Unraveling the interaction between pathological upper limb synergies and compensatory trunk movements during reach-to-grasp after stroke: a cross-sectional study

The aim of the present study was to identify how pathological limb synergies between shoulder and elbow movements interact with compensatory trunk movements during a functional movement with the paretic upper limb after stroke. 3D kinematic joint and trunk angles were measured during a reach-to-grasp movement in 46 patients with stroke and 12 healthy individuals. We used principal component analyses (PCA) to identify components representing linear relations between the degrees of freedom of the upper limb and trunk across patients with stroke and healthy participants. Using multivariate logistic regression analysis, we investigated whether component scores were related to the presence or absence of basic limb synergies as indicated by the arm section of the Fugl-Meyer motor assessment (FMA). Four and three principal components were extracted in patients with stroke and healthy individuals, respectively. Visual inspection revealed that the contribution of joint and trunk angles to each component differed substantially between groups. The presence of the flexion synergy (Shoulder Abduction and Elbow Flexion) was reflected by component 1, whereas the compensatory role of trunk movements for lack of shoulder and elbow movements was reflected by components 2 and 3 respectively. The presence or absence of basic limb synergies as determined by means of the FMA was significantly related to components 2 (p = 0.014) and 3 (p = 0.003) in patients with stroke. These significant relations indicate that PCA is a useful tool to identify clinically meaningful interactions between compensatory trunk movements and pathological synergies in the elbow and shoulder during reach-to-grasp after stroke.     

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.     

Workspace location influences joint coordination during reaching in post-stroke hemiparesis

The purpose of this study was to determine the influence of workspace location on joint coordination in persons with post-stroke hemiparesis when trunk motion was required to complete reaches beyond the arm’s functional reach length. Seven subjects with mild right hemiparesis following a stroke and seven age and gender matched control subjects participated. Joint motions and characteristics of hand and trunk movement were measured over multiple repetitions. The variance (across trials) of joint combinations was partitioned into two components at every point in the hand’s trajectory using the uncontrolled manifold approach; the first component is a measure of the extent to which equivalent joint combinations are used to control a given hand path, and reflects performance flexibility. The second component of joint variance reflects the use of non-equivalent joint combinations, which lead to hand path error. Compared to the control subjects, persons with hemiparesis demonstrated a significantly greater amount of non-equivalent joint variability related to control of the hand’s path and of the hand’s position relative to the trunk when reaching toward the hemiparetic side (ipsilaterally), but not when reaching to the less involved side. The relative timing of the hand and trunk was also altered when reaching ipsilaterally. The current findings support the idea that the previously proposed “arm compensatory synergy” may be deficient in subjects with hemiparesis. This deficiency may be due to one or a combination of factors: changes in central commands that are thought to set the gain of the arm compensatory synergy; a limited ability to combine shoulder abduction and elbow extension that limits the expression of an appropriately set arm compensatory synergy; or a reduction of the necessary degrees-of-freedom needed to adequately compensate for poor trunk control when reaching ipsilaterally.     

A kinematic comparison of single and multijoint pointing movements

Summary Rapid pointing movements (no accuracy or reaction time requirements) were performed under three conditions which limited motion to the shoulder, elbow or a combination of these two joints. Velocity profiles of the hand's trajectory differed during single and multijoint movements. For the same magnitude of displacement, the hand always had a higher peak velocity, shorter rise time (time to peak velocity) and shorter movement time during single joint movements. However, when the profiles were normalized with respect to amplitude and movement time, no significant differences were observed between these three movement conditions. The velocity profiles of the elbow and/or shoulder were also compared under single and multijoint movement conditions. Analysis of these profiles revealed that the relationships between peak velocity and displacement and between movement time and displacement remained the same at the shoulder joint during single and multijoint movements. In contrast, the elbow joint velocity profiles were significantly affected by movement conditions. These relationships (peak velocity/ displacement and movement time/displacement) changed during multijoint movements and became the same as those observed at the shoulder joint. The shape of the hand velocity profile and its invariance across movement conditions can best be explained by dynamic optimization theory and supports the notion that movement of the hand is of primary importance during rapid pointing. However, the consistency of the shoulder velocity profile and the highly significant relationships between the movement of the elbow and shoulder joints indicates that a subordinate joint planning strategy is also used. The purpose of this strategy is to functionally decrease the available degrees of freedom and to simplify coordination between the moving joints. Thus, the organization of arm movements is hierarchically structured with important, but different contributions being made on both the hand planning and joint planning levels.     

Vestibular contribution to combined arm and trunk motion

Recent studies have shown that the hand-pointing movements within arm's reach remain invariant whether the trunk is recruited or not or its motion is unexpectedly prevented. This suggests the presence of compensatory arm-trunk coordination minimizing the deflections of the hand from the intended trajectory. It has been postulated that vestibular signals elicited by the trunk motion and transmitted to the arm motor system play a major role in the compensation. One prediction of this hypothesis is that vestibular stimulation should influence arm posture and movement during reaching. It has been demonstrated that galvanic vestibular stimulation (GVS) can influence the direction of pointing movements when body motion is restrained. In the present study, we analyzed the effects of GVS on trunk-assisted pointing movements. Subjects either moved the hand to a target or maintained a steady-state posture near the target, while moving the trunk forward with the eyes closed. When GVS was applied, the final position of the hand was deviated in the lateral and sagittal direction in both tasks. This was the result of two independent effects: a deviation of the trunk trajectory and a modification of the arm position relative to the trunk. Thus, the vestibular system might be directly involved not only in the control of trunk motion but also in the arm-trunk coordination during trunk-assisted reaching movements. Electronic Publication     

How to extend the elbow with a weak or paralyzed triceps: control of arm kinematics for aiming in C6-C7 quadriplegic patients

This study aims to investigate how quadriplegic patients with a C6-C7 spinal lesion coordinate their upper limb to extend the elbow despite the paralysis or weakness of the triceps brachii, and what is the effect of a surgical musculotendinous transfer. For this purpose, aiming movements in a wide workspace were recorded in seven healthy subjects and in patients with incomplete (five cases) or complete (eight cases) triceps paralysis and after musculotendinous transfer (eight cases). We used four electromagnetic field sensors to quantify hand trajectory and to compute the angles describing the rotations at the scapula, glenohumeral joint, elbow and wrist (10 degrees of freedom). Extent and smoothness of the hand trajectories and hand velocity profiles were surprisingly similar between healthy subjects and quadriplegic patients. The reduction of elbow extension observed in patients was compensated by rotations distributed across several degrees of freedom including the scapula. Principal components analysis showed that the joint rotations could be summarized by an additive combination of two synergies, respectively orientating and stretching out the limb, which explained similar amounts of variance in healthy subjects and in patients. The participations of degrees of freedom in the synergies were roughly similar in the different groups of subjects, the main difference concerning scapular medial-lateral rotation, which seems to be critical in patients with a complete triceps paralysis. This demonstrates that elbow extension in quadriplegic patients is due to anticipated mechanical interaction coupling between upper limb segments. We propose that the persisting (incomplete paralysis) or restored (musculotendinous transfer) elbow extensor strength may act by stabilizing the elbow. This counterintuitive preservation of limb kinematics for horizontal aiming movements in quadriplegic patients despite the drastic changes in muscle action provoked by paralysis and/or by surgery strongly suggests that the motor system does not primarily control forces but the morphological aspects of movement, via joint rotation synergies.     

Motor synergies during manual tracking differ between familiar and unfamiliar trajectories

Synergistic control of the effector space allows high precision in task-relevant degrees of freedom, while noise is limited to task-irrelevant degrees of freedom. The present study investigates whether this typical structure of the variance–covariance matrix of the joint angles during manual tracking differs between familiar and unfamiliar trajectories. Subjects tracked a target moving in 2D on a graphics tablet with a hand-held pen, while their arm movements were not restricted. Subjects familiarized themselves with one target trajectory during an initial training block with 40 periodic trials. In the following test block, this familiar trajectory and several unfamiliar trajectories were presented in a mixed-block design to study prediction effects at the level of endpoint and joint trajectories. The differences in the synergistic control of arm movements were analyzed using the “uncontrolled manifold method.” The results showed smaller variances and weaker motor synergies during tracking of familiar trajectories than during tracking of unfamiliar trajectories. The decrease in the synergy index was due to a stronger decrease in the variance irrelevant than of the variance relevant for pen position. In the context of motor control theory, these results suggest that tracking movements on familiar and unfamiliar target trajectories do not only differ in the available knowledge about target location but also apply different strategies to control the effector space.     

Limb position drift: implications for control of posture and movement

In the absence of visual feedback, subject reports of hand location tend to drift over time. Such drift has been attributed to a gradual reduction in the usefulness of proprioception to signal limb position. If this account is correct, drift should degrade the accuracy of movement distance and direction over a series of movements made without visual feedback. To test this hypothesis, we asked participants to perform six series of 75 repetitive movements from a visible start location to a visible target, in time with a regular, audible tone. Fingertip position feedback was given by a cursor during the first five trials in the series. Feedback was then removed, and participants were to continue on pace for the next 70 trials. Movements were made in two directions (30 degrees and 120 degrees ) from each of three start locations (initial shoulder angles of 30 degrees, 40 degrees, 50 degrees, and initial elbow angles of 90 degrees ). Over the 70 trials, the start location of each movement drifted, on average, 8 cm away from the initial start location. This drift varied systematically with movement direction, indicating that drift is related to movement production. However, despite these dramatic changes in hand position and joint configuration, movement distance and direction remained relatively constant. Inverse dynamics analysis revealed that movement preservation was accompanied by substantial modification of joint muscle torque. These results suggest that proprioception continues to be a reliable source of limb position information after prolonged time without vision, but that this information is used differently for maintaining limb position and for specifying movement trajectory.     

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.     

Differences in control of limb dynamics during dominant and nondominant arm reaching

This study compares the coordination patterns employed for the left and right arms during rapid targeted reaching movements. Six right-handed subjects reached to each of three targets, designed to elicit progressively greater amplitude interaction torques at the elbow joint. All targets required the same elbow excursion (20 degrees ), but different shoulder excursions (5, 10, and 15 degrees, respectively). Movements were restricted to the shoulder and elbow and supported on a horizontal plane by a frictionless air-jet system. Subjects received visual feedback only of the final hand position with respect to the start and target locations. For motivation, points were awarded based on final position accuracy for movements completed within an interval of 400-600 ms. For all subjects, the right and left hands showed a similar time course of improvement in final position accuracy over repeated trials. After task adaptation, final position accuracy was similar for both hands; however, the hand trajectories and joint coordination patterns during the movements were systematically different. Right hand paths showed medial to lateral curvatures that were consistent in magnitude for all target directions, whereas the left hand paths had lateral to medial curvatures that increased in magnitude across the three target directions. Inverse dynamic analysis revealed substantial differences in the coordination of muscle and intersegmental torques for the left and right arms. Although left elbow muscle torque contributed largely to elbow acceleration, right arm coordination was characterized by a proximal control strategy, in which movement of both joints was primarily driven by the effects of shoulder muscles. In addition, right hand path direction changes were independent of elbow interaction torque impulse, indicating skillful coordination of muscle actions with intersegmental dynamics. In contrast, left hand path direction changes varied directly with elbow interaction torque impulse. These findings strongly suggest that distinct neural control mechanisms are employed for dominant and non dominant arm movements. However, whether interlimb differences in neural strategies are a consequence of asymmetric use of the two arms, or vice versa, is not yet understood. The implications for neural organization of voluntary movement control are discussed.     

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.     

Arm reaching improvements with short-term practice depend on the severity of the motor deficit in stroke

The effects of short-term, constant practice on the kinematics of a multi-joint pointing movement were studied in the hemiparetic arm of 20 chronic patients with unilateral left cerebro-vascular accident (CVA) and in 10 age- and sex-matched healthy individuals. Practice consisted of a single session of 70 pointing movements made with the right arm. Movements were made from a target located beside the body to one in the contralateral workspace, in front of the body. Vision of the final hand position was allowed after every 5^th trial. At the beginning of practice, stroke patients made slower, less precise and more segmented movements, characterised by smaller active ranges of elbow and shoulder motion, disrupted elbow–shoulder coordination, as well as greater trunk movement compared with healthy subjects. With practice, healthy subjects and some patients made faster and more precise movements. These tendencies were revealed only after many repetitions (up to 55 for those with severe hemiparesis), whereas changes in healthy individuals occurred after fewer trials (approximately 20). In addition, the patients decreased movement segmentation with practice. In healthy subjects, faster movement times may be attributed to better shoulder/elbow movement timing in the first half of the reach, whereas improvement of precision was not correlated with any changes in the movement variables. In patients, improvements were accomplished differently depending on arm motor severity. For some patients with mild-to-moderate clinical symptoms, practice resulted in better timing of shoulder/elbow movements with less trunk rotation in middle to late reach. Patients with more severe impairment also improved shoulder/elbow movement timing in mid-reach but used more compensatory trunk rotation. The results suggest that even one session of repetitive practice of a multi-joint pointing task leads to improvements in movement performance-based outcome measures, but the mechanisms of improvement may vary with the individual's level of motor impairment.     

Recruitment and sequencing of different degrees of freedom during pointing movements involving the trunk in healthy and hemiparetic subjects

 Previous studies have shown that in neurologically normal subjects the addition of trunk motion during a reaching task does not affect the trajectory of the arm endpoint. Typically, the trunk begins to move before the onset and continues to move after the offset of the arm endpoint displacement. This observation shows that the potential contribution of the trunk to the motion of the arm endpoint toward a target is neutralized by appropriate compensatory movements of the shoulder and elbow. We tested the hypothesis that cortical and subcortical brain lesions may disrupt the timing of trunk and arm endpoint motion in hemiparetic subjects. Eight hemiparetic and six age-matched healthy subjects were seated on a stool with the right (dominant) arm in front of them on a table. The tip of the index finger (the arm endpoint) was initially at a distance of 20 cm from the midline of the chest. Wrist, elbow, and upper body positions as well as the coordinates of the arm endpoint were recorded with a three-dimensional motion analysis system (Optotrak) by infrared light-emitting diodes placed on the tip of the finger, the styloid process of the ulna, the lateral epicondyle of the humerus, the acromion processes bilaterally, and the sternal notch. In response to a preparatory signal, subjects lifted their arm 1–2 cm above the table and in response to a ”go” signal moved their endpoint as fast as possible from a near to a far target located at a distance of 35 cm and at a 45° angle to the right or left of the sagittal midline of the trunk. After a pause (200– 500 ms) they moved the endpoint back to the near target. Pointing movements were made without trunk motion (control trials) or with a sagittal motion of the trunk produced by means of a hip flexion or extension (test trials). In one set of test trials, subjects were required to move the trunk forward while moving the arm to the target (”in-phase movements”). In the other set, subjects were required to move the trunk backward when the arm moved to the far target (”out-of-phase movements”). Compared with healthy subjects, movements in hemiparetic subjects were segmented, slower, and characterized by a greater variability and by deflection of the trajectory from a straight line. In addition, there was a moderate increase in the errors in movement direction and extent. These deficits were similar in magnitude whether or not the trunk was involved. Although hemiparetic subjects were able to compensate the influence of the trunk motion on the movement of the arm endpoint, they accomplished this by making more segmented movements than healthy subjects. In addition, they were unable to stabilize the sequence of trunk and arm endpoint movements in a set of trials. It is concluded that recruitment and sequencing of different degrees of freedom may be impaired in this population of patients. This inability may partly be responsible for other deficits observed in hemiparetic subjects, including an increase in movement segmentation and duration. The lack of stereotypic movement sequencing may imply that these subjects had deficits in learning associated with short-term memory. Received: 9 September 1997 / Accepted: 9 October 1998     

Path constraints on point-to-point arm movements in three-dimensional space

In this paper data are presented concerning the kinematic and dynamic characteristics of point-to-point arm movements which are inwardly or outwardly directed in three-dimensional space. Elbow and wrist position as well as elbow angle of extension were measured. From these data, other angles were computed trigonometrically and elbow and shoulder torques were calculated. Some of the angles describing arm and forearm motion were found to be linearly related for any given movement. Changes in shoulder and elbow torque were found to be similar to those described for movements restricted to one degree of freedom. Shoulder and elbow motions were not affected when it was required that the orientation of the hand in space remain constant. These observations were taken to indicate that shoulder and elbow motions are tightly coupled for movements in three-dimensional space and that wrist motion has no influence on this coupling. Linear relations between angles express such coupling. They are taken to result from functional constraints and may facilitate the mapping between extrinsic and intrinsic coordinate systems. Some of the observations pertaining to the torque lead to the hypothesis of a further constraint limiting the number of possible trajectories in a point-to-point movement.     

Coordination of multi-joint arm movements in cerebellar ataxia: Analysis of hand and angular kinematics

Kinematic abnormalities of fast multijoint movements in cerebellar ataxia include abnormally increased curvature of hand trajectories and an increased hand path and are thought to originate from an impairment in generating appropriate levels of muscle torques to support normal coordination between shoulder and elbow joints. Such a mechanism predicts that kinematic abnormalities are pronounced when fast movements are performed and large muscular torques are required. Experimental evidence that systematically explores the effects of increasing movement velocities on movement kinematics in cerebellar multijoint movements is limited and to some extent contradictory. We, therefore, investigated angular and hand kinematics of natural multijoint pointing movements in patients with cerebellar degenerative disorders and healthy controls. Subjects performed self-paced vertical pointing movements with their right arms at three different target velocities. Limb movements were recorded in three-dimensional space using a two-camera infrared tracking system. Differences between patients and healthy subjects were most prominent when the subjects performed fast movements. Peak hand acceleration and deceleration were similar to normals during slow and moderate velocity movements but were smaller for fast movements. While altering movement velocities had little or no effect on the length of the hand path and angular motion of elbow and shoulder joints in normal subjects, the patients exhibited overshooting motions (hypermetria) of the hand and at both joints as movement velocity increased. Hypermetria at one joint always accompanied hypermetria at the neighboring joint. Peak elbow angular deceleration was markedly delayed in patients compared with normals. Other temporal movement variables such as the relative timing of shoulder and elbow joint motion onsets were normal in patients. Kinematic abnormalities of multijoint arm movements in cerebellar ataxia include hypermetria at both the elbow and the shoulder joint and, as a consequence, irregular and enlarged paths of the hand, and they are marked with fast but not with slow movements. Our findings suggest that kinematic movement abnormalities that characterize cerebellar limb ataxia are related to an impairment in scaling movement variables such as joint acceleration and deceleration normally with movement speed. Most likely, increased hand paths and decomposition of movement during slow movements, as described earlier, result from compensatory mechanisms the patients may employ if maximum movement accuracy is required.     

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.     

Control of human arm movements in two dimensions: paths and joint control in avoiding simple linear obstacles

In order to examine path planning and the control of redundant degrees of freedom in the human arm, the movements of the shoulder, elbow and wrist were recorded as subjects moved a pointer to a target and avoided a simple obstacle. With respect to joint control, the results show that the extra degree of freedom provided by the wrist is incorporated into target movements in a systematic manner for both large and small obstacles; it is not used only when there is no geometrical alternative. For the wrist, two strategies are apparent, depending upon the length of the obstacle. Wrist extension predominates for shorter obstacles, while flexion or extension and flexion predominate for longer obstacles. These wrist movements shorten the effective length of the distal segments (lower arm plus hand and pointer) and thus reduce the excursion required at the proximal joints. In part, they correspond to assuming the most comfortable arm configuration at each point in the new path necessitated by the obstacle and can be described by static cost functions. However, wrist extension is also used to move the hand and pointer away from the obstacle as shoulder and elbow movements carry the wrist itself towards the obstacle. Wrist flexion is also used to move the pointer tip rapidly past the obstacle. These components, which cannot be explained by static cost functions alone, confirm for the human arm the hypothesized use of redundant degrees of freedom in obstacle avoidance. With respect to path planning, the results show that the minimum distance between pointer and obstacle remains fairly constant over a large range of obstacle lengths; this relative invariance is interpreted to support the hypothesis that workspace coordinates are important for movement planning. However, minimum distance and several other path parameters do depend significantly on the orientation and location of the movement in the workspace. This inhomogeneity implies that movement planning does not occur exclusively in workspace coordinates; it suggests an influence of joint space criteria. In frontal movements, for example, the systematic decline in the minimum distance with increasing obstacle length is interpreted as a compromise reducing the amount of extra joint movement and the discomfort of arm configurations.     

Spatial control of arm movements

Summary Human subjects were instructed to point one hand to different visual targets which were randomly sequenced, using a paradigm which allowed two degrees of freedom (shoulder, elbow). The time course of the hand trajectory and the joint angular curves were observed. The latter exhibited patterns which change markedly for different movements, whereas the former preserve similar characteristics (in particular, a single peaked tangential velocity curve). The hypothesis is then formulated that the central command for these movements is formulated in terms of trajectories of the hand in space.Research supported by the Italian Research Council (Bilateral Research Contract) and by N.I.H. Grants NS 09343 and AM 26710     

Modulation of phasic and tonic muscle synergies with reaching direction and speed

How the CNS masters the many degrees of freedom of the musculoskeletal system to control goal-directed movements is a long-standing question. We have recently provided support to the hypothesis that the CNS relies on a modular control architecture by showing that the phasic muscle patterns for fast reaching movements in different directions are generated by combinations of a few time-varying muscle synergies: coordinated recruitment of groups of muscles with specific activation profiles. However, natural reaching movements occur at different speeds and require the control of both movement and posture. Thus we have investigated whether muscle synergies also underlie reaching at different speeds as well as the maintenance of stable arm postures. Hand kinematics and shoulder and elbow muscle surface EMGs were recorded in five subjects during reaches to eight targets in the frontal plane at different speeds. We found that the amplitude modulation of three time-invariant synergies captured the variations in the postural muscle patterns at the end of the movement. During movement, three phasic and three tonic time-varying synergies could reconstruct the time-normalized muscle pattern in all conditions. Phasic synergies were modulated in both amplitude and timing by direction and speed. Tonic synergies were modulated only in amplitude by direction. The directional tuning of both types of synergies was well described by a single or a double cosine function. These results suggest that muscle synergies are basic control modules that allow generating the appropriate muscle patterns through simple modulation and combination rules.     

Torque response to external perturbation during unconstrained goal-directed arm movements

It is unclear to what extent control strategies of 2D reaching movements of the upper limbs also apply to movements with the full seven degrees of freedom (DoFs) including rotation of the forearm. An increase in DoFs may result in increased movement complexity and instability. This study investigates the trajectories of unconstrained reaching movements and their stability against perturbations of the upper arm. Reaching movements were measured using an ultrasound marker system, and the method of inverse dynamics was applied to compute the time courses of joint torques. In full DoF reaching movements, the velocity of some joint angles showed multiple peaks, while the bell-shaped profile of the tangential hand velocity was preserved. This result supports previous evidence that tangential hand velocity is an essential part of the movement plan. Further, torque responses elicited by external perturbation started shortly after perturbation, almost simultaneously with the perturbation-induced displacement of the arm, and were mainly observed in the same joint angles as the perturbation torques, with similar shapes but opposite signs. These results indicate that these torque responses were compensatory and contributed to system stabilization.