A model for a motor unit train recorded during constant force isometric contractions

A model for the motor-unit action-potential train is developed, based on previously obtained empirical information. The auto and cross-correlation functions are calculated. The autocorrelation function is used to derive the mean rectified value, the variance and the root-mean-squared value of a motor-unit action-potential train. These parameters are solved by using two approximations for the motor-unit action-potential; a piece-wise approximation and a Dirac Delta function approximation. The Dirac Delta function approximation sufficiently simplifies the mathematics so that the model can be extended to myoelectric signals. The cross-correlation function contains information about the synchronization of motorunit action-potential trains that may be useful as an objective indicator of muscle fatigue.     

Derivation of some parameters of myoelectric signals recorded during sustained constant force isometric contractions.

Mathematical expressions are derived for some parameters of the myoelectric (ME) signal recorded during a constant force isometric contraction. The expressions are developed from a stochastic model for the motor-unit action-potential trains obtained from empirical results. The following parameters: (a) the mean rectified value, (b) the mean integrated rectified value, (c) the root-mean-square value, and (d) the power density spectrum are described as functions of contraction time and constant force of an isometric muscle contraction. The calculated parameters are compared to their corresponding empirically obtained measurements which have been reported in the literature. A discussion on the behavior of the parameters during increasing contraction time is presented. Synchronization of the motor-unit action-potential trains is shown to have a pronounced effect on the parameters of the myoelectric signal. This result should be considered when analyzing long records of myoelectric signals.     

Experimentally verified model of mechanomyograms recorded during single motor unit contractions

The aim of the paper is to create a model which enables to observe the mechanomyographic (MMG) wave generated during single motor unit contractions in a muscle, while the muscle is immersed in paraffin oil. The muscle model is described as a rheological membrane. Both the muscle and the medium models have been built by using Stiff-Finite-Element-Method (SFEM), which allows one to simulate the muscle surface displacement and the acoustic propagation of this effect in the oil. Such a modelling enables one to determine the impact of the rheological properties of the liquid environment on the shape of the MMG wave. In order to verify the model, the MMG signals and the contraction forces have been recorded in vivo from the medial gastrocnemius muscle of a rat. In these experiments single motor units were stimulated with various stimulation frequencies. A piezotransducer, immersed in paraffin oil, has been used to record the MMG signal recording. The signals recorded during individual twitches of the motor units have been used to estimate the parameters of the model. Subsequently, the model has been experimentally verified. The signals recorded in experiments during unfused and fused tetani have been compared with the simulated model responses in the analogous stimulation program. It has been observed that the MMG signals obtained with the proposed linear model have been consistent with the results of in vivo experiments.     

Motor unit recruitment during prolonged isometric contractions

Motor unit recruitment patterns were studied during prolonged isometric contraction using fine wire electrodes. Single motor unit potentials were recorded from the brachial biceps muscle of eight male subjects, during isometric endurance experiments conducted at relative workloads corresponding to 10% and 40% of maximal voluntary contraction (MVC), respectively. The recordings from the 10% MVC experiment demonstrated a characteristic time-dependent recruitment. As the contraction progressed both the mean number of motor unit spikes counted and the mean amplitude of the spikes increased significantly (P<0.01). This progressive increase in spike activity was the result of a discontinuous process with periods of increasing and decreasing activity. The phenomenon in which newly recruited motor units replace previously active units is termed motor unit rotation and appeared to be an important characteristic of motor control during a prolonged low level contraction. In contrast to the 10% MVC experiment, there was no indication of de novo recruitment in the 40% MVC experiment. Near the point of exhaustion a marked change in action potential shape and duration dominated the recordings. These findings demonstrate a conspicuous difference in the patterns of motor unit recruitment during a 10% and a 40% MVC sustained contraction. It is suggested that there is a close relationship between intrinsic muscle properties and central nervous system recruitment strategies which is entirely different in fatiguing high and low level isometric contractions.     

Age-related decreases in motor unit discharge rate and force control during isometric plantar flexion

Aging is related to multiple changes in muscle physiology and function. Previous findings concerning the effects of aging on motor unit discharge rate (DR) and fluctuations in DR and force are somewhat contradictory. Eight YOUNG and nine OLD physically active males performed isometric ramp (RECR) and isotonic (ISO) plantar flexions at 10 and 20% of surface EMG at MVC. Motor unit (MU) action potentials were recorded with intramuscular fine-wire electrodes and decomposed with custom build software “Daisy”. DR was lower in OLD in RECR-10% (17.9%, p < 0.001), RECR-20% (15.8%, p < 0.05), ISO-10% (17.7%, p < 0.01) and ISO-20% (14%, n.s.). In YOUNG force fluctuations were smaller at ISO-10% (72.1%, p < 0.001) and ISO-20% (55.2%, p < 0.05) which were accompanied with a slight increase in DR variation (n.s.). The observed lower DR in OLD is in line with earlier findings in small distal muscles. Also the larger force fluctuation in OLD was in line with previous studies with smaller hand muscles. These findings suggest that the age-related changes in MU control do exist also in large leg extensors that play an important role in human locomotion and balance control.     

Discharge properties of motor units during steady isometric contractions performed with the dorsiflexor muscles

The purpose of this study was to record the discharge characteristics of tibialis anterior motor units over a range of target forces and to import these data, along with previously reported observations, into a computational model to compare experimental and simulated measures of torque variability during isometric contractions with the dorsiflexor muscles. The discharge characteristics of 44 motor units were quantified during brief isometric contractions at torques that ranged from recruitment threshold to an average of 22 ± 14.4% maximal voluntary contraction (MVC) torque above recruitment threshold. The minimal [range: 5.8-19.8 pulses per second (pps)] and peak (range: 8.6-37.5 pps) discharge rates of motor units were positively related to the recruitment threshold torque (R(2) ≥ 0.266; P < 0.001). The coefficient of variation for interspike interval at recruitment was positively associated with recruitment threshold torque (R(2) = 0.443; P < 0.001) and either decreased exponentially or remained constant as target torque increased above recruitment threshold torque. The variability in the simulated torque did not differ from the experimental values once the recruitment range was set to ～85% MVC torque, and the association between motor twitch contraction times and peak twitch torque was defined as a weak linear association (R(2) = 0.096; P < 0.001). These results indicate that the steadiness of isometric contractions performed with the dorsiflexor muscle depended more on the distributions of mechanical properties than discharge properties across the population of motor units in the tibialis anterior.     

Use of surface EMG for studying motor unit recruitment during isometric linear force ramp.

The aim of this work was to demonstrate the rank order of motor unit (MU) recruitment by surface EMG based on a Laplacian detection technique and to document the MU features at their recruitment threshold. Surface EMG signals were recorded on the biceps brachii of 10 healthy subjects during linear force ramps. When achievable, the signals were decomposed into MU action potential (MUAP) trains. MU inter-pulse interval (IPI), conduction velocity (MUCV) and amplitude were estimated on the first 12 MUAPs of each detectable train in order to characterize the MU features at their firing onset. A strong correlation was found between MU recruitment threshold and IPI, MUCV, and amplitude, showing that the size principle can be demonstrated by a fully non-invasive EMG technique. However, signal decomposition was not possible on seven subjects due to the effects of the volume conductor when the skinfold thickness was too large. When requirements for an optimal detection of MUAP trains are met, surface EMG may be used to improve our understanding of MU activity.     

Muscular sound and force relationship during isometric contraction in man

The contracting muscle generates a low frequency sound detectable at the belly surface, ranging from 11 to 40 Hz. To study the relationship between the muscular sound and the intensity of the contraction a sound myogram (SMG) was recorded by a contact sensor from the biceps brachii of seven young healthy males performing 4-s isometric contractions from 10% to 100% of the maximal voluntary contraction (MVC), in 10% steps. Simultaneously, the electromyogram (EMG) was recorded as an index of muscle activity. SMG and EMG were integrated by conventional methods (iSMG and iEMG). The relationship between iSMG and iEMG vs MVC% is described by parabolic functions up to 80% and 100% MVC respectively. Beyond 80% MVC the iSMG decreases, being about half of its maximal value at 100% MVC. Our results indicate that the motor unit recruitment and firing rate affect the iSMG and iEMG in the same way up to 80% MVC. From 80% to 100% MVC the high motor units' discharge rate and the muscular stiffness together limit the pressure waves generated by the dimensional changes of the active fibres. The muscular sound seems to reflect the intramuscular visco-elastic characteristics and the motor unit activation pattern of a contracting muscle.     

Sympathetic control of the forearm blood flow in man during brief isometric contractions

Experiments were performed to assess the possible neurally mediated constriction in active skeletal muscle during isometric hand-grip contractions. Forearm blood flow was measured by venous occlusion plethysmography on 5 volunteers who exerted a series of repeated contractions of 4 s duration every 12 s at 60% of their maximum strength of fatigue. The blood flows increased initially, but then remained constant at 20-24 ml X min(-1) X 100 ml(-1) throughout the exercise even though mean arterial blood pressure reached 21-23 kPa (160-170 mm Hg). When the same exercise was performed after arterial infusion of phentolamine, forearm blood flow increased steadily to near maximal levels of 38.7 +/- 1.4 ml X min(-1) X 100 ml(-1). Venous catecholamines, principally norepinephrine, increased throughout exercise, reaching peak values of 983 +/- 258 pg X ml(-1) at fatigue. Of the vasoactive substances measured, the concentration of K+ and osmolarity in venous plasma also increased initially and reached a steady-state during the exercise but ATP increased steadily throughout the exercise. These data indicate a continually increasing alpha-adrenergic constriction to the vascular beds in active muscles in the human forearm during isometric exercise, that is only partially counteracted by vasoactive metabolites.     

Distribution of motor unit potential velocities in the biceps brachii muscle of sprinters and endurance athletes during short static contractions at low force levels

In surface electromyography (sEMG), the distribution of motor unit potential (MUP) velocities has been shown to reflect the proportion of faster and slower propagating MUPs. This study investigated whether the distribution of MUP velocities could distinguish between sprinters and endurance athletes in not-specifically trained muscle (biceps brachii). sEMG results were acquired from 15 sprinters and 18 endurance athletes during short static contractions (3.8 s) at three force levels: unloaded, 10% and 20% of maximum voluntary contraction. The features extracted from the sEMG were: the mean muscle conduction velocity (CV) – estimated using the inter-peak latency and the cross-correlation methods, the within-subject skewness of MUP velocities (expressing the relative proportions of faster and slower propagating MUPs), and the within-subject standard deviation of MUP velocities. Sprinters had a higher CV than endurance athletes using both methods. Sprinters also demonstrated a greater proportion of fast propagating MUPs, as indicated by the skewness. Thus, the distribution of MUP velocities was able to demonstrate physiological differences between sprinters and endurance athletes during short contractions at low forces. The findings can be extrapolated to the motor unit level. Since the investigated muscle was not involved in specific training, the differences seem to reflect inherited properties.     

Mechanomyography and electromyography force relationships during concentric, isometric and eccentric contractions.

The purpose of this study was to investigate systematically if complementary knowledge could be obtained from the recordings of electromyography (EMG) and mechanomyography (MMG) signals. EMG and MMG activities were recorded from the first dorsal interosseous muscle during slow concentric, isometric, and eccentric contraction at 0, 25, 50, 75 and 100% of the maximal voluntary contraction (MVC). The combination of the EMG and MMG recordings during voluntary concentric-isometric-eccentric contraction showed significant different non-linear EMG/force and MMG/force relationships (P<0.001). The EMG root mean square (rms) values increased significantly from 0 to 50% MVC during concentric and isometric contraction and up to 75% MVC during eccentric contraction (P<0.05). The MMG rms values increased significantly from 0 to 50% MVC during concentric contraction (P<0.05). The non-linear relationships depended mainly on the type and the level of contraction together with the angular velocity. Furthermore, the type of contraction, the contraction level, and the angular velocity influenced the electromechanical efficiency evaluated as the MMG to EMG ratio (P<0.05). These results highlight that EMG and MMG provide complementary information about the electrical and mechanical activity of the muscle. Different activation strategies seem to be used during graded isometric and anisometric contraction.     

Motor unit firing rates during isometric voluntary contractions performed at different muscle lengths

Firing rates of motor units and surface EMG were measured from the triceps brachii muscles of able-bodied subjects during brief submaximal and maximal isometric voluntary contractions made at 5 elbow joint angles that covered the entire physiological range of muscle lengths. Muscle activation at the longest, midlength, and shortest muscle lengths, measured by twitch occlusion, averaged 98%, 97%, and 93% respectively, with each subject able to achieve complete activation during some contractions. As expected, the strongest contractions were recorded at 90 degrees of elbow flexion. Mean motor unit firing rates and surface EMG increased with contraction intensity at each muscle length. For any given absolute contraction intensity, motor unit firing rates varied when muscle length was changed. However, mean motor unit firing rates were independent of muscle length when contractions were compared with the intensity of the maximal voluntary contraction (MVC) achieved at each joint angle.     

Electromyogram and force fluctuation during different linearly varying isometric motor tasks

The purpose of this work was to verify if deviation from the mirror-like behaviour of the motor units activation strategy (MUAS) and de-activation strategy (MUDS) and the degree of the error of the motor control system, during consecutive linearly increasing–decreasing isometric tension tasks, depend on the maximum reached tension and/or on the rate of tension changes. In 12 male subjects the surface EMG and force produced by the first dorsal interosseus activity were recorded during two (a and b) trapezoid isometric contractions with different plateau (a: 50% maximal voluntary contraction (MVC) and b: 100% MVC) and rate of tension changes (a: 6.7% MVC/s and b: 13.3% MVC/s) during up-going (UGR) and down-going (DGR) ramps. Ten steps (ST) 6 s long at 5, 10, 20, 30, 40, 50, 60, 70, 80 and 90% MVC were also recorded. The root mean square (RMS) and mean frequency (MF) from EMG and the relative error of actual force output with respect to the target (% ERR) were computed. The EMG-RMS/% MVC and EMG-MF/% MVC relationships were not overlapped when the ST and DGR as well as the UGR and DGR data were compared. The % ERR/% MVC relationships during a and b contractions differed from ST data only below 20% MVC. It can be concluded that MUAS and MUDS are not mirroring one each other because MU recruitment or de-recruitment threshold may be influenced by the maximum effort and by the % MVC/s of UGR and DGR. The role of MUs mechanical and/or central nervous system hysteresis on force decrement control is discussed.     

Spatial reorganisation of muscle activity correlates with change in tangential force variability during isometric contractions

The aim of this study was to quantify the effects of spatial reorganisation of muscle activity on task-related and tangential components of force variability during sustained contractions. Three-dimensional forces were measured from isometric elbow flexion during submaximal contractions (50 s, 5–50% of maximal voluntary contraction (MVC)) and total excursion of the centre of pressure was extracted. Spatial electromyographic (EMG) activity was recorded from the biceps brachii muscle. The centroids of the root mean square (RMS) EMG and normalised mutual information (NMI) maps were computed to assess spatial muscle activity and spatial relationship between EMG and task-related force variability, respectively. Result showed that difference between the position of the centroids at the beginning and at the end of the contraction of the RMS EMG and the NMI maps were different in the medial–lateral direction (P < 0.05), reflecting that muscle regions modulate their activity without necessarily modulating the contribution to the task-related force variability over time. Moreover, this difference between shifts of the centroids was positively correlated with the total excursion of the centre of pressure at the higher levels of contractions (>30% MVC, R² > 0.30, P < 0.05), suggesting that changes in spatial muscle activity could impact on the modulation of tangential forces. Therefore, within-muscle adaptations do not necessarily increase force variability, and this interaction can be quantified by analysing the RMS EMG and the NMI map centroids.     

Muscle weakness following sustained and rhythmic isometric contractions in man

The effects of sustained and rhythmically performed isometric contractions on electrically evoked twitch and tetanic force generation of the triceps surae have been investigated in 4 healthy male subjects. The isometric contractions were performed separately and on different occasions at 30%, 60% and 100% of the force of maximal voluntary contraction (MVC). The area under the maximal voluntary contraction (MVC) force/time curve during the rhythmic and sustained contractions was the same for each experiment. The results showed that following rhythmic isometric exercise there was a small decrease in low (10 and 20 Hz) and high (40 Hz) frequency tetanic tension which was associated with % MVC. However, there was no change in the 20/40 ratio of tetanic forces, MVC or the contraction times and force of the maximal twitch. In contrast, following sustained isometric exercise tetanic forces were markedly reduced, particularly at low frequencies of stimulation. The 20/40 ratio decreased and the induced muscle weakness was greater at 30% than 60% or 100% MVC. The performance of sustained isometric contractions also effected a decrease in contraction time of the twitch and MVC. The results are in accord with previous findings for dynamic work (Davies and White 1982), and show that if isometric exercise is performed rhythmically the effect on tetanic tensions is small and there is no evidence of a preferential loss of electrically evoked force at either high or low frequencies of stimulation following the contractions. For sustained contractions, however, the opposite is true, the ratio of 20/40 Hz forces is markedly reduced and following 30% sustained MVC there is a significant (p less than 0.05) change in the time to peak tension (TPT) of the maximal twitch.     

Sodium and potassium currents recorded during an action potential

A simple method was used to measure directly sodium and potassium currents underlying the action potential in single nerve fibres of Xenopus laevis. A short rectangular stimulus under current-clamp conditions elicited an action potential which was digitally stored and later used as command when voltageclamping the same fibre. The currents thus obtained nearly reproduced the original rectangular stimulus. Adding first 100 nM TTX and subsequently 100 nM TTX plus 10 mM TEA to the extracellular Ringer solution revealed the sodium and the potassium currents during an action potential. They were converted to permeabilities by use of the constant-field equation and are in good agreement with the curves which had been calculated from conventional voltage-clamp data. Thus experimentally determined currents and permeabilities are shown as they are changing during an action potential.     

Maximal voluntary isometric elbow flexion force during unilateral versus bilateral contractions in individuals with chronic stroke

The purpose of this study was to determine whether the phenomenon of bilateral deficit in muscular force production observed in healthy subjects and mildly impaired stroke patients also exists in patients with more chronic and greater levels of stroke impairment. Ten patients with chronic hemiparesis resulting from stroke performed unilateral and bilateral maximal voluntary isometric contractions of the elbow flexors. When the total force produced by both arms was compared, 12% less force was produced in the bilateral compared with unilateral condition (p=0.01). However, studying the effect of task conditions on each arm separately revealed a significant decline in nonparetic (p=0.01) but not paretic elbow flexor force in the bilateral compared with unilateral condition. Results suggest that a significant bilateral force deficit exists in the nonparetic but not the paretic arm in individuals with chronic stroke. Bilateral task conditions do not seem to benefit or impair paretic arm maximal isometric force production in individuals with moderate-severity chronic stroke.     

Information theoretic analysis of action potential trains

A crosscorrelational method of action potential trains has been proposed, based on information theory. Two information theoretic quantities, mutual information and channel capacity, were calculated from a pair of action potential trains for detecting a crosscorrelation and estimating synaptic connectivity. The method was compared with conventional ones, using action potential trains obtained by the simulation of a neuron model. This method was shown to have the advantages to more easily find a weak but significant crosscorrelation and to give better estimation of synaptic connectivity independent of the firing probability of a presynaptic neuron.     

Computer simulation study of the shape of motor unit action potential

Motor unit action potentials (MUAPs) of brachial biceps were simulated. A simulated MUAP was obtained as a sum of single fibre action potentials (SFAPs) from all the muscle fibres of a motor unit (MU). The influence of the following factors on MUAP shape for different kinds of recording electrode was studied: fibre density, neuromuscular jitter, temporal dispersion and electrode displacements. The simulation confirms that typical MUAPs recorded with needle electrodes from muscles of low fibre density such as brachial biceps are usually triphasic. Increased fibre density produces MUAPs of more complex shape and higher amplitude. Normal neuromuscular jitter is responsible for the variability of shape of subsequent potentials from the same MU as well as for electromyographic shimmer. Pathologic (increased) jitter makes the shapes of subsequent potentials unrecognizable. The influence of temporal dispersion is interconnected with other factors but rather of minor importance. The simulation shows how big changes in MUAP shape can be expected due to electrode displacements during single experiment or during estimation of MU territory.