Investigation of motor unit recruitment during stimulated contractions of tibialis anterior muscle

This work investigated motor unit (MU) recruitment during transcutaneous electrical stimulation (TES) of the tibialis anterior (TA) muscle, using experimental and simulated data. Surface electromyogram (EMG) and torque were measured during electrically-elicited contractions at different current intensities, on eight healthy subjects.EMG detected during stimulation (M-wave) was simulated selecting the elicited MUs on the basis of: (a) the simulated current density distribution in the territory of each MU and (b) the excitation threshold characteristic of the MU. Exerted force was simulated by adding the contribution of each of the elicited MUs. The effects of different fat layer thickness (between 2 and 8 mm), different distributions of excitation thresholds (random excitation threshold, higher threshold for larger MUs or smaller MUs), and different MU distributions within the muscle (random distribution, larger MU deeper in the muscle, smaller MU deeper) on EMG variables and torque were tested.Increase of the current intensity led to a first rapid increase of experimental M-wave amplitude, followed by a plateau. Further increases of the stimulation current determined an increase of the exerted force, without relevant changes of the M-wave. Similar results were obtained in simulations.Rate of change of conduction velocity (CV) and leading coefficient of the second order polynomial interpolating the force vs. stimulation level curve were estimated as a function of increasing current amplitudes. Experimental data showed an increase of estimated CV with increasing levels of the stimulation current (for all subjects) and a positive leading coefficient of force vs. stimulation current curve (for five of eight subjects). Simulations matched the experimental results only when larger MUs were preferably located deeper in the TA muscle (in line with a histochemical study). Marginal effect of MU excitation thresholds was observed, suggesting that MUs closer to the stimulation electrode are recruited first during TES regardless of their excitability.     

Ultrasonographic quantification of architectural response in tibialis anterior to neuromuscular electrical stimulation

While muscle contraction in voluntary efforts has been widely investigated, little is known about contraction during neuromuscular electrical stimulation (NMES). The aim of this study was to quantify in vivo muscle architecture of agonist and antagonist muscles at the ankle joint during NMES. Muscle fascicle lengths and pennation angles of the tibialis anterior (TA) and lateral gastrocnemius muscles were assessed via ultrasonography in 8 healthy young males. Measures were obtained during maximal NMES and torque-matched voluntary dorsiflexion contractions. In the TA, NMES induced a shorter fascicle length (67.2 ± 8.1 mm vs 74.6 ± 11.4 mm; p = 0.04) and a greater pennation angle (11.0 ± 2.4° vs 9.3 ± 2.5°; p = 0.03) compared with voluntary torque-matched dorsiflexion contractions. Architectural responses in the antagonist lateral gastrocnemius muscle did not significantly differ from rest or between voluntary and electrically induced contractions (p > 0.05). Contraction of the antagonist muscle was not a contributing factor to a greater fascicle shortening and increased pennation angle in the TA during NMES. TA architectural response during NMES likely arose from the contribution of muscle synergists during voluntary contractions coupled with a potentially localized contractile activity under the stimulation electrodes during NMES induced contractions.     

M-wave properties during progressive motor unit activation by transcutaneous stimulation.

The aim of this study was to interpret changes in experimentally recorded M waves with progressive motor unit (MU) activation based on simulation of the surface electromyogram. Activation order during transcutaneous electrical stimulation was analyzed by investigating M-wave average rectified value, spectral properties, and conduction velocity (CV) during electrically elicited contractions. M-waves were detected from the biceps brachii muscle of 10 healthy male subjects by a linear adhesive array of eight electrodes. Electrical stimulation was delivered to the motor point at either constant current intensity (40, 60, 80, and 100% of the supramaximal stimulation current) or with linearly increasing current. A model of surface electromyogram generation that varied activation order based on MU size and location was used to interpret the experimental results. From the experimental and model analysis, it was found that 1) MUs tended to be activated from low to high CV and from the superficial to the deep muscle layers with increasing transcutaneous electrical stimulation of the biceps brachii muscle, and 2) characteristic spectral frequencies of the M-wave were affected by many factors other than average CV (such as the activation order by MU location or the spread of the MU innervation zones and CVs), thus decreasing with a concomitant increase in CV during progressive MU activation.     

Variability of successive contractions subtracted from unfused tetanus of fast and slow motor units

Stimulation of motor units (MUs) with repeated pulses evokes tetanic contractions, which consist of overlapping mechanical responses. The summation of these responses into tetanus is a nonlinear process due to the dynamic changes in the amplitudes and time parameters of the successive components. In order to study these changes, two MUs (one fast and one slow) of rat medial gastrocnemius muscle were stimulated with a progressively increasing number of pulses, from one (i = 1) to sixteen (i = 16) at a frequency of 15 Hz for the slow MU and 60 Hz for the fast MU. The individual responses were calculated by subtracting the (i)th from the (i + 1)th tetanus recording. The contractions obtained following subtraction were modeled using a novel 6-parameter analytical function. The main conclusions of this study are (1) the newly presented analytical function is able to precisely describe the variable shape of all subtracted experimental contractions; (2) the shapes of successive contractions are variable and the subtracted contractions differ from the individual twitches; (3) as the pulse number increases, the parameters of the subtracted contractions change in a different manner for the slow and fast MUs: for the slow MU, the maximal forces and the time parameters increase considerably up to the 4th response, after which they remain nearly constant or show only a slight increase; for the fast MU, the maximal forces and durations also increase, whereas the remaining time parameters initially increase and then maintain a constant level or decrease, which explains the sag phenomenon visible in the unfused tetanus of fast MUs.     

Phantom reflexes: muscle contractions at a frequency not physically present in the input stimuli

In the motor system, the periodic stimulation of one Ia-afferent input produces reflex muscle contractions at the input frequency. However, we observed that when two Ia monosynaptic reflex-afferent inputs are involved the periodic muscle contractions may occur at a frequency physically not present in the afferent inputs even when these inputs are sub-threshold. How can the muscles respond with such phantom reflex contractions at a frequency physically absent in the sub-threshold Ia-afferent input stimuli? Here we provide an explanation for this phenomenon in the cat spinal cord, that we termed “ghost motor response”. We recorded monosynaptic reflexes in the L7 ventral root, intracellular potentials in the motoneurons, and the associated muscular contractions elicited by stimulation of the lateral and medial gastrocnemius nerves. By stimulating with periodic pulses of sub-threshold intensities and distinct frequencies of 2 and 3 Hz the lateral and medial gastrocnemius nerves, respectively, we observed monosynaptic responses and phantom reflex muscle contractions occurring at the fundamental frequency (1 Hz), which was absent in the input stimuli. Thus we observed a reflex ghost motor response at a frequency not physically present in the inputs. We additionally studied the inharmonic case for sub-threshold stimuli and observed muscular contractions occurring at much lower frequencies, which were also conspicuously absent in the inputs. This is the first experimental evidence of a phantom reflex response in the nervous system. The observed behavior was modeled by numerical simulations of a pool of neurons subjected to two different input pulses.     

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.     

A myocontrolled neuroprosthesis integrated with a passive exoskeleton to support upper limb activities

This work aimed at designing a myocontrolled arm neuroprosthesis for both assistive and rehabilitative purposes.The performance of an adaptive linear prediction filter and a high-pass filter to estimate the volitional EMG was evaluated on healthy subjects (N = 10) and neurological patients (N = 8) during dynamic hybrid biceps contractions. A significant effect of filter (p = 0.017 for healthy; p < 0.001 for patients) was obtained. The post hoc analysis revealed that for both groups only the adaptive filter was able to reliably detect the presence of a small volitional contribution.An on/off non-linear controller integrated with an exoskeleton for weight support was developed. The controller allowed the patient to activate/deactivate the stimulation intensity based on the residual EMG estimated by the adaptive filter. Two healthy subjects and 3 people with Spinal Cord Injury were asked to flex the elbow while tracking a trapezoidal target with and without myocontrolled-NMES support. Both healthy subjects and patients easily understood how to use the controller in a single session. Two patients reduced their tracking error by more than 60% with NMES support, while the last patient obtained a tracking error always comparable to the healthy subjects performance (<4°).This study proposes a reliable and feasible solution to combine NMES with voluntary effort.     

Electromyographic assessment of isometric and dynamic activation characteristics of the latissimus dorsi muscle

The aim of the current study was to analyze the activation characteristics and potential compartmentalization of the latissimus dorsi (LD) muscle during common maximal voluntary isometric contractions (MVICs) and functional dynamic tasks. Surface electromyography (sEMG) was used to measure activation magnitudes from four electrode sites (referenced to the T10, T12, L1 & L4 LD vertebral origins) across the fanning muscle belly of the LD. In addition, EMG waveforms were cross-correlated to study temporal activation timing between electrode sites (T10-T12, T12-L1, L1-L4 & T10-L4). The MVICs that were tested included a humeral adduction, humeral adduction with internal rotation, a chest-supported row and a humeral extension. Dynamic movements included sagittal lift/lowers from the floor to knee, knee to hip and hip to shoulder. No magnitude-based (p = 0.6116) or temporal-based differences were observed between electrode sites during the MVIC trials. During dynamic movements no temporal-based, but some magnitude-based differences between electrode sites were observed to be present; these differences were small in magnitude and were observed for both the maximum (p = 0.0002) and mean (p = 0.0002) EMG magnitudes. No clear pattern of compartmentalization was uncovered in the contractions studied here. In addition to these findings, it was determined that the most effective MVIC technique for LD EMG normalization purposes was a chest-supported row MVIC, paired with a T12 electrode site.     

Firing rate and conduction velocity of single motor units in the trapezius muscle in fibromyalgia patients and healthy controls

Fibromyalgia is a common chronic pain condition in the population (2–4%), which often is associated with prominent negative consequences with respect to participation in daily activities. There are several reports in the literature concerning the effects of acute experimental pain on motor control. However, a more heterogeneous picture exists in the literature with respect to whether chronic pain conditions affect motor control.This study compares firing rate and conduction velocity (CV) of single motor units (MUs) in the trapezius muscle of fibromyalgia patients (FM) and healthy controls (CON). Multi-channel surface electromyography was used to estimate both MU firing rate and CV because this technique allows simultaneous estimation of both these variables and the measurements are easy and non-invasive. In this study, 29 FM and 30 CON subjects participated and performed isometric shoulder elevations using weights up to 4 kg. No significant differences in the firing rate of MUs in the trapezius muscle were found between the FM and CON groups (95% confidence interval was −1.9 and 1.3 pulses per second). There were no significant differences in CV between the groups at 1 and 2 kg load. However, the FM group had significantly higher CV in contractions without external load (p = 0.004).We were unable to confirm the pain-adaptation model since no differences in firing rate between the two groups were found. CV was significantly higher in FM than in healthy controls; this might be due to alterations in histopathology and microcirculation.     

Effect of pain location on spatial reorganisation of muscle activity

IntroductionWe aimed to determine whether the changes in muscle activity (in terms of both gross electromyography (EMG) and motor unit (MU) discharge characteristics) observed during pain are spatially organized with respect to pain location within a muscle which is the main contributor of the task.MethodsSurface and fine-wire EMG was recorded during matched low-force isometric plantarflexion from soleus (from four quadrants with fine-wire EMG and from the medial/lateral sides with surface EMG), both gastrocnemii heads, peroneus longus, and tibialis anterior. Four conditions were tested: two control conditions that each preceded contractions with pain induced in either the lateral (Pain_L) or medial (Pain_M) side of soleus.ResultsNeither the presence (p = 0.28) nor location (p = 0.19) of pain significantly altered gross muscle activity of any location (lateral/medial side of soleus, gastrocnemii, peroneus longus and tibialis anterior). Group data from 196 MUs show redistribution of MU activity throughout the four quadrants of soleus, irrespective of pain location. The significant decrease of MU discharge rate during pain (p < 0.0001; Pain_L: 7.3 ± 0.9–6.9 ± 1.1 Hz, Pain_M: 7.0 ± 1.1 to 6.6 ± 1.1 Hz) was similar for all quadrants of the soleus (p = 0.43), regardless of the pain location (p = 0.98). There was large inter-participant variation in respect to the characteristics of the altered MU discharge with pain.ConclusionResults from both surface and fine-wire EMG recordings do not support the hypothesis that muscle activity is reorganized in a simple systematic manner with respect to pain location.     

Motor unit recruitment when neuromuscular electrical stimulation is applied over a nerve trunk compared with a muscle belly: triceps surae

Neuromuscular electrical stimulation (NMES) can be delivered over a nerve trunk or muscle belly and can generate contractions by activating motor (peripheral pathway) and sensory (central pathway) axons. In the present experiments, we compared the peripheral and central contributions to plantar flexion contractions evoked by stimulation over the tibial nerve vs. the triceps surae muscles. Generating contractions through central pathways follows Henneman's size principle, whereby low-threshold motor units are activated first, and this may have advantages for rehabilitation. Statistical analyses were performed on data from trials in which NMES was delivered to evoke 10-30% maximum voluntary torque 2-3 s into the stimulation (Time(1)). Two patterns of stimulation were delivered: 1) 20 Hz for 8 s; and 2) 20-100-20 Hz for 3-2-3 s. Torque and soleus electromyography were quantified at the beginning (Time(1)) and end (Time(2); 6-7 s into the stimulation) of each stimulation train. H reflexes (central pathway) and M waves (peripheral pathway) were quantified. Motor unit activity that was not time-locked to each stimulation pulse as an M wave or H reflex ("asynchronous" activity) was also quantified as a second measure of central recruitment. Torque was not different for stimulation over the nerve or the muscle. In contrast, M waves were approximately five to six times smaller, and H reflexes were approximately two to three times larger during NMES over the nerve vs. the muscle. Asynchronous activity increased by 50% over time, regardless of the stimulation location or pattern, and was largest during NMES over the muscle belly. Compared with NMES over the triceps surae muscles, NMES over the tibial nerve produced contractions with a relatively greater central contribution, and this may help reduce muscle atrophy and fatigue when NMES is used for rehabilitation.     

The putative somatostatin antagonist cyclo-somatostatin has opioid agonist effects in gastrointestinal preparations

AimsSpecificity of receptor antagonists used is crucial for clarifying physiological/pathophysiological roles of the respective endogenous agonist. We studied the effects (somatostatin antagonist and possibly other actions) of cyclo-somatostatin (CSST), a putative somatostatin receptor antagonist on the guinea-pig small intestine, a preparation where somatostatin causes inhibition of nerve-mediated contractions.Main methodsIn isolated organ experiments, half-maximal cholinergic “twitch” contractions of the guinea-pig small intestine were evoked or tonic contractions of the rat stomach fundus strip (in the presence of physostigmine) were elicited by electrical field stimulation. The effects of somatostatin (somatostatin-14), CSST, naloxone, as well as of direct smooth muscle stimulants were examined.Key findingsSomatostatin (10 nM–1 μM) caused transient inhibition of the twitch contraction, in a naloxone-insensitive manner. Surprisingly, CSST (0.3–1 μM) also inhibited twitch contractions (more than 50% reduction at 1 μM). This effect was prevented by the opioid receptor antagonist naloxone. Responses to acetylcholine or histamine were not or only minimally inhibited by CSST (up to 3 μM). CSST (0.3 μM in the absence or 1–10 μM in the presence of naloxone) failed to inhibit the effect of somatostatin. The SST₂ receptor antagonist CYN-154806 (3 μM) attenuated the effect of somatostatin and failed to evoke naloxone-sensitive inhibition of the twitch response. The naloxone-sensitive inhibitory effect of CSST on cholinergic contractions was also confirmed in the rat stomach fundus preparation.SignificanceCyclo-somatostatin exerts opioid agonist activity in the two preparations tested, while it does not behave as a somatostatin-receptor antagonist in the guinea-pig intestine.     

Spatial distribution of surface action potentials generated by individual motor units in the human biceps brachii muscle

This study analyses the spatial distribution of individual motor unit potentials (MUPs) over the skin surface and the influence of motor unit depth and recording configuration on this distribution. Multichannel surface (13 × 5 electrode grid) and intramuscular (wire electrodes inserted with needles of lengths 15 and 25 mm) electromyographic (EMG) signals were concurrently recorded with monopolar derivations from the biceps brachii muscle of 10 healthy subjects during 60-s isometric contractions at 20% of the maximum torque. Multichannel monopolar MUPs of the target motor unit were obtained by spike-triggered averaging of the surface EMG. Amplitude and frequency characteristics of monopolar and bipolar MUPs were calculated for locations along the fibers’ direction (longitudinal), and along the direction perpendicular (transverse) to the fibers. In the longitudinal direction, monopolar and bipolar MUPs exhibited marked amplitude changes that extended for 16–32 mm and 16–24 mm over the innervation and tendon zones, respectively. The variation of monopolar and bipolar MUP characteristics was not symmetrical about the innervation zone. Motor unit depth had a considerable influence on the relative longitudinal variation of amplitude for monopolar MUPs, but not for bipolar MUPs. The transverse extension of bipolar MUPs ranged between 24 and 32 mm, whereas that of monopolar MUPs ranged between 72 and 96 mm. The mean power spectral frequency of surface MUPs was highly dependent on the transverse electrode location but not on depth. This study provides a basis for the interpretation of the contribution of individual motor units to the interference surface EMG signal.     

Transcutaneous nerve stimulation. Frequency and waveform specificity in humans

The effects of transcutaneous electrical nerve stimulation (TNS) of a constant alternating current administered at various frequencies and waveforms to volunteer human subjects were investigated. The TNS was found to evoke noncutaneous subjective sensations in all the subjects. Only with a sinusoid waveform of TNS were distinct frequency ranges of the stimulation associated with specific noncutaneous subjective sensations. Our findings suggest that nervous tissue is capable of discriminating the waveform parameters of an electrical stimulus.     

Neuromuscular electrical stimulation attenuates thigh skeletal muscles atrophy but not trunk muscles after spinal cord injury

The current study examined the effects of 12 weeks of surface neuromuscular electrical stimulation (NMES) and ankle weights on the cross-sectional areas (CSAs) of three thigh [Gracilis (Gra), Sartorious (Sar) and Adductor (Add)] as well as two trunk [hip flexor (HF) and back extensor (BE)] muscle groups in men with spinal cord injury (SCI). Seven individuals with chronic motor complete SCI were randomly assigned into a resistance training + diet (RT + diet; n = 4) or diet control (n = 3) groups. The RT + diet group underwent twice weekly training with surface NMES and ankle weights for 12 weeks. Training composed of four sets of 10 repetitions of leg extension exercise while sitting in their wheelchairs. Both groups were asked to monitor their dietary intake. Magnetic resonance images were captured before and after 12 weeks of interventions. Gra muscle CSA showed no change before and after interventions. A significant interaction (P = 0.001) was noted between both groups as result of 9% increase and 10% decrease in the Gra muscle CSA of the RT + diet and diet groups, respectively. Sar muscle CSA increased [1.7 ± 0.4–2.5 ± 0.5 cm²; P = 0.029] in the RT + diet group with no change [2.9 ± 1.4–2.6 ± 1.3 cm²] in the diet group; with interaction noted between both groups (P = 0.002). Analysis of covariance indicated that Add muscle CSA was 38% greater in the RT + diet compared to the diet group (P = 0.025) after 12 weeks; a trend of interaction was also noted between both groups (P = 0.06). HF and BE muscle groups showed no apparent changes in CSA in both groups. The results suggested that surface NMES can delay the process of progressive skeletal muscle atrophy after chronic SCI. However, the effects are localized to the trained thigh muscles and do not extend to the proximal trunk muscles.     

Feasibility study of detecting surface electromyograms in severely obese patients

The aims of this study were to examine if surface EMG signals can be detected from the quadriceps femoris muscle of severely obese patients and to investigate if differences exist in quadriceps force and myoelectric manifestations of fatigue between obese patients and lean controls.Fourteen severely obese patients (body mass index, BMI, mean ± SD: 44.9 ± 6.3 kg/m²) and fourteen healthy controls (BMI: 23.7 ± 2.5 kg/m²) were studied. The vastus medialis and lateralis of the dominant thigh were concurrently investigated during voluntary isometric contractions (10-s long at submaximal and maximal intensities and intermittent submaximal contractions until exhaustion) and sustained (120-s long) electrically elicited contractions.We found that the detection of surface EMG signals from the quadriceps is feasible also in severely obese subjects presenting increased thickness of the subcutaneous fat tissue. In addition, we confirmed and extended previous findings showing that the volume conductor properties determine the amplitude and spectral features of the detected surface EMG signals: the lower the subcutaneous tissue thickness, the higher the amplitude and mean frequency estimates. Further, we found no differences in the mechanical and myoelectric manifestations of fatigue during intermittent voluntary and sustained electrically elicited contractions between obese patients and lean controls.     

The impact of altered task mechanics on timing and duration of eccentric bi-articular muscle contractions during cycling

In order to understand muscle adaptations to altered task mechanics during cycling, this study investigated the impact of altered seat height and cadence on timing and duration of gastrocnemius (GAST), biceps femoris (BF) and vastus lateralis (VL) eccentric contractions and muscle activation patterns, and cycling economy. Ten male cyclists completed 9 × 5 min of cycling at 3 seat heights and 3 cadences. Three-dimensional leg kinematics and muscle activation patterns were recorded to estimate timing of eccentric muscle contractions. Onset, offset and duration of eccentric contractions and, onset, offset and duration of muscle activation were calculated, along with cycling economy. Duration of GAST and VL eccentric contractions decreased with increasing seat height due to earlier offset of eccentric muscle contractions. Duration of BF eccentric contractions significantly increased with seat height due to a later eccentric contraction offset. Offset of GAST and BF muscle activation occurred earlier with increasing cadence. Cycling economy was significantly affected by cadence but not seat height. The results suggest that as a consequence of altered seat height, proprioceptive feedback is used to fine-tune the timing of bi-articular eccentric muscle contractions. These results may have implications for seat height self-selection.     

Adaptations of upper trapezius muscle activity during sustained contractions in women with fibromyalgia

The study compared the distribution of electromyographic (EMG) signal amplitude in the upper trapezius muscle in 10 women with fibromyalgia and in 10 healthy women before and after experimentally-induced muscle pain. Surface EMG signals were recorded over the right upper trapezius muscle with a 10 × 5 grid of electrodes during 90° shoulder abduction sustained for 60 s. The control subjects repeated the abduction task following injections of isotonic and hypertonic (painful) saline into the upper trapezius muscle. The EMG amplitude was computed for each electrode pair and provided a topographical map of the distribution of muscle activity. The pain level rated by the patients at the beginning of the sustained contraction was 5.9 ± 1.5. The peak pain intensity for the control group following the injection of hypertonic saline was 6.0 ± 1.6. During the sustained contractions, the EMG amplitude increased relatively more in the cranial than caudal region of the upper trapezius muscle for the control subjects (shift in the distribution of EMG amplitude: 2.3 ± 1.3 mm; P < 0.01). The patient group showed lower average EMG amplitude than the controls during the contraction (P < 0.05) and did not show different changes in EMG amplitude between different regions of the upper trapezius. A similar behavior was observed for the control group following injection of hypertonic saline. The results indicate that muscle pain prevents the adaptation of upper trapezius activity during sustained contractions as observed in non-painful conditions, which may induce overuse of similar muscle compartments with fatigue.     

Effect of aging on properties of motor unit action potentials in the rat medial gastrocnemius muscle

The purpose of this study was to investigate whether age-related changes in motor unit (MU) contractile properties are reflected in parameters of motor unit action potentials (MUAPs). MUs of the medial gastrocnemius muscle were functionally isolated in anaesthetized Wistar rats. A control group of young animals (5–10 mo) was compared to two groups of old rats (24–25 mo and 28–30 mo). The basic contractile properties of MUs as well as the amplitude, total duration, peak-to-peak time, and number of turns within MUAPs were measured. Effects of aging were mainly observed for fast fatigable MUs (a prolongation of MUAPs and increased number of turns). The MUAP amplitude did not change significantly with aging in either MU type, but it correlated to the twitch or tetanic forces, which tended to increase with age, especially for slow MUs. We concluded that the prolongation of MUAPs and the greater incidence of signal turns was probably a result of a decrease in muscle fiber conduction velocity and/or an increase in their dispersion, and enlargement of MU territories – presumably caused by axonal sprouting of surviving motoneurons. The latter might also be responsible for the observed age-related tendency for a increase in MUAP amplitudes in slow MUs.