Activation of human quadriceps femoris during isometric, concentric, and eccentric contractions.

Maximal and submaximal activation level of the right knee-extensor muscle group were studied during isometric and slow isokinetic muscular contractions in eight male subjects. The activation level was quantified by means of the twitch interpolation technique. A single electrical impulse was delivered, whatever the contraction mode, on the femoral nerve at a constant 50 degrees knee flexion (0 degrees = full extension). Concentric, eccentric (both at 20 degrees /s velocity), and isometric voluntary activation levels were then calculated. The mean activation levels during maximal eccentric and maximal concentric contractions were 88.3 and 89.7%, respectively, and were significantly lower (P < 0.05) with respect to maximal isometric contractions (95.2%). The relationship between voluntary activation levels and submaximal torques was linearly fitted (P < 0.01): comparison of slopes indicated lower activation levels during submaximal eccentric compared with isometric or concentric contractions. It is concluded that reduced neural drive is present during 20 degrees /s maximal concentric and both maximal and submaximal eccentric contractions. These results indicate a voluntary activation dependency on both tension levels and type of muscular actions in the human knee-extensor muscle group.     

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.     

Mechanical behavior of the quadriceps femoris muscle tendon unit during low-load contractions.

We examined the relationships between morphology and muscle-tendon dynamics of the quadriceps femoris muscle of 11 men using velocity-encoded phase-contrast magnetic resonance imaging (MRI). Thigh muscle electromyography and joint range of motion were first measured outside the MRI scanner during knee extension-flexion tasks that were performed at a rate of 40 times/min with elastic bands providing peak resistance of 5.2 kp (SD 0.4) to the extension. The same movement was repeated inside the MRI scanner bore where tissue velocities and muscle morphology were recorded. The average displacement in the proximal and distal halves of the rectus femoris and vastus intermedius aponeuroses was different (P = 0.049), reflecting shortening (1.6%), but the tensile strain along the length of the aponeuroses was uniform. The aponeurosis behavior varied among individuals, and these individual patterns were best explained by the differences in relative cross-sectional area of rectus femoris to vastus muscles (r = 0.71, P = 0.014). During dynamic contraction, considerable deformation of muscles in the axial plane caused an anatomic measure such as muscle thickness to change differently (decrease or increase) in different sites of measurement. For example, when analyzed from the axial images, the vastus lateralis thickness did not change (P = 0.946) in the frontal plane through femur but increased in a 45 degrees oblique plane between the frontal and sagittal planes (P = 0.004). The present observations of the heterogeneity and individual behavior emphasize the fact that single-point measurements do not always reflect the overall behavior of muscle-tendon unit.     

Effect of quadriceps femoris muscle length on neural activation during isometric and concentric contractions.

The effect of muscle length on neural drive (here termed "neural activation") was investigated from electromyographic activities and activation levels (twitch interpolation). The neural activation was measured in nine men during isometric and concentric (30 and 120 degrees /s) knee extensions for three muscle lengths (35, 55, and 75 degrees knee flexion, i.e., shortened, intermediate, and lengthened muscles, respectively). Long (76 degrees ), medium (56 degrees ), and short (36 degrees ) ranges of motion were used to investigate the effect of the duration of concentric contraction. Neural activation was found to depend on muscle length. Reducing the duration of contraction had no effect. Neural activation was higher with short muscle length during isometric contractions and was weaker for shortened than for intermediate and lengthened muscles performing 120 degrees /s concentric contractions. Muscle length had no effect on 30 degrees /s concentric neural activation. Peripheral mechanisms and discharge properties of the motoneurons could partly explain the observed differences in the muscle length effect. We thus conclude that muscle length has a predominant effect on neural activation that would modulate the angular velocity dependency.     

Force depression in human quadriceps femoris following voluntary shortening contractions.

The purpose of this study was to investigate whether the isometric muscle force, redeveloped following maximal-effort voluntary shortening contractions in human skeletal muscle, is smaller than the purely isometric muscle force at the corresponding length. Isometric knee extensor moments, surface electromyographic (EMG) signals of quadriceps femoris, and interpolated twitch moments (ITMs) were measured while 10 subjects performed purely isometric knee extensor contractions at a 60 degrees knee angle and isometric knee extensor contractions at a 60 degrees knee angle preceded by maximal-effort voluntary shortening of the quadriceps muscles. It was found that the knee extensor moments were significantly decreased for the isometric-shortening-isometric contractions compared with the isometric contractions for the group as a whole, whereas the corresponding EMG and ITM values were the same. This study is the first to demonstrate force depression following muscle shortening for voluntary contractions. We concluded that force depression following muscle shortening is an actual property of skeletal muscle rather than a stimulation artifact and that force depression during voluntary contraction is not accompanied by systematic changes in muscle activation as evaluated by EMG and ITM.     

Knee extensor torque and quadriceps femoris EMG during perceptually-guided isometric contractions.

The aim of this study was to examine superficial quadriceps femoris (QF) EMG and torque at perceived voluntary contraction efforts. Thirty subjects (15 males, 15 females) performed 9, 5 s, sub-maximal contractions at prescribed levels of perceived voluntary effort at points 1-9 on an 11-point scale (0-10), in a random order. Surface electromyograms (EMG) of the vastus medialis (VM), vastus lateralis (VL), and rectus femoris (RF) muscles, as well as QF peak torque (PT), average torque (AT), and torque coefficient of variation (C.V.), were sampled. The raw EMG signals were full-wave rectified and integrated over the middle three s of each contraction. The sampled EMG signals, and PT and AT at each perceived exertion level were normalized to the average of three maximal voluntary contractions. The normalized EMG and torque values at each perceived exertion level were then compared to equivalent percent values (i.e., 10% at a perceived level of 1). The results demonstrated that at all perceived exertion levels, with the exception of the RF at a level of 2 which was equivalent to 20%, and the VL and RF muscles at a level 1 in which activation was greater than 10%, activation was significantly less than the equivalent percent value at each point on the scale. VM EMG was found to be less than the VL and RF from contraction levels 3-9. PT was shown to be less than the equivalent percent values at contraction levels 6-9. The AT was found to be lower than the expected percent value at perceived effort levels 2-9. Torque C.V. was not found to be different across the range of perceived effort. The major findings of this study suggested that humans over-estimate voluntary QF muscle torque when guided by perceptual sensations. It is also suggested that the produced EMG signals revealed a reliance on the VL muscle for knee extensor torque generation at sub-maximal levels.     

Time-frequency methods applied to muscle fatigue assessment during dynamic contractions.

This paper discusses the assessment of the electrical manifestations of muscle fatigue during dynamic contractions. In the past, the study of muscle fatigue was restricted to isometric constant force contractions because, in this contraction paradigm, the myoelectric signal may be considered as wide sense stationary over epochs lasting up to two or three seconds, and hence classic spectral estimation techniques may be applied. Recently, the availability of spectral estimation techniques specifically designed for nonstationary signal analysis made it possible to extend the employment of muscle fatigue assessment to cyclic dynamic contractions, thus increasing noticeably its possible clinical applications. After presenting the basics of time-frequency distributions, we introduce instantaneous spectral parameters well suited to tracking spectral changes due to muscle fatigue, discuss the issues of quasi-stationarity and quasi-cyclostationarity, and present different strategies of signal analysis to be utilized with cyclic dynamic contractions. We present preliminary results obtained by analyzing data collected from paraspinal muscles during repetitive lift movements, from the first dorsal interosseus during abduction-adduction movements of the index finger, and from knee flexors and extensors during isokinetic exercise. In conclusion, data herein reported demonstrate that the described techniques allow for evidencing the electrical manifestations of muscle fatigue in different paradigms of cyclic dynamic contractions. We believe that the extension of the objective assessment of the electrical manifestations of muscle fatigue from static to dynamic contractions may increase considerably the interest of researchers and clinicians and open new application fields, as ergonomics and sports medicine.     

A comparison of EMG-based muscle fatigue assessments during dynamic contractions

Several EMG-based approaches to muscle fatigue assessment have recently been proposed in the literature. In this work, two multivariate fatigue indices developed by the authors: a generalized mapping index (GMI) and the first component of principal component analysis (PCA) were compared to three univariate indices: Dimitrov’s normalized spectral moments (NSM), Gonzalez-Izal’s waveletbased indices (WI), and Talebinejad’s fractal-based Hurst Exponent (HE). Nine healthy participants completed two repetitions of fatigue tests during isometric, cyclic and random fatiguing contractions of the biceps brachii. The fatigue assessments were evaluated in terms of a modified sensitivity to variability ratio yielding the following scores (mean ± std.dev.): PCA: (12.6 ± 5.6), GMI: (11.5 ± 5.4), NSM: (10.3 ± 5.4), WI: (8.9 ± 4.6), HE: (8.0 ± 3.3). It was shown that PCA statistically outperformed WI and HE (p < 0.01) and that GMI outperformed HE (p < 0.02). There was no statistical difference among NSM, WI and HE (p > 0.2). It was found that taking the natural logarithm of NSM and WI, although reducing the parameters’ sensitivity to fatigue, increased SVR scores by reducing variability.     

EMG spectral indices and muscle power fatigue during dynamic contractions

The purpose of this study was to examine acute exercise-induced changes on muscle power output and surface electromyography (sEMG) parameters (amplitude and spectral indices of muscle fatigue) during a dynamic fatiguing protocol. Fifteen trained subjects performed five sets consisting of 10 leg presses (10RM), with 2 min rest between sets. Surface electromyography was recorded from vastus medialis (VM) and lateralis (VL) and biceps femoris (BF) muscles. A number of EMG-based parameters were compared for estimation accuracy and sensitivity to detect peripheral muscle fatigue. These were: Mean Average Voltage, median spectral frequency, Dimitrov spectral index of muscle fatigue (FI_nsm5), as well as other parameters obtained from a time–frequency analysis (Choi–Williams distributions) such as mean and variance of the instantaneous frequency and frequency variance. The log FI_nsm5 as a single parameter predictor accounted for 37% of the performance variance of changes in muscle power and the log FI_nsm5 and MFM as a two factor combination predictor accounted for 44%. Peripheral impairments assessed by sEMG spectral index FI_nsm5 may be a relevant factor involved in the loss of power output after dynamic high-loading fatiguing task.     

A dynamometer for the measurement of the extension torque of the lower leg during static and dynamic contractions of the quadriceps femoris muscle

A dynamometer which makes an angular movement is described. The dynamometer enables the measurement of the extension torque of the lower leg at different knee angles during static and slow concentric and eccentric contractions of the quadriceps femoris muscle. The influence of gravity on the measured torque signal can be compensated for by another signal representing the angular movement. The application of the dynamometer is demonstrated by giving an example of measurement.     

Voluntary activation level and muscle fiber recruitment of human quadriceps during lengthening contractions.

Voluntary activation levels during lengthening, isometric, and shortening contractions (angular velocity 60 degrees/s) were investigated by using electrical stimulation of the femoral nerve (triplet, 300 Hz) superimposed on maximal efforts. Recruitment of fiber populations was investigated by using the phosphocreatine-to-creatine ratio (PCr/Cr) of single characterized muscle fibers obtained from needle biopsies at rest and immediately after a series of 10 lengthening, isometric, and shortening contractions (1 s on/1 s off). Maximal voluntary torque was significantly higher during lengthening (270 +/- 55 N.m) compared with shortening contractions (199 +/- 47 N.m, P < 0.05) but was not different from isometric contractions (252 +/- 47 N.m). Isometric torque was higher than torque during shortening (P < 0.05). Voluntary activation level during maximal attempted lengthening contractions (79 +/- 8%) was significantly lower compared with isometric (93 +/- 5%) and shortening contractions (92 +/- 3%, P < 0.05). Mean PCr/Cr values of all fibers from all subjects at rest were 2.5 +/- 0.6, 2.0 +/- 0.7, and 2.0 +/- 0.7, respectively, for type I, IIa, and IIax fibers. After 10 contractions, the mean PCr/Cr values for grouped fiber populations (regardless of fiber type) were all significantly different from rest (1.3 +/- 0.2, 0.7 +/- 0.3, and 0.8 +/- 0.6 for lengthening, isometric, and shortening contractions, respectively; P < 0.05). The cumulative distributions of individual fiber populations after either contraction mode were significantly different from rest (P < 0.05). Curves after lengthening contractions were less shifted compared with curves from isometric and shortening contractions (P < 0.05), with a smaller shift for the type IIax compared with type I fibers in the lengthening contractions. The results indicate a reduced voluntary drive during lengthening contractions. PCr/Cr values of single fibers indicated a hierarchical order of recruitment of all fiber populations during maximal attempted lengthening contractions.     

EMG-based muscle fatigue assessment during dynamic contractions using principal component analysis

A novel approach to fatigue assessment during dynamic contractions was proposed which projected multiple surface myoelectric parameters onto the vector connecting the temporal start and end points in feature-space in order to extract the long-term trend information. The proposed end to end (ETE) projection was compared to traditional principal component analysis (PCA) as well as neural-network implementations of linear (LPCA) and non-linear PCA (NLPCA). Nine healthy participants completed two repetitions of fatigue tests during isometric, cyclic and random fatiguing contractions of the biceps brachii. The fatigue assessments were evaluated in terms of a modified sensitivity to variability ratio (SVR) and each method used a set of time-domain and frequency-domain features which maximized the SVR. It was shown that there was no statistical difference among ETE, PCA and LPCA (p > 0.99) and that all three outperformed NLPCA (p < 0.0022). Future work will include a broader comparison of these methods to other new and established fatigue indices.     

EMG frequency content changes with increasing force and during fatigue in the quadriceps femoris muscle of men and women.

The purpose of this study was to determine the effect of gender on changes in electromyographic (EMG) signal characteristics of the quadriceps muscles with increasing force and with fatigue. A total of fourteen healthy adults (seven men, seven women) participated in the study. Subjects had to perform isometric ramp contractions in knee extension with the force gradually increasing from 0 to 100% of the maximal voluntary contraction (MVC) in a 6-s period. Subjects then performed a fatigue task, consisting of a sustained maximum isometric knee extension contraction held until force decreased below 50% of the pre-fatigue MVC. Subjects also performed a single ramp contraction immediately after the fatigue task. The Root Mean Square (RMS) amplitude, mean power frequency (MPF) and median frequency (MF) of EMG signals obtained from the vastus lateralis, vastus medialis and rectus femoris were calculated at nine different force levels from the ramp contractions (10, 20, 30, 40, 50, 60, 70, 80 and 90% MVC), as well as every 5 s during the fatigue task. The main results were a more pronounced increase in EMG RMS amplitude for the three muscles and in MPF for the VL muscle with force in men compared with women. No significant effect of gender was found with regards to fatigue. These observations most likely reflect a moderately greater type II fiber content and/or area in the VL muscle of men compared to that of women.     

Contraction mode shift in quadriceps femoris muscle activation during dynamic knee extensor exercise with increasing loads

The objective of the present study was to examine the superficial quadriceps femoris (QF) muscle electromyogram (EMG) during dynamic sub-maximal knee extension exercise between young adult men and women. Thirty subjects completed, in a random order, 2 sub-maximal repetitions of single-leg knee extensions at 20-90% of their one-repetition maximum (1RM). Vastus medialis (VM), vastus lateralis (VL) and rectus femoris (RF) muscle integrated EMG (IEMG) during each sub-maximal lift was normalized to the respective 1RM for concentric, isometric and eccentric modes. The EMG median frequency (f(med)) was determined over the isometric mode. Men attained a significantly (p<0.05) greater knee angular velocity than the women during the concentric mode (83.6+/-19.1 degrees /s and 67.4+/-19.8 degrees /s, respectively). RF IEMG was significantly lesser than the VM (p=0.014) and VL (p<0.001) muscles, when collapsed across all contraction modes, loads, and sex. Overall IEMG was significantly greater during the concentric (p<0.001) and isometric (p<0.001) modes, than the eccentric mode. Men generated significantly (p=0.03) greater VL muscle IEMG than the women, while the opposite pattern emerged for the RF muscle. VM f(med) (105.1+/-11.1Hz) was significantly lesser than the VL (180.3+/-19.5Hz) and RF (127.7+/-13.9Hz) muscles across all lifting intensities, while the men (137.7+/-10.7Hz) generated greater values than the women (129.0+/-11.4Hz). The findings demonstrate a reduction in QF muscle activation across the concentric to eccentric transition, which may be related to the mode-specific velocity pattern.     

Electromyographic activity and rate of muscle fatigue of the quadriceps femoris during cycling exercise in the severe domain

This study compared the activation pattern and the fatigue rate among the superficial muscles of the quadriceps femoris (QF) during severe cycling exercise. Peak oxygen consumption (VO(2)peak) and maximal accumulated oxygen Deficit (MAOD) were established by 10 well-trained male cyclists (27.5 ± 4.1 years, 71.0 ± 10.3 kg, 173.4 ± 6.6 cm, mean VO(2)peak 56.7 ± 4.4 ml·kg·min(-1), mean MAOD 5.7 ± 1.1 L). Muscle activity (electromyographic [EMG] signals) was obtained during the supramaximal constant workload test (MAOD) and expressed by root mean square (RMS) and median frequency (MF slope). The RMS of the QF, vastus lateralis (VL) and vastus medialis (VM) muscles were significantly higher than at the beginning after 75% of exercise duration, whereas for the rectus femoris (RF), this was observed after 50% of exercise duration (p ≤ 0.05). The slope of the MF was significantly higher in the RF, followed by the VL and VM (-3.13 ± 0.52 vs. -2.61 ± 0.62 vs. -1.81 ±0.56, respectively; p < 0.05). We conclude that RF may play an important role in limiting performance during severe cycling exercise.     

Gender and muscle differences in EMG amplitude and median frequency, and variability during maximal voluntary contractions of the quadriceps femoris.

The purpose of this study was to evaluate gender and muscle differences in electromyographic (EMG) amplitude and median frequency mean and standard deviation during maximal voluntary contractions of the quadriceps femoris. Thirty recreationally active volunteers were assessed for isometric EMG activity of the vastus medialis (VM), vastus lateralis (VL), and rectus femoris (RF) muscles during three 5-s maximal isometric voluntary contractions (MVCs). Median frequency of the three muscles was assessed through a power spectral analysis (fast Fourier transformation, Hanning window processing, 512 points). The power spectral analysis was performed during the middle 3 s of each contraction over 11 consecutive, 512 ms epochs overlapping each other by half their length (256 ms). The median frequency (F(med)) for each of the 11 windows was determined for each muscle. The mean and standard deviation of the F(med) across the 11 overlapping windows were then calculated for each contraction and muscle. EMG amplitude was determined by calculating the root mean square (RMS-50 ms time constant) over the same contraction period for each muscle. The mean amplitude and standard deviation about the mean value were then determined. A three-factor ANOVA with repeated measures was performed on the calculated F(med) mean and standard deviation values, and RMS standard deviations, to assess any gender, muscle, or trial differences, or interactions. A two-factor (gender by muscle) ANOVA with repeated measures was performed on the RMS mean amplitude for each muscle. Intraclass correlation coefficients (ICCs-2,1), standard errors of measurement (SEMs), and associated 95% confidence intervals were then calculated for maximal quadriceps torque and F(med) for each muscle. The results from this study demonstrated that the VL muscle displayed significantly higher F(med) values than the RF and VM muscles. The RF muscle showed significantly higher F(med) values (mean of 11 overlapping windows) than the VM muscle. Intrasession reliability was found to be high for the calculated mean values (ICC=0.85-0.96), but was shown to be low for variability (ICC=0.13-0.45). The major findings of this study support the notion that the EMG signal is "quasi-random" in nature, as demonstrated by the reproducible F(med) means and unreliable variability.     

Comparison between voluntary and stimulated contractions of the quadriceps femoris for growth hormone response and muscle damage.

This study aimed to compare voluntary and stimulated exercise for changes in muscle strength, growth hormone (GH), blood lactate, and markers of muscle damage. Nine healthy men had two leg press exercise bouts separated by 2 wk. In the first bout, the quadriceps muscles were stimulated by biphasic rectangular pulses (75 Hz, duration 400 mus, on-off ratio 6.25-20 s) with current amplitude being consistently increased throughout 40 contractions at maximal tolerable level. In the second bout, 40 voluntary isometric contractions were performed at the same leg press force output as the first bout. Maximal voluntary isometric strength was measured before and after the bouts, and serum GH and blood lactate concentrations were measured before, during, and after exercise. Serum creatine kinase (CK) activity and muscle soreness were assessed before, immediately after, and 24, 48, and 72 h after exercise. Maximal voluntary strength decreased significantly (P < 0.05) after both bouts, but the magnitude of the decrease was significantly (P < 0.05) greater for the stimulated contractions (-22%) compared with the voluntary contractions (-9%). Increases in serum GH and lactate concentrations were significantly (P < 0.05) larger after the stimulation compared with the voluntary exercise. Increases in serum CK activity and muscle soreness were also significantly (P < 0.05) greater for the stimulation than voluntary exercise. It was concluded that a single bout of electrical stimulation exercise resulted in greater GH response and muscle damage than voluntary exercise.     

Strength training reduces force fluctuations during anisometric contractions of the quadriceps femoris muscles in old adults.

The greater fluctuations in motor output that are often exhibited by old adults can be reduced with strength training. The purpose of the study was to determine the effect of strength and steadiness training by old adults on fluctuations in force and position during voluntary contractions with the quadriceps femoris muscle. Healthy old adults (65-80 yr) completed 16 wk of heavy-load (80% of maximum, n = 11) strength training, heavy-load steadiness training (n = 6), or no training (n = 9). Steadiness training required subjects to match the angular displacement about the knee joint to a constant-velocity template. The Heavy-Load group experienced a 5.5% increase in muscle volume, a 25% increase in maximal voluntary contraction force, and a 26% increase in the one-repetition (1-RM) load. The Heavy-Load Steady group experienced increases of 11.5, 31, and 36%, respectively. The maximal electromyogram signal of quadriceps femoris increased by 51% in the two training groups. The coefficient of variation (CV) for force during submaximal isometric contractions did not change with training for any group. Although both training groups also experienced a reduction in CV for force during anisometric contractions with a 50% 1-RM load, the standard deviation of position did not change with time for any group. The Heavy-Load Steady group also experienced a reduction in CV for force during the training contractions performed with the 80% 1-RM load. Thus strength training reduced the force fluctuations of the quadriceps femoris muscles during anisometric contractions but not during isometric contractions.     

Characteristics of the muscle mechanoreflex during quadriceps contractions in humans

We examined muscle sympathetic nerve activity (MSNA) in thenonexercising lower limb during repetitive static quadricepscontraction paradigm at 25% maximal voluntary contraction in eightmen. Subjects performed 20-s contractions with 5-s rest periods for upto 12 contractions. Although the workload was constant, we found that MSNA amplitude rose as a function of contraction number [0.6 ln (amplitude/min)/contraction]; this suggests chemicalsensitization of the muscle reflex response. We employedsignal-averaging techniques and then integrated the data to examine theonset latency of the MSNA response as a function of the 25-scontraction-rest period. We observed an onset latency of ~4-6 s.Moreover, although the onset latency did not appear to vary as afunction of contraction number, the rate of MSNA increase tookapproximately four contractions to reach a steady-state rate of rise;this suggests contraction-induced sensitization. The onset latencyreported here is similar to findings in recent animal studies, but itis at odds with latencies determined in prior human handgripcontraction studies. We believe our data suggest that1) mechanically sensitive afferentscontribute importantly to the MSNA response to the paradigm employedand 2) these afferents may besensitized by the chemical products of muscle contraction.