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Low-frequency and high-frequency wide-pulse neuromuscular electrical stimulation (NMES) can generate extra torque (ET) via afferent pathways. Superimposing tendon vibration (TV) to NMES can increase the activation of these afferent pathways and favor ET generation. Knowledge of the characteristics of ET is essential to implement these stimulation paradigms in clinical practice. Thus, we aimed to investigate the effects of frequency and TV superimposition on the occurrence and magnitude of ET in response to wide-pulse NMES. NMES-induced isometric plantar flexion torque was recorded in 30 healthy individuals who performed five NMES protocols: wide-pulse low-frequency (1 ms; 20 Hz; WPLF) and wide-pulse high-frequency (1 ms; 100 Hz; WPHF) without and with superimposed TV (1 mm; 100 Hz) and conventional NMES (50 µs; 20 Hz; reference protocol). Each NMES protocol consisted of three 20-s trains interspersed by 90 s of rest, with NMES intensity being adjusted to reach 10% of maximal voluntary contraction. The ET occurrence was similar for WPLF and WPHF ( P = 0.822). In the responders, the ET magnitude was greater for WPHF than WPLF ( P < 0.001). There was no effect of superimposed TV on ET characteristics. This study reported an effect of NMES frequency on ET magnitude, whereas TV superimposition did not affect this parameter. In the context of our experimental design decisions, the present findings question the clinical use of wide-pulse NMES and its combination with superimposed TV. Yet, further research is needed to maximize force production through the occurrence and magnitude of ET. NEW & NOTEWORTHY This study is the first to assess the effect of stimulation frequency and superimposed tendon vibration on extra torque characteristics generated by wide-pulse neuromuscular electrical stimulation. The percentage of subjects showing extra torque (i.e., considered as responders) was similar for low-frequency and high-frequency wide-pulse neuromuscular electrical stimulation. In the responders, the extra torque was greater for high-frequency than for low-frequency wide-pulse neuromuscular electrical stimulation. The superimposition of tendon vibration had no effect on extra torque occurrence or magnitude.

The interaction between the somatosensory and motor systems is important for normal human motor function and learning. Enhancing somatosensory input using somatosensory electrical stimulation (SES) can increase motor performance, but the neuronal mechanisms underlying these effects are largely unknown. With EEG, we examined whether skill acquisition, consolidation, and interlimb transfer after SES was related to increased activity in sensorimotor regions, as assessed by the N30 somatosensory evoked potential or rather increased connectivity between these regions, as assessed by the phase slope index (PSI). Right- and left-hand motor performance and EEG measures were taken before, immediately after, and 24 h ( day 2) after either SES ( n = 12; 5 men) or Control ( n = 12; 5 men). The results showed skill acquisition and consolidation in the stimulated right hand immediately after SES (6%) and on day 2 (9%) and interlimb transfer to the nonstimulated left hand on day 2 relative to Control (8%, all P < 0.05). Increases in N30 amplitudes correlated with skill acquisition while PSI from electrodes that represent the posterior parietal and primary somatosensory cortex to the electrode representing the primary motor cortex correlated with skill consolidation. In contrast, interlimb transfer did not correlate with the EEG-derived neurophysiological estimates obtained in the present study, which may indicate the involvement of subcortical structures in interlimb transfer after SES. In conclusion, weak peripheral somatosensory inputs in the form of SES improve skill acquisition, consolidation, and interlimb transfer that coincide with different cortical adaptations, including enhanced N30 amplitudes and PSI. NEW & NOTEWORTHY The relationship between adaptations in synaptic plasticity and motor learning following somatosensory electrical stimulation (SES) is incompletely understood. Here, we used for the first time a multifactorial approach that examined skill acquisition, consolidation, and interlimb transfer following 20 min of SES. In addition, we quantified sensorimotor integration and the magnitude and direction of connectivity with EEG. Following artificial electrical stimulation, increases in sensorimotor integration and connectivity were found to correlate with skill acquisition and consolidation, respectively.

We compared the extent and origin of muscle fatigue induced by short-pulse-low-frequency [conventional (CONV)] and wide-pulse-high-frequency (WPHF) neuromuscular electrical stimulation. We expected CONV contractions to mainly originate from depolarization of axonal terminal branches (spatially determined muscle fiber recruitment) and WPHF contractions to be partly produced via a central pathway (motor unit recruitment according to size principle). Greater neuromuscular fatigue was, therefore, expected following CONV compared with WPHF. Fourteen healthy subjects underwent 20 WPHF (1 ms-100 Hz) and CONV (50 μs-25 Hz) evoked isometric triceps surae contractions (work/rest periods 20:40 s) at an initial target of 10% of maximal voluntary contraction (MVC) force. Force-time integral of the 20 evoked contractions (FTI) was used as main index of muscle fatigue; MVC force loss was also quantified. Central and peripheral fatigue were assessed by voluntary activation level and paired stimulation amplitudes, respectively. FTI in WPHF was significantly lower than in CONV (21,717 ± 11,541 vs. 37,958 ± 9,898 N·s P<0,001). The reductions in MVC force (WPHF: -7.0 ± 2.7%; CONV: -6.2 ± 2.5%; P < 0.01) and paired stimulation amplitude (WPHF: -8.0 ± 4.0%; CONV: -7.4 ± 6.1%; P < 0.001) were similar between conditions, whereas no change was observed for voluntary activation level (P > 0.05). Overall, our results showed a different motor unit recruitment pattern between the two neuromuscular electrical stimulation modalities with a lower FTI indicating greater muscle fatigue for WPHF, possibly limiting the presumed benefits for rehabilitation programs., (Copyright © 2014 the American Physiological Society.)

The extent and characteristics of muscle fatigue of different muscle groups when subjected to a similar fatiguing task may differ. Thirteen healthy young men performed sustained contractions at 50% maximal voluntary contraction (MVC) force until task failure, with four different muscle groups, over two sessions. Per session, one upper limb and one lower limb muscle group were tested (knee extensors and thumb adductor, or plantar and elbow flexors). Changes in voluntary activation level and contractile properties were derived from doublet responses evoked during and after MVCs before and after exercise. Time to task failure differed (P < 0.05) between muscle groups (220 ± 64 s for plantar flexors, 114 ± 27 s for thumb adductor, 77 ± 25 s for knee extensors, and 72 ± 14 s for elbow flexors). MVC force loss immediately after voluntary task failure was similar (-30 ± 11% for plantar flexors, -37 ± 13% for thumb adductor, -34 ± 15% for knee extensors, and -40 ± 12% for elbow flexors, P > 0.05). Voluntary activation was decreased for plantar flexors only (from 95 ± 5% to 82 ± 9%, P < 0.05). Potentiated evoked doublet amplitude was more depressed for upper limb muscles (-59.3 ± 14.7% for elbow flexors and -60.1 ± 24.1% for thumb adductor, P < 0.05) than for knee extensors (-28 ± 15%, P < 0.05); no reduction was found in plantar flexors (-7 ± 12%, P > 0.05). In conclusion, despite different times to task failure when sustaining an isometric contraction at 50% MVC force for as long as possible, diverse muscle groups present similar loss of MVC force after task failure. Thus the extent of muscle fatigue is not affected by time to task failure, whereas this latter determines the etiology of fatigue.

The aim of the present study was to define the chronic effects of neuromuscular electrical stimulation (NMES) on the neuromuscular properties of human skeletal muscle. Eight young healthy male subjects were subjected to 25 sessions of isometric NMES of the quadriceps muscle over an 8-wk period. Needle biopsies were taken from the vastus lateralis muscle before and after training. The training status, myosin heavy chain (MHC) isoform distribution, and global protein pattern, as assessed by proteomic analysis, widely varied among subjects at baseline and prompted the identification of two subgroups: an "active" (ACT) group, which performed regular exercise and had a slower MHC profile, and a sedentary (SED) group, which did not perform any exercise and had a faster MHC profile. Maximum voluntary force and neural activation significantly increased after NMES in both groups (+∼30% and +∼10%, respectively). Both type 1 and 2 fibers showed significant muscle hypertrophy. After NMES, both groups showed a significant shift from MHC-2X toward MHC-2A and MHC-1, i.e., a fast-to-slow transition. Proteomic maps showing ∼500 spots were obtained before and after training in both groups. Differentially expressed proteins were identified and grouped into functional categories. The most relevant changes regarded 1) myofibrillar proteins, whose changes were consistent with a fast-to-slow phenotype shift and with a strengthening of the cytoskeleton; 2) energy production systems, whose changes indicated a glycolytic-to-oxidative shift in the metabolic profile; and 3) antioxidant defense systems, whose changes indicated an enhancement of intracellular defenses against reactive oxygen species. The adaptations in the protein pattern of the ACT and SED groups were different but were, in both groups, typical of both resistance (i.e., strength gains and hypertrophy) and endurance (i.e., a fast-to-slow shift in MHC and metabolic profile) training. These training-induced adaptations can be ascribed to the peculiar motor unit recruitment pattern associated with NMES.

This study aimed to 1) compare electrical and magnetic stimulations for quadriceps muscle function assessment, and 2) ascertain whether the ratios of the second twitch elicited by supramaximal electrical and magnetic femoral nerve stimulation at 10 and 100 Hz (T2(10:100)) and the total twitch force elicited by the same types of stimulations (Fpaired(10:100)) are equivalent to the standard low- to high-frequency force ratio associated with submaximal electrical tetanic stimulations (Ftet(10:100)). Quadriceps force and vastus lateralis EMG were recorded at rest (n = 21 subjects), immediately after, and 30 min after a 30-min downhill run (n = 10) when 1) supramaximal electrical nerve stimulation (ENS), 2) magnetic nerve stimulation (MNS) and 3) submaximal electrical muscle stimulation (EMS) were delivered in random order at 1 (single stimulation), 10, and 100 Hz (paired stimulations). Ten- and 100-Hz 500-ms tetani were also evoked with EMS to determine Ftet(10:100). Before exercise, contractile properties with single and paired stimulations were similar for ENS and MNS (all intraclass correlation coefficients k > 0.90), but smaller for EMS (P < 0.001). M-wave characteristics were also similar for ENS and MNS (all k > 0.90). After exercise, changes in all parameters did not differ between methods. With fatigue, the changes in Ftet(10:100) were inconsistently correlated with the changes in T2(10:100) (r(2) = 0.24-0.73, P = 0.002-0.15) but better correlated with the changes in Fpaired(10:100) (immediately after exercise: r(2) = 0.80-0.83, P < 0.001; 30 min after exercise: r(2) = 0.46-0.82, P = 0.001-0.03). We conclude that ENS and MNS provide similar quadriceps muscle function assessment, while Fpaired(10:100) is a better index than T2(10:100) of low- to high-frequency fatigue of the quadriceps in vivo.

This study aimed to compare voluntary and stimulated exercise for changes in muscle strength, growth hormone (GH), blood lactate, and markers of muscle damage. Nine healthy men had two leg press exercise bouts separated by 2 wk. In the first bout, the quadriceps muscles were stimulated by biphasic rectangular pulses (75 Hz, duration 400 mus, on-off ratio 6.25-20 s) with current amplitude being consistently increased throughout 40 contractions at maximal tolerable level. In the second bout, 40 voluntary isometric contractions were performed at the same leg press force output as the first bout. Maximal voluntary isometric strength was measured before and after the bouts, and serum GH and blood lactate concentrations were measured before, during, and after exercise. Serum creatine kinase (CK) activity and muscle soreness were assessed before, immediately after, and 24, 48, and 72 h after exercise. Maximal voluntary strength decreased significantly (P < 0.05) after both bouts, but the magnitude of the decrease was significantly (P < 0.05) greater for the stimulated contractions (-22%) compared with the voluntary contractions (-9%). Increases in serum GH and lactate concentrations were significantly (P < 0.05) larger after the stimulation compared with the voluntary exercise. Increases in serum CK activity and muscle soreness were also significantly (P < 0.05) greater for the stimulation than voluntary exercise. It was concluded that a single bout of electrical stimulation exercise resulted in greater GH response and muscle damage than voluntary exercise.

Endurance time of a submaximal sustained contraction is longer when the muscle is fatigued in a shortened position. The aim of the present study was to compare central and peripheral mechanisms of fatigue after an isometric contraction of the knee extensor muscles performed at 20% maximal voluntary contraction (MVC) at two knee angles (35 degrees , short length vs. 75 degrees , long length; 0 degrees = full extension) until exhaustion. Eleven men (24 +/- 4 yr) attended two experimental randomized sessions. Endurance time was greater at 35 degrees compared with 75 degrees (974 +/- 457 vs. 398 +/- 144 s; P < 0.001) despite a similar reduction in knee extensor MVC (-28.4 +/- 16.0%, P < 0.001 vs. -27.6 +/- 18.8%, P < 0.001, respectively). Voluntary activation level was similarly depressed after the fatiguing contraction performed at the two muscle lengths (-19 +/- 16.7% at 35 degrees , P < 0.01 vs. -13.7 +/- 14.5% at 75 degrees , P < 0.01). After the fatiguing contraction, peak twitch potentiation was observed only at the short length (+31.8 +/- 17.6% at 35 degrees , P < 0.01 vs. +6.4 +/- 21.3% at 75 degrees , P > 0.05), whereas M-wave properties were similarly altered for the two angles. These results suggest that 1) central fatigue at task failure for a sustained isometric contraction was not dependent on the muscle length, and 2) the longer endurance time of a sustained isometric contraction performed at a shortened length is related to potentiation. It is suggested that the greater endurance time of a sustained isometric contraction observed at 35 degrees is related to the occurrence of potentiation at this short length, because central fatigue is similar at task failure for both tasks.

Neuromuscular adaptations of the plantar flexor muscles were assessed before and subsequent to short-term electromyostimulation (EMS) training. Eight subjects underwent 16 sessions of isometric EMS training over 4 wk. Surface electromyographic (EMG) activity and torque obtained under maximal voluntary and electrically evoked contractions were analyzed to distinguish neural adaptations from contractile changes. After training, plantar flexor voluntary torque significantly increased under isometric conditions at the training angle (+8.1%, P < 0.05) and at the two eccentric velocities considered (+10.8 and +13.1%, P < 0.05). Torque gains were accompanied by higher normalized soleus EMG activity and, in the case of eccentric contractions, also by higher gastrocnemii EMG (P < 0.05). There was an 11.9% significant increase in both plantar flexor maximal voluntary activation (P < 0.01) and postactivation potentiation (P < 0.05), whereas contractile properties did not change after training. In the absence of a change in the control group, it was concluded that an increase in neural activation likely mediates the voluntary torque gains observed after short-term EMS training.

The effects of prolonged cycling on neuromuscular parameters were studied in nine endurance-trained subjects during a 5-h exercise sustained at 55% of the maximal aerobic power. Torque during maximal voluntary contraction (MVC) of the quadriceps muscle decreased progressively throughout the exercise (P < 0.01) and was 18% less at the end of exercise compared with the preexercise value. Peak twitch torque, contraction time, and total area of mechanical response decreased significantly (P < 0.05) after the first hour of exercise. In contrast, changes in M-wave characteristics were significant only after the fourth hour of the exercise. Significant reductions (P < 0.05) in electromyographic activity normalized to the M wave occurred after the first hour for the vastus lateralis muscle but only at the end of the exercise for the vastus medialis muscle. Muscle activation level, assessed by the twitch interpolation technique, decreased by 8% (P < 0.05) at the end of the exercise. The results suggest that the time course is such that the contractile properties are significantly altered after the first hour, whereas excitability and central drive are more impaired toward the latter stages of the 5-h cycling exercise.

Neuromuscular fatigue of the knee extensor (KE) and plantar flexor (PF) muscles was characterized after a 65-km ultramarathon race in nine well-trained runners by stimulating the femoral and tibial nerves, respectively. One week before and immediately after the ultramarathon, maximal twitches were elicited from the relaxed KE and PF. Electrically evoked superimposed twitches of the KE were also elicited during maximal voluntary contractions (MVCs) to determine maximal voluntary activation. MVC and maximal voluntary activation decreased significantly after the ultramarathon (-30.2 +/- 18.0% and -27.7 +/- 13.0%, respectively; P < 0.001). Surprisingly, peak twitch increased after the ultramarathon from 15.8 +/- 6.3 to 19.7 +/- 3.3 N. m for PF (P < 0.01) and from 131.9 +/- 21.2 to 157.1 +/- 35.9 N for KE (P < 0.05). Also, shorter contraction and half-relaxation times were observed for both muscles. The compound muscle action potentials (M wave) were not significantly altered by the ultramarathon with the exception of the soleus, which showed a slightly higher M-wave amplitude after the running. From these results, it can be concluded that 65 km of running 1) severely depressed the maximal voluntary force capacity mainly because of a decrease in maximal voluntary activation, 2) potentiated the twitch mechanical response, and 3) did not change significantly the M-wave characteristics.

The aim of this study was to compare the mechanical and electromyographic (EMG) characteristics of soleus motor units activated during maximal H reflex and direct M response among subjects with different histories of physical activity. Power-trained athletes produced stronger twitches, with a higher rate of twitch tension buildup and relaxation, than their endurance counterparts for both maximal H-reflex and maximal M-wave responses. The maximal H-reflex-to-maximal M-wave ratios for both force output (twitch) and EMG wave amplitude were significantly lower in power-trained than endurance-trained athletes. However, power-trained athletes exhibited a significantly greater twitch-to-EMG ratio for the reflexly activated motor units with respect to the entire motor pool, whereas endurance-trained athletes had comparable twitch-to-EMG ratios for both reflexly and directly activated units. Power training increases the force output of the whole ensemble of the motor units, thereby compensating for the lower efficacy of the reflex transmission between Ia spindle afferent input and soleus alpha-motoneuron. On the other hand, the lower level of force evoked by the reflexly activated units in endurance-trained athletes is associated with a greater motor pool reflex excitability. Therefore, endurance-trained athletes produce the necessary force by recruitment of more slow-twitch units than do other subjects for comparable levels of force and type of task.