Influence of motor units recruitment and firing rate on the soundmyogram and EMG characteristics in cat gastrocnemius

The separate contributions of the recruitment level and of the firing rate of the motor units on the soundmyogram and electromyogram time domain parameters were investigated during stimulation of the motor nerve of the cat gastrocnemius muscle. Upon orderly increase in the number of active motor units at a fixed firing rate, both the peak to peak amplitude (P-P_max) and the root mean square (RMS) of the sound myogram increased. At full recruitment the increase in firing rate from 2.5 to 50 Hz induced an exponential decline in the P-P_max. The RMS, however, followed this trend only from 15 to 50 Hz while showing an increase from 2.5 to 10 Hz. During simultaneous changes of recruitment and firing rate, the effect of increasing the number of motor units on the P-P_max and RMS is dampened by the increasing firing rate. The peak to peak amplitude of the EMG compound action potential increased with the number of active motor units. Moreover, its amplitude was not influenced by the firing rate. The EMG RMS, however, increases as a function of the firing rate. The results indicate that both the number and the firing rate of the active motor units contribute to the determination of the soundmyogram characteristics. Moreover, the peculiar changes of the soundmyogram time domain properties, compared to the ones of the EMG, allow one to differentiate the influence of the motor units number and firing rate on the electrical and mechanical performance of the muscle when stimulated.     

Ultrastructural diversity in motor units of crustacean stomach muscles

The physiological and ultrastructural properties of muscle fiber.s comprising three motor units in the gastric mill of blue crabs are described. In their contractile properties muscle fibers in all motor units are similar and resemble the slow type fibers in crustacean limb muscles. The majority of fibers generate large excitatory post-synaptic potentials which do not facilitate strongly. Structurally two types of fibers are found. The one type has long sarcomeres (greater than 6 mum), thin to thick myofilament ratios of 5-6:1 and diads located near the ends of the A-band. The other type has shorter sarcomeres (less than 6 mum), thin to thick myofilament ratios of 3:1 and diads located at mid sarcomere level. Both types of fibers occur within a single motor unit and this differs from the vertebrate situation. Furthermore, the finding of fibers with a low thin to thick myofilament ratio of 3:1 demonstrates that they are not exclusive to fast type crustacean muscle but also occur in slow stomach muscles.     

A simulation method for the firing sequences of motor units.

The firing sequences of motoneurons contain important information with regard to the underlying neural processes. Several methods have been proposed in the literature to simulate these sequences, however, one of the limitations is that they are not capable of simulating the complex neural dynamics of motor neurons, especially those of concurrently active ones, such as motor unit synchrony and motor unit common drive. In this paper, a novel model based on the Hodgkin-Huxley (HH) system is proposed, which has the ability to simulate the complex neurodynamics of the firing sequences of motor neurons. The model is presented at the cellular level and network level, and some simulation results from a simple 3-neuron network are presented to demonstrate its applications.     

Comparison of methods for estimating motor unit firing rate time series from firing times

The central nervous system regulates recruitment and firing of motor units to modulate muscle tension. Estimation of the firing rate time series is typically performed by decomposing the electromyogram (EMG) into its constituent firing times, then lowpass filtering a constituent train of impulses. Little research has examined the performance of different estimation methods, particularly in the inevitable presence of decomposition errors. The study of electrocardiogram (ECG) and electroneurogram (ENG) firing rate time series presents a similar problem, and has applied novel simulation models and firing rate estimators. Herein, we adapted an ENG/ECG simulation model to generate realistic EMG firing times derived from known rates, and assessed various firing rate time series estimation methods. ENG/ECG-inspired rate estimation worked exceptionally well when EMG decomposition errors were absent, but degraded unacceptably with decomposition error rates of ⩾1%. Typical EMG decomposition error rates—even after expert manual review—are 3–5%. At realistic decomposition error rates, more traditional EMG smoothing approaches performed best, when optimal smoothing window durations were selected. This optimal window was often longer than the 400 ms duration that is commonly used in the literature. The optimal duration decreased as the modulation frequency of firing rate increased, average firing rate increased and decomposition errors decreased. Examples of these rate estimation methods on physiologic data are also provided, demonstrating their influence on measures computed from the firing rate estimate.     

Study of firing pattern in human soleus motor units in tonic vibration reflex

Tonic vibration reflex was produced in the human soleus muscle by vibrating the tendon at the rate of 30–180 Hz and motor unit potentials were recorded. A correlation was found between the points at which these potentials occurred and vibratory stimuli over lower ranges of vibration rates (of up to 70–80 Hz) in all motor units, indicative of discrete bursts in the synaptic inflow to the motoneuron matching the vibratory stimuli. The correlation disappeared with an increase in vibration rate and manifested at high as well as low vibration rates in voluntarily contracted muscle. Since vibration is known to (presynaptically) depress monosynaptic reflexes induced by activating primary spindle endings, it is suggested and maintained that the correlation found at low vibration rate ranges could result from activating vibrational stimuli of secondary spindle endings which act on motoneurons via short pathways, thus evoking discrete motoneuronal EPSP.Information Transmission Research Institute, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 21, No. 6, pp. 765–772, November–December, 1989.     

Digit flexor muscles in the cat: their action and motor units

The relation between muscle action and the mechanical properties of motor units has been explored in the main digit flexors of the cat hind limb: plantaris (PL); flexor digitorum brevis (FDB); flexor hallucis longus (FHL); and, flexor digitorum longus (FDL). General observations on muscle action revealed that PL is an ankle extensor as well as a digit flexor. PL and FHL were shown to be the major force contributors to digit flexion with FDL playing a lesser but still significant role. The mechanical properties of PL, FHL and FDB motor units were studied by noting twitch and tetanic tensions produced by electrical stimulation of single alpha axons, functionally isolated from the ventral root filaments. These data were compared to similar data reported by Olson and Swett (1966) for flexor digitorum longus (FDL). Our sample (114 PL, 60 FDB and 124 FHL units) disclosed that PL, FDB and FHL have units of uniformly fast contraction times (means 22, 27 and 27 msec respectively). PL units developed the most tetanic tension (3 to 160, mean 62 gm-wt) followed by FHL (2 to 87, mean 31 gm-wt) with FDB units producing very little tension (1 to 20, mean 6 gm-wt). Swett and Olson's FDL sample (108 units) showed tensions ranging from 0.3 to 100 gm-wt (mean 10 gm-wt). A division of labor among the four muscles is proposed. The large PL units are advantageous for forceful phasic inputs to the digits during the locomotion and in keeping with PL's additional role as an ankle exstensor. The low output forces of FDB units are optimal for discrete input to the digits during subtle adjustments of posture. We propose that the larger fast contracting units of FHL are used primarily for forceful digit flexions required in locomotion and for phasic protrusion of the claws while the predominately small and slow contracting units of FDL are used for sustained claw protrusion.     

Innervation and motor patterns of the abdominal superficial flexor muscles in larval lobsters

The pattern of innervation and motor program of the abdominal superficial flexor muscle was investigated electrophysiologically in larval lobsters (Homarus americanus). The muscle receives both excitatory and inhibitory innervation in the larval as well as in the embryonic stages. Individual muscle fibers receive a single inhibitory neuron (f5) and a maximum of three excitors. Based on spike heights these axons belong to either the small (f1 or f2) or large (f3, f4) motoneurons. While the small axons preferentially innervate the medial muscle fibers the large axons innervate medial as well as lateral fibers. This larval pattern of innervation resembles the pattern in the adult lobster. The resemblance extends to the firing patterns as well with both large and small excitors firing spontaneously. Furthermore, evoked activity in the larvae produces reciprocal (and occasionally cyclical) bursts of excitor and inhibitor neurons denoting abdominal extension and flexion and resembling the firing patterns in adults. Consequently motor programs employed in steering the pelagic larvae are reminiscent of the programs for maintaining posture in the benthic adult lobsters.     

Aminergic control of neuronal firing rate in thalamic motor nuclei of the rat

The effects induced on neuronal firing by microiontophoretic application of the biological amines noradrenaline (NA) and 5-hydroxytryptamine (5-HT) were studied "in vivo" in ventral-anterior (VA) and ventrolateral (VL) thalamic motor nuclei of anaesthetized rats. In both nuclei the amines had a mostly depressive action on neuronal firing rate, the percentage of units responsive to NA application (88%) being higher than to 5-HT (72%). Short-lasting (less than 2 min) and long lasting (up to 20 min) inhibitory responses were recorded, the former mostly evoked by NA and the latter by 5-HT ejection. In some cases 5-HT application had no effect on the firing rate but modified the firing pattern. NA-evoked responses were significantly more intense in VL than in VA neurons. Short-lasting inhibitory responses similar to NA-induced effects were evoked by the alpha2 adrenergic receptor agonist clonidine and to a lesser extent by the beta adrenergic receptor agonist isoproterenol. Inhibitory responses to 5-HT were partially mimicked by application of the 5-HT(1A) receptor agonist 8-hydroxy-2(di-n-propylamino)tetralin (8-OH-DPAT) and of the 5-HT2 receptor agonist alpha-methyl-5-hydroxytryptamine (ALPHA-MET-5-HT). The latter evoked excitatory responses in some cases. Both 5-HT agonists were more effective on VA than on VL neurons. The effects evoked by agonists were at least partially blocked by respective antagonists. These results suggest that although both 5-HT and NA depress neuronal firing rate, their effects differ in time course and in the amount of inhibition; besides aminergic modulation is differently exerted on VA and VL.     

Short term bed-rest reduces conduction velocity of individual motor units in leg muscles

Space permanence simulations such as prolonged bed-rest can mimic some of the physiological modifications in the human body and provide study conditions that are more accessible than during space flight. A short term bed-rest experiment was organized to simulate the effects of weightlessness for studying the adaptation to this condition. Eight healthy young volunteers were studied before and immediately after the 14 day periods of strict bed-rest.Surface EMG signals were detected with linear electrode arrays from vastus medialis, vastus lateralis and tibialis anterior muscle during isometric voluntary contractions at 20% MVC. Motor unit action potentials (MUAPs) of individual motor units were extracted from the interference EMG signals with a partial decomposition algorithm and averaged.MUAP templates generated by the same motor unit could be retrieved before and after bed-rest period. Muscle fiber conduction velocity (CV) was estimated from each averaged MUAP template and from the global EMG signal. Both global and single MU conduction velocity was observed to decrease by about 10% after the bed-rest period (p < 0.05). Amplitude and power spectral parameters did not significantly change after the bed-rest period.It is concluded that a short term bed-rest reduces the CV of individual motor units without a significant effect on muscle force or on other electrophysiological parameters.     

Activity of single motor units in attention-demanding tasks: firing pattern in the human trapezius muscle

Activity of single motor units in relation to surface electromyography (EMG) was studied in 11 subjects in attention-demanding work tasks with minimal requirement of movement. In 53 verified firing periods, single motor units fired continuously from 30 s to 10 min (duration of the experiment work task) with a stable median firing rate in the range of 8–13 Hz. When the integrated surface EMG were stable, the motor units identified as a rule were continuously active with only small modulations of firing rate corresponding to low-amplitude fluctuations in surface EMG. Marked changes in the surface EMG, either sudden or gradual, were caused by recruitment or derecruitment of motor units, and not by modulations of the motor unit firing rate. Motor unit firing periods (duration 10 s-35 s) in low-level voluntary contractions (approximately 1%–5% EMG_max) performed by the same subjects showed median firing rates (7–12 Hz) similar to the observations in attention-related activation.     

Discharge frequencies of single motor units in human diaphragm and parasternal muscles in lying and standing

Single motor unitdischarge was measured directly in diaphragm and parasternalintercostal muscles to determine whether neural drive to humaninspiratory muscles changes between lying and standing. The finaldischarge frequency of diaphragmatic motor units increased slightly, by1 Hz (12%; P < 0.01), when subjects were standing [182 units, median 9.1 Hz (interquartile range 7.6-11.3 Hz)]compared with lying supine [159 units, 8.1 Hz (6.6-10.3Hz)]. However, this increase with standing occurred in onlytwo of six subjects, in one of whom tidal volume increasedsignificantly during standing. Parasternal intercostal motor unitfinal discharge frequencies did not differ between standing [116units, 8.0 Hz (6.6-9.6 Hz)] and lying [124 units, 8.4 Hz(7.0-10.3 Hz)]. The discharge frequencies at the onset ofinspiration did not differ between lying and standing for eithermuscle. A larger proportion of motor units in both inspiratory muscleshad postinspiratory or tonic expiratory activity for lying comparedwith standing (15 vs. 4%; P < 0.05). We conclude thatthere is no major difference in the phasic inspiratory drive to thediaphragm with the change in posture.

     

Discharge properties of motor units during steady isometric contractions performed with the dorsiflexor muscles

The purpose of this study was to record the discharge characteristics of tibialis anterior motor units over a range of target forces and to import these data, along with previously reported observations, into a computational model to compare experimental and simulated measures of torque variability during isometric contractions with the dorsiflexor muscles. The discharge characteristics of 44 motor units were quantified during brief isometric contractions at torques that ranged from recruitment threshold to an average of 22 ± 14.4% maximal voluntary contraction (MVC) torque above recruitment threshold. The minimal [range: 5.8-19.8 pulses per second (pps)] and peak (range: 8.6-37.5 pps) discharge rates of motor units were positively related to the recruitment threshold torque (R(2) ≥ 0.266; P < 0.001). The coefficient of variation for interspike interval at recruitment was positively associated with recruitment threshold torque (R(2) = 0.443; P < 0.001) and either decreased exponentially or remained constant as target torque increased above recruitment threshold torque. The variability in the simulated torque did not differ from the experimental values once the recruitment range was set to ～85% MVC torque, and the association between motor twitch contraction times and peak twitch torque was defined as a weak linear association (R(2) = 0.096; P < 0.001). These results indicate that the steadiness of isometric contractions performed with the dorsiflexor muscle depended more on the distributions of mechanical properties than discharge properties across the population of motor units in the tibialis anterior.     

Decreases in motor unit firing rate during sustained maximal-effort contractions in young and older adults

Previous studies have suggested that older adults may be more resistant to muscular fatigue than young adults. We sought to determine whether motor unit firing rate might be a factor that determines the response to fatiguing exercise in young and older subjects. Motor unit recordings and muscular forces were obtained from the tibialis anterior (TA) muscle of 11 young and 8 older individuals. Maximal voluntary force was first measured during maximal-effort dorsiflexion contractions. Each subject then performed a series of 15 maximal isometric contractions, with each contraction lasting 30 s. A 10-s rest period separated the fatiguing contractions. As a result of the fatiguing exercise, both subject groups demonstrated a significant loss in maximal force. The force decline was less in the older adults (20.4%) than in the young adults (33.8%). As expected, prior to muscle fatigue, maximal firing rates in the TA muscle were greater in the young (28.1 ± 5.8 imp/s) than in the older adults (22.3 ± 4.8 imp/s). The decrease in motor unit firing rate with fatigue was also greater in the young adults (34.9%), than in the older adults (22.0%). These results suggest that the greater fatigue-resistance exhibited by older individuals might be explained by the fact that the decline in motor unit firing rate during fatigue is greater in young persons than it is in older adults.     

Properties of twitch motor units of the ankle extensor muscles in the bullfrog Rana catesbeiana

A study of the mechanical properties of the twitch motor units in the ankle extensor muscles of bullfrogs was undertaken to expand our view of the diversity of motor unit properties among vertebrates. Two muscles were chosen that represent a wide range of extensor function: the plantaris longus (PL) is a large muscle providing most of the force for ankle extension in hopping and swimming, and the tibialis posticus (TP) is relatively small and may act as an ankle stabilizer or be primarily postural in function. Both muscles have highly fatigable motor units, but also some (especially in TP) low or non-fatigable ones. Mean tetanic tensions of motor units in both muscles are relatively large as compared with those of mammals but are especially large in PL, No clear correlations were found between contraction times and either motor unit tetanic tensions or fatigability, nor did the motor units fall into clearly defined types based on any functional parameters. Overall contraction and relaxation times are slow compared with those of mammals and are somewhat slow compared to those of other frogs; unlike results from earlier studies, the large units of PL are slower than the small units of TP. This results in PL units reaching fused tetani at lower stimulus frequencies. The twitch/tetanus and force/frequency ratios in PL motor units are much larger than those of TP, giving PL units greater relative forces at lower stimulus rates. These results are discussed in the context of motor unit function. © 1994 Wiley-Liss, Inc.