Estimation of the muscle fibre semi-length under varying joint positions on the basis of non-invasively extracted motor unit action potentials.

Changes in muscle fibre length and surface electrode position with respect to the muscle fibres affect the amplitude and frequency characteristics of surface electromyography (SEMG) in different ways. Knowledge of changes in muscle fibre length would help towards a better interpretation of the signals. The possibility of estimating the length through SEMG during voluntary contractions was checked in this study. The fibres' semi-length was estimated from the product of the conduction velocity and conduction time during which the wave of excitation propagated from the end-plate region to the ends of the fibres. Short (10 s), moderate (30% of maximum voluntary contraction) isometric contractions were performed by 10 subjects at different elbow joint angles (80-140 degrees in steps of 20 degrees ). Monopolar signals were detected non-invasively, using a two-dimensional electrode array. High spatial resolution EMG and a decomposition technique were utilised to extract single motor unit activities for triggered averaging and to estimate conduction velocity. A significant increase with joint angle was found in conduction time and estimated fibre semi-length. Changes in conduction velocity with joint angle were found to be not significant. The methodology described allows the relative changes in fibres' semi-length to be estimated from SEMG data.     

Sensitivity of the cross-correlation between simulated surface EMGs for two muscles to detect motor unit synchronization.

The purpose of the study was to evaluate the use of cross-correlation analysis between simulated surface electromyograms (EMGs) of two muscles to quantify motor unit synchronization. The volume conductor simulated a cylindrical limb with two muscles and bone, fat, and skin tissues. Models of two motor neuron pools were used to simulate 120 s of surface EMG that were detected over both muscles. Short-term synchrony was established using a phenomenological model that aligned the discharge times of selected motor units within and across muscles to simulate physiological levels of motor unit synchrony. The correlation between pairs of surface EMGs was estimated as the maximum of the normalized cross-correlation function. After imposing four levels of motor unit synchrony across muscles, five parameters were varied concurrently in the two muscles to examine their influence on the correlation between the surface EMGs: 1) excitation level (5, 10, 15, and 50% of maximum); 2) muscle size (350 and 500 motor units); 3) fat thickness (1 and 4 mm); 4) skin conductivity (0.1 and 1 S/m); and 5) mean motor unit conduction velocity (2.5 and 4 m/s). Despite a constant and high level of motor unit synchronization among pairs of motor units across the two muscles, the cross-correlation index ranged from 0.08 to 0.56, with variation in the five parameters. For example, cross-correlation of EMGs from pairs of hand muscles, each having thin layers of subcutaneous fat and mean motor unit conduction velocities of 4 m/s, may be relatively insensitive to the level of synchronization across muscles. In contrast, cross-correlation of EMGs from pairs of leg muscles, with larger fat thickness, may exhibit a different sensitivity. These results indicate that cross correlation of the surface EMGs from two muscles provides a limited measure of the level of synchronization between motor units in the two muscles.     

Physiological characteristics of motor units in the brachioradialis muscle across fatiguing low-level isometric contractions.

The purpose of this study was to determine (i) if decomposition-based quantitative electromyography (DQEMG) could detect changes in motor unit potential (MUP) morphology and motor unit (MU) firing pattern statistics associated with muscle fatigue, (ii) if any detected changes are correlated with surface electromyographic (SEMG) signs of fatigue, and (iii) if significant fatigue-dependent changes are repeatable within individuals. Mean MU firing rates and the morphology of MUPs detected using needle and surface electrodes during constant-torque isometric contractions held until exhaustion were investigated in the brachioradialis (BR) muscle in 10 healthy volunteers (mean age=28.6 yr, SD+/-3.9). Time dependant changes were investigated using an analysis of variance with normalized time as a main effect. Partial correlation coefficients were computed using a repeated measures analysis of covariance to determine if changes in MU firing rates, needle-detected MUPs and surface-detected MUPs (SMUPs) were related to changes in SEMG signal amplitude and frequency parameters. Intraclass correlation coefficients (ICCs) were used to determine the within-subject repeatability of changes in MU firing rates, and MUP and SMUP parameters. Significant decreases in mean MU firing rates were found along with significant increases in various duration and area related parameters in both MUPs and SMUPs across the fatiguing contraction. The SEMG signal demonstrated the expected changes with fatigue: an increase in amplitude and a decrease in frequency content. SEMG amplitude was significantly positively correlated with SMUP peak-to-peak voltage (r=0.85, p<0.05), and SMUP area (r=0.86, p<0.05). Mean power frequency was significantly negatively correlated with SMUP negative peak duration (r=-0.74, p<0.05). The significant time-dependent changes were reliably observed (ICCs were 0.94 for MUP peak to peak amplitude, 0.97 for MUP area and 0.95 for MUP area to amplitude ratio, 0.95 for SMUP peak-to-peak voltage, 0.83 for SMUP area, 0.99 for SMUP negative peak amplitude and 0.88 for SMUP negative peak area). The decreases in mean MU firing rates measured along with the increases in amplitude, duration and area parameters of MUPs and SMUPs and their partial correlation with SEMG amplitude during submaximal fatiguing contractions of the BR, suggest that recruitment is a main cause of increased SEMG amplitude parameters with fatigue. We conclude that DQEMG can be effectively and reliably used to detect changes in physiological characteristics of MUs that accompany fatigue.     

Effect of electrocardiographic contamination on surface electromyography assessment of back muscles

The purpose of this study was to demonstrate the relative effect of electrocardiography (ECG) on back muscle surface electromyography (SEMG) parameters and their corresponding sensitivity in low back pain (LBP) assessment.Back muscle SEMG activities were recorded from 17 healthy subjects and 18 chronic LBP patients under static postures (straight sitting and upright standing), and dynamic action (flexion–extension). ECG cancellation based on independent component analysis (ICA) method was performed. Root mean square (RMS) and median frequency (MF) of raw and denoised SEMG data were computed respectively. Multiple comparisons were then performed.A consistent trend of change (increased MF and decreased RMS) followed ECG removal was noticed. In particular, in SEMG measurements under static postures, a significant decrease in RMS (p < 0.05) and increase in MF (p < 0.05) were found in all recording muscle groups. Level of corruption by ECG artifacts on SEMG measurements was found to be more serious and prominent in static postures than that in dynamic action. After ECG removal, significant improvements in the ability of SEMG to discriminate LBP patients from healthy subjects were seen in RMS amplitude recorded while standing (p < 0.05) and MF in all measuring conditions (p < 0.05).This study provides a more complete understanding on the relative effect of ECG contamination on back muscles SEMG parameters and LBP assessment.     

Rectification of SEMG as a tool to demonstrate synchronous motor unit activity during vibration

The use of surface electromyography (SEMG) in vibration studies is problematic since motion artifacts occupy the same frequency band with the SEMG signal containing information on synchronous motor unit activity. We hypothesize that using a harsher, 80–500 Hz band-pass filter and using rectification can help eliminate motion artifacts and provide a way to observe synchronous motor unit activity that is phase locked to vibration using SEMG recordings only. Multi Motor Unit (MMU) action potentials using intramuscular electrodes along with SEMG were recorded from the gastrocnemius medialis (GM) of six healthy male volunteers. Data were collected during whole body vibration, using vibration frequencies of 30 Hz, 35 Hz, 40 Hz or 50 Hz. A computer simulation was used to investigate the efficacy of filtering under different scenarios: with or without artifacts and/or motor unit synchronization. Our findings indicate that motor unit synchronization took place during WBV as verified by MMU recordings. A harsh filtering regimen along with rectification proved successful in demonstrating motor unit synchronization in SEMG recordings. Our findings were further supported by the results from the computer simulation, which indicated that filtering and rectification was efficient in discriminating motion artifacts from motor unit synchronization. We suggest that the proposed signal processing technique may provide a new methodology to evaluate the effects of vibration treatments using only SEMG. This is a major advantage, as this non-intrusive method is able to overcome movement artifacts and also indicate the synchronization of underlying motor units.     

Pectoralis muscle morphology in the little brown bat,Myotis lucifugus: A non-convergence with birds

Recent studies of muscle architecture demonstrate that many mammalian muscles are composed of short, interdigitating fibers. In addition, the avian pectoralis, a muscle capable of producing high frequency oscillations has been shown to possess a serially arranged pattern of muscle endplate in all sizes of birds studied. The pectoralis muscle of the little brown bat, Myotis lucifugus (Chiroptera: Vespertilionidae), is composed of fairly uniform fibers that span the length of the muscle and is characterized by a zone of motor endplates within the middle third of the muscle. The homogeneous fiber architecture of the bat pectoralis muscle is in contrast to the serial arrangement of endplates (and presumably muscle muscle fibers) in the avian pectoralis in species equivalent in size to Myotis. The short fiber organization and motor endplate pattern observed in most birds is thus not a requisite design for flying vertebrates. © 1994 Wiley-Liss, Inc.     

Trunk and hip muscle activity in early walkers with and without cerebral palsy – A frequency analysis

Poor control of postural muscles is a primary impairment in cerebral palsy (CP), yet core trunk and hip muscle activity has not been thoroughly investigated. Frequency analysis of electromyographic (EMG) signals provides insight about the intensity and pattern of muscle activation, correlates with functional measures in CP, and is sensitive to change after intervention. The objective of this study was to investigate differences in trunk and hip muscle activation frequency in children with CP compared to children with similar amounts of walking experience and typical development (TD). EMG data from 31 children (15 with CP, 16 with TD) were recorded from 16 trunk and hip muscles bilaterally. A time–frequency pattern was generated using the continuous wavelet transform and instantaneous mean frequency (IMNF) was calculated at each interval of the gait cycle. Functional principal component analysis (PCA) revealed that IMNF was significantly higher in the CP group throughout the gait cycle for all muscles. Additionally, stride-to-stride variability was higher in the CP group. This evidence demonstrated altered patterns of trunk and hip muscle activation in CP, including increased rates of motor unit firing, increased number of recruited motor units, and/or decreased synchrony of motor units. These altered muscle activation patterns likely contribute to muscle fatigue and decreased biomechanical efficiency in children with CP.     

CHOLINESTERASE ACTIVITY OF THE MOTOR ENDPLATE IN ISOLATED MUSCLE MEMBRANE

Abstract— The cholinesterase activity of motor endplates in tibialis anterior muscle of rats accounted for about 20 per cent of the total cholinesterase activity of the muscle. In the isolated muscle membrane preparation of rat intercostal muscle, the cholinesterase activity was localized solely in the motor endplate, as shown by cholinesterase staining. The cholinesterase activity of the membrane per unit of nitrogen was 26·9 times that of the muscle homogenate. The membrane (endplate) cholinesterase had an optimal pH of 8, K_m value of 3·1 m m , and was stable at 4° for at least 13 days. Cholinesterase of a motor endplate hydrolysed 2·69 × 10⁸ acetylcholine molecules in 1 msec. Since it is estimated that 10⁸ cholinesterase active sites are present in a motor endplate, the turnover time (time necessary for one enzyme site to hydrolyse one acetylcholine molecule) is calculated to be 372 μ sec, and the turnover number (molecules of acetylcholine hydrolysed by one enzyme site/min) to be 1·61 × 10⁵. From studies with cholinesterase inhibitors, cholinesterase activity was estimated to be due mostly to acetylcholinesterase, and only a minor part to pseudocholinesterase. The muscle membrane preparation seems to be useful for the study of other properties of the motor endplate.     

Influence of motor unit firing statistics on the median frequency of the EMG power spectrum

Changes in the EMG power spectrum during static fatiguing contractions are often attributed to changes in muscle fibre action potential conduction velocity. Mathematical models of the EMG power spectrum, which have been empirically confirmed, predict that under certain conditions a distinct maximum occurs in the low-frequency part of the spectrum, indicating the dominant firing rate of the motor units. The present study investigated the influence of this firing rate peak on the spectral changes during a static fatiguing contraction at 50% of maximum EMG amplitude in the frontalis and corrugator supercilii muscles. An exponential decrease of the median frequency (MF) of the EMG power spectrum was observed when the firing rate peak was absent. When the firing rate peak was present, an exaggerated decrease of MF in the beginning of the contraction was found, which was associated with an increase in firing rate peak magnitude. In later stages of the contraction, a partial recovery of MF occurred, concomitant with a decrease in firing rate peak magnitude. The influence of the firing rate peak on MF was also investigated during nonfatiguing contractions of the frontalis muscle at 20, 40, 60, and 80% of maximum EMG amplitude. A curvilinear relationship between MF and contraction strength was found, whether firing rate peaks were present or absent. The presence of firing rate peaks, however, was associated with a decrease in MF which was inversely related to contraction strength, due to the inverse relationship between firing rate peak magnitude and contraction strength.     

Computer simulation of the motoneuron pool–muscle complex. I. Input system and motoneuron pool

 The aim of the present study was to simulate the input system and the motoneuron (MN) pool of the MN pool–muscle complex (MNPMC). Input fibers, which can originate from command centers in the central nervous system or from sensory organs, activate the MN pool. They generate sequences of action potentials, the frequency of which is proportional to a time-dependent activation factor (which is an input to the model). Different connection patterns between the input fibers and motor units (MUs) are allowed. For simplicity and since no precise experimental data are available, 70 input fibers and 4 boutons per fiber and MN are simulated (this corresponds approximately to the monosynaptic group-Ia input of the cat medial gastrocnemius muscle). Each bouton generates the same conductance change in the postsynaptic membrane. The MNs are modeled with a single compartment and a homogenous membrane. According to experimental data, the membrane leakage conductance and capacitance are MU dependent. Since the precise relation is unknown: (a) the computed relation between MU contraction force and the MN leakage conductance was taken from a steady-state MNPMC model, and (b) the capacitance was assumed to be proportional to the leakage conductance. The MN membrane includes time- and voltage-dependent ionic channels (fast and slow K⁺ and low- and high-threshold Ca²⁺ channels). The density and time constant of the slow K⁺ channels and the density of the Ca²⁺ channels were fitted to approximate afterhyperpolarization characteristics and frequency-injected current relations of type-identified cat MNs. If the membrane reaches a voltage threshold the MNs generate action potentials, which were simulated by voltage pulses. The activation of the MN pool of the human first dorsal interosseus muscle was simulated with injected and synaptic currents in order to illustrate the size principle, synaptic noise, and other features of muscle activation. It is concluded that the present model reproduces the main properties of the input–output relations of different MN types within a muscle. Together with the simulation of the muscle force and the surface EMG, which will be published in subsequent papers, it will be a powerful tool for reproducing experiments on the motor system and investigating functional mechanisms of motor control. Received: 17 April 2001 / Accepted in revised form: 6 November 2001     

The establishment and the subsequent elimination of polyneuronal innervation of developing muscle: theoretical considerations

An analysis is given of the polyneuronal innervation of embryonic skeletal muscle and its subsequent elimination during development. The amount of polyneuronal innervation that has been observed is consistent with the notion that initially each motor neuron distributes its contacts at random among the available fibres of a particular muscle. The idea that the elimination of excess innervation proceeds through interactions between terminals is placed on quantitative basis. Each motor neuron is presumed to have a finite capacity for maintaining the structure and activity of its terminals, which is shared out among them; a survival strength can be assigned to each terminal. Survival strengths undergo a process of continual adjustment. A terminal with above average strength for its endplate is strengthened at the expense of the weaker terminals, subject to the total survival strength available to each motor neuron remaining constant. It is proved that this scheme will transform the initial pattern of innervation into one in which each muscle fibre has contact from a single axon. Interpretations of the following results are given: the decrease in the spread of motor unit size during the development of innervation of the rat soleus muscle; the time course of superinnervation; the effects of neonatal partial denervation. Various suggestions are made for future experimental approaches.     

Relation of mechanical noise in the rat papillary muscle to the level of its contraction

The existence of mechanical noise (MN) has been demonstrated in isolated papillary muscles of rats at rest. The mean amplitude of the MN was about 1 mg, the mean frequency 1.5 Hz (t 22 degrees C). A good agreement was found between the MN amplitude and the contracture level of the muscle. However, during long contractures, the correlation between the noise and contracture magnitude was disturbed. There was no relationship between the MN amplitude and contracture magnitude during exposures inducing metabolic alterations (hypoxia, NaCN) and upsetting the work of the sarcoplasmic reticulum (caffeine). It is believed that the MN amplitude is in a good agreement with the contracture magnitude and, therefore, with the concentration of intracellular Ca2+, if the sarcoplasmic reticulum and contractile elements of the cells are intact.     

Limitations of the surface electromyography technique for estimating motor unit synchronization

Motor unit synchronization was estimated from the surface electromyograms (EMG) of the first dorsal interosseus muscle of human volunteers by a simplified surface-EMG technique (Milner-Brown et al. 1973, 1975). Single motor units were identified from intramuscular recordings and were used to obtain a spike-triggered average of the surface-EMG. The discharge rate of a reference motor unit was controlled at two levels (high and low), and the effect of motor unit activity on the surface-EMG estimate of synchronization was studied in 56 motor units. The surface-EMG estimate of motor unit synchronization was significantly higher when the reference motor unit discharged at the high rate than when it discharged at the low rate. A regression analysis indicated that the synchronization ratio calculated from the surface EMG was significantly correlated with the level of EMG activity in the muscle. Motor unit synchronization was also estimated from surface-EMG measurements that were derived by computer simulation. The simulation permitted manipulation of motor unit activity (discharge rate and recruitment) with a complete absence of synchrony among the units in the pool. The stimulated surface-EMG index was influenced by an artifact associated with signal rectification, and this effect changed non-monotonically with motor unit activity. Furthermore, the increase in the motor unit activity reduced the signal-to-noise ratio of the spike-triggered surface EMG average, and consequently decreased the sensitivity of the surface-EMG index as an estimate of motor unit synchronization. We conclude that the simplified surface-EMG method (Milner-Brown et al. 1973, 1975) does not provide a useful index of motor unit synchronization due to its inability to accurately distinguish the synchronization from methodological effects related to a rectification artifact and variation in the signal-to-noise ratio.     

Characterization of Levels of Extremely Low Frequency Magnetic Fields Emitted From Portable Hand‐Held Fans

This study aims to assess the levels of extremely low frequency magnetic fields (ELF‐MF) emitted from portable hand‐held fans (HHFs) and their principal frequency and to identify factors influencing these levels. We collected a total of eleven models of HHF and monitored the ELF‐MF as a function of fan speed and distance from the fan. EMDEX II was used to monitor the ELF‐MF. An SMP2 EMF‐meter equipped with a P400 field probe was used to determine the levels of ELF‐MF and the frequency spectrum. Ten of the fans, excluding only one bladeless‐fan model, emitted a high level of ELF‐MF near the source of the HHF direct‐current motor. The maximum measured level of ELF‐MF ranged from 14.07 to 218.7 µT. All measurements of the ELF‐MF taken within 10 cm from the HHFs showed values higher than 1.0 µT. ELF‐MF levels were found to decrease markedly with distance, regardless of the HHF product. The level of ELF‐MF rose noticeably with increased fan speed. The speed of and distance from the HHF significantly influenced the level of ELF‐MF. All principal frequencies ranged from 1 to 300 Hz, which falls in the typical range of ELF. Bioelectromagnetics. 2019;40:569–577. © 2019 Bioelectromagnetics Society.     

Recurrent inhibition of human firing motoneurons (experimental and modeling study)

Recurrent inhibition between tonically activated single human motoneurons was studied experimentally and by means of a computer simulation. Motor unit activity was recorded during weak isometric constant-force muscle contractions of brachial biceps (BB) and soleus (SOL) muscles. Three techniques (cross correlogram, frequencygram, and interspike interval analysis) were used to gauge the relations between single motor unit potential trains. Pure inhibition was detected in 5.6% of 54 BB motoneuron pairs and in 5.2% of 43 SOL motoneuron pairs. In 27.8% (BB) and 23.7% (SOL) presumed inhibition symptoms were accompanied by a synchrony peak; 37% (BB) and 48.8% (SOL) exhibited synchrony alone. The demonstrated inhibition was very weak, at the edge of detectability. Computer simulations were based on the threshold-crossing model of a tonically firing motoneuron. The model included synaptic noise as well as threshold and postsynaptic potential (PSP) amplitude change within interspike interval. Inhibition efficiency of the model neurons increased with IPSP amplitude and duration, and with increasing source firing rate. The efficiency depended on target motoneuron interspike interval in a manner similar to standard deviation of ISI. The minimum detectable amplitude estimated in the simulations was about 50V, which, compared with the experimental results, suggests that amplitudes of detectable recurrent IPSPs in human motoneurons during weak muscle contractions do not exceed this magnitude. Since recurrent inhibition is known to be progressively depressed with an increase in the force of voluntary contraction, it is concluded that the recurrent inhibition hardly plays any important role in the isometric muscle contractions of constant force.     

Short conduction delays cause inhibition rather than excitation to favor synchrony in hybrid neuronal networks of the entorhinal cortex

How stable synchrony in neuronal networks is sustained in the presence of conduction delays is an open question. The Dynamic Clamp was used to measure phase resetting curves (PRCs) for entorhinal cortical cells, and then to construct networks of two such neurons. PRCs were in general Type I (all advances or all delays) or weakly type II with a small region at early phases with the opposite type of resetting. We used previously developed theoretical methods based on PRCs under the assumption of pulsatile coupling to predict the delays that synchronize these hybrid circuits. For excitatory coupling, synchrony was predicted and observed only with no delay and for delays greater than half a network period that cause each neuron to receive an input late in its firing cycle and almost immediately fire an action potential. Synchronization for these long delays was surprisingly tight and robust to the noise and heterogeneity inherent in a biological system. In contrast to excitatory coupling, inhibitory coupling led to antiphase for no delay, very short delays and delays close to a network period, but to near-synchrony for a wide range of relatively short delays. PRC-based methods show that conduction delays can stabilize synchrony in several ways, including neutralizing a discontinuity introduced by strong inhibition, favoring synchrony in the case of noisy bistability, and avoiding an initial destabilizing region of a weakly type II PRC. PRCs can identify optimal conduction delays favoring synchronization at a given frequency, and also predict robustness to noise and heterogeneity.     

SEMG power spectrum changes during a sustained 50% Maximum Voluntary Isometric Torque do not depend upon the prior knowledge of the exercise duration.

Endurance time (Tlim) is a relevant indicator of muscular resistance to fatigue. It has been recently shown that SEMG changes computed during shorter periods (sub-maximal durations) than the whole test duration could serve to predict Tlim. The aim of the present study was to test whether the prior knowledge of the sub-maximal duration had any influence on Tlim prediction throughout SEMG changes. For this purpose, we compared myoelectric changes estimated over a 30-s isometric contraction whose duration was known by the subjects, to those changes measured during the first 30 s of a test prolonged until exhaustion. The effort intensity was set at 50% of the maximal voluntary isometric torque (MVIT). The myoelectric manifestations of muscle fatigue appeared to change in a similar way during both sessions for temporal and spectral analyses. In conclusion, the prior knowledge of the duration of sustained isometric contraction did not influence the motor unit recruitment strategy throughout surface EMG evaluation. These results confirmed that the use of SEMG changes computed over shorter periods than expected endurance time may be applied to investigate the capability of the initial rate of SEMG changes to predict muscle endurance capacity at 50%MVIT. This may be of particular interest for patient evaluation in the clinical field.     

Does the frequency content of the surface mechanomyographic signal reflect motor unit firing rates? A brief review.

The purpose of this review is to examine the literature that has investigated the potential relationship between mechanomyographic (MMG) frequency and motor unit firing rates. Several different experimental designs/methodologies have been used to address this issue, including: repetitive electrical stimulation, voluntary muscle actions in muscles with different fiber type compositions, fatiguing and non-fatiguing isometric or dynamic muscle actions, and voluntary muscle actions in young versus elderly subjects and healthy individuals versus subjects with a neuromuscular disease(s). Generally speaking, the results from these investigations have suggested that MMG frequency is related to the rate of motor unit activation and the contractile properties (contraction and relaxation times) of the muscle fibers. Other studies, however, have reported that MMG mean power frequency (MPF) does not always follow the expected pattern of firing rate modulation (e.g. motor unit firing rates generally increase with torque during isometric muscle actions, but MMG MPF may remain stable or even decrease). In addition, there are several factors that may affect the frequency content of the MMG signal during a voluntary muscle action (i.e. muscle stiffness, intramuscular fluid pressure, etc.), independent of changes in motor unit firing rates. Despite the potential influences of these factors, most of the evidence has suggested that the frequency domain of the MMG signal contains some information regarding motor unit firing rates. It is likely, however, that this information is qualitative, rather than quantitative in nature, and reflects the global motor unit firing rate, rather than the firing rates of a particular group of motor units.     

Organized variability in the neuromuscular system: a survey of task-related adaptations.

This survey concerns the physiology of the neuromuscular system, as studied at the level of the single mammalian limb muscle and its motoneurones (MNs). Particular attention is devoted to the ways in which the properties and the organization of spinal MNs are adapted for the control of muscle (unit) force. These questions are discussed in relation to: a) The general and basic task of the system: providing a smooth and finely gradeable force by the mechanisms of rate- and recruitment-modulation of MN activity. b) Gradation problems in relation to specific peripheral requirements in connection with: (i) tasks of different duty-time (i.e. problems related to fatigue and endurance); (ii) tasks of different speed; (iii) task using different muscle lengths; (iv) tasks requiring different adjustment-gains. c) The adaptational properties of the neuromuscular system as it is subjected to long-term changes in its motor tasks. These matters are largely discussed in relation to experiments for studying the responses of the neuromuscular system to different patterns of chronic electrical stimulation. d) The manner in which the neuromuscular system, at the level of a single unidirectional muscle, is used for different motor programs. Evidence is summarized which shows the presence of task-related variations in MN recruitment patterns, and it is pointed out that such variations may be related to the intraspinal topography of the respective MNs. It is suggested that these task-related variations in MN recruitment behaviour might largely reflect topographic (and other) differences in the organization of spinal interneuronal systems responsible for the execution of different motor programs.     

Inference of motor unit recruitment order in voluntary and electrically elicited contractions

The relationship between surface myoelectric signal parameters and the level of voluntary or electrically elicited contractions was studied in 32 experiments on the tibialis anterior muscle of 22 healthy human subjects. Contractions were performed at 20 and 80% of the maximum voluntary contraction torque. Two levels of stimulation current were used, yielding, respectively, a maximum M wave and an M wave approximately 30% of the maximum. A four-bar electrode probe was used to detect single- and double-differential signals from which mean and median frequency of the power spectrum and average muscle fiber conduction velocity were estimated. Measurements obtained from voluntary contractions showed a positive correlation between contraction levels and both conduction velocity and spectral parameters. Conduction velocity increased by 21.2 +/- 10.9% when voluntary contraction level increased from 20 to 80% of the maximal value. Spectral parameters increased by similar amounts. Tetanic electrical stimulation was applied to a muscle motor point for 20 s via surface electrodes. Rectangular current pulses with 0.1-ms width and frequencies of 20, 25, 30, 35, and 40 Hz were used. Four types of behavior were observed with increasing stimulation level: 1) the two spectral parameters and conduction velocity both increased with stimulation in 15 experiments, 2) the two spectral parameters decreased and conduction velocity increased in 8 experiments, 3) the two spectral parameters and conduction velocity both decreased in 6 experiments, and 4) the two spectral parameters increased and conduction velocity decreased in 3 experiments. Conduction velocity increased with increasing stimulation current in 72% of the experiments, indicating a recruitment order similar to that of voluntary contractions, although it decreased in the other 28% of the cases, indicating a reverse order of recruitment. Contrary to what is observed in direct stimulation of nerves, motor units are not in general recruited in reverse order of size during electrical stimulation of a muscle motor point. This discrepancy may be the result of geometric factors or a lack of correlation between axonal branch diameter and the diameter of the parent motoneuron axon. Changes of conduction velocity and spectral parameters in opposite directions may be the result of the combined effect of the motor unit recruitment order and of the different tissue filtering function associated with the geometric location of the recruited motor units within the muscle.