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.     

Repeatability of maximal voluntary force and of surface EMG variables during voluntary isometric contraction of quadriceps muscles in healthy subjects

The repeatability of initial values and rate of change of EMG signal mean spectral frequency (MNF), average rectified values (ARV), muscle fiber conduction velocity (CV) and maximal voluntary contraction (MVC) was investigated in the vastus medialis obliquus (VMO) and vastus lateralis (VL) muscles of both legs of nine healthy male subjects during voluntary, isometric contractions sustained for 50 s at 50% MVC. The values of MVC were recorded for both legs three times on each day and for three subsequent days, while the EMG signals have been recorded twice a day for three subsequent days. The degree of repeatability was investigated using the Fisher test based upon the ANalysis Of VAriance (ANOVA), the Standard Error of the Mean (SEM) and the Intraclass Correlation Coefficient (ICC).

Data collected showed a high level of repeatability of MVC measurement (normalized SEM from 1.1% to 6.4% of the mean). MNF and ARV initial values also showed a high level of repeatability (ICC>70% for all muscles and legs except right VMO). At 50% MVC level no relevant pattern of fatigue was observed for the VMO and VL muscles, suggesting that other portions of the quadriceps might have contributed to the generated effort. These observations seem to suggest that in the investigation of muscles belonging to a multi-muscular group at submaximal level, the more selective electrically elicited contractions should be preferred to voluntary contractions.     

Pain-induced changes in cervical muscle activation do not affect muscle fatigability during sustained isometric contraction

This study investigated whether pain-induced changes in cervical muscle activation affect myoelectric manifestations of cervical muscle fatigue. Surface EMG signals were detected from the sternocleidomastoid and splenius capitis muscles bilaterally from 14 healthy subjects during 20-s cervical flexion contractions at 25% of the maximal force. Measurements were performed before and after the injection of 0.5 ml of hypertonic (painful) or isotonic (control) saline into either the sternocleidomastoid or splenius capitis in two experimental sessions. EMG average rectified value and mean power spectral frequency were estimated throughout the sustained contraction. Sternocleidomastoid or splenius capitis muscle pain resulted in lower sternocleidomastoid EMG average rectified value on the side of pain (P < 0.01). However, changes over time of sternocleidomastoid EMG average rectified value and mean frequency (myoelectric manifestations of fatigue) during sustained flexion were not changed during muscle pain. These results demonstrate that pain-induced modifications of cervical muscle activity do not change myoelectric manifestations of fatigue. This finding has implications for interpreting the mechanisms underlying greater cervical muscle fatigue in people with neck pain disorders.     

Non-uniform electromyographic activity during fatigue and recovery of the vastus medialis and lateralis muscles

The aim of the study was to investigate EMG signal features during fatigue and recovery at three locations of the vastus medialis and lateralis muscles. Surface EMG signals were detected from 10 healthy male subjects with six 8-electrode arrays located at 10%, 20%, and 30% of the distance from the medial (for vastus medialis) and lateral (vastus lateralis) border of the patella to the anterior superior spine of the pelvic. Subjects performed contractions at 40% and 80% of the maximal force (MVC) until failure to maintain the target force, followed by 20 2-s contractions at the same force levels every minute for 20 min (recovery). Average rectified value, mean power spectral frequency, and muscle fiber conduction velocity were estimated from the EMG signals in 10 epochs from the beginning of the contraction to task failure (time to task failure, mean ± SD, 70.7 ± 25.8 s for 40% MVC; 27.4 ± 16.8 s for 80% MVC) and from the 20 2 s time intervals during recovery. During the fatiguing contraction, the trend over time of EMG average rectified value depended on location for both muscles (P < 0.05). After 20-min recovery, mean frequency and conduction velocity of both muscles were larger than in the beginning of the fatigue task (P < 0.05) (supernormal values). Moreover, the trend over time of mean frequency during recovery was affected by location and conduction velocity values depended on location for both muscles (P < 0.05). The results indicate spatial dependency of EMG variables during fatigue and recovery and thus the necessity of EMG spatial sampling for global muscle assessment.     

Surface EMG signal processing during isometric contractions

This paper provides an overview of techniques suitable for the estimation, interpretation and understanding of time variations that affect the surface electromyographic (EMG) signal during sustained voluntary or electrically elicited contractions. These variations concern amplitude variables, spectral variables and muscle fiber conduction velocity, are interdependent and are referred to as the ‘fatigue plot'. The fatigue plot provides information suitable for the classification of muscle behavior. In addition, the information obtainable by means of linear electrode arrays is discussed, and applications of mathematical models for the interpretation of array signals are presented. The model approach provides additional ways for the classification of muscle behavior.     

High frequency vibration conditioning stimulation centrally reduces myoelectrical manifestation of fatigue in healthy subjects

ObjectiveVibration conditioning has been adopted as a tool to improve muscle force and reduce fatigue onset in various rehabilitation settings. This study was designed to asses if high frequency vibration can induce some conditioning effects detectable in surface EMG (sEMG) signal; and whether these effects are central or peripheral in origin.Design300 Hz vibration was applied for 30 min during 5 consecutive days, to the right biceps brachii muscle of 10 healthy males aged from 25 to 50 years. sEMG was recorded with a 16 electrode linear array placed on the skin overlying the vibrated muscle. The test protocol consisted of 30% and 60% maximal voluntary contraction (MVC) as well as involuntary (electrically elicited) contractions before and after treatment.ResultsNo statistically significant differences were found between PRE and POST vibration conditioning when involuntary stimulus-evoked contraction and 30% MVC were used. Significant differences in the initial values and rates of change of muscle fibre conduction velocity were found only at 60% MVC.Conclusions300 Hz vibration did not induce any peripheral changes as demonstrated by the lack of differences when fatigue was electrically induced. Differences were found only when the muscle was voluntarily fatigued at 60% MVC suggesting a modification in the centrally driven motor unit recruitment order, and interpreted as an adaptive response to the reiteration of the vibratory conditioning.     

Variations in the relationship between the frequency content of EMG signals and the rate of torque development in voluntary and elicited contractions.

Our purpose was to characterize the relationship between EMG mean power frequency (MPF) or median frequency (MF) and rate of torque development in voluntary ballistic and electrically elicited isometric contractions. Twenty-three healthy adults participated in two sets of experiments performed on elbow flexor muscles. For Experiment 1, subjects were asked to generate voluntary ballistic contractions by reaching four different target torque levels (20, 40, 60 and 100% of the maximal voluntary contraction (MVC)) as fast as they could. For Experiment 2, electrical (M-waves) and mechanical (twitches) responses to electrical stimulation of the nerves supplying the biceps brachii and brachioradialis muscles were recorded with the subjects at rest and with a background isometric contraction of 15% MVC. MPF, MF and rate of torque development (% MVC/s) were calculated for both voluntary and elicited contractions. Significant positive correlations were observed between MPF and rate of torque development for the voluntary contractions, whereas significant negative correlations were observed between the two variables for elicited contractions. This suggests that factors other than muscle fiber composition influence the frequency content of EMG signals and/or the rate of torque development, and that the effect of these factors will vary between voluntary and elicited contractions.     

Muscle weakness following sustained and rhythmic isometric contractions in man

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

Effect of caffeine on skeletal muscle function before and after fatigue

We studied the effect of caffeine on voluntary and electrically stimulated contractions of the adductor pollicis muscle in five adult volunteers. Caffeine (500 mg) was administered orally in a double-blind fashion. Electrical stimulation of the ulnar nerve was performed at 10, 20, 30, 50, and 100 Hz before and after a sustained voluntary contraction held at 50% of the maximal voluntary contraction (MVC). A brief tetanus at 30 Hz was also performed to calculate relaxation rate in the fresh muscle. Contractile properties, relaxation rate, and endurance were then assessed after caffeine and placebo, as well as the response of the fatigued muscle to different frequencies of stimulation. There was no difference in the maximal tension obtained with electrical stimulation (T100) or in the MVC between placebo and caffeine. The tensions developed with electrical stimulation at lower frequencies increased significantly with caffeine ingestion, shifting the frequency-force curve to the left, both before and after fatigue. Mean plasma caffeine concentration associated with these responses was 12.2 +/- 4.9 mg/l. We conclude that caffeine has a direct effect on skeletal muscle contractile properties both before and after fatigue as demonstrated by electrical stimulation.     

Mechanical and myoelectric manifestations of fatigue in subjects with anorexia nervosa.

PURPOSE: Aim of this work is to compare mechanical and myoelectric manifestations of fatigue during an isometric contraction at 80% of maximal voluntary contraction (MVC) in a population of eight anorexic female patients (AN: 24.9+/-6.5 years, mean+/-SD) with respect to a group of seven healthy female subjects (CO: 30.0+/-6.6 years, mean+/-SD). METHODS: sEMG signals were recorded, using a linear electrode array (eight channels, 10mm apart), from vastus lateralis muscle of the dominant side. MVC, endurance time, initial value and rate of change of the EMG variables [conduction velocity (CV), mean power frequency (MNF), average rectified value (ARV)] were studied during the fatiguing contractions. RESULTS: Absolute and relative (normalized with respect to the body weight) knee torque values and endurance times were found not statistically different between the two groups. Similarly, EMG amplitude initial values and rate of change and MNF initial values were found not significantly different between the two groups. CV initial values and CV rate of change were found greater in healthy than in pathological subjects (AN CV: 3.74+/-0.86m/s, CO CV: 4.96+/-0.64m/s, p=0.004; AN CV rate of change: 0.006+/-0.015m/s2, CO CV rate of change: -0.006+/-0.007m/s2, p=0.006, mean+/-SD). Contrary to expectations MNF rate of changes in the AN group (-0.35+/-0.16Hz/s) was found greater than in the CO group (-0.17+/-0.13Hz/s, p=0.004, Mann-Whitney U test, mean+/-SD). CONCLUSIONS: CV values and their rate of change were compatible with a predominance of type I fibres (and/or with an hypotrophy of type II fibres) and/or with a lower sub-cutaneous tissue thickness with respect to CO group, as described in the literature with this pathology. The behaviour of MNF during sustained contractions, opposite to that of CV, suggests an altered central control strategy aimed to increase mechanical force output increasing the level of synchronization of motor units. This study confirms the capability of sEMG to assess muscle condition during severe malnutrition suggesting further studies to assess if sEMG can be used to monitor the effect of re-feeding and rehabilitation treatments.     

Effects of aging on EMG variables during fatiguing isometric contractions

The purpose of this study was to evaluate the neuromuscular adaptation that occurred with aging, by comparing young and aged subjects with respect to changes in surface EMG from the tibialis anterior muscle during fatiguing contractions. EMG variables such as the averaged rectified value (ARV), median frequency (MDF), and muscle fiber conduction velocity (MFCV) were calculated during maximal (MVC, 3 sec) and submaximal (60% MVC, 60 sec) isometric contractions. Muscular force, ARV, MDF, and MFCV during MVC were significantly greater in the young than in the elderly (p < 0.05). EMG amplitude increased and the waveform slowed in all subjects during submaximal contractions, indicating the development of local muscle fatigue. As fatigue progressed, the ARV increased and the MDF and MFCV decreased significantly (p < 0.01). The fatigue-induced changes in the MDF and MFCV were significantly smaller in aged than in young subjects (p < 0.05), a trend also seen in the ARV change, which means that the elderly cannot be fatigued as much as the young with contractions of the same relative intensity. These results as a whole suggest that the aged subjects hold an adaptive motor strategy to cope with age-related neuromuscular deteriorations, due to the decline of motor unit activation and selective atrophy of fast twitch muscle fibers.     

The muscle sound properties of different muscle fiber types during voluntary and electrically induced contractions.

Soundmyogram (SMG) and electromyogram signals were recorded simultaneously from the relatively fast medial gastrocnemius (MG) and slow soleus (SOL) during voluntary and electrically induced contractions. Using a spike-triggered averaging technique, the averaged elementary sound and corresponding MU spikes were also obtained from about 35 different MUs identified. The rms-SMG of MG increased as a function of force (P < 0.01). On the contrary, these values for SOL increased up to 60% MVC (P < 0.01), but decreased at 80% MVC. The relationship between the peak to peak amplitude of SMG and MU spike indicated significant positive correlations (r = 0.631 to approximately 0.657, P < 0.01). During electrical stimulation at 5 Hz, the SMG power spectral peak frequency (PF) was matched with stimulation frequency in both muscles. At higher stimulation frequencies, e.g., > 15 Hz, only in the MG was SMG-PF synchronized with stimulation frequency; the slow SOL did not show such synchronization. Our data suggest that the SMG frequency components might reflect active motor unit firing rates, and that the SMG amplitude depends upon mechanical properties of contraction, muscle fiber composition, and firing rate during voluntary and electrically induced contractions.     

Pulse charge and not waveform affects M-wave properties during progressive motor unit activation

The aim of this study was to investigate changes in experimentally recorded M-waves with progressive motor unit (MU) activation induced by transcutaneous electrical stimulation with different pulse waveforms. In 10 subjects, surface electromyographic signals were detected with a linear electrode array during electrically elicited contractions of the biceps brachii muscle. Three different monophasic waveforms of 304-μs duration were applied to the stimulation electrode on the main muscle motor point: triangular, square, and sinusoidal. For each waveform, increasing stimulation current intensities were applied in 10 s (frequency: 20 Hz). It was found that: (a) the degree of MU activation, as indicated by M-wave average rectified value, was a function of the injected charge and not of the stimulation waveform, and (b) MUs tended to be recruited in order of increasing conduction velocity with increasing charge of transcutaneous stimulation. Moreover, the subjects reported lower discomfort during the contractions elicited by the triangular waveform with respect to the others. Since subject tolerance to the stimulation protocol must be considered as important as MU recruitment in determining the effectiveness of neuromuscular electrical stimulation (NMES), we suggest that both charge and waveform of the stimulation pulses should be considered relevant parameters for optimizing NMES protocols.     

Feasibility study of detecting surface electromyograms in severely obese patients

The aims of this study were to examine if surface EMG signals can be detected from the quadriceps femoris muscle of severely obese patients and to investigate if differences exist in quadriceps force and myoelectric manifestations of fatigue between obese patients and lean controls.Fourteen severely obese patients (body mass index, BMI, mean ± SD: 44.9 ± 6.3 kg/m²) and fourteen healthy controls (BMI: 23.7 ± 2.5 kg/m²) were studied. The vastus medialis and lateralis of the dominant thigh were concurrently investigated during voluntary isometric contractions (10-s long at submaximal and maximal intensities and intermittent submaximal contractions until exhaustion) and sustained (120-s long) electrically elicited contractions.We found that the detection of surface EMG signals from the quadriceps is feasible also in severely obese subjects presenting increased thickness of the subcutaneous fat tissue. In addition, we confirmed and extended previous findings showing that the volume conductor properties determine the amplitude and spectral features of the detected surface EMG signals: the lower the subcutaneous tissue thickness, the higher the amplitude and mean frequency estimates. Further, we found no differences in the mechanical and myoelectric manifestations of fatigue during intermittent voluntary and sustained electrically elicited contractions between obese patients and lean controls.     

Repeatability of surface EMG variables during voluntary isometric contractions of the biceps brachii muscle.

The repeatability of initial value and rate of change of mean spectral frequency (MNF), average rectified values (ARV) and muscle fiber conduction velocity (CV) was investigated in the dominant biceps brachii of ten normal subjects during sustained isometric voluntary contractions. Four levels of contraction were studied: 10%, 30%, 50% and 70% of the maximal voluntary contraction level (MVC). Each contraction was repeated three times in each of three different days for a total of nine contractions/level/subject and 90 contractions per level across the ten subjects. Repeatability was investigated using the Intraclass Correlation Coefficient (ICC) and the standard error of the mean (SEM) of the estimates for each subject. Contrary to observations in other muscles, CV estimates appeared to be very repeatable both within and between subjects. CV showed a small but significant increase when contraction force increased from 10% to 50% MVC but no change for further increase of force. As force increased, MNF showed a slight decrease possibly related to a wider spreading of the CV values. The rate of time decrement of MNF and CV increased with the level of contraction. The normalized decrement (% of initial value per second) was in general higher for MNF than for CV and was more repeatable between subjects at 10% MVC than at 70% MVC. A final observation is that a resting time of 5 minutes may not be sufficient after a contraction at 50% or 70% MVC.     

Differential changes in muscle oxygenation between voluntary and stimulated isometric fatigue of human dorsiflexors.

The purpose of this study was to compare fatigue and recovery of maximal voluntary torque [maximal voluntary contraction (MVC)] and muscle oxygenation after voluntary (Vol) and electrically stimulated (ES) protocols of equal torque production. On 1 day, 10 male subjects [25 yr (SD 4)] completed a Vol fatigue protocol and, on a separate day, an ES fatigue protocol of the right dorsiflexors. Each task involved 2 min of intermittent (2-s on, 1-s off) isometric contractions at 50% of MVC. For the ES protocol, stimulation was delivered percutaneously to the common peroneal nerve at a frequency of 25 Hz. Compared with the Vol protocol, the ES protocol caused a greater impairment in MVC (75 vs. 83% prefatigue value; Pre) and greater increase in 50-Hz half relaxation time (165 vs. 117% Pre) postexercise. After acute (1 min) recovery, MVC impairment was similar for both protocols, whereas 50- Hz half relaxation time was still greater in the ES than Vol protocol. Total hemoglobin decreased to a similar extent in both protocols during exercise, but it was elevated above the resting value to a significantly greater extent for the ES protocol in recovery (18 vs. 11 microM). Oxygen saturation was significantly lower in the ES than Vol protocol during exercise (46 vs. 57% Pre), but it was significantly greater during recovery (120 vs. 105% Pre). These findings suggest that despite, equal torque production, ES contractions impose a greater metabolic demand on the muscle that leads to a transient greater impairment in MVC. The enforced synchronization and fixed frequency of excitation inherent to ES are the most likely causes for the exacerbated changes in the ES compared with the Vol protocol.     

Reliability of burst superimposed technique to assess central activation failure during fatiguing contraction.

Recording a superimposed electrically-induced contraction at the limit of endurance during voluntary contraction is used as an indicator of failure of muscle activation by the central nervous system and discards the existence of peripheral muscle fatigue. We questioned on the reliability of this method by using other means to explore peripheral muscle failure. Fifteen normal subjects sustained handgrip at 60% of maximal voluntary contraction (MVC) until exhaustion. During sustained contraction, the power spectrum analysis of the flexor digitorum surface electromyogram allowed us to calculate the leftward shift of median frequency (MF). A superimposed 60 Hz 3 s pulse train (burst superimposition) was delivered to the muscle when force levelled off close to the preset value. Immediately after the fatigue trial had ended, the subject was asked to perform a 5 s 60% MVC and we measured the peak contractile response to a 60 Hz 3 s burst stimulation. Recordings of the compound evoked muscle action potential (M-wave) allowed us to explore an impairment of neuromuscular propagation. A superimposed contraction was measured in 7 subjects in their two forearms, whereas it was absent in the 8 others. Despite these discrepancies, all subjects were able to reproduce a 3 s 60% MVC immediately after the fatigue trial ended and there was no post-fatigue decrease of contraction elicited by the 60 Hz 3 s burst stimulation, as well as no M-wave decrease in amplitude and conduction time. Thus, there was no indication of peripheral muscle fatigue. MF decrease was present in all individuals throughout the fatiguing contraction and it was not correlated with the magnitude of superimposed force. These observations indicate that an absence of superimposed electrically-induced muscle contraction does not allow us to conclude the existence of a sole peripheral muscle fatigue in these circumstances.     

M-wave properties during progressive motor unit activation by transcutaneous stimulation.

The aim of this study was to interpret changes in experimentally recorded M waves with progressive motor unit (MU) activation based on simulation of the surface electromyogram. Activation order during transcutaneous electrical stimulation was analyzed by investigating M-wave average rectified value, spectral properties, and conduction velocity (CV) during electrically elicited contractions. M-waves were detected from the biceps brachii muscle of 10 healthy male subjects by a linear adhesive array of eight electrodes. Electrical stimulation was delivered to the motor point at either constant current intensity (40, 60, 80, and 100% of the supramaximal stimulation current) or with linearly increasing current. A model of surface electromyogram generation that varied activation order based on MU size and location was used to interpret the experimental results. From the experimental and model analysis, it was found that 1) MUs tended to be activated from low to high CV and from the superficial to the deep muscle layers with increasing transcutaneous electrical stimulation of the biceps brachii muscle, and 2) characteristic spectral frequencies of the M-wave were affected by many factors other than average CV (such as the activation order by MU location or the spread of the MU innervation zones and CVs), thus decreasing with a concomitant increase in CV during progressive MU activation.     

Rate of fatigue during repeated submaximal contractions of human quadriceps muscle.

Our purpose was to determine the effect of eight different combinations of contraction intensity, duration, and rest on the rate of fatigue in vastus lateralis muscle. A single combination consisted of contractions at 30 or 70% maximal voluntary contraction (MVC), held for 3 or 7 s with 3- or 7-s rest intervals. Contractions were repeated until the subject could not hold the force for the requisite duration. At regular intervals during each experiment, a brief MVC, a single twitch, and the response to eight stimulation pulses at 50 Hz were elicited. The rate of fatigue was the rate of decline of MVC calculated from regression analysis. Mean rate of fatigue (n = 8) ranged from 0.3 to 25% MVC/min and was closely related (r = 0.98) to the product of the relative force and the duty cycle. Force from 50 Hz stimulation fell linearly and in parallel with MVC. Twitch force was first potentiated and then fell twice as fast as 50 Hz stimulation and MVC (p less than 0.05). Differentiated twitch contraction and relaxation rates were higher at potentiation and lower at the limit of endurance, compared with control values (p less than 0.05). The maximal electromyogram decreased 25% and the submaximal EMG increased to maximal by the end of the protocol, indicating that the entire motor unit pool had been recruited. The close relation between rate of fatigue and the force x time product probably reflects the off-setting interaction of contraction amplitude, duration, and rest interval. This occurs despite the changes in twitch characteristics and the apparent recruitment of fast fatiguing motor units.     

Torque potentiation and myosin light-chain phosphorylation in human muscle following a fatiguing contraction

Indices of electrically stimulated and maximal voluntary isometric muscle torgue and the phosphate content of myosin phosphorylatable light chains (P light chains) were studied during recovery following a 60-s maximal voluntary isometric contraction (MVC) in 21 human subjects. Analysis of muscle biopsy samples revealed that immediately after the 60-s MVC there were significant decreases in ATP (-15%) and phosphocreatine (-82%), and lactate concentration increased by 17-fold. All indices of muscle torque production were reduced by the 60-s MVC, but the twitch torque and torque at 10 Hz were relatively less reduced compared with the torque at 20 and 50 Hz or a 1-s MVC. Between 3 and 6 min of recovery, twitch torque and torque at 10 Hz stimulation were significantly potentiated, reaching peak values of 125 and 134%, respectively, compared with rest. Phosphate content of the fast and two slow P light chains was significantly increased over rest levels immediately after and 4 min after the 60-s MVC. These results suggest that myosin P light-chain phosphorylation could provide a mechanism to increase human muscle torque under conditions of submaximal contractile element activation following fatigue.