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Movements of the crab gill bailer were entrained to an alternating current applied to the thoracic ganglion. There was little distortion of the muscle recruitment cycle, both absolute and relative coordination were observed, and the phase of the driven system in the driving cycle was a function of the difference between free running and driving frequencies.
In ten spastic patients and in an equal number of healthy controls, the relaxation phase occurring at the end of an isotonic voluntary contraction has been studied on soleus muscle using EMG and H-reflex methods. In the spastic group, the duration of motor unit-decruitment was consistently prolonged (from two to six times the control values). Moreover, the decrease in the excitability of the H-reflex arc, which normally accompanies the end of muscle contraction was delayed in time, reduced in amplitude or in some cases even absent. As a rule, patients in whom such release-associated inhibition (RAI) was lacking or severely reduced exhibited the longest motor unit-decruitment times, due to the interference of sustained clonic sequences. It is proposed that a lowered effectiveness of RAI might explain the clonic activity which frequently hinders voluntary muscle relaxation in spasticity.
The role of the differentiation of motor units (MU's) in the relation between integrated surface electromydgram and force was studied here for the biceps brachii, in the course of static contraction in man. The global EMG of the biceps brachii was recorded by two silver electrodes fixed to the skin. The MU's activity was simultaneously lead off by three bipolar wire electrodes inserted in three points of the muscle. MU firing frequency and recruitment were estimated by counting the action potentials on the three elementary records. The comparison between the increase of the integrated surface EMG and MU recruitment as a function of force shows that MU recruitment in itself cannot account for the increase in the integrated global EMG, particularly for high values of force. The difference between MU recruitmentand integrated global EMG, which can neither come from a possible MU's synchronisation as was shown in the discussion, is being suggested to be connected to the progressive firing of "phasic" MU's. This type of recruitment may also play an important role in the gradation of isometric force in normal man.
The paper reviews the fundamental components of stochastic and motor-unit-based models of the surface electromyogram (SEMG). Stochastic models used in ergonomics and kinesiology consider the SEMG to be a stochastic process whose amplitude is related to the level of muscle activation and whose power spectral density reflects muscle conduction velocity. Motor-unit-based models for describing the spatio-temporal distribution of individual motor-unit action potentials throughout the limb are quite robust, making it possible to extract precise information about motor-unit architecture from SEMG signals recorded by multi-electrode arrays. Motor-unit-based models have not yet been proven as successful, however, for extracting information about recruitment and firing rates throughout the full range of contraction. The relationship between SEMG and force during natural dynamic movements is much too complex to model in terms of single motor units.
Motor-unit activity in m. biceps brachii, m. brachialis and m. brachioradialis during isometric contractions has been compared with motor-unit activity during slow voluntary (extension and flexion) movements made against external loads. During these slow movements the recruitment threshold of m. biceps motor units is considerably lower than it is during isometric contractions but the recruitment threshold of both m. brachialis and m. brachioradialis motor units is considerably higher. For all three elbow flexor muscles the motor-unit firing frequency seems to depend on the direction of movement: the firing frequency is higher during flexion movements (3 deg/s) and lower during extension movements (-3 deg/s) than during isometric contractions. The relative contribution of the biceps to the total exerted flexion torque during slow voluntary movements is estimated to increase from 36% to about 48% and that of the brachialis/brachioradialis is estimated to decrease from 57% to about 45% compared to the relative contribution of these muscles during isometric contractions. This difference in the relative contribution of the three major elbow flexor muscles is shown to be caused by differences in the central activation in force tasks and movement tasks.
The relationship between nerve stimulation, pulse amplitude and isometric muscle force was measured to investigate recruitment of motor units. Force addition experiments were performed to obtain insight in the intersection of motor unit groups recruited by different electrodes. Intrafascicular and extraneural multielectrode configurations were used for nerve stimulation. Experiments were performed on rats. The common peroneal nerve was stimulated and the forces of the tibial anterior and extensor digitorum longus muscles were measured isometrically. Recruitment was more stable for intrafascicular electrodes than for extraneural electrodes. Especially for intrafascicular electrodes no strict inverse recruitment was observed. Force addition experiments indicated that small overlap of recruited motor unit groups occurred more often for intrafascicular than for extraneural electrodes.
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An important measure of the performance of a myoelectric control channel for powered artificial limbs is the myoelectric signal processor output signal-to-noise ratio (SNR). The signal and noise in this context are, respectively, the mean and variance of the estimate of some signal parameter to be used for control purposes. These quantities are determined by the signal processor, motor unit recruitment and motor unit firing statistics. The paper investigates, through analytical, simulation and experimental work, the role and significance of recruitment and firing statistics in channel performance. Equations are derived which express, for the single and multi-unit cases, channel SNR as a function of the number of active units, firing rates, action potential amplitude variation and action potential moments. A computer-simulated myoelectric signal is generated in which these variables can be controlled and SNR measured. The simulation results are compared with the theoretical and found to agree very well. Limited experiments with wire intramuscular electrodes and surface electrodes are performed to measure in vivo SNR from the biceps brachii muscle. The results of the experiments agree well with those of the simulation and theoretical work. The significance of this work is that it provides insight into the roles of important physiological parameters in myoelectric channel performance. It will also provide data necessary for the development of SNR enhancement techniques.
Intraneural stimulation with multi-electrodes in principle offers the best possibilities to reach selectivity at motor unit level and to improve recruitment order. The selectivity of stimulation in the peroneal nerve of the rat is explored in the paper by calculations and measurements, using a linear 12-electrode array and a newly devised selectivity test method. With analytical models for potential field distributions, areas of excitation can be calculated for arbitrary electrode configurations. It is demonstrated, using tripolar electrode combinations, how selectivity can be further enhanced and recruitment order improved.
Animal experiments and model simulations of monopolar, intrafascicular nerve stimulation are presented to study force-current relationships (recruitment curves). The conductivity of the extraneural medium is of prime importance to the resulting recruitment cures: an insulating extraneural medium generally leads to steeper curves with lower threshold currents than a well-conducting extraneural medium. Extensive statistical comparison of experimental and model results suggests the occurrence of clustering of alpha-motoneurons within the fascicle, manifesting itself mainly by an increased spread in threshold currents, as opposed to the situation where the fibres are distributed uniformly throughout the entire fascicle.
The recruitment order of motoneurons of first dorsal interosseous (1DI) muscle has been shown to be the same during either isometric abduction or flexion of the index finger. However, it is not known whether this stereotyped recruitment is the same for all functional tasks. This question was addressed by using the 1DI muscle, which contributes to many functional tasks involving the hand and fingers. Single motor units and surface electromyographic activity were recorded from 1DI in three human subjects. Each subject performed tasks of abduction of the index finger, rotation to unscrew a threaded knob and pincer to close a spring-loaded clip. The same motor units were activated for each of the three tasks, providing no evidence for the existence of task groups. The order of recruitment of pairs of selected motor units was the same for each of the three tasks.
The recruitment order of motor units (MU) was compared during voluntary and electrically induced contractions. With the use of spike-triggered averaging, a total of 302 MUs with recruitment thresholds ranging from 1% to 88% of maximal voluntary contraction were recorded in the human tibialis anterior muscle in five subjects. The mean (+/-SD) MU force was 98.3+/-93.3 mN (mean torque 16.8+/-15.9 mNm) and the mean contraction time (CT) 46.2+/-12.7 ms. The correlation coefficients (r) between MU twitch force and CT versus the recruitment threshold in voluntary contractions were +0.68 and -0.38 (P<0.001), respectively. In voluntary contractions, MUs were recruited in order of increasing size except for only 6% of the cases; whereas, during transcutaneous electrical stimulation (ES) at the muscle motor point, MU pairs showed a reversal of recruitment order in 28% and 35% of the observations, respectively, when the pulse durations were 1.0 ms or 0.1 ms. This recruitment reversal during ES was not related to the magnitude of the difference in voluntary recruitment thresholds between MUs. It is concluded that if the reversal of MU recruitment observed during ES is biophysically controlled by differences in their nerve axon input impedance, in percutaneous stimulation at the motor point, other factors such as the size and the morphological organisation of the axonal branches can also influence the order of activation.
Laryngeal adduction for swallowing chiefly involves contraction of the thyroarytenoid and lateral cricoarytenoid muscles to seal the glottic chink. Vocal cord elongation supplements closure through cricoarytenoid activation. Relaxation of the posterior cricoarytenoid muscle is also involved in the swallowing process. Recent interest has focused on stimulating the laryngeal nerves to protect the lower airway from conditions where normal muscular coordination may be disrupted (e.g., in aspiration following stroke). Unfortunately, electrical stimulation results in a generalized contraction of all the dependent intrinsic laryngeal muscles because the larger, more excitable axons fire before their smaller counterparts can be activated. In the physiological state, however, the smaller fibers are recruited first. The current study focuses on electronic manipulation of force in the glottic muscles involved in deglutition. We used a stimulator that could selectively activate the intrinsic laryngeal muscles based on their specific motor unit architectures. In 5 dogs, the circuit recruited the axons in the recurrent and superior laryngeal nerves from small to large. The muscles were identified according to the differential recruitment rates of their compound muscle action potentials as they appeared on the graph. The smaller axons in the thyroarytenoid recruited faster than the large ones found in the lateral cricoarytenoid muscles, with intermediate figures observed with the cricothyroid. The posterior cricoarytenoid presented with the slowest recruitment rates, as expected from this muscle's highest contingent of larger motor units. Latencies between the onsets of stimulations and muscle saturations also appeared stable. This approach to manipulating glottic force saves energy because it allows stimulating the adductory muscles with minimal interference from their abductor antagonist.
Glottic adduction couples with relaxation of the tonically contracted upper esophageal sphincter (UES) to constitute the end point of pharyngeal swallowing. Together with deglutitive laryngeal elevation, this reciprocal relationship contributes to protecting the lungs from aspiration. Degrees of uncoordination between glottic and upper esophageal sphincters can be seen under diverse circumstances of neurologic damage such as stroke, gastroesophageal reflux, and in the growing elderly population presenting with weaker musculature. We hypothesized that reciprocal coupling between glottic closure and UES relaxation may be artificially reestablished through vagal stimulation if the appropriate neural centers and their leading pathways remain capable of exciting a critical number of motor units. Orderly recruitment of the vagus nerve was produced in three dogs with a circuit superimposing 600-Hz, 3800-0-microA blocking over 10-70-Hz, 0-2300-microA stimulating currents. Amplitudes of motor unit or compound muscle action potentials were recorded from the thyroarytenoideus and cricopharyngeus via surface electromyographic electrodes. Stimulation was accompanied by a drop in UES intraluminal pressure. The reciprocal relationship between the two muscles could be time locked within certain stimulus parameters. We submit that electronic coupling between the glottic and UESs may show promise in preventing aspiration under selected circumstances.
Many studies have identified changes in trunk muscle recruitment in clinical low back pain (LBP). However, due to the heterogeneity of the LBP population these changes have been variable and it has been impossible to identify a cause-effect relationship. Several studies have identified a consistent change in the feedforward postural response of transversus abdominis (TrA), the deepest abdominal muscle, in association with arm movements in chronic LBP. This study aimed to determine whether the feedforward recruitment of the trunk muscles in a postural task could be altered by acute experimentally induced LBP. Electromyographic (EMG) recordings of the abdominal and paraspinal muscles were made during arm movements in a control trial, following the injection of isotonic (non-painful) and hypertonic (painful) saline into the longissimus muscle at L4, and during a 1-h follow-up. Movements included rapid arm flexion in response to a light and repetitive arm flexion-extension. Temporal and spatial EMG parameters were measured. The onset and amplitude of EMG of most muscles was changed in a variable manner during the period of experimentally induced pain. However, across movement trials and subjects the activation of TrA was consistently reduced in amplitude or delayed. Analyses in the time and frequency domain were used to confirm these findings. The results suggest that acute experimentally induced pain may affect feedforward postural activity of the trunk muscles. Although the response was variable, pain produced differential changes in the motor control of the trunk muscles, with consistent impairment of TrA activity.
In the present study we investigated the extent to which individuals suffering from spastic tetraparesis as a consequence of cerebral palsy tune their trunk involvement to accuracy demands in a spoon-handling task. Twenty-two participants (ten adolescents with spastic tetraparesis and 12 control participants) had to transport a spoon filled with water or sugar to a small or a large bowl that was placed within reach. Even though trunk displacement was larger in the tetraparetic participants than it was in the control participants, the effects of the imposed accuracy constraints were remarkably similar. Participants in both groups increased trunk displacement with increasing precision requirements. Furthermore, in both groups the largest trunk involvement was found in the initial and final part of the substance-transporting phase, when wrist velocity was lowest. We propose several explanations for these findings and conclude that the large trunk involvement in individuals with tetraparetic cerebral palsy should, in any case, not be regarded as a primary symptom of the disorder, but rather as an adaptive reaction to increased task demands.
In addition to local biomechanical and reflex factors influencing muscle activation, global factors may be used by the nervous system to control all muscles in a coherent and task-specific way. It has been hypothesized that a virtual or referent (R) configuration of the body determined by muscle recruitment thresholds specified by neural control levels is such a factor. Due to the threshold nature of the R configuration, the activity of each muscle depends on the difference between the actual (Q) and the R configuration of the body. The nervous system modifies the R configuration to produce movement. One prediction of this hypothesis is that the Q and R configurations may match each other, most likely in movements with reversals in direction, resulting in a minimum in the electromyographic (EMG) activity level of muscles involved. The depth of the minima is constrained by the degree of coactivation of opposing muscle groups. Another prediction is that EMG minima in the activity of multiple muscles may occur not only when the movement is assisted but also when it is opposed by external forces (e.g., gravity). To verify these predictions, we analyzed EMG patterns of 16-21 functionally diverse muscles of the legs, trunk, and arms during jumping and stepping in place. One EMG minimum in the activity of all muscles regularly occurred near the apex of the jump. A minimum was also observed near the point of transition of the body from flexion to extension leading to a jump. During stepping in place, the activity of muscles of each side of the body was usually minimized near the beginning and near the end of the stance phase as well as during the maximum elevation of the foot. Since EMG minima occurred not only during gravity-assisted but also gravity-opposed movement reversals, it is concluded that neural factors (such as matching between the Q and R) rather than mechanical factors are responsible for minimizing the EMG activity in these movements.
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 (d F/d t) 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.