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     

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.     

Compensation for interaction torques during single- and multijoint limb movement

During multijoint limb movements such as reaching, rotational forces arise at one joint due to the motions of limb segments about other joints. We report the results of three experiments in which we assessed the extent to which control signals to muscles are adjusted to counteract these "interaction torques." Human subjects performed single- and multijoint pointing movements involving shoulder and elbow motion, and movement parameters related to the magnitude and direction of interaction torques were manipulated systematically. We examined electromyographic (EMG) activity of shoulder and elbow muscles and, specifically, the relationship between EMG activity and joint interaction torque. A first set of experiments examined single-joint movements. During both single-joint elbow (experiment 1) and shoulder (experiment 2) movements, phasic EMG activity was observed in muscles spanning the stationary joint (shoulder muscles in experiment 1 and elbow muscles in experiment 2). This muscle activity preceded movement and varied in amplitude with the magnitude of upcoming interaction torque (the load resulting from motion of the nonstationary limb segment). In a third experiment, subjects performed multijoint movements involving simultaneous motion at the shoulder and elbow. Movement amplitude and velocity at one joint were held constant, while the direction of movement about the other joint was varied. When the direction of elbow motion was varied (flexion vs. extension) and shoulder kinematics were held constant, EMG activity in shoulder muscles varied depending on the direction of elbow motion (and hence the sign of the interaction torque arising at the shoulder). Similarly, EMG activity in elbow muscles varied depending on the direction of shoulder motion for movements in which elbow kinematics were held constant. The results from all three experiments support the idea that central control signals to muscles are adjusted, in a predictive manner, to compensate for interaction torques-loads arising at one joint that depend on motion about other joints.     

A novel shoulder–elbow mechanism for increasing speed in a multijoint arm movement

The speed of arm movements is normally increased by increasing agonist muscle activity, but in overarm throwing, an additional effect on speed may come from exploitation of interaction torques (a passive torque associated with motion at adjacent joints). We investigated how the central nervous system (CNS) controls interaction torques at the shoulder and elbow to increase speed in 2-D overarm throwing. Twelve experienced throwers made slow, medium, and fast 2-D throws in a parasagittal plane. Joint motions were computed from recordings made with search coils; joint torques were calculated using inverse dynamics. For slow and medium-speed throws, elbow extension was primarily produced by elbow muscle torque. For fast throws, there was an additional late-occurring elbow extensor interaction torque. Parceling out this elbow extension interaction torque revealed that it primarily arose from shoulder extension deceleration. Surprisingly, shoulder deceleration before ball release was not caused by shoulder flexor (antagonist) muscle torque. Rather, shoulder deceleration was produced by passive elbow-to-shoulder interaction torques that were primarily associated with elbow extension acceleration and velocity. It is concluded that when generating fast 2-D throws, the CNS utilized the arm’s biomechanical properties to increase ball speed. It did this by coordinating shoulder and elbow motions such that an instantaneous mechanical positive feedback occurred of interaction torques between shoulder and elbow before ball release. To what extent this mechanism is utilized in other fast multijoint arm movements remains to be determined.     

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.     

Handedness: dominant arm advantages in control of limb dynamics

Recent findings from our laboratory suggest that a major factor distinguishing dominant from nondominant arm performance is the ability by which the effects of intersegmental dynamics are controlled by the CNS. These studies indicated that the dominant arm reliably used more torque-efficient patterns for movements made with similar speeds and accuracy than nondominant arm movements. Whereas, nondominant hand-path curvatures systematically varied with the amplitude of the interaction torques transferred between the segments of the moving limb, dominant hand-path curvatures did not. However, our previous studies did not distinguish whether dominant arm coordination advantages emerged from more effective control of dynamic factors or were simply a secondary effect of planning different kinematics. The purpose of this study was to further investigate interlimb differences in coordination through analysis of inverse dynamics and electromyography recorded during the performance of reaching movements. By controlling the amplitude of intersegmental dynamics in the current study, we were able to assess whether systematic differences in torque-efficiency exist, even when differences in hand-path shape were minimal. Subject's arms were supported in the horizontal plane by a frictionless air-jet system and were constrained to movements about the shoulder and elbow joints. Two targets were designed, such that the interaction torques elicited at the elbow were either large or small. Our results showed that the former produced large differences in hand-path curvature, whereas the latter did not. Additionally, the movements with small differences in hand-path kinematics showed substantial differences in torque patterns and corresponding EMG profiles which implied a more torque-efficient strategy for the dominant arm. In view of these findings we propose that distinct neural control mechanisms are employed for dominant and nondominant arm movements.     

Coordinated turn-and-reach movements. II. Planning in an external frame of reference

The preceding study demonstrated that normal subjects compensate for the additional interaction torques generated when a reaching movement is made during voluntary trunk rotation. The present paper assesses the influence of trunk rotation on finger trajectories and on interjoint coordination and determines whether simultaneous turn-and-reach movements are most simply described relative to a trunk-based or an external reference frame. Subjects reached to targets requiring different extents of arm joint and trunk rotation at a natural pace and quickly in normal lighting and in total darkness. We first examined whether the larger interaction torques generated during rapid turn-and-reach movements perturb finger trajectories and interjoint coordination and whether visual feedback plays a role in compensating for these torques. These issues were addressed using generalized Procrustes analysis (GPA), which attempts to overlap a group of configurations (e.g., joint trajectories) through translations and rotations in multi-dimensional space. We first used GPA to identify the mean intrinsic patterns of finger and joint trajectories (i.e., their average shape irrespective of location and orientation variability in the external and joint workspaces) from turn-and-reach movements performed in each experimental condition and then calculated their curvatures. We then quantified the discrepancy between each finger or joint trajectory and the intrinsic pattern both after GPA was applied individually to trajectories from a pair of experimental conditions and after GPA was applied to the same trajectories pooled together. For several subjects, joint trajectories but not finger trajectories were more curved in fast than slow movements. The curvature of both joint and finger trajectories of turn-and-reach movements was relatively unaffected by the vision conditions. Pooling across speed conditions significantly increased the discrepancy between joint but not finger trajectories for most subjects, indicating that subjects used different patterns of interjoint coordination in slow and fast movements while nevertheless preserving the shape of their finger trajectory. Higher movement speeds did not disrupt the arm joint rotations despite the larger interaction torques generated. Rather, subjects used the redundant degrees of freedom of the arm/trunk system to achieve similar finger trajectories with differing joint configurations. We examined finger movement patterns and velocity profiles to determine the frame of reference in which turn-and-reach movements could be most simply described. Finger trajectories of turn-and-reach movements had much larger curvatures and their velocity profiles were less smooth and less bell-like in trunk-based coordinates than in external coordinates. Taken together, these results support the conclusion that turn-and-reach movements are controlled in an external frame of reference.     

Fast corrective responses are evoked by perturbations approaching the natural variability of posture and movement tasks

A wealth of studies highlight the importance of rapid corrective responses during voluntary motor tasks. These studies used relatively large perturbations to evoke robust muscle activity. Thus it remains unknown whether these corrective responses (latency 20-100 ms) are evoked at perturbation levels approaching the inherent variability of voluntary control. To fill this gap, we examined responses for large to small perturbations applied while participants either performed postural or reaching tasks. To address multijoint corrective responses, we induced various amounts of single-joint elbow motion with scaled amounts of combined elbow and shoulder torques. Indeed, such perturbations are known to elicit a response at the unstretched shoulder muscle, which reflects an internal model of arm intersegmental dynamics. Significant muscle responses were observed during both postural control and reaching, even when perturbation-related joint angle, velocity, and acceleration overlapped in distribution with deviations encountered in unperturbed trials. The response onsets were consistent across the explored range of perturbation loads, with short-latency onset for the muscles spanning the elbow joints (20-40 ms), and long-latency for shoulder muscles (onset > 45 ms). In addition, the evoked activity was strongly modulated by perturbation magnitude. These results suggest that multijoint responses are not specifically engaged to counter motor errors that exceed a certain threshold. Instead, we suggest that these corrective processes operate continuously during voluntary motor control.     

Horizontal-plane arm movements with direction reversals performed by normal individuals and individuals with down syndrome

We examined the systematic variation in shoulder and elbow torque, as well as movement kinematics, for horizontal-plane arm movements with direction reversals performed by normal individuals and individuals with Down syndrome. Eight neurologically normal individuals and eight individuals with Down syndrome performed horizontal, planar reversal movements to four different target locations. The four locations of the targets were chosen such that there is a systematic increase in elbow interaction torque for each of the four different target locations. This systematic increase in interaction torque has previously been shown to lead to progressively larger movement reversal errors, and trajectories that do not show a sharp reversal of direction, for movements to and from the target in patients who have proprioceptive abnormalities. We computed joint torques at the elbow and shoulder and found a high correlation between elbow and shoulder torque for the neurologically normal subjects. The ratio of joint torques varied systematically with target location. These findings extend previously reported findings of a linear synergy between shoulder and elbow joints for a variety of point-to-point movements. There was also a correlation between elbow and shoulder torque in individuals with Down syndrome, but the magnitude of the correlation was less. The ratio of joint torques changed systematically with target direction in individuals with Down syndrome but was slightly different from the ratio observed for neurologically normal individuals. The difference in the ratio was caused by the generation of proportionately more elbow torque than shoulder torque. The fingertip path of individuals with Down syndrome showed a sharp reversal in moving toward and then away from the target. In this respect, they were similar to neurologically normal individuals but dissimilar to individuals with proprioceptive deficits. Finally, we observed that individuals with Down syndrome spend proportionately more time in the vicinity of the target than normal individuals. Collectively these results show that there is a systematic relationship between joint torques at the elbow and shoulder. This relationship is present for reversal movements and is also present in individuals with Down syndrome.     

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.     

Interjoint coordination dynamics during reaching in stroke

A technique is described that characterizes the dynamics of the interjoint coordination of arm reaching movements in healthy subjects (n=10) and in patients who had sustained a left-sided cerebrovascular accident (n=18). All participants were right-handed. Data from the affected right arm of patients with stroke were compared with those from the right arm of healthy subjects. Seated subjects made 25 pointing movements in a single session. Movements were made from an initial target located ipsilaterally to the right arm beside the body, to a final target located in front of the subject in the contralateral arm workspace. Kinematic data from the finger, wrist, elbow, both shoulders and sternum were recorded in three dimensions at 200 Hz with an optical tracking system. Analysis of interjoint coordination was based on the patterns of temporal delay between rotations at two adjacent joints (shoulder and elbow). The data were reduced to a single graph (Temporal Coordination or TC index) integrating the essential temporal characteristics of joint movement (the angular displacements, velocities and timing). TC segments, duration and amplitude, were analysed. The analysis was sensitive to the differences in interjoint coordination between healthy subjects and patients with arm motor deficits. In patients, the temporal coordination between elbow and shoulder movements was disrupted from the middle to the end of the reach. More specifically, in mid-reach, all patients had difficulty coordinating elbow flexion with shoulder horizontal adduction. In addition, patients with severe arm hemiparesis had difficulty changing elbow movement direction from flexion to extension and in coordinating this change with shoulder movement. At the end of the reach, patients with severe hemiparesis had deficits in the execution of elbow extension while all patients had impaired coordination of elbow extension and shoulder horizontal adduction. In addition, active ranges of joint motions were significantly decreased in the stroke compared to the healthy subjects. Finally, TC analysis revealed significant relationships between specific aspects of disrupted interjoint coordination and the level of motor impairment, suggesting that it may be a useful tool in the identification of specific movement coordination deficits in neurological impaired populations that can be targeted in treatment for arm motor recovery.     

Multijoint movement control in Parkinson's disease

Impairments in the performance of complex actions in Parkinson's disease (PD) patients are well documented. The aim of the present study was to investigate potential mechanisms that may be contributing to impaired movement performance in PD patients. PD patients and age-matched control subjects performed rapid pointing movements to a series of four tabletop targets. The height of the table was adjusted until the targets could be achieved with arm movements in the horizontal plane. The targets were arranged such that target 1 required elbow extension only and targets 2–4 required increasing amounts of horizontal shoulder flexion in addition to the elbow extension. While the control subjects accelerated and decelerated the elbow and shoulder joints simultaneously regardless of the target location, the PD patients decomposed motion during the acceleration phase by accelerating first the shoulder and then the elbow joint. For PD patients this decomposition of arm segments was associated with greater coactivation of the muscles about the elbow when elbow extension and shoulder flexion were simultaneously required (targets 2–4), in contrast to the single joint action. The control subjects decreased elbow joint coactivation while the patients increased it across the four targets. The resulting peak interaction torques at both the elbow and shoulder joints occurred relatively later for the PD patients. The coactivation patterns observed in PD patients may reduce the ability to take advantage of interaction torques and may also contribute to joint motion decomposition. Electronic Publication     

Control of 3D limb dynamics in unconstrained overarm throws of different speeds performed by skilled baseball players

This study investigated how the human CNS organizes complex three-dimensional (3D) ball-throwing movements that require both speed and accuracy. Skilled baseball players threw a baseball to a target at three different speeds. Kinematic analysis revealed that the fingertip speed at ball release was mainly produced by trunk leftward rotation, shoulder internal rotation, elbow extension, and wrist flexion in all speed conditions. The study participants adjusted the angular velocities of these four motions to throw the balls at three different speeds. We also analyzed the dynamics of the 3D multijoint movements using a recently developed method called "nonorthogonal torque decomposition" that can clarify how angular acceleration about a joint coordinate axis (e.g., shoulder internal rotation) is generated by the muscle, gravity, and interaction torques. We found that the study participants utilized the interaction torque to generate larger angular velocities of the shoulder internal rotation, elbow extension, and wrist flexion. To increase the interaction torque acting at these joints, the ball throwers increased muscle torque at the shoulder and trunk but not at the elbow and wrist. These results indicates that skilled ball throwers adopted a hierarchical control in which the proximal muscle torques created a dynamic foundation for the entire limb motion and beneficial interaction torques for distal joint rotations.     

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.     

Kinematics and kinetics of multijoint reaching in nonhuman primates

The present study identifies the mechanics of planar reaching movements performed by monkeys (Macaca mulatta) wearing a robotic exoskeleton. This device maintained the limb in the horizontal plane such that hand motion was generated only by flexor and extensor motions at the shoulder and elbow. The study describes the kinematic and kinetic features of the shoulder, elbow, and hand during reaching movements from a central target to peripheral targets located on the circumference of a circle: the center-out task. While subjects made reaching movements with relatively straight smooth hand paths and little variation in peak hand velocity, there were large variations in joint motion, torque, and power for movements in different spatial directions. Unlike single-joint movements, joint kinematics and kinetics were not tightly coupled for these multijoint movements. For most movements, power generation was predominantly generated at only one of the two joints. The present analysis illustrates the complexities inherent in multijoint movements and forms the basis for understanding strategies used by the motor system to control reaching movements and for interpreting the response of neurons in different brain regions during this task.     

Utilization and compensation of interaction torques during ball-throwing movements

The manner in which the CNS deals with interaction torques at each joint in ball throwing was investigated by instructing subjects to throw a ball at three different speeds, using two (elbow and wrist) or three joints (shoulder, elbow, and wrist). The results indicated that the role of the muscle torque at the most proximal joint was to accelerate the most proximal joint and to produce the effect of interjoint interaction on the distal joints. In the three-joint throwing, shoulder muscle torque produced the assistive interaction torque for the elbow, which was effectively utilized to generate large elbow angular velocity when throwing fast. However, at the wrist, the muscle torque always counteracted the interaction torque. By this kinetic mechanism, the wrist angular velocity at the ball-release time was kept relatively constant irrespective of ball speed, which would lead to an accurate ball release. Thus it was concluded that humans can adjust the speed and accuracy of ball-throwing by utilizing interaction torque or compensating for it.     

The leading joint hypothesis for spatial reaching arm motions

The leading joint hypothesis (LJH), developed for planar arm reaching, proposes that the interaction torques experienced by the proximal joint are low compared to the corresponding muscle torques. The human central nervous system could potentially ignore these interaction torques at the proximal (leading) joint with little effect on the wrist trajectory, simplifying joint-level control. This paper investigates the extension of the LJH to spatial reaching. In spatial motion, a number of terms in the governing equation (Euler’s angular momentum balance) that vanish for planar movements are non-trivial, so their contributions to the joint torque must be classified as net, interaction or muscle torque. This paper applies definitions from the literature to these torque components to establish a general classification for all terms in Euler’s equation. This classification is equally applicable to planar and spatial motion. Additionally, a rationale for excluding gravity torques from the torque analysis is provided. Subjects performed point-to-point reaching movements between targets whose locations ensured that the wrist paths lay in various portions of the arm’s spatial workspace. Movement kinematics were recorded using electromagnetic sensors located on the subject’s arm segments and thorax. The arm was modeled as a three-link kinematic chain with idealized spherical and revolute joints at the shoulder and elbow. Joint torque components were computed using inverse dynamics. Most movements were ‘shoulder-led’ in that the interaction torque impulse was significantly lower than the muscle torque impulse for the shoulder, but not the elbow. For the few elbow-led movements, the interaction impulse at the elbow was low, while that at the shoulder was high, and these typically involved large elbow and small shoulder displacements. These results support the LJH and extend it to spatial reaching motion.     

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.     

Evidence for a dynamic-dominance hypothesis of handedness

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

Proprioceptive control of multijoint movement: unimanual circle drawing

The present experiments addressed whether proprioception is used by the central nervous system (CNS) to control the spatial and temporal characteristics of unimanual circle drawing. Circle drawing is a multijoint movement, in which the muscles crossing the elbow and the shoulder are sequentially activated. The spatial and temporal characteristics of circle drawing depend on the precise coordination of these sequential activation patterns, and proprioception is ideally suited to support this coordination. Blindfolded human subjects produced a counterclockwise circular drawing motion (diameter = 16 cm) with the dominant arm at a repetition rate of 1/s. In some trials, 60–70 Hz vibration was applied to the tendons of the biceps brachii and/or the anterior deltoid. Spatial parameters measured from hand-movement data included the x- and y-axis diameters, circularity, and drift of the hand in the workspace. Vibration of either the biceps brachii or the anterior deltoid caused subjects to draw circles with decreased diameter, with changes in circularity, and with a systematic drift of the hand. These distortions to circle drawing by tendon vibration demonstrate that the CNS uses proprioceptive information to accomplish the spatial characteristics of this motor task. Simultaneous vibration of both muscles produced a drift that exceeded the individual vibration effects, which suggests that the CNS combined proprioceptive information related to elbow and shoulder rotation to control the movement of the hand. The temporal characteristics of circle drawing were quantified from joint angle data. While vibration did not significantly influence the relative phase between elbow and shoulder rotation, the variability of the phase relationship increased significantly, which suggests that proprioception contributes to phase stabilization. During circle drawing, elbow flexion-extension movements were produced with limited activation of the biceps. Nevertheless, biceps vibration distorted the circle metrics, suggesting that a muscle’s significance as a sensory transducer is independent of its activity level. Received: 29 November 1997 / Accepted: 16 February 1999