Motor Unit Discharge Frequency Research Articles

Real-time finger force prediction via parallel convolutional neural networks: a preliminary study.

Continuous and accurate decoding of intended motions is critical for human-machine interactions. Here, we developed a novel approach for real-time continuous prediction of forces in individual fingers using parallel convolutional neural networks (CNNs). We extracted populational motor unit discharge frequency using CNNs in a parallel structure without spike sorting. The CNN parameters were trained based on two features from high-density electromyogram (HD-EMG), namely temporal energy heatmaps and frequency spectrum maps. The populational motor unit discharge frequency was then used to continuously predict finger forces based on a linear regression model. The force prediction performance was compared with a motor unit decomposition method and the conventional EMG amplitude-based method. Our results showed that the correlation coefficient between the predicted and the recorded forces of the CNN approach was on average 0.91, compared with the offline decomposition method of 0.89, the online decomposition method of 0.82, and the EMG amplitude method of 0.81. Additionally, the CNN based approach showed generalizable performance, with CNN trained on one finger applicable to a different finger. The outcomes suggest that our CNN based algorithm can offer an accurate and efficient force decoding method for human-machine interactions.

Causes of, and countermeasures to, increased ventilatory drive in tetraplegia.

Causes of, and countermeasures to, increased ventilatory drive in tetraplegia.

Increased diaphragm motor unit discharge frequencies during quiet breathing in people with chronic tetraplegia.

Respiratory muscle strength is compromised in people with tetraplegia, which may be compensated for by an increase in neural drive to the diaphragm. We found that the discharge frequencies of diaphragm motor units are higher in people with chronic tetraplegia compared with able-bodied people during quiet breathing. Furthermore, we found that the area of single motor unit potentials was increased in people with tetraplegia. These results suggest an increased motoneurone output to the diaphragm and remodelling of diaphragm motor units to maintain ventilation in tetraplegia. People with tetraplegia have reduced inspiratory muscle strength, ∼40% of able-bodied individuals. Paralysed or partially paralysed respiratory muscles as a result of tetraplegia compromise lung function, increase the incidence of respiratory infections and can cause dyspnoea. We hypothesised that reduced inspiratory muscle strength in tetraplegia may increase neural drive to the inspiratory muscles to maintain ventilation. We recorded the discharge properties of single motor units from the diaphragm in participants with chronic tetraplegia (8 males, 42-78 years, C3-C6 injury, AIS A-C) and able-bodied control participants (6 males matched for age and body mass index). In each group, 117 and 166 single motor units, respectively, were discriminated from recordings in the costal diaphragm using a monopolar electrode. A linear mixed-effects model analysis showed higher peak discharge frequencies of motor units during quiet breathing in tetraplegia (17.8 ± 4.9Hz; mean ± SD) compared with controls (12.4 ± 2.2Hz) (P < 0.001). There were no differences in tidal volume, inspiratory time or mean air flow between groups. Motor unit potentials in tetraplegia, compared with controls, were larger in amplitude (1.1 ± 0.7mV and 0.5 ± 0.3mV, respectively, P = 0.007) and area (1.83 ± 1.49µV ms and 0.69 ± 0.52µV ms, respectively, P = 0.003). The findings indicate that diaphragm motor unit remodelling is likely to have occurred in people with chronic tetraplegia and that there is an increase in diaphragm motor unit discharge rates during quiet breathing. These neural changes ensure that ventilation is maintained in people with chronic tetraplegia.

Genioglossus motor unit activity in supine and upright postures in obstructive sleep apnea.

This study investigated whether a change in posture affected the activity of the upper-airway dilator muscle genioglossus in participants with and without obstructive sleep apnea (OSA). During wakefulness, a monopolar needle electrode was used to record single motor unit activity in genioglossus in supine and upright positions to alter the gravitational load that causes narrowing of the upper airway. Activity from 472 motor units was recorded during quiet breathing in 17 males, nine of whom had OSA. The mean number of motor units for each participant was 11.8 (SD 3.4) in the upright and 16.0 (SD 4.2) in the supine posture. For respiratory-modulated motor units, there were no significant differences in discharge frequencies between healthy controls and participants with OSA. Within each breath, genioglossus activity increased through the recruitment of phasic motor units and an increase in firing rate, with an overall increase of ~6 Hz (50%) across both postures and participant groups. However, the supine posture did not lead to compensatory increases in the peak discharge frequencies of inspiratory and expiratory motor units, despite the increase in gravitational load on the upper airway. Posture also had no significant effect on the discharge frequency of motor units that showed no respiratory modulation during quiet breathing. We postulate that, in wakefulness, any increase in genioglossus activity to compensate for the gravitational effects on the upper airway is achieved primarily through the recruitment of additional motor units in both healthy controls and participants with OSA.

Absence of hyperexcitability of spinal motoneurons in patients with amyotrophic lateral sclerosis.

•Amyotrophic lateral sclerosis (ALS) motoneurons become hypoexcitable with disease progression in experimental models, raising questions about the neural hyperexcitability supported by clinical observations. •A variant of the ∆F method, based on motor unit discharge frequency modulations during recruitment and derecruitment, has been developed to investigate the motoneuron capacity to self-sustained discharge in patients. •The modulation of motor unit firing rate during ramp contraction and vibration-induced recruitment are modified in ALS, suggesting lower motoneuron capacity to self-sustained discharge, which is a sign of hypoexcitability. •∆F-D decreases with functional impairment and its reduction is more pronounced in fast progressors. •In patients with ALS, motoneurons exhibit hypoexcitability, which increases with disease progression. Experimental models have primarily revealed spinal motoneuron hypoexcitability in amyotrophic lateral sclerosis (ALS), which is contentious considering the role of glutamate-induced excitotoxicity in neurodegeneration and clinical features rather supporting hyperexcitability. This phenomenon was evaluated in human patients by investigating changes in motor unit firing during contraction and relaxation. Twenty-two ALS patients with subtle motor deficits and 28 controls performed tonic contractions of extensor carpi radialis, triceps brachialis, tibialis anterior and quadriceps, aiming to isolate a low-threshold unit (U1) on the electromyogram (EMG). Subsequently, they performed a stronger contraction or tendon vibration was delivered, to recruit higher threshold unit (U2) for 10s before they relaxed progressively. EMG and motor unit potential analyses suggest altered neuromuscular function in all muscles, including those with normal strength (Medical Research Council score at 5). During the preconditioning tonic phase, U1 discharge frequency did not differ significantly between groups. During recruitment, the increase in U1 frequency (∆F-R) was comparable between groups both during contraction and tendon vibration. During derecruitment, the decrease in U1 frequency (∆F-D) was reduced in ALS regardless of the recruitment mode, particularly for ∆F-R<8Hz in the upper limbs, consistent with the muscle weakness profile of the group. ∆F-D was associated with functional disability and its reduction was more pronounced in patients with more rapid disease progression rate. This in vivo study has demonstrated reduced motoneuron capacity for self-sustained discharge, and further supports that motoneurons are normo- to hypoexcitable in ALS patients, similar to observations in experimental models.

Comparisons of voluntary and evoked rate of torque development and rate of velocity development during isokinetic muscle actions

BACKGROUND: The rate of torque development (RTD) and the rate of velocity development (RVD) have previously been described as related; however, a direct comparison has not been performed. OBJECTIVE: The purposes of this study were to compare voluntary and evoked RVD and RTD during the same maximal isokinetic leg extensions muscle actions and to indirectly explore the influence of motor unit discharge frequency on these variables. METHODS: Sixteen men completed three maximal voluntary and three maximal evoked isokinetic leg extension muscle actions at 60 ◦ ·s −1 . Peak RVD, general RVD, peak RTD, and electromechanical delay (EMD) were calculated from the voluntary and evoked muscle actions. Voluntary and evoked RTD and RVD were also calculated for each 10 ms epoch up to 200 ms of the muscle actions. RESULTS:There was no interaction between voluntary and evoked RVD across time (p = 0.12), but there was an interaction for RTD ( p< 0.01). However, peak RTD occurred prior to the isokinetic load range. Peak RTD ( p< 0.001), peak RVD ( p< 0.01), general RVD ( p< 0.01), isokinetic load range ( p< 0.001), EMD ( p< 0.001), and PT ( p< 0.001) were greater for voluntary than evoked muscle actions, which was expected due to the influence of voluntary motor unit firing rates. CONCLUSIONS: Overall, these results suggested that the calculation of RTD during the acceleration phase of an isokinetic muscle action may not be valid due to the unknown load and increasing velocity. Furthermore, the RVD may be influenced by motor unit firing rate, but to a lesser extent than RTD.

Distribution of electrical activation to the external intercostal muscles during high frequency spinal cord stimulation in dogs

In contrast to previous methods of electrical stimulation of the inspiratory muscles, high frequency spinal cord stimulation (HF-SCS) results in more physiological activation of these muscles. The spatial distribution of activation to the external intercostal muscles by this method is unknown. In anaesthetized dogs, multiunit and single motor unit (SMU) EMG activity was monitored in the dorsal portion of the 3rd, 5th and 7th interspaces and ventral portion of the 3rd interspace during spontaneous breathing and HF-SCS following C2 spinal section. Stimulus amplitude during HF-SCS was adjusted such that inspired volumes matched spontaneous breathing (Protocol 1). During HF-SCS, mean peak SMU firing frequency was highest in the 3rd interspace (dorsal) (18.8 ± 0.3 Hz) and significantly lower in the 3rd interspace (ventral) (12.2 ± 0.2 Hz) and 5th interspace (dorsal) (15.3 ± 0.3 Hz) (P <0.05 for each comparison). Similar rostrocaudal and dorsoventral gradients of activity were observed during spontaneous breathing prior to C2 section. No significant activity was observed in the 7th interspace during either spontaneous breathing or HF-SCS. Since peak discharge frequencies of the SMUs were higher and rib cage movement greater during HF-SCS compared to spontaneous breathing, stimulus amplitude during HF-SCS was adjusted such that rib cage movement matched (Protocol 2). Under these conditions, mean peak SMU frequencies and rostrocaudal and dorsoventral gradients of activity during HF-SCS were not significantly different compared to spontaneous breathing. These results indicate that (a) the topographic pattern of electrical activation of the external intercostal muscles during HF-SCS is similar to that occurring during spontaneous breathing and (b) differential descending synaptic input from supraspinal centres is not a required component of the differential spatial distribution of external intercostal muscle activation. HF-SCS may provide a more physiological method of inspiratory muscle pacing.

Interplay Between the Inspiratory and Postural Functions of the Human Parasternal Intercostal Muscles

The parasternal intercostal muscles are obligatory inspiratory muscles. To test the hypothesis that they are also involved in trunk rotation and to assess the effect of any postural role on inspiratory drive to the muscles, intramuscular electromyographic (EMG) recordings were made from the parasternal intercostals on the right side in six healthy subjects during resting breathing in a neutral posture ("neutral breaths"), during an isometric axial rotation effort of the trunk to the right ("ipsilateral rotation") or left ("contralateral rotation"), and during resting breathing with the trunk rotated. The parasternal intercostals were commonly active during ipsilateral rotation but were consistently silent during contralateral rotation. In addition, with ipsilateral rotation, peak parasternal inspiratory activity was 201 +/- 19% (mean +/- SE) of the peak inspiratory activity in neutral breaths (P < 0.001), and activity commenced earlier relative to the onset of inspiratory flow. These changes resulted from an increase in the discharge frequency of motor units (14.3 +/- 0.3 vs. 11.0 +/- 0.3 Hz; P < 0.001) and the recruitment of new motor units. The majority of units that discharged during ipsilateral rotation were also active in inspiration. However, with contralateral rotation, parasternal inspiratory activity was delayed relative to the onset of inspiratory flow, and peak activity was reduced to 72 +/- 4% of that in neutral breaths (P < 0.001). This decrease resulted from a decrease in the inspiratory discharge frequency of units (10.5 +/- 0.2 vs. 12.0 +/- 0.2 Hz; P < 0.001) and the derecruitment of units. These observations confirm that in addition to an inspiratory function, the parasternal intercostal muscles have a postural function. Furthermore the postural and inspiratory drives depolarize the same motoneurons, and the postural contraction of the muscles alters their output during inspiration in a direction-dependent manner.

Skeletal Muscle Fatigue

The article by Jane Kent-Braun, Ph.D., FACSM, (6) combines a muscular component of aging with a neural component to hypothesize a silver lining - as people age, their muscles get weaker, but also less fatigable. The muscular component combines consistent evidence of a shift in fiber-type distribution with age and her own data showing reduced accumulation of H+ and Pi in aged compared with younger subjects' muscles during successive maximum voluntary contractions to make a compelling case for increased metabolic economy of aged muscle. To support the neural component, she reviews a small but consistent literature reporting reduced maximal motor unit discharge rates in aged subjects and hypothesizes a consequent reduced metabolic demand. Together, these mechanisms are hypothesized to increase metabolic economy and decrease accumulation of metabolic products, an adaptation that decreases fatigability in aged muscle. The literature reviewed in (6) comparing fatigability of younger with that of older people's muscles, shows mixed results attributed in this article to dynamic versus static fatiguing protocols. It is known that fatigue can be caused by many different mechanisms. Kent-Braun's technique identifies fatigue that is associated with less acidosis and lower accumulation of Pi and H2PO4-. Cairns et al. (2) pointed out that various protocols might selectively stress different mechanisms among the manifold possible causes of fatigue, the protocol effectively selecting the mechanism of fatigue. In this context, it is important to ask whether protocols that select for fatigue caused by accumulation of metabolites are the most appropriate to study fatigue mechanisms relevant to problems associated with aging. We also suggest that the reduction in maximal motor unit discharge frequency among aged subjects may not be a neural adaptation to aging, but rather a corollary to the selective cell death of larger motor neurons that occurs with aging. Motor neurons serving Type I muscle units have lower discharge rates, increased metabolic economy because of both less calcium pumping and lower myosin ATPase rates (1,8), less production of metabolic products, less sensitivity to metabolic products (4), and lower susceptibility to deficits in excitation-contraction coupling (7). We therefore suggest an alternative hypothesis: a larger proportion of Type I motor unit cross-sectional area confers resistance to some important types of fatigue. Our hypothesis is consistent with that of Kent-Braun (6), but may make the age advantage also a gender advantage, and a general advantage to all of us with slower fiber-type distributions - likely a large population, as the variance in fiber-type distribution in human populations is very large (5). Type I fibers have long been recognized as less fatigable than Type II under a restricted definition of fatigue. Future studies might relax this restriction. There are emerging experimental approaches that focus on identifying mechanisms of fatigue that are task specific (3). Selected comparisons of tasks, groups of individuals, and interventions with a task-failure approach have potential to delve into the mechanisms that limit performance of many different activities of daily living, providing a translational approach. Steven L. Lehman Department of Integrative Biology University of California-Berkeley Berkeley, CA Ladora V. Thompson Department of Physical Medicine and Rehabilitations University of Minnesota Minneapolis, MN

Age-related decline in rate of torque development is accompanied by lower maximal motor unit discharge frequency during fast contractions

The aim of this study was to investigate the association between the rate of torque development and maximal motor unit discharge frequency in young and elderly adults as they performed rapid submaximal contractions with the ankle dorsiflexors. Recordings were obtained of the torque exerted by the dorsiflexors during the isometric contractions and the surface and intramuscular electromyograms (EMGs) from the tibialis anterior. The maximal rate of torque development and integrated EMG (percentage of total EMG burst) at peak rate of torque development during fast contractions were lower in elderly than young adults by 48% (P < 0.05) and 16.5% (P < 0.05), respectively. The young adults, but not the elderly adults, exhibited a positive association (r2 = 0.33; P < 0.01) between the integrated EMG computed up to the peak rate of torque development and the maximal rate of torque development achieved during the fast contractions. These age-related changes during fast voluntary contractions were accompanied by a decline (P < 0.001) in motor unit discharge frequency (19, 28, and 34% for first 3 interspike intervals, respectively) and in the percentage of units (45%; P < 0.05) that exhibited double discharges (doublets) at brief intervals (<5 ms). Because aging decreased the maximal rate of torque development of fast voluntary contractions to a greater extent ( approximately 10%) than that of an electrically evoked twitch, collectively the results indicate that the age-related decline in maximal motor unit discharge frequency likely limit, in addition to the slowing of muscle contractile properties, the performance of fast voluntary contractions.

Slower conduction velocity and motor unit discharge frequency are associated with muscle fatigue during isometric exercise in type 1 diabetes mellitus

Type 1 diabetes mellitus (T1DM) is associated with a peripheral neuropathy that reduces nerve conduction velocity. This may impair high motor-unit discharge frequencies (MUDF), decrease muscle activation, and curtail the ability to sustain repetitive contractile tasks. We examined (1) whether MUDF, the contractile properties of the knee extensors, and the conduction velocity of persons with T1DM differed from controls; (2) whether persons with T1DM can maintain adequate MUDF during a fatigue protocol; and (3) the relationship between these parameters and impaired glycemic control. We studied male and female subjects with T1DM and controls matched for age, height, weight, and gender. Single motor unit recordings were made from vastus lateralis during maximal and submaximal contractions and during a fatigue protocol. Glycemic control was assessed from blood glucose concentration and glycosylated hemoglobin (HbA1c). Control femoral conduction velocities were comparable to literature values and those of the T1DM subjects were slower. These values correlated with plasma glucose and HbA1c. T1DM subjects fatigued 45% sooner than controls, and time to fatigue and conduction velocity were correlated (r = 0.54, P < 0.05). Discharge frequencies tended to be slower during 50% maximal voluntary contractile force in the T1DM subjects at task failure. Persons with T1DM had slower conduction velocities and lower MUDF than their controls, which apparently leads to impaired activation of muscle and decreased endurance during isometric fatigue.

Spatial distribution of inspiratory drive to the parasternal intercostal muscles in humans

The human parasternal intercostal muscles are obligatory inspiratory muscles with a diminishing mechanical advantage from cranial to caudal interspaces. This study determined whether inspiratory neural drive to these muscles is graded, and whether this distribution matches regional differences in inspiratory mechanical advantage. To determine the neural drive, intramuscular EMG was recorded from the first to the fifth parasternal intercostals during resting breathing in six subjects. All interspaces showed phasic inspiratory activity but the onset of activity relative to inspiratory flow in the fourth and fifth spaces was delayed compared with that in cranial interspaces. Activity in the first, second and third interspaces commenced, on average, within the first 10% of inspiratory time, and sometimes preceded inspiratory airflow. In contrast, activity in the fourth and fifth interspaces began after an average 33% of inspiratory time. The peak inspiratory discharge frequency of motor units in the first interspace averaged 13.4 +/- 1.0 Hz (mean +/- s.e.m.) and was significantly greater than in all other interspaces, in particular in the fifth space (8.0 +/- 1.0 Hz). Phasic inspiratory activity was sometimes superimposed on tonic activity. In the first interspace, only 3% of units had tonic firing, but this proportion increased to 34% in the fifth space. In five subjects, recordings were also made from the medial and lateral extent of the second parasternal intercostal. Both portions showed phasic inspiratory activity which began within the first 6% of inspiratory time. Motor units from the lateral and medial portions fired at the same peak discharge rate (10.4 +/- 0.7 versus 10.7 +/- 0.6 Hz). These observations indicate that the distribution of neural drive to the parasternal intercostals in humans has a rostrocaudal gradient, but that the drive is uniform along the mediolateral extent of the second interspace. The distribution of inspiratory neural drive to the parasternal intercostals parallels the spatial distribution of inspiratory mechanical advantage, while tonic activity was higher where mechanical advantage was lower.

Responses of constantly firing motor units to afferent stimulation.

Constantly firing motor units of the short abductor muscles of the first finger in the human hand and the abdominal wall muscle in immobilized rats responded to afferent stimulation of the median and sciatic nerves respectively with changes in the nature of spike activity. In the first 250 msec of the post-stimulus period, the frequency of motor unit spike activity became unstable and peri-stimulus histograms were individually quite distinct. This was followed by relative stabilization of motor unit discharge frequencies, and the subsequently (750 msec) determined motor unit spike frequency depended on most cases on the background spike frequency.

Differential activation of motor units in the wrist extensor muscles during the tonic vibration reflex in man.

1. Single motor unit activity was recorded in the extensor carpi radialis longus and extensor carpi radialis brevis muscles of five healthy human subjects, using metal microelectrodes. 2. Motor units were characterized on the basis of their twitch contraction times and their force recruitment thresholds during voluntary imposed-ramp contractions. 3. The discharge patterns of forty-three motor units were studied during tonic vibration reflex elicited by prolonged (150 s) trains of vibration (30 Hz) applied to the distal tendons of the muscles. The temporal relationships between the individual small tendon taps of the vibratory stimulus and the motor unit impulses were analysed on dot raster displays and post-stimulus time histograms. 4. After tendon taps, the impulses of motor units with long twitch contraction times (mean +/- S.D., 47.2 +/- 10.7 ms) and low recruitment thresholds (0.88 +/- 0.6 N) formed a single narrow peak (P1) with a latency (22.7 +/- 1.4 ms) which was comparable to that of the tendon jerk in the extensor carpi radialis muscles. These motor units were named 'P1 units'. On the other hand, the response of motor units with shorter twitch contraction times (31.1 +/- 3.3 ms) and higher recruitment thresholds (3.21 +/- 1.3 N) showed two peaks: a short latency (23.4 +/- 1.3 ms) P1 peak similar to the previous one and a P2 peak occurring 9.4 +/- 1.2 ms later. These motor units were named 'P1-P2 units'. 5. When the reflex contraction increased slowly, the P1 peaks of 'P1-P2 units' were clearly predominant at the beginning of the contraction, during the rising phase of the motor unit discharge frequency, while the P2 peaks became predominant when the units had reached their maximal discharge frequency. 6. Increasing the tendon vibration frequency (35, 55, 75, 95 Hz) did not modify the 'P1 unit' discharge pattern. Due to interference between vibration period and peak latencies, increasing the vibration frequency caused the P1 and P2 peaks of 'P1-P2 units' to overlap. 7. Superficial cutaneous stimulation of the dorsal side of the forearm during tendon vibration noticeably decreased the P1 peaks in both types of motor units. In the P2 peaks it could result in either a decrease or an increase but the average effect was a slight increase. 8. When applied 10 s before tendon vibration, cutaneous stimulation considerably suppressed the tonic vibration reflex.(ABSTRACT TRUNCATED AT 400 WORDS)

こう筋における持続的張力発現のメカニズム

The purpose of this paper is to investigate the motor unit activities of the human masseter muscle during sustained the bite force at a constant level. The electrical activities recorded with surface and inserted electrodes were studied, with the following results. 1. The masseter muscle had the changes of activities in two phases as a contraction progressed. 2. In the first phase, surface EMG activities decreased and discharge frequency of motor units also decreased. 3. In the second phase, surface EMG activities increased and discharge frequency of motor units also increased. 4. In the first phase, it was suggested that the bite force was maintained by an increase in the twitch tension produced by a motor unit and that there were no recruitment of additional motor units. 5. In the second phase, it was indicated that the bite force was maintained by the recruitment of new motor units and an increase in the discharge frequency of motor units to compensate a loss of force resulted from the contractile element fatigue.

The effect of dantrolene sodium on the discharge of alpha and gamma motoneurones to the soleus muscle in the decerebrate rat.

1. The effects of intravenous infusion of the direct acting muscle relaxant, dantrolene sodium (5 mg kg-1), on tension, integrated EMG, soleus muscle motor unit discharge frequency and gamma nerve fibre discharge were measured in the decerebrate rat. 2. Dantrolene sodium did not have any detectable direct effect upon the discharge of the gamma nerve fibres. 3. The soleus muscle of the decerebrate preparations exhibited spontaneous tension and reflex responses. 4. With the muscle held at constant length, dantrolene sodium caused an increase in the integrated EMG in 15 out of 18 experiments and a decrease in muscle tension in 15 out of 17. The results from these experiments showed great variability. 5. Dantrolene sodium caused a slight reduction in the tension response to tonic stretch; this was accompanied by an increase in the integrated EMG. 6. Dantrolene sodium also caused a shift in the relationship between tension and integrated EMG during the phasic component of the stretch reflex, with a greater integrated EMG being associated with a reduced tension. 7. Motor unit discharge frequencies were found to increase but not sufficiently to overcome the action of dantrolene sodium. It is concluded that motor unit recruitment must play an important role in the compensation for the muscle weakening action of dantrolene sodium.

Behavior of single motor units of human tibialis anterior muscle during voluntary shortening contraction under constant load torque

The recruitment and discharge frequency of motor units during voluntary shortening contraction with constant load torque were studied in tibialis anterior muscle of 10 human subjects trained to dorsiflex the tibiopedal joint as linearly as possible from 30° plantar flexion from the neutral position (tibiopedal joint angle, 90°) by 20° in 1, 2, 3, 5, 8, and 10 s. A constant load torque of 0, 5, 10, 20, and 25% of maximum voluntary torque of the muscle was applied. When speed of shortening contraction was constant there was considerable degree of constancy in the muscle length at which individual motor units were recruited. However, the recruitment order was not rigidly fixed among the units whose length range of recruitment overlapped. When the contraction was performed faster the units were recruited earlier. The discharge frequencies of the motor units changed little with a given ramp shortening of the muscle. There were also few changes in frequency among different contraction speeds. With increasing load torque to the muscle, the discharge frequency increased slightly at a rate of 1 to 2 Hz by increase of the load. The results suggest that the speed of shortening was coded by recruitment of motor units and that discharge frequency did not play any major role when a muscle was shortened under constant load torque.

Electromechanical changes during electrically induced and maximal voluntary contractions: Surface and intramuscular EMG responses during sustained maximal voluntary contraction

Changes in the electrical activity of the human gastrocnemius and soleus muscles during fatiguing maximal plantar flexions were studied with computer-aided EMG frequency power spectral analysis and intramuscular spike amplitude-frequency histogram analysis. In some experiments, brief supramaximal nerve stimulations of 80 Hz were given at 15-s intervals during sustained maximal voluntary contractions (MVCs). Multiple muscle biopsy samples were also obtained from the gastrocnemius muscle for fiber type determination. The surface EMG frequency spectral analysis showed a highly significant reduction in mean power frequency and root mean square EMG amplitude during sustained MVCs. The intramuscular spike amplitude-frequency histograms showed that the gastrocnemius muscle had a progressive reduction in the motor unit discharge frequency, particularly those with a relatively high amplitude, whereas the soleus muscle hardly showed a reduction in motor unit activity. Reduction in motor unit activity was also found to be more pronounced in gastrocnemius muscles with higher proportions of type II fibers. Brief maximal tetanic stimulations initially matching the MVC failed to increase the contraction force. Similarly, the evoked compound mass action potentials showed little change in the amplitude in subjects with different muscle fiber compositions. Results of this study suggest that (i) during sustained MVCs, force fatigue could not be attributed to a failure of muscle membrane electrical propagation; (ii) a progressive reduction in motor unit activation does not result in a functional disadvantage, but may optimize excitation-contraction coupling by avoiding a muscle electrical conduction failure; and (iii) the extent of the reduction in motor unit activation seems to be muscle-fiber-type-dependent which may account for the reduction in amplitude and frequency of the surface EMG.

Steps in production of motoneuron spikes during rhythmic firing.

Steps in production of motoneuron spikes during rhythmic firing.