Does the activity of ankle plantar flexors differ between limbs while healthy,young subjects stand at ease?

Inferences on the active contribution of plantar flexors to the stabilisation of human standing posture have been drawn from surface electromyograms (EMGs). Surface EMGs were however often detected unilaterally, presuming the myoelectric activity from muscles in a single leg reflects the pattern of muscle activation in both legs. In this study we question whether surface EMGs detected from plantar flexor muscles in both legs provide equal estimates of the duration of activity. Arrays of surface electrodes were used to collect EMGs from gastrocnemius and soleus muscles while twelve, young male participants stood at ease for 60 s. Muscles in each leg were deemed active whenever the Root Mean Square amplitude of EMGs (40 ms epochs) detected by any channel in the arrays exceeded the noise level, defined from EMGs detected during rest. The Chi-Square statistics revealed significant differences in the relative number of active periods for both muscles in 10 out of 12 participants tested, ranging from 2% to 65% (χ² > 17.90; P < 0.01). Pearson correlation analysis indicated side differences in the duration of gastrocnemius though not soleus activity were associated with the centre of pressure mean, lateral position (R = 0.60; P = 0.035). These results suggest therefore that surface EMGs may provide different estimates of the timing of plantar flexors’ activity if collected unilaterally during standing and that asymmetric activation may be not necessarily associated with weight distribution between limbs. Depending on the body side from which EMGs are collected, the active contribution of plantar flexors to standing stabilization may be either under- or over-valued.     

Electrical and mechanical activity of respiratory muscles during hypercapnia

In nine anesthetized supine spontaneously breathing dogs, we compared moving average electromyograms (EMGs) of the costal diaphragm and the third parasternal intercostal muscles with their respective respiratory changes in length (measured by sonomicrometry). During resting O2 breathing the pattern of diaphragm and intercostal muscle inspiratory shortening paralleled the gradually incrementing pattern of their moving average EMGs. Progressive hypercapnia caused progressive increases in the amount and velocity of respiratory muscle inspiratory shortening. For both muscles there were linear relationships during the course of CO2 rebreathing between their peak moving average EMGs and total inspiratory shortening and between tidal volume and total inspiratory shortening. During single-breath airway occlusions, the electrical activity of both the diaphragm and intercostal muscles increased, but there were decreases in their tidal shortening. The extent of muscle shortening during occluded breaths was increased by hypercapnia, so that both muscles shortened more during occluded breaths under hypercapnic conditions (PCO2 up to 90 Torr) than during unoccluded breaths under normocapnic conditions. These results suggest that for the costal diaphragm and parasternal intercostal muscles there is a close relationship between their electrical and mechanical behavior during CO2 rebreathing, this relationship is substantially altered by occluding the airway for a single breath, and thoracic respiratory muscles do not contract quasi-isometrically during occluded breaths.     

How should we normalize electromyograms obtained from healthy participants? What we have learned from over 25 years of research

Electromyograms (EMGs) need to be normalized if comparisons are sought between trials when electrodes are reapplied, as well as between different muscles and individuals. The methods used to normalize EMGs recorded from healthy individuals have been appraised for more than a quarter of a century. Eight methods were identified and reviewed based on criteria relating to their ability to facilitate the comparison of EMGs. Such criteria included the magnitude and pattern of the normalized EMG, reliability, and inter-individual variability. If the aim is to reduce inter-individual variability, then the peak or mean EMG from the task under investigation should be used as the normalization reference value. However, the ability of such normalization methods to facilitate comparisons of EMGs is questionable. EMGs from MVCs can be as reliable as those from submaximal contractions, and do not appear to be affected by contraction mode or joint kinematics, particularly for the elbow flexors. Thus, the EMG from an isometric MVC is endorsed as a normalization reference value. Alternatively the EMG from a dynamic MVC can be used, although it is recognized that neither method is guaranteed to be able to reveal how active a muscle is in relation to its maximal activation capacity.     

An ensemble analysis of electromyographic activity during whole body pointing with the use of support vector machines

We explored the use of support vector machines (SVM) in order to analyze the ensemble activities of 24 postural and focal muscles recorded during a whole body pointing task. Because of the large number of variables involved in motor control studies, such multivariate methods have much to offer over the standard univariate techniques that are currently employed in the field to detect modifications. The SVM was used to uncover the principle differences underlying several variations of the task. Five variants of the task were used. An unconstrained reaching, two constrained at the focal level and two at the postural level. Using the electromyographic (EMG) data, the SVM proved capable of distinguishing all the unconstrained from the constrained conditions with a success of approximately 80% or above. In all cases, including those with focal constraints, the collective postural muscle EMGs were as good as or better than those from focal muscles for discriminating between conditions. This was unexpected especially in the case with focal constraints. In trying to rank the importance of particular features of the postural EMGs we found the maximum amplitude rather than the moment at which it occurred to be more discriminative. A classification using the muscles one at a time permitted us to identify some of the postural muscles that are significantly altered between conditions. In this case, the use of a multivariate method also permitted the use of the entire muscle EMG waveform rather than the difficult process of defining and extracting any particular variable. The best accuracy was obtained from muscles of the leg rather than from the trunk. By identifying the features that are important in discrimination, the use of the SVM permitted us to identify some of the features that are adapted when constraints are placed on a complex motor task.     

Altered patterns of muscle activation during performance of four functional tasks in patients with shoulder disorders: interpretation from voluntary response index.

Altered motor control of the shoulder muscles during performance of a specific motor task in patients with shoulder disorders (SDs) has been an interesting subject to researchers. This study compared shoulder muscle activation patterns by surface electromyography (sEMG), including the upper trapezius (UT), lower trapezius (LT), and serratus anterior (SA) muscles, during four functional tasks in 25 patients with SDs and controls. A voluntary response index (VRI) was calculated, including magnitude and similarity index (SI), to quantify sEMG patterns during four functional tasks. Responsiveness and clinically meaningful levels of discrimination between patients and control for EMG magnitude and SI were determined. An altered pattern of motor control during four functional tasks was evident in the patients, in which greater EMG amplitude and abnormal EMG patterns were found. For SI among four functional tasks, normal subjects ranged from 0.80 to 1.00 while patients ranged from 0.70 to 0.99. High probabilities (97%) of discrimination between patients and normal subjects were found by SI method during an overhead height task (patients: 0.85-0.96, normal subjects: 0.95-1.00). Our results also suggest that an individual can be estimated to be abnormal when lower SI values are observed during the four functional tasks.     

Automatic segmentation of surface EMG images: Improving the estimation of neuromuscular activity

Surface electromyograms (EMGs) recorded with a couple of electrodes are meant to comprise representative information of the whole muscle activation. Nonetheless, regional variations in neuromuscular activity seem to occur in numerous conditions, from standing to passive muscle stretching. In this study, we show how local activation of skeletal muscles can be automatically tracked from EMGs acquired with a bi-dimensional grid of surface electrodes (a grid of 8 rows and 15 columns was used). Grayscale images were created from simulated and experimental EMGs, filtered and segmented into clusters of activity with the watershed algorithm. The number of electrodes on each cluster and the mean level of neuromuscular activity were used to assess the accuracy of the segmentation of simulated signals. Regardless of the noise level, thickness of fat tissue and acquisition configuration (monopolar or single differential), the segmentation accuracy was above 60%. Accuracy values peaked close to 95% when pixels with intensity below ～70% of maximal EMG amplitude in each segmented cluster were excluded. When simulating opposite variations in the activity of two adjacent muscles, watershed segmentation produced clusters of activity consistently centered on each simulated portion of active muscle and with mean amplitude close to the simulated value. Finally, the segmentation algorithm was used to track spatial variations in the activity, within and between medial and lateral gastrocnemius muscles, during isometric plantar flexion contraction and in quiet standing position. In both cases, the regionalization of neuromuscular activity occurred and was consistently identified with the segmentation method.     

Comparison of electromyographic signals from monopolar current and potential amplifiers derived from a penniform muscle,the gastrocnemius medialis

Electromyograms (EMGs) are measured by bipolar surface electrodes that quantify potential differences. Bipolar potentials over penniform muscles may be associated with errors. Our assumption was that muscle activity can be quantified more reliably and with a higher spatial resolution using current measurements.The purpose of this work is: (a) to introduce the concept of current measurements to detect muscle activity, (b) to show the coherences observed over a segment of a typical penniform muscle, the gastrocnemius medialis where one would expect a synchronicity of the activation, and (c) to show the amount of mixing that is caused by the finite inter electrode resistance.A current amplifier was developed. EMGs were recorded at 40% of maximum voluntary contraction during isometric contractions of the gastrocnemius medialis. EMGs of twelve persons were recorded with an array of four peripheral and one central electrode. Monopolar EMGs were recorded for “all-potential”, “center at current” and “all-current” conditions. Coherence revealed the similarity of signals recorded from neighboring electrodes.Coherence was high for the “all-potential”, significant for the “current at center” condition and disappeared in the “all-current” condition.It was concluded that EMG array recordings strongly depends on the measurement configuration. The proposed current amplifier significantly improves spatial resolution of EMG array recordings because the inter-electrode cross talk is reduced.     

Neuromuscular activation of vastus intermedius muscle during fatiguing exercise

The purpose of this study was to investigate neuromuscular activation of the vastus intermedius (VI) muscle during fatiguing contraction. Seven healthy men performed sustained isometric knee extension exercise at 50% of maximal voluntary contraction until exhaustion. During the fatiguing task, surface electromyograms (EMGs) were recorded from four muscle components of the quadriceps femoris muscle group: VI; vastus lateralis (VL); vastus medialis (VM); and rectus femoris (RF) muscles. For the VI muscle, our recently developed technique was used. Root mean square (RMS) and median frequency (MF) of the surface EMG signal were calculated and these variables were then normalized by the value at the beginning of the task. Normalized RMS of the VI muscle resembled those of the other three muscles at all given times. At 95% of exhaustion time, normalized MF of the VI muscle was significantly higher than that of the VL muscle (p < 0.05). These results suggested that neuromuscular activation is not consistent between the VI and VL muscles at the exhaustion for isometric submaximal contraction and this could reflect the dissimilar intramuscular metabolism between these muscles.     

Insights gained into the interpretation of surface electromyograms from the gastrocnemius muscles: A simulation study

Interpretation of surface electromyograms (EMG) is usually based on the assumption that the surface representation of action potentials does not change during their propagation. This assumption does not hold for muscles whose fibers are oblique to the skin. Consequently, the interpretation of surface EMGs recorded from pinnate muscles unlikely prompts from current knowledge. Here we present a complete analytical model that supports the interpretation of experimental EMGs detected from muscles with oblique architecture. EMGs were recorded from the medial gastrocnemius muscle during voluntary and electrically elicited contractions. Preliminary indications obtained from simulated and experimental signals concern the spatial localization of surface potentials and the myoelectric fatigue. Specifically, the spatial distribution of surface EMGs was localized about the fibers superficial extremity. Strikingly, this localization increased with the pinnation angle, both for the simulated EMGs and the recorded M-waves. Moreover, the average rectified value (ARV) and the mean frequency (MNF) of interference EMGs increased and decreased with simulated fatigue, respectively. The degree of variation in ARV and MNF did not depend on the pinnation angle simulated. Similar variations were observed for the experimental EMGs, although being less evident for a higher fiber inclination. These results are discussed on a physiological context, highlighting the relevance of the model proposed here for the interpretation of gastrocnemius EMGs and for conceiving future experiments on muscles with pinnate geometry.     

Muscle activation patterns of selected lower extremity muscles during stepping and cutting tasks

Lower extremity muscle activations during crossover and side step cut tasks are hypothesized to play an important role in controlling knee motion, and therefore, impact the design of knee injury prevention and rehabilitation programs. However, the contribution of lower extremity muscles to frontal and transverse plane moments during cutting tasks is unclear. The purpose of this study was to compare the muscle activation patterns of selected lower extremity muscles (vastus lateralis, medial/lateral hamstrings and medial/lateral gastrocnemius) of subjects performing a stepping down and side step cut, a stepping down and crossover cut and an equivalent straight ahead task. Ground reaction force was used to determine the cut angle, stance time and compare the lower limb loading during each task. Electromyography data during all tasks were normalized to the average activation during the straight ahead tasks to determine relative changes in muscle activation between the straight ahead and different cut styles (crossover and side step). There were no differences in the pattern of muscle activation of the vastus lateralis, or lateral hamstring muscles when comparing the cutting tasks to the equivalent straight ahead task. However, the crossover cut task resulted in significantly higher muscle activation of the medial hamstrings and lateral gastrocnemius muscles relative to both the side step cut and straight ahead tasks. These results suggest the medial/lateral hamstrings and medial/lateral gastrocnemius play a role in transverse and frontal plane control during cut tasks.     

Responses of the anterolateral abdominal muscles during cough and expiratory threshold loading in the cat.

The present study was conducted to determine the pattern of activation of the anterolateral abdominal muscles during the cough reflex. Electromyograms (EMGs) of the rectus abdominis, external oblique, internal oblique, transversus abdominis, and parasternal muscles were recorded along with gastric pressure in anesthetized cats. Cough was produced by mechanical stimulation of the lumen of the intrathoracic trachea or larynx. The pattern of EMG activation of these muscles during cough was compared with that during graded expiratory threshold loading (ETL; 1-30 cmH(2)O). ETL elicited differential recruitment of abdominal muscle EMG activity (transversus abdominis > internal oblique > rectus abdominis congruent with external oblique). In contrast, both laryngeal and tracheobronchial cough resulted in simultaneous activation of all four anterolateral abdominal muscles with peak EMG amplitudes 3- to 10-fold greater than those observed during the largest ETL. Gastric pressures during laryngeal and tracheobronchial cough were at least eightfold greater than those produced by the largest ETL. These results suggest that, unlike their behavior during expiratory loading, the anterolateral abdominal muscles act as a unit during cough.     

Electromyogram refinement using muscle synergy based regulation of uncertain information

Electromyogram signal (EMG) measurement frequently experiences uncertainty attributed to issues caused by technical constraints such as cross talk and maximum voluntary contraction. Due to these problems, individual EMGs exhibit uncertainty in representing their corresponding muscle activations. To regulate this uncertainty, we proposed an EMG refinement, which refines EMGs with regulating the contribution redundancy of the signals from EMGs to approximating torques through EMG-driven torque estimation (EDTE) using the muscular skeletal forward dynamic model. To regulate this redundancy, we must consider the synergistic contribution redundancy of muscles, including “unmeasured” muscles, to approximating torques, which primarily causes redundancy of EDTE. To suppress this redundancy, we used the concept of muscle synergy, which is a key concept of analyzing the neurophysiological regulation of contribution redundancy of muscles to exerting torques. Based on this concept, we designed a muscle-synergy-based EDTE as a framework for EMG refinement, which regulates the abovementioned uncertainty of individual EMGs in consideration of unmeasured muscles. In achieving the proposed EMG refinement, the most considerable point is to suppress a large change such as overestimation attributed to enhancement of the contribution of particular muscles to estimating torques. Therefore it is reasonable to refine EMGs by minimizing the change in EMGs. To evaluate this model, we used a Bland-Altman plot, which quantitatively evaluates the proportional bias of refined signals to EMGs. Through this evaluation, we showed that the proposed EDTE minimizes the bias while approximating torques. Therefore this minimization optimally regulates the uncertainty of EMGs and thereby leads to optimal EMG refinement.     

Electromyographic assessment of trunk muscle activation amplitudes during a simulated lifting task using pattern recognition techniques

This study sought to determine the patterns of neuromuscular response from 24-trunk muscle sites during a symmetrical lift and replace task. Surface electromyograms (EMG) and kinematic variables were recorded from 29 healthy subjects. Pattern recognition techniques were used to examine how activation amplitude patterns changed with the different physical demands of the task (reach, phase of movement). The results indicated that there was very little trunk and pelvis motion during the task. Three principal patterns accounted for 95% of the total variation suggesting that the measured data had a simple underlying structure of variance. ANOVA results revealed significant differences in principal pattern scores. These differences captured subtle changes in muscle recruitment strategies that most likely reflect different stability and biomechanical demands. More balanced activations (bracing) between the abdominal and back sites were observed during the lighter demands, whereas differential recruitment among the back extensor sites was more predominant in the more demanding conditions. A pattern recognition technique offers a novel method to examine the relationships among a large number of muscles and test how different work characteristics change the relationships among the muscle sites.     

Motor pattern control for increasing crushing force in the striped burrfish (Chilomycterus schoepfi)

The relationship between muscular force modulation and the underlying nervous system control signals has been difficult to quantify for in vivo animal systems. Our goal was to understand how animals alter muscle activation patterns to increase bite forces and to evaluate how accurate these patterns are in predicting crushing forces. We examined the relationship between commonly used measures of cranial muscle activity and force production during feeding events of the striped burrfish (Chilomycterus schoepfi), a mollusc crushing specialist. We quantified the force required to crush a common gastropod prey item (Littorina irrorata) of burrfish using a materials testing device. Burrfish were fed these calibrated prey items while we recorded electromyograms (EMGs) from the main jaw closing muscles (adductor mandibulae A1beta, A2alpha, and A2beta). We quantified EMG activity by measuring the burst duration, rectified integrated area, and then calculated the intensity of activity from these two variables. Least squares regressions relating force to crush (Fcrush) and all EMG variables were calculated for each fish. Multiple regression analyses were used to determine how much of the variation in Fcrush could be explained by muscle activation patterns. We found that 20 cm burrfish are capable of generating extremely high crushing forces (380 N peak force) primarily by increasing the duration of muscle activity. EMG variables explained 71% of the total variation in force production. After accounting for the inherent variation in Fcrush of snails, EMGs do a very good job of predicting bite forces for these fish.     

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.     

Assessment of upper extremity muscle function in persons with tetraplegia

The objective was to investigate the actual level of muscle function impairment in tetraplegic persons and, inextricably related to this, the possibilities to compensate function loss using new muscle coalitions. In this cross-sectional group study, 20 persons with a cervical spinal cord injury (SCI) at segmental levels C5C6 or C7C8 and 10 control persons participated. Activity from 21 upper extremity and trunk muscles was recorded during standardised gross upper extremity task performance. No substantial differences in main activation patterns were found between C7C8 and control subjects. In contrast, main activation patterns in C5C6 persons showed an absence of selectivity, which may be explained by the participants activating every controllable muscle in an attempt to perform maximally. In order to identify more intricate differences in muscle activation between control and C7C8 persons a fine motor function task may be necessary. Muscle activation patterns during arm task performance were stable in all three groups.     

Activation of human arm muscles during flexion/extension and supination/pronation tasks: A theory on muscle coordination

Generally the number of muscles acting across a joint exceeds the number of degrees of freedom available to the joint. This redundancy raises a problem regarding the ratio in which these muscles are activated during a particular motor task. In this paper we present a theory to explain the activation patterns of muscles used during voluntary and reflex induced contractions. The basic assumptions underlying the theory are that 1) coordination of muscles is based on synergistic muscle activities, 2) the synergisms involved satisfy certain transformations of muscle spindle signals to muscle activation signals and 3) muscle spindle output is proportional to the ratio of muscle stretch and muscle length in lengthening muscles, and is zero in shortening muscles. The theory is used to predict the recruitment threshold of motor units in six arm muscles during voluntary isometric contractions. All theoretical predictions are in reasonable agreement with the experimentally observed behavior of a large population of motor units within each muscle. However, within a single muscle sometimes motor-unit populations have been found to have different types of recruitment behavior. This deviating behavior is discussed in the light of the theory presented here.     

Upper airway muscle and diaphragm responses to hypoxia in the piglet

The neonatal ventilatory response to hypoxia is characterized by initial transient stimulation and subsequent respiratory depression. It is unknown, however, whether this response is also exhibited by the upper airway muscles that regulate nasal, laryngeal, and pharyngeal patency. We therefore compared electromyogram (EMG) amplitudes and minute EMGs for the diaphragm (DIA), alae nasi (AN), posterior cricoarytenoid (PCA), and genioglossus (GG) muscles in 12 anesthetized spontaneously breathing piglets during inhalation of 12% O2 over 10 min. Minute EMG for the DIA responded to hypoxia with an initial transient increase and subsequent return to prehypoxia levels by 10 min. Hypoxia also stimulated all three upper airway muscles. In contrast to the DIA EMG, however, AN, PCA, and GG EMGs all remained significantly above prehypoxia levels after 10 min of hypoxia. We have thus demonstrated that the initial stimulation and subsequent depression of the DIA EMG after 12% O2 inhalation contrast with the sustained increase in AN, PCA, and GG EMGs during hypoxia. We speculate that 1) central inhibition during neonatal hypoxia is primarily distributed to the motoneuron pools regulating DIA activation and 2) peripheral chemoreceptor stimulation and/or central disinhibition induced by hypoxia preferentially influence those motoneuron pools that regulate upper airway muscle activation, causing the different hypoxic responses of these muscle groups in the young piglet.     

Comparison of Trunk Activity during Gait Initiation and Walking in Humans

To understand the role of trunk muscles in maintenance of dynamic postural equilibrium we investigate trunk movements during gait initiation and walking, performing trunk kinematics analysis, Erector spinae muscle (ES) recordings and dynamic analysis. ES muscle expressed a metachronal descending pattern of activity during walking and gait initiation. In the frontal and horizontal planes, lateroflexion and rotation occur before in the upper trunk and after in the lower trunk. Comparison of ES muscle EMGs and trunk kinematics showed that trunk muscle activity precedes corresponding kinematics activity, indicating that the ES drive trunk movement during locomotion and thereby allowing a better pelvis mobilization. EMG data showed that ES activity anticipates propulsive phases in walking with a repetitive pattern, suggesting a programmed control by a central pattern generator. Our findings also suggest that the programs for gait initiation and walking overlap with the latter beginning before the first has ended.     

New Data on Axial Locomotion in Fishes: How Speed Affects Diversity of Kinematics and Motor Patterns

Despite considerable recent progress in understanding the functionof the axial muscles and skeleton in fishes, generalizing fromthese results has been hindered by the great phylogenetic diversityof taxa, the lack of quantitative morphometric data on axialmusculoskeletal structure, and limited analysis of the fullrange of locomotor behaviors exhibited within any one taxon.This paper reviews novel results from our studies of two taxawithin a single monophyletic clade, the sunfish family Centrarchidae.Integrated analyses of lateral displacement, lateral bending,and axial muscle activity reveal widespread effects of swimmingspeed both within a particular mode of swimming and among differentbehavioral modes. The longitudinal position along the body ofthe fish also commonly affects kinematics, muscle activity andthe timing of electromyograms (EMGs) relative to kinematics.EMGs and kinematic events propagate from head to tail for bothsteady and kick and glide swimming. In contrast, during theescape response, the onset of EMGs forms a standing wave pattern,whereas kinematic events are propagated. Several novel featuresof the axial motor pattern are summarized for the kick and glidemode of unsteady swimming. For example, the onset of white fiberEMGs lags significantly behind that of the red fibers at thesame longitudinal position, and red fibers are inactivated athigher unsteady swimming speeds. Muscle activity propagatesposteriorly via the sequential activation of myomeres, but thereare statistically significant differences in the timing of EMGsfrom the contractile tissue opposite a single vertebra. Duringrelatively slow kick and glide swimming, the extreme dorsaland ventral portions of myomeres are not active. Estimates ofthe longitudinal extent of the fish with simultaneous muscleactivity indicate that EMGs from an individual myomere usuallyhave temporal overlap with more than 20 neighboring myomereson the same side of the fish. Consequently, the functional unitsfor axial locomotion of fishes do not correspond simply to theanatomical units of individual myomeres. Rather the in vivomotor pattern is a primary determinant of the functional unitsinvolved in swimming.