Hand trajectory invariance in reaching movements involving the trunk

Movements of different body segments may be combined in different ways to achieve the same motor goal. How this is accomplished by the nervous system was investigated by having subjects make fast pointing movements with the arm in combination with a forward bending of the trunk that was unexpectedly blocked in some trials. Subjects moved their hand above the surface of a table without vision from an initial position near the midline of the chest to remembered targets placed within the reach of the arm in either the ipsi- or contralateral workspace. In experiment 1, subjects were instructed to make fast arm movements to the target without corrections whether or not the trunk was arrested. Only minor changes were found in the hand trajectory and velocity profile in response to the trunk arrest, and these changes were seen only late in the movement. In contrast, the patterns of the interjoint coordination substantially changed in response to the trunk arrest, suggesting the presence of compensatory arm-trunk coordination minimizing the deflections from the hand trajectory regardless of whether the trunk is recruited or mechanically blocked. Changes in the arm interjoint coordination in response to the trunk arrest could be detected kinematically at a minimal latency of 50 ms. This finding suggests a rapid reflex compensatory mechanism driven by vestibular and/or proprioceptive afferent signals. In experiment 2, subjects were required, as soon as they perceived the trunk arrest, to change the hand motion to the same direction as that of the trunk. Under this instruction, subjects were able to initiate corrections only after the hand approached or reached the final position. Thus, centrally mediated compensatory corrections triggered in response to the trunk arrest were likely to occur too late to maintain the observed invariant hand trajectory in experiment 1. In experiment 3, subjects produced similar pointing movements, but to a target that moved together with the trunk. In these body-oriented pointing movements, the hand trajectories from trials in which the trunk was moving or arrested were substantially different. The same trajectories represented in a relative frame of reference moving with the trunk were virtually identical. We conclude that hand trajectory invariance can be produced in an external spatial (experiment 1) or an internal trunk-centered (experiment 3) frame of reference. The invariance in the external frame of reference is accomplished by active compensatory changes in the arm joint angles nullifying the influence of the trunk motion on the hand trajectory. We suggest that to make a transition to the internal frame of reference, control systems suppress this compensation. One of the hypotheses opened to further experimental testing is that the integration of additional (trunk) degrees of freedom into movement is based on afferent (proprioceptive, vestibular) signals stemming from the trunk motion and transmitted to the arm muscles.     

Superposition of independent units of coordination during pointing movements involving the trunk with and without visual feedback

Previous studies addressing the problem of the control of multiple degrees of freedom have examined the influence of trunk movement on pointing movements within the arm's reach. Such movements may be controlled by two functionally independent units of coordination (synergies): one involving only arm joints and producing the hand trajectory to the target (the transport synergy), and the other coordinating trunk and arm movements leaving the hand trajectory unchanged (the compensatory synergy). The question of whether or not this functional subdivision depends on visual feedback was addressed in the present study. We also tested whether or not the motor effects of different synergies are summated as independent components, a control strategy called "superposition." Finally, we investigated whether or not the relationship between different degrees of freedom within each synergy could be considered linear resulting in proportional changes in different joint angles. Seated subjects produced fast, uncorrected arm movements to an ipsi- or a contralateral target in the direction of +/-45 degrees to the sagittal midline of the trunk. Targets could be reached using the arm alone (control trials) or by combining the arm motion with a forward or backward trunk motion produced by hip flexion or extension (test trials), with and without visual feedback. The shape of the hand trajectory, its direction and tangential velocity, movement precision, joint angles and the sequence of the trunk and hand recruitment and de-recruitment were measured. In both visual conditions, the direction of the hand trajectory observed in control trials was generally preserved in test trials. In terms of sequencing, even in the absence of vision, the trunk movement was initiated before the onset of and outlasted the hand shift, indicating that the potential influence of the trunk on the hand movement was compensated by rotations in the elbow and shoulder joint. The analysis of other variables also implied that the effects of trunk recruitment on the hand trajectory were minor compared to those which could be observed if these effects were not compensated by appropriate changes in the arm joint angles. It was concluded that an arm-trunk compensatory synergy is present in pointing movements regardless of visual feedback. Principal component analysis showed that the relationship between elbow, shoulder and hip joint angles in individual arm and combined arm-trunk movements cannot be considered linear, implying that this relationship is adjusted according to the changing arm geometry. The changes in each arm joint angle (elbow, shoulder) elicited by a forward trunk bending in one block of trials were compared with those elicited by a backward bending in another block, whereas the hand moved to the same target in both blocks. These changes were opposite but of similar magnitude. As a result, for each moment of movement, the mean joint angle obtained by averaging across two directions of trunk motion was practically identical to that in control trials in which the trunk was motionless. It is concluded that the transport and arm-trunk compensatory synergies are combined as independent units, according to the principle of superposition. This principle may simplify the control of the coordination of a redundant number of degrees of freedom.     

Coordination deficits during trunk-assisted reach-to-grasp movements in Parkinson’s disease

The present study investigated how Parkinson’s disease (PD) affects temporal coordination among the trunk, arm, and fingers during trunk-assisted reach-to-grasp movements. Seated participants with PD and healthy controls made prehensile movements. During the reach to the object, the involvement of the trunk was altered based on the instruction; the trunk was not involved, moved forward (flexion), or moved backward (extension) in the sagittal plane. Each of the trunk movements was combined with an extension or flexion motion of the arm during the reach. For the transport component, the individuals with PD substantially delayed the onset of trunk motion relative to that of arm motion in conditions where the trunk was moved in the direction opposite from the arm reaching toward the object. At the same time, variability of intervals between the onsets and intervals between the velocity peaks of the trunk and wrist movements was increased. The magnitudes of the variability measures were significantly correlated with the severity of PD. Regarding the grasp component, the individuals with PD delayed the onset of finger movements during reaching. These results imply that PD impairs temporal coordination between the axial and distal body segments during goal-directed skilled actions. When there is a directional discrepancy between the trunk and wrist motions, individuals with PD appear to prioritize wrist motion that is tied to the task goal over the trunk motion. An increase in disease severity magnifies the coordination deficits.     

Dystonia is predictive of subsequent altered dopaminergic responsiveness in a chronic 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine+3-nitropropionic acid model of striatonigral degeneration in monkeys

Using galvanic vestibular stimulation (GVS), we tested whether a change in vestibular input at the onset of goal-directed arm movements induces deviations in arm trajectory. Eight head-fixed standing subjects were instructed to reach for memorized visual targets in complete darkness. In half of the trials, randomly-selected, a 3 mA bipolar binaural galvanic stimulation of randomly alternating polarity was triggered by the movement onset. Results revealed significant GVS-induced directional shifts of reaching movements towards the anode side. The earliest significant deviations of hand path occurred 240 ms after stimulation onset. The likely goal of these online deviations of arm trajectory was to compensate for a vestibular-evoked apparent change in the spatial relationship between the target and the hand.     

The coordination between trunk and arm motion during pointing movements

The coordination between the trunk and arm of six subjects was examined during unrestrained pointing movements to five target locations. Two targets were within arm's length, three were beyond. The trunk participated in reaching primarily when the target could not be attained by arm and scapular motion. When the trunk did contribute to hand transport, its motion started simultaneously with arm movement and continued until target contact. Redundancy in the degrees of freedom used to execute the movement had no effect on the configuration of joints and segments used to attain a specified target; no difference in variability was noted regardless of whether redundancy existed. However, different configurations were used to achieve the same wrist coordinates along a common endpoint path, depending on the final position of the hand. The addition of trunk flexion, rotation and scapular motion did not alter the coupling between the elbow and shoulder joints and had no effect on the path of the hand or the smoothness of its velocity profile. Thus, trunk motion was integrated smoothly into the transport phase of the hand. As the trunk's contribution to hand transport increased, it played a progressively greater role in positioning the hand close to the target during the terminal stage of the reach. Of the movement components measured, trunk flexion was the last component to complete its motion when target reaches were made beyond arm's length. Hence, the trunk not only acts as a postural stabilizer during reaching, but becomes an integral component in positioning the hand close to the target.     

Coordination between equilibrium and hand trajectories during whole body pointing movements

We examined the coordination between equilibrium and voluntary pointing movements executed from the standing position, using the whole body. It has previously been shown that trunk movement has little effect upon kinematic characteristics of hand pointing when movements are executed in the sitting position. The present study asked if elements of hand trajectory are modified by requirements of large trunk displacements and fine equilibrium control when pointing movements are executed from the standing position. To achieve this, center of pressure (CoP) and center of mass (CoM) displacements were analyzed along with the kinematics of the pointing hand. Results showed that the CoM was not stabilized (it displaced between 23% and 61+/-21% of the foot's length), confirming that instead of a compensation of mechanical perturbations due to arm and trunk movements, the present equilibrium strategy consisted of controlling CoM acceleration towards the target. Hand paths were curved and were not distance or speed invariant. Rather than simple inefficiencies in programming or execution, path curvature suggested that different hand movement strategies were chosen as a function of equilibrium constraints. In light of these results, we hypothesize that postural stability may play a role in the generation of hand trajectory for complex, whole-body pointing movements, in addition to constraints placed upon end-effector kinematics or the dynamic optimization of upper-limb movements. A dependent regulation of equilibrium and spatial components of the movement is proposed.     

Arm–trunk coordination in the absence of proprioception

During trunk-assisted reaching to targets placed within arms length, the influence of trunk motion on the hand trajectory is compensated for by changes in the arm configuration. The role of proprioception in this compensation was investigated by analyzing the movements of 2 deafferented and 12 healthy subjects. Subjects reached to remembered targets (placed ~80° ipsilateral or ~45° contralateral to the sagittal midline) with an active forward movement of the trunk produced by hip flexion. In 40% of randomly selected trials, trunk motion was mechanically blocked. No visual feedback was provided during the experiment. The hand trajectory and velocity profiles of healthy subjects remained invariant whether or not the trunk was blocked. The invariance was achieved by changes in arm interjoint coordination that, for reaches toward the ipsilateral target, started as early as 50 ms after the perturbation. Both deafferented subjects exhibited considerable, though incomplete, compensation for the effects of the perturbation. Compensation was more successful for reaches to the ipsilateral target. Both deafferented subjects showed invariance between conditions (unobstructed or blocked trunk motion) in their hand paths to the ipsilateral target, and one did to the contralateral target. For the other deafferented subject, hand paths in the two types of trials began to deviate after about 50% into the movement, because of excessive elbow extension. In movements to the ipsilateral target, when deafferented subjects compensated successfully, the changes in arm joint angles were initiated as early as 50 ms after the trunk perturbation, similar to healthy subjects. Although the deafferented subjects showed less than ideal compensatory control, they compensated to a remarkably large extent given their complete loss of proprioception. The presence of partial compensation in the absence of vision and proprioception points to the likelihood that not only proprioception but also vestibulospinal pathways help mediate this compensation.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Deficits in adaptive upper limb control in response to trunk perturbations in Parkinson’s disease

The ability of patients with Parkinsons disease (PD) to compensate for unexpected perturbations remains relatively unexplored. To address this issue PD subjects were required to compensate at the arm for an unexpected mechanical perturbation of the trunk while performing a trunk-assisted reach. Twelve healthy and nine PD subjects (off medication) performed trunk-assisted reaching movements without vision or knowledge of results to a remembered target in the ipsilateral (T1) or contralateral (T2) workspace. On 60% of the trials trunk motion was unrestrained (free condition). On the remaining 40% of randomly selected trials trunk motion was arrested at movement onset (blocked condition). If subjects appropriately changed arm joint angles to compensate for the trunk arrest, there should be spatial and temporal invariance in the hand trajectories and in the endpoint errors across conditions. The control group successfully changed their arm configuration in a context-dependent manner which resulted in invariant hand trajectory profiles across the free and blocked conditions. More so, they initiated these changes rapidly after the trunk perturbation (group mean 70 ms). Some PD subjects were unable to maintain invariant hand paths and movement errors across conditions. Their hand velocity profiles were also more variable relative to those of the healthy subjects in the blocked-trunk trials but not in the free-trunk trials. Furthermore, the latency of compensatory changes in arm joint angles in movements toward T1 was longer in the PD group (group mean 153 ms). Finally, PD subjects arm and trunk were desynchronized at movement onset, confirming our previous findings and consistent with PD patients known problems in the sequential or parallel generation of different movement components. The findings that individual PD subjects were unsuccessful or delayed in producing context-dependent responses at the arm to unexpected perturbations of the trunk suggests that the basal ganglia are important nodes in the organization of adaptive behavior.     

Vestibular system may provide equivalent motor actions regardless of the number of body segments involved in the task

The vestibulospinal system likely plays an essential role in motor equivalence—the ability to reach the desired motor goal despite intentional or imposed changes in the number of body segments involved in the task. To test this hypothesis, we compared the ability of healthy subjects and patients with unilateral vestibular lesions (surgical acoustic neuroma resection 0.6 to 6.7 yr before the study) to maintain either the same hand position or the same trajectory of within arm reach movements while flexing the trunk, in the absence of vision. In randomly selected trials, the trunk motion was prevented by an electromagnetic device. Healthy subjects were able to preserve the hand position or trajectory by modifying the elbow and shoulder joint rotations in a condition-dependent way, at a minimal latency of about 60 ms after the trunk movement onset. In contrast, six of seven patients showed deficits in the compensatory angular modifications at least in one of two tasks so that 30–100% of the trunk displacement was not compensated and thus influenced the hand position or trajectory. Results suggest that vestibular influences evoked by the head motion during trunk flexion play a major role in maintaining the consistency of arm motor actions in external space despite changes in the number of body segments involved. Our findings also suggest that despite long-term plasticity in the vestibular system and related neural structures, unilateral vestibular lesion may reduce the capacity of the nervous system to achieve motor equivalence.     

Postural adjustments for online corrections of arm movements in standing humans

The aim of this study was to investigate how humans correct ongoing arm movements while standing. Specifically, we sought to understand whether the postural adjustments in the legs required for online corrections of arm movements are predictive or rely on feedback from the moving limb. To answer this question we measured online corrections in arm and leg muscles during pointing movements while standing. Nine healthy right-handed subjects reached with their dominant arm to a visual target in front of them and aligned with their midline. In some trials, the position of the target would switch from the central target to one of the other targets located 15°, 30°, or 45° to the right of the central (midline) target. For each target correction, we measured the time at which arm kinematics, ground reaction forces, and arm and leg muscle electromyogram significantly changed in response to the target displacement. Results show that postural adjustments in the left leg preceded kinematic corrections in the limb. The corrective postural muscle activity in the left leg consistently preceded the corrective reaching muscle activity in the right arm. Our results demonstrate that corrections of arm movements in response to target displacement during stance are preceded by postural adjustments in the leg contralateral to the direction of target shift. Furthermore, postural adjustments preceded both the hand trajectory correction and the arm-muscle activity responsible for it, which suggests that the central nervous system does not depend on feedback from the moving arm to modify body posture during voluntary movement. Instead, postural adjustments lead the online correction in the arm the same way they lead the initiation of voluntary arm movements. This suggests that forward models for voluntary movements executed during stance incorporate commands for posture that are produced on the basis of the required task demands.     

Hemispheric specialization in the co-ordination of arm and trunk movements during pointing in patients with unilateral brain damage

During pointing movements involving trunk displacement, healthy subjects perform stereotypically, selecting a strategy in which the movement is initiated with either the hand or trunk, and where the trunk continues after the end of the hand movement. In a previous study, such temporal co-ordination was not found in patients with left-hemispheric brain lesions reaching with either their dominant paretic or with their non-dominant non-paretic arm. This co-ordination deficit may be associated in part with the presence of a lesion in the dominant left hemisphere. If so, then no deficit should be observed in patients with stroke-related damage in their non-dominant right hemisphere moving with their ipsilesional arm. To verify this, 21 right-hand dominant adults (7 who had had a stroke in the right hemisphere, 7 who had had a stroke in the left hemisphere and 7 healthy subjects) pointed to two targets located on a table in front of them in the ipsilateral and contralateral workspace. Pointing was done under three movement conditions: while not moving the trunk, while bending the trunk forward and while bending the trunk backwards. The experiment was repeated with the non-paretic arm of patients with stroke and for the right and left arms of healthy subjects. Kinematic data were recorded (Optotrak). Results showed that, compared to healthy subjects, arm-trunk timing was disrupted in patients with stroke for some conditions. As in patients with lesions in the dominant hemisphere, arm-trunk timing in those with lesions in the non-dominant hemisphere was equally more variable than movements in healthy subjects. However, patients with dominant hemisphere lesions used significantly less trunk displacement than those with non-dominant hemisphere lesions to accomplish the task. The deficit in trunk displacement was not due to problems of trunk control or sitting balance since, in control experiments, all subjects were able to move the trunk the required distance, with and without the added weight of the limb. Results support the hypothesis that the temporal co-ordination of trunk and arm recruitment during pointing movements is mediated bilaterally by each hemisphere. However, the difference in the range of trunk displacement between patients with left and right brain lesions suggests that the left (dominant) hemisphere plays a greater role than the right in the control of movements involving complex co-ordination between the arm and trunk. Electronic Publication     

Interaction of visual and proprioceptive feedback during adaptation of human reaching movements

People tend to make straight and smooth hand movements when reaching for an object. These trajectory features are resistant to perturbation, and both proprioceptive as well as visual feedback may guide the adaptive updating of motor commands enforcing this regularity. How is information from the two senses combined to generate a coherent internal representation of how the arm moves? Here we show that eliminating visual feedback of hand-path deviations from the straight-line reach (constraining visual feedback of motion within a virtual, "visual channel") prevents compensation of initial direction errors induced by perturbations. Because adaptive reduction in direction errors occurred with proprioception alone, proprioceptive and visual information are not combined in this reaching task using a fixed, linear weighting scheme as reported for static tasks not requiring arm motion. A computer model can explain these findings, assuming that proprioceptive estimates of initial limb posture are used to select motor commands for a desired reach and visual feedback of hand-path errors brings proprioceptive estimates into registration with a visuocentric representation of limb position relative to its target. Simulations demonstrate that initial configuration estimation errors lead to movement direction errors as observed experimentally. Registration improves movement accuracy when veridical visual feedback is provided but is not invoked when hand-path errors are eliminated. However, the visual channel did not exclude adjustment of terminal movement features maximizing hand-path smoothness. Thus visual and proprioceptive feedback may be combined in fundamentally different ways during trajectory control and final position regulation of reaching movements.     

Separate adaptive mechanisms for controlling trajectory and final position in reaching

We examined control of the hand's trajectory (direction and shape) and final equilibrium position in horizontal planar arm movements by quantifying transfer of learned visuomotor rotations between two tasks that required aiming the hand to the same spatial targets. In a trajectory-reversal task ("slicing"), the hand reversed direction within the target and returned to the origin. In a positioning task ("reaching"), subjects moved the hand to the target and held it there; cursor feedback was provided only after movement ended to isolate learning of final position from trajectory direction. We asked whether learning acquired in one task would transfer to the other. Transfer would suggest that the hand's entire trajectory, including its endpoint, was controlled using a common spatial plan. Instead we found minimal transfer, suggesting that the brain used different representations of target position to specify the hand's initial trajectory and its final stabilized position. We also observed asymmetrical practice effects on hand trajectory, including systematic curvature of reaches made after rotation training and hypermetria of untrained slice reversals after reach training. These are difficult to explain with a unified control model, but were replicated in computer simulations that specified the hand's initial trajectory and its final equilibrium position. Our results suggest that the brain uses different mechanisms to plan the hand's initial trajectory and final position in point-to-point movements, that it implements these control actions sequentially, and that trajectory planning does not account for specific impedance values to be implemented about the final stabilized posture.     

From head orientation to hand control: evidence of both neck and vestibular involvement in hand drawing

This research investigated the effect of head to trunk relation in a sensorimotor drawing task. In the first experiment, seated participants were asked to reproduce with eyes closed geometric shapes (square or diamond) with the tip of their right index finger in the frontoparallel plane. Their head was either aligned with the trunk or tilted 25° towards the left or right shoulder. Results showed that drawings were subjected to an overall rotation of a few degrees in the opposite direction to the tilt. In two subsequent experiments, the respective contribution of both otoliths and neck receptors to this head tilt effect was investigated. In Experiment 2, seated participants kept their head straight but were subjected to 2.5 mA vestibular galvanic stimulation (GVS). Results indicated that GVS induced a small but significant deviation of the drawings towards the anode. Finally, in Experiment 3, subjects performed the drawing task either seated upright (seated condition) or lying on their back (supine condition). Unlike in the seated condition, tilting the head towards the shoulders in a supine posture does not modulate afferents from the otolith stimulation and therefore mainly stimulates neck receptors. Head tilt induced rotations of hand-drawn reproductions in both seated and supine conditions, suggesting a significant contribution of neck afferents in the control of hand motion in space in the absence of vision. Overall the data provided evidence for a strong head-hand linkage during kinaesthetically guided drawing movements. Electronic Publication     

Dynamic visual–vestibular integration during goal directed human locomotion

Normal visual input plays a very dominant role during locomotion. Functionally, it can assist the central nervous system to overcome a destabilizing effect of abnormal or perturbed vestibular information. However, a recent study has shown a directional effect of transmastoidal galvanic vestibular stimulation (GVS) on gait trajectory when visual information is unreliable. The purpose of this study was to investigate how inputs from the visual and vestibular systems are weighted to optimize locomotor performance under impoverished visual conditions during goal directed locomotion. For unimodal stimulation, the visual input was manipulated using displacing prisms that caused 14 degrees horizontal displacement of perceived target location to the right or left. In addition, GVS (0.8 mAmp) was applied to manipulate vestibular system information during bimodal stimulation conditions. Two bimodal stimulation conditions were defined by the polarity of the galvanic current (anode on congruent and incongruent sides of prismatic deviation). The center of mass (CoM) displacement, head and trunk yaw angles and trunk roll angles were computed to analyze the global output as well as segmental coordination, as the participants walked towards the target. Although the performance was primarily guided by visual information, both congruent and incongruent GVS significantly altered CoM displacement. Similarly, the basic pattern of segmental responses during steering was maintained; however, the magnitude of the responses was altered. Spatio-temporal analysis demonstrated that during bimodal stimulation, the effect of GVS on global output tapered off as the participants approached the target. Results suggest a dynamic visual-vestibular interaction in which the gain of the vestibular input is initially upregulated in the presence of insufficient or impoverished visual information. However, there is a gradual habituation and the visual information, although insufficient, primarily dominates during goal directed locomotion. The experimental trajectories resembled mathematically simulated trajectories with a decaying GVS gain as opposed to a constant gain, further supporting the dynamic nature of sensory integration.     

The influence of motion signals in hand movements

It has been shown that motion after-effects (MAE) may affect the perceived position of moving objects and, more recently, that MAE signals can also affect pursuit eye movements: smooth pursuit eye movements are favoured by the illusory motion percept that is caused by motion adaptation. Here we investigated the relationship between MAE and arm movements. The objective of our research was: (1) to analyze possible effects of MAE when the arm tracks the changing position of a moving object, and (2) to investigate the influence of MAE on pointing movements to both static and moving targets. Our results show that the (unseen) hand position was trailing the target much less when target and MAE direction was the same. At the end of manual pursuit, subjects caught up with the moving target. However, when target direction was opposite the MAE, subjects’ hands moved more slowly, causing larger lags between the target and the hand position (Experiment 1). In Experiment 2, we found a similar effect of motion signals when subjects pointed to a moving target but found no effect of MAE when pointing to a static object (Experiment 3). We conclude that the effect of motion signals is only revealed when we need to update the changing position of a target.     

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.     

Visual–vestibular interaction during goal directed locomotion: effects of aging and blurring vision

Normal vision overrides perturbed vestibular information for the optimization of performance during goal directed locomotion, suggesting down-regulation of vestibular gain. However, it is not known if the responses to vestibular perturbation are accentuated when vision is impaired. Furthermore, both visual and vestibular systems deteriorate with age. It is not clear, however, how age-related decline in these sensory systems influences visual–vestibular interaction. Therefore, the dual purpose of the present study was to investigate the effects of aging and blurring vision, that simulated the consequences of cataracts, on visual–vestibular interaction. Young and healthy elderly walked to a target located straight ahead with either normal or blurring vision. On randomly selected trials vestibular system perturbation was achieved by applying transmastoidal galvanic vestibular stimulation (GVS). Two different galvanic stimulation intensities were used to provide insight into scaling effect of vestibular perturbation on locomotor performance and how age and vision influences this scaling effect. Maximum path deviation, frontal trunk tilt and postural coordination in the mediolateral direction were evaluated. The magnitude of the path deviation and the trunk tilt response were scaled to the magnitude of the vestibular perturbation in older adults independent of the visual condition. Older participants demonstrated increased coupling of the head and trunk segments irrespective of visual and vestibular perturbations. The results suggest that when visual information was available, the vestibular input reweighting was less effective in older individuals, as shown by the scaled responses to the GVS intensities and the inability to converge efficiently towards the target.     

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

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

Use of the trunk for reaching targets placed within and beyond the reach in adult hemiparesis

Multijoint movements such as reaching are impaired after brain lesions involving sensorimotor areas and pathways. However, the mechanisms by which such lesions affect motor control are not fully understood. Direct effects of the lesion may be partly compensated by both the system's redundancy and its plasticity. Indeed stroke patients with limited arm movement can reach objects placed within the reach of the arm by using a compensatory strategy involving trunk recruitment. A similar strategy is observed in healthy individuals reaching for objects placed beyond the reach of the arm. Determining the control mechanism(s) governing this compensatory strategy in stroke patients was the goal of this study. Kinematics of reaching movements in hemiparetic and healthy participants to targets placed within and beyond the length of the arm were analysed. Targets were placed sagittally in front of the midline of the body. Two targets (targets 1 and 2) were within reaching distance defined as the length of the stretched arm from axilla to wrist crease. Two others were beyond arm's reach so that one required a forward trunk inclination (target 3) and the other required body raising to a semi-standing position (target 4). Healthy participants used minimal trunk displacement for reaches to targets 1 and 2. For reaches to targets 3 and 4, trunk displacement increased with target distance. Whenever the trunk was involved, there was a stereotyped sequential recruitment of the arm and trunk in that the trunk began moving simultaneously with or before the hand and stopped moving after the end of hand movement. This suggested that the control system predicts that the trunk movement will be needed to extend the reach and includes the trunk, in an anticipatory way, into the reach. In contrast, most hemiparetic participants recruited their trunk for reaches to all four targets, even those placed close to the body. Similar to healthy individuals, the sequence of hand and trunk recruitment was stereotyped, suggesting that temporal planning aspects of the motor program underlying movement coordination were relatively unaffected. In contrast to healthy participants, the contribution of the trunk movement to the endpoint displacement was substantially higher in the hemiparetic group and occurred earlier in the reach. It is suggested that the target distance at which the trunk is integrated into the movement to extend the reach of the arm is attained around the limit of arm extension and that this limit is reduced in hemiparetic individuals.