Methodological aspects of SEMG recordings for force estimation – A tutorial and review

Insight into the magnitude of muscle forces is important in biomechanics research, for example because muscle forces are the main determinants of joint loading. Unfortunately muscle forces cannot be calculated directly and can only be measured using invasive procedures. Therefore, estimates of muscle force based on surface EMG measurements are frequently used. This review discusses the problems associated with surface EMG in muscle force estimation and the solutions that novel methodological developments provide to this problem. First, some basic aspects of muscle activity and EMG are reviewed and related to EMG amplitude estimation. The main methodological issues in EMG amplitude estimation are precision and representativeness. Lack of precision arises directly from the stochastic nature of the EMG signal as the summation of a series of randomly occurring polyphasic motor unit potentials and the resulting random constructive and destructive (phase cancellation) superimpositions. Representativeness is an issue due the structural and functional heterogeneity of muscles. Novel methods, i.e. multi-channel monopolar EMG and high-pass filtering or whitening of conventional bipolar EMG allow substantially less variable estimates of the EMG amplitude and yield better estimates of muscle force by (1) reducing effects of phase cancellation, and (2) adequate representation of the heterogeneous activity of motor units within a muscle. With such methods, highly accurate predictions of force, even of the minute force fluctuations that occur during an isometric and isotonic contraction have been achieved. For dynamic contractions, EMG-based force estimates are confounded by the effects of muscle length and contraction velocity on force producing capacity. These contractions require EMG amplitude estimates to be combined with modeling of muscle contraction dynamics to achieve valid force predictions.     

Seasonal changes in the effects of arginine vasotocin and stretch on Anolis uterine contractions in vitro

We determined the effects of acute stretch on spontaneous and arginine vasotocin (AVT)-driven contractions of the Anolis carolinensis uterus in vitro. Whole uteri from reproductively inactive females (October) were placed in a bath of oxygenated 32 degrees C Anolis "Ringer's." Two initial tensions were utilized, 1.5 g or 15 g, the latter being an estimate of the tension on the wall of a uterine compartment. Uteri were then exposed to either saline or AVT (50 ng/ml), and spontaneous or AVT-driven contractions were recorded for 20 min with the use of a strain gauge and physiograph. A similar experiment was performed on uteri from reproductively active females in the summer (June). Our results indicate that the effects of acute stretch and AVT on uterine contractility were qualitatively similar in summer and fall. That is, AVT induced a tonic contraction; stretch decreased the duration of the tonic contraction; the saline-treated uteri exhibited spontaneous rhythmic contractions; AVT increased the amplitude of the rhythmic contractions, but only at the lower tension; there were no effects of AVT on the timing (contraction interval, duration, rest interval) of the rhythmic contractions; and stretch increased the frequency of the rhythmic contractions. Season greatly influenced the magnitude of these contractile phenomena. Uteri tested during the breeding season exhibited greater distensibility, an increase in the amplitude and duration of the AVT-driven tonic contraction, and an increase in the frequency of both spontaneous and AVT-driven rhythmic contractions because of a decrease in both contraction duration and rest interval.(ABSTRACT TRUNCATED AT 250 WORDS)     

Modeling nonlinear errors in surface electromyography due to baseline noise: A new methodology

The surface electromyographic (EMG) signal is often contaminated by some degree of baseline noise. It is customary for scientists to subtract baseline noise from the measured EMG signal prior to further analyses based on the assumption that baseline noise adds linearly to the observed EMG signal. The stochastic nature of both the baseline and EMG signal, however, may invalidate this assumption. Alternately, “true” EMG signals may be either minimally or nonlinearly affected by baseline noise. This information is particularly relevant at low contraction intensities when signal-to-noise ratios (SNR) may be lowest. Thus, the purpose of this simulation study was to investigate the influence of varying levels of baseline noise (approximately 2–40% maximum EMG amplitude) on mean EMG burst amplitude and to assess the best means to account for signal noise. The simulations indicated baseline noise had minimal effects on mean EMG activity for maximum contractions, but increased nonlinearly with increasing noise levels and decreasing signal amplitudes. Thus, the simple baseline noise subtraction resulted in substantial error when estimating mean activity during low intensity EMG bursts. Conversely, correcting EMG signal as a nonlinear function of both baseline and measured signal amplitude provided highly accurate estimates of EMG amplitude. This novel nonlinear error modeling approach has potential implications for EMG signal processing, particularly when assessing co-activation of antagonist muscles or small amplitude contractions where the SNR can be low.     

Regulation of urinary bladder smooth muscle contractions by ryanodine receptors and BK and SK channels

This study examines the roles of voltage-dependent Ca(2+) channels (VDCC), ryanodine receptors (RyRs), large-conductance Ca(2+)-activated K(+) (BK) channels, and small-conductance Ca(2+)-activated K(+) (SK) channels in the regulation of phasic contractions of guinea pig urinary bladder smooth muscle (UBSM). Nisoldipine (100 nM), a dihydropyridine inhibitor of VDCC, abolished spontaneous UBSM contractions. Ryanodine (10 microM) increased contraction frequency and thereby integrated force and, in the presence of the SK blocker apamin, had a greater effect on integrated force than ryanodine alone. Blocking BK (iberiotoxin, 100 nM) or SK (apamin, 100 nM) channels increased contraction amplitude and duration but decreased frequency. The contractile response to iberiotoxin was more pronounced than to apamin. The increases in contraction amplitude and duration to apamin were substantially augmented with ryanodine pretreatment. These results indicate that BK and SK channels have prominent roles as negative feedback elements to limit UBSM contraction amplitude and duration. RyRs also appear to play a significant role as a negative feedback regulator of contraction frequency and duration, and this role is influenced by the activity of SK channels.     

Assessment of low back muscle fatigue by surface EMG signal analysis: methodological aspects.

This paper focuses on methodological issues related to surface electromyographic (EMG) signal detection from the low back muscles. In particular, we analysed (1) the characteristics (in terms of propagating components) of the signals detected from these muscles; (2) the effect of electrode location on the variables extracted from surface EMG; (3) the effect of the inter-electrode distance (IED) on the same variables; (4) the possibility of assessing fatigue during high and very low force level contractions. To address these issues, we detected single differential surface EMG signals by arrays of eight electrodes from six locations on the two sides of the spine, at the levels of the first (L1), the second (L2), and the fifth (L5) lumbar vertebra. In total, 42 surface EMG channels were acquired at the same time during both high and low force, short and long duration contractions. The main results were: (1) signal quality is poor with predominance of non-travelling components; (2) as a consequence of point (1), in the majority of the cases it is not possible to reliably estimate muscle fiber conduction velocity; (3) despite the poor signal quality, it was possible to distinguish the fatigue properties of the investigated muscles and the fatigability at different contraction levels; (4) IED affects the sensitivity of surface EMG variables to electrode location and large IEDs are suggested when spectral and amplitude analysis is performed; (5) the sensitivity of surface EMG variables to changes in electrode location is on average larger than for other muscles with less complex architecture; (6) IED influences amplitude initial values and slopes, and spectral variable initial values; (7) normalized slopes for both amplitude and spectral variables are not affected by IED and, thus, are suggested for fatigue analysis at different postures or during movement, when IED may change in different conditions (in case of separated electrodes); (8) the surface EMG technique at the global level of amplitude and spectral analysis cannot be used to characterize fatigue properties of low back muscles during very low level, long duration contractions since in these cases the non-stable MU pool has a major influence on the EMG variables. These considerations clarify issues only partially investigated in past studies. The limitations indicated above are important and should be carefully discussed when presenting surface EMG results as a means for low back muscle assessment in clinical practice.     

Comparison of electromyographic activity during eccentrically versus concentrically loaded isometric contractions

Electromyographic (EMG) amplitude and mechanical tension are directly related during isometric contraction. Maximal voluntary isometric contractions are typically elicited through two different procedures; resisting a load, which is eccentric in nature, and contracting against an immovable object, which is concentric in nature. A wealth of literature exists indicating that EMG amplitude during concentric contractions is greater than that of eccentric contractions of the same magnitude. However, the effects of different methods to elicit isometric contraction on EMG amplitude have yet to be investigated. The purpose of this study was to compare EMG amplitudes under different loading configurations designed to elicit isometric muscle contraction. Twenty healthy volunteers (10 males and 10 females, age = 23 ± 2 yrs, height = 1.7 ± 0.09 m, mass = 69.9 + 16.8 kg) performed a maximal voluntary plantarflexion effort for which the vertical ground reaction force (GRFv) sampled from a force plate and surface EMG of the soleus were recorded. Participants then performed isometric plantarflexion at 20%, 30%, 40%, and 50% GRFv_max in a seated position, from a neutral ankle position, under two different counterbalanced isometric loading conditions (concentric and eccentric). For concentric loading conditions, the subject contracted against an immovable resistance to the specified %GRFv identified via visual and auditory feedback. For eccentric loading conditions, subjects contracted against an applied load placed on the distal anterior thigh that produced the specified %GRFv. This applied load had the tendency to force the ankle into dorsiflexion. Therefore, plantarflexion force, in an attempt to maintain the ankle in a neutral position, resisted lengthening of the plantarflexor musculature, thus representing eccentric loading during an isometric contraction. Mean EMG amplitude was compared across loading levels and types using a 2 (loading type: concentric, eccentric) × 4 (loading level: 20%, 30%, 40%, 50% GRFv) repeated-measures ANOVA. The main effect for loading level was significant (p = 0.007). However, the main effect for loading type, and the loading type × loading level interaction were non-significant (p > 0.05). The present findings provide evidence that isometric muscle contractions loaded in either concentric or eccentric manners elicit similar EMG amplitudes, and are therefore comparable in research settings.     

Relationship between mechanomyogram signals and changes in force of human forefinger flexor muscles during voluntary contraction

In previous studies on mechanomyogram (MMG) signals no analysis of these signals accompanying force generation has been performed. Therefore, we have recorded MMG signals (previously referred to as muscle sound or acoustomyographic signals) during voluntary contractions of forefinger flexor muscles in 31 young subjects. These subjects made contractions to produce force records of triangular or trapeziform shape. The peak target force amounted to 10, 20 or 40 N which represented less than 40% of maximal voluntary contraction. The MMG signals during the transient phases of force generation at three different rates were analysed. The MMG intensity level calculated for MMG records and the peak-to-peak amplitude of MMG signals correlated with both the velocity of force increase and the contraction force. The occurrence of the strongest MMG signals corresponded to changes in contractile force. Therefore, it is suggested that measurements of these parameters could be a useful tool in studies of changes in contractile force. Accepted: 11 March 1998     

Recovery of the human biceps electromyogram after heavy eccentric, concentric or isometric exercise.

Five men performed submaximal isometric, concentric or eccentric contractions until exhaustion with the left arm elbow flexors at respectively 50%, 40% and 40% of the prefatigued maximal voluntary contraction force (MVC). Subsequently, and at regular intervals, the surface electromyogram (EMG) during 30-s isometric test contractions at 40% of the prefatigued MVC and the muscle performance parameters (MVC and the endurance time of an isometric endurance test at 40% prefatigued MVC) were recorded. Large differences in the surface EMG response were found after isometric or concentric exercise on the one hand and eccentric exercise on the other. Eccentric exercise evoked in two of the three EMG parameters [the EMG amplitude (root mean square) and the rate of shift of the EMG mean power frequency (MPF)] the greatest (P less than 0.001) and longest lasting (up to 7 days) response. The EMG response after isometric or concentric exercise was smaller and of shorter duration (1-2 days). The third EMG parameter, the initial MPF, had already returned to its prefatigued value at the time of the first measurement, 0.75 h after exercise. The responses of EMG amplitude and of rate of MPF shift were similar to the responses observed in the muscle performance parameters (MVC and the endurance time). Complaints of muscle soreness were most frequent and severe after the eccentric contractions. Thus, eccentric exercise evoked the greatest and longest lasting response both in the surface EMG signal and in the muscle performance parameters.     

Determinants of transpyloric fluid transport: a study using combined real-time ultrasound,manometry, and impedance recording

Intraluminal impedance recording has made it possible to record fluid transport across the pylorus during the interdigestive state without filling the stomach. During antral phase II, fluid transport occurs with and without manometrically detectable antral contraction. Our aim was to investigate the relationships between ultrasonographic patterns of antral contraction, manometric pressure waves, and transpyloric fluid transport during antral phase II. Antral wall movements were recorded by real-time ultrasound (US) in eight healthy volunteers (mean age 24 +/- 7 yr) during 17 +/- 5 min of antral phase II. Concomitantly, a catheter positioned across the pylorus, monitored by transmucosal potential difference measurement, recorded five impedance signals (1 antral, 1 pyloric, and 3 duodenal) and six manometric signals (2 antral, 1 pyloric, and 3 duodenal). Antral contractions detected by US at the level of the two antral impedance electrodes were classified according to their association with a pyloric opening or a duodenal contraction. Transpyloric liquid transport events (impedance drop of >40% of the baseline with an antegrade or retrograde propagation) and manometric pressure waves (amplitude and duration) were identified during the whole study and especially during each period of US antral contraction. A total of 110 antral contractions was detected by US. Of these, 79 were also recorded by manometry. Fluid transport across the pylorus was observed in 70.9% of the US-detected antral contractions. Pyloric opening was observed in 98.6% of the contractions associated with fluid transport compared with 50% in the absence of fluid transport (P < 0.05). Antral contractions associated with fluid transport were significantly (P < 0.05) more often propagated to the duodenum (92%) than those without fluid transport (53%). Pressure waves associated with fluid transport were of higher amplitude (208 mmHg, range 22-493) and longer duration (7 s, range 2.5-13.5 s) than those not associated with fluid transport (102 mmHg, range 18-329 mmHg, and 4.1 s, range 2-8.5 s; P < 0.05). The propagation of the antral contractions in the duodenum in US was always associated with a pyloric opening, whereas only 8 of the 25 contractions without duodenal propagation were associated with a pyloric opening (P < 0.05). The presence of duodenal contractile activity before the onset of an antral contraction in US was always accompanied by pyloric opening and with fluid transport in 93.3%, compared with 56.8% in its absence (P < 0.05). In antral phase II, US is the most sensitive technique to detect antral contractions. Transpyloric fluid transport observed in relation to antral contractions occurs mainly in association with contractions of high amplitude and long duration and is associated with pyloric opening and/or duodenal propagation.     

M-wave modulation at relative levels of maximal voluntary contraction

Frequency (mean and median power frequency, f and f _m) and amplitude (average rectified and root mean square values, ARV and rms), parameters of the M-wave, and the dorsiflexor force parameters of the anterior tibial muscles were measured in seven healthy human subjects. Intermittent, voluntary contractions at relative intensities (40%, 60%, and 80%) of maximal voluntary contraction (MVC) were performed in conjunction with electrical stimulation. The M-wave parameter changes were measured over the course of the isometric contractions. At higher force levels, M-wave potentiation was observed as increases in both ARV and rms. The ARV augmentation attained levels as high as 206.1 (SD 7.4)% of resting values after both initial and final contractions of 80% MVC, reaching statistical significance (P < 0.01). The f and f _m failed to show a significant difference at any level of contraction. It was surmised that potentiation of the M-wave was the result of an increased contribution of muscle fibre type IIb recruited during higher contraction levels, reflecting the change to larger, deeper innervating motoneurons as the intensity of contraction, as a percentage of MVC, rose. Recruitment of type IIb fibres, which have been reported to have a higher energy potential and frequency content, were thought to reflect changes in the local, excitability threshold of some motor units as the force intensity increased during the intermittent voluntary contractions. It is suggested that the M-wave elicited after contractions has the potential to reflect, to some extent, motor unit recruitment changes resulting from the preceding contractions, and that through comparisons of M-wave amplitude parameters, contributions of varying fibre types over the course of a contraction may be indicated.     

Predicting muscle force generation during fast-starts for the common carp Cyprinus carpio

Muscle contractile properties have been characterised for white myotomal muscle from the common carp Cyprinus carpio at 10, 15, and 20 °C. The time course of muscle force development was measured when one, two, or three stimuli were delivered at the onset of constant velocity shortening. As the shortening velocity increased several parameters decreased including the maximum force, the time course for the contraction and the relative duration of the deactivation compared to the activation. The maximum force and the relative rates of activation to deactivation for the contraction were relatively independent of temperature, whereas the duration of the contraction decreased with increasing temperature. A predictive model was developed which was based on fitting a modified Weibull distribution to these observations. The model was used to interpolate the expected contractile forces during cyclic length-changes. Measured and predicted values for force and power during such cyclic work-loop experiments showed an excellent agreement over the range of shortening regimes typically found during swimming behaviours. However, the predicted force was overestimated during the deactivation phase of the contractions when the shortening velocities exceeded those found during swimming. Accepted: 25 May 1999     

Reproducible measurement of voluntary activation of human elbow flexors with motor cortical stimulation.

Voluntary activation of muscle is commonly quantified by comparison of the extra force added by motor nerve stimulation during a contraction [superimposed twitch (SIT)] with that produced at rest by the same stimulus (resting twitch). An inability to achieve 100% voluntary activation implies that failure to produce maximal force output from the muscle must have occurred at a site at or above the level of the motoneurons. We have used cortical stimulation to quantify voluntary activation. Here, incomplete activation implies a failure at or above the level of motor cortical output. With cortical stimulation, it is inappropriate to compare extra force evoked during a contraction with the twitch evoked in resting muscle because motor cortical and spinal cord excitability both increase with activity. However, an appropriate "resting twitch" can be estimated. We previously estimated its amplitude by extrapolation of the linear relation between SIT amplitude and voluntary torque calculated from 35 contractions of >50% maximum (Todd G, Taylor JL, and Gandevia SC. J Physiol 551: 661-671, 2003). In this study, we improved the utility of this method to enable evaluation of voluntary activation when it may be changing over time, such as during the development of fatigue, or in patients who may be unable to perform large numbers of contractions. We have reduced the number of contractions required to only three. Estimation of the resting twitch from three contractions was reliable over time with low variability. Furthermore, its reliability and variability were similar to the resting twitch estimated from 30 contractions and to that evoked by conventional motor nerve stimulation.     

K+-induced twitch potentiation is not due to longer action potential

The objective of this study was to determine whether an increased duration of the action potential contributes to the K+-induced twitch potentiation at 37 degrees C. Twitch contractions were elicited by field stimulation, and action potentials were measured with conventional microelectrodes. For mouse extensor digitorum longus (EDL) muscle, twitch force was greater at 7-13 mM K+ than at 4.7 mM (control). For soleus muscle, twitch force potentiation was observed between 7 and 11 mM K+. Time to peak and half-relaxation time were not affected by the increase in extracellular K+ concentration in EDL muscle, whereas both parameters became significantly longer in soleus muscle. Decrease in overshoot and prolongation of the action potential duration observed at 9 and 11 mM K+ were mimicked when muscles were respectively exposed to 25 and 50 nM tetrodotoxin (TTX; used to partially block Na+ channels). Despite similar action potentials, twitch force was not potentiated by TTX. It is therefore suggested that the K+-induced potentiation of the twitch in EDL muscle is not due to a prolongation of the action potential and contraction time, whereas a longer contraction, especially the relaxation phase, may contribute to the potentiation in soleus muscle.     

Physiological characteristics of motor units in the brachioradialis muscle across fatiguing low-level isometric contractions.

The purpose of this study was to determine (i) if decomposition-based quantitative electromyography (DQEMG) could detect changes in motor unit potential (MUP) morphology and motor unit (MU) firing pattern statistics associated with muscle fatigue, (ii) if any detected changes are correlated with surface electromyographic (SEMG) signs of fatigue, and (iii) if significant fatigue-dependent changes are repeatable within individuals. Mean MU firing rates and the morphology of MUPs detected using needle and surface electrodes during constant-torque isometric contractions held until exhaustion were investigated in the brachioradialis (BR) muscle in 10 healthy volunteers (mean age=28.6 yr, SD+/-3.9). Time dependant changes were investigated using an analysis of variance with normalized time as a main effect. Partial correlation coefficients were computed using a repeated measures analysis of covariance to determine if changes in MU firing rates, needle-detected MUPs and surface-detected MUPs (SMUPs) were related to changes in SEMG signal amplitude and frequency parameters. Intraclass correlation coefficients (ICCs) were used to determine the within-subject repeatability of changes in MU firing rates, and MUP and SMUP parameters. Significant decreases in mean MU firing rates were found along with significant increases in various duration and area related parameters in both MUPs and SMUPs across the fatiguing contraction. The SEMG signal demonstrated the expected changes with fatigue: an increase in amplitude and a decrease in frequency content. SEMG amplitude was significantly positively correlated with SMUP peak-to-peak voltage (r=0.85, p<0.05), and SMUP area (r=0.86, p<0.05). Mean power frequency was significantly negatively correlated with SMUP negative peak duration (r=-0.74, p<0.05). The significant time-dependent changes were reliably observed (ICCs were 0.94 for MUP peak to peak amplitude, 0.97 for MUP area and 0.95 for MUP area to amplitude ratio, 0.95 for SMUP peak-to-peak voltage, 0.83 for SMUP area, 0.99 for SMUP negative peak amplitude and 0.88 for SMUP negative peak area). The decreases in mean MU firing rates measured along with the increases in amplitude, duration and area parameters of MUPs and SMUPs and their partial correlation with SEMG amplitude during submaximal fatiguing contractions of the BR, suggest that recruitment is a main cause of increased SEMG amplitude parameters with fatigue. We conclude that DQEMG can be effectively and reliably used to detect changes in physiological characteristics of MUs that accompany fatigue.     

Wide-pulse-width, high-frequency neuromuscular stimulation: implications for functional electrical stimulation.

Electrical stimulation (1-ms pulses, 100 Hz) produces more torque than expected from motor axon activation (extra contractions). This experiment investigates the most effective method of delivering this stimulation for neuromuscular electrical stimulation. Surface stimulation (1-ms pulses; 20 Hz for 2 s, 100 Hz for 2 s, 20 Hz for 3 s) was delivered to triceps surae and wrist flexors (muscle stimulation) and to median and tibial nerves (nerve stimulation) at two intensities. Contractions were evaluated for amplitude, consistency, and stability. Surface electromyograph was collected to assess how H-reflexes and M-waves contribute. In the triceps surae, muscle stimulation produced the largest absolute contractions (23% maximal voluntary contraction), evoked the largest extra contractions as torque increased by 412% after the 100-Hz stimulation, and was more consistent and stable compared with tibial nerve stimulation. Absolute and extra contraction amplitude, consistency, and stability of evoked wrist flexor torques were similar between stimulation types: torques reached 11% maximal voluntary contraction, and extra contractions increased torque by 161%. Extra contractions were 10 times larger in plantar flexors compared with wrist flexors with muscle stimulation but were similar with nerve stimulation. For triceps surae, H reflexes were 3.4 times larger than M waves during nerve stimulation, yet M waves were 15 times larger than H reflexes during muscle stimulation. M waves in the wrist flexors were larger than H reflexes during nerve (8.5 times) and muscle (18.5 times) stimulation. This is an initial step toward utilizing extra contractions for neuromuscular electrical stimulation and the first to demonstrate their presence in the wrist flexors.     

Objective assessment of the fusion frequency in functional electrical stimulation using the fast Fourier transform.

In functional electrical stimulation (FES) the dynamics of tetanic muscle contractions is often described by the fusion frequency (FF), as determined by palpation: contractions elicited by stimulation frequencies above the FF appear smooth. To contribute to a more objective assessment of this important FES parameter, we have developed a dedicated signal analysis method based on fast Fourier transformation (FFT). The ripple to peak ratio (R(rpFFT)) - the relation between ripple amplitude and peak force value of a recorded tetanic muscle force in relation to the applied stimulation frequency - was determined automatically by analysing a 0.2-s interval in the steady state of a stimulation burst. The method was tested on simulated data and on force recordings from isolated tibialis anterior muscles of six rabbits. The results were compared to manual estimates. The robustness of the method was tested by adding noise and hum. Simulated noise at 100% of the ripple force increased R(rpFFT) by 4%. Hum at 20 Hz away from the stimulation frequency caused changes of less than 0.5%. The results of the automated analysis of recorded signals matched the manual estimates sufficiently well, especially for stimulation frequencies near or above FF. R(rpFFT) therefore seems suitable for automated, objective and robust assessment of the ripple and the FF of electrically stimulated muscle.     

Force and EMG correlates of constant effort contractions

The relation between the force exerted by the left arm and the electromyographic (EMG) activity of the brachial biceps and triceps muscles was examined during constant effort contractions maintained for 120 s. The initial force levels were set at 35%, 50% and 65% of each subject's maximal strength, but thereafter no feedback was provided. In contrast to previous results it was found that the direction of the change in force and EMG during constant effort contractions was dependent on the level of force initially exerted. During the lowest initial force contraction the force remained constant, while for the other two force levels there was an exponential decline in the force exerted. These changes in force during the three contractions were well described by an exponential equation with two free parameters. The EMG also varied as a function of initial force. For the higher two forces the amplitude of the EMG fluctuated during the first 40 s but thereafter remained constant, while it increased steadily during the lowest initial force contraction. These results suggest that depending upon the initial level of exertion either peripheral sensory cues relating to the actual force exerted, or centrally-generated signals reflecting the magnitude of the descending motor command may be used by subjects to maintain a constant level of muscular effort.     

Recurrent inhibition of human firing motoneurons (experimental and modeling study)

Recurrent inhibition between tonically activated single human motoneurons was studied experimentally and by means of a computer simulation. Motor unit activity was recorded during weak isometric constant-force muscle contractions of brachial biceps (BB) and soleus (SOL) muscles. Three techniques (cross correlogram, frequencygram, and interspike interval analysis) were used to gauge the relations between single motor unit potential trains. Pure inhibition was detected in 5.6% of 54 BB motoneuron pairs and in 5.2% of 43 SOL motoneuron pairs. In 27.8% (BB) and 23.7% (SOL) presumed inhibition symptoms were accompanied by a synchrony peak; 37% (BB) and 48.8% (SOL) exhibited synchrony alone. The demonstrated inhibition was very weak, at the edge of detectability. Computer simulations were based on the threshold-crossing model of a tonically firing motoneuron. The model included synaptic noise as well as threshold and postsynaptic potential (PSP) amplitude change within interspike interval. Inhibition efficiency of the model neurons increased with IPSP amplitude and duration, and with increasing source firing rate. The efficiency depended on target motoneuron interspike interval in a manner similar to standard deviation of ISI. The minimum detectable amplitude estimated in the simulations was about 50V, which, compared with the experimental results, suggests that amplitudes of detectable recurrent IPSPs in human motoneurons during weak muscle contractions do not exceed this magnitude. Since recurrent inhibition is known to be progressively depressed with an increase in the force of voluntary contraction, it is concluded that the recurrent inhibition hardly plays any important role in the isometric muscle contractions of constant force.     

Catchlike-inducing train activation of human muscle during isotonic contractions: burst modulation.

Stimulation trains that exploit the catchlike property [catchlike-inducing trains (CITs)] produce greater forces and rates of rise of force than do constant-frequency trains (CFTs) during isometric contractions and isovelocity movements. This study examined the effect of CITs during isotonic contractions in healthy subjects. Knee extension was electrically elicited against a load of 10% of maximum voluntary isometric contraction. The stimulation intensity was set to produce 20% of maximum voluntary isometric contraction. The muscle was tested before and after fatigue with a 6-pulse CFT and 6-pulse CITs that contained an initial doublet, triplet, or quadruplet. For prefatigue responses, the greatest isotonic performance was produced by CITs with initial doublets. When the muscles were fatigued, triplet CITs were best. CITs produce greater excursion, work, peak power, and average power than do CFTs, because CITs produced more rapid rates of rise of force. Faster rates of rise of force enabled the preload on the muscle to be exceeded earlier during the stimulation train.