The vastus lateralis neuromuscular activity during all-out cycling exercise

ObjectiveThe objective of this work was to study modifications in motor control through surface electromyographic (sEMG) activity during a very short all-out cycling exercise.MethodsTwelve male cyclists (age 23 ± 4 years) participated in this study. After a warm-up period, each subject performed three all-out cycling exercises of 6 s separated by 2 min of complete rest. This protocol was repeated three times with a minimum of 2 days between each session. The braking torque imposed on cycling motion was 19 N m. The sEMG of the vastus lateralis was recorded during the first seven contractions of the sprint. Time–frequency analysis of sEMG was performed using continuous wavelet transform. The mean power frequency (MPF, qualitative modifications in the recruitment of motor units) and signal energy (a quantitative indicator of modifications in the motor units recruitment) were computed for the frequency range 10–500 Hz.ResultssEMG energy increased (P ? 0.05) between contraction number 1 and 2, decreased (P ? 0.05) between contraction number 2 and 3 then stabilized between contraction number 3 and 7 during the all-out test. MPF increased (P ? 0.05) during the all-out test. This increase was more marked during the first two contractions.ConclusionsThe decrease in energy and the increase in the sEMG MPF suggest a large spatial recruitment of motor units (MUs) at the beginning of the sprint followed by a preferential recruitment of faster MUs at the end of the sprint, respectively.     

The application of independent component analysis to the multi-channel surface electromyographic signals for separation of motor unit action potential trains: part II-modelling interpretation.

The purpose of this article was to investigate whether or not FastICA can separate identical motor unit action potential trains (MUAPTs) of the 8-channel surface electromyographic (sEMG) signals constructed by an sEMG model into the independent components. Firstly, we have examined how much the increase of motor units (MUs) in the simulated sEMG signals influenced the performance on the separation of MUAPTs by kurtosis. The decreased trend of mean kurtosis on both sEMG signals and their independent components were observed as MUs were increased. These data suggested that the separation performance decayed when MUs were increased. Secondary, the differences between the independent components and the principal components have been also applied to the simulated sEMG signals with or without time delay between the sEMG channels. FastICA could successfully separate identical MUAPTs with no time delay but principal component analysis (PCA) could not do so. Against it, both FastICA and PCA could not separate MUAPTs with some time delay. In conclusion, our results suggested that FastICA could separate identical MUAPTs with no time delay into the independent components by FastICA, which might offer a new technique for the separation of interfered MUAP waveforms based on statistical properties of sEMG signal distributions.     

A simulation study on the relation between muscle motor unit numbers and the non-Gaussianity/non-linearity levels of surface electromyography

Recent research has demonstrated that surface electromyography (sEMG) signals have non-Gaussianity and non-linearity properties. It is known that more muscle motor units are recruited and firing rates (FRs) increase as exertion increases. A hypothesis was proposed that the Gaussianity test (S _g) and linearity test (S _?) levels of sEMG signals are associated with the number of active motor units (nMUs) and the FR. The hypothesis has only been preliminarily discussed in experimental studies. We used a simulation sEMG model involving spatial (active MUs) and temporal (three FRs) information to test the hypothesis. Higher-order statistics (HOS) from the bi-frequency domain were used to perform S _g and S _?. Multivariate covariance analysis and a correlation test were employed to determine the nMUs-S _g relationship and the nMUs-S _? relationship. Results showed that nMUs, the FR, and the interaction of nMUs and the FR all influenced the S _g and S _? values. The nMUs negatively correlated to both the S _g and S _? values. That is, at the three FRs, sEMG signals tended to a more Gaussian and linear distribution as exertion and nMUs increased. The study limited experiment factors to the sEMG non-Gaussianity and non-linearity levels. The study quantitatively described nMUs and the FR of muscle that are not directly available from experiments. Our finding has guiding significance for muscle capability assessment and prosthetic control.     

Spectral properties of electromyographic and mechanomyographic signals during dynamic concentric and eccentric contractions of the human biceps brachii muscle

The purpose of this study was to describe and examine the variations in recruitment patterns of motor units (MUs) in biceps brachii (BB) through a range of joint motion during dynamic eccentric and concentric contractions. Twelve healthy participants (6 females, 6 males, age = 30 ± 8.5 years) performed concentric and eccentric contractions with constant external loading at different levels. Surface electromyography (EMG) and mechanomyography (MMG) were recorded from BB. The EMGs and MMGs were decomposed into their intensities in time–frequency space using a wavelet technique. The EMG and MMG spectra were then compared using principal component analysis. Variations in total intensity, first principal component (PCI), and the angle θ formed by first component (PCI) and second component (PCII) loading scores were explained in terms of MU recruitment patterns and elbow angles. Elbow angle had a significant effect on dynamic concentric and eccentric contractions. The EMG total intensity was greater for concentric than for eccentric contractions in the present study. MMG total intensity, however, was lower during concentric than during eccentric contractions. In addition, there was no significant difference in θ between concentric and eccentric contractions for both EMG and MMG. Selective recruitment of fast MUs from BB muscle during eccentric muscle contractions was not found in the present study.     

Assessment of the non-Gaussianity and non-linearity levels of simulated sEMG signals on stationary segments

The purpose of this paper was to evaluate the effects of the longitudinal single differential (LSD), the longitudinal double differential (LDD) and the normal double differential (NDD) spatial filters, the electrode shape, the inter-electrode distance (IED) on non-Gaussianity and non-linearity levels of simulated surface EMG (sEMG) signals when the maximum voluntary contraction (MVC) varied from 10% to 100% by a step of 10%. The effects of recruitment range thresholds (RR), the firing rate (FR) strategy and the peak firing rate (PFR) of motor units were also considered.A cylindrical multilayer model of the volume conductor and a model of motor unit (MU) recruitment and firing rate were used to simulate sEMG signals in a pool of 120 MUs for 5 s. Firstly, the stationarity of sEMG signals was tested by the runs, the reverse arrangements (RA) and the modified reverse arrangements (MRA) tests. Then the non-Gaussianity was characterised with bicoherence and kurtosis, and non-linearity levels was evaluated with linearity test.The kurtosis analysis showed that the sEMG signals detected by the LSD filter were the most Gaussian and those detected by the NDD filter were the least Gaussian. In addition, the sEMG signals detected by the LSD filter were the most linear. For a given filter, the sEMG signals detected by using rectangular electrodes were more Gaussian and more linear than that detected with circular electrodes. Moreover, the sEMG signals are less non-Gaussian and more linear with reverse onion-skin firing rate strategy than those with onion-skin strategy. The levels of sEMG signal Gaussianity and linearity increased with the increase of the IED, RR and PFR.     

Muscular strength and power are correlated with motor unit action potential amplitudes,but not myosin heavy chain isoforms in sedentary males and females

It remains unclear if the sizes of higher-threshold motor units (MU) are associated with muscular strength and power. Therefore, the purpose of this study was to examine sex-related differences in muscle cross-sectional area (mCSA), percent myosin heavy chain (%MHC) isoform expression, and the MU action potential amplitudes (MUAP_AMPS)-recruitment threshold (RT) relationships of the vastus lateralis and isometric peak torque, isokinetic peak torque and mean power at 1.05 rad·s⁻¹ of the leg extensors. Surface electromyographic decomposition techniques were used to quantify MUAP_AMPS recorded during isometric muscle actions at 70% of maximal voluntary contractions and regressed against RTs with the slopes calculated. Ultrasound images were used to measure mCSA. Males had greater slopes from the MUAP_AMP-RT relationship than the females (P < 0.05). The greater slopes likely reflected larger higher-threshold MUs for the males. The mCSAs and slopes from the relationships were strongly correlated with isometric and isokinetic peak torque and isokinetic mean power (r = 0.78–0.82), however, type I %MHC isoform was only moderately correlated with isometric peak torque (r = −0.54). The results indicated that sex-related differences in muscular strength and power were associated more so with the sizes of the higher-threshold MUs (slopes) and mCSA than MHC isoforms. The amount of cross-bridge activity within muscle fibers that comprise higher-threshold MUs may be the primary contributor to muscular strength and power rather than the contractile properties of the muscle.     

Distribution of motor unit potential velocities in the biceps brachii muscle of sprinters and endurance athletes during prolonged dynamic exercises at low force levels

In surface electromyography (sEMG), the distribution of motor unit potential (MUP) velocities has been shown to reflect the proportion of faster and slower propagating MUPs. This study investigated whether the distribution of MUP velocities could distinguish between sprinters (n = 11) and endurance athletes (n = 12) in not-specifically trained muscle (biceps brachii) during prolonged dynamic exercises at low forces. sEMG was acquired during 4 min’ exercises: unloaded, 5%, 10% and 20% of maximal voluntary contraction (MVC). The features extracted from the sEMG were: the mean muscle conduction velocity – estimated using the inter-peak latency and cross-correlation methods, the within-subject skewness (expressing the proportions of faster and slower propagating MUPs) and the within-subject standard deviation of MUP velocities (SD-mup). Sprinters showed a greater proportion of faster propagating MUPs than endurance athletes. During fatigue, the SD-mup of sprinters broadened progressively, whereas that of endurance athletes did not. The findings suggest that sprinters conveyed a greater proportion of faster motor units than endurance athletes and that motor unit behavior during fatigue differed between groups. Thus, the distribution of MUP velocities enables distinction between a muscle of sprinters and endurance athletes during prolonged dynamic exercises at low forces.     

The effect of fast and slow motor unit activation on whole-muscle mechanical performance: the size principle may not pose a mechanical paradox

The output of skeletal muscle can be varied by selectively recruiting different motor units. However, our knowledge of muscle function is largely derived from muscle in which all motor units are activated. This discrepancy may limit our understanding of in vivo muscle function. Hence, this study aimed to characterize the mechanical properties of muscle with different motor unit activation. We determined the isometric properties and isotonic force–velocity relationship of rat plantaris muscles in situ with all of the muscle active, 30% of the muscle containing predominately slower motor units active or 20% of the muscle containing predominately faster motor units active. There was a significant effect of active motor unit type on isometric force rise time (p < 0.001) and the force–velocity relationship (p < 0.001). Surprisingly, force rise time was longer and maximum shortening velocity higher when all motor units were active than when either fast or slow motor units were selectively activated. We propose this is due to the greater relative effects of factors such as series compliance and muscle resistance to shortening during sub-maximal contractions. The findings presented here suggest that recruitment according to the size principle, where slow motor units are activated first and faster ones recruited as demand increases, may not pose a mechanical paradox, as has been previously suggested.     

Human motor unit recruitment and derecruitment during long lasting intermittent contractions.

Seven healthy subjects were investigated in cyclic ramp-and-hold long lasting isometric contractions. Wire branched electrodes were used for selective recording of single motor unit (MU) potentials from m. biceps brachii. MU behaviour was defined in terms of recruitment/derecruitment thresholds (RT and DT) and the duration of interspike intervals (ISI). A total of 63 MUs was investigated: 40 units were active from the beginning of the task performance and another 23 were recruited later. There were no changes in the recruitment pattern of MUs with fatigue development - a short first ISI followed by a very long second one and an almost constant firing rate after this transient phase. The tendency of RT to gradually decrease dominates the results. Thus, the required constant rate of force increase with fatigue development was maintained mostly by the mechanisms of space coding (i.e., decrease of RT and recruitment of additional MUs). Oppositely, the time behaviour of the DT changes was not uniform and rate coding was an essential mechanism in the adaptation of MU activity to muscle fatigue during relaxation phases. The recruitment pattern and fatigue related behaviour of the additionally recruited MUs were similar to those of MUs active from the first cycle of the motor task performance.     

Effect of synchronization of firings of different motor unit types on the force variability in a model of the rat medial gastrocnemius muscle

The synchronized firings of active motor units (MUs) increase the oscillations of muscle force, observed as physiological tremor. This study aimed to investigate the effects of synchronizing the firings within three types of MUs (slow—S, fast resistant to fatigue–FR, and fast fatigable–FF) on the muscle force production using a mathematical model of the rat medial gastrocnemius muscle. The model was designed based on the actual proportion and physiological properties of MUs and motoneurons innervating the muscle. The isometric muscle and MU forces were simulated by a model predicting non-synchronized firing of a pool of 57 MUs (including 8 S, 23 FR, and 26 FF) to ascertain a maximum excitatory signal when all MUs were recruited into the contraction. The mean firing frequency of each MU depended upon the twitch contraction time, whereas the recruitment order was determined according to increasing forces (the size principle). The synchronization of firings of individual MUs was simulated using four different modes and inducing the synchronization of firings within three time windows (± 2, ± 4, and ± 6 ms) for four different combinations of MUs. The synchronization was estimated using two parameters, the correlation coefficient and the cross-interval synchronization index. The four scenarios of synchronization increased the values of the root-mean-square, range, and maximum force in correlation with the increase of the time window. Greater synchronization index values resulted in higher root-mean-square, range, and maximum of force outcomes for all MU types as well as for the whole muscle output; however, the mean spectral frequency of the forces decreased, whereas the mean force remained nearly unchanged. The range of variability and the root-mean-square of forces were higher for fast MUs than for slow MUs; meanwhile, the relative values of these parameters were highest for slow MUs, indicating their important contribution to muscle tremor, especially during weak contractions.     

Time-related changes of motor unit properties in the rat medial gastrocnemius muscle after spinal cord injury. I. Effects of total spinal cord transection

The contractile properties of motor units (MUs) were electrophysiologically investigated in the medial gastrocnemius (MG) muscle in 17 Wistar three-month-old female rats: 14, 30, 90 and 180 days after the total transection of the thoracic spinal cord and compared to those in intact (control) rats. A sag phenomenon, regularly observed in unfused tetani of fast units in intact animals at 40 Hz stimulation, almost completely disappeared in spinal rats. Therefore, the MUs of intact and spinal rats were classified as fast or slow types basing on 20 Hz tetanus index, the value of which was lower or equal 2.0 for fast and higher than 2.0 for slow MUs. The MUs composition of MG muscle changed with time after the spinal cord transection: an increasing proportion of fast fatigable (FF) units starting one month after injury and a disappearance of slow (S) units within the three months were observed. In all MUs investigated the twitch contraction and half-relaxation time were significantly prolonged after injury (p < 0.01, Mann–Whitney U-test). Moreover, a decrease of the fatigue index for fast resistant (FR) and slow MUs was observed in subsequent groups of spinal rats. No significant changes were found between twitch forces in all MU types of spinal animals (p > 0.05). However, due to a decrease of the maximal tetanic force, a significant rise of the twitch-to-tetanus ratio of all MUs in spinal rats was detected (p < 0.01). The considerable reduction of ability to potentiate the force was noticed for fast, especially FF type MUs. In conclusion, the spinal cord transection leads to changes in the proportion of the three MU types in rat MG muscle. The majority of changes in MUs’ contractile properties were developed progressively with time after the spinal cord injury. However, the most intensive alterations of twitch-time parameters were observed in rats one month after the transection.     

Orderly recruitment among motoneurons supplying different muscles.

Virtually all movements involve the recruitment of motor units from multiple muscles. Given the functional diversity of motor units (motoneurons and the muscle fibers they supply), the effective production of specific movements undoubtedly depends upon some principle(s) to organize the ensemble of active motor units. The principle acting to organize the recruitment of motor units within muscles is the size principle, whereby the first motor units to be recruited have the smallest values for axonal conduction velocity and contractile force, and are the slowest to contract and fatigue. Here we consider the possibility that the size principle applies in the recruitment of motor units across muscles, i.e., that regardless of their muscles of origin, active motor units are recruited in rank order, for example, from low to high conduction velocity. The benefits of orderly recruitment across muscles could be similar to the acknowledged advantages of orderly recruitment within muscles. One benefit is that the neural process involved in organizing active motor units would be simplified. In a muscle-based scheme, the size principle would organize only those motor units within individual muscles, leaving the nervous system with the additional task of coordinating the relative activities of motor units from different muscles. By contrast, in an ensemble-based scheme, orderly recruitment of all motor units according to the size principle would automatically coordinate motor units both within and across motor nuclei. Another potential benefit is the provision for movements with smooth trajectory, the result of interleaving the divergent torque contributions made by motor units from muscles that differ in their orientations about joints. Otherwise, if order were restricted within muscles, the torque trajectory of a joint would change unevenly as participating muscles begin contracting at different times and grade activity at different rates. These considerations support speculation that motor units recruited from co-contracting muscles are collectively recruited according to the size principle.     

The fatigability of two agonistic muscles in human isometric voluntary submaximal contraction: an EMG study. II. Motor unit firing rate and recruitment

The recruitment and firing rate of biceps brachii (BB) and brachioradialis (BR) motor units (MUs) were studied in the course of fatiguing isometric contractions at 20%-30% of maximal voluntary contraction (MVC). MU recruitment generally occurred throughout the maintained contraction and was similar for BB and BR muscles. Newly recruited MUs started to discharge in the form of bursts, the duration of which increased until a continuous rhythmical firing was achieved. Within each burst, the first interval between two consecutive discharges was usually the shortest. MU threshold was lowered just after the limit time of the maintained contraction. The MU's firing rate either increased or remained stable as a function of the elapsed time. It is concluded that (1) in fatiguing isometric contractions at 20%-30% MVC contractile failure is mainly compensated for by MU recruitment and a lowered MU threshold and (2) differences between in surface changes in the electromyogram of BB and BR muscles cannot easily be explained by related differences in MU firing rate and recruitment.     

Amplitude and spectral characteristics of biceps Brachii sEMG depend upon speed of isometric force generation.

In the present study the influence of speed of contraction on the interplay between recruitment and firing rate of motor units (MUs) was assessed. The surface electromyographic (sEMG) signal was recorded in nine healthy subjects from the right biceps brachii using a linear electrode array during ramp isometric contractions (from 0 to 100% of the maximal voluntary force, MVC) at 5, 10, and 20% MVC s(-1) (ramp phase), followed by 10 s of sustained MVC (hold phase). The median frequency (MDF), Root Mean Square (RMS) and conduction velocity (CV) of sEMG, were computed on adjacent epochs covering a force range of 5% MVC each. Full motor unit recruitment (FMUR) point was assessed as the force level at which MDF reached its maximum value; the MDF decay during the hold phase was taken as an index of localized muscle fatigue. At 5% MVC s(-1), FMUR was reached at 52.3% MVC. At 10%MVC s(-1) FMUR was achieved at 58% MVC; while at 20% MVC s(-1) FMUR point was located at 77% MVC, being statistically different from 5 and 10% MVCs(-1) ramps (p<0.05). The MDF decay was steeper at higher speed. CV modifications mirrored those reported for MDF. The RMS increased in a curvilinear fashion and the maximum value was always attained during the hold phase. Our findings suggest that MU recruitment strategies are significantly related to the speed of contraction even in a single muscle.     

Measuring the accuracies of motor unit firing times and action potential waveforms derived from surface electromyographic decomposition

This study included spike trigger averaging (STA) procedures to examine the acceptability of the Precision Decomposition (PD) III derived motor unit action potential (MUAP) trains that met the >90% accuracy criteria from the reconstruct-and-test. MUs met the >90% accuracy criteria from the reconstruct-and-test with STA procedures then applied. Y-intercepts and slopes were calculated for the firing rate- and MUAP amplitude-recruitment threshold relationships. Gaussian noise (1% of the SD of the mean interspike interval) was added to the firing times with the changes in MUAPs quantified. A total of 455 MUs were decomposed with 155 MUs removed as a result of the reconstruct-and-test. Five additional MUs were excluded via the STA criteria. The MUAP waveforms deteriorated with the inclusion of Gaussian noise. There were differences in the derived action potentials amplitudes of higher-threshold MUs between the PD III algorithm and the STA procedure. There was excellent agreement among the slopes and y-intercepts between the relationships that included or excluded MUs that did not meet the STA criteria. There was good agreement between the MUAP amplitude-recruitment threshold relationships derived from the PD III and STA procedure. The addition of the STA procedures did not alter the MU-derived relationships.     

Effects of whole body vibration on motor unit recruitment and threshold

Whole body vibration (WBV) has been suggested to elicit reflex muscle contractions but this has never been verified. We recorded from 32 single motor units (MU) in the vastus lateralis of 7 healthy subjects (34 ± 15.4 yr) during five 1-min bouts of WBV (30 Hz, 3 mm peak to peak), and the vibration waveform was also recorded. Recruitment thresholds were recorded from 38 MUs before and after WBV. The phase angle distribution of all MUs during WBV was nonuniform (P < 0.001) and displayed a prominent peak phase angle of firing. There was a strong linear relationship (r = -0.68, P < 0.001) between the change in recruitment threshold after WBV and average recruitment threshold; the lowest threshold MUs increased recruitment threshold (P = 0.008) while reductions were observed in the higher threshold units (P = 0.031). We investigated one possible cause of changed thresholds. Presynaptic inhibition in the soleus was measured in 8 healthy subjects (29 ± 4.6 yr). A total of 30 H-reflexes (stimulation intensity 30% Mmax) were recorded before and after WBV: 15 conditioned by prior stimulation (60 ms) of the antagonist and 15 unconditioned. There were no significant changes in the relationship between the conditioned and unconditioned responses. The consistent phase angle at which each MU fired during WBV indicates the presence of reflex muscle activity similar to the tonic vibration reflex. The varying response in high- and low-threshold MUs may be due to the different contributions of the mono- and polysynaptic pathways but not presynaptic inhibition.     

Motor unit recruitment strategies investigated by surface EMG variables.

During isometric contractions of increasing strength, motor units (MUs) are recruited by the central nervous system in an orderly manner starting with the smallest, with muscle fibers that usually show the lowest conduction velocity (CV). Theory predicts that the higher the velocity of propagation of the action potential, the higher the power at high frequencies of the detected surface signal. These considerations suggest that the power spectral density of the surface detected electromyogram (EMG) signal may give indications about the MU recruitment process. The purpose of this paper is to investigate the potential and limitations of spectral analysis of the surface EMG signal as a technique for the investigation of muscle force control. The study is based on a simulation approach and on an experimental investigation of the properties of surface EMG signals detected from the biceps brachii during isometric linearly increasing torque contractions. Both simulation and experimental data indicate that volume conductor properties play an important role as confounding factors that may mask any relation between EMG spectral variables and estimated CV as a size principle parameter during ramp contractions. The correlation between spectral variables and CV is thus significantly lower when the MU pool is not stable than during constant-torque isometric contractions. Our results do not support the establishment of a general relationship between spectral EMG variables and torque or recruitment strategy.     

Percutaneous electrical stimulation in strength training: an update

Numerous studies have used percutaneous electrical stimulation (PES) in the context of training programs to develop strength and physical performance in healthy populations (sedentary or trained). Significant increases in muscle and fiber cross-sectional area, isokinetic peak torque, maximal isometric and dynamic strength, and motor performance skills have been found after PES training. These strength gains are explained on the basis of the characteristics of PES motor units (MUs) recruitment: (a) a continuous and exhausting contractile activity in the same pool of MUs during the entire exercise period, (b) a supramaximal temporal recruitment imposed by the high frequency chosen (up to 40 Hz), and (c) a synchronous recruitment of neighboring fibers. The PES training method is complementary to voluntary training, mainly because the application of PES causes an unconventional spatial recruitment of MUs that, depending on the muscular topography, may entail the preferential recruitment of the fast-twitch MUs. In addition, the method does not specifically develop elasticity in skeletal muscle, and it must be accompanied by a technical workout.     

Motor unit recruitment for dynamic tasks: current understanding and future directions

Skeletal muscle contains many muscle fibres that are functionally grouped into motor units. For any motor task there are many possible combinations of motor units that could be recruited and it has been proposed that a simple rule, the ‘size principle’, governs the selection of motor units recruited for different contractions. Motor units can be characterised by their different contractile, energetic and fatigue properties and it is important that the selection of motor units recruited for given movements allows units with the appropriate properties to be activated. Here we review what is currently understood about motor unit recruitment patterns, and assess how different recruitment patterns are more or less appropriate for different movement tasks. During natural movements the motor unit recruitment patterns vary (not always holding to the size principle) and it is proposed that motor unit recruitment is likely related to the mechanical function of the muscles. Many factors such as mechanics, sensory feedback, and central control influence recruitment patterns and consequently an integrative approach (rather than reductionist) is required to understand how recruitment is controlled during different movement tasks. Currently, the best way to achieve this is through in vivo studies that relate recruitment to mechanics and behaviour. Various methods for determining motor unit recruitment patterns are discussed, in particular the recent wavelet-analysis approaches that have allowed motor unit recruitment to be assessed during natural movements. Directions for future studies into motor recruitment within and between functional task groups and muscle compartments are suggested.