Relationship between cocontraction, movement kinematics and phasic muscle activity in single-joint arm movement

Patterns of muscle coactivation provide a window into mechanisms of limb stabilization. In the present paper we have examined muscle coactivation in single-joint elbow and single-joint shoulder movements and explored its relationship to movement velocity and amplitude, as well as phasic muscle activation patterns. Movements were produced at several speeds and different amplitudes, and muscle activity and movement kinematics were recorded. Tonic levels of electromyographic (EMG) activity following movement provided a measure of muscle cocontraction. It was found that coactivation following movement increased with maximum joint velocity at each of two amplitudes. Phasic EMG activity in agonist and antagonist muscles showed a similar correlation that was observable even during the first 30 ms of muscle activation. All subjects but one showed statistically significant correlations on a trial-by-trial basis between tonic and phasic activity levels, including the phasic activity measure taken at the initiation of movement. Our findings provide direct evidence that muscle coactivation varies with movement velocity. The data also suggest that cocontraction is linked in a simple manner to phasic muscle activity. The similarity in the patterns of tonic and phasic activation suggests that the nervous system may use a simple strategy to adjust coactivation and presumably limb impedance in association with changes in movement speed. Moreover, since the pattern of tonic activity varies with the first 30 ms of phasic activity, the control of cocontraction may be established prior to movement onset. Electronic Publication     

Antagonist muscle activation preceding rapid flexion movements of the elbow joint in human subjects

Our study was designed to look for interactions between fast movements and pre-existing voluntary tonic motor activity when both motor acts employ the same muscles. Five normal subjects performed a continuous sequence of two motor tasks about their right elbow joint: A tonic isometric extension (slowly increasing or decreasing) against a force transducer, followed immediately after a "go" tone by a fast isotonic flexion. The position of the lower arm was recorded using a search coil system. Signals (force, position, and surface EMGs of triceps and biceps brachii muscles) were A/D converted and sampled at 1 kHz. A premovement silence in the tonically active triceps muscle (extensor) usually preceded the fast flexion movement if the triceps' tonic force was either constant or decreased slowly. If the tonic triceps activity had been increasing before the fast flexion began, this classical picture disappeared, and the premovement silence was replaced by a phasic premovement excitation. Subjects were unaware of this transient EMG and force increase in the unintended direction. Our results demonstrate unconscious reciprocal interactions between commands governing evolving movements (and tuning the motor system accordingly) and those concerned with ongoing motor acts.     

Independent coactivation of shoulder and elbow muscles

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

Responses of mono- and bi-articular muscles to load perturbations of the human arm

Summary We studied the behavior of muscles acting synergistically in elbow flexion in response to load perturbations. The perturbations were applied either proximally or distally to the elbow joint and consisted of single pulses or steps of torque and of pseudorandom sequences of torque pulses. They produced changes in angular position and torque at both the shoulder and elbow joints. The electromyographic (EMG) responses evoked in biceps, brachio-radialis and brachialis muscles were different when elbow and shoulder motion was in the same direction and when the two angular motions were oppositely directed. For example, elbow extension resulted both when a downward force perturbation was applied to the forearm as well as when a posteriorly directed force applied to the upper arm was released. Elbow flexors were activated at a short latency only in the former case and not in the latter. The modulation of EMG activity in elbow flexors evoked by the perturbations was related to the global motion of the limb, including the angular motions at both the shoulder and elbow joints. The time course of the EMG responses in biceps, which acts on both joints, differed from that of brachio-radialis and brachialis muscles, which act only at the elbow. The results are discussed in the context of the possible mechanisms responsible for the muscle responses to the perturbations.     

Arm muscle activation for static forces in three-dimensional space

1. Muscle activity was related to the direction of a static force at the human wrist. For each muscle the force direction of maximal activity and the directional tuning characteristics were determined. 2. Electromyographic (EMG) activity was recorded from nine superficial elbow and/or shoulder muscles while subjects held the right arm stationary in one of six postures. The direction of the force at the wrist was varied in two orthogonal planes. In each experiment a cable was attached to the subject's wrist, and a constant force magnitude was applied in various directions with the use of a pulley system. 3. The relationship between the averaged EMG level and the force direction was described for each muscle, in each posture, and in each plane. The EMG data were fit with a nonlinear, multiple cosine function, which allowed the identification of one, two, or sometimes three separate cosine peaks. 4. Two-cosine functions often provided the best fit to the EMG data. All nine muscles were best fit with a two-cosine function in at least two of the six postures. Four of the muscles had a second peak of activity in more than one-half of the experimental situations. The second peak was often in a direction that was nearly opposite the direction of the first peak and represented a negative contribution to the total force produced at the wrist ("coactivation"). We suggest that multimodal directional tuning results from the convergence of multiple sources of descending signals onto motoneurons. 5. The mechanical actions of nine elbow and/or shoulder muscles were estimated with the use of published data from a cadaver study by Wood and co-workers. Postural changes in the mechanical actions of muscles were substantial. A 45 degrees rotation of the shoulder, for example, might cause a 30-50 degrees change in the direction of force at the wrist that could be produced by the contraction of a given muscle. The magnitude of these postural changes suggests that arm position is an important determinate of EMG patterns. 6. Postural changes in the direction of maximal EMG activity usually paralleled the postural changes in mechanical pulling direction. Postural changes in EMG amplitudes usually covaried with postural changes in mechanical advantage. 7. The posterior deltoid (PD) was an exception to the general rule of covariation of mechanical actions and EMG activities. Instead of reflecting the muscle's mechanical action, the EMG activity of the PD closely resembled the EMG activity of the medial deltoid (MD).(ABSTRACT TRUNCATED AT 400 WORDS)     

Neural control of superficial and deep neck muscles in humans

Human neck muscles have a complex multi-layered architecture. The role and neural control of these neck muscles were examined in nine seated subjects performing three series of isometric neck muscle contractions: 50-N contractions in eight fixed horizontal directions, 25-N contractions, and 50-N contractions, both with a continuously changing horizontal force direction. Activity in the left sternocleidomastoid, trapezius, levator scapulae, splenius capitis, semispinalis capitis, semispinalis cervicis, and multifidus muscles was measured with wire electrodes inserted at the C(4)/C(5) level under ultrasound guidance. We hypothesized that deep and superficial neck muscles would function as postural and focal muscles, respectively, and would thus be controlled by different neural signals. To test these hypotheses, electromyographic (EMG) tuning curves and correlations in the temporal and frequency domains were computed. Three main results emerged from these analyses: EMG tuning curves from all muscles exhibited well-defined preferred directions of activation for the 50-N isometric forces, larger contractions (25 vs. 50 N) yielded more focused EMG tuning curves, and agonist neck muscles from all layers received a common neural drive in the range of 10-15 Hz. The current results demonstrate that all neck muscles can exhibit phasic activity during isometric neck muscle contractions. Similar oscillations in the EMG of neck muscles from different layers further suggest that neck motoneurons were activated by common neurons. The reticular formation appears a likely generator of the common drive to the neck motoneurons due to its widespread projections to different groups of neck motoneurons.     

Basic features of phasic activation for reaching in vertical planes

The purpose of this study was to fully characterize the timing and intensity of the phasic portion of the electromyographic (EMG) waveform for reaching movements in vertical planes. Electromyographic activity was simultaneously recorded from nine superficial elbow and/or shoulder muscles while human subjects made rapid arm movements. Hand paths comprised 20 directions in a sagittal plane and 20 directions in a frontal plane. In order to focus on the more phasic aspects of muscle activation, estimates of postural EMG activity were subtracted from the EMG traces recorded during rapid reaches. These postural estimates were obtained from activity recorded during very slow reaches to the same targets. After subtraction of this postural activity, agonist or antagonist burst patterns were often observed in the phasic EMG traces. For nearly all muscles and all subjects, the relation between phasic EMG intensity and movement direction was a function with multiple peaks. For all muscles, the timing of phasic EMG bursts varied as a function of movement direction: the data from each muscle exhibited a gradual temporal shift of activity over a certain range of directions. This gradual temporal shift has no obvious correspondence to the mechanical requirements of the task and might represent a neuromuscular control strategy in which burst timing contributes to the specification of movement direction.     

Compensation for interaction torques during single- and multijoint limb movement

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

Multijoint muscle regulation mechanisms examined by measured human arm stiffness and EMG signals

Stiffness properties of the musculo-skeletal system can be controlled by regulating muscle activation and neural feedback gain. To understand the regulation of multijoint stiffness, we examined the relationship between human arm joint stiffness and muscle activation during static force control in the horizontal plane by means of surface electromyographic (EMG) studies. Subjects were asked to produce a specified force in a specified direction without cocontraction or they were asked to keep different cocontractions while producing or not producing an external force. The stiffness components of shoulder, elbow, and their cross-term and the EMG of six related muscles were measured during the tasks. Assuming that the EMG reflects the corresponding muscle stiffness, the joint stiffness was predicted from the EMG by using a two-link six-muscle arm model and a constrained least-square-error regression method. Using the parameters estimated in this regression, single-joint stiffness (diagonal terms of the joint-stiffness matrix) was decomposed successfully into biarticular and monoarticular muscle components. Although biarticular muscles act on both shoulder and elbow, they were found to covary strongly with elbow monoarticular muscles. The preferred force directions of biarticular muscles were biased to the directions of elbow monoarticular muscles. Namely, the elbow joint is regulated by the simultaneous activation of monoarticular and biarticular muscles, whereas the shoulder joint is regulated dominantly by monoarticular muscles. These results suggest that biarticular muscles are innervated mainly to control the elbow joint during static force-regulation tasks. In addition, muscle regulation mechanisms for static force control tasks were found to be quite different from those during movements previously reported. The elbow single-joint stiffness was always higher than cross-joint stiffness (off-diagonal terms of the matrix) in static tasks while elbow single-joint stiffness is reported to be sometimes as small as cross-joint stiffness during movement. That is, during movements, the elbow monoarticular muscles were occasionally not activated when biarticular muscles were activated. In static tasks, however, monoarticular muscle components in single-joint stiffness were increased considerably whenever biarticular muscle components in single- and cross-joint stiffness increased. These observations suggest that biarticular muscles are not simply coupled with the innervation of elbow monoarticular muscles but also are regulated independently according to the required task. During static force-regulation tasks, covariation between biarticular and elbow monoarticular muscles may be required to increase stability and/or controllability or to distribute effort among the appropriate muscles.     

Effect of combined variation of force amplitude and rate of force development on the modulation characteristics of muscle activation during rapid isometric aiming force production

Studies of rapid target-directed limb movements have suggested that various control schemes can be defined by the modulation pattern of the muscle activity. The present study was aimed to address the question regarding the extent to which a simultaneous control of force amplitude, and rate of force development influences the modulation characteristics of muscle activation associated with producing rapid isometric aiming forces at the elbow joint. The subjects were instructed to produce rapid isometric force pulses to three different force amplitudes (15, 35, and 55% of their maximal voluntary contractions) under systematically varied force-rate conditions ranging from a fast and accurate force-rate to the fastest force-rate possible. The results showed that larger force amplitudes were achieved by increasing the rate of force development (dF/dt) while the time to peak force remained relatively constant. The magnitude of the electromyographic (EMG) burst systematically increased as a function of force amplitude at all force-rate conditions. The primary finding was that the characteristic of the EMG burst duration associated with different force amplitudes showed a significant difference among force-rate conditions. Under a fast and accurate force-rate condition, the duration of the agonist burst increased linearly with force amplitude. A gradual transition into a fixed duration of the agonist burst then was observed over the remaining three force-rate requirements. With increasingly faster force-rates, there were no changes in the agonist burst duration over three force amplitudes. These results indicate that the combined variations in force amplitude and force-rate examined relative to the most rapid force-rate influence the control patterns for the muscle activation during the fast isometric force production. Changes in the EMG modulation patterns observed are likely due to the constraints imposed by muscle contractile properties.     

Robustness of muscle synergies underlying three-dimensional force generation at the hand in healthy humans

Previous studies using advanced matrix factorization techniques have shown that the coordination of human voluntary limb movements may be accomplished using combinations of a small number of intermuscular coordination patterns, or muscle synergies. However, the potential use of muscle synergies for isometric force generation has been evaluated mostly using correlational methods. The results of such studies suggest that fixed relationships between the activations of pairs of muscles are relatively rare. There is also emerging evidence that the nervous system uses independent strategies to control movement and force generation, which suggests that one cannot conclude a priori that isometric force generation is accomplished by combining muscle synergies, as shown in movement control. In this study, we used non-negative matrix factorization to evaluate the ability of a few muscle synergies to reconstruct the activation patterns of human arm muscles underlying the generation of three-dimensional (3-D) isometric forces at the hand. Surface electromyographic (EMG) data were recorded from eight key elbow and shoulder muscles during 3-D force target-matching protocols performed across a range of load levels and hand positions. Four synergies were sufficient to explain, on average, 95% of the variance in EMG datasets. Furthermore, we found that muscle synergy composition was conserved across biomechanical task conditions, experimental protocols, and subjects. Our findings are consistent with the view that the nervous system can generate isometric forces by assembling a combination of a small number of muscle synergies, differentially weighted according to task constraints.     

Two components of muscle activation: scaling with the speed of arm movement.

1. The temporal waveform of muscle activity was related to the speed of arm movement. Speed was expressed in terms of the duration of a fixed amplitude movement or the "movement time." 2. Human subjects moved their arms to targets in three-dimensional space. The right arm started at a standard initial position and moved directly to the target in a single stroke. The targets were placed in various directions in a vertical plane. The arm movements consisted of shoulder and elbow rotations. 3. Subjects were required to vary the speed of their movements. In most of the experiments, trials with different movement times were randomly ordered. One of the experiments also included randomly interspersed static trials, in which the subject held the arm still at the initial posture, the final posture, or halfway between the two extremes. 4. Electromyographic (EMG) activity was recorded from several superficial elbow and/or shoulder muscles. The time base of rectified EMG records was normalized for movement time such that records from movements with various speeds were compressed to align the ending times of the movements. 5. A principal component (PC) analysis revealed that the compressed EMG waveforms could be described by a summation of PC1 and PC2 waveforms; each individual EMG waveform was approximated by a weighted sum of these two components. 6. The PC1 weighting coefficients scaled down in a monotonic relationship with movement time such that the fastest movement corresponded to a large positive weighting coefficient and the slowest movement corresponded to a small positive weighting coefficient. The PC2 weighting coefficients exhibited a similar monotonic scaling, but the values ranged from positive to negative. Further analysis demonstrated that these two components can be mathematically transformed into a tonic waveform with a constant mathematically transformed into a tonic waveform with a constant weighting coefficient and a phasic waveform with positive weighting coefficients that scale down with movement time. 7. The amplitude scaling of EMG records cannot be described by a single component, but instead requires a summation of two separate components. The tonic component may correspond to the force element needed to counteract gravity, because the magnitude of this element does not scale with movement speed. The phasic component may correspond to the force element that scales quadratically to produce a linear increase in velocity.     

Hysteresis in corticospinal excitability during gradual muscle contraction and relaxation in humans

Many studies have demonstrated that the firing behavior of single motor units varies in a nonlinear manner to the exerted torque during gradual muscle contraction and relaxation. However, it is unclear whether corticospinal excitability has such a hysteresis-like feature. In this study, we examined corticospinal excitability using transcranial magnetic stimulation (TMS) during gradual muscle contraction and relaxation for torque regulation in elbow flexor muscles. Eight healthy male subjects performed two different isometric elbow flexion tasks, namely, sinusoidal and tonic torque exertion tasks. In the sinusoidal task, the subjects sinusoidally increased and decreased the isometric elbow flexion torque (range of 0–15% of maximum voluntary contraction) at three different frequencies (0.33, 0.17, and 0.08 Hz). For each ascending (contraction: CON) and descending (relaxation: REL) period of the exerted torque, a single TMS was applied at 5 phases. In the tonic task, the elbow flexion torque was tonically exerted at 7 levels in a similar range as that in the sinusoidal task. EMG activities were recorded from the agonists, the biceps brachii (BB) and brachioradialis (BRD) muscles, and an antagonist, the triceps brachii (TB) muscle. The results demonstrated that the EMG activities of both the agonists and antagonist were larger in the CON period than the REL period, even when the exerted torque was the same. However, there were no significant differences in EMG activation profiles among the different frequencies of contraction. In BB and BRD, the motor-evoked potential (MEP) elicited by the TMS was also greater in the CON period than in the REL period. This CON-REL difference of MEP amplitudes was still observed when corrections were made for the increased EMG activities; that is, the MEP amplitudes to the identical EMG activities were greater in the CON period than in the REL period, and this phenomenon was more pronounced at higher frequencies. In addition, the degree to which sinusoidal MEPs exceeded tonic MEPs in the CON period and were smaller than tonic MEPs in the REL period became more pronounced at higher frequencies. On the other hand, there were significant correlations between the BB and BRD MEP amplitudes and the rate of change of elbow flexion/extension torque. These results indicate that corticospinal excitability during muscle contraction and relaxation has a neural hysteresis to the muscle activity, i.e., spinal motoneuronal activity, according to the rate of change of the exerted torque, i.e., muscle tension. This suggests that corticospinal excitability modulation depends not only on concurrent spinal motoneuronal activity and muscle tension but also on the time-series pattern of their changes during muscle contraction and relaxation.     

Adaptation of the precentral cortical command to elbow muscle fatigue

The control exerted by individual motor cortical cells on their fatigued target muscles was assessed by analyzing the discharge patterns and electromyographic (EMG) postspike effects of cortical cells in monkeys making repeated forceful, but submaximal, isometric flexions of the elbow to produce fatigue. Two monkeys were trained to perform self-paced isometric contractions (for longer than 2 s) at forces greater than 35% maximal contraction, with three sets of 20 consecutive contractions; the first and last sets were at the same force level. Pairs of EMG electrodes were implanted in the biceps brachii, brachioradialis, and triceps brachii. The cortical cell discharges were modulated with the active and passive movements of the elbow and produced consistent EMG postspike effects during isometric contraction. Muscle fatigue was assessed as a statistically significant (P<0.05) drop in the mean power frequency of the EMG power spectrum in one or both flexors in the last set of contractions. Clear signs of muscular fatigue occurred in 20 different experimental sessions. Before fatigue, cortical cells were classified as phasic-tonic (18), phasicramp (three), or tonic (five). Twenty cells briskly fired to passive elbow extension, and 9 also responded to passive flexion. Only 6 cells showed a decreased discharge to passive extension. A 22–30% increase in the contraction force produced a higher discharge frequency in 13 cells, and a lower frequency in 5 cells. All cells exerted EMG postspike effects in their target muscles: 20 cells facilitated the flexors, and some of these also inhibited (3 cells) or cofacilitated (5 cells) the extensor; the other 6 cells had mixed effects: 5 of them inhibited at least one flexor, and 1 cell only facilitated the extensor. Most cells (24/26) still produced EMG postspike effects in their target muscles during fatigue, and the number of facilitated muscles increased: 21 cells facilitated the flexors, and 12 of them cofacilitated the extensor. Only 3 cells still inhibited the flexors and were tonic cells. The cortical cell firing frequency increased during fatigue in 13 cells and decreased in 8 cells. Increases involved 10 cells excited by passive elbow extension. Fourteen cells showed parallel changes in firing frequency with fatigue and force, and 9 of these cells facilitated both extensors and flexors in fatigue. Increases were found in 8 cells, decreases in 5 cells and no change in 1 cell. As muscle afferents provide substantial information to cortical cells, which in turn establish functional linkages with their target muscles before and during fatigue, the changes in cell firing frequencies during fatigue demonstrate the active participation of the motor cortex in the control of compensation for the peripheral adjustments concomitant with muscle fatigue.     

Discharge properties of primate forearm motor units during isometric muscle activity

Activity of single motor units (MUs) was recorded in forelimb muscles of rhesus macaques while they generated isometric ramp-and-hold torques about the wrist. Multiunit electromyographic (EMG) activity was recorded from 10-12 identified flexor and extensor muscles of the wrist and digits with implanted EMG wire electrodes. Single MUs from these muscles were recorded with a remotely controlled tripolar microelectrode array. The parent muscle of each MU was determined by compiling MU-triggered averages of multiunit EMGs. The MU firing patterns during the isometric task were determined from response histograms aligned with change in torque. At moderate torque levels, MUs (n = 86) exhibited four types of discharge patterns during the ramp-and-hold trajectory: phasic-tonic (23%), tonic (33%), decrementing (39%), and phasic (5%). Phasic-tonic MUs exhibited a phasic burst of activity during the torque ramp which exceeded the firing rate during the static hold period. Both phasic-tonic and tonic MUs exhibited a constant mean firing rate during the hold period; the discharge of decrementing MUs gradually decreased during the static hold. Phasic MUs fired only during the change in force. The relation between MU firing rate and torque was investigated as the monkeys generated responses of different levels of static torque during the hold period. Mean firing rate during the hold was found to be proportional to static torque up to moderate torque levels, where it reached a maximum. In the linear range, the mean rate-torque slope was 3.4 +/- 1.9 imp/s per 10(5) dyn . cm (n = 9).     

EMG responses to load perturbations of the upper limb: effect of dynamic coupling between shoulder and elbow motion

Summary Load perturbations were applied to the arm of human subjects under conditions where both limb segments (upper arm and forearm) were free to move. The perturbations consisted of pulses of torque 50 ms in duration and of pseudo-random sequences of such pulses. They were applied to either the forearm or the upper arm. Under all conditions, the perturbations resulted in angular motion at the shoulder and elbow joints and evoked consistent responses in muscles acting about these joints (biceps, triceps, anterior and posterior deltoid). Activity in biceps and triceps was not related simply to angular motion at the elbow joint. For example, activation of biceps could be evoked during elbow flexion (by applying a torque perturbation at the shoulder) as well as during elbow extension (by applying a torque perturbation at the elbow). The effect of varying degrees of dynamic coupling between upper arm and forearm on EMG responses was investigated by applying torque perturbations to the upper arm over a wide range of elbow angles. When the forearm is extended, such a perturbation induces a greater amount of elbow flexion than when the forearm is in a flexed position. The results of these experiments showed that the larger was the amount of flexion of the forearm induced by the perturbation, the larger was the activation of biceps. The results are incompatible with the notion of a negative feedback of total muscle length as being responsible for the EMG activity following the load perturbations. It is suggested that the EMG responses can best be interpreted functionally in terms of parameters more global than muscle length. Among such global parameters, changes in net torque at a joint resulting from the perturbation gave the best correlation with the pattern of EMG activities observed.     

EMG patterns in antagonist muscles during isometric contraction in man: Relations to response dynamics

Summary We studied the EMG activity of biceps and triceps in human subjects during isometric force adjustments at the elbow. Rapid targeted force pulses exhibited stereotyped trajectories in which peak force was a linear function of the derivatives of force and the time to peak force was largely independent of its amplitude. These responses were associated with an alternating triphasic pattern of EMG bursts in agonist and antagonist muscles similar to that previously described for rapid limb movements. When the instructions demanded rapid force pulses, initial agonist bursts were of constant duration, and their magnitude was strongly related to peak force achieved. The timing of EMG bursts in antagonist pairs was closely coupled to the dynamics of the force trajectory, and the rising phase of the force was determined by both agonist and antagonist bursts. When peak force was kept constant and rise time systematically varied, the presence and magnitude of antagonist and late agonist bursts were dependent on the rate of rise of force, appearing at a threshold value and then increasing in proportion to this parameter. It is proposed that antagonist activity compensates for nonlinearity in muscle properties to enable the linear scaling of targeted forces which characterizes performance in this task.Supported by the Dystonia Medical Research Foundation and NS 19205     

Characteristics of synergic relations during isometric contractions of human elbow muscles

We studied the patterns of EMG activity in elbow muscles in three normal human subjects. The myoelectrical activity of 7-10 muscles that act across the human elbow joint was simultaneously recorded with intramuscular electrodes during isometric joint torques exerted over a range of directions. These directions included flexion, extension, varus (internal humeral rotation), valgus (external humeral rotation), and several intermediate directions. The forces developed at the wrist covered a range of 360 degrees, all orthogonal to the long axis of the forearm. The levels of EMG activity were observed to increase with increasing joint torque in an approximately linear manner. All muscles were active for ranges less than 360 degrees and most were active for less than 180 degrees. The EMG activity was observed to vary in a systematic manner with changes in torque direction and, when examined over the full angular range at a variety of torque levels, is simply scaled with increasing torque magnitude. There were no torque directions or torque magnitudes for which a single muscle was observed to be active alone. In all cases, joint torque appeared to be produced by a combination of muscles. The direction for which the EMG of a muscle reached a maximum value was observed to correspond to the direction of greatest mechanical advantage as predicted by a simple mechanical model of the elbow and relevant muscles. Muscles were relatively inactive during varus torques. This implies that the muscles were not acting to stabilize the joint in this direction and could have been allowing ligaments to carry the load. Plots of EMG activity in one muscle against EMG activity in another demonstrate some instances of pure synergies, but patterns of coactivation for most muscles are more complicated and vary with torque direction. The complexity of these patterns raises the possibility that synergies are determined by the task and may have no independent existence. Activity in two heads of triceps brachii (medial head--a single-joint muscle and long head--a two-joint muscle) covaried closely for a range of torque magnitudes and directions, though shoulder torque and hence the forces experienced by the long head of the triceps undoubtedly varied. The similarity of activation patterns indicates that elbow torque was the principal determining factor. The origins of muscle synergies are discussed. It is suggested that they are best understood on the basis of a model which encodes limb torque in premotor neurons.(ABSTRACT TRUNCATED AT 400 WORDS)     

Trajectory control in targeted force impulses

The functional role of opposing muscles in the production of isometric force trajectories was studied in six adult subjects producing impulses and steps of elbow flexor force, with different rise times and amplitudes. Rapidly rising forces were invariably associated with an alternating pattern of EMG activity in agonist and antagonist muscles: an agonist burst (AG1) initiated the development of force in the desired direction while a reciprocal burst in the antagonist (ANT-R) led to the deceleration of the force trajectory prior to the peak force. The temporal pattern of agonist and antagonist activation was dependent on force rise time. Force trajectories with long rise times (greater than 200 ms) were entirely controlled by the agonist, and EMG activity closely followed the contours of the rising force trajectory. For rise times of about 120 to 200 ms, agonist activation formed a discrete EMG burst, and force continued to rise during the subsequent silent period. For brief force rise times (less than 120 ms), reciprocal activation of the antagonist muscle occurred at about the time of the peak dF/dt. The integrated magnitude of AG1 was dependent on peak force but was independent of force rise time. AG1 duration varied directly with both peak force and force rise time. The integrated value of ANT-R varied as an inverse function of force rise time and was minimally influenced by peak force. ANT-R was present with the same magnitude and timing in both force impulses and steps when rise times were equal; therefore it did not serve to return force to baseline. Rather it served to truncate the rising force when very brief rise times were required, thus compensating for the low-pass filter properties of the agonist muscle. Subjects were able to voluntarily suppress ANT-R in rapidly accelerated force trajectories, indicating that the linkage between the commands controlling agonist and antagonist is not obligatory; however AG1 was then prolonged. Our findings emphasize that neuronal commands to opposing muscles acting at a joint must be adapted to constraints imposed by the properties of the neuromuscular plant.     

Task differences with the same load torque alter the endurance time of submaximal fatiguing contractions in humans

Endurance time, muscle activation, and mean arterial pressure were measured during two types of submaximal fatiguing contractions that required each subject to exert the same net muscle torque in the two tasks. Sixteen men and women performed isometric contractions at 15% of the maximum voluntary contraction (MVC) force with the elbow flexor muscles, either by maintaining a constant force while pushing against a force transducer (force task) or by supporting an equivalent inertial load while maintaining a constant elbow angle (position task). The endurance time for the force task (1402 +/- 728 s) was twice as long as that for the position task (702 +/- 582 s, P < 0.05), despite a similar reduction in the load torque at exhaustion for each contraction. The rate of increase in average electromyographic activity (EMG, % peak MVC value) for the elbow flexor muscles was similar for the two tasks. However, the average EMG was greater at exhaustion for the force task (22.4 +/- 1.2%) compared with the position task (14.9 +/- 1.0%, P < 0.05). In contrast, the rates of increase in the mean arterial pressure, the rating of perceived exertion, anterior deltoid EMG, and fluctuations in motor output (force or acceleration) were greater for the position task compared with the force task (P < 0.05). Furthermore, the rate of bursts in EMG activity, which corresponded to the transient recruitment of motor units, was greater for the brachialis muscle during the position task. These results indicate that the briefer endurance time for the position task was associated with greater levels of excitatory and inhibitory input to the motor neurons compared with the force task.