Effects of fatigue duration and muscle type on voluntary and evoked contractile properties

Behm, D. G., and D. M. M. St-Pierre. Effects of fatigueduration and muscle type on voluntary and evoked contractile properties. J. Appl. Physiol. 82(5):1654-1661, 1997.The effects of fatigue duration and muscle typeon voluntary and evoked contractile properties were investigated withan isometric, intermittent, submaximal fatigue protocol. Four groupsperformed contractions of the plantar flexors and quadriceps at variousintensities to produce long (LDF; 19 min 30 s)- and short-durationfatigue (SDF; 4 min 17 s). The LDF group had a significantly greaterdecrease in muscle activation than did the SDF group (12 vs. 5.8%)during recovery, although there was no difference in the impairment of maximum voluntary contraction force beyond 30 s of recovery. The significant decrease in the compound muscle action potential of the LDFgroup (M-wave amplitude; 14.7%) contrasted with the M-wave potentiation of the SDF group (15.7%), suggesting changes in membrane excitation may affect LDF. The quadriceps group performing contractions at 50% MVC experienced a smaller decrease in agonist electromyograph activity than did other groups, indicating both muscle and fatigue duration specificity. Impairments in excitation-contraction coupling were indicated by changes in quadriceps peak twitch and time to peaktwitch while decreases in PF M-wave amplitudes suggested a disruptionof membrane potentials. Results suggest that fatigue mechanisms may beduration (activation, half relaxation time) or muscle specific(electromyograph, twitch torque) or a combination of both (M wave, timeto peak twitch torque).

     

Human neuromuscular fatigue is associated with altered Na+-K+-ATPase activity following isometric exercise.

The purpose of this study was to investigate the hypothesis that reductions in Na+-K+- ATPase activity are associated with neuromuscular fatigue following isometric exercise. In control (Con) and exercised (Ex) legs, force and electromyogram were measured in 14 volunteers [age, 23.4 +/- 0.7 (SE) yr] before and immediately after (PST0), 1 h after (PST1), and 4 h after (PST4) isometric, single-leg extension exercise at ~60% of maximal voluntary contraction for 30 min using a 0.5 duty cycle (5-s contraction, 5-s rest). Tissue was obtained from vastus lateralis muscle before exercise in Con and after exercise in both the Con (PST0) and Ex legs (PST0, PST1, PST4), for the measurements of Na+-K+-ATPase activity, as determined by the 3-O-methylfluorescein phosphatase (3-O-MFPase) assay. Voluntary (maximal voluntary contraction) and elicited (10, 20, 50, 100 Hz) force was reduced 30-55% (P < 0.05) at PST0 and did not recover by PST4. Muscle action potential (M-wave) amplitude and area (measured in the vastus medialis) and 3-O-MFPase activity at PST0-Ex were less than that at PST0-Con (P < 0.05) by 37, 25, and 38%, respectively. M-wave area at PST1-Ex was also less than that at PST1-Con (P < 0.05). Changes in 3-O-MFPase activity correlated to changes in M-wave area across all time points (r = 0.38, P < 0.05, n = 45). These results demonstrate that Na+-K+- ATPase activity is reduced by sustained isometric exercise in humans from that in a matched Con leg and that this reduction in Na+-K+-ATPase activity is associated with loss of excitability as indicated by M-wave alterations.     

Rate of fatigue during repeated submaximal contractions of human quadriceps muscle.

Our purpose was to determine the effect of eight different combinations of contraction intensity, duration, and rest on the rate of fatigue in vastus lateralis muscle. A single combination consisted of contractions at 30 or 70% maximal voluntary contraction (MVC), held for 3 or 7 s with 3- or 7-s rest intervals. Contractions were repeated until the subject could not hold the force for the requisite duration. At regular intervals during each experiment, a brief MVC, a single twitch, and the response to eight stimulation pulses at 50 Hz were elicited. The rate of fatigue was the rate of decline of MVC calculated from regression analysis. Mean rate of fatigue (n = 8) ranged from 0.3 to 25% MVC/min and was closely related (r = 0.98) to the product of the relative force and the duty cycle. Force from 50 Hz stimulation fell linearly and in parallel with MVC. Twitch force was first potentiated and then fell twice as fast as 50 Hz stimulation and MVC (p less than 0.05). Differentiated twitch contraction and relaxation rates were higher at potentiation and lower at the limit of endurance, compared with control values (p less than 0.05). The maximal electromyogram decreased 25% and the submaximal EMG increased to maximal by the end of the protocol, indicating that the entire motor unit pool had been recruited. The close relation between rate of fatigue and the force x time product probably reflects the off-setting interaction of contraction amplitude, duration, and rest interval. This occurs despite the changes in twitch characteristics and the apparent recruitment of fast fatiguing motor units.     

Localized muscular fatigue duration, EMG parameters and accuracy of rapid limb movements

While much is known about the physiological basis of local muscular fatigue, little is known about the kinematic and electromyographic (EMG) consequences of brief fatiguing isometric contractions. Five male subjects performed a horizontal elbow flexion-extension reversal movement over 90° in 250 ms to reversal before and after one of five single maximal isometric elbow flexions ranging in duration from 15–120 s. Surface EMG signals were recorded from the biceps brachii, the long head of the triceps, the clavicular portion of the pectoralis major, and the posterior deltoid. Spatial and temporal errors were computed from potentiometer output. During the fatiguing bouts, maximum voluntary force dropped linearly an average of 4% in the 15 s condition and 58% in the 120 s condition relative to maximum force. The associated biceps rectified-integrated EMG signal increased from the onset of each fatigue bout for 15–30 s, then decreased over the remainder of the longer bouts. Following the fatigue bout, subjects undershot the target distance on the first movement trial in all conditions. Following short fatigue durations (i.e. 15–30 s), the peak biceps EMG amplitude was disrupted and movement velocity decreased, but both measures recovered within seconds. As fatigue duration increased, progressive decreases in peak velocity occurred with increased time to reversal, reduced EMG amplitude, and longer recovery times. However, the relative timing of the EMG pattern was maintained suggesting the temporal structure was not altered by fatigue. The findings suggest that even short single isometric contractions can disrupt certain elements of the motor control system.     

Supraspinal fatigue during intermittent maximal voluntary contractions of the human elbow flexors.

Responses to transcranial magnetic stimulation in human subjects (n = 9) were studied during series of intermittent isometric maximal voluntary contractions (MVCs) of the elbow. Stimuli were given during MVCs in four fatigue protocols with different duty cycles. As maximal voluntary torque fell during each protocol, the torque increment evoked by cortical stimulation increased from approximately 1.5 to 7% of ongoing torque. Thus "supraspinal" fatigue developed in each protocol. The motor evoked potential (MEP) and silent period in the elbow flexor muscles also changed. The silent period lengthened by 20-75 ms (lowest to highest duty cycle protocol) and recovered significantly with a 5-s rest. The MEP increased in area by >50% in all protocols and recovered significantly with 10 s, but not 5 s, of rest. These changes are similar to those during sustained MVC. The central fatigue demonstrated by the torque increments evoked by the stimuli did not parallel the changes in the electromyogram responses. This suggests that part of the fatigue developed during intermittent exercise is "upstream" of the motor cortex.     

Electromyogram and force during stimulated fatigue tests of muscles in dominant and non-dominant hands

Mechanical and electrical properties were studied for the first dorsal interosseous muscle of the dominant (d-FDI) and non-dominant hand (nd-FDI). Observations were made before, during and after a fatigue test, fatigue being evoked by percutaneous electrical stimulation of the ulnar nerve. The test consisted of 30 Hz bursts of ten supramaximal 0.1 ms pulses, repeated once a second for 5 min. The measurements included the amplitude of the first and fifth compound muscle action potentials (M-waves) within bursts, the peak burst force and the amplitude and time course of single twitches. At the end of the fatigue test, burst force had decreased to about the same extent in the FDI of both hands. The final decline in first M-wave amplitude was, however, significantly more pronounced for the nd-FDI than for the d-FDI. There were no longer any significant discrepancies between the two muscles after a subsequent recovery-period of 15 min. Comparisons among nd-FDI of various individuals demonstrated the presence of significant inter-individual differences in fatigue-related force-drop without any associated differences in M-wave decline. Intra-individual variability was similar for fatigue-related force-drop and M-wave decline.     

Interpretation of EMG changes with fatigue: facts, pitfalls, and fallacies.

Failure to maintain the required or expected force, defined as muscle fatigue, is accompanied by changes in muscle electrical activity. Although studied for a long time, reasons for EMG changes in time and frequency domain have not been clear until now. Many authors considered that theory predicted linear relation between the characteristic frequencies and muscle fibre propagation velocity (MFPV), irrespective of the fact that spectral characteristics can drop even without any changes in MFPV, or in proportion exceeding the MFPV changes. The amplitude changes seem to be more complicated and contradictory since data on increased, almost unchanged, and decreased amplitude characteristics of the EMG, M-wave or motor unit potential (MUP) during fatigue can be found in literature. Moreover, simultaneous decrease and increase in amplitude of MUP and M-wave, detected with indwelling and surface electrodes, were referred to as paradoxical. In spite of this, EMG amplitude characteristics are predominantly used when causes for fatigue are analysed. We aimed to demonstrate theoretical grounds for pitfalls and fallacies in analysis of experimental results if changes in intracellular action potential (IAP), i.e. in peripheral factors of muscle fatigue, were not taken into consideration. We based on convolution model of potentials produced by a motor unit and detected by a point or rectangular plate electrode in a homogeneous anisotropic infinite volume conductor. Presentation of MUP in the convolution form gave us a chance to consider power spectrum (PS) of MUP as a product of two terms. The first one, PS of the input signal, represented PS of the first temporal derivative of intracellular action potential (IAP). The second term, PS of the impulse response, took into account MFPV, differences in instants of activation of each fibre, MU anatomy, and MU position in the volume conductor in respect to the detecting electrode. PS presentation through product means that not only changes in MFPV could be responsible for PS shift as is usually assumed. Changes in IAP duration and IAP after-potential magnitude, affecting the first term of the product, influence the product and thus MUP PS. Moreover, the interrelations between the two spectra and thus sensitivity of spectrum to different parameters change with MU-electrode distance because the second term depends on it. Thus, we have demonstrated that theory does not predict a linear relation between the characteristic frequencies (maximum, mean and median) and MFPV. IAP duration and after-potential magnitude are among parameters affecting MUP or M-wave PS and thus, EMG PS detected by monopolar and bipolar electrodes. Usage of single fibre action potential models instead of MUP ones can result in false dependencies of frequency characteristics. The MUP amplitude characteristics are determined not only by amplitude of IAP, but also by the length of the IAP profile and source-electrode distance. Due to the IAP profile lengthening and an increase in the negative after-potential, surface detected EMG amplitude characteristics can increase even when IAP amplitude decreases considerably during fatigue. Increase in surface detected MUP or M-wave amplitude should not be attributed to a weaker attenuation of the low-frequency components with distance. Simultaneous decrease and increase in amplitude of MUP and M-wave detected with indwelling and surface electrodes are regular, not paradoxical. Corner frequency of the high pass filter should be 0.5 or 1 Hz when muscle fatigue is analyzed. The area of MUP or M-wave normalized in respect of the amplitude of the terminal phase (that is produced during extinction of the depolarized zones at the ends of the fibres) could be useful as a fatigue index. Analysing literature data on IAP changes due to Ca(2+) increasing, we hypothesised that the ability of muscle fibres to uptake Ca(2+) back into the sarcoplasmic reticulum could be the limiting site for fatigue. If this hypothesis is valid, IAP changes are not a cause of fatigue; they are due to it.     

Migration of cell surface concanavalin A receptors in pulsed electric fields.

Concanavalin A (con A) receptors on the surface of cultured Xenopus myoblasts redistributed in response to monopolar, pulsed electric fields. The prefield uniform distribution of the receptors became asymmetrical, and was polarized toward the cathodal pole, in the same way as in DC fields. The extent of asymmetry depended on the duration of field exposure, pulse width (or alternatively, interpulse interval), frequency, and intensity. This relationship was most conveniently expressed by using duty cycle, a quantity determined by both pulse width and frequency. Pulses of average intensity 1.5 V/cm induced detectable asymmetry within 5 min. At the lowest average field intensity used, 0.8 V/cm, significant asymmetry was detected at 150 min. For pulses of high duty cycle (greater than 25%), steady state was reached after 30 min exposure and the steady state asymmetry was dependent on average field intensity. For low duty cycle fields, the time required to reach steady state was prolonged (greater than 50 min). Before reaching a steady state, effectiveness of the pulses, as compared with DC fields of equivalent intensity, was a function of duty cycle. A low duty cycle field (fixed number of pulses at low frequency or long interpulse interval) was less effective than high duty cycle fields or DC.     

Cycle time influences the development of muscle fatigue at low to moderate levels of intermittent muscle contraction

Localized muscle fatigue (LMF) during a repetitive task can be influenced by several aspects such as the level and duration of exertions. Among these aspects, though, the influence of cycle time remains unclear. Here, the effect of cycle time on LMF and performance was examined for a simple biomechanical system during repetitive static efforts. Participants performed 1-h trials of intermittent isometric index finger abduction with a duty cycle of 50% in all combinations of two cycle times (30 and 60 s) and two exertion levels (15% and 25% of maximum voluntary capacity). Measures of discomfort, performance (force fluctuations), and muscle capacity (voluntary strength and low-frequency twitch responses) were obtained, all of which demonstrated a beneficial effect of the 30 s cycle time. Specifically, the shorter cycle time led to lower rates of increase in perceived discomfort, lower rates of increase in force fluctuations, lower rates of decrease in voluntary capacity, and smaller changes in twitch responses. These benefits, reflecting less LMF development in the shorter cycle time, were quite consistent between genders and the two levels of effort. Results of this study can be used to modify current models predicting work–rest allowance and/or LMF, helping to enhance performance and reduce the risk of adverse musculoskeletal outcomes.     

Limiting mechanisms of force production after repetitive dynamic contractions in human triceps surae.

The influence of repetitive dynamic fatiguing contractions on the neuromuscular characteristics of the human triceps surae was investigated in 10 subjects. The load was 50% of the torque produced during a maximal voluntary contraction, and the exercise ended when the ankle range of motion declined to 50% of control. The maximal torque of the triceps surae and the electromyographic (EMG) activities of the soleus and medial gastrocnemius were studied in response to voluntary and electrically induced contractions before and after the fatiguing task and after 5 min of recovery. Reflex activities were also tested by recording the Hoffmann reflex (H reflex) and tendon reflex (T reflex) in the soleus muscle. The results indicated that whereas the maximal voluntary contraction torque, tested in isometric conditions, was reduced to a greater extent (P < 0.05) at 20 degrees of plantar flexion (-33%) compared with the neutral position (-23%) of the ankle joint, the EMG activity of both muscles was not significantly reduced after fatigue. Muscle activation, tested by the interpolated-twitch method or the ratio of the voluntary EMG to the amplitude of the muscle action potential (M-wave), as well as the neuromuscular transmission and sarcolemmal excitation, tested by the M-wave amplitude, did not change significantly after the fatiguing exercise. Although the H and T reflexes declined slightly (10-13%; P < 0.05) after fatigue, these adjustments did not appear to have a direct deleterious effect on muscle activation. In contrast, alterations in the mechanical twitch time course and postactivation potentiation indicated that intracellular Ca(2+)-controlled excitation-contraction coupling processes most likely played a major role in the force decrease after dynamic fatiguing contractions performed for short duration.     

Twitch potentiation is greater after a fatiguing submaximal isometric contraction performed at short vs. long quadriceps muscle length.

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.     

Force output during fatigue with progressively increasing stimulation frequency

There is currently a controversy over whether stimulation frequencies should increase or decrease to optimize force output over time. This study compared changes in thenar muscle force and M-wave amplitude during progressively increasing (20–40 Hz), decreasing (40–20 Hz) and constant (20 Hz) frequency stimulation of the median nerve continuously for 3 min. Twenty-three individuals participated in three sets of experiments. There was no significant difference in the force–time integrals between the three fatigue tasks. The rate of fatigue was not correlated to the number of stimulation pulses delivered (20 Hz: 3600, 20–40 and 40–20 Hz: 5400). All fatigue tasks caused a significant reduction in M-wave amplitude and the reduction was largest for the 20–40 Hz protocol. However, multiple linear regression analysis revealed that the M-wave amplitude could not predict the changes in force over time for the 20 Hz or 20–40 Hz protocols. Thus during sustained evoked contractions with stimulation frequencies within the physiological range, frequencies can vary significantly without changing the overall force–time integral.     

Up-regulation of matrix in bovine articular cartilage explants by electric fields

Electric stimulation has long been used as a tool to promote connective tissue healing, but the mechanism(s) by which this is accomplished are not yet known. We have previously determined, using mass cultures of fetal bovine articular chondrocytes, a specific set of capacitively coupled electrical stimulation parameters (e.g., duration of stimulation, response time, amplitude, frequency, and duty cycle) that significantly elevated production of collagen and proteoglycan, and up-regulated type II collagen and aggrecan mRNA expression in vitro. In the present study, we applied our best signal parameters (30-min continuous stimulation (100% duty cycle) followed by a pulsed (1h on, 5h off, 4x/day) 50% (1 min on, 1 min off) duty cycle) to cultures of adult bovine articular cartilage explants and obtained similar results. Since the latter system more closely mimics the in vivo chondrocyte environment, these data argue for the utility of electric stimulation for the maintenance of adult cartilage matrix in situ.     

Variability of the fatigue response of paralyzed skeletal muscle in relation to the time after spinal cord injury: mechanical and electrophysiological characteristics

The aim of this study was to determine the effect of the time after spinal cord injury (less than and greater than 10 months) on the mechanical and electrophysiological characteristics of muscle fatigue of the paralyzed electrically stimulated quadriceps muscle. Morphologically and histochemically, a relationship was observed between muscle fatigue and the delay from injury, revealing a critical period of enzymatic turning and a maximum peak of atrophy around the 10th month after the injury, followed by a long-term stabilization. Knee-torque output and M-wave variables (amplitude, latency, duration, and root mean square, RMS) of two muscular heads of the quadriceps were recorded in 19 paraplegic patients during a 120-s isometric contraction. The fatiguing muscle contraction was elicited by supramaximal continuous 20-Hz electrical stimulation. Compared to the chronic group, the acutely paralyzed group showed a greater resistance to fatigue (amount and rate of force decline, P < or = 0.01), smaller alterations of the M-wave amplitude and RMS, and a limited decrease of the muscle fiber conduction velocity (P < 0.05). Mechanical and electrophysiological changes during fatigue provided a clear functional support of the transformation of skeletal muscle under the lesion and of the existence of a critical period of muscular turn. In conclusion, when considering the artificial restoration of motor function, the evolution of the endurance and force-generating capabilities of the muscle actuator must be taken into account, particularly when tasks require important safety conditions (e.g., standing and walking).     

Sub-MHz bursts of nanosecond pulses excite neurons at paradoxically low electric field thresholds without membrane damage

Neuromodulation applications of nanosecond electric pulses (nsEP) are hindered by their low potency to elicit action potentials in neurons. Excitation by a single nsEP requires a strong electric field which injures neurons by electroporation. We bypassed the high electric field requirement by replacing single nsEP stimuli with high-frequency brief nsEP bursts. In hippocampal neurons, excitation thresholds progressively decreased at nsEP frequencies above 20–200 kHz, with up to 20–30-fold reduction at sub-MHz and MHz rates. For a fixed burst duration, thresholds were determined by the duty cycle, irrespective of the specific nsEP duration, rate, or number of pulses per burst. For 100-μs bursts of 100-, 400-, or 800-ns pulses, the threshold decreased as a power function when the duty cycle exceeded 3–5 %. nsEP bursts were compared with single “long” pulses whose duration and amplitude matched the duration and the time-average amplitude of the burst. Such pulses deliver the same electric charge as bursts, within the same time interval. High-frequency nsEP bursts excited neurons at the time-average electric field 2–3 times below the threshold for a single long pulse. For example, the excitation threshold of 139 ± 14 V/cm for a single 100-μs pulse decreased to 57 ± 8 V/cm for a 100-μs burst of 100-ns, 0.25-MHz pulses (p < 0.001). Applying nsEP in bursts reduced or prevented the loss of excitability in multiple stimulation attempts. Stimulation by high-frequency nsEP bursts is a powerful novel approach to excite neurons at paradoxically low electric charge while also avoiding the electroporative membrane damage.     

Effect of ambient temperature on human skeletal muscle metabolism during fatiguing submaximal exercise.

To examine the effect of ambient temperature on metabolism during fatiguing submaximal exercise, eight men cycled to exhaustion at a workload requiring 70% peak pulmonary oxygen uptake on three separate occasions, at least 1 wk apart. These trials were conducted in ambient temperatures of 3 degrees C (CT), 20 degrees C (NT), and 40 degrees C (HT). Although no differences in muscle or rectal temperature were observed before exercise, both muscle and rectal temperature were higher (P < 0.05) at fatigue in HT compared with CT and NT. Exercise time was longer in CT compared with NT, which, in turn, was longer compared with HT (85 +/- 8 vs. 60 +/- 11 vs. 30 +/- 3 min, respectively; P < 0.05). Plasma epinephrine concentration was not different at rest or at the point of fatigue when the three trials were compared, but concentrations of this hormone were higher (P < 0.05) when HT was compared with NT, which in turn was higher (P < 0.05) compared with CT after 20 min of exercise. Muscle glycogen concentration was not different at rest when the three trials were compared but was higher at fatigue in HT compared with NT and CT, which were not different (299 +/- 33 vs. 153 +/- 27 and 116 +/- 28 mmol/kg dry wt, respectively; P < 0.01). Intramuscular lactate concentration was not different at rest when the three trials were compared but was higher (P < 0.05) at fatigue in HT compared with CT. No differences in the concentration of the total intramuscular adenine nucleotide pool (ATP + ADP + AMP), phosphocreatine, or creatine were observed before or after exercise when the trials were compared. Although intramuscular IMP concentrations were not statistically different before or after exercise when the three trials were compared, there was an exercise-induced increase (P < 0.01) in IMP. These results demonstrate that fatigue during prolonged exercise in hot conditions is not related to carbohydrate availability. Furthermore, the increased endurance in CT compared with NT is probably due to a reduced glycogenolytic rate.     

Mechanical and metabolic alterations in rat diaphragm during electrical stimulation

To investigate the hypothesis that the rate of fatigue development is not influenced by the absolute duration of contraction (train duration) and relaxation (off-phase of duty cycle) at constant duty cycle, strips of the diaphragm from 36 male adult rats (mean +/- SD wt 152 +/- 21 g) were stimulated directly for periods of 180, 250, and 320 ms at a constant duty cycle of 50%. The frequency of stimulation was adjusted to produce 40% of maximal tetanic tension at supramaximal voltages. After 30 min of stimulation, analysis of twitch characteristics between control and experimental groups indicated a prolongation of contraction time of 9% (P less than 0.05), an increase in relaxation time of 75% (P less than 0.05), and a decrease in twitch tension by 78% (P less than 0.05). Similarly, reductions (P less than 0.05) in isometric force output at high stimulation frequency (100 Hz) of 58% and at low frequency (20 Hz) of 67% were also noted. These changes were accompanied by an approximately 60% reduction in the maximal velocity of shortening. No difference was observed for any of the mechanical measures between experimental conditions. After 30-min stimulation, decreases of between 43 and 46% were noted for ATP (P less than 0.05) and increases of between three- and fourfold noted for IMP (P less than 0.05). No changes were found for either ADP or AMP. Total adenine nucleotide concentrations declined (P less than 0.05) an average of 24%. As with the mechanical data, no differences were found between the different stimulation conditions. It is concluded that for the conditions studied, fatigue mechanisms become manifest early in the stimulation period and are only minimally altered by the duration of specific contractions provided the relaxation period is of equal duration.     

Amplitude-related characteristics of motor unit and M-wave potentials during fatigue. A simulation study using literature data on intracellular potential changes found in vitro.

To realize possible reasons for changes in EMG amplitude characteristics with fatigue, we analyzed motor unit potentials (MUPs) and M-waves under simultaneous variations of the intracellular action potential (IAP) amplitude, duration, and shape as well as of the muscle fiber propagation velocity and desynchronization in activation of individual muscle fibers. Analysis was performed through computer simulation of MUPs and M-waves detected at different distances from active fibers in infinite anisotropic volume conductor. Changes in the IAP spike and negative after-potential were taken from in vitro experiments reported in the literature. It was shown that the amplitudes of MUP and M-wave detected simultaneously at different distances could decrease close to the active fibers, be almost unchanged at middle distances, and increase far from the fibers even under IAP amplitude decreasing. This reflected the distance-dependent effects of changes in the IAP profile along the fiber. Electrode position affected sensitivity of MUP and M-wave durations to changes in the IAP duration and propagation velocity. Thus, the signal area and RMS depended on electrode position and could change with fatigue in a way different from that of signal amplitude. The results can help to avoid misleading interpretation of EMG changes.     

Low frequency fatigue in human quadriceps is fatigue dependent and not task dependent.

It is well accepted that a low intensity/long duration isometric contraction induces more low frequency fatigue (LFF) compared to a high-intensity/short-duration contraction. However, previous reports examined the intensity/duration of the contraction but did not control the level of fatigue when concluding fatigue is task dependent. The purpose of this study was to determine whether a long duration/low intensity fatiguing contraction would induce greater LFF than a short duration/high-intensity contraction when the quadriceps muscle was fatigued to similar levels. Eighteen healthy male subjects performed quadriceps contractions sustained at 35% and 65% of maximal voluntary contraction (MVC) on separate days, until the tasks induced a similar amount of fatigue (force generating capacity=45% MVC). Double pulse torque to single pulse torque ratio (D/S ratio) was obtained before, immediately and 5min after fatigue along with the electromyographic (EMG) signal from vastus medialis (VM) and rectus femoris (RF). The D/S ratio significantly (p<0.05) increased by 8.7+/-8.5% (mean+/-SD) and 10.2+/-9.2% after 35% and 65% tasks, respectively, and remained elevated 5min into recovery; however, there was no significant difference in ratio between the two sessions immediately or 5min post-fatigue (p>0.05) even though the endurance time for the 35% fatigue task (124+/-39.68s) was significantly longer (p=0.05) than that of the 65% task (63+/-17.73s). EMG amplitude and median power frequency (MPF) analysis also did not reveal any significant differences between these two sessions after fatigue. These findings indicate that LFF fatigue is fatigue dependent as well as task intensity/duration dependent. These findings assist us in understanding task dependency and muscle fatigue.     

Neuromuscular electrical stimulation for skeletal muscle function

Lack of neural innervation due to neurological damage renders muscle unable to produce force. Use of electrical stimulation is a medium in which investigators have tried to find a way to restore movement and the ability to perform activities of daily living. Different methods of applying electrical current to modify neuromuscular activity are electrical stimulation (ES), neuromuscular electrical stimulation (NMES), transcutaneous electrical nerve stimulation (TENS), and functional electrical stimulation (FES). This review covers the aspects of electrical stimulation used for rehabilitation and functional purposes. Discussed are the various parameters of electrical stimulation, including frequency, pulse width/duration, duty cycle, intensity/amplitude, ramp time, pulse pattern, program duration, program frequency, and muscle group activated, and how they affect fatigue in the stimulated muscle.