Conduction velocity of low-threshold motor units during ischemic contractions performed with surface EMG feedback.

The aim of this study was to analyze the effect of ischemia on low-threshold motor unit conduction velocity. Nine subjects were trained to isolate the activity of a single motor unit (target motor unit) in the abductor pollicis brevis muscle with feedback on surface EMG signals recorded with a 16-electrode linear array. After training, the subjects activated the target motor unit at approximately 8 pulses per second (pps) for five 3-min-long contractions. During the third and fourth contractions, a cuff inflated at 180 mmHg around the forearm induced ischemia of the hand. The exerted force (mean +/- SE, 4.6 +/- 2.1% of the maximal voluntary contraction force), discharge rate (8.6 +/- 0.4 pps), interpulse interval variability (34.8 +/- 2.5%), and peak-to-peak amplitude of the target motor unit action potentials (176.6 +/- 18.2 microV) were not different among the five contractions. Conduction velocity, mean power spectral frequency, and action potential duration were the same in the beginning of the five contractions (2.8 +/- 0.2 m/s, 195.2 +/- 10.5 Hz, and 5.4 +/- 0.3 ms, respectively) and changed over the 3 min of sustained activation only during the fourth contraction. Conduction velocity and mean power spectral frequency decreased (10.05 +/- 1.8% and 8.50 +/- 2.18% during the 3 min, respectively) and action potential duration increased (8.2 +/- 4.6% in the 3 min) during the fourth contraction. In conclusion, 1) subjects were able to isolate the activity of a single motor unit with surface EMG visual feedback in ischemic conditions maintained for 16 min, and 2) the activation-induced decrease in single motor unit conduction velocity was significantly larger with ischemia than with normal circulation, probably due to the alteration of mechanisms of ion exchange across the fiber membrane.     

Spike-triggered average torque and muscle fiber conduction velocity of low-threshold motor units following submaximal endurance contractions.

The motor unit twitch torque is modified by sustained contraction, but the association to changes in muscle fiber electrophysiological properties is not fully known. Thus twitch torque, muscle fiber conduction velocity, and action potential properties of single motor units were assessed in 11 subjects following an isometric submaximal contraction of the tibialis anterior muscle until endurance. The volunteers activated a target motor unit at the minimum discharge rate in eight 3-min-long contractions, three before and five after an isometric contraction at 40% of the maximal torque, sustained until endurance. Multichannel surface electromyogram signals and joint torque were averaged with the target motor unit potential as trigger. Discharge rate (mean +/- SE, 6.6 +/- 0.2 pulses/s) and interpulse interval variability (33.3 +/- 7.0%) were not different in the eight contractions. Peak twitch torque and recruitment threshold increased significantly (93 +/- 29 and 12 +/- 5%, P <0.05) in the contraction immediately after the endurance task with respect to the preendurance values (0.94 +/- 0.26 mN.m and 3.7 +/- 0.5% of the maximal torque), whereas time to peak of the twitch torque did not change (74.4 +/- 10.1 ms). Muscle fiber conduction velocity decreased and action potential duration increased in the contraction after the endurance (6.3 +/- 1.8 and 9.8 +/- 1.8%, respectively, P <0.05; preendurance values, 3.9 +/- 0.2 m/s and 11.1 +/- 0.8 ms), whereas the surface potential peak-to-peak amplitude did not change (27.1 +/- 3.1 microV). There was no significant correlation between the relative changes in muscle fiber conduction velocity or surface potential duration and in peak twitch torque (R2= 0.04 and 0.10, respectively). In conclusion, modifications in peak twitch torque of low-threshold motor units with sustained contraction are mainly determined by mechanisms not related to changes in action potential shape and in its propagation velocity.     

Practice reduces motor unit discharge variability in a hand muscle and improves manual dexterity in old adults.

A steadiness-improving intervention was used to determine the contribution of variability in motor unit discharge rate to the fluctuations in index finger acceleration and manual dexterity in older adults. Ten healthy and sedentary old adults (age 72.9 +/- 5.8 yr; 5 men) participated in the study involving abduction of the left index finger. Single motor unit activity was recorded in the first dorsal interosseus muscle before, after 2 wk of light-load training (10% maximal load), and after 4 wk of heavy-load training (70% maximal load). As expected, the light-load training was effective in reducing the fluctuations in index finger acceleration during slow shortening (0.25 +/- 0.12 to 0.13 +/- 0.08 m/s(2)) and lengthening contractions (0.29 +/- 0.10 to 0.14 +/- 0.06 m/s(2)). Along with the decline in the magnitude of the fluctuations, there was a parallel decrease in the coefficient of variation for discharge rate during both contraction types (33.8 +/- 6.8 to 25.0 +/- 5.9%). The heavy-load training did not further improve either the fluctuations in acceleration or discharge rate variability. Furthermore, the manual dexterity of the left hand improved significantly with training (Purdue pegboard test: 11 +/- 3 to 14 +/- 1 pegs). Bivariate correlations indicated that the reduction in fluctuations in motor output during shortening (r(2) = 0.24) and lengthening (r(2) = 0.14) contractions and improvement in manual dexterity (r(2) = 0.26) was directly associated with a decline in motor unit discharge rate variability. There was a strong association between the fluctuations in motor output and manual dexterity (r(2) = 0.56). These results indicate that practice of a simple finger task was accompanied by a reduction in the discharge rate variability of motor units, a decrease in the fluctuations in motor output of a hand muscle, and an improvement in the manual dexterity of older adults.     

Effect of temperature on spike-triggered average torque and electrophysiological properties of low-threshold motor units.

The aim of the study was to jointly analyze temperature-induced changes in low-threshold single motor unit twitch torque and action potential properties. Joint torque, multichannel surface, and intramuscular electromyographic signals were recorded from the tibialis anterior muscle of 12 subjects who were instructed to identify the activity of a target motor unit using intramuscular electromyographic signals as feedback. The target motor unit was activated at the minimum stable discharge rate in seven 3-min-long contractions. The first three contractions (C1-C3) were performed at 33 degrees C skin temperature. After 5 min, the subject performed three contractions at 33 degrees C (T1), 39 degrees C (T2), and 45 degrees C (T3), followed by a contraction at 33 degrees C (C4) skin temperature. Twitch torque and multichannel surface action potential of the target motor unit were obtained by spike-triggered averaging. Discharge rate (mean +/- SE, 7.1 +/- 0.5 pulses/s), interpulse interval variability (35.8 +/- 9.2%), and recruitment threshold (4.5 +/- 0.4% of the maximal voluntary torque) were not different among the seven contractions. None of the investigated variables were different among C1-C3, T1, and C4. Conduction velocity and peak twitch torque increased with temperature (P < 0.05; T1: 3.53 +/- 0.21 m/s and 0.82 +/- 0.23 mN x m, T2: 3.93 +/- 0.24 m/s and 1.17 +/- 0.36 mN x m, T3: 4.35 +/- 0.25 m/s and 1.46 +/- 0.40 mN x m, respectively). Twitch time to peak and surface action potential peak-to-peak amplitude were smaller in T3 (61.8 +/- 2.0 ms and 27.4 +/- 5.1 microV, respectively) than in T1 (71.9 +/- 4.1 ms and 35.0 +/- 6.5 microV, respectively) (P < 0.05). The relative increase in conduction velocity between T1 and T3 was positively correlated (P < 0.05) with the increase in twitch peak amplitude (r2 = 0.48), with the decrease in twitch time to peak (r2 = 0.43), and with the decrease in action potential amplitude (r2 = 0.50). In conclusion, temperature-induced modifications in fiber membrane conduction properties may have a direct effect on contractile motor unit properties.     

Prolonged muscle vibration increases stretch reflex amplitude, motor unit discharge rate, and force fluctuations in a hand muscle.

The purpose of this study was to compare the influence of prolonged vibration of a hand muscle on the amplitude of the stretch reflex, motor unit discharge rate, and force fluctuations during steady, submaximal contractions. Thirty-two young adults performed 10 isometric contractions at a constant force (5.0 +/- 2.3% of maximal force) with the first dorsal interosseus muscle. Each contraction was held steady for 10 s, and then stretch reflexes were evoked. Subsequently, 20 subjects had vibration applied to the relaxed muscle for 30 min, and 12 subjects received no vibration. The muscle vibration induced a tonic vibration reflex. The intervention (vibration or no vibration) was followed by 2 sets of 10 constant-force contractions with applied stretches (After and Recovery trials). The mean electromyogram amplitude of the short-latency component of the stretch reflex increased by 33% during the After trials (P < 0.01) and by 38% during the Recovery trials (P < 0.01). The standard deviation of force during the steady contractions increased by 21% during the After trials (P < 0.05) and by 28% during the Recovery trials (P < 0.01). The discharge rate of motor units increased from 10.3 +/- 2.7 pulses/s (pps) before vibration to 12.2 +/- 3.1 pps (P < 0.01) during the After trials and to 11.9 +/- 2.6 pps during the Recovery trials (P < 0.01). There was no change in force fluctuations or stretch reflex magnitude for the subjects in the Control group. The results indicate that prolonged vibration increased the short-latency component of the stretch reflex, the discharge rate of motor units, and the fluctuations in force during contractions by a hand muscle. These adjustments were necessary to achieve the target force due to the vibration-induced decrease in the force capacity of the muscle.     

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.     

Motor unit firing patterns of lower leg muscles during isometric plantar flexion with flexed knee joint position

The ankle flexor and extensor muscles are essential for pedal movements associated with car driving. Neuromuscular activation of lower leg muscles is influenced by the posture during a given task, such as the flexed knee joint angle during car driving. This study aimed to investigate the influence of flexion of the knee joint on recruitment threshold-dependent motor unit activity in lower leg muscles during isometric contraction. Twenty healthy participants performed plantar flexor and dorsiflexor isometric ramp contractions at 30 % of the maximal voluntary contraction (MVC) with extended (0°) and flexed (130°) knee joint angles. High-density surface electromyograms were recorded from medial gastrocnemius (MG), soleus (SOL), and tibialis anterior (TA) muscles and decomposed to extract individual motor units. The torque-dependent change (Δpps /Δ%MVC) of the motor unit activity of MG (recruited at 15 %MVC) and SOL (recruited at 5 %MVC) muscles was higher with a flexed compared with an extended knee joint (p < 0.05). The torque-dependent change of TA MU did not different between the knee joint angles. The motor units within certain limited recruitment thresholds recruited to exert plantar flexion torque can be excited to compensate for the loss of MG muscle torque output with a flexed knee joint.     

Motor unit activation patterns during concentric wrist flexion in humans with different muscle fibre composition

Muscle activity was recorded from the flexor carpi radialis muscle during static and dynamic-concentric wrist flexion in six subjects, who had exhibited large differences in histochemically identified muscle fibre composition. Motor unit recruitment patterns were identified by sampling 310 motor units and counting firing rates in pulses per second (pps). During concentric wrist flexion at 30% of maximal exercise intensity the mean firing rate was 27 (SD 13) pps. This was around twice the value of 12 (SD 5) pps recorded during sustained static contraction at 30% of maximal voluntary contraction, despite a larger absolute force level during the static contraction. A similar pattern of higher firing rates during dynamic exercise was seen when concentric wrist flexion at 60% of maximal exercise intensity [30 (SD 14) pps] was compared with sustained static contraction at 60% of maximal voluntary contraction [19 (SD 8) pps]. The increase in dynamic exercise intensity was accomplished by recruitment of additional motor units rather than by increasing the firing rate as during static contractions. No difference in mean firing rates was found among subjects with different muscle fibre composition, who had previously exhibited marked differences in metabolic response during corresponding dynamic contractions. It was concluded that during submaximal dynamic contractions motor unit firing rate cannot be deduced from observations during static contractions and that muscle fibre composition may play a minor role. Accepted: 5 May 1998     

Time course of vasodilation at the onset of repetitive skeletal muscle contractions

To characterize the vasodilatory response in the transition from a single skeletal contraction to a series of contractions, we measured the response of hamster cremaster muscle arterioles associated with four to five skeletal muscle fibers stimulated to contract for one, two, three, or four contractions (250-ms train duration) at 4-s intervals [15 contractions per minute (CPM)] for up to 12 s, at stimulus frequencies of 4, 10, 20, 30, 40, 60, and 80 Hz. To investigate the contribution of contraction frequency, we stimulated muscle fiber bundles at 30 or 60 CPM for 12 s at stimulus frequencies of 4, 20, and 60 Hz. Arteriolar diameters at the site of overlap with the stimulated muscle fibers were measured before and after each contraction. At 15 CPM at 4, 20, and 60 Hz, we observed a peak change in diameter following the first contraction of 1.1 +/- 0.1, 1.6 +/- 0.2, and 2.1 +/- 0.2 mum that almost doubled in response to the second contraction (2.0 +/- 0.1, 3.0 +/- 0.1, and 3.8 +/- 0.1 mum, respectively), but there was no further dilation following the third or fourth contraction. A similar response occurred at all stimulus and contraction frequencies tested. At 30 and 60 CPM at 60 Hz, the plateau after two contractions was followed by a further increase in diameter to a second plateau at 7-8 s. Therefore, the vasodilatory response in the transition from single to multiple contractions had components that were stimulation parameter dependent and independent and showed a plateauing behavior indicative of rapid changes in either the nature and/or concentration of vasodilators released or changes in vascular reactivity.     

Ia-afferent input to motoneurons during shortening and lengthening muscle contractions in humans.

The central nervous system employs different strategies to execute specific motor tasks. Because afferent feedback during shortening and lengthening muscle contractions differs, the neural strategy underlying these tasks may be quite distinct. Cortical drive may be adjusted or afferent input regulated. The exact mechanisms are not clear. Here, we examine the control of synaptic transmission across the Ia synapse during shortening and lengthening muscle contractions. Subjects were instructed to maintain isolated activity in a single tibialis anterior (TA) motor unit while muscle length was varied from flexion to extension and back. At a fixed interval after a firing of the active motor unit, a single electrical stimulus was applied to the common peroneal nerve to activate Ia afferents from the TA muscle. We investigated the stimulus-induced change in firing probability of 19 individual low-threshold TA motor units during shortening and lengthening contractions. Any change in firing probability depends on both pre- and postsynaptic mechanisms. In this experiment, motoneuron firing rate was similar during both contraction types. There was no difference in the firing probability between shortening and lengthening contractions (0.23 +/- 0.03 and 0.20 +/- 0.02, respectively). We suggest that there is no contraction type-specific control of Ia input to the motoneurons during shortening and lengthening muscle contractions. Cortical adjustments may have occurred.     

P2 antagonist PPADS attenuates responses of thin fiber afferents to static contraction and tendon stretch

Injection into the arterial supply of skeletal muscle of pyridoxal phosphate-6-azophenyl-2',4'-disulfonic acid (PPADS), a P2 receptor antagonist, has been shown previously to attenuate the reflex pressor responses to both static contraction and to tendon stretch. In decerebrated cats, we tested the hypothesis that PPADS attenuated the responses of groups III and IV muscle afferents to static contraction as well as to tendon stretch. We found that injection of PPADS (10 mg/kg) into the popliteal artery attenuated the responses of both group III (n = 16 cats) and group IV afferents (n = 14 cats) to static contraction. Specifically, static contraction before PPADS injection increased the discharge rate of the group III afferents from 0.1 +/- 0.05 to 1.6 +/- 0.5 impulses/s, whereas contraction after PPADS injection increased the discharge of the group III afferents from 0.2 +/- 0.1 to only 1.0 +/- 0.5 impulses/s (P < 0.05). Likewise, static contraction before PPADS injection increased the discharge rate of the group IV afferents from 0.3 +/- 0.1 to 1.0 +/- 0.3 impulses/s, whereas contraction after PPADS injection increased the discharge of the group IV afferents from 0.2 +/- 0.1 to only 0.3 +/- 0.1 impulses/s (P < 0.05). In addition, PPADS significantly attenuated the responses of group III afferents to tendon stretch but had no effect on the responses of group IV afferents. Our findings suggest that both groups III and IV afferents are responsible for evoking the purinergic component of the exercise pressor reflex, whereas only group III afferents are responsible for evoking the purinergic component of the muscle mechanoreflex that is evoked by tendon stretch.     

Changes in motor unit discharge rate are not associated with the amount of twitch potentiation in old men.

This study examined, in nine old men (82 +/- 4 yr), whether there is an association between the magnitude of change in motor unit discharge rate and the amount of twitch potentiation after a conditioning contraction (CC). The evoked twitch force and motor unit discharge rate during isometric ramp-and-hold contractions (10-18 s) of the triceps brachii muscle at 10, 20, and 30% of the maximal voluntary contraction were determined before and 10 s, 2 min, 6 min, and 11 min after a 5-s CC at 75% maximal voluntary contraction. After the CC, there was a potentiation of twitch force (approximately twofold), and the discharge rate of the 47 sampled motor units declined (P < 0.05) an average of 1 Hz 10 s after the CC, compared with the control condition. The CC had no effect on the variability (coefficient of variation) of both force and discharge rate, as well as the electromyographic activity recorded over the triceps brachii and biceps brachii muscles. In contrast to our earlier study of young men (Klein CS, Ivanova TD, Rice CL, and Garland SJ, Neurosci Lett 316: 153-156, 2001), the magnitude of the reduction in discharge rate after the CC was not associated (r = 0.06) with the amount of twitch potentiation. These findings suggest an age-related alteration in the neural strategy for adjusting motor output to a muscle after a CC.     

Effect of different ankle- and knee-joint positions on gastrocnemius medialis fascicle length and EMG activity during isometric plantar flexion

The purpose of this study was to provide evidence on the fact that the observed decrease in EMG activity of the gastrocnemius medialis (GM) at pronounced knee flexed positions is not only due to GM insufficiency, by examining muscle fascicle lengths during maximal voluntary contractions at different positions. Twenty-two male long distance runners (body mass: 78.5+/-6.7 kg, height: 183+/-6 cm) participated in the study. The subjects performed isometric maximal voluntary plantar flexion contractions (MVC) of their left leg at six ankle-knee angle combinations. To examine the resultant ankle joint moments the kinematics of the left leg were recorded using a Vicon 624 system with 8 cameras operating at 120 Hz. The EMG activity of GM, gastrocnemius lateralis (GL), soleus (SOL) and tibialis anterior (TA) were measured using surface electromyography. Synchronously, fascicle length and pennation angle values of the GM were obtained at rest and at the plateau of the maximal plantar flexion using ultrasonography. The main findings were: (a) identifiable differences in fascicle length of the GM at rest do not necessarily imply that these differences would also exist during a maximal isometric plantar flexion contraction and (b) the EMG activity of the biarticular GM during the MVC decreased at a pronounced flexed knee-joint position (up to 110 degrees ) despite of no differences in GM fascicle length. It is suggested that the decrease in EMG activity of the GM at pronounced knee flexed positions is due to a critical force-length potential of all three muscles of the triceps surae.     

Motor unit firing rates and contractile properties in tibialis anterior of young and old men.

The effects of aging on motoneuron firing rates and muscle contractile properties were studied in tibialis anterior muscle by comparing results from six young (20.8 +/- 0.8 yr) and six old men (82.0 +/- 1.7 yr). For each subject, data were collected from repeated tests over a 2-wk period. Contractile tests included maximal voluntary contraction (MVC) with twitch interpolation and stimulated twitch contractions. The old men had 26% lower MVC torque (P < 0.01) than did the young men, but percent activation was not different (99.1 and 99.3%, respectively). Twitch contraction durations were 23% longer (P < 0.01) in the old compared with the young men. During a series of repeated brief steady-state contractions at 10, 25, 50, 75, and 100% MVC, motor unit firing rates were recorded. Results from approximately 950 motor unit trains in each subject group indicated that at all relative torque levels mean firing rates were 30-35% lower (P < 0.01) in the old subjects. Comparisons between young and old subjects' mean firing rates at each of 10%, 50%, and MVC torques and their corresponding mean twitch contraction duration yielded a range of moderate-to-high correlations (r = -0.67 to -0.84). That lower firing rates were matched to longer twitch contraction durations in the muscle of old men, and relatively higher firing rates were matched with shorter contraction times from the young men, indirectly supports the neuromuscular age-related remodeling principle.     

Apamin-sensitive nitric oxide- and ATP-mediated motor effects on the guinea pig small intestine

The involvement of nitric oxide and ATP in both spontaneous and electrically-induced nonadrenergic noncholinergic (NANC) motor activity with special interest in the apamin-sensitive mechanisms was studied in a guinea pig ileum model. Depending on the concentration (0.1 or 1 micromol/l), apamin, a blocker of the calcium-activated potassium channels and antagonist of ATP action, induced either TTX (0.1 micromol/l)-resistant increase in tone or contractions. SNP, a nitric oxide donor, applied cumulatively (0.1-100 micromol/l) evoked a concentration-dependent relaxation, the EC50 value being 0.39 +/- 0.12 micromol/l. At concentrations of 0.1 or 1 micromol/l, apamin decreased the SNP effects and shifted the concentration-response curves for SNP to the right. The EC50 value for SNP in the presence of apamin at a concentration of 0.1 micromol/l increased to 59.34 +/- 36.53 micromol/l. ATP (1 or 50 micromol/l) induced TTX-resistant contractions. The effects of ATP were reduced by apamin (1 micromol/l). The contractile effect of ATP occurred in the presence of SNP. SNP provoked relaxation on the background of ATP. The NANC responses to electrical stimulation (0.8 ms, 40 V, 2 or 20 Hz, 20 s) consisted of an initial relaxation phase followed by a phase of contractions, twitch-like and tonic. L-NNA (0.5 mmol/l), an inhibitor of nitric oxide syntheses, abolished the relaxation phase. L-arginine (0.5 mmol/l) restored it. Apamin (0.1 or 1 micromol/l) completely eliminated the relaxation phase and concentration-dependently inhibited the tonic contraction of the phase of contractions. The present findings indicate that the apamin-sensitive nitric oxide-evoked relaxation could be realized by calcium-activated potassium channels and that the apamin-sensitive ATP-induced contraction is mediated via contraction-producing P2 purinoceptors.     

Motor unit firing behavior during prolonged 50% MVC dorsiflexion contractions in young and older adults

The purpose of this study was to investigate changes in motor unit firing behavior during prolonged contractions in young and older adults. Motor unit activity was recorded from the tibialis anterior of 16 subjects (8 young and 8 older), while they performed isometric dorsiflexion at 50% MVC until task failure. Mean motor unit firing rate, the standard deviation (SD), and coefficient of variation (CV) of the interspike intervals, and number of doublet discharges were calculated for a total of 52 motor units, tracked for an average of 92.9 ± 68.6 s. There was no age-related difference in the time to task failure. A modest decline in firing rate was observed in 71% of the motor units, with no significant age-related difference. The SD and CV of the interspike interval had a positive slope in 65% and 69% of the motor units, respectively, with no significant age-related differences. The number of doublet discharges remained stable throughout the contraction. Both groups exhibited motor unit dropout (discharge cessation) during the contraction. Thus, a fatiguing task producing modest changes in firing rate in young and older adults is accompanied by an appreciable increase in firing rate variability. The incidence of doublet discharges is not increased during fatiguing contractions.     

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.     

The role of motor unit rate modulation versus recruitment in repeated submaximal voluntary contractions performed by control and spinal cord injured subjects.

The relative roles of motor unit firing rate modulation and recruitment were evaluated when individuals with cervical spinal cord injury (SCI) and able-bodied controls performed a brief (6 s), 50% maximal voluntary contraction (50% MVC; target contraction) of triceps brachii every 10 s until it required maximal effort to achieve the target force. Mean (+/-SD) endurance times for SCI and control subjects were 34+/-26 and 15+/-5 min, respectively, at which point significant reductions in maximal triceps force had occurred. Twitch occlusion analysis in controls indicated that force declines resulted largely from peripheral contractile failure. In SCI subjects, triceps surface EMG and motor unit potential amplitude declined in parallel suggesting failure at axon branch points and/or alterations in muscle membrane properties. The force of low threshold units, measured by spike-triggered averaging, declined in SCI but not control subjects, suggesting that higher threshold units fatigued in controls. Central fatigue was also obvious after SCI. Mean (+/-SD) MVC motor unit firing rates declined significantly with fatigue for control (24.6+/-7.1 to 17.3+/-5.1Hz), but not SCI subjects (25.9+/-12.7 to 20.1+/-9.7Hz). Unit firing rates were unchanged during target contractions for each subject group, but with the MVC rate decreases, units of SCI and control subjects were activated intensely at endurance time (88% and 99% MVC rates, respectively). New unit recruitment also maintained the target contractions although it was limited after SCI because many descending inputs to triceps motoneurons were disrupted. This resulted in sparse EMG, even during MVCs, but allowed the same unit to be recorded throughout. These EMG data showed that both unit recruitment and rate modulation were important for maintaining force during repeated submaximal intermittent contractions of triceps brachii muscles performed by SCI subjects. Similar results were found for control subjects. Muscles weakened by SCI may therefore provide a useful model in which to directly study motor unit rate modulation and recruitment during weak or strong voluntary contractions.     

Impulse propagation and muscle activation in long maximal voluntary contractions

With fatigue, force generation may be limited by several factors, including impaired impulse transmission and/or reduced motor drive. In 5-min isometric maximal voluntary contraction, no decline was seen in the peak amplitude of the tibialis anterior compound muscle mass action potential (M wave) either during or immediately after the voluntary effort, provided maximal nerve stimulation was retained. For first dorsal interosseous (FDI) muscle, M wave amplitudes declined by 19.4 +/- 1.6% during the first 2 min but did not change significantly thereafter, despite the continued force reduction (up to 94% in 5 min for both muscles). The duration of the FDI M waves increased (greater than 30%), suggesting that the small decline in amplitude was the result of increased dispersion between the responses of different motor units. Some subjects kept FDI maximally activated throughout, but when they used tibialis anterior, twitch occlusion and tetanic muscle stimulation showed that most subjects were usually only able to do so for the first 60 s and thereafter only during brief "extra efforts." Thus force loss during isometric voluntary contractions sustained at the highest intensities results mainly from failure of processes within the muscle fibers.     

Voluntary activation level and muscle fiber recruitment of human quadriceps during lengthening contractions.

Voluntary activation levels during lengthening, isometric, and shortening contractions (angular velocity 60 degrees/s) were investigated by using electrical stimulation of the femoral nerve (triplet, 300 Hz) superimposed on maximal efforts. Recruitment of fiber populations was investigated by using the phosphocreatine-to-creatine ratio (PCr/Cr) of single characterized muscle fibers obtained from needle biopsies at rest and immediately after a series of 10 lengthening, isometric, and shortening contractions (1 s on/1 s off). Maximal voluntary torque was significantly higher during lengthening (270 +/- 55 N.m) compared with shortening contractions (199 +/- 47 N.m, P < 0.05) but was not different from isometric contractions (252 +/- 47 N.m). Isometric torque was higher than torque during shortening (P < 0.05). Voluntary activation level during maximal attempted lengthening contractions (79 +/- 8%) was significantly lower compared with isometric (93 +/- 5%) and shortening contractions (92 +/- 3%, P < 0.05). Mean PCr/Cr values of all fibers from all subjects at rest were 2.5 +/- 0.6, 2.0 +/- 0.7, and 2.0 +/- 0.7, respectively, for type I, IIa, and IIax fibers. After 10 contractions, the mean PCr/Cr values for grouped fiber populations (regardless of fiber type) were all significantly different from rest (1.3 +/- 0.2, 0.7 +/- 0.3, and 0.8 +/- 0.6 for lengthening, isometric, and shortening contractions, respectively; P < 0.05). The cumulative distributions of individual fiber populations after either contraction mode were significantly different from rest (P < 0.05). Curves after lengthening contractions were less shifted compared with curves from isometric and shortening contractions (P < 0.05), with a smaller shift for the type IIax compared with type I fibers in the lengthening contractions. The results indicate a reduced voluntary drive during lengthening contractions. PCr/Cr values of single fibers indicated a hierarchical order of recruitment of all fiber populations during maximal attempted lengthening contractions.