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Deep brain stimulation of the subthalamic nucleus (STN DBS) significantly improves clinical motor symptoms, as well as intensive aspects of movement like velocity and amplitude in patients with Parkinson's disease (PD). However, the effects of bilateral STN DBS on integrative and coordinative aspects of motor control are equivocal. The aim of this study was to investigate the effects of bilateral STN DBS on integrative and coordinative aspects of movement using a memory-guided sequential reaching task. The primary outcomes were eye and finger velocity and end-point error. We expected that bilateral STN DBS would increase reaching velocity. More importantly, we hypothesized that bilateral STN DBS would increase eye and finger end-point error and this would not simply be the result of a speed accuracy trade-off. Ten patients with PD and bilaterally implanted subthalamic stimulators performed a memory-guided sequential reaching task under four stimulator conditions (DBS-OFF, DBS-LEFT, DBS-RIGHT, and DBS-BILATERAL) over 4 days. DBS-BILATERAL significantly increased eye velocity compared to DBS-OFF, DBS-LEFT, and DBS-RIGHT. It also increased finger velocity compared to DBS-OFF and DBS-RIGHT. DBS-BILATERAL did not change eye end-point error. The novel finding was that DBS-BILATERAL increased finger end-point error compared to DBS-OFF, DBS-LEFT, and DBS-RIGHT even after adjusting for differences in velocity. We conclude that bilateral STN DBS may facilitate basal ganglia-cortical networks that underlie intensive aspects of movement like velocity, but it may disrupt selective basal ganglia-cortical networks that underlie certain integrative and coordinative aspects of movement such as spatial accuracy.

Priming can be described as behavior change generated by preceding stimuli. Although various types of priming have been long studied in the field of psychology, priming that targets motor cortex is a relatively new topic of research in the fields of motor control and rehabilitation. In reference to a rehabilitation intervention, priming is categorized as a restorative approach. There are a myriad of possible priming approaches including noninvasive brain stimulation, motor imagery, and sensory-based priming, to name a few. The authors report on movement-based priming which, compared to other priming types, is less frequently examined and under reported. Movement-based priming includes, but is not limited to, bilateral motor priming, unilateral priming, and aerobic exercise. Clinical and neural mechanistic aspects of movement-based priming techniques are explored.

Eye–hand coordination is a crucial element of goal-directed movements. However, few studies have looked at the extent to which unconstrained movements of the eyes and hand made to targets influence each other. We studied human participants who moved either their eyes or both their eyes and hand to one of three static or flashed targets presented in 3D space. The eyes were directed, and hand was located at a common start position on either the right or left side of the body. We found that the velocity and scatter of memory-guided saccades (flashed targets) differed significantly when produced in combination with a reaching movement than when produced alone. Specifically, when accompanied by a reach, peak saccadic velocities were lower than when the eye moved alone. Peak saccade velocities, as well as latencies, were also highly correlated with those for reaching movements, especially for the briefly flashed targets compared to the continuous visible target. The scatter of saccade endpoints was greater when the saccades were produced with the reaching movement than when produced without, and the size of the scatter for both saccades and reaches was weakly correlated. These findings suggest that the saccades and reaches made to 3D targets are weakly to moderately coupled both temporally and spatially and that this is partly the result of the arm movement influencing the eye movement. Taken together, this study provides further evidence that the oculomotor and arm motor systems interact above and beyond any common target representations shared by the two motor systems.

Simple movements can be seen as building blocks for complex action sequences, and neural control of an action sequence can be expected to preserve some control features of its constituent blocks. It was previously found that during single-joint elbow movements to a single target, the proprioceptive feedback control is initially suppressed, and we tested this feedback suppression in a two-segment sequence during which subjects momentarily slowed down at an intermediate target at a 30° distance (first segment) and then immediately moved another 30° to the final target (second segment). Either the first or second segment was unexpectedly perturbed; the latency of the earliest response to the perturbation in the muscle surface electromyogram was analyzed. The perturbations were delivered either at the onset of each segment or about 0.1 s later. We found that in both segments, the response latency to the late perturbation was shorter than the latency to the early perturbation, which suggests that the proprioceptive feedback control is suppressed in the beginning of each segment. Next, we determined the latency of the response to unexpected perturbations in 30° movements to a single target. We found that the response latency was not significantly different in the movement to a single target and in each segment in the sequence. This result suggests that the initial suppression of the proprioceptive feedback control in movements to single targets is preserved in movements through intermediate targets and supports the idea of modular organization of neural control of movement sequences.

Objective: To examine whether behavioral and electrophysiological measures of motor performance accurately differentiate Parkinson's disease (PD) and essential tremor (ET)., Methods: Twenty-four patients (12 PD; 12 ET) performed isometric force, ballistic movements, and tremor tasks. Receiver operating characteristic (ROC) analyses were conducted on all dependent measures that were significantly different between the two patient groups., Results: Patients with PD were more impaired on measures of movement deceleration than ET. Patients with ET were more impaired on measures of force variability than PD. ROC analyses revealed that sensitivity and specificity were excellent when combining measures during the isometric force task (torque rise time and force variability; 92% sensitivity and 92% specificity; AUC = 0.97). When combining measures across the force and movement tasks, the ROC analysis revealed improved sensitivity and specificity (force variability and peak deceleration; 92% sensitivity and 100% specificity; AUC = 0.99)., Conclusions: Combining measures of force variability and movement deceleration accurately differentiate patients with PD from those with ET with high sensitivity and specificity., Significance: If validated in a larger sample, these measures can serve as markers to confirm the diagnosis of PD or ET and thus, enhance decision making for appropriate treatments for patients with these respective diseases., (Copyright © 2011 International Federation of Clinical Neurophysiology. Published by Elsevier Ireland Ltd. All rights reserved.)

Reaching movements to a target usually have stereotypical kinematics. Although this suggests that the desired kinematics of a movement might be planned, does it also mean that deviations from the planned kinematics are corrected by proprioceptive feedback control? To answer this question, we designed a task in which the subjects made center-forward movements to a target while holding the handle of a robot. Subjects were instructed to make movements at a peak velocity of 1 m/s. No further instructions were given with respect to the movement trajectory or the velocity time profile. In randomly chosen trials the robot imposed servo-controlled deviations from the previously computed unperturbed velocity and position time profiles. The duration of the velocity deviations and the magnitude of accumulated position deviations were manipulated. The subjects were instructed to either "Attempt to correct" or "Do not correct" the movement. The responses to the imposed deviations in the surface electromyograms in the elbow and shoulder agonist muscles consisted of an initial burst followed by a sharp decrease in the "Do not correct" condition or by sustained activity in the "Attempt to correct" condition. The timing and magnitude of the initial response burst reflected those of the velocity deviations and were not affected by the instruction. The timing and magnitude of the late response activity reflected position feedback control and were strongly affected by the instruction. We suggest that proprioceptive feedback control is suppressed in the beginning of the movement, then velocity feedback control is activated in the middle of the movement to control a desired velocity, whereas position feedback control is facilitated late in the movement to acquire the final position.

This study examined the long-term effects of chronic subthalamic nucleus (STN) deep brain stimulation (DBS) using both clinical evaluation and laboratory motor control measures. Over a 5-year time period, changes in the motor section of the Unified Parkinson's Disease Rating Scale (UPDRS) and movement speed and strength at the ankle joint were evaluated on and off STN DBS in eight patients with Parkinson's disease (PD). Four patients were also studied at the elbow joint. Patients with PD originally received unilateral STN DBS between years 2001 and 2003. They were re-evaluated after 5 years of long-term STN DBS between years 2006-2008. At baseline (year 0) and after 5 years, patients with PD were tested off treatment and on STN DBS. In each testing condition, patients performed ballistic, single degree of freedom ankle dorsiflexion and ankle plantarflexion movements and peak velocity was calculated. Patients also performed maximal voluntary contractions at the ankle joint in both directions, and peak torque was calculated. Results showed increased motor UPDRS scores from year 0 to year 5, but STN DBS was efficacious in reducing them. In contrast to the increase in motor UPDRS scores, motor control results showed a marked improvement in peak velocity and peak torque over the 5-year time period in the off treatment condition, and STN DBS was efficacious by improving both peak velocity and peak torque. The current findings suggest that 5 years of chronic STN DBS can have beneficial effects on the motor system over the long term in discrete motor tasks in which maximal effort and maximal neural output is required.

It has previously been found that in fast point-to-point arm movements, proprioceptive feedback is centrally suppressed at the beginning of movement and is facilitated at a time that is correlated with temporal parameters of the planned movement. Here, we show that this correlation holds when subjects are explicitly instructed to move at less than maximal speed. We studied elbow flexion movements made at maximal speed and at 70% of maximal speed over a short distance against a light inertial load and over a long distance against a heavy inertial load. A small number of trials were unexpectedly perturbed by using a servo-controlled motor to decrease the movement velocity. The servo control was turned on early in the movement. The main novel finding is that responses in the surface EMG in the elbow muscles to the perturbation occurred later in the slow-speed conditions than fast-speed conditions. When viewed across all conditions, the onset of the EMG responses to the perturbation increased with the time to peak acceleration in unperturbed movements. In the inertial loaded movements, the time of peak acceleration coincides with the time of peak inertial torque, and so the observed correlation can be interpreted as reflecting the relation between either the planned movement kinematics or the planned movement dynamics. These results are compatible with a hypothesis that a descending command suppresses the proprioceptive feedback control at the movement onset and facilitates it at a time that depends on the time parameters of the planned movement.

It has previously been found that in point-to-point movements against inertial loads, proprioceptive feedback is centrally suppressed in the beginning of movement and is facilitated at a time that is correlated with the expected time of peak velocity. This suggests that the modulation of proprioceptive feedback is governed by the desired movement kinematics. Here we show that in movements against inertial and viscous loads, the correlation of the time when the feedback is facilitated is strongest with the time when the joint torque is expected to be maximal. This suggests that the modulation of proprioceptive feedback is governed by the desired movement dynamics. We applied unexpected perturbations in point-to-point elbow flexion movements against known light and heavy inertial and viscous loads and determined the time and magnitude of responses in the electromyogram (EMG) of the biceps and triceps muscles. In movements against the inertial and viscous loads, the time of the EMG responses was better predicted by the time of the peak joint torque in the unperturbed movement than by the time of peak velocity or the time of peak acceleration or by measures related to the agonist EMG. Moreover, the EMG response changed from a reciprocal pattern in the inertial load conditions to a co-contraction pattern in the viscous load conditions. Our results suggest that during movements against known stable dynamic loads, proprioceptive feedback is tuned to the expected task dynamics and is facilitated so as to maintain muscle stiffness at a time when the muscles are expected to generate maximal force.

Different investigators have proposed that multi-joint arm movements are planned with respect to either the path of the hand or the forces and torques acting about the moving joints. In this experiment, we examined the kinematic and kinetic response of the motor system when a load was applied to the forearm, which reduced the natural anisotropy of the arm. We asked two questions: (1) when the movement path changes upon the introduction of the novel load, do muscle torques at the shoulder and elbow remain the same as they were before the load was applied? and (2) when the path is restored partially as the novel load is learned, do changes in muscle torque occur preferentially at one or the other joint? Participants performed rapid arm movements to a target with and without the novel load attached to their arm. Changes in hand path and muscle torque profiles were examined immediately after the application of the load and again following 30 practice trials. The introduction of the load increased the curvature of hand paths for each participant and resulted in changes in the magnitude and time course of muscle torque at both joints, although to a greater extent at the shoulder. After practice with the load, hand paths and elbow muscle torques resembled those produced with no load. Muscle torques produced at the shoulder, however, did not return to pre-load patterns. These observations provide support for the idea that movements are initiated by planned muscle torques and that as the movement proceeds muscle torques are regulated in order to produce hand paths that conform approximately to a kinematic plan.

In the context of target-reaching movements that involve the arms, trunk and legs, we have delineated rules for apportioning motions amongst body segments, which would be valid for a range of target locations that require forward bending. We further attempted to determine whether the rules are altered when motion is restricted at the knee, obliging a re-apportioning of segment motions. For each participant moving with unrestricted joints to nine target locations, principal component analysis of the changes in orientation (i.e., excursions) of six chosen segments revealed that their coupling can be described by two effective degrees of freedom (DoFs), whose weighted combinations account for the segmental excursions. Investigating the similarities and differences among individuals, we found that a set of two effective DoFs could account for the segment excursions among the group of participants who flex their knees significantly, and another set of two for the group who do not. Comparing the motions with and without the knee joints braced, we found that for each individual participant a set of two (or in some cases three) effective DoFs derived from the unrestricted segment excursions could account well for the altered segment excursions when the participant reached for the targets with the knees restricted. Our findings imply that the redundancy of kinematic DoFs can be resolved by reliance on a small number of couplings of segmental excursions, and, in light of the robustness of these couplings against mechanical restriction of joint motion, suggest a neural rather than mechanical origin for them.

The authors examined the relationship between movement velocity and distance and the associated muscle activation patterns in 18 individuals with focal hand dystonia (FHD) compared with a control group of 18 individuals with no known neuromuscular condition. Participants performed targeted voluntary wrist and elbow flexion movements as fast as possible across 5 movement distances. Individuals with FHD were slower than controls across all distances, and this difference was accentuated for longer movements. Muscle activation patterns were triphasic in the majority of individuals with FHD, and muscle activation scaled with distance in a similar manner to controls. Cocontraction did not explain movement slowing in individuals with dystonia, but there was a trend toward underactivation of the 1st agonist burst in the dystonic group. The authors concluded that slowness is a consistent feature of voluntary movement in FHD and is present even in the absence of dystonic posturing. Underactivation of the 1st agonist burst appears to be the most likely reason to explain slowing.

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 cerebellum, parietal cortex, and premotor cortex are integral to visuomotor processing. The parameters of visual information that modulate their role in visuomotor control are less clear. From motor psychophysics, the relation between the frequency of visual feedback and force variability has been identified as nonlinear. Thus we hypothesized that visual feedback frequency will differentially modulate the neural activation in the cerebellum, parietal cortex, and premotor cortex related to visuomotor processing. We used functional magnetic resonance imaging at 3 Tesla to examine visually guided grip force control under frequent and infrequent visual feedback conditions. Control conditions with intermittent visual feedback alone and a control force condition without visual feedback were examined. As expected, force variability was reduced in the frequent compared with the infrequent condition. Three novel findings were identified. First, infrequent (0.4 Hz) visual feedback did not result in visuomotor activation in lateral cerebellum (lobule VI/Crus I), whereas frequent (25 Hz) intermittent visual feedback did. This is in contrast to the anterior intermediate cerebellum (lobule V/VI), which was consistently active across all force conditions compared with rest. Second, confirming previous observations, the parietal and premotor cortices were active during grip force with frequent visual feedback. The novel finding was that the parietal and premotor cortex were also active during grip force with infrequent visual feedback. Third, right inferior parietal lobule, dorsal premotor cortex, and ventral premotor cortex had greater activation in the frequent compared with the infrequent grip force condition. These findings demonstrate that the frequency of visual information reduces motor error and differentially modulates the neural activation related to visuomotor processing in the cerebellum, parietal cortex, and premotor cortex.

During movement, the intrinsic muscle force-velocity property decreases the net force for the shortening muscle (agonist) and increases it for the lengthening muscle (antagonist). The authors present a quantitative analysis of the effect of that muscle property on activation and force output of the same muscle acting as agonist and antagonist in fast and medium speed goal-oriented movements. They compared biceps activation and force output when that muscle was the agonist in a series of elbow flexions and when it was the antagonist in a series of elbow extensions. They performed the same analysis for the lateral, long, and medial heads of the triceps muscle. Muscle EMG was about 2 times larger and the angular impulse developed by the modeled contractile torque was up to 3 times larger when the muscle or muscles acted as the agonist than when the same muscle or muscles acted as the antagonist in movements with similar kinematics. The large effect of the muscle force-velocity property strongly suggests that the neural controller must account for intrinsic muscle properties to generate movements with a commonly observed bell-shaped velocity profile.

We studied target reaching tasks involving not only the arms but also the trunk and legs, which necessitated some trunk flexion. Such tasks can be successfully completed using an infinite number of combinations of segment motions due to the inherent kinematic redundancy with the excessive degrees of freedom (DOFs). Sagittal plane motions of six segments (shank, thigh, pelvis, trunk, humerus, and forearm) and dynamic torques of six joints (ankle, knee, hip, lumbar, shoulder, and elbow) were analyzed separately by principal component (PC) analyses to determine if there was a commonality among the shapes of the respective waveforms. Additionally, PC analyses were used to probe for constraining relationships among the 1) relative magnitudes of segment excursions and 2) the peak-to-peak dynamic joint torques. In summary, at the kinematic level, the tasks are simplified by the use of a single common waveform for all segment excursions with 89.9% variance accounted for (VAF), but with less fixed relationships among the relative scaling of the magnitude of segment excursions (62.2% VAF). However, at the kinetic level, the time course of the dynamic joint torques are not well captured by a single waveform (72.7% VAF), but the tasks are simplified by relatively fixed relationships among the scaling of dynamic joint torque magnitudes across task conditions (94.7% VAF). Taken together, these results indicate that, while the effective DOFs in a multi-joint task are reduced differently at the kinematic and kinetic levels, they both contribute to simplifying the neural control of these tasks.

When young, healthy subjects perform rapid point-to-point and reversal movements over a range of distances, the patterns of muscle activation associated with accelerating the limb toward the target are modulated in the same way for both movement tasks. Differences in patterns of muscle activation for these two movement types are not observed until the deceleration phase of the movements. In this study, we first test the hypothesis that healthy, older subjects and subjects with Parkinson's disease will modulate the pattern of muscle activation in the same way during the acceleration phase of point-to-point and reversal elbow movements. Second, we test the hypothesis that healthy, older subjects and subjects with Parkinson's disease exhibit the same relationship in muscle activation patterns between the two movement types that have been observed for the young in the deceleration phase of the movements. Subjects performed point-to-point and reversal movements initiated in the direction of flexion over three distances (36, 54 and 72 degrees) "as fast as possible". Angle, velocity, acceleration and surface EMGs from biceps and triceps were recorded. With respect to the first hypothesis, the EMG, kinetic, and kinematic measures related to the acceleration phase of the movements were modulated in the same way for both movement types in the healthy older subjects. In the Parkinson's disease group, the kinematic and kinetic measures during the acceleration phase of the movements were the same in both movement types; however, the flexor and extensor EMG activation was smaller during reversal movements than during point-to-point movements. With respect to the second hypothesis, in contrast to that found in young subjects, in healthy older subjects, there was no significant difference between the movement types in the flexor EMG activity immediately after the time of peak velocity. This difference between younger and older subjects may be attributed to the fact that older subjects perform both movement types more slowly than do younger subjects. Although subjects with Parkinson's disease also move slowly, the flexor EMG shuts off more abruptly and more completely just after the time of peak velocity during reversal movements than during point-to-point movements. These results show that (1) for healthy subjects, when the task requirements are the same for the two movement types (acceleration phase), muscle activation patterns are modulated in the same way, and (2) both age and disease alter the relationship of muscle activation, kinetics and kinematics between point-to-point and reversal movements.

Research on isometric contractions in subjects with Parkinson's disease (PD) has shown that anti-parkinsonian medication results in a greater increase in extensor strength than flexor strength. This finding is consistent with the hypothesis that there is a greater impairment in neural activation of extensor muscles as compared to flexor muscles in subjects with PD. Such a hypothesis is physiologically feasible given the known differences in the neural control of flexor and extensor muscles. If the above hypothesis is true for both phasic and tonic muscle activation, then differences between performance of rapid single-joint flexion and extension movements should exist in subjects with PD. Twelve subjects with PD, "off" and "on" medication, and 12 age-and sex-matched healthy control subjects performed rapid single-joint movements in flexion and extension over three distances. For neurologically healthy subjects, we did not identify any significant differences in either kinematic or EMG parameters between flexion and extension movements. In contrast, in the PD subjects extension movements were slower and associated with more agonist bursts when compared to flexion movements. The results are consistent with the hypothesis that there is a differential impairment of neural activation of extensor muscles of the arm as compared to flexor muscles in subjects with PD.

When moving an object, the motor system estimates the dynamic properties of the object and then controls the movement using a combination of predictive feedforward control and proprioceptive feedback. In this study, we examined how the feedforward and proprioceptive feedback processes depend on the expected movement task. Subjects made fast elbow flexion movements from an initial position to a target. The experimental protocol included movements made over a short and a long distance against an expected light or heavy inertial load. In each task in a few randomly chosen trials, a motor applied an unexpected viscous load that produced a velocity error, defined as the difference between the expected and unexpected velocities, and electromyographic (EMG) responses. The EMG responses appeared not earlier than 170-250 ms from the agonist EMG onset. Our main finding is that the onset of the EMG responses was correlated with the expected time of peak velocity, which increased for longer distances and larger loads. An analysis of the latency of the EMG responses with respect to the velocity error suggested that the EMG responses were due to segmental reflexes. We conclude that segmental reflex gains are centrally modulated with the time course dependent on the expected movement task. According to this view, the control of fast point-to-point movement is feedforward from the agonist EMG onset until the expected time of peak velocity after which the segmental reflex feedback is briefly facilitated.

It is known that muscle activation patterns are changed when the force requirements of a task are increased (e.g., moving a heavier inertial load) or when the available muscle force is reduced (e.g., by inducing muscle fatigue). It is not known whether this is true when the torque-producing capability of a muscle is altered. Eight neurologically healthy subjects performed flexion and extension maximum voluntary isometric contractions (MVC) at five different joint positions (10 degrees, 40 degrees, 70 degrees, 100 degrees, and 130 degrees, where 0 degrees is full elbow extension). Flexion MVC increased by 138% and extension MVC increased by 74% as the elbow joint position changed from the most extended to the most flexed position tested. The same subjects then made rapid, 30 degrees elbow flexion movements from each of four starting elbow positions (10 degrees, 40 degrees, 70 degrees, and 100 degrees). Muscle activation patterns for movements made from the more extended positions showed an increased first agonist burst duration and increased latency of the antagonist burst. There was no change in the initial rate of rise of the agonist burst across starting joint positions. Movements made from the most extended starting position were significantly slower and had longer acceleration and deceleration times than did movements made from the more flexed starting positions. The changes in muscle activation patterns were consistent with those seen when the force requirements of a task are increased or the available muscle force is reduced. We hypothesize that a fall in the ratio of available to required muscle forces causes the nervous system to change muscle activation patterns, to increase the ratio. Our results are consistent with this hypothesis.

The performance of a standing reaching task that necessitates some forward bending requires: (1) the coordination of multiple joints (i.e., the trunk and limb segments) to reach the target, and (2) the preservation of postural stability. It has been proposed that the neural control of multijoint reaching tasks can be simplified by time scaling of joint motions while keeping joint excursions the same. To determine if time scaling of joint motions was used in this more complex reaching task, we had 20 healthy subjects (10 male and 10 female) reach for two targets located in a parasagittal plane while standing on a force platform. Subjects reached for the targets at a comfortable speed and a fast paced speed. Sagittal plane motions of the right shank, thigh, pelvis, trunk, humerus, and forearm were measured. At the fast paced movement speeds subjects had significantly larger excursions of the thigh, pelvis, humerus, and forearm compared to the comfortable speed. Thus, segment motions are not simply time scaled for standing multijoint reaches. We explored three possible reasons for not obeying time scaling: (1) to reduce scaling of peak kinetic energy, (2) to reduce scaling of peak horizontal ground reaction force, and (3) a convergence of movement strategies at faster speeds. While subjects modified their movement strategy in relationship to movement speed, these changes had no significant effect on the expected scaling of peak kinetic energy, or peak horizontal ground reaction forces. Given the intersubject differences in movement strategies used to perform these reaching tasks at the fast speeds, a convergence of movement strategies was ruled out. We propose that the increase in segment motions with speed may be a consequence of rules underlying motor output, the increases being greater for segments in which viscoelastic resistance to movement is more significant compared to inertial resistance.

This study investigated the role of inertial anisotropy at the hand in causing distortions in movement. Subjects drew circles in the horizontal plane at four locations in the workspace at three instructed paces using elbow and shoulder movements. Specifically, we tested two hypotheses, which we would expect if the anisotropy of inertia were not completely accounted for by the CNS when generating circle-drawing movements: 1) speed will affect the circularity of figures, with faster movements associated with greater elongation into an oval shape, irrespective of workspace location for configurations with a similar angle between the forearm and upper arm. 2) The elongation of the circle at fast speeds will be in the direction of least inertia. The results showed that despite individual differences in the speed dependence of the relative motions at the elbow and the shoulder, the circularity decreased (distortion increased) with increased speed, and workspace location had no effect on circularity. We also found that the elongation of the circles at fast speeds was in a direction close to but significantly different from the direction of least inertia for three workspace locations and was in the direction of least inertia for the fourth location. We suggest that the elongation results from lack of full accounting by the CNS of the anisotropy of viscosity and inertia.

This study examined the effects of unexpected loading on muscle activation during fast goal-oriented movements. We tested the hypothesis that the electromyographic (EMG) response to an unexpected load occurs at a short latency after the difference between the expected and the unexpected movement velocity exceeds a fixed threshold. Subjects performed two movement tasks as follows: 1) 30 degrees fast elbow flexion movement with an inertial load added by a torque motor; and 2) 50 degrees fast elbow flexion movement with no added load. These movement tasks were chosen to have similar timing parameters, such as movement time, time-to-peak velocity, and duration of the first agonist burst, while the magnitudes of the angular displacement, velocity, and acceleration were different. In task 1, in random trials a viscous load was substituted for the inertial load at movement onset. In task 2, the same viscous load was added in random trials. The earliest consistent response to the unexpected load was detected in the agonist (biceps) EMG at the same time, about 200 ms from the EMG onset, in both tasks. However, the velocity errors were different in the two tasks and no velocity error threshold dependency could be found. Therefore we reject the hypothesis that the timing of the EMG response to an unexpected load is related to a velocity error threshold. Instead, we suggest that the timing of the EMG response is primarily determined by descending regulation of segmental reflex gain.

The present study investigated how muscle fatigue influences single degree-of-freedom elbow flexion movements and their associated patterns of phasic muscle activation. Maximal unfatigued voluntary isometric elbow flexor and extensor joint torque was measured at the beginning of the experiment. Subjects then performed elbow flexion movements over two distances as fast as possible, and movements over the longer distance at an intentionally slower speed. The slower speed was close to what would become the maximal speed in the fatigued state. Subjects then performed a fatiguing protocol of 20 sustained isometric flexion contractions of 25 s duration with 5 s rest at 50% maximal unfatigued voluntary force. After a recovery period they repeated the movements. The fatigue protocol was successful in inducing muscle fatigue, the evidence being decreased isometric maximal joint torque of over 20%. Fatigued movements had lower peak muscle torque and speed. Our principal finding was of changes in the timing of the phasic patterns of fatigued muscle activation. There was an increase in the duration of the agonist burst and a delay in the timing of the antagonist muscle as measured by the centroid of the EMG signals. We conclude that these changes serve as partial but incomplete, centrally driven compensation for fatigue induced changes in muscle function. An additional, unexpected finding was how small an effect fatigue had on movement performance when using a recovery time of 10 min that is long enough to allow muscle membrane conduction velocity to return to normal. This raises questions concerning the behavioral significance of classical laboratory studies of human fatigue mechanisms.

Studies of electromyographic (EMG) patterns during movements in Parkinson's disease (PD) have often yielded contradictory results, making it impossible to derive a set of rules to explain how muscles are activated to perform different movement tasks. We sought to clarify the changes in modulation of EMG parameters associated with control of movement distance during fast movements in patients with PD. Specifically, we studied surface EMG activity during rapid elbow flexion movements over a wide range of distances (5-72 degrees) in 14 patients with relatively mild symptoms of PD and 14 control subjects of similar age, sex, height, and weight. The PD group exhibited several changes in EMG modulation including impaired modulation of agonist burst duration; increased number of agonist bursts; reduced scaling of agonist EMG magnitude in the more severely impaired subjects; and increased temporal overlap of the antagonist and agonist signals in the most severely impaired subjects. These findings suggest that progressive motor dysfunction in PD is accompanied by increasing deficits in modulating muscle activation. These results help clarify previous disparate and sometimes contradictory results of EMG patterns in subjects with PD., (Copyright 2001 Movement Disorder Society.)

Learning a motor task is associated with specific changes in movement kinematics. Recently, it has been shown that changes in different kinematic parameters occurred with different time courses for subjects who practiced simple, single-joint elbow movements. For example, movement time was seen to decrease and level off in a shorter time than peak velocity, which increased and plateaued later. What is not known, however, is whether the time course and temporal order of these learning-related changes seen at the elbow are similar for movements learned at other joints and with different instructions. In this study, neurologically normal subjects practiced 50 degrees -flexion movements made at the wrist, with the instruction to be both "fast and accurate" (same instruction used in the earlier elbow study). A different group of subjects practiced wrist movements of the same amplitude, but with instructions to make movements that were "always accurate;" only as movement skill developed could subjects increase their speed (but without ever sacrificing accuracy). We measured time-related parameters (duration of acceleration, duration of deceleration, and total movement duration) and magnitude-related parameters (peak velocity, peak acceleration, and peak deceleration). We found that the time course of changes in kinematic parameters for subjects instructed to be "fast and accurate" was similar to that reported at the elbow. When the instruction was changed to be "always accurate," the time for changes in kinematic parameters to level off was found to be longer. However, regardless of instruction, time-related parameters plateaued before magnitude-related parameters. Thus, our results indicate that motor learning mechanisms may operate in a similar way at different joints.

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.

Learning of a motor task, such as making accurate goal-directed movements, is associated with a number of changes in limb kinematics and in the EMG activity that produces the movement. Some of these changes include increases in movement velocity, improvements in end-point accuracy, and the development of a biphasic/triphasic EMG pattern for fast movements. One question that has remained unanswered is whether the time course of the learning-related changes in movement parameters is similar for all parameters. The present paper focuses on this question and presents evidence that different parameters evolve with a specific temporal order. Neurologically normal subjects were trained to make horizontal, planar movements of the elbow that were both fast and accurate. The performance of the subjects was monitored over the course of 400 movements made during experiments lasting approximately 1.5 h. We measured time-related parameters (duration of acceleration, duration of deceleration, and movement duration) and amplitude-related parameters (peak acceleration, peak deceleration, peak velocity), as well as movement distance. In addition, each subject's reaction time and EMG activity was monitored. We found that reaction time was the parameter that changed the fastest and that reached a steady baseline earliest. Time-related parameters decreased at a somewhat slower rate and plateaued next. Amplitude-related parameters were slowest in reaching steady-state values. In subjects making the fastest movements, a triphasic EMG patterns was observed to develop. Our findings reveal that movement parameters change with different time courses during the process of motor learning. The results are discussed in terms of the neural substrates that may be responsible for the differences in this aspect of motor learning and skill acquisition.

Background and Objective: The scientific rationale for pallidotomy as a treatment for PD is that the lesion will reduce excessive tonic inhibition of the thalamus, thereby allowing movement to proceed more normally. If true, then PD patients who move slowly while on medication should increase movement speed following pallidotomy. To test this we used a simple motor task to determine if pallidotomy leads to an improvement in "on" motor performance when those movements are impaired before surgery., Methods: Nine patients with PD performed elbow flexion movements "as fast as possible" while they were "on" before and 1 month after pallidotomy. Patients with mild PD and healthy control subjects were also tested., Results: The clinical effects of pallidotomy were typical of those found in other studies. "Off" Unified Parkinson's Disease Rating Scale scores improved and dyskinesias were reduced. Although before surgery the patients were far slower while they were "on" than the groups of mild PD patients and healthy control subjects, there was no change in mean peak velocity while they were "on" after pallidotomy. There was no change in other mean "on" motor performance measures such as peak acceleration, peak deceleration, initiation time, and symmetry. There was a decrease in the variability of peak acceleration, symmetry, and initiation time., Conclusion: Despite the clinical efficacy of pallidotomy while patients were "off," bradykinesia of elbow flexion movements while patients were "on" is not affected by pallidotomy. Therefore, we conclude that the bradykinesia observed in this experiment is due to a mechanism other than excessive tonic inhibition of the motor thalamus. Our results are consistent with the idea that pallidotomy reduces the noise from the abnormally functioning basal ganglia.

Five subjects made rapid, discrete elbow flexion movements over different distances, against different inertial loads, as well as under distance and load combinations that kept movement time constant. The results demonstrated that an increase in peak movement velocity was associated with an increase in the temporal symmetry ratio of the movement (acceleration time divided by deceleration time), as well as with an increase in both agonist electromyographic (EMG) burst duration and antagonist EMG latency. Since an increase in peak movement velocity is associated with faster agonist muscle shortening, as well as with faster stretching of the antagonist muscle, we hypothesize that the velocity-related changes in movement symmetry can be viewed as, at least partially, a consequence of muscle viscosity. Viscosity increasingly resists the shortening agonist and assists the lengthening antagonist when movement velocity increases. Therefore, the agonist muscles require more time to produce the required impulse, while the antagonist muscle can brake the movement in a shorter period of time. In order to test the hypothesis that viscosity is responsible for the velocity-associated changes in the symmetry ratio, we performed a second experiment with distance and load combinations identical to those of the first experiment, but with different external viscous loads, which resisted the slower and assisted the faster movements. The results demonstrated that the movements became more symmetrical in the presence of the viscous load. There were also changes in agonist duration and antagonist latency. We conclude that changes in the symmetry associated with changes in movement velocity may be due to the effects of either muscle viscosity or changes in how muscles are activated to account for differences in viscous force.

Studies of rapid, single degree-of-freedom movements have shown different changes in electromyographic patterns for movement tasks that appear very similar (e.g., movements over different ranges of distance). However, it is not clear whether these differences are a result of joint-specific control schemes or whether they are instead due to the limited range of task parameters studied relative to the mechanical constraints of each joint (e.g., short compared with long movements relative to the range of motion of a particular joint). In this study, we measured and compared the kinematic trajectories and electromyograms recorded during various movement tasks at the wrist, elbow, and ankle. Subjects performed movements over a wide range of distances "as fast as possible," "at a comfortable speed," and against two inertial loads (at the elbow only), and they performed movements over a fixed distance at three different speeds at the wrist and ankle. For fast movements we show that, in spite of some joint-specific differences, the basic pattern of electromyographic (EMG) modulation is similar at all three joints; for example, the agonist EMG burst transitions from a fixed duration to an increasing duration with increasing movement distance at all three joints. Moreover, the distance at which this transition occurs in one joint relative to the distance at which this transition occurs in the other two joints is consistent across subjects. The transition occurs at the shortest distance at the ankle and the longest distance at the wrist. In general we suggest that the data are consistent with a single set of control rules applied at all three joints, with the biomechanical constraints at each joint accounting for the differences in the EMG and kinematic patterns observed across joints.

1. Eight subjects performed three series of pointing tasks with the unconstrained arm. Series one and two required subjects to move between two fixed targets as quickly as possible with different weights attached to the wrist. By specifying initial and final positions of the finger tip, the first series was performed by flexion of both shoulder and elbow and the second by shoulder flexion and elbow extension. The third series required flexion at both joints, and subjects were instructed to vary movement speed. We examined how variations in load or intended speed were associated with changes in the amount and timing of the electromyographic (EMG) activity and the net muscle torque production. 2. EMG and torque patterns at the individual joints varied with load and speed according to most of the same rules we have described for single-joint movements. 1) Movements were produced by biphasic torque pulses and biphasic or triphasic EMG bursts at both joints. 2) The accelerating impulse was proportional to the load when the subject moved "as fast and accurately as possible" or to speed if that was intentionally varied. 3) The area of the EMG bursts of agonist muscles varied with the impulse. 4) The rates of rise of the net muscle torques and of the EMG bursts were proportional to intended speed and insensitive to inertial load. 5) The areas of the antagonist muscle EMG bursts were proportional to intended movement speed but showed less dependence on load, which is unlike what is observed during single-joint movements. 3. Comparisons across joints showed that the impulse produced at the shoulder was proportional to that produced at the elbow as both varied together with load and speed. The torques at the two joints varied in close synchrony, achieving maxima and going through zero almost simultaneously. 4. We hypothesize that "coordination" of the elbow and shoulder is by the planning and generation of synchronized, biphasic muscle torque pulses that remain in near linear proportionality to each other throughout most of the movement. This linear synergy produces movements with the commonly observed kinematic properties and that are preserved over changes in speed and load.

The kinematic, kinetic, and electromyographic (EMG) patterns observed during fast, single-joint flexion movement have been widely studied as a paradigm for understanding voluntary movement. Several patterns have been described that depend upon the movement task (e.g., distance, speed, and load). A previous model that interpreted differences in EMG patterns in terms of pulse-height or pulse-width modulation of rectangular pulses of motoneuron pool excitation cannot explain all the EMG patterns reported in the literature. We proposed a more general version of that model, consisting of a set of four equations, which specify the parameters of the excitation pulses for a wide variety of movement tasks. Here we report experiments in which subjects performed fast elbow flexions over a range of distances from 2.8 degrees to 45 degrees. The EMG patterns that we observe are consistent with this more general model. We conclude that this model is sufficient to specify muscle excitation patterns that will launch a movement toward and stop it in the neighborhood of a target. This model operates on the basis of prior knowledge about the task rather than feedback received during the task.

The control of distance and speed during single-joint human elbow flexion is accomplished by different models of activating the motoneuron pools. Distance is controlled by modulating the duration of activation, while speed is controlled by modulating the intensity. The experiments reported on here compare movements of different distances under two sets of instructions: subjects moved either as fast and accurately as possible or in a specified time. The first task showed duration modulation, whereas the second, which required simultaneous control of distance and speed, showed both duration and intensity modulation. These results are interpreted in the context of a model for motor control, predicated on the existence of movement plans that use prior knowledge of the dynamics of the movement task to generate muscle activation patterns that produce joint torques. These plans use a simple algorithm based upon parameters of the task such as distance, load, and speed. From this plan, the kinematic trajectory emerges.

Predictions of two views on single-joint motor control, namely programming of muscle force patterns and equilibrium-point control, were compared with the results of experiments with reproduction of movement distance and final location during fast unidirectional elbow flexions. Two groups of subjects were tested. The first group practiced movements over a fixed distance (36 degrees), starting from seven different initial positions (distance group, DG). The second group practiced movements from the same seven initial positions to a fixed final location (location group, LG). Later, all the subjects were tested at the practiced task with their eyes closed, and then, unexpectedly for the subjects, they were tested at the other, unpracticed task. In both groups, the task to reproduce final position had lower indices of final position variability than the task to reproduce movement distance. Analysis of the linear regression lines between initial position and final position (or movement distance) also demonstrated a better (more accurate) performance during final position reproduction than during distance reproduction. The data are in a good correspondence with the predictions of the equilibrium-point hypothesis, but not with the predictions of the force-pattern control approach.

The previous paper in this series showed that changes both within and between experimental sessions can be understood in the framework of the dual-strategy hypothesis of motor control, with a modification sometimes required for the timing of the antagonist muscle. The present paper extends these findings by determining how practicing movements at one distance generalizes to changes in performance at other distances. Five subjects made elbow flexion movements over five different distances (pretest). They then performed 1400 movements (seven sessions of ten blocks of 20 trials) at only one of those distances. The subjects then repeated the flexion movements over the five different distances (posttest). On the posttest, subjects decreased their average movement time by 20 ms. In addition, their movements became less variable. The electromyographic pattern of the faster movements was characterized by a more rapidly rising electromyogram, for three of the subjects, and an antagonist latency that decreased.

This study investigated changes in myoelectric and mechanical variables for movements made "as fast as possible" as a function of practice in the context of the dual-strategy hypothesis of motor control (Gottlieb et al. 1989b). Five male subjects made 1400 rapid elbow flexion movements in ten blocks of 20 trials over seven experimental sessions. Improved performance was defined as increased peak movement velocity, decreased peak velocity variability, increased acceleration and deceleration, a proportionately greater increase in peak deceleration than peak acceleration, and greater consistency in terminal location. The changes observed over experimental sessions were very similar to (but larger and more consistent than) those seen for the first experimental session, with the partial exception of the timing of the antagonist electromyogram (EMG). In general, the increases in the values of the measured mechanical variables covary with myoelectric measures in the same way as when subjects are asked to intentionally change speed in accordance with the rules of the speed-sensitive strategy (Corcos et al. 1989). However, there are differences between subjects in the extent to which speed changes can be attributable to the agonist muscle, the antagonist muscle, or in the timing between the muscles. In one of the five subjects, the latency of the antagonist EMG decreased over blocks on the 1st day but increased over experimental sessions and was consequently activated proportionately later in the movement. This suggests that extended practice can give at least some subjects flexibility in modifying the motor programs that underlie movement.

1. Normal human subjects made discrete elbow flexions in the horizontal plane under different task conditions of initial or final position, inertial loading, or instruction about speed. We measured joint angle, acceleration, and electromyographic signals (EMGs) from two agonist and two antagonist muscles. 2. For many of the experimental tasks, the latency of the antagonist EMG burst was strongly correlated with parameters of the first agonist EMG burst defined by a single equation, expressed in terms of the agonist's hypothetical excitation pulse. Latency is proportional to the ratio of pulse duration to pulse intensity, making it proportional to movement distance and inertial load and inversely proportional to planned movement speed. However, these rules are not sufficient to define the timing of every possible single joint movement. 3. For movements described by the speed-insensitive strategy, the quantity of both antagonist and agonist muscle activity can be uniformly associated with selected kinetic measures that incorporate muscle force-velocity relations. 4. For movements collectively described by the speed-sensitive strategy, (i.e., that have direct or indirect constraints on speed), no single rule can describe all the combinations of agonist-antagonist coordination that are used to perform these diverse tasks. 5. Estimates of joint viscosity were made by calculating the amount of velocity-dependent torque used to terminate movements on target. These estimates are similar to those that have previously been made of limb viscosity during postural maintenance. They imply that a significant component of muscle activity must be used to overcome these forces. 6. These and previous results are all consistent with a dual-strategy hypothesis for those single-joint movements that are sufficiently fast to require pulse-like muscle activation patterns. The major features of such patterns (pulse intensities, durations, and latencies) are determined by central commands programmed in advance of movement initiation. The selection between speed-insensitive or speed-sensitive rules of motoneuron pool excitation is implicitly specified by the nature of speed constraints of the movement task.

Three subjects practiced fast, accurate 36 degrees elbow flexion movements to a 2.5 degrees target for 14 sessions of 100 trials (total, 1400 trials). Subjects then returned for a 15th experimental session in which they were asked to perform 15 movements under identical conditions to the practice condition. They were then tested under three experimental conditions without visual feedback: (1) identical to the practice conditions, (2) with small shifts in starting position (+/- 3 degrees of the practiced starting position), that were insufficient for subjective discrimination and, therefore, subjects were instructed to repeat the practiced movements; and (3) with a large shift in starting position (range, +/- 15 degrees of the practiced starting position), under the instruction to move to the same target. Experimental conditions 2 and 3 demonstrated that shifts in starting position were partially correlated with shifts in final position. These results are interpreted from the point of view of the equilibrium-point hypothesis of motor control.

1. Normal human subjects made isometric pulse and step contractions about the elbow to visually defined target torques of different amplitudes and at different rates. We measured joint torque and electromyograms (EMG) from two agonist and two antagonist muscles. 2. When the task specification requires that the subject explicitly alter the rate at which torque is increased, the rates of rise of the agonist and antagonist EMG bursts covary with the rate of rise of the torque. For pulses of torque the duration of motoneuron excitation varies with the duration of the task-defined contractile event. 3. When a subject is asked to generate torques of different amplitudes without specifying a time interval, torque amplitude is positively correlated with how long, and therefore how high, the EMG rose. Subjects usually proportionately covary the strength of the agonist and antagonist contractions but are not constrained to do so. Some subjects use a strategy of varying the antagonist inversely with the agonist contraction. 4. We extend the organizing principles for the control of movement about a single joint to the control of isometric torque. These rules state that control of torque about a single joint is exercised by one of two strategies: the speed-sensitive strategy modulates the rate at which contraction rises by varying the intensity of motoneuron-pool excitation. The speed-insensitive strategy varies the duration over which contraction rises but does not change the rate. These two respective patterns of torque emerge from pulse-height and pulse-width modulation of motoneuron-pool excitation. 5. The rules defining speed-sensitive and speed-insensitive strategies for movements are broadened for isometric contractions because of the wider range of torque patterns that we observe under these conditions. We propose a step-excitation component for prolonged isometric step contractions and slowly rising ramp patterns of excitation for contractions that develop over several hundreds of milliseconds. 6. The choice of strategies is based on task-specific torque requirements. The same two strategies that control torque to produce movement apply to the control of isometric torque. Unlike movements, however, isometric tasks are more often controlled by a blending of the two patterns. Possible reasons for this are discussed.

1. Human subjects made discrete elbow flexions in a horizontal plane over different distances, from a stationary initial position to a visually defined stationary target 9 degrees wide. We measured joint angle, acceleration, and electromyograms (EMGs) from two agonist and two antagonist muscles. 2. Subjects made movements over four different distances following one of four different instructions. The first instructed the subject simply to choose a comfortable speed. The other three explicitly emphasized either speed, accuracy, or maintenance of the "same" speed over different distances. These instructions produced a wide range of movement velocities. 3. The initial rises of the acceleration (and therefore of the inertial torque), as well as the initial slope of the agonist EMG, were all invariant over changes in the target distance for any single instruction but were all sensitive to the given instruction. 4. Our results demonstrate that the speed-insensitive strategy is a standard or default pattern for performing movements that may be carried out for different instructions over a wide range of speeds. A uniform intensity of excitation pulse is not a byproduct of moving at maximal speed. Submaximal intensities are associated with submaximal speeds and are a selected feature of the pattern of movement control.

Spastic patients were instructed to make accurate, rapid ankle dorsiflexion and plantarflexion movements over different distances to a target. Ankle position and surface electromyograms (EMGs) from tibialis anterior (TA) and soleus (SOL) muscles were recorded. In 3 of 8 spastic patients tested, dorsiflexion evoked velocity-dependent activation of the antagonist (SOL) muscle which impeded the movement to the extent that the limb unintentionally reversed movement direction. We propose that this activation is reflex in origin since it is tightly synchronized, has a large peak amplitude, occurs about 50 ms after the initiation of the movement, and is velocity dependent. One of the 3 patients who had reflex-induced antagonist activation in dorsiflexion also demonstrated sustained clonus during plantarflexion. This usually occurred only if the target had been overshot so that the return of the limb stretched the soleus muscle and triggered clonus. We conclude that in some patients, hyperactive stretch reflexes cause movement deficits.

1. Normal human subjects made discrete elbow flexions and extensions in the horizontal plane from a stationary initial position to visually defined targets at different distances with a constant inertial load or made flexions to a visually defined target with different inertial loads. We measured joint angle, acceleration, and electromyograms (EMGs) from two agonist and two antagonist muscles. 2. Subjects were instructed to move their limbs accurately but quickly to the targets. Movements of greater distances or lesser loads were performed at higher velocities. 3. Peak inertial torque, acceleration and velocity, movement time, and integrated, rectified EMG were all highly correlated with the task variables, distance and inertial load. We show that peak inertial torque can be used as a linking variable that is almost sufficient to explain all correlations between the tasks, the EMG, and movement kinematics. 4. The rate at which subjects initially developed torque to accelerate their movements was invariant over changes in the value of either task variable. The rising phase of the agonist EMG was also independent of the distance or load moved. 5. Two components were distinguished in the antagonist EMG. The first had a relatively constant latency and amplitude. It terminated on the onset of the second and larger component at a latency that was delayed as both distance and load increased. 6. The integrated, rectified antagonist EMG was proportional to inertial load and peak decelerating torque for changes in inertial load. When target distance varied, proportionality between peak decelerating torque and antagonist EMG could be found if correction was made for the effects of muscle length on the torque-EMG relationship. 7. We propose organizing principles for the control of single-joint human movements in which tasks are performed by one of two strategies. These are called speed-insensitive and speed-sensitive strategies. 8. A model is described in which movements made under a speed-insensitive strategy are executed by controlling the duration and the relative timing of amplitude invariant patterns of activation to the spinal motoneuron pools.

1. Normal human subjects made discrete flexions of the elbow over a fixed distance in the horizontal plane from a stationary initial position to a visually defined target. We measured joint angle, acceleration, and electromyograms (EMGs) from two agonist and two antagonist muscles. 2. Changes in movement speed were elicited either by explicit instruction to the subject or by adjusting the target width. Instructions always required accurately stopping in the target zone. 3. Peak inertial torques and accelerations, movement times, and integrated EMGs were all highly correlated with speed. We show that inertial torque can be used as a linking variable that is almost sufficient to explain all correlations between the task, the EMG, and movement kinematics. 4. When subjects perform tasks that require control of movement speed, they adjust the rate at which torque is developed by the muscles. This rate is modulated by the way in which the muscles are activated. The rate at which joint torque develops is correlated with the rate at which the agonist EMG rises as well as with integrated EMG. 5. The antagonist EMG shows two components. The latency of the first is 30-50 ms and independent of movement dynamics. The latency of the second component is proportional to movement time. The rate of rise and area of both components scale with torque. 6. We propose organizing principles for the control of single-joint movements in which tasks are performed by one of two strategies. These are called speed-insensitive and speed-sensitive strategies. 7. A model is proposed in which movements made under a speed-sensitive strategy are executed by controlling the intensity of an excitation pulse delivered to the motoneuron pool. The effect is to regulate the rate at which joint torque, and consequently acceleration, increases. 8. Movements of variable distance, speed, accuracy, and load are shown to be controlled by one of two consistent sets of rules for muscle activation. These rules apply to the control of both the agonist and antagonist muscles. Rules of activation lead to distinguishable patterns of EMG and torque development. All observable changes in movement kinematics are explained as deterministic consequences of these effects.

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.

Three subjects practiced accurate, fast elbow flexions of 54 degrees to a 3 degrees wide target. Movements of 36 degrees, 54 degrees and 72 degrees were then tested. Comparison over the three distances showed that the normally monotonic relationship between movement distance and movement time is alterable by specific training. Subjects learn to go faster over the practiced distance by refining their neural commands to the muscles. The benefits of practice only partially transfer to other distances. We conclude that many of the relationships seen among movement variables in simple tasks are plastic in nature and affected by prior experience.