Time-dependent adaptations to posture and movement characteristics during the development of repetitive reaching induced fatigue

Repetitive movements are common to many daily activities but often lead to the development of fatigue. We have previously shown that fatigue leads to changes in tridimensional spatial characteristics of the whole body. However, temporal aspects of these posture and movement adaptations have yet to be investigated. Healthy subjects (N = 14) performed a continuous reaching task by pointing between two targets placed at shoulder height, at 100 and 30% arm’s length, anterior to the subject’s midline until fatigue (assessed using the Borg CR-10 scale). Whole body kinematics and upper Trapezius EMG were recorded and analyzed at 1-min intervals to document the progression of fatigue on outcome variables. For all upper limb and postural variables analyzed, changes began to occur approximately midway to fatigue and were followed by an increase in Trapezius activity from baseline. Reach-to-reach variability of joint average positions and range of motion (ROM) increased in multiple directions for shoulder and elbow parameters. Reach-to-reach variability of the center-of-mass ROM also increased in several directions. Changes were also observed in within-movement inter-segmental timing. The peak velocities of elbow and endpoint occurred closer together in time during fatigue while the shoulder peak velocity occurrence showed a greater reach-to-reach variability. Our results suggest that the effects of fatigue on repetitive movement kinematics can be observed across three temporal dimensions of the task: (1) within individual movements, (2) from one movement to the next, and (3) as fatigue develops. Each observed change is discussed as a potential contributor to task-specific control strategies to prolong task performance.     

Repetitive arm motion-induced fatigue affects shoulder but not endpoint position sense

Neck/shoulder pain has previously been linked to repetitive work and muscle fatigue. We have shown that asymptomatic people performing repetitive upper limb tasks display signs of shoulder fatigue and of whole-body compensatory strategies. However, the role played by the proprioceptive system in the production of these compensatory strategies has not been studied. A group of asymptomatic adults (n = 18) performed a repetitive pointing task at shoulder height to fatigue. Before and after fatigue, they performed two position sense tasks, eyes closed: a single-joint task where they abducted their shoulder to the perceived horizontal and a multi-joint task, where they stood and placed their finger at the perceived location of a target in front of them at shoulder height. After fatigue, subjects made larger shoulder errors by raising their elbow higher above the horizontal (~+1.3 cm) than before fatigue; however, their finger position accuracy was not changed, despite all subjects performing the movement in less time (~−0.18 s) while fatigued. There were no gender differences in shoulder or finger position accuracy before or after fatigue; however, there were gender differences in the perceived finger-target location and in the temporal characteristics of the finger movement toward the target. Results suggest that healthy individuals are able to develop strategies to compensate for fatigue-induced deficits at one joint to maintain the endpoint accuracy of a multi-joint task constant. Gender differences in movement strategies and perception of endpoint location may play parts in the previously reported gender differences in work-related neck/shoulder symptoms.     

Movement timing and reach to reach variability during a repetitive reaching task in persons with chronic neck/shoulder pain and healthy subjects

Neck/shoulder pain is linked to movement repetition, awkward postures, and muscular fatigue. Studies have examined the influence of pain and fatigue on movement characteristics, but few report spatial and temporal characteristics within- and between-dynamic movements. The combined influences of fatigue and pain on these parameters are examined here. A shoulder-injured group (PAIN) (intensity >3/10, duration >3 consecutive months) and an age-sex-matched control group (CTRL) (n = 16 in each) performed a repetitive reaching task to voluntary termination. Kinematics, heart rate, and muscle activity were recorded. Group comparisons were made at the beginning and end of task. Both pain and fatigue changed movement parameters with CTRL subjects adapting to fatigue with increased arm movement, while the PAIN group increased center of mass (CoM) and minimized shoulder movement. Fatigue was associated with decreased arm relative variability, while pain caused increased arm and decreased CoM relative variability. Time to peak velocity tended to indicate increased joint coupling in the CTRL group only. Together, this suggests that despite initial differences in movement strategies, both groups tend to move toward more fixed movement strategies at the end of the task. Those of CTRL are more fixed temporally and spatially (in the arm), whereas the PAIN group adaptations are primarily spatial in nature and appear to focus on controlling CoM.     

The effect of movement direction on joint torque covariation

It has been proposed that unconstrained upper limb movements are coordinated via a kinetic constraint that produces dynamic muscle torques at each moving joint that are a linear function of a single torque command. This constraint has been termed linear synergy (Gottlieb et al. J Neurophysiol 75:1760–1764, 1996). The current study tested two hypotheses: (1) that the extent of covariation between dynamic muscle torques at the shoulder and elbow varied with the direction of movement and (2) that the extent to which muscle torques deviated from linear synergy would be reproduced by a simulation of pointing movements in which the path of the hand was constrained to be straight. Dynamic muscle torques were calculated from sagittal plane pointing movements performed by 12 participants to targets in eight different directions. The results of principal component analyses performed on the muscle torque data demonstrated direction-dependent variation in the extent to which dynamic muscle torques covaried at the shoulder and elbow. Linear synergy was deviated from substantially in movement directions for which the magnitude of muscle torque was low at one joint. A simulation of movements with straight hand paths was able to accurately estimate the amount of covariation between muscle torques at the two joints in many directions. These results support the idea that a kinematic constraint is imposed by the central nervous system during unconstrained pointing movements. Linear synergy may also be applied as a coordinating constraint in circumstances where its application allows the path of the moving endpoint to remain close to a straight line.     

The relationship between muscle kinetic parameters and kinematic variables in a complex movement

Summary Kinematic variables of the vertical jump (jumping height, jump phase durations and joint angles) were measured on 39 male physical education students. In addition, kinetic parameters of the hip and knee extensors, and of the plantar flexors (maxima voluntary force and its rate of development) were recorded on the same subjects, in isometric conditions. The results demonstrated significant positive correlations between kinetic parameters of the active muscle groups and jumping height (r=0.217−0.464). The dominant effect on these correlations was due to the knee extensors. Correlations between these parameters and the duration of the jump phases were much weaker. Correlation coefficients between kinetic parameters and limb angles in the lowest body position showed that fast force production in one muscle group was related to a significant decrease in the joint angles of distant body segments. Multiple correlation coefficients between leg extensor parameters and kinematic variables (ranging between 0.256 for the duration of the counter-movement phase and 0.616 for jump height) suggested that kinetic parameters could explain more than a quarter of the variability of this complex human movement. Therefore, the conclusion was drawn that an extended set of measurements of the relevant musculo-skeletal system parameters could predict a considerable amount of the variability of human movement. However, high correlation coefficients between the same kinetic parameters of different muscle groups suggest that not all active muscle groups have to be included in the measurements.     

The effects of neuromuscular fatigue on task performance during repetitive goal-directed movements

Proper movement timing is essential to the successful execution of many motor tasks and may be adversely affected by muscle fatigue. This study quantified how muscle fatigue affected task performance during a repetitive upper extremity task. A total of 14 healthy young adults pushed a low load back and forth along a low-friction horizontal track in time with a metronome until volitional exhaustion. Kinematic, force, and electromyography (EMG) data were measured continuously throughout the task. The first and last 3.5 min were analyzed to represent “early” and “late” fatigue. Means and standard deviations of movement distance, speed, and timing errors were computed. We also decomposed variations in movement distance and speed into deviations that directly affected achieving the task goal and those that did not, by identifying the goal equivalent manifold (GEM) of all valid solutions to this task. Detrended fluctuation analysis was used to quantify the temporal persistence in each time series. Principle components analysis provided a direct measure of alignment with the GEM. Median power frequencies of the EMG significantly decreased in six of the nine muscles tested indicating that subjects did fatigue. However, there were no differences in the means or variability of movement distance, speed, or timing errors. Thus, subjects maintained overall performance despite fatigue. Subjects applied slightly higher peak handle forces when they were fatigued (P = 0.032). Muscle fatigue caused significant reductions in the temporal persistence of movement speed (P = 0.037) and timing errors (P = 0.046), indicating that subjects corrected errors more quickly when fatigued. Mean deviations and variability perpendicular to the GEM were much smaller than variability along the GEM (P < 0.001). Deviations perpendicular to the GEM were also corrected much more rapidly than those along the GEM (P < 0.001). Subjects aligned themselves very closely (<±7°), but not exactly (P < 0.001), with the GEM. These measures were not significantly affected by muscle fatigue. Overall, these results indicated that subjects altered their biomechanical movement patterns in response to muscle fatigue, but did so in a way that specifically preserved the goal relevant features of task performance.     

Patterns of coordinated multi-joint movement

In multi-joint reaching movements, the motor system may choose any one of an infinite set of possible joint rotations to move the hand between given start and target positions. In order to find out whether reaching movements are represented in Cartesian hand coordinates or in joint coordinates, it is necessary to measure whether hand paths or joint paths have lower variability. We have measured hand paths and rotations of shoulder, elbow and wrist joints simultaneously in five subjects reaching in four orientations in the horizontal plane. As in earlier studies, we found a preference for nearly straight hand paths, despite different patterns of joint rotation for different orientations of movement. However, movements in three of four orientations showed a single principal joint, which rotated essentially without reversals. This may reflect optimisation in the motor system, preferring the simplest pattern of joint control for a desired hand path. We used generalised Procrustes analysis to quantify the variability in shape of repeated paths in hand space and joint space. Results showed that hand paths were less variable than the joint angles used to realise them, due to the kinematic redundancy of the limb, suggesting that hand paths, rather than joint angles, are directly represented by the motor system. Nevertheless, movements with straighter hand paths, on average, and those requiring coordinated activity at both shoulder and elbow joints also showed more variability in the shape of the hand path. Other orientations such as movement across the body use primarily a single joint and are less variable at the cost of a slightly curved path. These results suggest that coordinating multiple joints to produce a straight hand path has a definite computational cost. The motor system may perform a trade-off between the benefits of planning reaching movements as straight hand paths and the computational simplicity of executing them using patterns of joint rotation which simplify multi-joint coordination.     

Independent coactivation of shoulder and elbow muscles

 The aim of this study was to examine the possibility of independent muscle coactivation at the shoulder and elbow. Subjects performed rapid point-to-point movements in a horizontal plane from different initial limb configurations to a single target. EMG activity was measured from flexor and extensor muscles acting at the shoulder (pectoralis clavicular head and posterior deltoid) and elbow (biceps long head and triceps lateral head) and flexor and extensor muscles acting at both joints (biceps short head and triceps long head). Muscle coactivation was assessed by measuring tonic levels of electromyographic (EMG) activity after limb position stabilized following the end of the movements. It was observed that tonic EMG levels following movements to the same target varied as a function of the amplitude of shoulder and elbow motion. Moreover, for the movements tested here, the coactivation of shoulder and elbow muscles was found to be independent – tonic EMG activity of shoulder muscles increased in proportion to shoulder movement, but was unrelated to elbow motion, whereas elbow and double-joint muscle coactivation varied with the amplitude of elbow movement and were not correlated with shoulder motion. In addition, tonic EMG levels were higher for movements in which the shoulder and elbow rotated in the same direction than for those in which the joints rotated in opposite directions. In this respect, muscle coactivation may reflect a simple strategy to compensate for forces introduced by multijoint limb dynamics. Received: 7 July 1998 / Accepted: 28 July 1998     

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.     

Shoulder and elbow muscle activity in goal-directed arm movements

 This research examined the electromyographic (EMG) activity of shoulder and elbow muscles during reaching movements of the upper limb. Subjects performed goal-directed arm movements in the horizontal plane. Movements which varied in amplitude, speed, and direction were performed in different sections of the workspace. EMG activity was recorded from the pectoralis major, posterior deltoid, biceps brachii short head, brachioradialis, triceps brachii long head, and triceps brachii lateral head; motion recordings were obtained with an optoelectric system. The analysis focused on the magnitude and timing of opposing muscle groups at the shoulder and elbow joints. For hand movements within any given direction of the workspace direction, kinematic manipulations changed agonist and antagonist EMG magnitude and intermuscle timing in a manner consistent with previous single-joint findings. To produce reaching movements in different directions and areas of the workspace, shoulder and elbow agonist EMG magnitude increased for those hand motions which required higher angular velocities, while the timing between opposing muscle groups at each joint was invariant. Received: 11 January 1996 / Accepted: 24 February 1997     

Pointing in 3D space to remembered targets II: Effects of movement speed toward kinesthetically defined targets

The accuracy of visually guided pointing movements decreases with speed. We have shown that for movements to a visually defined remembered target, the variability of the final arm endpoint position does not depend on movement speed. We put forward a hypothesis that this observation can be explained by suggesting that movements directed at remembered targets are produced without ongoing corrections. In the present study, this hypothesis was tested for pointing movements in 3D space to kinesthetically defined remembered targets. Passive versus active acquisition of kinesthetic information was contrasted. Pointing errors, movement kinematics, and joint-angle coordination were analyzed. The movements were performed at a slow speed (average peak tangential velocity of about 1.2 m/s) and at a fast speed (2.7 m/s). No visual feedback was allowed during the target presentation or the movement. Variability in the final position of the arm endpoint did not increase with speed in either the active or the passive condition. Variability in the final values of the arm-orientation angles determining the position of the forearm and of the upper arm in space was also speed invariant. This invariance occurred despite the fact that angular velocities increased by a factor of two for all the angles involved. The speed-invariant variability supports the hypothesis that there is an absence of ongoing corrections for movements to remembered targets: in the case of a slower movement, where there is more time for movement correction, the final arm endpoint variability did not decrease. In contrast to variability in the final endpoint position, the variability in the peak tangential acceleration increased significantly with movement speed. This may imply that the nervous system adopts one of two strategies: either the final endpoint position is not encoded in terms of muscle torques or there is a special on-line mechanism that adjusts movement deceleration according to the muscle-torque variability at the initial stage of the movement. The final endpoint position was on average farther from the shoulder than the target. Constant radial-distance errors were speed dependent in both the active and the passive conditions. In the fast speed conditions, the radial distance overshoots of the targets increased. This increase in radial-distance overshoot with movement speed can be explained by the hypothesis that the final arm position is not predetermined in these experimental conditions, but is defined during the movement by a feedforward or feedback mechanism with an internal delay.     

The Ia afferent feedback of a given movement evokes the illusion of the same movement when returned to the subject via muscle tendon vibration

The aim of the present study was to further investigate the contribution of primary muscle spindle feedback to proprioception and higher brain functions, such as movement trajectory recognition. For this purpose, complex illusory movements were evoked in subjects by applying patterns of muscle tendon vibration mimicking the natural Ia afferent pattern. Ia afferent messages were previously recorded using microneurographic method from the six main muscle groups acting on the ankle joint during imposed “writing like” movements. The mean Ia afferent pattern was calculated for each muscle group and used as a template to pilot each vibrator. Eleven different vibratory patterns were applied to ten volunteers. Subjects were asked both to copy the perceived illusory movements by hand on a digitizing tablet and to recognize and name the corresponding graphic symbol. The results show that the Ia afferent feedback of a given movement evokes the illusion of the same movement when it is applied to the subject via the appropriate pattern of muscle tendon vibration. The geometry and the kinematic parameters of the imposed and illusory movements are very similar and the so-called “two-thirds power law” is present in the reproduction of the vibration-induced illusory movements. Vibrations within the “natural” frequency range of Ia fibres firing (around 30 Hz) produce clear illusions of movements in all the tested subjects. In addition, increasing the mean frequency of the vibration patterns resulted in a linear increase in the size of the illusory movements. Lastly, the subjects were able to recognize and name the symbols evoked by the vibration-induced primary muscle spindle afferent patterns in 83% of the trials. These findings suggest that the “proprioceptive signature” of a given movement is associated with the corresponding “perceptual signature”. The neural mechanisms possibly underlying the sensory to perceptual transformation are discussed in the general framework of “the neuronal population vector model”.     

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     

Multi-joint limbs permit a flexible response to unpredictable events

The human arm is kinematically redundant, which may allow flexibility in the execution of reaching movements. We have compared reaching movements with and without kinematic redundancy to unpredictable double-step targets. Subjects sat in front of a digitising tablet and were able to view an arc of four targets reflected in the mirror as virtual images in the plane of the tablet. They were instructed to move, from a central starting point, in as straight a line as possible to a target. In one-third of trials, the target light switched to one of its neighbours during the movement. Subjects made 60 movements using shoulder, elbow and wrist and then another 60 movements in which only shoulder and elbow movement were allowed. By restraining the wrist, the limb was made non-redundant. The path length was calculated for each movement. In single-step trials, there was no significant difference between path lengths performed with and without wrist restraint. As expected there was a significant increase in path length during double-step trials. Moreover this increase was significantly greater when the wrist was restrained. The variability across both single- and double-step movements was significantly less while the wrist was restrained. Importantly the performance time of the movements did not alter significantly for single-step, double-step or restrained movements. These results suggest that the nervous system exploits the intrinsic redundancy of the limb when controlling voluntary movements and is therefore more effective at reprogramming movements to double-step targets. Received: 24 March 1997 / Accepted: 7 July 1997     

Effects of altering initial position on movement direction and extent

The purpose of this study was to examine the relative influence of initial hand location on the direction and extent of planar reaching movements. Subjects performed a horizontal-plane reaching task with the dominant arm supported above a table top by a frictionless air-jet system. A start circle and a target were reflected from a horizontal projection screen onto a horizontally positioned mirror, which blocked the subject's view of the arm. A cursor, representing either actual or virtual finger location, was only displayed between each trial to allow subjects to position the cursor in the start circle. Prior to occasional "probe trials," we changed the start location of the finger relative to the cursor. Subjects reported being unaware of the discrepancy between cursor and finger. Our results indicate that regardless of initial hand location, subjects did not alter the direction of movement. However, movement distance was systematically adjusted in accord with the baseline target position. Thus when the hand start position was perpendicularly displaced relative to the target direction, neither the direction nor the extent of movement varied relative to that of baseline. However, when the hand was displaced along the target direction, either anterior or posterior, movements were made in the same direction as baseline trials but were shortened or lengthened, respectively. This effect was asymmetrical such that movements from anterior displaced positions showed greater distance adjustment than those from posterior displaced positions. Inverse dynamic analysis revealed substantial changes in elbow and shoulder muscle torque strategies for both right/left and anterior/posterior pairs of displacements. In the case of right/left displacements, such changes in muscle torque compensated changes in limb configuration such that movements were made in the same direction and to the same extent as baseline trials. Our results support the hypothesis that movement direction is specified relative to an origin at the current location of the hand. Movement extent, on the other hand, appears to be affected by the workspace learned during baseline movement experience.     

Shoulder muscle strength in paraplegics before and after kayak ergometer training

The purpose was to investigate if shoulder muscle strength in post-rehabilitated persons with spinal cord injury (SCI) was affected by kayak ergometer training and to compare shoulder strength in persons with SCI and able-bodied persons. Ten persons with SCI (7 males and 3 females, injury levels T3–T12) performed 60 min kayak ergometer training three times a week for 10 weeks with progressively increased intensity. Maximal voluntary concentric contractions were performed during six shoulder movements: flexion and extension (range of motion 65°), abduction and adduction (65°), and external and internal rotation (60°), with an angular velocity of 30° s⁻¹. Position specific strength was assessed at three shoulder angles (at the beginning, middle and end of the range of motion) in the respective movements. Test–retests were performed for all measurements before the training and the mean intraclass correlation coefficient was 0.941 (95% CI 0.928–0.954). There was a main effect of kayak ergometer training with increased shoulder muscle strength after training in persons with SCI. The improvements were independent of shoulder movement, and occurred in the beginning and middle positions. A tendency towards lower shoulder muscle strength was observed in the SCI group compared to a matched reference group of able-bodied persons. Thus, it appears that post-rehabilitated persons with SCI have not managed to fully regain/maintain their shoulder muscle strength on a similar level as that of able-bodied persons, and are able to improve their shoulder muscle strength after a period of kayak ergometer training.     

Unimanual and bimanual voluntary movement in Huntington's disease

Unimanual and bimanual cyclical forearm movements were studied in 15 Huntington's disease (HD) patients and 15 healthy, gender- and age-matched controls. Whereas the unimanual task was only performed at maximal speed, the bimanual movements were performed according to the in-phase and anti-phase mode at different cycling frequencies. The HD patients also performed the tasks after 12 months of follow-up. Findings revealed that maximal cycling frequency during unimanual movement was significantly lower in HD patients as compared with controls. In addition, measures of relative phasing established that bimanual cyclical movements were performed with lower accuracy and higher variability in HD patients. The differential variability between both groups was magnified by increasing the cycling frequency and coordinative complexity whereas only coordinative complexity differentially affected the accuracy of relative phasing. The obtained performance measures were found to be significantly correlated with disease duration (unimanual) and with the score on the total motor scale, the Mini-Mental State Examination and the Stroop Interference Test (uni- and bimanual). After 12 months, maximal cycling frequency of unimanual elbow flexion–extension was significantly decreased in HD patients whereas the quality of the in-phase and anti-phase movement patterns remained stable. Electronic Publication     

Directional invariance during loading-related modulations of muscle activity: evidence for motor equivalence

In the present study, we investigated the influence of external force manipulations on movements in different directions, while keeping the amplitude invariant. Subjects (n=10) performed a series of cyclical anteroposterior, mediolateral, and oblique line-drawing movements (star drawing task) with their dominant limb in the horizontal plane. To dissociate kinematics from the underlying patterns of muscle activation, spring loading was applied to the forearm of the moving limb. Whereas spring loading of the arm resulted in considerable changes in the overall amount of muscle activation in the elbow and shoulder muscles, invariance was largely maintained at the kinematic level. Subjects produced the required movement directions and amplitudes of the star drawing largely successfully, irrespective of the force bias induced by the spring. These observations demonstrate motor equivalence and strengthen the notion that the spatial representation of drawing movements is encoded in the higher brain regions in a rather abstract form that is dissociated from the concrete muscle activation patterns underlying a particular movement direction. To achieve this goal, the central nervous system shifted between two or more muscle grouping strategies to overcome modulations in the interaction among posture-dependent (joint stiffness), dynamic (inertial), and elastic (spring) torque components in the joints. Spring loading induced general changes in the overall amount of EMG activity, which was largely muscle but not direction specific, presumably to represent the posture-dependent biasing force of the spring. Loading was mainly shown to increase muscle coactivation in the elbow joint. This indicates that the subjects tended to increase stiffness in the elbow to compensate for changes in the spring bias forces in order to minimize trajectory errors. Changes in muscle grouping of the shoulder antagonists were mainly a consequence of movement direction but were also affected partly by loading, presumably reflecting the influence of dynamic force components. Taken together, the results confirmed the hypothesis that changes of movement direction and direction of force in the end-effector generated specific sets of muscle grouping to overcome the dynamic requirements in the joints while keeping the kinematics largely unchanged. This suggests that directional tuning in muscle activity and changes in muscle grouping reflects the formation of appropriate internal models in the CNS that give rise to motor equivalence. Electronic Publication     

Kinematic variability of grasp movements as a function of practice and movement speed

Summary Grasp movements were studied in six female subjects to determine the effects of practice and movement speed on kinematics and movement variability. Subjects performed four-joint pinch movements of the index finger and thumb, with 200 repetitions at each of three durations (100, 200, and 400 ms). As observed previously, movements of high velocity were performed with bell-shaped, single-peaked velocity profiles. In contrast, slower movements (200, 400 ms) were performed as a series of two to four submovements with multiple peaks in the associated joint angular velocity profiles. With practice, only the slowest movements (400 ms duration) showed significant reductions in variability of joint end-positions. Surprisingly, variability of finger and thumb joint end-positions did not increase with increasing movement speed as has been observed for arm pointing movements. This was apparently due to reductions in positional variability during deceleration of the movement which offset increases in positional variability during acceleration. Neither practice nor movement speed affected variability of the location of fingertip contact on the thumb, which always occurred on the thumb distal pulpar surface.     

Movement reorganization to compensate for fatigue during sawing

Peripheral (muscle) aspects of fatigue are well documented. However, little is known about the central aspects of fatigue that could influence, in particular, multijoint coordination. To investigate the central aspects of fatigue, we compared the multijoint kinematics of non-fatigued and fatigued individuals while sawing. Muscle fatigue was associated with decreases in sawing force and movement amplitude at the elbow whereas the basic characteristics of the saw trajectory, including the movement direction, extent and duration, remained invariant. This invariance was maintained by increasing the movement amplitude at the wrist, shoulder and trunk. The system thus takes advantage of the redundancy of the motor apparatus to maintain the endpoint trajectory despite fatigue.