Reprogramming of muscle activation patterns at the wrist in compensation for elbow reaction torques during planar two-joint arm movements

The relationship between wrist kinematics, dynamics and the pattern of muscle activation were examined during a two-joint planar movement in which the two joints moved in opposite directions, i.e. elbow flexion/wrist extension and elbow extension/wrist flexion. Elbow movements (ranging from 10 to 70 deg) and wrist movements (ranging from 10 to 50 deg) were performed during a visual, step-tracking task in which subjects were required to attend to the initial and final angles at each joint. As the elbow amplitude increased, wrist movement duration increased and the wrist movement trajectories became quite variable. Analysis of the torques acting at the wrist joint showed that elbow movements produced reaction torques acting in the same direction as the intended wrist movement. Distinct patterns of muscle activation were observed at the wrist joint that were dependent on the relative magnitude of the elbow reaction torque in relation to the net wrist torque. When the magnitude of the elbow reaction torque was quite small, the wrist agonist was activated first. As the magnitude of the elbow reaction torque increased, activity in the wrist agonist decreased significantly. In conditions where the elbow reaction torque was much larger than the net wrist torque, the wrist muscle torque reversed direction to oppose the intended movement. This reversal of wrist muscle torque was directly associated with a change in the pattern of muscle activation where the wrist antagonist was activated prior to the wrist agonist. Our findings indicate that motion of the elbow joint is an important consideration in planning wrist movement. Specifically, the selection of muscle activation patterns at the wrist is dependent on the relative magnitude and direction of the elbow reaction torque in relation to the direction of wrist motion.     

General coordination of shoulder,elbow and wrist dynamics during multijoint arm movements

Studies of multijoint arm movements have demonstrated that the nervous system anticipates and plans for the mechanical effects that arise from motion of the linked limb segments. The general rules by which the nervous system selects appropriate muscle activities and torques to best deal with these intersegmental effects are largely unknown. In order to reveal possible rules, this study examined the relationship of muscle and interaction torques to joint acceleration at the shoulder, elbow and wrist during point-to-point arm movements to a range of targets in the horizontal plane. Results showed that, in general, dynamics differed between the joints. For most movements, shoulder muscle torque primarily determined net torque and joint acceleration, while interaction torque was minimal. In contrast, elbow and wrist net torque were determined by a combination of muscle and interaction torque that varied systematically with target direction and joint excursion. This "shoulder-centered pattern" occurred whether subjects reached targets using straight or curved finger paths. The prevalence of a shoulder-centered pattern extends findings from a range of arm movement studies including movement of healthy adults, neurological patients, and simulations with altered interaction effects. The shoulder-centered pattern occurred for most but not all movements. The majority of the remaining movements displayed an "elbow-centered pattern," in which muscle torque determined initial acceleration at the elbow and not at the shoulder. This occurred for movements when shoulder excursion was <50% of elbow excursion. Thus, both shoulder- and elbow-centered movements displayed a difference between joints but with reversed dynamics. Overall, these findings suggest that a difference in dynamics between joints is a general feature of horizontal plane arm movements, and this difference is most commonly reflected in a shoulder-centered pattern. This feature fits well with other general shoulder-elbow differences suggested in the literature on arm movements, namely that: (a) agonist muscle activity appears more closely related to certain joint kinematics at the shoulder than at the elbow, (b) adults with neurological damage display less disruption of shoulder motion than elbow motion, and (c) infants display adult-like motion first in the shoulder and last at the wrist.     

Control of the wrist in three-joint arm movements to multiple directions in the horizontal plane

In a reaching movement, the wrist joint is subject to inertial effects from proximal joint motion. However, precise control of the wrist is important for reaching accuracy. Studies of three-joint arm movements report that the wrist joint moves little during point-to-point reaches, but muscle activities and kinetics have not yet been described across a range of movement directions. We hypothesized that to minimize wrist motion, muscle torques at the wrist must perfectly counteract inertial effects arising from proximal joint motion. Subjects were given no instructions regarding joint movement and were observed to keep the wrist nearly motionless during center-out reaches to directions throughout the horizontal plane. Consistent with this, wrist muscle torques exactly mirrored interaction torques, in contrast to muscle torques at proximal joints. These findings suggest that in this reaching task the nervous system chooses to minimize wrist motion by anticipating dynamic inertial effects. The wrist muscle torques were associated with a direction-dependent choice of muscles, also characterized by initial reciprocal activation rather than initial coactivation to stiffen the wrist joint. In a second experiment, the same pattern of muscle activities persisted even after many trials reaching with the wrist joint immobilized. These results, combined with similar features at the three joints, such as cosine-like tuning of muscle torques and of muscle onsets across direction, suggest that the nervous system uses similar rules for muscles at each joint, as part of one plan for the arm during a point-to-point reach.     

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.     

Effects of inactivation of the anterior interpositus nucleus on the kinematic and dynamic control of multijoint movement

We previously showed that inactivating the anterior interpositus nucleus in cats disrupts prehension; paw paths, normally straight and accurate, become curved, hypometric, and more variable. In the present study, we determined the joint kinematic and dynamic origins of this impairment. Animals were restrained in a hammock and trained to reach and grasp a cube of meat from a narrow food well at varied heights; movements were monitored using the MacReflex analysis system. The anterior interpositus nucleus was inactivated by microinjection of the GABA agonist muscimol (0.25-0.5 microgram in 0.5 microliter saline). For each joint, we computed the torque due to gravity, inertial resistance (termed self torque), interjoint interactions (termed interaction torque), and the combined effects of active muscle contraction and passive soft tissue stretch (termed generalized muscle torque). Inactivation produced significant reductions in the amplitude, velocity, and acceleration of elbow flexion. However, these movements continued to scale normally with target height. Shoulder extension was reduced by inactivation but wrist angular displacement and velocity were not. Inactivation also produced changes in the temporal coordination between elbow, shoulder, and wrist kinematics. Dynamic analysis showed that elbow flexion both before and during inactivation was produced by the combined action of muscle and interaction torque, but that the timing depended on muscle torque. Elbow interaction and muscle torques were scaled to target height both before and during inactivation. Inactivation produced significant reductions in elbow flexor interaction and muscle torques. The duration of elbow flexor muscle torque was prolonged to compensate for the reduction in flexor interaction torque. Shoulder extension was produced by extensor interaction and muscle torques both before and during inactivation. Inactivation produced a reduction in shoulder extension, primarily by reduced interaction torque, but without compensation. Wrist plantarflexion, which occurred during elbow flexion, was driven by plantarflexor interaction and gravitational torques both before and during inactivation. Muscle torque acted in the opposite direction with a phase lead to restrain the plantarflexor interaction torque. During inactivation, there was a reduction in plantarflexor interaction torque and a loss of the phase lead of the muscle torque. Our findings implicate the C1/C3 anterior interpositus zone of the cerebellum in the anticipatory control of intersegmental dynamics during reaching, which zone is required for coordinating the motions of the shoulder and wrist with those of the elbow. In contrast, this cerebellar zone does not play a role in scaling the movement to match a target.     

Trained slow tracking. I. Muscular production of wrist movement

Electromyographic (EMG) activity was recorded from those forearm muscles that act across the wrist as highly trained monkeys tracked slow hold-ramp-hold target trajectories with angular wrist position. During performance of this task, the forearm flexors and extensors had a common "basic pattern" of EMG activity. Flexor digitorum sublimis (FDS) and extensor digitorum communis (EDC), though commonly classified as prime movers of the fingers, were the most active flexor and extensor muscles during these movements at the wrist. The basic pattern of EMG activity was analyzed by varying independently 1) the movement direction, 2) the initial and final held wrist positions, 3) the ramp-movement velocity, and 4) the direction and magnitude of maintained external torque load. Most of the modulation of the basic pattern was related to wrist position: EMG amplitude was greatest at the extreme of muscle shortening. There was a slight difference in EMG activity between flexion and extension ramps that was related purely to the direction of movements, independent of wrist position, velocity, and external load; EMG amplitude was greater when a muscle was shortening and less when it was lengthening. During ramp movement, there was little or no observed EMG activity related to velocity (8-28 degrees/s). The magnitude of EMG activity varied in proportion to the external torque load, but this load-related component was additive, and the basic pattern of activity (related to direction and position) did not change with load. From these results we infer that a muscle's EMG activity was determined by 1) passive elastic properties of the wrist and the active length-tension characteristics of the muscle itself (position), 2) asymmetries in the muscle's contractile force depending on whether it was lengthening or shortening (direction), and 3) magnitude of the external torque load (force). By contrast, since no EMG activity was related to velocity in these slow movements, passive viscous properties and velocity-related cross-bridge kinetics were apparently so slight as to make undetectable the small additional EMG activity and contractile force presumably required to overcome them. A model of the muscle forces acting at the wrist incorporates these experimental observations.     

Cerebellar ataxia: torque deficiency or torque mismatch between joints?

Prior work has shown that cerebellar subjects have difficulty adjusting for interaction torques that occur during multi-jointed movements. The purpose of this study was to determine whether this deficit is due to a general inability to generate sufficient levels of phasic torque inability or due to an inability to generate muscle torques that predict and compensate for interaction torques. A second purpose was to determine whether reducing the number of moving joints by external mechanical fixation could improve cerebellar subjects' targeted limb movements. We studied control and cerebellar subjects making elbow flexion movements to touch a target under two conditions: 1) a shoulder free condition, which required only elbow flexion, although the shoulder joint was unconstrained and 2) a shoulder fixed condition, where the shoulder joint was mechanically stabilized so it could not move. We measured joint positions of the arm in the sagittal plane and electromyograms (EMGs) of shoulder and elbow muscles. Elbow and shoulder torques were estimated using inverse dynamics equations. In the shoulder free condition, cerebellar subjects made greater endpoint errors (primarily overshoots) than did controls. Cerebellar subjects' overshoot errors were largely due to unwanted flexion at the shoulder. The excessive shoulder flexion resulted from a torque mismatch, where larger shoulder muscle torques were produced at higher rates than would be appropriate for a given elbow movement. In the shoulder fixed condition, endpoint errors of cerebellar subjects and controls were comparable. The improved accuracy of cerebellar subjects was accompanied by reduced shoulder flexor muscle activity. Most of the correct cerebellar trials in the shoulder fixed condition were movements made using only muscles that flex the elbow. Our findings suggest that cerebellar subjects' poor shoulder control is due to an inability to generate muscle torques that predict and compensate for interaction torques, and not due to a general inability to generate sufficient levels of phasic torque. In addition, reducing the number of muscles to be controlled improved cerebellar ataxia.     

Counteractive relationship between the interaction torque and muscle torque at the wrist is predestined in ball-throwing

Many investigators have demonstrated that in swing motions such as ball-throwing, the motion of the proximal joint (shoulder) produced assistive interaction torque for the distal joint (elbow). In line with these studies, the shoulder and elbow motions would be expected to produce the assistive interaction torque for the wrist joint as well. However, we recently showed that the interaction torque at the wrist was always counteractive to the wrist muscle torque during ball-throwing. The purpose of this study is to clarify, by means of computer simulation, whether the counteractive relationship at the wrist during ball-throwing is caused by the neural contribution or the musculoskeletal mechanical properties of the human arm. First, we simulated the throwing motions of the normal forearm-hand model by systematically changing the proximal-to-distal delay of muscle activities and could line up two candidates for the determinant of the counteractive relationship: the rest angle (neutral angle) of the wrist and the length and mass of the hand. Second, we simulated the throwing motions of the virtual forearm-hand models, showing that only nonrealistic elongation of these two parameters produced the assistive relationship between the interaction torque and muscle torque. These results suggested that the mechanical properties of the human wrist are the main determinant of the counteractive relationship, which is advantageous for keeping the state of the wrist joint stable in multi-joint upper-limb movements and would lead to avoidance of excessive wrist extension or flexion and simplification of extrinsic finger control.     

Compensation for mechanically unstable loading in voluntary wrist movement

In order to study the roles of muscle mechanics and reflex feedback in stabilizing movement, experiments were conducted in which healthy human subjects performed targeted wrist movements under conditions where the damping of the wrist was reduced with a load having the property of negative viscosity. If the movement speed and negative viscosity were sufficiently high, the wrist oscillated for several hundred milliseconds about the final target position. Subjects increased the activation of both wrist flexor and extensor muscles to increase joint stiffness to damp the oscillations. With practice, both the tendency to oscillate and the level of muscle activation were reduced. A small bias torque in either direction, added to the negative viscosity, enhanced the oscillations as well as the amount of flexor and extensor muscle activation during the stabilization phase of fast movements. The tendency for the wrist to oscillate was also seen during slow movements where the oscillations were superimposed upon the voluntary movement. We suggest that this reduction in mechanical stability is primarily of reflex origin. As wrist stiffness increases, the natural frequency of the wrist also increases, which in turn produces an increase in the phase lag of the torque generated by the myotatic reflex with respect to wrist angular velocity, effectively reducing damping. The oscillation frequency was often close to a critical frequency for stability at which torque, due to the myotatic reflex, lagged angular velocity by 180° (6–7.5 Hz). Nevertheless, subjects were able to damp these oscillations, probably because the torque due to intrinsic muscle stiffness (in phase with position and hence lagging velocity by only 90°) dominated the torque contribution of the myotatic reflex. Increasing stiffness with declining oscillation amplitude may also have contributed significantly to damping.     

Activity patterns of upper arm muscles in relation to direction of rapid wrist movement in man

Summary Adjustment of arm posture associated with rapid wrist movements was studied by EMG analysis. Seven healthy adults, seated and holding their right arm with the shoulder in a neutral position with the elbow in 90° flexion and the wrist position neutral, were instructted to flex or extend the wrist as fast as possible. To examine whether the activity patterns of the upper arm muscles were related to the prime mover or the direction of the movement in space, the forearm was in two postures, supinate and pronate. The surface EMGs of biceps brachii, brachialis, triceps brachii and the prime movers were recorded along with the angular displacement of the wrist. The sequences of the upper arm muscle activities changed in relation to the direction of the movement. The earliest activities of the upper arm muscles were considered to counteract the dynamic perturbation induced by the rapid wrist movement. The onsets of the earliest activity of the upper arm muscles preceded the movement onset by 50–60 ms. These results revealed that the activity patterns of the arm muscles associated with the rapid wrist movements were functionally compatible with the anticipatory postural adjustment and were controlled according to the direction of the movement in space.     

Disruptions in joint control during drawing arm movements in Parkinson’s disease

Impairments in control of multi-joint arm movements in Parkinsons Disease (PD) were investigated. The PD patients and age-matched elderly participants performed cyclical arm movements, tracking templates of a large circle and four differentially oriented ovals on a horizontal table. The wrist was immobilized and the movements were performed with shoulder and elbow rotations. The task was performed with and without vision at a cycling frequency of 1.5 Hz. Traces of the arm endpoint, joint-motion parameters represented by range of motion and relative phase, and joint-control characteristics represented by amplitude and timing of muscle torque were analyzed. The PD patients provided deformations of the template shapes that were not observed in movements of elderly controls. The deformations were consistent for each shape but differed across the shapes, making quantification of impairments in the endpoint movement difficult. In contrast, the characteristics of joint control and motion demonstrated systematic changes across all shapes in movements of PD patients, although some of these changes were observed only without vision. A specification of the PD influence was observed at the level of joint control and it was not distinguishable in joint and endpoint motion, because of the property of multi-joint movements during which control at each joint influences motion at the other joints. The results suggest that inability of PD patients to provide fine muscle torque regulation coordinated across the joints contributes to the altered endpoint trajectories during multi-joint movements. The study emphasizes the importance of the torque analysis when deficits in multi-joint movements are investigated, because specific impairments that can be detected in joint-control characteristics are difficult to trace in characteristics of joint and endpoint kinematics, because of interactions between joint motions.     

Joint-specific disruption of control during arm movements in Parkinson’s disease

The leading joint hypothesis (LJH) suggests distinct types of control (leading and subordinate) at different joints during multi-joint movements. Taking into account specific features of movements in Parkinson’s disease (PD), the LJH predicts distinct effect of PD on control of leading and subordinate joints: impaired interaction torque (INT) regulation should be emphasized at the subordinate joints, and impaired generation of muscle torque (MUS) magnitude should be more pronounced at the leading joint. This prediction was tested by studying three tasks of horizontal shoulder-elbow movements in PD patients and age-matched controls: cyclic line drawing, cyclic point-to-point, and discrete pointing movements. Each task included movements in different directions, providing both shoulder-lead and elbow-lead control patterns. Torque analysis supported the prediction, specifically for Tasks 2 and 3 in which movement targets were chosen to emphasize the shoulder- and elbow-lead control patterns. Patients did not exploit INT for motion generation as successfully as controls did, but only at the subordinate joint. Underproduction of MUS by PD patients was more apparent at the leading than subordinate joint. The results support joint-specific effect of PD on movement control. They also suggest that dyscoordination of joint motions in PD stems predominantly from impaired control of subordinate joints, while bradykinesia is associated more with control of the leading than subordinate joint. Possible contribution of the revealed impairments in joint control to some other movement features in PD is discussed. The study demonstrates the efficiency of the LJH application for revealing changes in joint control caused by motor disorders.     

Human motor compensations for thixotropy‐dependent changes in muscular resting tension after moderate joint movements

Aim: This study on healthy subjects explores history‐dependent changes in the resting tension of relaxed wrist muscles after moderate joint excursions and the motor control consequences of these changes during voluntary wrist joint position maintenance. Methods: Integrated surface electromyogram (IEMG) was recorded from wrist extensor/flexor muscles. Angular position and torque were recorded from the wrist joint. Changes in wrist flexor muscle resting tension were sensed by a force transducer pressed against the tendons. Results: Consecutive stepwise changes (7.5°) in wrist joint position (within the dorsiflexed range) were either imposed on relaxed subjects or actively performed while the subjects under visual guidance tried to mimic the passive movements. In relaxed subjects, passive joint torque resistance at a given steady dorsiflexed position either gradually declined or rose depending on the direction of the previous transition movements. In corresponding voluntary contraction experiments, the IEMG amplitude from position holding wrist extensors was found to vary in a similar way as the passive torque resistance. Further, there was a strong correlation between history‐dependent changes in extensor IEMG amplitude and stress alterations exhibited by the relaxed antagonist flexors. The above described, slowly subsiding post‐movement mechanical and motor adaptations were accelerated by brief forceful cocontractions of the forearm muscles. Conclusion: Moderate stepwise changes in joint position are sufficient to induce history‐dependent after‐effects in passive muscular resting tension, after‐effects which during voluntary position holding are effectively compensated for by the motor control system.     

Cerebellar subjects show impaired coupling of reach and grasp movements

We examined how cerebellar deficits in isolated reaching or grasping movements contribute to abnormalities in a combined reach and grasp movement, and whether people with cerebellar damage show abnormalities in the spatiotemporal relationships of reach and grasp movements. We studied subjects with cerebellar damage and matched controls as they performed a combined reach and grasp, an isolated reach, and an isolated grasp. These movements were performed under slow-accurate and fast speed conditions. Subjects were also tested for their ability to correctly estimate the target size based on visual information. We measured the three-dimensional position of the index finger, thumb and wrist joint during all tasks. Results showed that cerebellar subjects overestimated the target size to a greater extent than did controls. During movement testing, cerebellar subjects were impaired on isolated reach and isolated grasp. However, they did not worsen parameters of reach or grasp movements during the combined reach and grasp. Instead there were distinct deficits in the coupling of the reach and grasp movement. Compared with controls, cerebellar subjects showed abnormalities in the sequence of the reach and grasp movement and highly variable timing of peak grip aperture. In the slow-accurate condition, cerebellar subjects decomposed the reach and grasp movement into separate reach then grasp components, and produced multiple peaks in grip aperture. In the fast condition, cerebellar subjects did not decompose, produced a single peak grip aperture, and dropped the target more often. These results indicate that cerebellar damage can cause a specific breakdown in the coupling of reach and grasp movements. The cerebellum may be involved in combining reach and grasp movements into a single motor program.     

Electromyographic and biomechanical characteristics of segmental postural adjustments associated with voluntary wrist movements

This study re-investigates the characteristics of segmental postural adjustments associated with rapid mono-articular movements and analyses their dependence on initial postural conditions. Subjects performed rapid voluntary wrist flexions and extensions while maintaining their upper limb posture as stable as possible, with or without an elbow support. Surface electromyographic activity (EMG) was recorded from Flexor carpi ulnaris, Extensor carpi radialis, Biceps brachii, Triceps brachii and Deltoideus anterior. The kinematics of the three joints and kinetics in the support condition were also recorded. A planar mechanical model was used to determine the muscle torque required to keep the upper limb posture constant while performing wrist movements. All subjects showed anticipatory postural adjustments (APA) which, unlike those described for whole-body postural control, could not counteract in advance the perturbing inter-segmental forces created by the movement. Postural muscles were activated before the wrist movement with a chronology specific to the direction of the wrist movement. Some postural muscular activities anticipated that of the prime-movers in accordance with muscle torque, which had to be applied to the joints to keep the upper limb posture constant. These results reveal that the central nervous system (CNS) uses the same organization of the motor command for the control of both segmental and whole-body posture: APA and corrective postural adjustments (CPA), which are based on well-organized anticipatory postural muscle activities (APMA), except that APA can be non-efficient in segmental postural control. The presence or absence of an elbow support influenced the level of activation of postural muscle but not their chronology. This result suggests that the CNS uses a sequence of APMA: a postural muscle synergy which is predetermined as a function of the intended direction of the movements and modulates the gain towards certain muscles, in accordance with the gravitational effects, and supports reaction changes.     

Commonalities and differences in control of various drawing movements

Characteristics of control at the shoulder and elbow during nine types of drawing movements were studied in the present work. The task was to repetitively track a template, depicted on a horizontal table, with the index finger at a cyclic frequency of 1.5 Hz. The templates were a circle, four ovals and four lines of different orientations. The wrist was immobilized and the movement consisted of rotations at the shoulder and elbow joints. The studied movements varied in a wide range with respect to the amplitude of elbow and shoulder movements and relative phase between them. Kinetic analysis included analysis of torque signs, impulses, and timing. It demonstrated that the role of muscle torque in movement production was different at the two joints. During eight out of the nine movement types, the muscle torque at the shoulder accelerated and decelerated this joint and almost completely coped with the influence of the interactive torque arising from elbow motion. Conversely, interactive torque generated by shoulder motion played a dominant role in elbow acceleration and deceleration, whereas muscle torque at the elbow adjusted passive elbow movement to the various template shapes. EMG data were in agreement with the conclusions made from the kinetic analysis. Collectively, these data support the hypothesis that the two joints have different functions in movement production. The shoulder creates a foundation for motion of the entire arm through the interactive torque, and the elbow serves as a fine-tuner of the end-point movement. Control of the shoulder was similar across the eight movement types and the differences in the end-point path were provided by variations in elbow control. The two joints exchanged roles during one movement type, namely, drawing the line tilted right. During this movement, the elbow musculature generated motion at this joint and the shoulder musculature counteracted mechanical influence of this motion on the shoulder position. The findings suggest that during drawing movements, the control strategy exploits intersegmental dynamics of the shoulder-elbow mechanical linkage.     

Multijoint arm movements in cerebellar ataxia: Abnormal control of movement dynamics

In cerebellar ataxia, kinematic aberrations of multijoint movements are thought to originate from deficiencies in generating muscular torques that are adequate to control the mechanical consequences of dynamic interaction forces. At this point the exact mechanisms that lead to an abnormal control of interaction torques are not known. In principle, the generation of inadequate muscular torques may result from an impairment in generating sufficient levels of torques or from an inaccurate assessment and prediction of the mechanical consequences of movements of one limb segment on adjacent joints. We sought to differentiate the relative contribution of these two mechanisms and, therefore, analyzed intersegmental dynamics of multijoint pointing movements in healthy subjects and in patients with cerebellar degeneration. Unrestrained vertical arm movements were performed at three different target movement velocities and recorded using an optoelectronic tracking system. An inverse dynamics approach was employed to compute net joint torques, muscular torques, dynamic interaction torques and gravitational torques acting at the elbow and shoulder joint. In both groups, peak dynamic interaction forces and peak muscular forces were largest during fast movements. In contrast to normal subjects, patients produced hypermetric movements when executing fast movements. Hypermetric movements were associated with smaller peak muscular torques and smaller rates of torque change at elbow and shoulder joints. The patients’ deficit in generating appropriate levels of muscular force were prominent during two different phases of the pointing movement. Peak muscular forces at the elbow were reduced during the initial phase of the movement when simultaneous shoulder joint flexion generated an extensor influence upon the elbow joint. When attempting to terminate the movement, gravitational and dynamic interaction forces caused overshooting extension at the elbow joint. In normal subjects, muscular torque patterns at shoulder and elbow joint were synchronized in that peak flexor and extensor muscular torques occurred simultaneously at both joints. This temporal pattern of muscular torque generation at shoulder and elbow joint was preserved in patients. Our data suggest that an impairment in generating sufficient levels of phasic muscular torques significantly contributes to the patients’ difficulties in controlling the mechanical consequences of dynamic interaction forces during multijoint movements. Received: 28 October 1996 / Accepted: 30 September 1997     

Deficits in movements of the wrist ipsilateral to a stroke in hemiparetic subjects

We examined step-tracking movements of the wrist and associated EMG activity in seven patients (age range, 27-73 yr) and in seven normal subjects that were matched to patients in age, sex, and handedness. All patients exhibited a hemiparesis that resulted from a unilateral cerebrovascular accident (CVA) that included motor areas in the frontal lobe or their efferents. The lesion in three patients was in their dominant hemisphere. The patients were tested 1-48 mo following their CVA. They had great difficulty in performing or were unable to perform step-tracking movements with the contralesional wrist. In addition, the patients displayed striking deficits in wrist movements and muscle activity of the ipsilesional wrist. These movements were >50% slower than those of controls. The initial movement step routinely undershot the target and was only 63% as large as that of controls. The patients made wrist movements with marked directional errors requiring corrective responses. These errors were due largely to inappropriate temporal sequencing of muscle activity. The deficits in movement and muscle activity in the wrist ipsilesional to a CVA were marked, regardless of whether the lesion was in the dominant or nondominant hemisphere. These observations indicate that unilateral lesions can have significant bilateral effects on the generation and control of distal limb movements.     

Influence of joint interactional effects on the coordination of planar two-joint arm movements

We have examined EMG-movement relations in two-joint planar arm movements to determine the influence of interactional torques on movement coordination. Explicitly defined combinations of elbow movements (ranging from 20 to 70°) and wrist movements (ranging from 20 to 40°) were performed during a visual, step-tracking task in which subjects were specifically required to attend to the initial and final angles at each joint. In all conditions the wrist and elbow rotated in the same direction, that is, flexion-flexion or extension-extension. Elbow movement kinematics were only slightly influenced by motion about the wrist. In contrast, the trajectory of the wrist movement was significantly influenced by uncompensated reaction torques resulting from movement about the elbow joint. At any given wrist amplitude, wrist movement duration increased and peak velocity decreased as elbow amplitude increased. In addition, as elbow amplitude increased, wrist movement on-set was progressively delayed relative to this elbow movement. Surprisingly, the changes between joint movement onsets were not accompanied by corresponding changes between agonist EMG onsets at the elbow and wrist joints. The mean difference in onset times between elbow and wrist agonists (22–30 ms) remained unchanged across conditions. In addition, a basic pattern of muscle activation that scaled with movement amplitude was observed at each joint. Phasic agonist activity at the wrist and elbow joints remained remarkably similar across conditions and thus the changes in joint movement onset could not be attributed to changes in the motor commands. Rather, the calculated torques from the averaged data showed that the difference in timing of joint movement onsets was influenced by joint interactional torques. These findings suggest that during simple two-joint planar movements of the elbow and the wrist joint, the central nervous system does not alter the basic motor commands at each joint and as a result the actual trajectory of each joint is determined by interactional torques.     

Adaptation to destabilizing dynamics by means of muscle cocontraction

Adaptive control of wrist mechanics was investigated by means of destabilizing dynamics created by a torque motor. Subjects performed a 20 degrees movement to a 3 degrees target under the constraint that no motion should occur outside of the target zone once 800 ms had elapsed from movement onset. This constraint served as the minimum acceptable level of postural stability. The ability of subjects to modify their muscle activation patterns in order to successfully achieve this stability was investigated by creating three types of destabilizing dynamics with markedly different features: negative stiffness, negative damping, and square-wave vibration. Subjects performed sets of trials with the first type of destabilizing dynamics and were then required to adapt to the second and third. The adaptive response was quantified in terms of the rms electromyographic (EMG) activity recorded during various phases of the task. Surface EMG activity was recorded from three muscles contributing to wrist flexion and three muscles contributing to wrist extension. With negative stiffness, a significant compensatory increase in cocontraction of wrist flexor and extensor muscles was observed for slow movements, but there was little change in the muscle activity for rapid movements. With negative damping, muscle cocontraction was elevated to stabilize rapid movements, declining only gradually after the target was reached. For slow movements, cocontraction occurred only when negative damping was high. The response to square-wave vibration (10 Hz, +/-0.5 Nm), beginning at movement onset, was similar to that of negative damping, in that it resulted in elevated cocontraction. However, because the vibration persisted after the target was reached, there was no subsequent decrease in muscle activity. When the frequency was reduced to 5.5 Hz, but with the same torque impulse, cocontraction increased. This is consistent with greater mechanical instability. In summary, agonist-antagonist cocontraction was adapted to the stability of the task. This generally resulted in less of a change in muscle activity during the movement phase, when the task was performed quickly compared with slowly. On the other hand, the change in muscle activity during stabilization depended more on the nature of the instability than the movement speed.