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.     

Different plantar flexors neuromuscular and mechanical characteristics depending on the preferred running form

Two main types of endurance runners have been identified: aerial runners (AER), who have a larger flight time, and terrestrial runners (TER), who have a longer ground contact time. The purpose of this study was to assess the neuromuscular characteristics of plantar flexors between AER and TER runners. Twenty-four well-trained runners participated in the experiment. They were classified either in a TER or AER group according to the Volodalen® scale. Plantar flexors’ maximal rate of force development (RFD) and maximal voluntary contraction force (MVC) were assessed. Percutaneous electrical stimulation was delivered to the posterior tibial nerve to evoke maximal M-waves and H-reflexes of the triceps surae muscles. These responses, as well as voluntary activation, muscle potentiation, and V-waves, were recorded by superimposing stimulations to MVCs. RFD was significantly higher in AER than in TER, while MVC remained unchanged. This was accompanied by higher myoelectrical activity recorded in the soleus muscle. While M-waves and other parameters remained unchanged, maximal H-reflex was significantly higher in AER than in TER, still in soleus only. The present study raised the possibility of different plantar flexors’ neuromuscular characteristics according to running profile. These differences seemed to be focused on the soleus rather than on the gastrocnemii.     

Turning on the central contribution to contractions evoked by neuromuscular electrical stimulation.

Neuromuscular electrical stimulation can generate contractions through peripheral and central mechanisms. Direct activation of motor axons (peripheral mechanism) recruits motor units in an unnatural order, with fatigable muscle fibers often activated early in contractions. The activation of sensory axons can produce contractions through a central mechanism, providing excitatory synaptic input to spinal neurons that recruit motor units in the natural order. Presently, we quantified the effect of stimulation frequency (10-100 Hz), duration (0.25-2 s of high-frequency bursts, or 20 s of constant-frequency stimulation), and intensity [1-5% maximal voluntary contraction (MVC) torque generated by a brief 100-Hz train] on the torque generated centrally. Electrical stimulation (1-ms pulses) was delivered over the triceps surae in eight subjects, and plantar flexion torque was recorded. Stimulation frequency, duration, and intensity all influenced the magnitude of the central contribution to torque. Central torque did not develop at frequencies < or = 20 Hz, and it was maximal at frequencies > or = 80 Hz. Increasing the duration of high-frequency stimulation increased the central contribution to torque, as central torque developed over 11 s. Central torque was greatest at a relatively low contraction intensity. The largest amount of central torque was produced by a 20-s, 100-Hz train (10.7 +/- 5.5 %MVC) and by repeated 2-s bursts of 80- or 100-Hz stimulation (9.2 +/- 4.8 and 10.2 +/- 8.1% MVC, respectively). Therefore, central torque was maximized by applying high-frequency, long-duration stimulation while avoiding antidromic block by stimulating at a relatively low intensity. If, as hypothesized, the central mechanism primarily activates fatigue-resistant muscle fibers, generating muscle contractions through this pathway may improve rehabilitation applications.     

Role of NO in modulating neuronal activity in superficial dorsal horn of spinal cord during exercise pressor reflex

Static contraction of hindlimb skeletal muscle in cats induces a reflex pressor response. The superficial dorsal horn of the spinal cord is the major site of the first synapse of this reflex. In this study, static contraction of the triceps surae muscle was evoked by electrical stimulation of the tibial nerve for 2 min in anesthetized cats (stimulus parameters: two times motor threshold at 30 Hz, 0.025-ms duration). Ten stimulations were performed and 1-min rest was allowed between stimulations. Muscle contraction caused a maximal increase of 32 +/- 5 mmHg in mean arterial pressure (MAP), which was obtained from the first three contractions. Activated neurons in the superficial dorsal horn were identified by c-Fos protein. Distinct c-Fos expression was present in the L6-S1 level of the superficial dorsal horn ipsilateral to the contracting leg (88 +/- 14 labeled cells per section at L7), whereas only scattered c-Fos expression was observed in the contralateral superficial dorsal horn (9 +/- 2 labeled cells per section, P < 0.05 compared with ipsilateral section). A few c-Fos-labeled cells were found in control animals (12 +/- 5 labeled cells per section, P < 0.05 compared with stimulated cats). Furthermore, double-labeling methods demonstrated that c-Fos protein coexisted with nitric oxide (NO) synthase (NOS) positive staining in the superficial dorsal horn. Finally, an intrathecal injection of an inhibitor of NOS, N-nitro-L-arginine methyl ester (5 mM), resulted in fewer c-Fos-labeled cells (58 +/- 12 labeled cells per section) and a reduced maximal MAP response (20 +/- 3 mmHg, P < 0.05). These results suggest that the exercise pressor reflex induced by static contraction is mediated by activation of neurons in the superficial dorsal horn and that formation of NO in this region is involved in modulating the activated neurons and the pressor response to contraction.     

Comparison of the effects of botulinum toxin in adult and neonatal rats: neuromuscular blockade and toxicity

A single dose of botulinum toxin (BoTX) was injected subcutaneously to induce neuromuscular blockade in the triceps surae muscles of the hindlimbs of neonatal and adult rats. The efficacy of the toxin in producing complete neuromuscular blockade of the lower limb muscles, assessed by blockade of (a) postural and flexor reflexes and (b) muscle contraction in response to nerve stimulation, was dose dependent at all ages over a BoTX dose range of 10-60 ng/kg. However, BoTX was dramatically more toxic in adult animals resulting in a decline in body weight and lethal consequences in 25% of adult animals 1 week after administration of BoTX doses as low as 40 ng/kg. In contrast, neonatal animals, given the same dosage, continued to grow and no mortalities were observed. The differences in toxicity of BoTX in adult and neonatal rats are readily accounted for by the short duration of effect in the younger animals, which, in turn, is probably the result of more rapid generation of new and functional nerve terminals.     

The pressor response to voluntary and electrically evoked isometric contractions in man

Blood pressure and heart rate changes during sustained isometric exercise were studied in 11 healthy male volunteers. The responses were measured during voluntary and involuntary contractions of the biceps brachii at 30% of maximal voluntary contraction (MVC), and the triceps surae at 30% and 50% MVC. Involuntary contractions were evoked by percutaneous electrical stimulation of the muscle. Measurements of the time to peak tension of maximal twitch showed the biceps brachii (67.0 +/- 7.9 ms) muscle to be rapidly contracting, and the triceps surae (118.0 +/- 10.5 ms) to be slow contracting. The systolic and diastolic blood pressures increased linearly throughout the contractions, and systolic blood pressure increased more rapidly than diastolic. There was no significant difference in response to stimulated or voluntary contractions, nor was there any significant difference between the responses to contractions of the calf or arm muscles at the same relative tension. In contrast the heart rate rose to a higher level (P less than 0.01) in the biceps brachii than the triceps surae at given % MVC, and during voluntary compared with the electrically evoked contractions in the two muscle groups. It was concluded that the arterial blood pressure response to isometric contractions, unlike heart rate, is primarily due to a reflex arising within the active muscles (cf. Hultman and Sj?holm 1982) which is associated with relative tension but independent of contraction time and muscle mass.     

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.     

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

Neuromuscular electrical stimulation (NMES) can be delivered over a nerve trunk or muscle belly and both can generate contractions through peripheral and central pathways. Generating contractions through peripheral pathways is associated with a nonphysiological motor unit recruitment order, which may limit the efficacy of NMES rehabilitation. Presently, we compared recruitment through peripheral and central pathways for contractions of the knee extensors evoked by NMES applied over the femoral nerve vs. the quadriceps muscle. NMES was delivered to evoke 10 and 20% of maximum voluntary isometric contraction torque 2-3 s into the NMES (time(1)) in two patterns: 1) constant frequency (15 Hz for 8 s); and 2) step frequency (15-100-15 Hz and 25-100-25 Hz for 3-2-3 s, respectively). Torque and electromyographic activity recorded from vastus lateralis and medialis were quantified at the beginning (time(1)) and end (time(2); 6-7 s into the NMES) of each pattern. M-waves (peripheral pathway), H-reflexes, and asynchronous activity (central pathways) during NMES were quantified. Torque did not differ regardless of NMES location, pattern, or time. For both muscles, M-waves were ～7-10 times smaller and H-reflexes ～8-9 times larger during NMES over the nerve compared with over the muscle. However, unlike muscles studied previously, neither torque nor activity through central pathways were augmented following 100 Hz NMES, nor was any asynchronous activity evoked during NMES at either location. The coefficient of variation was also quantified at time(2) to determine the consistency of each dependent measure between three consecutive contractions. Torque, M-waves, and H-reflexes were most variable during NMES over the nerve. In summary, NMES over the nerve produced contractions with the greatest recruitment through central pathways; however, considering some of the limitations of NMES over the femoral nerve, it may be considered a good complement to, as opposed to a replacement for, NMES over the quadriceps muscle for maintaining muscle quality and reducing contraction fatigue during NMES rehabilitation.     

Muscle weakness following sustained and rhythmic isometric contractions in man

The effects of sustained and rhythmically performed isometric contractions on electrically evoked twitch and tetanic force generation of the triceps surae have been investigated in 4 healthy male subjects. The isometric contractions were performed separately and on different occasions at 30%, 60% and 100% of the force of maximal voluntary contraction (MVC). The area under the maximal voluntary contraction (MVC) force/time curve during the rhythmic and sustained contractions was the same for each experiment. The results showed that following rhythmic isometric exercise there was a small decrease in low (10 and 20 Hz) and high (40 Hz) frequency tetanic tension which was associated with % MVC. However, there was no change in the 20/40 ratio of tetanic forces, MVC or the contraction times and force of the maximal twitch. In contrast, following sustained isometric exercise tetanic forces were markedly reduced, particularly at low frequencies of stimulation. The 20/40 ratio decreased and the induced muscle weakness was greater at 30% than 60% or 100% MVC. The performance of sustained isometric contractions also effected a decrease in contraction time of the twitch and MVC. The results are in accord with previous findings for dynamic work (Davies and White 1982), and show that if isometric exercise is performed rhythmically the effect on tetanic tensions is small and there is no evidence of a preferential loss of electrically evoked force at either high or low frequencies of stimulation following the contractions. For sustained contractions, however, the opposite is true, the ratio of 20/40 Hz forces is markedly reduced and following 30% sustained MVC there is a significant (p less than 0.05) change in the time to peak tension (TPT) of the maximal twitch.     

Eccentric exercise increases EMG amplitude and force fluctuations during submaximal contractions of elbow flexor muscles.

The purpose of this study was to determine the effect of eccentric exercise on the ability to exert steady submaximal forces with muscles that cross the elbow joint. Eight subjects performed two tasks requiring isometric contraction of the right elbow flexors: a maximum voluntary contraction (MVC) and a constant-force task at four submaximal target forces (5, 20, 35, 50% MVC) while electromyography (EMG) was recorded from elbow flexor and extensor muscles. These tasks were performed before, after, and 24 h after a period of eccentric (fatigue and muscle damage) or concentric exercise (fatigue only). MVC force declined after eccentric exercise (45% decline) and remained depressed 24 h later (24%), whereas the reduced force after concentric exercise (22%) fully recovered the following day. EMG amplitude during the submaximal contractions increased in all elbow flexor muscles after eccentric exercise, with the greatest change in the biceps brachii at low forces (3-4 times larger at 5 and 20% MVC) and in the brachialis muscle at moderate forces (2 times larger at 35 and 50% MVC). Eccentric exercise resulted in a twofold increase in coactivation of the triceps brachii muscle during all submaximal contractions. Force fluctuations were larger after eccentric exercise, particularly at low forces (3-4 times larger at 5% MVC, 2 times larger at 50% MVC), with a twofold increase in physiological tremor at 8-12 Hz. These data indicate that eccentric exercise results in impaired motor control and altered neural drive to elbow flexor muscles, particularly at low forces, suggesting altered motor unit activation after eccentric exercise.     

Use of motor cortex stimulation to measure simultaneously the changes in dynamic muscle properties and voluntary activation in human muscles.

Force responses to transcranial magnetic stimulation of motor cortex (TMS) during exercise provide information about voluntary activation and contractile properties of the muscle. Here, TMS-generated twitches and muscle relaxation during the TMS-evoked silent period were measured in fresh, heated, and fatigued muscle. Subjects performed isometric contractions of elbow flexors in two studies. Torque and EMG were recorded from elbow flexor and extensor muscles. One study (n = 6) measured muscle contraction times and relaxation rates during brief maximal and submaximal contractions in fresh and fatigued muscle. Another study (n = 7) aimed to 1) assess the reproducibility of muscle contractile properties during brief voluntary contractions in fresh muscle, 2) validate the technique for contractile properties in passively heated muscle, and 3) apply the technique to study contractile properties during sustained maximal voluntary contractions. In both studies, muscle contractile properties during voluntary contractions were compared with the resting twitch evoked by motor nerve stimulation. Measurement of muscle contractile properties during voluntary contractions is reproducible in fresh muscle and reveals faster and slower muscle relaxation rates in heated and fatigued muscle, respectively. The technique is more sensitive to altered muscle state than the traditional motor nerve resting twitch. Use of TMS during sustained maximal contractions reveals slowing of muscle contraction and relaxation with different time courses and a decline in voluntary activation. Voluntary output from the motor cortex becomes insufficient to maintain complete activation of muscle, although slowing of muscle contraction and relaxation indicates that lower motor unit firing rates are required for fusion of force.     

Repetitive Hops Induce Postactivation Potentiation in Triceps Surae as well as an Increase in the Jump Height of Subsequent Maximal Drop Jumps

Postactivation potentiation (PAP) has been defined as the increase in twitch torque after a conditioning contraction. The present study aimed to investigate the effectiveness of hops as conditioning contractions to induce PAP and increase performance in subsequent maximal drop jumps. In addition, we wanted to test if and how PAP can contribute to increases in drop jump rebound height. Twelve participants performed 10 maximal two-legged hops as conditioning contractions. Twitch peak torques of triceps surae muscles were recorded before and after the conditioning hops. Then, subjects performed drop jumps with and without 10 conditioning hops before each drop jump. Recordings included ground reaction forces, ankle and knee angles and electromyographic activity in five leg muscles. In addition, efferent motoneuronal output during ground contact was estimated with V-wave stimulation. The analyses showed that after the conditioning hops, twitch peak torques of triceps surae muscles were 32% higher compared to baseline values (P < 0.01). Drop jumps performed after conditioning hops were significantly higher (12%, P < 0.05), but V-waves and EMG activity remained unchanged. The amount of PAP and the change in drop jump rebound height were positively correlated (r² = 0.26, P < 0.05). These results provide evidence for PAP in triceps surae muscles induced by a bout of hops and indicate that PAP can contribute to the observed performance enhancements in subsequent drop jumps. The lack of change in EMG activity and V-wave amplitude suggests that the underlying mechanisms are more likely intramuscular than neural in origin.     

Reflex effect of skeletal muscle mechanoreceptor stimulation on the cardiovascular system

To determine the potential for mechanical stimulation of skeletal muscle to contribute to the reflex cardiovascular response to static contraction (exercise reflex), we examined the cardiovascular effects caused by either passive stretch or external pressure applied to the triceps surae muscles. First, the triceps surae were stretched to an average developed tension of 4.8 +/- 0.3 kg. This resulted in increases in mean arterial pressure (MAP) of 28 +/- 7 mmHg, dP/dt of 1,060 +/- 676 mmHg/s, and heart rate (HR) of 6 +/- 2 beats/min (P less than 0.05). Additionally, increments of 0.3, 0.5, 1.0, 2.0, 4.0, and 8.0 kg of tension produced by passive stretch elicited pressor responses of -6 +/- 1, 7 +/- 1, 16 +/- 3, 21 +/- 8, 28 +/- 6, and 54 +/- 9 mmHg, respectively. External pressure, applied with a cuff to the triceps surae to produce intramuscular pressures (125-300 mmHg) that were similar to those seen during static contraction, also elicited small increases in MAP (4 +/- 1 to 10 +/- 1 mmHg) but did not alter HR. Transection of dorsal roots L5-L7 and S1 abolished the responses to passive stretch and external pressure. Moreover, when the triceps surae were stretched passively to produce a pattern and amount of tension similar to that seen during static hindlimb contraction, a significant reflex cardiovascular response occurred. During this maneuver, the pressor response averaged 51% of that seen during contraction.(ABSTRACT TRUNCATED AT 250 WORDS)     

Muscle activation and the isokinetic torque-velocity relationship of the human triceps surae

The influence of muscle activation and the time allowed for torque generation on the angle-specific torque-velocity relationship of the triceps surae was studied during plantar flexion using supramaximal electrical stimulation and a release technique on six male subjects [mean (SD) age 25 (4) years]. Torque-velocity data were obtained under different levels of constant muscle activation by varying the stimulus frequency and the time allowed for isometric torque generation prior to release and isokinetic shortening. To eliminate the effects of the frequency response on absolute torque the isokinetic data were normalized to the maximum isometric torque values at 0.44 rad. There were no significant differences in the normalized torques generated at any angular velocity using stimulus frequencies of 20, 50 or 80 Hz. When the muscle was stimulated at 50 Hz the torques obtained after a 400 ms and 1 s pre-release isometric contraction did not differ significantly. However, with no pre-release contraction significantly less torque was generated at all angular velocities beyond 1.05 rad · s^–1 when compared with either the 200, 400 ms or 1 s condition. With a 200 ms pre-release contraction significantly less torque was generated at angular velocities beyond 1.05 rad · s^–1 when compared with the 400 ms or 1 s conditions. It would seem that the major factor governing the shape of the torque-velocity curve at a constant level of muscle activation is the time allowed for torque generation.     

Motor unit firing rates of the gastrocnemii during maximal brief steady-state contractions in humans

The human triceps surae (soleus, medial (MG) and lateral (LG) gastrocnemii) is complex and important for posture and gait. The soleus exhibits markedly lower motor unit firing rates (MUFRs; ∼16 Hz) during maximal voluntary isometric contraction (MVC) than other limb muscles, but this information is unknown for the MG and LG. During multiple visits, subjects performed a series of 5–7, ∼7-s plantar flexor MVCs with tungsten microelectrodes inserted into the MG and LG. During a separate testing session, another group of subjects performed submaximal isometric contractions at 25%, 50%, and 75% MVC with inserted fine-wires in the MG, LG and soleus. Maximum steady-state MUFRs for MG and LG (∼23 Hz) were not different, but faster than prior reports for the soleus. No differences between the three triceps surae components were detected for 25% or 50% MVC, but at 75% MVC, the MG MUFRs were 31% greater than soleus. The triceps surae exhibit similar torque modulation strategies at <75% MVC, but to achieve higher contraction intensities (>75% MVC) the gastrocnemii rely on faster rates to generate maximal torque than the soleus. Therefore, the MG and LG exhibit a larger range of MUFR capacities.     

Active recovery has a positive and acute effect on recovery from fatigue induced by repeated maximal voluntary contractions of the plantar flexors

This study investigated the acute effect of active recovery (AR) following fatigue induced by 80 three-second maximal voluntary isometric plantar flexion contractions (MVICs) in 12 young men. AR consisted of a total of 180 voluntary isometric ramp contractions of the plantar flexors (0.75-s contraction/relaxation) targeting 10% of MVIC torque. MVIC torque, voluntary activation and root mean square values of electromyographic signals for the triceps surae normalized by each peak-to-peak amplitude of compound motor action potential were determined before, and immediately, 10, 20 and 30 min after the fatiguing task. Evoked torques were similarly assessed except for immediately after it. The AR and passive recovery were randomly performed on two days by each participant between 5 min and 10 min after the fatiguing task. For all the parameters other than MVIC torque, there was no significant difference between the conditions at any time point. MVIC torque decreased significantly immediately after the fatiguing task regardless of condition (P < 0.05), and the corresponding decrease in MVIC torque recovered 30 min after the fatiguing task only in AR (P < 0.05). These results suggest an acute positive effect of AR on recovery of neuromuscular function and/or contractile properties after fatigue.     

Rapid biphasic arteriolar dilations induced by skeletal muscle contraction are dependent on stimulation characteristics

To test the hypothesis that measurable changes in microvasculature dilation occur in response to a single short-duration tetanic contraction, we contracted three to five skeletal muscle fibres of the hamster cremaster muscle microvascular preparation (in situ) and evaluated the response of an arteriole overlapping the active muscle fibres. Arteriolar diameter (baseline diameter = 16.4 +/- 0.9 micro m, maximum diameter = 34.7 +/- 1.2 micro m) was measured before and after a single contraction resulting from a range of stimulus frequencies (4, 10, 20, 30, 40, 60, and 80 Hz) within a 250- or 500-ms train. Four and 10 Hz produced a significant dilation at 2.9 +/- 0.4 and 6.5 +/- 2.8 s, respectively, within a 250-ms train and 3.0 +/- 0.2 and 6.1 +/- 1.3 s, respectively, within a 500-ms train. Biphasic dilations were observed within a 250-ms train at 20 Hz (at 3.9 +/- 0.9 and 22.1 +/- 4.3 s), 30 Hz (at 2.7 +/- 0.3 and 17.5 +/- 2.9 s), and 40 Hz (at 3.8 +/- 0.4 and 23.2 +/- 2.6 s) and within a 500-ms train at 20 Hz (at 4.8 +/- 0.4 and 31.9 +/- 3.8 s) and 30 Hz (at 3.4 +/- 0.3 and 27.6 +/- 3.0 s). A single dilation was observed within a 250-ms train at 60 Hz (at 5.1 +/- 0.7 s) and 80 Hz (at 14.2 +/- 3.3 s) and within a 500-ms train at 40 Hz (at 9.9 +/- 3.2 s), 60 Hz (at 7.9 +/- 2.1 s), and 80 Hz (at 13.4 +/- 4.0 s). We have shown that a single contraction ranging from a single twitch (4 Hz, 250 ms) to fused tetanic contractions produces significant arteriolar dilations and that the pattern of dilation is dependent on the stimulus frequency and train duration.     

Electromyography and fatigue during prolonged, low-level static contractions

Findings from five separate studies of EMG changes and muscle fatigue during prolonged low-level static contractions are summarized, and the possible mechanisms behind the changes are briefly discussed. Sustained static contractions (10%, 7% and 5% MVC) of up to 1 h duration were performed by finger flexors, elbow flexors and extensors, and knee extensors. In one experiment, intermittent static arm pulling (triceps) (10 s contraction and 5 s rest, average work load 14% and 10% MVC) was performed for 7 h. The endurance time for the sustained contractions was around one hour for 10% MVC, and it was shown--all in all--that the concept of "indefinite" endurance times at contractions below 15-20% MVC cannot be maintained. After 5% MVC sustained contractions for one hour a 12% reduction in MVC was seen, and significant increases in EMG amplitude and decreases in the mean spectral frequency of the EMG-power spectrum were found. Marked differences were also seen in the EMG changes in the elbow flexors and extensors, and transcutaneous electrical stimulation of the knee extensors showed that low frequency fatigue was present after the contraction. With intermittent contractions similar changes in the EMG parameters were seen after 2-3 h of contractions at 14% MVC. On average, during contractions of 10% MVC no EMG changes were detected. Increased extracellular potassium concentration in the contracting muscles is suggested as a possible explanation of these findings.     

MLR stimulation and exercise pressor reflex activate different renal sympathetic fibers in decerebrate cats.

Although mesencephalic locomotor region (MLR) stimulation and the exercise pressor reflex have been shown to increase whole nerve renal sympathetic activity, it is not known whether these mechanisms converge onto the same population of renal sympathetic postganglionic efferents. In decerebrate cats, we examined the responses of single renal sympathetic postganglionic efferents to stimulation of the MLR and the exercise pressor reflex (i.e., static contraction of the triceps surae muscles). We found that, in most instances (24 of 28 fibers), either MLR stimulation or the muscle reflex, but not both, increased the discharge of renal postganglionic sympathetic efferents. In addition, we found that renal sympathetic efferents that responded to static contraction while the muscles were freely perfused responded more vigorously to static contraction during circulatory arrest. Moreover, stretch of the calcaneal (Achilles) tendon stimulated the same renal sympathetic efferents as did static contraction. These findings suggest that MLR stimulation and the exercise pressor reflex do not converge onto the same renal sympathetic postganglionic efferents.     

Velocity-dependent muscle strategy during plantarflexion in humans

This work examines the relative contribution of the triceps surae heads and the tibialis anterior (TA) to tension development with reference to voluntary plantarflexion at various velocities and at two articular positions of the knee joint (extended and flexed at 90 °). Subjects were instructed to perform plantarflexion at various submaximal and maximal velocities with no intention of stopping the movement. Voluntary electromyographic (EMG) activity was recorded and the amplitude, duration and integral were analysed. Integrated EMG (IEMG) was normalized with respect to duration. The maximal M wave and the Hoffmann (H) reflex elicited by electrical stimulation of the tibial nerve were recorded in the triceps surae to estimate the effects in gastrocnemii (G) length and motoneuron excitability differences, respectively, in the two knee positions. The results indicate that: (a) although the largest EMG activity was recorded in the extended limb, the greatest maximal velocities were performed in the flexed knee position; (b) with increasing velocity of movement, all triceps surae muscles showed enhanced IEMG activities; (c) at a low velocity of movement the soleus (So1)/G IEMG ratio was larger in the flexed compared to the extended knee; and (d) with increasing velocity, co-activation of agonist and antagonist muscles appeared. It is concluded that the larger maximal velocity of movement observed in the flexed compared to the extended knee was not primarily related to the neural command of the different triceps surae components, but rather to their mechanical properties. Furthermore, co-activation of agonist and antagonist muscles may contribute to the performance of the contractile strategy during rapid movements.