Non-invasive assessment of single motor unit mechanomyographic response and twitch force by spike-triggered averaging

A method for non-invasive assessment of single motor unit (MU) properties from electromyographic (EMG), mechanomyographic (MMG) and force signals is proposed. The method is based on the detection and classification of single MU action potentials from interference multichannel surface EMG signals and on the spike-triggered average of the MMG (detected by an accelerometer) and force signals. The first dorsal interosseous (FDI) and abductor digiti minimi (ADM) muscles were investigated at contraction levels of 2% and 5% of the maximum voluntary contraction (MVC) force. A third contraction was performed by selective activation of a single MU with surface MU action potential visual feedback provided to the subject. At 5% MVC, the mean (±standard error) single MU MMG peak-to-peak value was 11.0±1.8 mm s⁻² (N=17) and 32.3±6.5 mm s⁻² (N=20) for the FDI and AMD muscles, respectively. The peak of the twitch force was, at the same contraction livel, 7.41±1.34 mN and 14.42±2.92 mN, for the FDI and ADM muscles, respectively. The peak-to-peak value of the MMG was significantly different for the same MU at different contraction levels, indicating a non-linear summation of the single MU contributions. For the FDI muscle, the MMG peak-to-peak value of individual MUs was 21.5±7.8 mm s⁻², when such MUs were activated with visual feedback provided to the subject, whereas, for the same MUs, it was 11.8±3.8 mm s⁻², when the subject maintained a constant force level of 2% MVC. The method proposed allows the non-invasive assessment of single MU membrane and contractile properties during voluntary contractions.     

Electromyographic and mechanomyographic responses.

The purpose of the present study was to determine the electromyographic (EMG) and mechanomyographic (MMG) responses to cycle ergometry at critical power (CP). Seven moderately active males (25 +/- 3 years) completed a 60-min trial at their CP estimated from a nonlinear, 3 parameter regression model. EMG and MMG amplitudes were recorded from the vastus lateralis during 60-min continuous rides at CP. The mean CP was 175 +/- 25 W, which represented 56 +/- 5% of the subjects' peak power outputs. The results indicated that the slope coefficient for the EMG amplitude versus time relationship was not significantly different from zero; however, MMG amplitude decreased significantly over the 60 min. This dissociation between the electrical (EMG) and mechanical (MMG) aspects of muscular activity during cycle ergometry may be due to neuromuscular changes associated with "muscle wisdom" or changes in muscular compliance.     

Passive hyperthermia reduces voluntary activation and isometric force production

It has been suggested that a critically high body core temperature may impair central neuromuscular activation and cause fatigue. We investigated the effects of passive hyperthermia on maximal isometric force production (MVC) and voluntary activation (VA) to determine the relative roles of skin (T_sk) and body core temperature (T_c) on these factors. Twenty-two males [O_2max=64.2 (8.9) ml kg^–1 min^–1, body fat=8.2 (3.9)%] were seated in a knee-extension myograph, then passively heated from 37.4 to 39.4°C rectal temperature (T_re) and then cooled back to 37.4^oC using a liquid conditioning garment. Voluntary strength and VA (interpolated twitch) were examined during an isometric 10-s MVC at 0.5°C intervals during both heating and cooling. Passive heating to a T_c of 39.4^oC reduced VA by 11 (11)% and MVC by 13 (18)% (P<0.05), but rapid skin cooling, with a concomitant reduction in cardiovascular strain [percentage heart rate reserve decreased from 64 (11)% to 29 (11)%] and psychophysical strain did not restore either of these measures to baseline. Only when cooling lowered T_c back to normal did VA and MVC return to baseline (P<0.05). We conclude that an elevated T_c reduces VA during isometric MVC, and neither T_sk nor cardiovascular or psychophysical strain modulates this response. Results are given as mean (SD) unless otherwise stated.     

Relationships between voluntary activation and motor unit firing rate during maximal voluntary contractions in young and older adults

The extent to which an individual can voluntarily produce maximal muscular force can be estimated using the interpolated twitch technique. Incompleteness of activation is typically attributed to either incomplete recruitment, suboptimal firing rates or both of these mechanisms. The purpose of this study was to assess the relationship between muscle activation and maximal motor unit firing rates. Measures of muscle activation and motor unit firing rates during maximal effort contractions were obtained from 15 subjects (8 young, 7 older) throughout a 6-week strength training program for the knee extensors. High resolution interpolated twitch responses were obtained using a circuit that removes the force level of the maximal voluntary contraction prior to amplification of the additional evoked force. Maximal firing rates were obtained using a four-wire needle electrode. Electrical stimulation of the knee extensors during maximal effort contractions evoked a transient increase in the force and the amplitude of this additional force was correlated with maximal firing rates at r = -0.62 (p < 0.05). Central activation ratio and activation level, two indexes of activation, were correlated with maximal firing rates at r = 0.58 (p < 0.05) and r = 0.68 (p < 0.05), respectively. The training program elicited parallel increases in muscular activation and maximal firing rates. These results provide direct support for maximal firing rate as a significant factor limiting maximal force production.     

Hand digit control in children: motor overflow in multi-finger pressing force vector space during maximum voluntary force production

The aim of this study was to investigate the contralateral motor overflow in children during single-finger and multi-finger maximum force production tasks. Forty-five right handed children, 5–11 years of age produced maximum isometric pressing force in flexion or extension with single fingers or all four fingers of their right hand. The forces produced by individual fingers of the right and left hands were recorded and analyzed in four-dimensional finger force vector space. The results showed that increases in task (right) hand finger forces were linearly associated with non-task (left) hand finger forces. The ratio of the non-task hand finger force magnitude to the corresponding task hand finger force magnitude, termed motor overflow magnitude (MOM), was greater in extension than flexion. The index finger flexion task showed the smallest MOM values. The similarity between the directions of task hand and non-task hand finger force vectors in four-dimensional finger force vector space, termed motor overflow direction (MOD), was the greatest for index and smallest for little finger tasks. MOM of a four-finger task was greater than the sum of MOMs of single-finger tasks, and this phenomenon was termed motor overflow surplus. Contrary to previous studies, no single-finger or four-finger tasks showed significant changes of MOM or MOD with the age of children. We conclude that the contralateral motor overflow in children during finger maximum force production tasks is dependent upon the task fingers and the magnitude and direction of task finger forces.     

Longitudinal and transverse propagation of surface mechanomyographic waves generated by single motor unit activity

Multi-channel surface mechanomyographic (MMG) signals generated by individual motor units were analyzed to investigate whether the surface mechanical waves induced by fiber contraction propagate over the skin surface. The MMG signals were recorded from the tibialis anterior muscle of ten healthy subjects with 13 uniaxial accelerometers, located both along and transverse to the fiber direction. Intramuscular electromyographic signals served to identify individual motor units whose action potentials were used to trigger the averaging of the MMG signals. The spike-triggered averaged MMG had similar characteristics in locations along the longitudinal direction; however, its amplitude decreased along the transverse direction. Moreover, the time-to-positive peak increased along the transverse direction, indicating a transverse wave propagation with a velocity of 2.4 +/- 1.1 m/s in the linear direction. The results support the hypothesis that the MMG signal mainly originates from muscle fiber displacement underlining a bending mode due to contraction and provide the basis for interpreting the interference MMG in relation to motor unit activity.     

Electromyographic signal and force comparisons during maximal voluntary isometric contraction in water and on dry land

This study was designed to compare surface electromyographic (sEMG) signal and force production during maximal voluntary isometric contractions (MVCs) in water and on dry land. The reproducibility of sEMG and isometric force measurements between water and dry land environments was also assessed. Nine women performed MVC for elbow flexion and extension, hip flexion, and extension against identical fixed resistance in both environments. The sEMG signal from biceps brachii, triceps brachii, rectus femoris, and biceps femoris was recorded with waterproof adhesives placed over each electrode. The sEMG and force production showed no significant difference between water and dry land, except for HEX (p = 0.035). In addition, intraclass correlation coefficient values were significant and ranged from moderate to high (0.66–0.96) for sEMG and force production between environments. These results showed that the environment did not influence the sEMG and force in MVC.     

Relationship between motor activity-related cortical potential and voluntary muscle activation

The purpose of this study was to investigate the relationship between EEG-derived motor activity-related cortical potential (MRCP) and voluntary muscle activation. Eight healthy volunteers participated in two experimental sessions. In one session, subjects performed isometric elbow-flexion contractions at four intensity levels [10%, 35%, 60%, and 85% maximal voluntary contraction (MVC)]. In another session, a given elbow-flexion force (35% MVC) was generated at three different rates (slow, intermediate, and fast). Thirty to 40 contractions were performed at each force level or rate. EEG signals were recorded from the scalp overlying the supplementary motor area (SMA) and contralateral sensorimotor cortex, and EMG signals were recorded from the skin surface overlying the belly of the biceps brachii and brachioradialis muscles during all contractions. In each trial, the force was used as the triggering signal for MRCP averaging. MRCP amplitude was measured from the beginning to the peak of the negative slope. The magnitude of MRCP from both EEG recording locations (sensorimotor cortex and SMA) was highly correlated with elbow-flexion force, rate of rising of force, and muscle EMG signals. These results suggest that MRCP represents cortical motor commands that scale the level of muscle activation.     

Conventionally assessed voluntary activation does not represent relative voluntary torque production

The ability to voluntarily activate a muscle is commonly assessed by some variant of the twitch interpolation technique (ITT), which assumes that the stimulated force increment decreases linearly as voluntary force increases. In the present study, subjects (n = 7) with exceptional ability for maximal voluntary activation (VA) of the knee extensors were used to study the relationship between superimposed and voluntary torque. This includes very high contraction intensities (90–100%VA), which are difficult to consistently obtain in regular healthy subjects (VA of ∼90%). Subjects were tested at 30, 60, and 90° knee angles on two experimental days. At each angle, isometric knee extensions were performed with supramaximal superimposed nerve stimulation (triplet: three pulses at 300 Hz). Surface EMG signals were obtained from rectus femoris, vastus lateralis, and medialis muscles. Maximal VA was similar and very high across knee angles: 97 ± 2.3% (mean ± SD). At high contraction intensities, the increase in voluntary torque was far greater than would be expected based on the decrement of superimposed torque. When voluntary torque increased from 79.6 ± 6.1 to 100%MVC, superimposed torque decreased from 8.5 ± 2.6 to 2.8 ± 2.3% of resting triplet. Therefore, an increase in VA of 5.7% (from 91.5 ± 2.6 to 97 ± 2.3%) coincided with a much larger increase in voluntary torque (20.4 ± 6.1%MVC) and EMG (33.9 ± 6.6%max). Moreover, a conventionally assessed VA of 91.5 ± 2.6% represented a voluntary torque of only 79.6 ± 6.1%MVC. In conclusion, when maximal VA is calculated to be ∼90% (as in regular healthy subjects), this probably represents a considerable overestimation of the subjects’ ability to maximally drive their quadriceps muscles.     

运动单位数目测定在神经源性疾病中的价值

目的：探讨运动单位数目测定(MUNE)对神经肌肉疾病的诊断价值。方法：应用Viking IV型肌电图仪中的“统计学方法”对85例健康人和56例神经源性病变患者的趾短伸肌和鱼际肌进行测定。记录和刺激电极均为表面电极。结果：对照组的结果显示可重复性好,≤60岁时,运动单位(MU)数与年龄、性别和左右侧无关;＞60岁时,MU数显著减少。56例患者均表现MU数减少,可重复性高于对照组,并且随着病情的好转,功能性MU数增加。结论：MUNE是显示肌肉MU丧失的一种简单、非创伤性的快速方法,是检测神经源性疾病的理想手段之一。     

Comparison of F-waves of motor unit action potentials activated during voluntary contraction

The system for classifying F-waves was developed to study the properties of F-wave and to compare single motor unit (MU) F-waves with motor unit action potentials (MUAPs) activated during voluntary contraction. The F-waves evoked by submaximal stimulation as well as the EMG signals during voluntary contraction at 6 levels of 10-100% of maximum voluntary contraction (MVC) were measured in the tibialis anterior muscles of 3 healthy volunteers. Nine channel F-wave waveforms in a selected electrode array were classified using a template-matching method. After the detection procedure of MUAPs in voluntary EMG signals, the MUAPs were also classified by the same method. Most of the F-waves (88.4%) were composed of a single MUAP. The numbers of MU classified from single MU F-waves in 3 subjects were 12, 12 and 15, and the numbers of MU classified from the voluntary EMG signals at 6 contraction levels were 20, 27 and 24, respectively. A total of 26 single MU F-waves were identified with the MUs extracted from the data during voluntary contractions. The results suggest that the F-waves are composed of a population of the MUs, which are recruited at a wide range of contraction levels. The classification procedures of F-waves and voluntary EMG signals have made it possible to recognize the same MU in both signals and to analysis the firing thresholds of F-waves.     

Surface electromyogram spectral characterization and motor unit activity during voluntary ramp contraction in men

Summary The relationships were investigated between the surface electromyographic (SEMG) power spectrum analysed by the 20 order autoregressive model (AR spectrum) and underlying motor unit (MU) activity during isometric contractions increasing linearly from 0% to 80% maximal voluntary contraction. Intramuscular spikes and SEMG signals were recorded simultaneously from biceps brachii muscle; the former were analysed by a computer-aided intramuscular MU spike amplitude-frequency (ISAF) histogram and the latter subjected to AR spectral analysis. Results indicated that there was a positive correlation between the force output and the mean amplitude of the ISAF histogram but not with the mean frequency. These changes were accompanied by changes in relative power of the high frequency (100–200 Hz) peak (HL) in the AR spectrum. It was also found that there was a positive correlation between the mean amplitude of the ISAF histogram and the HL value. These data suggested that the power of the high frequency peak in the AR spectrum of the SEMG signal preferentially reflected the progressive recruitment of underlying MU according to their size. Differences between the AR spectrum and the spectrum estimated by fast Fourier transform algorithm have also been discussed.     

Cessation of human motor unit discharge during sustained maximal voluntary contraction.

The purpose of this study was to determine whether cessation of motor unit discharge contributes to fatigue in human subjects. Multiple fine-wire and tungsten microelectrodes were inserted into the extensor digitorum or extensor indicis muscles of the forearm in an attempt to record the activity of the same motor unit from different locations within either muscle while subjects maintained a maximal voluntary contraction of the finger extensors until force dropped by approximately 50%. The activities of 13 motor units were followed for extended periods during the fatigue task. Of these, six appeared to cease discharging prior to the end of the task, which could not be attributed to electrode movement. These findings suggest that some motor neurons may not be able to discharge continuously in the presence of sustained volitional synaptic drive or that excitatory drive may diminish during maximal voluntary effort.     

EMG activation patterns during force production in precision grip. III. Synchronisation of single motor units

Motor unit (MU) synchronisation during isometric force production in the precision grip was analysed in five subjects performing a visually guided steptracking motor task with three different force levels. With this aim multi-unit electromyographic (EMG) activity of 14 intrinsic and extrinsic finger muscles from 15 experimental sessions was decomposed into the potentials of single MUs. The behaviour of 62 intrinsic and 30 extrinsic MUs in the motor task was quantified. Most MUs displayed a positive correlation between firing rate and grip force. Compared to MUs in extrinsic muscles, intrinsic MUs had steeper regression lines with negative intercepts indicating higher force sensitivity and higher recruitment thresholds. A cross-correlation analysis was performed for 69 intra- and 166 intermuscular MU pairs while steady grip force was exerted at the three force levels. Synchronisation, for at least one force level, was found in 78% of the intra- and 45% of the intermuscular pairs. The occurrence of synchronisation was not stable over the force range tested. Factors influencing the fluctuations in occurrence and strength of synchronisation were investigated. Force increase was not paralleled by increased synchronisation; in contrast, in most MU pairs, especially intermuscular pairs, synchronisation occurred preferentially at the lower force levels. The recruitment threshold appeared to play a determining role in synchronisation: the more similar the thresholds of two MUs, the greater the probability of them being synchronised at this force level. Synchronised MUs fired on average at a lower frequency than non-synchronised ones. Finally, synchronisation at the multi-unit EMG level does not indicate that all underlying MUs are synchronised, nor does the absence of temporal coupling at the multi-unit level indicate that none of the MUs is synchronised.     

Influence of handedness on motor unit discharge properties and force tremor

Discharge properties of motor units (MUs) in the first dorsal interosseous muscle (FDI) were studied in the dominant and non-dominant hands of six right-handed (RH) and six left-handed (LH) individuals. MU discharge rates and variability were similar in each hand in RH (186 MUs) and LH (160 MUs) subjects. MU synchronization was less prominent in the dominant hand of RH subjects, with 51% (45/88) of cross-correlograms of MU discharge having significant central peaks, compared with 81% (90/111) for the non-dominant hand. The strength of MU synchronization (expressed as the frequency of extra synchronous discharges above chance) was weaker in the dominant hand of right-handers (0.23 ± 0.03 s^-1 vs 0.39 ± 0.03 s^-1), and synchronous peaks from that hand were slightly broader. Four of six RH subjects had significant differences in synchronization between hands (weaker in dominant hand). In contrast, left-handers had similar incidence (80 vs 82%, n = 161) and strength (0.41 ± 0.03 s^-1 vs 0.37 ± 0.03 s^-1) of MU synchrony in dominant and non-dominant hands. No LH subject had a significant difference in synchronization between hands. Force tremor was quantified in each hand in the same subjects during isometric abduction of FDI at 0.5 N and 3.5 N, and directly correlated with the extent of MU synchronization in the muscle. Tremor root mean square amplitude was similar in dominant and non-dominant hands. Power spectral analysis of the tremor force revealed that the peak frequency in the power spectrum was not influenced by handedness, but power at the peak frequency was higher in the non-dominant hand of RH subjects. Correlations between MU discharge variability and synchrony with measures of tremor amplitude were weak. The reduced MU synchronization in the dominant hand of right-handers may reflect a more restricted distribution of direct projections from motor cortical neurons within the FDI motoneuron pool, or reduced excitability of the cortical neurons during the task. These differences in MU synchronization, however, had an insignificant influence on the magnitude of physiological tremor in the FDI.     

Effector-specific motor activation modulates verb production

Language comprehension studies have demonstrated that effector-specific activation of the motor system supports the representation of word meaning. The aim of the present study was to test whether motor activation is also relevant for verb production. In the first part of the experiment, participants named photographs of actions either in effector-homogeneous blocks, with all actions involving the same effector, or effector-heterogeneous blocks, with actions involving different effectors. Action-naming latencies were longer in homogeneous blocks, indicating the activation of effector information. In the second part of the experiment, the same participants named action pictures in random order, while performing a motor task with either their hand or foot. The motor task caused interference for action-picture naming: latencies were longer when the effector of the depicted action was congruent with the effector of the action used in the motor task. While these results do not exclude the existence of abstract semantic representations, they indicate that effector-specific effects found in language comprehension extend to language production.     

Reductions in recruitment force thresholds in human single motor units by successive voluntary contractions

Summary Recruitment force thresholds of biceps brachii single motor units were studied in 4 male subjects before and after an isometric muscle contraction, passive muscle stretch, or following successive muscle contractions, muscle stretches or during alternations between muscle stretches and muscle contractions. Isometric muscle contractions of 5 s duration decreased subsequent single motor unit force thresholds. These force thresholds could usually be reset at or near precontraction force threshold values by passive muscle stretch induced by elbow extension. Single motor units showing reduced force thresholds following contraction were momentarily derecruited during and/or after muscle stretch. Successive muscle stretches alone did not significantly alter single motor unit force thresholds. In contrast, single motor unit recruitment force thresholds during successive weaker contractions were progressively lowered. Intercontraction muscle stretches maintained the single motor unit force thresholds at or near the initial force threshold level. The mechanism(s) underlying a muscle contraction-induced lowering of single motor unit force thresholds may reside in stretch reflex pathways.     

Independence of motor unit recruitment and rate modulation during precision force control

The vertebrate motor system chiefly employs motor unit recruitment and rate coding to modulate muscle force output. In this paper, we studied how the recruitment of new motor units altered the firing rate of already-active motor units during precision force production in the first dorsal interosseous muscle. Six healthy adults performed linearly increasing isometric voluntary contractions while motor unit activity and force output were recorded. After motor unit discharges were identified, motor unit firing rates were calculated before and after the instances of new motor unit recruitment. Three procedures were applied to compute motor unit firing rate, including the mean of a fixed number of inter-spike intervals and the constant width weighted Hanning window filter method, as well as a modified boxcar technique. In contrast to previous reports, the analysis of the firing rates of over 200 motor units revealed that reduction of the active firing rates was not a common mechanism used to accommodate the twitch force produced by the recruitment of a new motor unit. Similarly, during de-recruitment there was no tendency for motor unit firing rates to increase immediately following the cessation of activity in other motor units. Considerable consistency in recruitment behavior was observed during repeated contractions. However, firing rates during repeated contractions demonstrated considerably more fluctuation. It is concluded that the neuromuscular system does not use short-term preferential motor unit disfacilitation to effect precise regulation of muscular force output.     

Reliability of maximal muscle force and voluntary activation as markers of exercise-induced muscle damage

The loss of the ability of skeletal muscle to generate force is one of the most appropriate and valid means to quantify muscle damage. Routine measurements of maximal muscle force, however, include many potential sources of error, the most important of which may be a possible lack of central drive to the muscles. The aim of the present study was to determine the reliability of maximal isometric quadriceps muscle force and voluntary activation over a timescale that is typically employed to examine the aetiology of exercise-induced muscle damage. We also attempted to characterise the reliability of several twitch interpolation variables including the size of the interpolated twitch and the state (i.e. unpotentiated vs potentiated) and size of the resting twitch. Over a 7-day period, eight healthy active males performed repeated maximal voluntary isometric contractions (MVC) of the quadriceps (baseline and 2 h, 6 h, 24 h, 48 h, 72 h and 7 days post). Systematic variations in maximal muscle force, voluntary activation, interpolated twitch, unpotentiated twitch and potentiated twitch were not statistically significant (P>0.05) and 95% repeatability coefficients of ±76.03 N, ±4.42%, ± 8.44 N, ±25.92 N and ±43.58 N were observed, respectively. These data indicate that young healthy well-familiarised male subjects can reproduce their perceived maximal efforts both within and between days where activation levels of >90% are routinely achieved. Providing activation remains within these limits in the 7 days following an acute bout of exercise, the researcher would be 95% certain that exercise-induced muscle damage is present in individual subjects (taken from similar subject populations) if MVC force falls outside these limits.     

Correlates of measures of voluntary force with the functional state of the motor system

Oscillation spectra were analyzed for prolonged isometric force recorded in healthy subjects of three age groups. Changes in the distributions of spectral components of the oscillations in force were noted, along with differences in the distributions of spectral density as exhaustion developed in the age groups. The amplitude-frequency ranges of changes in the spectral densities of oscillations in force characterized the activity at the suprasegmental and segmental levels of the motor system which support the voluntary control and automatic regulation of posture during the performance of movements. Correlates of the functional state of the motor system are discussed in terms of the voluntary and involuntary components of control. A significant increase in activity in the central structures of the movement control system was seen with the development of exhaustion, along with decreases in the frequency range of the activity of subcortical structures with age. __________ Translated from Rossiiskii Fiziologicheskii Zhurnal imeni I. M. Sechenova, Vol. 91, No. 5, pp. 488–501, May, 2005.