reaching demonstrated dominant arm advantages in controlling limb segment inertial dynamics (Sainburg and Kalakanis 2000). Based on these findings, I now propose the dynamic-dominance hypothesis, which states that the essential factor that distinguishes dominant from nondominant arm performance is the facility governing the control of limb dynamics. The purpose of this study is to test two predictions of this hypothesis: 1) adaptation to novel intersegmental dynamics, requiring the development of new dynamic transforms, should be more effective for the dominant arm; 2) there should be no difference in adapting to visuomotor rotations performed with the dominant as compared with the nondominant arm. The latter prediction is based on the idea that visual information about target position is translated into an internal reference frame prior to transformation of the movement plan into dynamic properties, which reflect the forces required to produce movement. To test these predictions, dominant arm adaptation is compared to nondominant arm adaptation during exposure to novel inertial loads and to novel visuomotor rotations. The results indicate substantial interlimb differences in adaptation to novel inertial dynamics, but equivalent adaptation to novel visuomotor rotations. Inverse dynamic analysis revealed better coordination of dominant arm muscle torques across both shoulder and elbow joints, as compared with nondominant arm muscle torques. As a result, dominant arm movements were produced with a fraction of the mean squared muscle torque computed for nondominant arm movements made at similar speeds. These results support the dynamic-dominance hypothesis, indicating that interlimb asymmetries in control arise downstream to visuomotor transformations, when dynamic variables that correspond to the forces required for motion are specified.     

Visuomotor learning generalizes between bilateral and unilateral conditions despite varying degrees of bilateral interference

Bilateral interference, referring to the tendency of movements of one arm to disrupt the intended movements made simultaneously with the other arm, is often observed in a task that involves differential planning of each arm movement during sensorimotor adaptation. In the present study, we examined two questions: 1) how does the compatibility between visuomotor adaptation tasks performed with both arms affect bilateral interference during bimanual performance? and 2) how do variations in bilateral interference affect transfer of visuomotor adaptation between bilateral and unilateral conditions? To examine these questions, we manipulated visuomotor compatibility using two kinematic variables (direction of required hand motion, direction of an imposed visual rotation). Experiment 1 consisted of two conditions in which the direction of visual rotations for both arms was either in the same or opposing directions, whereas the target direction for both arms was always the same. In experiment 2, we examined the pattern of generalization between the bilateral and unilateral conditions when both the target and rotation directions were opposing between the arms. In both experiments, subjects first adapted to a 30° visual rotation with one arm (preunilateral), then with both arms (bilateral), and finally with the arm that was not used in the first session (postunilateral). Our results show that bilateral interference was smallest when both variables were the same between the arms. Our data also show extensive transfer of visuomotor adaptation between bilateral and unilateral conditions, regardless of degree of bilateral interference.     

Effects of roll visual motion on online control of arm movement: reaching within a dynamic virtual environment

Reaching toward a visual target involves the transformation of visual information into appropriate motor commands. Complex movements often occur either while we are moving or when objects in the world move around us, thus changing the spatial relationship between our hand and the space in which we plan to reach. This study investigated whether rotation of a wide field-of-view immersive scene produced by a virtual environment affected online visuomotor control during a double-step reaching task. A total of 20 seated healthy subjects reached for a visual target that remained stationary in space or unpredictably shifted to a second position (either to the right or left of its initial position) with different inter-stimulus intervals. Eleven subjects completed two experiments which were similar except for the duration of the target’s appearance. The final target was either visible throughout the entire trial or only for a period of 200 ms. Movements were performed under two visual field conditions: the virtual scene was matched to the subject’s head motion or rolled about the line of sight counterclockwise at 130°/s. Nine additional subjects completed a third experiment in which the direction of the rolling scene was manipulated (i.e., clockwise and counterclockwise). Our results showed that while all subjects were able to modify their hand trajectory in response to the target shift with both visual scenes, some of the double-step movements contained a pause prior to modifying trajectory direction. Furthermore, our findings indicated that both the timing and kinematic adjustments of the reach were affected by roll motion of the scene. Both planning and execution of the reach were affected by roll motion. Changes in proportion of trajectory types, and significantly longer pauses that occurred during the reach in the presence of roll motion suggest that background roll motion mainly interfered with the ability to update the visuomotor response to the target displacement. Furthermore, the reaching movement was affected differentially by the direction of roll motion. Subjects demonstrated a stronger effect of visual motion on movements taking place in the direction of visual roll (e.g., leftward movements during counterclockwise roll). Further investigation of the hand path revealed significant changes during roll motion for both the area and shape of the 95% tolerance ellipses that were constructed from the hand position following the main movement termination. These changes corresponded with a hand drift that would suggest that subjects were relying more on proprioceptive information to estimate the arm position in space during roll motion of the visual field. We conclude that both the spatial and temporal kinematics of the reach movement were affected by the motion of the visual field, suggesting interference with the ability to simultaneously process two consecutive stimuli.     

Cerebral cortical dynamics during visuomotor transformation: adaptation to a cognitive-motor executive challenge

EEG was employed during cognitive-motor adaptation to a visuomotor transformation that required inhibition of an established motor plan. Performance was positively related to frontal alpha and theta power during both planning and execution of reaching movements to visual targets. EEG changes suggest initial involvement of frontal executive functioning to suppress established visuomotor mappings followed by progressive idling (i.e., alpha synchrony). Also, progressive idling of the temporal and parietal sites over the trials was observed, suggesting a decreasing role of working memory and encoding of the new visuomotor map, respectively. The regional changes in the cortical dynamics translated into the quality of motor behavior. This study expands our understanding of the role of frontal executive processes beyond the cognitive domain to the cognitive-motor domain.     

Multimodal reference frame for the planning of vertical arms movements

In this study we investigated the reference frames used to plan arm movements. Specifically, we asked whether the body axis, visual cues and graviception can each play a role in defining "up" and "down" in the planning and execution of movements along the vertical axis. Horizontal and vertical pointing movements were tested in two postures (upright and reclined) and two visual conditions (with and without vision) to identify possible effects of each of these cues on kinematics of movement. Movements were recorded using an optical 3D tracking system and analysis was conducted on velocity profiles of the hand. Despite a major effect of gravity, our analysis shows an effect of the movement direction with respect to the body axis when subjects were reclined with eyes closed. These results suggest that our CNS takes into account multimodal information about vertical in order to compute an optimal motor command that anticipates the effects of gravity.     

Clonidine as adjuvant for oxybuprocaine, bupivacaine or dextrorphan has a significant peripheral action in intensifying and prolonging analgesia in response to local dorsal cutaneous noxious pinprick in rats

The roles of visual and somatosensory information in arm movement planning remain enigmatic. Previous studies have examined these roles by dissociating visual and somatosensory cues about limb position prior to movement onset and examining the resulting effects on movements performed in the horizontal plane. Here we examined the effects of misaligned limb position cues prior to movement onset as reaches were planned and executed along different directions in the vertical plane. Movements were planned with somatosensory and visual feedback aligned at the starting position of the reach or with visual feedback displaced horizontally (Experiment 1) or vertically (Experiment 2). As in the horizontal plane, changes in movement directions induced by misaligned feedback indicated that vision and proprioception were both generally taken into account when planning vertical plane movements. However, we also found evidence that the contributions of vision and proprioception differed across target directions and between directions of displaced visual feedback. These findings suggest that the contributions of vision and proprioception to movement planning in the vertical plane reflect the unique multisensory and biomechanical demands associated with moving against gravity.     

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.     

Effect of visuomotor-map uncertainty on visuomotor adaptation

Vision and proprioception contribute to generating hand movement. If a conflict between the visual and proprioceptive feedback of hand position is given, reaching movement is disturbed initially but recovers after training. Although previous studies have predominantly investigated the adaptive change in the motor output, it is unclear whether the contributions of visual and proprioceptive feedback controls to the reaching movement are modified by visuomotor adaptation. To investigate this, we focused on the change in proprioceptive feedback control associated with visuomotor adaptation. After the adaptation to gradually introduce visuomotor rotation, the hand reached the shifted position of the visual target to move the cursor to the visual target correctly. When the cursor feedback was occasionally eliminated (probe trial), the end point of the hand movement was biased in the visual-target direction, while the movement was initiated in the adapted direction, suggesting the incomplete adaptation of proprioceptive feedback control. Moreover, after the learning of uncertain visuomotor rotation, in which the rotation angle was randomly fluctuated on a trial-by-trial basis, the end-point bias in the probe trial increased, but the initial movement direction was not affected, suggesting a reduction in the adaptation level of proprioceptive feedback control. These results suggest that the change in the relative contribution of visual and proprioceptive feedback controls to the reaching movement in response to the visuomotor-map uncertainty is involved in visuomotor adaptation, whereas feedforward control might adapt in a manner different from that of the feedback control.     

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.     

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.     

Preferred directions of arm movements are independent of visual perception of spatial directions

Directional preferences have previously been demonstrated during horizontal arm movements. These preferences were characterized by a tendency to exploit interaction torques for movement production at the shoulder or elbow, indicating that the preferred directions depend on biomechanical, and not on visual perception-based factors. We directly tested this hypothesis by systematically dissociating visual information from arm biomechanics. Sixteen subjects performed a free-stroke drawing task that required performance of fast strokes from the circle center toward the perimeter, while selecting stroke directions in a random order. Hand position was represented by a cursor displayed in the movement plane. The free-stroke drawing was performed twice, before and after visuomotor adaptation to a 30° clockwise rotation of the perceived hand path. The adaptation was achieved during practicing pointing movements to eight center-out targets. Directional preferences during performance of the free-stroke drawing task were revealed in ten out of the sixteen subjects. The orientation and strength of these preferences were largely the same in both conditions, showing no significant effect of the visuomotor adaptation. In both conditions, the major preferred directions were characterized by higher contribution of interaction torque to net torque at the shoulder as well as by relatively low inertial resistance and the sum of squared shoulder and elbow muscle torques. These results support the hypothesis that directional preferences are largely determined by biomechanical factors. However, this biomechanical effect can decrease or even disappear in some subjects when movements are performed in special conditions, such as the virtual environment used here.     

Motor planning of arm movements is direction-dependent in the gravity field

In the present study we analyzed kinematic and dynamic features of arm movements in order to better elucidate how the motor system integrates environmental constraints (gravity) into motor planning and control processes. To reach this aim, we experimentally manipulated the mechanical effects of gravity on the arm while maintaining arm inertia constant (i.e. the distribution of the mass around the shoulder joint). Six subjects performed single-joint arm movements (rotation around the shoulder joint) in both sagittal (upward, U, versus downward, D) and horizontal (left, L, versus right, R) planes, at different amplitudes and from different initial positions. Under these conditions, shoulder gravitational torques (SGTs) significantly varied when arm movements were performed in the sagittal but not in the horizontal plane. Contrary to SGTs, arm inertia remained constant and similar for both horizontal and sagittal planes since subjects performed arm movements with only one degree of freedom. All subjects, whatever the movement direction, appropriately scaled shoulder joint kinematic parameters according to movement amplitude. Furthermore, peak velocity and movement duration were equivalent for both horizontal and sagittal planes. Interestingly, some kinematic parameters significantly differed according to U/D but not L/R directions. Specifically, acceleration duration was greater for D than U movements, while the opposite was true for peak acceleration. Consequently, although vertical and horizontal arm movements shared a general common strategy (i.e. scaling law), the kinematic asymmetries between U and D arm movements, especially those that reflect central planning process (i.e. peak acceleration), indicated different motor intentions regarding the direction of the upcoming movement. These findings indicate that the interaction of the arm with the dynamics of the environment is internally represented during the generation of arm trajectories.     

Interlimb transfer of visuomotor rotations: independence of direction and final position information

Previous findings from our laboratory support the idea that the dominant arm is more proficient than the non-dominant arm in coordinating intersegmental dynamics for specifying trajectory direction and shape during multijoint reaching movements. We also showed that adaptation of right and left arms to novel visuomotor rotations was equivalent, suggesting that this process occurs upstream to processes that distinguish dominant and non-dominant arm performance. Because of this, we speculate that such visuomotor adaptations might transfer to subsequent performance during adaptation with the other arm. We now examine whether opposite arm training to novel visuomotor rotations transfers to affect adaptation using the right and left arms. Two subject groups, RL and LR, each comprising seven right-handed subjects, adapted to a 30 degrees counterclockwise rotation in the visual display during a center-out reaching task performed in eight directions. Each group first adapted using either the right (RL) or left (LR) arm, followed by opposite arm adaptation. In order to assess transfer, we compared the same side arm movements (either right or left) following opposite arm adaptation to those performed prior to opposite arm adaptation. Our findings indicate unambiguous transfer of learning across the arms. Different features of movement transferred in different directions: Opposite arm training improved the initial direction of right arm movements under the rotated visual condition, whereas opposite arm training improved the final position accuracy, but not the direction of left arm movements. These findings confirm that transfer of training was not due to a general cognitive strategy, since such an effect should influence either hand equally. These findings support the hypothesis that each arm controller has access to information learned during opposite arm training. We suggest that each controller uses this information differently, depending on its proficiency for specifying particular features of movement. We discuss evidence that these two aspects of control are differentially mediated by the right and left cerebral hemispheres.     

Task-specific internal models for kinematic transformations

Numerous studies of motor learning have focused on how people adapt their reaching movements to novel dynamic and visuomotor perturbations that alter the actual or visually perceived motion of the hand. An important finding from this work is that learning of novel dynamics generalizes across different movement tasks. Thus adaptation to an unusual force field generalizes from center-out reaching movements to circular movements (Conditt et al. 1997). This suggests that subjects acquired an internal model of the dynamic environment that could be used to determine the motor commands needed for untrained movements. Using a task interference paradigm, we investigated whether transfer across tasks is also observed when learning visuomotor transformations. On day 1, all subjects adapted to a +30 degrees rotation while making center-out-and-back reaching movements. After a delay of 5 min, different groups of subjects then adapted to a -30 degrees rotation while performing either a continuous tracking task, a figure-eight drawing task, or the center-out-and-back reaching task. All subjects were then retested the next day on the +30 degrees rotation in the reaching task. As expected, subjects who experienced the opposing rotations while performing the same reaching tasks showed no retention of learning for the first rotation when tested on day 2 (Krakauer et al. 1999). In contrast, such retrograde interference was not observed in the two groups of subjects who experienced the opposing rotations while performing different tasks. In fact, their performance on day 2 was similar to that of control subjects who never experienced the opposite rotation. This lack of interference suggests that memory resources for visuomotor rotations are task specific.     

The role of proprioception and attention in a visuomotor adaptation task

The role of proprioception in the control and adaptation of visuomotor relationships is still unclear. We have studied a deafferented subject, IW, and control subjects in a task in which they used single joint elbow extension to move to a visual target, with visual feedback of the terminal position provided by a cursor displayed in the plane of their movements. We report the differences in movement accuracy between the deafferented subject and controls in the normal task and when challenged with a cognitive load, counting backwards. All subjects were less accurate when counting; this was a small effect for the controls (<10% change) but much greater for the deafferented subject (>60% change). We also examined changes in movement kinematics when the instructed amplitude was altered via a changed gain between final arm position and presentation of the feedback cursor. The deafferented subject maintained temporal movement parameters stable and altered amplitude by scaling force (i.e. changed peak velocity), whereas the controls scaled both movement velocity and duration. Finally, we compared the subjects' adaptation of movement amplitude after a period of exposure to the changed visuomotor gain. The deafferented subject was able to adapt, but his adaptation was severely impaired by the counting task. These results suggest that proprioception is not an absolute requirement for adaptation to occur. Instead, proprioception has a more subtle role to play in the adjustment to visuomotor perturbations. It has an important role in the control of reaching movements, while in the absence of proprioception, attention appears necessary to monitor movements.     

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.     

Abnormal Purkinje cell activity in vivo in experimental allergic encephalomyelitis

It has been shown that learning visuomotor rotations with multiple target directions, compared with a single target direction, leads to greater generalization to untrained targets within the same limb. This implies that multiple direction learning results in a more complete internal model of the visuomotor transform. It has also been documented that the extent of transfer of movement information regarding visuomotor adaptations between the limbs is limited, relative to that between different configurations of the same limb. The present study thus investigated the origin of this restriction in interlimb transfer, by comparing the effects of eight-direction and one-direction training conditions with one arm on the subsequent performance with the other arm. It was hypothesized that if multiple direction learning leads to a more complete model of the novel visuomotor transform, interlimb transfer should be enhanced relative to that following single direction training. However, if no differences are observed between single and multiple direction training conditions, this would suggest that such learning is effector dependent. We also tested the hypothesis that interlimb transfer of visuomotor adaptation is not obligatory, by examining the effects of visual rotation direction (same or oppositely directed visuomotor rotations for the two arms). All subjects first adapted to a 30° rotation, either clockwise or counterclockwise, in the visual display during reaching movements. Following this, they adapted to a 30° rotation in either the same or opposing direction with the other arm. Results showed that initial training with the non-dominant arm facilitated subsequent performance with the dominant arm in terms of initial direction control, but only under the same rotation condition. Both single and eight direction training conditions led to substantial transfer in subsequent performance with the other arm, but multiple direction training was no more beneficial than single direction training. This finding suggests that the previously reported intralimb advantages of multiple direction learning are effector specific. Our findings are discussed in the context of hierarchical models of motor control to explain the intralimb advantages of multiple direction training.     

A dissociation between visual and motor workspace inhibits generalization of visuomotor adaptation across the limbs

We have previously shown that the pattern of interlimb transfer following visuomotor adaptation depends on whether the two arms share task-space at a given workspace location: when the two arms adapted to a novel visuomotor rotation in unshared, lateral workspaces, transfer of movement direction information occurred symmetrically (i.e., from dominant to nondominant arm, and vice versa). When the two arms shared the same task-space, however, transfer of the same information became asymmetric (i.e., only from dominant to nondominant arm). In the present study, I investigated the effect of a conflict between visual and proprioceptive information of task-space on the pattern of interlimb transfer, by dissociating visual and motor workspaces. I hypothesized that the pattern of interlimb transfer would be determined by the way the motor control system uses available sensory information, and predicted that depending on whether the system relied more on vision or proprioception, transfer would occur either symmetrically or asymmetrically. Surprisingly, the results indicated that despite substantial adaptation to a novel visuomotor rotation, no transfer occurred across the arms when the visual and motor workspaces were dissociated in space. Based on this finding, I suggest that when a conflict exists between visual and proprioceptive information with respect to the sharing of the given task-space by the two arms, it interferes with executive decisions made by the motor control system in determining hand dominance at a given workspace, which results in a lack of transfer across the arms.     

Sensory prediction errors drive cerebellum-dependent adaptation of reaching

The cerebellum is an essential part of the neural network involved in adapting goal-directed arm movements. This adaptation might rely on two distinct signals: a sensory prediction error or a motor correction. Sensory prediction errors occur when an initial motor command is generated but the predicted sensory consequences do not match the observed values. In some tasks, these sensory errors are monitored and result in on-line corrective motor output as the movement progresses. Here we asked whether cerebellum-dependent adaptation of reaching relies on sensory or on-line motor corrections. Healthy controls and people with hereditary cerebellar ataxia reached during a visuomotor perturbation in two conditions: "shooting" movements without on-line corrections and "pointing" movements that allowed for on-line corrections. Sensory (i.e., visual) errors were available in both conditions. Results showed that the addition of motor corrections did not influence adaptation in control subjects, suggesting that only sensory errors were needed for learning. Cerebellar subjects were comparably impaired in both adaptation conditions relative to controls, despite abnormal and inconsistent on-line motor correction. Specifically, poor on-line motor corrections were unrelated to cerebellar subjects' adaptation deficit (i.e., adaptation did not worsen), further suggesting that only sensory prediction errors influence this process. Therefore adaptation to visuomotor perturbations depends on the cerebellum and is driven by the mismatch between predicted and actual sensory outcome of motor commands.