EMG changes in human thenar motor units with force potentiation and fatigue

Few studies have analyzed activity-induced changes in EMG activity in individual human motor units. We studied the changes in human thenar motor unit EMG that accompany the potentiation of twitch force and fatigue of tetanic force. Single motor unit EMG and force were recorded in healthy subjects in response to selective stimulation of their motor axons within the median nerve just above the elbow. Twitches were recorded before and after a series of pulse trains delivered at frequencies that varied between 5 and 100 Hz. This stimulation induced significant increases in EMG amplitude, duration, and area. However, in relative terms, all of these EMG changes were substantially smaller than the potentiation of twitch force. Another 2 min of stimulation (13 pulses at 40 Hz each second) induced additional potentiation of EMG amplitude, duration, and area, but the tetanic force from every unit declined. Thus activity-induced changes in human thenar motor unit EMG do not indicate the alterations in force or vice versa. These data suggest that different processes underlie the changes in EMG and force that occur during human thenar motor unit activity.     

Force-frequency relationships of human thenar motor units

1. Force-frequency relationships were examined in 30 human thenar single motor units. The technique of intraneural stimulation was used to stimulate the motor axon in the median nerve proximal to the elbow with a tungsten microelectrode. 2. The stimulation consisted of either single shocks or trains of pulses (1 or 2 s duration) at constant rates varying between 5 and 100 Hz. To control various mechanical artifacts, the stimuli were delivered after electronically resetting the force baseline, and the stimuli were phased to the pulse pressure wave. Thumb flexion and abduction force components were recorded and the magnitude and direction of the resultant force calculated. Electromyographic responses (EMG) were recorded from both the proximal and distal thenar muscle surfaces. 3. For all units, twitch force began to fuse between 5 and 8 Hz and maximum tetanic force was achieved between 30 and 100 Hz. Half-maximum tetanic force was produced at 12 +/- 4 (SD) Hz, as assessed by interpolation using the nearly linear force-frequency relationship between 8 and 30 Hz (on logarithmic frequency coordinates). 4. For the majority of units (n = 19), the strongest force changes in response to variations in stimulation frequency occurred between 5 and 10 Hz (sensitivity 6 +/- 1 mN/Hz). Fewer units showed highest force-frequency sensitivity between 8 and 15 Hz (n = 7; 4 +/- 3 mN/Hz) or 10 and 20 Hz (n = 4; 5 +/- 2 mN/Hz).(ABSTRACT TRUNCATED AT 250 WORDS)     

Experimental simulation of cat electromyogram: evidence for algebraic summation of motor-unit action-potential trains.

Prompted by the observation that the slope of the relationship between average rectified electromyography (EMG) and the ensemble activation rate of a pool of motor units progressively decreased (showing a downward nonlinearity), an experimental study was carried out to test the widely held notion that the EMG is the simple algebraic sum of motor-unit action-potential trains. The experiments were performed on the cat soleus muscle under isometric conditions, using electrical stimulation of alpha-motor axons isolated in ventral root filaments. The EMG signals were simulated experimentally under conditions where the activation of nearly the entire pool of motor units or of subsets of motor units was completely controlled by the experimenter. Sets of individual motor units or of small groups of motor units were stimulated independently, using stimulation profiles that were strictly repeatable between trials. This permitted a rigorous quantitative comparison of EMGs that were recorded during combined activation of multiple motor filaments with EMGs that were synthesized from the algebraic summation of motor unit action potential trains generated by individual nerve filaments. These were recorded separately by individually stimulating the same filaments with the same activation profiles that were employed during combined stimulation. During combined activation of up to 10 motor filaments, experimentally recorded and computationally synthesized EMGs were virtually identical. This indicates that EMG signals indeed are the outcome of the simple algebraic summation of motor-unit action-potential trains generated by concurrently active motor units. For both recorded and synthesized EMGs, it was confirmed that EMG magnitude increased nonlinearly with the ensemble activation rate of a pool of motor units. The nonlinearity was largely abolished when EMG magnitude was estimated as the sum of rectified, instead of raw, motor-unit action-potential trains. This suggests that the downward nonlinearity in the EMG-ensemble activation rate relation is due to signal cancellation arising from the perfectly linear summation of positive and negative components of action-potential waveforms. The findings provide a much needed post hoc validation of the concept of EMG generation by strict algebraic summation of motor unit action potentials that is generally relied on in theoretical modeling studies of EMG and in EMG decomposition algorithms.     

Motor-unit synchronization increases EMG amplitude and decreases force steadiness of simulated contractions

The purpose of the study was to determine the effect of motor-unit synchronization on the surface electromyogram (EMG) and isometric force using a computer model of muscle contraction. The EMG and force were simulated by generating muscle fiber action potentials, defining motor-unit mechanical characteristics and territories, estimating motor-unit action potentials, specifying motor-unit discharge times, and imposing various levels of motor-unit synchronization. The output (EMG and force) was simulated at 11 levels of excitation, ranging from 5 to 100% of maximum. To synchronize motor-unit activity, selected motor-unit discharge times were adjusted; however, the number of motor units recruited and the average discharge rate of each unit was constant across synchronization conditions for a given level of excitation. Two levels of synchronization were imposed on the discharge times: a moderate and a high level, which approximated the experimentally observed range of motor-unit synchronization. The moderate level of synchrony caused the average EMG to increase by approximately 65%, whereas the high level caused a 130% increase in the EMG with respect to the no-synchrony condition. Neither synchrony condition influenced the magnitude of the average force. However, motor-unit synchronization did increase the amplitude of the fluctuations in the simulated force, especially at intermediate levels of excitation. In conclusion, motor-unit synchronization increased the amplitude of the average rectified EMG and decreased the steadiness of the force exerted by the muscle in simulated contractions.     

Twitch and tetanic properties of human thenar motor units paralyzed by chronic spinal cord injury

Little is known about how human motor units respond to chronic paralysis. Our aim was to record surface electromyographic (EMG) signals, twitch forces, and tetanic forces from paralyzed motor units in the thenar muscles of individuals (n = 12) with chronic (1.5-19 yr) cervical spinal cord injury (SCI). Each motor unit was activated by intraneural stimulation of its motor axon using single pulses and trains of pulses at frequencies between 5 and 100 Hz. Paralyzed motor units (n = 48) had small EMGs and weak tetanic forces (n = 32 units) but strong twitch forces, resulting in half-maximal force being achieved at a median of only 8 Hz. The distributions for cumulative twitch and tetanic forces also separated less for paralyzed units than for control units, indicating that increases in stimulation frequency made a smaller relative contribution to the total force output in paralyzed muscles. Paralysis also induced slowing of conduction velocities, twitch contraction times and EMG durations. However, the elevated ratios between the twitch and the tetanic forces, but not contractile speed, correlated significantly with the extent to which unit force summated in response to different frequencies of stimulation. Despite changes in the absolute values of many electrical and mechanical properties of paralyzed motor units, most of the distributions shifted uniformly relative to those of thenar units obtained from control subjects. Thus human thenar muscles paralyzed by SCI retain a population of motor units with heterogeneous contractile properties because chronic paralysis influenced all of the motor units similarly.     

Detecting the unique representation of motor-unit action potentials in the surface electromyogram

This study investigated the relative proportion of motor-unit action potentials that are uniquely represented in the simulated and experimental surface electromyogram (EMG). Two hundred motor units were simulated in a cylindrical anatomical system. Action potentials for each motor unit were generated with a model and then compared with those of other motor units. Pairs of motor units were considered indistinguishable and the motor units not uniquely represented in the surface EMG, when the difference in the mean energy for the pair of potentials was <5%. The anatomical conditions and recording configurations had a substantial influence on the percentage of motor units that could be uniquely identified in the simulated EMG. For example, a single monopolar channel could discriminate only 3.4% of motor units in the simulated population, whereas a system with 81 Laplacian channels arranged in a grid could discriminate 83.8% of the motor units under the same conditions. The simulation results were confirmed with populations of motor units recorded experimentally from the abductor digiti minimi muscle of eight healthy men. Furthermore, the relative proportion of uniquely identified motor units in the simulated signal was only moderately related to motor-unit size and distance from the electrodes. These results indicate the upper limit for detection of individual motor units from the surface EMG and show that a few channels of surface EMG recordings are not sufficient to study single motor units. The noninvasive identification of motor units from the surface EMG requires the use of multiple channels of information.     

Ordered motor-unit firing behavior in acute cerebellar stroke

It is known that at any given force level, the lower-threshold motor units generally fire at greater rates than the higher-threshold units during isometric tasks of extremity muscles. In addition to this hierarchical arrangement, firing rates of motor units fluctuate in unison with nearly no time delay; an observation that has led to the concept of common drive, a basic motoneuronal rule. Although it is established that the cerebellum plays a critical function in motor control, its role in the genesis, triggering, selection, and monitoring of motor-unit firing pattern discharges during isometric tasks is unknown. We applied an electromyographic (EMG) decomposition technique, known as precision decomposition, to accurately identify motor-unit firing times from the EMG signal recorded from the first dorsal interosseous muscle to unravel the features of motor-unit firings in three patients presenting a unilateral cerebellar stroke and exhibiting an acute cerebellar syndrome. We observed ataxic isometric force during visually guided abduction of the index finger on the affected side. However, the hierarchical response of individual motor units was spared. Furthermore, acute cerebellar ataxia was not associated with a loss of the common drive.     

Functional coupling of motor units is modulated during walking in human subjects

Time- and frequency-domain analysis of the coupling between pairs of electromyograms (EMG) recorded from leg muscles was investigated during walking in healthy human subjects. For two independent surface EMG signals from the tibialis anterior (TA) muscle, coupling estimated from coherence measurements was observed at frequencies 相似文献   

Fatigue properties of human thenar motor units paralysed by chronic spinal cord injury

Human muscles paralysed chronically by spinal cord injury (SCI) fatigue excessively. Whether these reductions in force reflect a decrease in the fatigue resistance of the motor units is unknown. Our aim was to determine the fatigability of thenar motor units paralysed chronically (10 ± 2 years) by cervical SCI. Surface electromyographic activity (EMG) and force were recorded from 17 paralysed motor units ( n = 7 subjects) in response to intraneural motor axon stimulation (13 pulses at 40 Hz, 1 s⁻¹ for 2 min). Unit force decreased progressively, reaching 8–60% of initial after 2 min, whereas both the amplitude and area of the first EMG potentials in the trains increased significantly (both P < 0.05). Thus, transmission of neural signals to the sarcolemma was effective and the reduction in force must reflect impaired processes in the muscle fibres. The median fatigue index for paralysed units (0.31), the ratio of the force at 2 min compared to the initial force, was significantly lower than that for units from control subjects (0.85, P < 0.05), but the distribution of fatigue indices for each population had a similar shape (ranges: 0.08–0.60 and 0.41–0.95, respectively). Hence, chronic paralysis did not limit the range of fatigability typically found for thenar units, only its magnitude. These findings suggest that all paralysed units underwent similar reductions in fatigue resistance. After fatigue, paralysed unit forces were reduced at all frequencies (1–100 Hz, P < 0.05). Twitch contraction and half-relaxation times were increased, as was the frequency needed to produce half maximal force ( P < 0.05). Thus, stimulation protocols used to produce functional movements in paralysed muscles need to accommodate the significant and rapid fatigue of the motor units.     

Common drive in motor units of a synergistic muscle pair

The interaction among the motor units of the extensor carpi radialis longus (ECRL) and the extensor carpi ulnaris (ECU) muscles in man was studied during wrist extensions in which the two muscles acted as synergists. Intramuscular recordings were obtained using special quadrifilar needle electrodes. Isometric wrist extensions at 20-30% of the maximal effort were studied. The electromyographic (EMG) signals were decomposed into the individual motor-unit action potential trains comprising the signal. The interaction among motor units were characterized by the estimated time-varying mean firing rate and the cross-correlation between the time-varying mean firing rates of pairs of motor units. Pairs of motor units within each muscle as well as pairs of motor units across the muscles were considered. In-phase common fluctuations, termed common drive, were observed in the mean firing rates of motor units within each muscle, consistent with earlier work on other muscles. Common fluctuations were also observed between the firing rates of ECU and ECRL motor units albeit with a variable phase shift. The existence of common drive across synergistic muscles was interpreted as implying that the CNS considers the muscles as a functional unit when they act as synergists.     

Amplitude cancellation of motor-unit action potentials in the surface electromyogram can be estimated with spike-triggered averaging

The study presents analytical, simulation, and experimental analyses of amplitude cancellation of motor-unit action potentials (APs) in the interference electromyogram (EMG) and its relation to the size of the spike-triggered average (STA) EMG. The amount of cancellation of motor-unit APs decreases monotonically as a function of the ratio between the root mean square (RMS) of the motor-unit AP and the RMS of the interference EMG signal. The theoretical derivation of this association indicates a method to measure cancellation in individual motor units by STA of the interference and squared EMGs. The theoretical relation was examined in both simulated EMG signals generated by populations of 200 motor units and experimental recordings of 492 and 184 motor-unit APs in the vastus medialis and abductor digiti minimi muscles, respectively. Although the theoretical relation predicted (R2 = 0.95; P < 0.001) the amount of cancellation in the simulated EMGs, the presence of motor-unit synchronization decreased the strength of the association for small APs. The decrease in size of the STA obtained from the squared EMG relative to that extracted from the interference EMG was predicted by the experimental measure of cancellation (R2 = 0.65; P < 0.001, for vastus medialis; R2 = 0.26; P < 0.05, for abductor digiti minimi). The results indicate that cancellation of APs in the interference EMG can be analytically predicted and experimentally measured with STA from the discharge times of the motor units into the surface EMG.     

Recording and identification of single motor units in the free-to-move primate hand

Summary A new technique is described for recording the activity of single motor units in human or monkey hand muscles. A pair of microwire electrodes is introduced into the muscle using a fine needle. After insertion, the needle can be completely removed, leaving the recording microwires in situ. The method allows stable recording of a motor unit during natural movement of the hand and fingers. The identity of a given single motor unit was reflected in the form and amplitude of the motor unit-triggered average (MU-TA), derived by averaging the unrectified surface EMG recorded from the muscle with discharges of the motor unit. The MU-TA of a given unit remained constant despite variations in the form and size of its action potential. Inspection of successive MU-TAs increased confidence that records were taken from one and the same unit over long recording periods. Control experiments in human first dorsal interosseous showed that the peak-to-peak amplitude of the MU-TA was highly correlated with both the twitch force (r= 0.65–0.92, mean 0.82, six subjects) and force threshold (r=0.62–0.93, mean 0.83) of a given unit. Similar findings were obtained for human abductor pollicis brevis (AbPB) motor units. In the monkey, AbPB motor units which were recruited early in a precision grip task and which discharged steadily during the grip had smaller MU-TAs than laterecruited, phasic units. The combination of methods described in this paper enable a single motor unit to be identified and recognised. The relative size of the unit, which is an important parameter in most motor unit studies, can be reliably estimated from the amplitude of the MU-TA. This allows indirect assessment of motor unit size in a free-to-move animal.     

The effect of muscle length on motor-unit recruitment during isometric plantar flexion in humans

The triceps surae muscle group, consisting of the mono-articular soleus (SOL) and bi-articular gastrocnemius (GAS) muscles, primarily generates plantar flexor torque. Since the GAS muscle crosses the knee joint, flexion of the knee reduces the length of this muscle, thus limiting its contribution to torque output. However, it is not clearly understood how the central nervous system activates muscles that are at inefficient or non-optimal force-producing lengths. Therefore, the present study was designed to determine the effect of muscle length on motor-unit recruitment in the medial GAS muscle. Single motor-unit activity was recorded from the medial GAS muscle while electromyographic (EMG) activity was recorded from the SOL muscle in nine male subjects. With the ankle angle held constant at 90 degrees, the knee angle was changed from 180 degrees to 90 degrees, corresponding to a long and short GAS muscle length, respectively. Levels of voluntary plantar flexor torque were produced at a rate of 2 Nm.s-1 until motor-unit activity was detected. A total of 229 motor units were recorded, of which 121 and 108 were obtained at the long and short muscle lengths, respectively. At the short length, onset of motor-unit activity occurred at significantly higher levels of plantar flexor torque and SOL EMG. Onset of motor-unit activity occurred at 2.97 +/- 7.78 Nm and 32.14 +/- 10.25 Nm, corresponding to 0.045 +/- 0.075 mV and 0.231 +/- 0.129 mV of SOL EMG in the long and short positions, respectively. No individual GAS motor unit could be recorded at both muscle lengths. Motor units in the shortened GAS muscle may be influenced by peripheral afferents capable of reducing the excitability of the motoneurone pool. This may also reflect a specific inhibition of motor units having shortened, non-optimal fascicle lengths, and they are thereby incapable of contributing to plantar flexor torque.     

Subthreshold electrical stimulation reduces motor unit discharge variability and decreases the force fluctuations of plantar flexion

The purpose of this study was to examine the influence of subthreshold electrical stimulation on the force fluctuations and motor-unit discharge variability during low-level, steady contraction of the plantar flexor muscles. Seven subjects performed a force-matching task of isometric plantar flexion at 5% of maximal voluntary contraction with and without random electrical stimulation applied to the tibial nerve. During the task, the motor unit action potential was continuously recorded with fine-wire electrodes, and the inter-spike intervals of a single motor unit were calculated. The coefficient of variation (CV) of the force fluctuations and the inter-spike intervals of the motor unit discharge were significantly decreased by the intervention of subthreshold electrical stimulation, although there were no changes in the mean values. These results suggest that subthreshold stimulation reduced the motor-unit discharge variability, which in turn, increased the steadiness of the force.     

Motor unit recruitment and derecruitment induced by brief increase in contraction amplitude of the human trapezius muscle

The activity pattern of low-threshold human trapezius motor units was examined in response to brief, voluntary increases in contraction amplitude ('EMG pulse') superimposed on a constant contraction at 4–7% of the surface electromyographic (EMG) response at maximal voluntary contraction (4–7% EMG_max). EMG pulses at 15–20% EMG_max were superimposed every minute on contractions of 5, 10, or 30 min duration. A quadrifilar fine-wire electrode recorded single motor unit activity and a surface electrode recorded simultaneously the surface EMG signal. Low-threshold motor units recruited at the start of the contraction were observed to stop firing while motor units of higher recruitment threshold stayed active. Derecruitment of a motor unit coincided with the end of an EMG pulse. The lowest-threshold motor units showed only brief silent periods. Some motor units with recruitment threshold up to 5% EMG_max higher than the constant contraction level were recruited during an EMG pulse and kept firing throughout the contraction. Following an EMG pulse, there was a marked reduction in motor unit firing rates upon return of the surface EMG signal to the constant contraction level, outlasting the EMG pulse by 4 s on average. The reduction in firing rates may serve as a trigger to induce derecruitment. We speculate that the silent periods following derecruitment may be due to deactivation of non-inactivating inward current ('plateau potentials'). The firing behaviour of trapezius motor units in these experiments may thus illustrate a mechanism and a control strategy to reduce fatigue of motor units with sustained activity patterns.     

Effect of task on the degree of synchronization of intrinsic hand muscle motor units in man.

1. Recordings were made of the firing of pairs of intrinsic hand muscle motor units active under different task conditions in man. The different tasks were defined as isometric contractions producing force in one of three different directions: finger abduction, finger extension, or finger flexion. The degree of motor-unit synchronization associated with each of these task conditions was compared with the use of cross-correlation analysis. 2. The average amount of synchronization between the firing of motor units recorded from within first dorsal interosseous muscle (IDI) was greater during index finger extension than during index finger abduction (n = 8 motor-unit pairings, 3 subjects). In addition, for another sample population of motor units, the average amount of synchronization was greater during index finger abduction than during index finger flexion (n = 11 motor-unit pairings, 4 subjects). 3. In a further series of experiments, one motor unit of each pair was recorded from second dorsal interosseous muscle (2DI), whereas the other motor unit of each pair was recorded from 1DI. The average amount of synchronization for these motor-unit pairings was greater during extension of the index and middle fingers than during abduction of the index and middle fingers (n = 8, 4 subjects). For another sample population of such motor-unit pairings, the average amount of synchronization was found to be greater during abduction of the index and middle fingers than during flexion of the index and middle fingers (n = 11, 4 subjects). 4. In approximately one-third of cases, it was not found possible to maintain the same firing rates from two motor units in 1DI when active under different task conditions. For instance, the "reference" motor unit might consistently fire at a faster rate than the "response" motor unit when active during index finger extension but consistently fire at a slower rate than the response motor unit when active during index finger abduction. Where such motor-unit pairs have been studied in detail, the pattern of task dependence in their synchronization was found to be similar to that described above for motor-unit pairs in which the firing rates remained constant under the different task conditions.(ABSTRACT TRUNCATED AT 400 WORDS)     

Motor-unit synchronization alters spike-triggered average force in simulated contractions

The purpose of the study was to quantify the effect of motor-unit synchronization on the spike-triggered average forces of a population of motor units. Muscle force was simulated by defining mechanical and activation characteristics of the motor units, specifying motor neuron discharge times, and imposing various levels of motor-unit synchronization. The model comprised 120 motor units. Simulations were performed for motor units 5-120 to compare the spike-triggered average responses in the presence and absence of motor-unit synchronization with the motor-unit twitch characteristics defined in the model. To synchronize motor-unit activity, selected motor-unit discharge times were adjusted; this kept the number of action potentials constant across the three levels of synchrony for each motor unit. Because there was some overlap of motor-unit twitches even at minimal discharge rates, the simulations indicated that spike-triggered averaging underestimates the twitch force of all motor units and the contraction time of motor units with contraction times longer than 49 ms. Although motor-unit synchronization increased the estimated twitch force and decreased the estimated contraction time of all motor units, spike-triggered average force changed systematically with the level of synchrony in motor units 59-120 (upper 90% of the range of twitch forces). However, the reduction in contraction time was similar for moderate and high synchrony. In conclusion, spike-triggered averaging appears to provide a biased estimate of the distribution of twitch properties for a population of motor units because twitch fusion causes an underestimation of twitch force for slow units and motor-unit synchronization causes an overestimation of force for fast motor units.     

Pattern of pulses that maximize force output from single human thenar motor units

We assessed the sequence of nerve impulses that maximize force output from individual human thenar motor units. When these motor units were stimulated intraneurally by a variable sequence of seven pulses, the pattern of pulses that elicited maximum force always started with a short (5-15 ms) interpulse interval termed a "doublet. " The twitch force summation caused by this "doublet" elicited, on average, 48 +/- 13% (SD) of the maximum tetanic force. The peak amplitude of "doublet" forces was 3.5 times that of the initial twitches, and twitch potentiation appeared to have little influence on twitch force summation elicited by the "doublets." For some units, the second optimal interpulse interval was also short. Peak forces elicited by the third to sixth interpulse intervals did not change substantially when the last interpulse interval was varied between 5 to 55 ms, so maximum force could not be attributed to any unique interpulse interval. Each successive pulse contributed a smaller force increment. When five to seven pulses were delivered in an optimal sequence, the evoked force was close to that recorded during maximal tetanic stimulation. In contrast, maximal force-time integral was evoked with one short interpulse interval (5-15 ms) then substantially longer interpulse intervals (>100 ms). Maximum force and force-time integrals were therefore elicited by different patterns of stimuli. We conclude that a brief initial interpulse interval (5-15 ms) is required to elicit maximum "doublet" force from human thenar motor units and that near-maximal tetanic forces can be elicited by only five or six additional post-"doublet" pulses if appropriately spaced in time. However, the rate at which these post-"doublet" stimuli must be provided is fairly uncritical. In contrast, maximum post-"doublet" force-time integrals were obtained at intervals corresponding to motoneuronal firing rates of approximately 7 Hz, rates close to that typically used to recruit motor units and to maintain weak voluntary contractions.     

Motor unit conduction velocity during sustained contraction of the vastus medialis muscle

The aim of the study was to analyze motor unit conduction velocity at varying force of the vastus medialis muscle during sustained contraction. Surface (8-electrode array) and intramuscular (two wire electrodes) EMG signals were recorded from the distal part of the dominant vastus medialis muscle of ten healthy male subjects. The subjects sat on a chair with the knee 90° flexed and performed seven 180-s long contractions at forces in the range 2.5–30% of the maximal voluntary contraction force. For each force level, the discharge patterns of the newly recruited motor units with respect to the previous force level were identified from the intramuscular recordings and used as trigger for averaging the surface EMG signals. Motor unit conduction velocity was estimated from the averaged surface EMG. Average discharge rate at which motor units were analyzed was the same for each force level (mean ± SD, 8.3 ± 0.8 pulses per second). Motor unit conduction velocity at the beginning of the contraction and its rate of change over time increased with force (P < 0.05). Conduction velocity at the beginning of the contraction estimated from the interference surface EMG (4.44 ± 0.66 m/s) and from single motor units (4.75 ± 0.56 m/s) were positively correlated (R ² = 0.46; P < 0.0001) but significantly different (P < 0.05). The results indicate that single motor unit conduction velocity and its rate of change during sustained contraction, assessed at a fixed discharge rate, depend on force level.     

Entrainment of motor-unit discharges as a neuronal mechanism of synchronization.

The neuronal mechanism which gives rise to the synchronization of motor-unit discharges has been inferred from an analysis of the interspike intervals of individual motor-unit discharges recorded from the soleus muscle. The motor units were divided into two groups on the basis of characteristic changes in their spike trains. The first group maintained a stationary discharge pattern throughout the process of synchronization with a firing rate of approximately 10 spikes/s. Small unidentified units simultaneously recorded gradually grouped around the individual spikes of the first group motor unit and with this process, high-frequency force oscillation appeared phase-locked with each of grouped discharges. The mean period of force oscillation was almost identical to the mean discharge interval. Therefore, the first group motor unit was considered as a pacemaker of this force-oscillation. The second group motor unit underwent from its initially stationary process to a transitional process characterized by spike dropouts from an otherwise regular spike train. When both groups of motor units were recorded by the same electrode, it was found that the firing rate of the second group motor unit discharges gradually approached that of the first group, and the spikes of the first and the second group motor units occurred near or at the same time. The number of double intervals decreased in a highly predictable fashion with an increase in a firing rate. It was furthermore observed that the spikes of a given motor unit whose discharges interval-to-period ratio is smaller at the beginning of transitional process was entrained to the first group motor-unit discharges with a faster time course than the unit whose discharge interval-to-period ratio is larger. The synchronizing process was described from the relations between shorter discharge interval-to-period ratios and the longer-to-shorter interval ratios obtained at several stages from the beginning of transitional process to the final synchronization. Their relations were best drawn by the second-order regression lines. The faster time course of synchronization was reflected in the larger value of coefficient a in the equation. The results of this and previous study (23) further provided evidence to justify that interaction of motor-unit discharges is responsible for the synchronization. Although the neuronal limiting device of the firing-rate control to approximately 10 spikes/s still remains unsolved, the possibility was considered that a disinhibitory neuronal network first acts to synchronize independently firing motoneurons and leads to the oscillation of the stretch reflex loop. This closed-loop system was considered as a site for the stored motor program and the use of disinhibitory neuronal network was discussed in relation to the Harmon's model of neuromimes.