The cross-correlation and phase-difference methods are not equivalent under noninvasive estimation of the motor unit propagation velocity.

Noninvasive estimation of motor unit propagation velocity (MUPV) was reduced to that of the time delay between signals detected by two surface EMG electrodes placed along the muscle fibres. When the cross-correlation function between the signals was used, the problem with temporal resolution arose. Estimation of the time delay in the frequency domain was proposed to overcome this problem. To check whether the cross-correlation and phase-difference methods give the same estimates, the results obtained by both methods were compared through simulation. A different sensitivity of the two methods to the effects of the excitation origin and extinction was found. Besides, the quality of the estimate depended on the electrode arrangement. The longitudinal double difference electrodes were preferable with the phase-difference method, while the MUPV estimates obtained by the cross-correlation technique were more correct when the longitudinal single difference or bipolar transversal double difference electrodes were used. In addition, the estimates obtained by the phase-difference method were more sensitive to the longitudinal scattering of motor end-plates and ends of the fibres, to the fibre lengths and to the negative after-potential magnitude. Such sensitivity could make MUPV estimates incorrect even under a relatively small distance between the motor unit axis and electrode.     

Estimation of monopolar signals from sphincter muscles and removal of common mode interference

The detection of surface electromyogram (EMG) by multi-electrode systems is applied in many research studies. The signal is usually recorded by means of spatial filters (linear combination of the potential under at least two electrodes) with vanishing sum of weights. Nevertheless, more information could be extracted from monopolar signals measured with respect to a reference electrode away from the muscle. Under certain conditions, surface EMG signal along a curve parallel to the fibre path has zero mean (property approximately satisfied when EMG is sampled by an array of electrodes that covers the entire support of the signal in space). This property allows estimating monopolar from single differential (SD) signals by pseudoinversion of the matrix relating monopolar to SD signals. The method applies to EMG signals from the external anal sphincter muscle, recorded using a specific cylindrical probe with an array of electrodes located along the circular path of the fibres. The performance of the algorithm for the estimation of monopolar from SD signals is tested on simulated signals. The estimation error of monopolar signals decreases by increasing the number of channels. Using at least 12 electrodes, the estimation error is negligible. The method applies to single fibre action potentials, single motor unit action potentials, and interference signals.The same method can also be applied to reduce common mode interference from SD signals from muscles with rectilinear fibres. In this case, the last SD channel defined as the difference between the potentials of the last and the first electrodes must be recorded, so that the sum of all the SD signals vanishes. The SD signals estimated from the double differential signals by pseudoinvertion are free of common mode.     

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.     

The linear electrode array: a useful tool with many applications.

In this review we describe the basic principles of operation of linear electrode arrays for the detection of surface EMG signals, together with their most relevant current applications. A linear array of electrodes is a system which detects surface EMG signals in a number of points located along a line. A spatial filter is usually placed in each point for signal detection, so that the recording of EMG signals with linear arrays corresponds to the sampling in one spatial direction of a spatially filtered version of the potential distribution over the skin. Linear arrays provide indications on motor unit (MU) anatomical properties, such as the locations of the innervation zones and tendons, and the fiber length. Such systems allow the investigation of the properties of the volume conductor and its effect on surface detected signals. Moreover, linear arrays allow to estimate muscle fiber conduction velocity with a very low standard deviation of estimation (of the order of 0.1-0.2 m/s), thus providing reliable indications on muscle fiber membrane properties and their changes in time (for example with fatigue or during treatment). Conduction velocity can be estimated from a signal epoch (global estimate) or at the single MU level. In the latter case, MU action potentials are identified from the interference EMG signals and conduction velocity is estimated for each detected potential. In this way it is possible, in certain conditions, to investigate single MU control and conduction properties with a completely non-invasive approach. Linear arrays provide valuable information on the neuromuscular system properties and appear to be promising tools for applied studies and clinical research.     

Spatial distribution of active muscle fibre characteristics in the upper trapezius muscle and its dependency on contraction level and duration

The aim of this study was to provide direct in vivo information of the physiological and structural characteristics of active muscle fibres from a large part of the upper trapezius muscle. Two-dimensional (2-D) multi-channel surface electromyography recordings were used, with 13 × 10 electrodes covering 6 × 4.5 cm of the skin’s surface. A previously developed method was applied to detect individual propagating motor unit action potentials and to estimate their corresponding muscle fibre conduction velocity (MFCV) and muscle fibre orientation (MFO). Using these estimates, spatial distributions of MFCV and MFO were examined for five male subjects performing isometric shoulder elevation at different force levels. The main results were: (1) the general relationship between MFCV and force generation was non-systematic, with a positive relationship at the inferior part of the muscle, (2) the spatial distribution of MFCV at different force levels and fatigue was inhomogeneous and (3) the MFO was slightly different (6°) of the muscle fibres with origin superior compared to inferior to the C7 vertebra. These findings provide new information of the MFO of contracting muscle fibres and knowledge of the physiological characteristics of a large part of the upper trapezius muscle that previously was based on observations from human cadavers only.     

Detection of motor unit action potentials with surface electrodes: influence of electrode size and spacing

A model of the motor unit action potential was developed to investigate the amplitude and frequency spectrum contributions of motor units, located at various depths within muscle, to the surface detected electromyographic (EMG) signal. A dipole representation of the transmembrane current in a three-dimensional muscle volume was used to estimate detected individual muscle fiber action potentials. The effects of anisotropic muscle conductance, innervation zone location, propagation velocity, fiber length, electrode area, and electrode configuration were included in the fiber action potential model. A motor unit action potential was assumed to be the sum of the individual muscle fiber action potentials. A computational procedure, based on the notion of isopotential layers, was developed which substantially reduced the calculation time required to estimate motor unit action potentials. The simulations indicated that: 1) only those motor units with muscle fibers located within 10–12 mm of the electrodes would contribute significant signal energy to the surface EMG, 2) variation in surface area of electrodes has little effect on the detection depth of motor unit action potentials, 3) increased interelectrode spacing moderately increases detection depth, and 4) the frequency content of action potentials decreases steeply with increased electrode-motor unit territory distance.     

A study of the motor unit action potential by means of computer simulation

In order to study the motor unit action potential a computer simulation model was developed. It is based on the superposition of single muscle fibre potentials of the fibres belonging to the motor unit. The parameters which characterize each fibre (spatial position, diameter, and a dispersion of arrival time of the potential at the electrode) are chosen from statistical distributions which can be derived from anatomical and physiological data. The electrode type, position and dimensions can be specified. Simulated motor unit action potentials are presented in the time and frequency domain. The simulation results refer to (1) the influence of the electrode position and dimensions with respect to the motor unit territory, (2) the meaning of this model for the study of pathological phenomena, (3) the variability of some parameters characterizing the motor unit, (4) the selectivity of uni- and bipolar electrodes and finally (5) the influence of the geometrical situation of the motor end-plates within the muscle, on the shape of motor unit action potentials.     

Fibre conduction velocity and fibre composition in human vastus lateralis

The relationship between muscle fibre composition and fibre conduction velocity was investigated in 19 male track athletes, 12 sprinters and 7 distance runners, aged 20-24 years, using needle biopsy samples from vastus lateralis. Cross sectional areas of the fast twitch (FT) and slow twitch (ST) fibres were determined by histochemical analysis. The percentage of FT fibre areas ranged from 22.6 to 93.6%. Sprinters had a higher percentage of FT fibres than distance runners. Muscle fibre conduction velocity was measured with a surface electrode array placed along the muscle fibres, and calculated from the time delay between 2 myoelectric signals recorded during a maximal voluntary contraction. The conduction velocity ranged from 4.13 to 5.20 m.s-1. A linear correlation between conduction velocity and the relative area of FT fibres was statistically significant (r = 0.84, p less than 0.01). This correlation indicates that muscle fibre composition can be estimated from muscle fibre conduction velocity measured noninvasively with surface electrodes.     

Separation of propagating and non propagating components in surface EMG

Surface electromyogram (EMG) detected by electrode arrays along the muscle fibre direction can be approximated by the sum of propagating and non propagating components. A technique to separate propagating and non propagating components in surface EMG signals is developed. The first step is an adaptive filter, which allows obtaining an estimation of the delay between signals detected at different channels and a first estimate of propagating and non propagating components; the second step is used to optimise the estimation of the two components. The method is applicable to signals with one propagating and one non propagating component. It was optimised on simulated signals, and then applied to single motor unit action potentials (MUAP) and to electrically elicited EMG (M-waves).

The new method was first tested on phenomenological signals constituted by the sum of a propagating and a non propagating signal and then applied to simulated and experimental EMG signals. Simulated signals were generated by a cylindrical, layered volume conductor model. Experimental signals were monopolar surface EMG signals collected from the abductor pollicis brevis muscle and M-waves recorded during transcutaneous electrical stimulation of the biceps muscle. The technique may find different applications: in single motor unit (MU) studies (a) for decreasing the variability and bias of CV estimates due to the presence of the non propagating components, (b) for estimating automatically the length of the muscle fibres from only three detected channels and (c) for removal of the stimulation artifact M-waves.     

Probabilistic model of the spatial distribution of muscle fibres in human muscles

An analytical model and computer simulation model for measuring fibre density in motor units of human skeletal muscles have been described. The model was developed for Gaussian distribution of the fibres in the motor unit territory. It has been shown that fibre density measurement using a triggering fibre was a biased estimate of the actual density of the fibres in the territory. The effects of varying the standard deviation of the spatial distribution on the estimate of fibre density has been investigated, and it has been shown that for high values of standard deviation a uniform distribution of the fibres in the territory was a good first order approximation.     

Neuromuscular organization of avian flight muscle: architecture of single muscle fibres in muscle units of the pectoralis (pars thoracicus) of pigeon (Columba livia)

The M. pectoralis (pars thoracicus) of pigeons (Columba livia) is comprised of short muscle fibres that do not extend from muscle origin to insertion but overlap ''in-series''. Individual pectoralis motor units are limited in territory to a portion of muscle length and are comprised of either fast twitch, oxidative and glycolytic fibres (FOG) or fast twitch and glycolytic fibres (FG). FOG fibres make up 88 to 90% of the total muscle population and have a mean diameter one-half of that of the relatively large FG fibres. Here we report on the organization of individual fibres identified in six muscle units depleted of glycogen, three comprised of FOG fibres and three comprised of FG fibres. For each motor unit, fibre counts revealed unequal numbers of depleted fibres in different unit cross-sections. We traced individual fibres in one unit comprised of FOG fibres and a second comprised of FG fibres. Six fibres from a FOG unit (total length 15.45 mm) ranged from 10.11 to 11.82 mm in length and averaged (± s.d.) 10.74 ± 0.79 mm. All originated bluntly (en mass) from a fascicle near the proximal end of the muscle unit and all terminated intramuscularly. Five of these ended in a taper and one ended bluntly. Fibres coursed on average for 70% of the muscle unit length. Six fibres from a FG unit (total length 34.76 mm) ranged from 8.97 to 18.38 mm in length and averaged 15.32 ± 3.75 mm. All originated bluntly and terminated intramuscularly; one of these ended in a taper and five ended bluntly. Fibres coursed on average for 44% of the muscle unit length. Because fibres of individual muscle units do not extend the whole muscle unit territory, the effective cross-sectional area changes along the motor unit length. These non-uniformities in the distribution of fibres within a muscle unit emphasize that the functional interactions within and between motor units are complex.     

Organization of skeletal muscle in the urodele Triturus cristatus: muscle fibre types and motor units

Four muscle fibre types are described in the biceps and extensor digitorum communis muscles of the newt forelimb. The histological criteria forming the basis for the distinctions include differential staining with p-phenylenediamine and succinate dehydrogenase histochemistry and electron microscopy. In addition, three distinctive motor unit types are described for the biceps muscle. These are fast units, slow units and intermediate units. The structure of muscle fibre and the physiological characteristics of muscle fibres belonging to each motor unit, have been correlated by using iontophoretic passage of Lucifer yellow into muscle fibres belonging to physiologically characterized motor units and their subsequent histological identification by the succinate dehydrogenase reaction. The three motor unit types correspond to slow muscle fibres, intermediate muscle fibres and two classes of fast muscle fibres.     

Simulation analysis of the ability of different types of multi-electrodes to increase selectivity of detection and to reduce cross-talk.

Selectivity of different one- and two-dimensional multi-electrodes and their ability to reduce cross-talk were analyzed. Signals from an individual motor unit (MU) were calculated as a single convolution of intracellular action potential (IAP) first temporal derivative and spatially filtered MU impulse response. It was shown that the uptake area (irrespective of the way it was defined) could not characterize electrode properties reliably because its estimate depended on the source parameters. Due to the different decline of individual phases of MU signals with depth, electrode should provide higher spatial and temporal resolution of the main phases for better selectivity and greater suppression of the terminal phases for cross-talk reduction. A two-dimensional normal double differentiating (NDD) electrode provided almost the same or slightly lower selectivity but weaker reduction of cross-talk than a longitudinal double differentiating (LDD) electrode. A transversal double differentiating (TDD) electrode provided a lower selectivity and weaker reduction of cross-talk than a LDD electrode. A new, BiTDD multi-electrode (performing difference between signals detected by two TDD electrodes) provided the best selectivity and reduction of cross-talk. To obtain the smallest cross-talk, a BiTDD electrode should be positioned above the end-plate region, while LDD, TDD, or NDD electrodes-above the ends of muscle that produced it. Signal differentiation improved selectivity but increased cross-talk.     

Local development of action potentials in slow muscle fibres after complete or partial denervation.

Pyriformis muscles of Rana temporaria were completely or partially denervated by cutting the sciatic nerve or some of the small nerve branches entering the muscle. One stimulating and one to three recording microelectrodes were inserted along the fibres in order to compare the electrical activity at these points. In an early period following denervation action potentials of variable size and shape could be observed; these action potentials were often composed of two, sometimes of three or four, components. The size of individual components depended on the position of the recording microelectrode. Individual components could occasionally be triggered separately by adjusting the strength of the stimulating current pulse; propagation of these "all or none" responses was absent. In other fibres one component of the action potential could trigger another one several millimetres apart, thus indicating propagation. Conduction velocities were approximately 0.4 m/s. In partially denervated slow fibres, endplate potentials were confined to one lateral segment of the fibres, while the action potential occupied the denervated part of the membrane. The amplitudes of endplate and action potentials varied inversely with distance. Rough estimates of the length constant of the slow fibre membrane were calculated from the spatial decay of action potentials, endplate potentials and hyperpolarizing electrotonic potentials; mean values obtained were 2.5, 4.8 and 7.7 mm respectively. The results suggest that following denervation Na channels are built into discrete areas of the slow fibre membrane and that this process depends on the amount of denervation in individual fibres.     

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.     

Estimation of muscle fiber conduction velocity of doublet discharges

A doublet is defined as two consecutive discharges of a motor unit occurring at short time interval between each other (e.g., <20 ms). In this paper, we propose a method for the estimation of muscle fiber conduction velocity (CV) from two partly overlapping action potentials generated by the same motor units. The method is based on the minimization of the mean square error between time-filtered versions of two surface EMG signals recorded along the direction of muscle fibers. The minimization is performed over the filter parameters that define the two propagation delays. The method was tested on simulated and experimental signals. Simulation results showed that the method is only in some cases superior to the simpler peak approach, due to limitations in the ideal model used for the algorithm development. However, application to experimental signals that mimic doublet motor unit discharges showed a substantial improvement in estimation quality of the new method with respect to the peak method.     

Extracellular recording of localized electrical activity in denervated frog slow muscle fibres

Pyriformis muscles of Rana temporaria were denervated by cutting the sciatic nerve in the pelvis. Slow muscle fibres were depolarized with intracellular current pulses, and the electrical activity was recorded simultaneously with intracellular and extracellular recording electrodes. When the extracellular electrode was moved along the fibre surface, outward and inward currents of variable amplitude were recorded. Inward currents coincided with the fast rising phase of the intracellularly recorded action potential; up to four inward current peaks could be detected in single fibres investigated over 3.4--8 mm of their length. The distance between inward current peaks was generally 1--2 mm, but greater distances were also observed. Composite action potentials could be shown to be due to inward currents arising in separate areas of the slow fibre membrane. It is concluded that after denervation Na-channels are incorporated into spatially limited areas of the membrane of slow muscle fibres.     

Recent progress in the diagnostic use of surface EMG for neurological diseases.

The different techniques to measure and analyze surface EMG are summarized with an emphasis on the clinician's point of view. The application of surface EMG in neurological disease is hampered by many inherent problems, especially the difficulties in extracting features of single motor units. However, the evolution of surface EMG from single bipolar recordings via a linear array of multiple electrodes to densely packed, multi-channel electrode arrays could in principle solve this problem. The added value of using multiple channels (up to 128) with an interelectrode distance of a few millimetres to obtain more spatial information is emphasized. At least for some muscles it is now possible to extract information from the surface EMG, conventionally thought to belong to the domain of needle EMG (for example the "electrical size" of motor units). The use of analysis techniques such as the estimation of muscle fiber conduction velocity has already proven to be of diagnostic value in several myopathies characterized by a disturbed membrane function and in metabolic myopathies with abnormal fatigue profiles. Future research should be directed at the development of analysis techniques enabling the extraction of more relevant motor unit variables from surface EMG signals.     

Spectral and time domain characteristics of single muscle fibre action potentials during continuous activity extracted from model considerations

A model of the muscle fibre extracellular action potentials (ECAPs) calculation using experimentally recorded intracellular action potentials (ICAPs) has been applied to investigate the effect of repetitive stimulation on the electrical activity of isolated frog muscle fibres. The ECAPs were calculated both at small (0.01 mm) and at large (5 mm) radial distances to the fibre axis, and their relationship with the original ICAP parameters has been inferred. Fourier transformation of the calculated ECAPs in order to obtain the spectral characteristics and to trace out their behaviour during continuous fibre activity was performed. Stimulation frequency dependence on the ECAP time characteristics and on the shift of the maximum spectral density towards low frequencies at small and large radial distance were observed. The spectral density peak frequency is propagation velocity (PV)-dependent. The advantage of the presented method over the available experimental extracellular recording techniques from isolated muscle fibers is the possibility to show the effect of continuous muscle fibre activity on the parameters of the ECAPs and their spectral characteristics at large radial distance, which is not experimentally accessible. Our results are in agreement with those experimentally obtained. The results from the model prove the role of changes in PV of excitation along the muscle fibres (representing the last link in the complex organized motor system) in the development of fatigue. Received: 24 July 1997 / Accepted in revised form: 2 July 1998     

Reliability of muscle fibre conduction velocity distribution estimation from surface EMG

Most of the neuromuscular diseases induce changes in muscle fibre characteristics. For example, Duchenne dystrophy is characterized by a specific loss of fast fibres, and an increase in small diameter fibres. These morphological changes may lead to large modifications in the distribution of fibre diameters, possibly producing bimodal distributions. It has already been shown that it is possible to reveal these morphological modifications through the distribution of muscle fibre conduction velocity (MFCV) as estimated from needle electromyography (EMG) recordings. In this paper, we investigate whether such changes can be extracted from surface EMG signals.

Simulation allows generation of surface EMG signals in which features are well described especially at a morphological level. Therefore, we generated a database of simulated signals both in voluntary and electrically elicited contraction conditions using a bimodal distribution of muscle fibre diameters. MFCV distributions were computed using two short-term methods based on cross-correlation and peak-to-peak techniques for voluntary contraction signals, and using a deconvolution method in time domain for electrically elicited signals. MFCV distributions were compared with true ones, as generated from modelling.

This work reveals that estimating MFCV distribution through these methods does not appear yet as precise and robust enough to accurately characterize changes in redistribution of various muscle fibre diameters. However, it opens to new experimental protocols that can be explored in order to improve the robustness of MFCV distribution estimation for the follow-up of patients suffering from neuromuscular disorders.