Influence of proprioceptive feedback on the firing rate and recruitment of motoneurons.

We investigated the relationships of the firing rate and maximal recruitment threshold of motoneurons recorded during isometric contraction with the number of spindles in individual muscles. At force levels above 10% of maximal voluntary contraction, the firing rate was inversely related to the number of spindles in a muscle, with the slope of the relationship increasing with force. The maximal recruitment threshold of motor units increased linearly with the number of spindles in the muscle. Thus, muscles with a greater number of spindles had lower firing rates and a greater maximal recruitment threshold. These findings may be explained by a mechanical interaction between muscle fibres and adjacent spindles. During low-level (0% to 10%) voluntary contractions, muscle fibres of recruited motor units produce force twitches that activate nearby spindles to respond with an immediate excitatory feedback that reaches maximal level. As the force increases further, the twitches overlap and tend towards tetanization, the muscle fibres shorten, the spindles slacken, their excitatory firings decrease, and the net excitation to the homonymous motoneurons decreases. Motoneurons of muscles with greater number of spindles receive a greater decrease in excitation which reduces their firing rates, increases their maximal recruitment threshold, and changes the motoneuron recruitment distribution.

A model of motoneuron behavior and muscle-force generation for sustained isometric contractions.

A model for the simulation of motoneuron firing behavior and muscle force during sustained constant-force isometric contractions was developed. It provides a non-linear relationship between the excitation to the motoneuron pool of a muscle and the firing behavior of motor units; it implements muscle mechanical changes induced by fatigue and it comprises a feedback loop to maintain the muscle force at a given target level. We simulated a series of repeated force contractions sustained at 20% MVC with the first dorsal interosseous muscle of the hand and the vastus lateralis muscle of the thigh. The model generates force and firing behaviors which are consistent with experimental findings and underscores the influence of muscle mechanical changes on the control behavior of motor units during sustained contractions. The model predicts the increase of force fluctuation with fatigue in both muscles, likely due to recruitment of high-threshold high-amplitude twitch motor units. Force variability is greater in the first dorsal interosseous muscle than in the vastus lateralis muscle at any time during the contraction series, due to the different electrical and mechanical properties of the muscles.

Electro-mechanical stability of surface EMG sensors.

This study compared the performance of surface electromyographic (sEMG) sensors for different detection conditions affecting the electro-mechanical stability between the sensor and its contact with the skin. These comparisons were made to gain a better understanding of how specific characteristics of sensor design and use may alter the ability of sEMG sensors to detect signals with high fidelity under conditions of vigorous activity. The first part of the study investigated the effect of different detection surface contours and adhesive tapes on the ability of the sensor to remain in electrical contact with the skin. The second part of the study investigated the effects of different skin preparations and hydrophilic gels on the production of movement artifact resulting from sinusoidal and impact mechanical perturbations. Both parts of the study evaluated sensor performance under dry skin and wet skin (from perspiration) conditions. We found that contouring the detection surface and adding a more adhesive double-sided tape were effective in increasing the forces needed to disrupt the electrical contact between the electrodes and the skin for both dry skin and wet skin conditions. The mechanical perturbation tests demonstrated that hydrophilic gel applied to the detection surface of the sensor produced greater movement artifacts compared to sensors without gel, particularly when the sensors were tested under conditions in which perspiration was present on the skin. The use of a surfactant skin preparation did not influence the amount of movement artifacts that resulted from either the sinusoidal or impact perturbations. The importance of these findings is discussed in terms of their implications for improving sEMG signal fidelity through sensor design modifications and procedures for interfacing them with the skin.

A sEMG-based method for assessing the design of computer mice.

Computer users who experience repetitive wrist movements and awkward hand positions are prone to developing upper extremity disorders. Manufacturers have designed various ergonomic mice in response to complaints of pain and discomfort related to computer mouse use. The objective of this work was to validate the use of surface electromyography (sEMG) in assessing the design of nonkeyboard input devices (computer mice). While holding the computer mouse in different grasp positions sEMG of the forearm and hand were recorded during a set of static tasks. The sEMG signal provided information regarding the level of muscle activity and the varied combinations of muscular effort needed to position the hand in a specified posture. A significant decrease in the level of sEMG activity was observed for the pronator muscles when subjects were tested using ergonomic computer mice. The sEMG-based method was validated to be sensitive to the impact of subtle differences in shape/design on the amplitude of the surface EMG data. We also proved a significant effect of hand size and grasp position on the level of muscle activity associated with different mice.

Motor unit recruitment and derecruitment induced by brief increase in contraction amplitude of the human trapezius muscle.

The activity pattern of low-threshold human trapezius motor units was examined in response to brief, voluntary increases in contraction amplitude ('EMG pulse') superimposed on a constant contraction at 4-7 % of the surface electromyographic (EMG) response at maximal voluntary contraction (4-7 % EMGmax). EMG pulses at 15-20 % EMGmax were superimposed every minute on contractions of 5, 10, or 30 min duration. A quadrifilar fine-wire electrode recorded single motor unit activity and a surface electrode recorded simultaneously the surface EMG signal. Low-threshold motor units recruited at the start of the contraction were observed to stop firing while motor units of higher recruitment threshold stayed active. Derecruitment of a motor unit coincided with the end of an EMG pulse. The lowest-threshold motor units showed only brief silent periods. Some motor units with recruitment threshold up to 5 % EMGmax higher than the constant contraction level were recruited during an EMG pulse and kept firing throughout the contraction. Following an EMG pulse, there was a marked reduction in motor unit firing rates upon return of the surface EMG signal to the constant contraction level, outlasting the EMG pulse by 4 s on average. The reduction in firing rates may serve as a trigger to induce derecruitment. We speculate that the silent periods following derecruitment may be due to deactivation of non-inactivating inward current ('plateau potentials'). The firing behaviour of trapezius motor units in these experiments may thus illustrate a mechanism and a control strategy to reduce fatigue of motor units with sustained activity patterns.

Time-frequency parameters of the surface myoelectric signal for assessing muscle fatigue during cyclic dynamic contractions.

The time-dependent shift in the spectral content of the surface myoelectric signal to lower frequencies has proven to be a useful tool for assessing localized muscle fatigue. Unfortunately, the technique has been restricted to constant-force, isometric contractions because of limitations in the processing methods used to obtain spectral estimates. A novel approach is proposed for calculating spectral parameters from the surface myoelectric signal during cyclic dynamic contractions. The procedure was developed using Cohen class time-frequency transforms to define the instantaneous median and mean frequency during cyclic dynamic contractions. Changes in muscle length, force, and electrode position contribute to the nonstationarity of the surface myoelectric signal. These factors, unrelated to localized fatigue, can be constrained and isolated for cyclic dynamic contractions, where they are assumed to be constant for identical phases of each cycle. Estimation errors for the instantaneous median and mean frequency are calculated from synthesized signals. It is shown that the instantaneous median frequency is affected by an error slightly lower than that related to the instantaneous mean frequency. In addition, we present a sample application to surface myoelectric signals recorded from the first dorsal interosseous muscle during repetitive abduction/adduction of the index finger against resistance. Results indicate that the variability of the instantaneous median frequency is related to the repeatability of the biomechanics of the exercise.

Motor control of low-threshold motor units in the human trapezius muscle.

The firing pattern of low-threshold motor units was examined in the human trapezius and first dorsal interosseous (FDI) muscles during slowly augmenting, low-amplitude contractions that were intended to mimic contractile activity in postural muscles. The motor unit activity was detected with a special needle electrode and was analyzed with the assistance of computer algorithms. The surface electromyographic (EMG) signal was recorded. Its root-mean-square (RMS) value was calculated and presented to the subject who used it to regulate the muscle force level. In the trapezius, there was minimal, if any, firing rate modulation of early recruited motor units during slow contractions (< or =1% EMG(max)/s), and later recruited motor units consistently presented higher peak firing rates. As the force rate of the contraction increased (3% EMG(max)/s), the firing rates of the motor units in the trapezius approached an orderly hierarchical pattern with the earliest recruited motor units having the greatest firing rate. In contrast, and as reported previously, the firing rates of all motor units in the FDI always presented the previously reported hierarchical "onion-skin" pattern. We conclude that the low-threshold motor units in the postural trapezius muscle, that is the motor units that are most often called on to activate the muscle in postural activities, have different control features in slow and fast contractions. More detailed analysis revealed that, in the low force-rate contractions of the trapezius, recruitment of new motor units inhibited the firing rate of active motor units, providing an explanation for the depressed firing rate of the low-threshold motor units. We speculate that Renshaw cell inhibition contributes to the observed deviation of the low-threshold motor units from the hierarchical onion-skin pattern.

Effects of aging on motor-unit control properties.

It was hypothesized that the age-related alterations in the morphological properties of a motor unit would be accompanied by modifications in the control aspects of the motor unit, as either an adaptive or compensatory mechanism to preserve smooth force production. In specific, the objective of the study was to investigate the age-related alterations in the concurrent firing behavior of multiple motor units in the first dorsal interosseous (FDI) muscle in isometric contractions at 20 and 50% of the subject's voluntary contraction level. Analysis of the data collected from 10 young (24-37 yr of age) and 10 elderly (65-88 yr of age) subjects led to three novel observations regarding the firing behavior of aged motor units. 1) Among elderly subjects, there is a decrease in the common fluctuations that are observed among the firing rates of motor units in the young. 2) The relationship observed between the firing rate and recruitment threshold of young subjects is disturbed in the elderly. Although in young subjects, at any point in a given submaximal contraction, earlier recruited motor units have higher firing rates than later-recruited units; in aged subjects this dependency of firing rate on recruitment rank is compromised. 3) The progressive decrease observed in the firing rates of concurrently active motor units in constant-force contractions in the young is not seen in the aged. In addition to these original findings, this study provided support for earlier reports of 1) decreased average firing rates probably reflecting the slowing of the muscle, 2) a shift in recruitment thresholds toward lower force levels in line with the shift toward type I fibers, and 3) multiphasic action potential shapes indicative of the reinnervation process that takes place during aging. Taken as a whole, these findings indicate significant age-related modifications in the control properties of human motor units.

Motor unit substitution in long-duration contractions of the human trapezius muscle.

We examined the activity pattern of low-threshold motor units in the human trapezius muscle during contractions of 10 min duration. Three procedures were applied in sequence: 1) static contraction controlled by maintaining a constant low level of the surface electromyogram (EMG)-detected root-mean-square signal, 2) a manipulation task with mental concentration, and 3) copying a text on a word processor. A quadrifilar fine-wire electrode was used to record single motor unit activity. Simultaneously, surface electrodes recorded the surface EMG signal. During these contractions, low-threshold motor units showed periods of inactivity and were substituted by motor units of higher recruitment threshold. This phenomenon was not observed during the first few minutes of the contraction. In several cases the substitution process coincided with a short period of inactivity in the surface EMG pattern. Substitution was observed in five of eight experiments. These observations may be explained by a time-variant recruitment threshold of motor units, sensitive to their activation history and temporal variation in the activity patterns. We speculate that the substitution phenomenon protects motor units in postural muscles from excessive fatigue when there is a demand for sustained low-level muscle activity.

Electromyographic analysis of grand-plié in ballet and modern dancers.

PURPOSE: The purpose of this report is to describe lower extremity muscle activity in grand-plié, as determined by EMG analysis; to compare and contrast muscle function in grand-plié and demi-plié to support the hypothesis that grand-plié is not simply a deeper demi-plié, but rather a fundamentally different movement in terms of muscle use; and to present further evidence in support of the hypothesis that ballet dancers use muscles differently than modern dancers in dance movement. METHODS: Surface electromyography was used to analyze lower extremity muscle activity during grand-plié in first position with lower extremities turned out in five ballet and seven modern female professional dancers. RESULTS: Electromyographic (EMG) activity of tibialis anterior included continuous activity from heel-off during the lowering phase, through midcycle, and ending at heel-on during the rising phase in all grand-pliés; the majority of tibialis anterior EMG tracings in ballet dancers had additional activity at the end of the rising phase. All EMG tracings for vastus lateralis and medialis included a peak of activity during the lowering phase, a decrease (valley) at midcycle, followed by another peak during the rising phase; increased activity at the end of the rising phase was observed in most grand-plié in ballet, and not modern, dancers. Adductor EMG activity was also observed in all tracings with a peak during the lowering phase from heel-off to midcycle, a valley at midcycle, followed by a peak of activity in early rising phase; the midcycle valley was of lower, and the rising phase peak of higher, activity in ballet compared with modern dancers. Variation of EMG patterns was observed for lateral and medial gastrocnemius, gluteus maximus, and hamstrings. CONCLUSIONS: The data support the concept that lower extremity muscle activity in dance movement is comprised of three major types: (a) unique, characteristic activity required for the execution of the movement; (b) varied activity which is characteristic of dancers of different dance idioms; and (c) varied activity which may depend on factors such as balance, personal habit, and individual training background. Furthermore, EMG activity of vastus lateralis and medialis at the midcycle valley in grand-plié was significantly less in ballet dancers than in modern dancers despite similar degree of knee flexion, suggesting that ballet dancers may have lower patellofemoral joint reaction force at midcycle than modern dancers.

Hand dominance and motor unit firing behavior.

Daily preferential use was shown to alter physiological and mechanical properties of skeletal muscle. This study was aimed at revealing differences in the control strategy of muscle pairs in humans who show a clear preference for one hand. We compared the motor unit (MU) recruitment and firing behavior in the first dorsal interosseous (FDI) muscle of both hands in eight male volunteers whose hand preference was evaluated with the use of a standard questionnaire. Myoelectric signals were recorded while subjects isometrically abducted the index finger at 30% of the maximal voluntary contraction (MVC) force. A myoelectric signal decomposition technique was used to accurately identify MU firing times from the myoelectric signal. In MUs of the dominant hand, mean values for recruitment threshold, initial firing rate, average firing rate at target force, and discharge variability were lower when compared with the nondominant hand. Analysis of the cross-correlation between mean firing rate and muscle force revealed cross-correlation peaks of longer latency in the dominant hand than in the nondominant side. This lag of the force output with respect to fluctuations in the firing behavior of MUs is indicative of a greater mechanical delay in the dominant FDI muscle. MVC force was not significantly different across muscle pairs, but the variability of force at the submaximal target level was higher in the nondominant side. The presence of lower average firing rates, lower recruitment thresholds, and greater firing rate/force delay in the dominant hand is consistent with the notion of an increased percentage of slow twitch fibers in the preferentially used muscle, allowing twitch fusion and force buildup to occur at lower firing rates. It is suggested that a lifetime of preferred use may cause adaptations in the fiber composition of the dominant muscle such that the mechanical effectiveness of its MUs increased.

Classification of back muscle impairment based on the surface electromyographic signal.

A surface electromyographic (EMG) procedure for classifying muscle impairments in persons with low back pain (LBP) is described. The procedure was studied using a device, the Back Analysis System (BAS), to acquire and process EMG signals from six bilateral muscle sites during sustained isometric contractions designed to progressively fatigue the lower back. Back muscle impairment was determined on the basis of the different ways in which the EMG median frequency parameters change as a function of contraction duration and muscle site. The article describes a series of studies that have been useful in developing an automated procedure for identifying back muscle impairment by comparing individual test results to a normative database. To date, the research results have produced multivariate discriminant functions that have identified two muscle impairment categories associated with deconditioning and imbalances secondary to LBP. We have found that the functions can distinguish individuals with and without LBP with an accuracy of approximately 90%. Other studies are described in which the technique is applied to monitoring changes in muscle performance capability that occur following rehabilitation for LBP. Many of our findings here are also compared to the results of independent studies by others using similar procedures. The need for further research and development of the technique to improve its clinical applicability is also described.

Development of new protocols and analysis procedures for the assessment of LBP by surface EMG techniques.

Spectral parameters of the surface electromyographic (EMG) signal from lumbar back muscles assessed during a fatiguing isometric contraction can be used to classify different categories of low back pain (LBP) subjects and control subjects without LBP. In the test protocol currently used at the NeuroMuscular Research Center at Boston University, subjects contract their back muscles at 80% of their maximal voluntary contraction (MVC) force. This fatigue-based protocol has been successfully applied to persons with subacute or chronic LBP; those in acute pain, however, have not been included because of their inability to perform a maximal exertion. In this paper we will examine the force sensitivity of the currently used EMG parameters and also give an overview of some of our efforts to develop new test procedures. Our goal is to develop force-insensitive surface EMG parameters that can be used for classification purposes in populations of subjects who develop low trunk extension forces. In addition, the development of a model to predict MVC from anthropometrical measurements will be presented.

Motor unit control properties in constant-force isometric contractions.

1. The purpose of this study was 1) to characterize the decrease observed in mean firing rates of motor units in the first 8-15 s of isometric constant-force contractions and 2) to investigate possible mechanisms that could account for the ability to maintain force output in the presence of decreasing motor unit firing rates. 2. The decrease in mean firing rates was characterized by investigating myoelectric signals detected with a specialized quadrifilar needle electrode from the first dorsal interosseus (FDI) and the tibialis anterior (TA) muscles of 19 healthy subjects during a total of 85 constant-force isometric contractions at 30, 50, or 80% of maximal effort. The firing times of motor units were obtained from the myoelectric signals with the use of computer algorithms to decompose the signal into the constituent motor unit action potentials. Time-varying mean firing rates and recruitment thresholds were also calculated. 3. Motor units detected from the TA muscle were found to have a continual decrease in their mean firing rates in 36 of 44 trials performed during isometric ankle dorsiflexion at force values ranging from 30 to 80% of maximal effort and a duration of 8-15 s. Likewise, motor units detected in the FDI muscle displayed a decrease in firing rate in 32 of 41 trials performed during constant-force isometric index finger abduction for contractions ranging from 30 to 80% of maximal effort. In 14 contractions (16% of total), firing rates were essentially constant, whereas in 3 contractions (4%), firing rates appeared to increase. 4. Motor units with the higher recruitment thresholds and lower firing rates tended to display the greater decreases in firing rate over the constant-force interval, whereas motor units with lower recruitment thresholds and higher firing rates had lesser rates of decrease. Furthermore, increasing contraction levels tended to intensify the decrease in the motor unit firing rates. 5. Three possible mechanisms were considered as factors responsible for the maintaining of force output while motor units decreased their firing rates: motor unit recruitment, agonist/antagonist interaction, and twitch potentiation. Of these, motor unit recruitment was discarded first because none was observed during the 8-15 s duration of any of the 85 contractions. Furthermore, contractions outside the physiological range of motor unit recruitment (at 80% of maximal effort) revealed the same decreasing trend in firing rates, ruling out recruitment as the means of sustaining force output. 6. The role of agonist or antagonist muscle interaction was investigated with the use of the muscles controlling the wrist joint. Myoelectric signals were recorded with quadrifilar needle electrodes from the wrist extensor muscles while myoelectric activity in the wrist flexor muscles was concurrently monitored with surface electrodes during constant-force isometric wrist extension at 50% of maximal effort. Firing rates of the motor units in the wrist extensor muscles simultaneously decreased while the flexor muscles were determined to be inactive. 7. All the findings of this study regarding the behavior of the firing rates could be well explained by the reported characteristics of twitch potentiation that have been previously documented in animals and humans. 8. The results of this study, combined with the results of other investigators, provide the following scenario to explain how a constant-force isometric contraction is sustained. As the contraction progresses, the twitch force of the muscle fibers undergoes a potentiation followed by a decrease. Simultaneously, the "late adaptation" property of the motoneuron decreases the firing rate of the motor unit. Findings of this study suggest that voluntary reduction in firing rates also cannot be ruled out as a means to augment the adaptation in motoneurons. (ABSTRACT TRUNCATED)

Rank-ordered regulation of motor units.

Myoelectric signals were detected from the tibialis anterior muscle of 5 subjects with a quadrifilar needle electrode while the subjects generated isometric forces that increased linearly with time (10% of maximal voluntary contraction/s) up to maximal voluntary level. Motor unit firing rates were studied as a function of force throughout the full range of muscle force output. The relationship between force and firing rate was found to contain three distinct regions. At recruitment and near maximal force levels, firing rates increased more rapidly with force than in the intermediate region. Furthermore, in the regions with rapid increases, the rate of change of firing rate was correlated to the recruitment threshold, with higher recruitment threshold motor units displaying greater rates of change. In the intermediate region, all motor units had similar rates of change of firing rate. A weak positive correlation was found between initial firing rate and recruitment threshold. Firing rates of motor units at any instant were found to be ordered according to the recruitment order: at any given time in the contraction motor units with lower recruitment thresholds had higher firing rates than units with higher recruitment thresholds. Firing rates of all motor units were observed to converge to the same value at maximal forces. Mechanisms underlying motor unit recruitment and firing rate modulation are discussed in the context of a conceptual model.

Open-loop and closed-loop postural control mechanisms in Parkinson's disease: increased mediolateral activity during quiet standing.

Stabilogram-diffusion analysis was used to gain insights into how idiopathic Parkinson's disease (IPD) affects the postural control mechanisms involved in maintaining erect stance. Twenty-two subjects with IPD and twenty-four healthy elderly subjects were studied under eyes-open, quiet-standing conditions. The postural control mechanisms in the parkinsonian subjects, compared to the healthy elderly, were characterized by an increase in the effective stochastic activity in the mediolateral direction. Mediolateral posturographic measures were also associated with a history of falls and poor performance on clinical measures of balance. It is hypothesized that the increase in mediolateral activity in subjects with IPD may reflect an attempt to maintain potentially stabilizing movements during quiet standing in the face of impaired movement in the anteroposterior direction. This study supports the notion that mediolateral instability is an important posturographic marker of functional balance impairment in the elderly.

Effects of muscle fiber type and size on EMG median frequency and conduction velocity.

This paper describes an in vitro method for comparing surface-detected electromyographic median frequency (MF) and conduction velocity (CV) parameters with histochemical measurements of muscle fiber type composition and cross-sectional area (CSA). Electromyographic signals were recorded during electrically elicited tetanic contractions from rat soleus, extensor digitorum longus, and diaphragm muscles placed in an oxygenated Krebs bath. Fibers were typed as slow oxidative, fast oxidative glycolytic, and fast glycolytic based on histochemical enzyme stains. Muscles with a greater percentage of fast glycolytic and fast oxidative glycolytic fibers exhibited greater initial values of MF and CV as well as a greater reduction in these variables over the course of the contraction. Regression indicated that fiber type composition could be predicted based on two MF parameters. A weighted measure of muscle fiber CSA was found to be linearly related to both initial MF and CV. The results of this study suggest that MF and CV parameters recorded during a muscular contraction are related to muscle fiber type composition and muscle fiber CSA.

Upright, correlated random walks: A statistical-biomechanics approach to the human postural control system.

The task of maintaining erect stance involves a complex sensorimotor control system, the output of which can be highly irregular. Even when a healthy individual attempts to stand still, the center of gravity of his or her body and the center of pressure (COP) under his or her feet continually move about in an erratic fashion. In this study, we approach the problem of characterizing postural sway from the perspective of random-walk theory. Specifically, we analyze COP trajectories as one-dimensional and two-dimensional random walks. These analyses reveal that over short-term intervals of time during undisturbed stance the COP behaves as a positively correlated random walk, whereas over long-term intervals of time it resembles a negatively correlated random walk. We interpret this novel finding as an indication that during quiet standing the postural control system utilizes open-loop and closed-loop control schemes over short-term and long-term intervals, respectively. From this perspective, our approach, known as stabilogram-diffusion analysis, has the advantage that it leads to the extraction of COP parameters which can be directly related to the steady-state behavior and functional interaction of the neuromuscular mechanisms underlying the maintenance of erect stance. (c) 1995 American Institute of Physics.