Biomedical subjects

A G Feldman

Publications and source records attributed to A G Feldman.

At least 19 recordsLinked to original sources

PubMed2003

Interjoint coordination dynamics during reaching in stroke.

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

Adult↗

PubMed2001

The timing of control signals underlying fast point-to-point arm movements.

It is known that proprioceptive feedback induces muscle activation when the facilitation of appropriate motoneurons exceeds their threshold. In the suprathreshold range, the muscle-reflex system produces torques depending on the position and velocity of the joint segment(s) that the muscle spans. The static component of the torque-position relationship is referred to as the invariant characteristic (IC). According to the equilibrium-point (EP) hypothesis, control systems produce movements by changing the activation thresholds and thus shifting the IC of the appropriate muscles in joint space. This control process upsets the balance between muscle and external torques at the initial limb configuration and, to regain the balance, the limb is forced to establish a new configuration or, if the movement is prevented, a new level of static torques. Taken together, the joint angles and the muscle torques generated at an equilibrium configuration define a single variable called the EP. Thus by shifting the IC, control systems reset the EP. Muscle activation and movement emerge following the EP resetting because of the natural physical tendency of the system to reach equilibrium. Empirical and simulation studies support the notion that the control IC shifts and the resulting EP shifts underlying fast point-to-point arm movements are gradual rather than step-like. However, controversies exist about the duration of these shifts. Some studies suggest that the IC shifts cease with the movement offset. Other studies propose that the IC shifts end early in comparison to the movement duration (approximately, at peak velocity). The purpose of this study was to evaluate the duration of the IC shifts underlying fast point-to-point arm movements. Subjects made fast (hand peak velocity about 1.3 m/s) planar arm movements toward different targets while grasping a handle. Hand forces applied to the handle and shoulder/elbow torques were, respectively, measured from a force sensor placed on the handle, or computed with equations of motion. In some trials, an electromagnetic brake prevented movements. In such movements, the hand force and joint torques reached a steady state after a time that was much smaller than the movement duration in unobstructed movements and was approximately equal to the time to peak velocity (mean difference < 80 ms). In an additional experiment, subjects were instructed to rapidly initiate corrections of the pushing force in response to movement arrest. They were able to initiate such corrections only when the joint torques and the pushing force had practically reached a steady state. The latency of correction onset was, however, smaller than the duration of unobstructed movements. We concluded that during the time at which the steady state torques were reached, the control pattern of IC shifts remained the same despite the movement block. Thereby the duration of these shifts did not exceed the time of reaching the steady state torques. Our findings are consistent with the hypothesis that, in unobstructed movements, the IC shifts and resulting shifts in the EP end approximately at peak velocity. In other words, during the latter part of the movement, the control signals responsible for the equilibrium shift remained constant, and the movement was driven by the arm inertial, viscous and elastic forces produced by the muscle-reflex system. Fast movements may thus be completed without continuous control guidance. As a consequence, central corrections and sequential commands may be issued rapidly, without waiting for the end of kinematic responses to each command, which may be important for many motor behaviours including typing, piano playing and speech. Our study also illustrates that the timing of the control signals may be substantially different from that of the resulting motor output and that the same control pattern may produce different motor outputs depending on external conditions.

Adult↗

PubMed2001

Hand trajectory invariance in reaching movements involving the trunk.

Movements of different body segments may be combined in different ways to achieve the same motor goal. How this is accomplished by the nervous system was investigated by having subjects make fast pointing movements with the arm in combination with a forward bending of the trunk that was unexpectedly blocked in some trials. Subjects moved their hand above the surface of a table without vision from an initial position near the midline of the chest to remembered targets placed within the reach of the arm in either the ipsi- or contralateral workspace. In experiment 1, subjects were instructed to make fast arm movements to the target without corrections whether or not the trunk was arrested. Only minor changes were found in the hand trajectory and velocity profile in response to the trunk arrest, and these changes were seen only late in the movement. In contrast, the patterns of the interjoint coordination substantially changed in response to the trunk arrest, suggesting the presence of compensatory arm-trunk coordination minimizing the deflections from the hand trajectory regardless of whether the trunk is recruited or mechanically blocked. Changes in the arm interjoint coordination in response to the trunk arrest could be detected kinematically at a minimal latency of 50 ms. This finding suggests a rapid reflex compensatory mechanism driven by vestibular and/or proprioceptive afferent signals. In experiment 2, subjects were required, as soon as they perceived the trunk arrest, to change the hand motion to the same direction as that of the trunk. Under this instruction, subjects were able to initiate corrections only after the hand approached or reached the final position. Thus, centrally mediated compensatory corrections triggered in response to the trunk arrest were likely to occur too late to maintain the observed invariant hand trajectory in experiment 1. In experiment 3, subjects produced similar pointing movements, but to a target that moved together with the trunk. In these body-oriented pointing movements, the hand trajectories from trials in which the trunk was moving or arrested were substantially different. The same trajectories represented in a relative frame of reference moving with the trunk were virtually identical. We conclude that hand trajectory invariance can be produced in an external spatial (experiment 1) or an internal trunk-centered (experiment 3) frame of reference. The invariance in the external frame of reference is accomplished by active compensatory changes in the arm joint angles nullifying the influence of the trunk motion on the hand trajectory. We suggest that to make a transition to the internal frame of reference, control systems suppress this compensation. One of the hypotheses opened to further experimental testing is that the integration of additional (trunk) degrees of freedom into movement is based on afferent (proprioceptive, vestibular) signals stemming from the trunk motion and transmitted to the arm muscles.

Abdomen↗

PubMed2000

Multi-muscle control of head movements in monkeys: the referent configuration hypothesis.

It is suggested that the nervous system may specify a referent configuration (R) of the body determined by the set of the threshold joint angles at which all skeletal muscles may be silent. At the same time, electromyographic (EMG) activity and forces are generated to resist deflections of the body from this configuration. The R configuration may thus be considered an internal geometric image with which the actual body configuration (Q) is compared. Thereby the difference between the R and Q is a major factor determining the recruitment and gradation of the activity of each skeletal muscle. Control systems may produce movements by changing the R configuration according to task demands. The referent hypothesis predicts that when the R and Q configurations match each other, a global minimum in the EMG activity of all muscles involved should occur, an event most likely observed in movements with reversal in direction. To test the validity of the R hypothesis for head movements, three-dimensional kinematics and EMG activity of 14 functionally diverse neck muscles were analysed in monkeys during head rotations to and from fruit targets placed beyond the oculomotor range. Despite the functional and anatomical diversity of the neck muscles, the activity of all muscles was minimised at a reversal point of the movement trajectory, as predicted by the R hypothesis. This study thus illustrates the notion that a change in the internal geometric image of a biomechanical system may underlie movement production.

Animals↗

PubMed2000

The timing of arm-trunk coordination is deficient and vision-dependent in Parkinson's patients during reaching movements.

The role of the basal ganglia in the coordination of different body segments and utilization of motor synergies was investigated by analyzing reaching movements to remembered three-dimensional (3D) targets in patients with Parkinson's disease (PD). Arm movements were produced alone or in combination with a forward bending of the trunk, with or without visual feedback. Movements in PD patients were more temporally segmented, as evidenced by irregular changes in tangential velocity profiles. In addition, the relative timing in the onsets and offsets of fingertip and trunk motions were substantially different in PD patients than in control subjects. While the control subjects synchronized both onsets and offsets, the PD patients had large mean intervals between the onsets and offsets of the fingertip and trunk motions. Moreover, PD patients showed substantially larger trial-to-trial variability in these intervals. The degree of synchronization in PD patients gradually increased during the movement under the influence of visual feedback. The mean and variability of the intersegmental intervals decreased as the fingertip approached the target. This improvement in timing occurred even though the separate variability in the timing of arm and trunk motions was not reduced by vision. In combined movements, even without vision, the PD patients were able to achieve normal accuracy, suggesting they were able to use the same movement synergies as normals to control the multiple degrees of freedom involved in the movements and to compensate for the added trunk movement. However, they were unable to recruit these synergies in the stereotyped manner characteristic of healthy subjects. These results suggest that the basal ganglia are involved in the temporal coordination of movement of different body segments and that related timing abnormalities may be partly compensated by vision. Abnormal intersegmental timing may be a highly sensitive indicator of a deficient ability to assemble complex movements in patients with basal-ganglia dysfunction. This abnormality may be apparent even when the overall movement goal of reaching a target is preserved and normal movement synergies appear to be largely intact.

Aged↗

PubMed2000

Superposition of independent units of coordination during pointing movements involving the trunk with and without visual feedback.

Previous studies addressing the problem of the control of multiple degrees of freedom have examined the influence of trunk movement on pointing movements within the arm's reach. Such movements may be controlled by two functionally independent units of coordination (synergies): one involving only arm joints and producing the hand trajectory to the target (the transport synergy), and the other coordinating trunk and arm movements leaving the hand trajectory unchanged (the compensatory synergy). The question of whether or not this functional subdivision depends on visual feedback was addressed in the present study. We also tested whether or not the motor effects of different synergies are summated as independent components, a control strategy called "superposition." Finally, we investigated whether or not the relationship between different degrees of freedom within each synergy could be considered linear resulting in proportional changes in different joint angles. Seated subjects produced fast, uncorrected arm movements to an ipsi- or a contralateral target in the direction of +/-45 degrees to the sagittal midline of the trunk. Targets could be reached using the arm alone (control trials) or by combining the arm motion with a forward or backward trunk motion produced by hip flexion or extension (test trials), with and without visual feedback. The shape of the hand trajectory, its direction and tangential velocity, movement precision, joint angles and the sequence of the trunk and hand recruitment and de-recruitment were measured. In both visual conditions, the direction of the hand trajectory observed in control trials was generally preserved in test trials. In terms of sequencing, even in the absence of vision, the trunk movement was initiated before the onset of and outlasted the hand shift, indicating that the potential influence of the trunk on the hand movement was compensated by rotations in the elbow and shoulder joint. The analysis of other variables also implied that the effects of trunk recruitment on the hand trajectory were minor compared to those which could be observed if these effects were not compensated by appropriate changes in the arm joint angles. It was concluded that an arm-trunk compensatory synergy is present in pointing movements regardless of visual feedback. Principal component analysis showed that the relationship between elbow, shoulder and hip joint angles in individual arm and combined arm-trunk movements cannot be considered linear, implying that this relationship is adjusted according to the changing arm geometry. The changes in each arm joint angle (elbow, shoulder) elicited by a forward trunk bending in one block of trials were compared with those elicited by a backward bending in another block, whereas the hand moved to the same target in both blocks. These changes were opposite but of similar magnitude. As a result, for each moment of movement, the mean joint angle obtained by averaging across two directions of trunk motion was practically identical to that in control trials in which the trunk was motionless. It is concluded that the transport and arm-trunk compensatory synergies are combined as independent units, according to the principle of superposition. This principle may simplify the control of the coordination of a redundant number of degrees of freedom.

Adult↗

PubMed1999

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

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

Adult↗

PubMed1999

[Simulation of lateral bending tests using a musculoskeletal model of the trunk].

INTRODUCTION: The lateral bending test is used for the preoperative evaluation of scoliotic patients in order to determine the type of spinal curvatures as well as to assess spine flexibility and possible corrections. However, very few biomechanical studies have been dedicated to the analysis of lateral bending. In this article, a biomechanical model of the human trunk has been used in order to evaluate the possibility of simulating lateral bending tests. METHODS: This model includes elements representing the osseo-ligamentous structures of the spine, rib cage and pelvis, as well as 160 muscle fascicles represented by bilinear cable elements. For 4 scoliotic patients (right thoracic and left lumbar curvatures), 3D upright standing and bending reconstructions were generated from calibrated x-rays and used to calculate the displacements of the vertebrae T1 and L5. These displacements were applied to the model in standing position in order to simulate lateral bending. The resulting geometry of the deformed model was compared to the reconstructed geometry in lateral bending for the other vertebral levels (T2 to L4). RESULTS: The model allows the reproduction of the thoracic Cobb angle modifications with an accuracy superior to 2 degrees, as well as the vertebral rotations in the frontal plane (agreement greater than 85%). The positions of the vertebral body centroids following the simulations showed an agreement of over 77% with reconstructed positions. The direction of the axial angulation for the thoracic and lumbar apical vertebrae is correctly reproduced by the model. The axial rotation for these vertebrae does not result in a common pattern for the 4 patients, which is consistent with the diversity of published data concerning the direction of this coupling. CONCLUSIONS: This study shows the feasibility of simulating lateral bending tests using a 3D biomechanical model integrating muscles. The effect of muscle forces on trunk stiffness and intersegmental mobility can also be assessed using this approach. Future developments should enable the evaluation of the biomechanical properties of scoliotic deformities, thus providing a useful tool for preoperative surgical planning.

Biomechanical Phenomena↗

PubMed1998

Compensatory arm-trunk coordination in pointing movements is preserved in the absence of visual feedback.

This study examined the influence of trunk recruitment on the kinematic characteristics of pointing movements. The distribution of final positions of the hand, the extent and direction of the hand trajectory was basically preserved when trunk movement was combined with arm pointing. These effects were observed during pointing not only with but also without vision. The results imply that two functionally independent units of coordination are used in pointing regardless of visual feedback-one producing arm movement to the target (the reaching synergy) and the other coordinating trunk and arm movements diminishing the influence of the trunk on the arm endpoint trajectory (the compensatory synergy).

Adaptation, Physiological↗

PubMed1998

1998 ISEK Congress Keynote Lecture: Multi-muscle control in human movements. International Society of Electrophysiology and Kinesiology.

It has been suggested that the coordination of the activity of multiple muscles results from the comparison of the actual configuration of the body with a referent configuration specified by the nervous system so that the recruitment and gradation of the activity of each skeletal muscle depend on the difference between these two configurations. Active movements may be produced by the modification of the referent configuration. The hypothesis predicts the existence of a global minimum in electromyographic (EMG) activity of multiple muscles during movements involving reversals in direction. This prediction was tested in five subjects by analysing movements resembling the act of reaching for an object placed beyond one's reach from a sitting position. In such movements, initially sitting subjects raise their body to a semi-standing position and then return to sitting. Consistent with the hypothesis is the observation of a global minimum in the surface EMG activity of 16 muscles of the arm, trunk and leg at a specific phase of the movement. When the minimum occurred, EMG activity of each muscle did not exceed 2-7% of its maximal activity during the movement. As predicted, global EMG minima occurred at the phase corresponding to the reversal in movement direction, that is, during the transition from raising to lowering of the body. The global EMG minimum may represent the point at which temporal matching occurs between the actual and the referent body configurations. This study implies a specific link between motor behavior and the geometric shape of the body modified by the brain according to the desired action.

Abdominal Muscles↗

PubMed1998

Recent tests of the equilibrium-point hypothesis (lambda model).

The lambda model of the equilibrium-point hypothesis (Feldman & Levin, 1995) is an approach to motor control which, like physics, is based on a logical system coordinating empirical data. The model has gone through an interesting period. On one hand, several nontrivial predictions of the model have been successfully verified in recent studies. In addition, the explanatory and predictive capacity of the model has been enhanced by its extension to multimuscle and multijoint systems. On the other hand, claims have recently appeared suggesting that the model should be abandoned. The present paper focuses on these claims and concludes that they are unfounded. Much of the experimental data that have been used to reject the model are actually consistent with it.

Electromyography↗

PubMed1997

Phasic and tonic stretch reflexes in muscles with few muscle spindles: human jaw-opener muscles.

We investigated phasic and tonic stretch reflexes in human jaw-opener muscles, which have few, if any, muscle spindles. Jaw-unloading reflexes were recorded for both opener and closer muscles. Surface electromyographic (EMG) activity was obtained from left and right digastric and superficial masseter muscles, and jaw orientation and torques were recorded. Unloading of jaw-opener muscles elicited a short-latency decrease in EMG activity (averaging 20 ms) followed by a short-duration silent period in these muscles and sometimes a short burst of activity in their antagonists. Similar behavior in response to unloading was observed for spindle-rich jaw-closer muscles, although the latency of the silent period was statistically shorter than that observed for jaw-opener muscles (averaging 13 ms). Control studies suggest that the jaw-opener reflex was not due to inputs from either cutaneous or periodontal mechanoreceptors. In the unloading response of the jaw openers, the tonic level of EMG activity observed after transition to the new jaw orientation was monotonically related to the residual torque and orientation. This is consistent with the idea that the tonic stretch reflex might mediate the change in muscle activation. In addition, the values of the static net joint torque and jaw orientation after the dynamic phase of unloading were related by a monotonic function resembling the invariant characteristic recorded in human limb joints. The torque-angle characteristics associated with different initial jaw orientations were similar in shape but spatially shifted, consistent with the idea that voluntary changes in jaw orientation might be associated with a change in a single parameter, which might be identified as the threshold of the tonic stretch reflex. It is suggested that functionally significant phasic and tonic stretch reflexes might not be mediated exclusively by muscle spindle afferents. Thus, the hypothesis that central modifications in the threshold of the tonic stretch reflex underlie the control of movement may be applied to the jaw system.

Electromyography↗

PubMed1997

Control processes underlying elbow flexion movements may be independent of kinematic and electromyographic patterns: experimental study and modelling.

Using a non-linear dynamic model based on the lambda version of the equilibrium-point hypothesis, we investigated the shape and duration of the control patterns underlying discrete elbow movements. The model incorporates neural control variables, time-, position- and velocity-dependent intrinsic muscle and reflex properties. Two control variables (R and C) specify a positional frame of reference for activation of flexor and extensor motoneurons. The variable R (reciprocal command) specifies the referent joint angle (R) at which the transition of net flexor to extensor active torque or vice versa can be observed during changes in the actual joint angle elicited by an external force. The variable C (coactivation command) surrounds the transition angle by an angular range in which flexor and extensor muscles may be simultaneously active (if C > 0) or silent (if C < or = 0). An additional, time-dimensional control variable (mu command) influences the dependency of the threshold of the stretch reflex on movement velocity. This control variable is responsible for the reflex damping. Changes in the R command result in shifts in the equilibrium state of the system, a dynamical process leading to electromyographic modifications and movement production. Commands C and mu provide movement stability and effective energy dissipation preventing oscillations at the end of movement. A comparison of empirical and model data was carried out. A monotonic ramp-shaped pattern of the R command can account for the empirical kinematic and electromyographic patterns of the fastest elbow flexion movements made with or without additional inertia, as well as of self-paced movements. The rate of the shifts used in simulation was different for the three types of movements but independent of movement distance (20-80 degrees). This implies that, for a given type of movement, the distance is encoded by the duration of shift in the equilibrium state. The model also reproduces the kinematic and electromyographic patterns of the fastest uncorrected movements opposed in random trials by a high load (80-90% of the maximal) generated by position feedback to a torque motor. The following perturbation effects were simulated: a substantial decrease in the arm displacement (from 60-70 degrees to 5-15 degrees) and movement duration (to about 100 ms) so that these movements ended near the peak velocity of those which were not perturbed; a prolongation of the first agonist electromyographic burst as long as the load was applied; the suppression of the antagonist burst during the dynamic and static phases of movements: the reappearance of the antagonist burst in response to unloading accompanied by a short-latency suppression of agonist activity. These kinematic and electromyographic features of both perturbed and non-perturbed movements were reproduced by using the same control patterns which elicited a monotonic shift in the equilibrium state of the system ending before the peak velocity of non-perturbed movements. Our results suggest that the neural control processes underlying the fastest unopposed changes in the arm position are completed long before the end of the movement and phasic electromyographic activity. Neither the timing nor the amplitude of electromyographic bursts are planned but rather they represent the long-lasting dynamic response of central, reflex and mechanical components of the system to a monotonic, short-duration shift in the system's equilibrium state.

Adult↗

PubMed1997

Central modifications of reflex parameters may underlie the fastest arm movements.

Descending and reflex pathways usually converge on common interneurons and motoneurons. This implies that active movements may result from changes in reflex parameters produced by control signals conveyed by descending systems. Specifically, according to the lambda-model, a fast change in limb position is produced by a rapid change in the threshold of the stretch reflex. Consequently, external perturbations may be ineffective in eliciting additional reflex modifications of electromyographic (EMG) patterns unless the perturbations are relatively strong. In this way, the model accounts for the relatively weak effects of perturbations on the initial agonist EMG burst (Ag1) usually observed in fast movements. On the other hand, the same model permits robust reflex modifications of the timing and shape of the Ag1 in response to strong perturbations even in the fastest movements. To test the model, we verified the suggestion that the onset time of the Ag1, even in the fastest movements, depends on proprioceptive feedback in a manner consistent with a stretch reflex. In control trials, subjects (n = 6) made fast unopposed elbow flexion movements of approximately 60 degrees (peak velocity 500-700 degrees/s) in response to an auditory signal. In random test trials, a brief (50 ms) torque of 8-15 Nm either assisting or opposing the movement was applied 50 ms after this signal. Subjects had no visual feedback and were instructed not to correct arm deflections in case of perturbations. In all subjects, the onset time of the Ag1 depended on the direction of perturbation: it was 25-60 ms less in opposing compared with assisting load conditions. Assisting torques caused, at a short latency of 37 ms, an additional antagonist EMG burst preceding the Ag1. The direction-dependent effects of the perturbation persisted when cutaneous feedback was suppressed. It was concluded that the direction-dependent changes in the onset time and duration of the Ag1 as well as the antagonist activation preceding the Ag1 resulted from stretch reflex activity elicited by the perturbations rather than from a change in the control strategy or cutaneous reflexes. The results support the hypothesis on the hierarchical scheme of sensorimotor integration in which EMG patterns and movement emerge from the modification of the thresholds and other parameters of proprioceptive reflexes by control systems.

Adult↗

PubMed1996

The control of multi-muscle systems: human jaw and hyoid movements.

A model is presented of sagittal plane jaw and hyoid motion based on the lambda model of motor control. The model, which is implemented as a computer simulation, includes central neural control signals, position- and velocity-dependent reflexes, reflex delays, and muscle properties such as the dependence of force on muscle length and velocity. The model has seven muscles (or muscle groups) attached to the jaw and hyoid as well as separate jaw and hyoid bone dynamics. According to the model, movements result from changes in neurophysiological control variables which shift the equilibrium state of the motor system. One such control variable is an independent change in the membrane potential of alpha-motoneurons (MNs); this variable establishes a threshold muscle length (lambda) at which MN recruitment begins. Motor functions may be specified by various combinations of lambda s. One combination of lambda s is associated with the level of coactivation of muscles. Others are associated with motions in specific degrees of freedom. Using the model, we study the mapping between control variables specified at the level of degrees of freedom and control variables corresponding to individual muscles. We demonstrate that commands can be defined involving linear combinations of lambda change which produce essentially independent movements in each of the four kinematic degrees of freedom represented in the model (jaw orientation, jaw position, vertical and horizontal hyoid position). These linear combinations are represented by vectors in lambda space which may be scaled in magnitude. The vector directions are constant over the jaw/hyoid workspace and result in essentially the same motion from any workspace position. The demonstration that it is not necessary to adjust control signals to produce the same movements in different parts of the workspace supports the idea that the nervous system need not take explicit account of musculo-skeletal geometry in planning movements.

Biophysical Phenomena↗

PubMed1996

Moment arms and lengths of human upper limb muscles as functions of joint angles.

Modeling of musculoskeletal structures requires accurate data on anatomical parameters such as muscle lengths (MLs), moment arms (MAs) and those describing the upper limb position. Using a geometrical model of planar arm movements with three degrees of freedom, we present, in an analytical form, the available information on the relationship between MAs and MLs and joint angles for thirteen human upper limb muscles. The degrees of freedom included are shoulder flexion/extension, elbow flexion/extension, and either wrist flexion/extension (the forearm in supination) or radial/ulnar deviation (the forearm in mid-pronation). Previously published MA/angle curves were approximated by polynomials. ML/angle curves were obtained by combining the constant values of MLs (defined by the distance between the origin and insertion points for a specific upper limb position) with a variable part obtained by multiplying the MA (joint radius) and the joint angle. The MAs of the prime wrist movers in radial/ulnar deviation were linear functions of the joint angle (R2 > or = 0.9954), while quadratic polynomials accurately described their MAs during wrist flexion/extensions. The relationship between MAs and the elbow angle was described by 2nd, 3rd or 5th-order polynomials (R2 > or = 0.9904), with a lesser quality of fit for the anconeus (R2 = 0.9349). In the full range of angular displacements, the length of wrist, elbow and shoulder muscles can change by 8.5, 55 and 200%, respectively.

Anthropometry↗

PubMed1996

One-trial adaptation of movement to changes in load.

1. We analyzed the rapid adaptation of elbow movement to unexpected changes in external load conditions at the elbow joint. The experimental approach was based on the lambda model, which defines control variables (CVs) setting the positional frames of reference for recruitment of flexor and extensor motoneurons. CVs may be specified by the nervous system independently of the current values of output variable such as electromyographic (EMG) activity, muscle torques, and kinematics. The CV R specifies the referent joint angle (R) at which the transition of flexor to extensor activity or vice versa can be observed during changes in the actual joint angle, theta, elicited by an external force. The other CV, the coactivation (C) command, instead of a single transition angle, defines an angular range in which flexor and extensor muscles may be simultaneously active (if C > 0) or silent (if C < 0). Changes in the R command result in shifts in the equilibrium state of the system, a dynamic process leading to EMG modifications resulting in movement or isometric force production if movement is obstructed. Fast movements are likely produced by combining the R command with a positive C command, which provides movement stability and effective energy dissipation, diminishing oscillations at the end of movement. 2. According to the model, changes in the load characteristic (e.g., from a 0 to a springlike load) influence the system's equilibrium state, leading to a positional error. This error may be corrected by a secondary movement produced by additional changes in R and C commands. In subsequent trials, the system may reproduce the CVs specified after correction in the previous trial. This behavior is called the recurrent strategy. It allows the system to adapt to the new load condition in the subsequent trials without corrections (1-trial adaptation). Alternatively, the system may reproduce the CVs specified before correction (invariant strategy). If the movement was perturbed only in a single trial, the invariant strategy allows the system to reach the target in the subsequent trials without corrections. 3. To test the assumption on the dominant role of the recurrent strategy in rapid adaptation of movement to new load conditions, we performed experiments in which subjects (n = 6) used a pivoting manipulandum and made fast 60 degrees movements to a target. After a random number of trials (5-10) with no load, we introduced opposing (experiment 1), assisting (experiment 2), or randomly varied opposing or assisting loads (experiment 3) for 5-10 trials before unexpectedly switching loads again (14-18 switches in total). The opposing or assisting torque was created by position feedback to a torque motor and was a linear function of the displacement of the manipulandum form the initial position (springlike load). Subjects were instructed to correct positional errors as soon as possible to reach the target. The EMG activity of two elbow flexors (biceps brachii and brachioradialis) and two elbow extensors (triceps brachii and anconeus), elbow position, velocity, and torque were recorded. Kinematic and EMG patterns were compared with those obtained in similar experiments in which subjects were instructed not to correct errors. 4. In 94% of the trials in which a change in the load occurred, the primary movement was in error and was followed by a corrective secondary movement. In primary movements, both the phasic and tonic levels of EMG activity as well as the kinematics were load dependent, implicating reflex and intramuscular mechanisms in the adaptation of muscle forces counteracting external loads. These mechanisms, however, were not sufficient to eliminate positional errors. 5. An undershoot error occurred in trials with an opposing load after those with no load or in trials with no load after those with an assisting load. After adaptation to a new load condition, a sudden return to the previous load condition resulted in an error of the oppo

Adult↗

PubMed1995

The relationship between control, kinematic and electromyographic variables in fast single-joint movements in humans.

Two versions of the hypothesis that discrete movements are produced by shifts in the system's equilibrium point are considered. The first suggests that shifts are monotonic and end near the peak velocity of movement, and the second presumes that they are nonmonotonic ("N-shaped") and proceed until the end of movement. The first version, in contrast to the second, predicts that movement time may be significantly reduced by opposing loads without changes in the control pattern. The purpose of the present study was to test the two hypotheses about the duration and shape of the shift in the equilibrium point based on their respective predictions concerning the effects of perturbations on kinematic and EMG patterns in fast elbow flexor movements. Subjects performed unopposed flexions of about 55-70 degrees (control trials) and, in random test trials, movements were opposed by spring-like loads generated by a torque motor. Subjects had no visual feedback and were instructed not to correct arm deflections in case of perturbations. After the end of the movement, the load was removed leading to a secondary movement to the same final position as that in control trials (equifinality). When the load was varied, the static arm positions before unloading and associated joint torques (ranging from 0 to 80-90% of maximum voluntary contraction) had a monotonic relationship. Test movements opposed by a high load (80-90% of maximal voluntary contraction) ended near the peak velocity of control movements. Phasic and tonic electromyographic patterns were load-dependent. In movements opposed by high loads, the first agonist burst was significantly prolonged and displayed a high level of tonic activity for as long as the load was maintained. In the same load conditions, the antagonist burst was suppressed during the dynamic and static phases of movement. The findings of suppression of the antagonist burst does not support the hypothesis of an N-shaped control signal. Equally, the substantial reduction in movement time by the introduction of an opposing load cannot be reconciled in this model. Instead, our data indicate that the shifts in the equilibrium point underlying fast flexor movements are of short duration, ending near the peak velocity of unopposed movement. This suggests that kinematic and electromyographic patterns represent a long-lasting oscillatory response of the system to the short-duration monotonic control pattern, external forces and proprioceptive feedback.

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