Moment dependency of the series elastic stiffness in the human plantar flexors measured in vivo.

The moment dependency of the series elastic stiffness (SES) in the human plantar flexors was investigated in vivo with the quick release method. At an ankle moment of 100 N m produced with either voluntary or electrical stimulation we found non-significantly different SES of 506+/-72 and 529+/-125 N m rad(-1), respectively. It has recently been proposed that the amount of series elastic tissue involved in plantar flexion changes with the moment level produced by the plantar flexors (Hof, J. Biomech 31 (1998) 793). However, our results indicate that the amount of series elastic tissue involved in plantar flexions remained constant with changing moment levels. We therefore propose that the series elastic component (SEC) in human plantar flexors act as one structure or rather one combination of anatomical structures which is engaged at all muscle activation levels, and that the mechanical properties (i.e. the stress-strain function) are determined by the combined tissue mechanical properties. Additionally, our results demonstrated that the SES in the human plantar flexors at moments levels up to about isometric maximum did not reach an asymptote where the stiffness is independent of moment, i.e. SEC of the plantar flexors is, during many daily activities, loaded for the greatest part in the non-linear part of the stress-strain function.     

Stimulation-induced damage in rabbit fast-twitch skeletal muscles: a quantitative morphological study of the influence of pattern and frequency

The aim of this study was to determine whether muscle fibre degeneration brought about by chronic lowfrequency electrical stimulation was related to the pattern and frequency of stimulation. Rabbit fast-twitch muscles, tibialis anterior and extensor digitorum longus, were stimulated for 9 days with pulse trains ranging in frequency from 1.25 Hz to 10 Hz. Histological data from these muscles were analysed with multivariate statistical techniques. At the lower stimulation frequencies there was a significantly lower incidence of degenerating muscle fibres. Fibres that reacted positively with an antineonatal antibody were most numerous in the sections that revealed the most degeneration. The dependence on frequency was generally similar for the two muscles, but the extensor digitorum longus muscles showed more degeneration than the tibialis anterior at every frequency. Muscles subjected to 10 Hz intermittent stimulation showed significantly less degeneration than muscles stimulated with 5 Hz continuously, although the aggregate number of impulses delivered was the same. The incidence of degeneration in the extensor digitorum longus muscles stimulated at 1.25 Hz was indistinguishable from that in control, unstimulated muscles; for the tibialis anterior muscles, this was also true for stimulation at 2.5 Hz. We conclude that damage is not an inevitable consequence of electrical stimulation. The influence of pattern and frequency on damage should be taken into account when devising neuromuscular stimulation régimes for clinical use.     

Spinal reflexes and coactivation of ankle muscles during a submaximal fatiguing contraction.

This study examined the involvement of spinal mechanisms in the control of coactivation during a sustained contraction of the ankle dorsiflexors at 50% of maximal voluntary contraction. Changes in the surface electromyogram (EMG) of the tibialis anterior and of two antagonist muscles, the soleus and lateral gastrocnemius, were investigated during and after the fatigue task. Concurrently, the compound action potential (M-wave) and the Hoffmann reflex of the soleus and lateral gastrocnemius were recorded. The results showed that the torque of the ankle dorsiflexors and the average EMG of the tibialis anterior during maximal voluntary contraction declined by 40.9 +/- 17.7% (mean +/- SD; P < 0.01) and 37.0 +/- 19.9% (P < 0.01), respectively, at task failure. During the submaximal fatiguing contraction, the average EMG of both the agonist and antagonist muscles increased, leading to a nearly constant ratio at the end of the contraction when normalized to postfatigue values. In contrast to the monotonic increase in average EMG of the antagonist muscles, the excitability of their spinal reflex pathways exhibited a biphasic modulation. The amplitude of the Hoffman reflexes in the soleus and lateral gastrocnemius increased to 147.5 +/- 52.9% (P < 0.05) and 166.7 +/- 74.9% (P < 0.01), respectively, during the first 20% of the contraction and then subsequently declined to 66.3 +/- 44.8 and 74.4 +/- 44.2% of their initial values. In conclusion, the results show that antagonist coactivation did not contribute to task failure. The different changes in voluntary EMG activity and spinal reflex excitability in the antagonist muscles during the fatiguing contraction support the concept that the level of coactivation is controlled by supraspinal rather than spinal mechanisms. The findings indicate, however, that antagonist coactivation cannot simply be mediated by a central descending "common drive" to the motor neuron pools of the agonist-antagonist muscle pairs. Rather, they suggest a more subtle regulation of the drive, possibly through presynaptic mechanisms, to the motoneurons that innervate the antagonist muscles.     

Effect of ankle joint position and electrode placement on the estimation of the antagonistic moment during maximal plantarflexion.

During maximal efforts, antagonistic activity can significantly influence the joint moment. During maximal voluntary "isometric" contractions, certain joint rotation can not be avoided. This can influence the estimation of the antagonistic moment from the EMG activity. Our study aimed to quantify the influence on the calculated agonistic moment produced during maximal voluntary isometric plantarflexions (a) when estimating antagonistic moments at different ankle angles and (b) when placing the EMG electrodes at different portions over the m. tibialis anterior. Ten subjects performed maximal voluntary isometric plantarflexions at 90 degrees ankle angle. In order to estimate the antagonistic moment, submaximal isometric dorsiflexions were performed at various ankle angles. Moment and EMG signals from mm. triceps surae and tibialis anterior were measured. The RMS differences between plantarflexors moment calculated considering the antagonistic cocontraction estimated at the same ankle angle at which the maximal plantarflexion moment was achieved and at different ankle angles ranged from 0.10 to 2.94 Nm. The location of the electrodes led to greater RMS differences (2.35-5.18 Nm). In conclusion, an angle 10 degrees greater than the initial plantarflexion angle is enough to minimize the effect of the change in length of the m. tibialis anterior during the plantarflexion on the estimation of the plantarflexors moment. The localisation of the electrodes over the m. tibialis anterior can influence the estimation of its cocontraction during maximal plantarflexion efforts.     

Quantitative morphology of stimulation-induced damage in rabbit fast-twitch skeletal muscles

Summary The purpose of this study was to examine the contention that stimulation-induced damage, resulting in degeneration with subsequent regeneration, plays a major role in the transformation of fibre type brought about by chronic electrical stimulation. Data from histological and histochemical sections of 9-day-stimulated rabbit fast-twitch muscles were analysed with multivariate statistical techniques. Fibre degeneration and regeneration varied non-systematically between sample areas at any given cross-sectional level. In the extensor digitorum longus muscle, but not in the tibialis anterior, there was more degeneration in proximal than in distal portions of the muscle. The extensor digitorum longus muscle consistently showed more degeneration than the tibialis anterior muscle. Degeneration was less extensive for an intermittent pattern of stimulation that delivered half the aggregate number of impulses of continuous stimulation. Degeneration and regeneration varied markedly between individual rabbits in each of the groups. Sections that revealed the most degeneration and regeneration also had more fibres that reacted positively with an anti-neonatal antibody. Rigorous analysis of different sources of variation has helped to explain apparent conflicts in the literature. The incidence of muscle fibre damage in the stimulated tibialis anterior muscle is low, showing that the contribution of degenerative-regenerative phenomena to fibre type conversion in this muscle is insignificant.     

Estimation of instantaneous moment arms of lower-leg muscles

Muscle moment arms at the human knee and ankle were estimated from muscle length changes measured as a function of joint flexion angle in cadaver specimens. Nearly all lower-leg muscles were studied: extensor digitorum longus, extensor hallucis longus, flexor digitorum longus, flexor hallucis longus, gastrocnemius lateralis, gastrocnemius medialis, peroneus brevis, peroneus longus, peroneus tertius, plantaris, soleus, tibialis anterior, and tibialis posterior. Noise in measured muscle length was filtered by means of quintic splines. Moment arms of the mm. gastrocnemii appear to be much more dependent on joint flexion angles than was generally assumed by other investigators. Some consequences for earlier analyses are mentioned.     

Aging does not affect voluntary activation of the ankle dorsiflexors during isometric, concentric, and eccentric contractions.

This study examines the age-related deficit in force of the ankle dorsiflexors during isometric (Iso), concentric (Con), and eccentric (Ecc) contractions. More specifically, the contribution of neural and muscular mechanisms to the loss of voluntary force was investigated in men and women. The torque produced by the dorsiflexors and the surface electromyogram (EMG) from the tibialis anterior and the soleus were recorded during maximal Iso contractions and during Con and Ecc contractions performed at constant angular velocities (5-100 degrees/s). Central activation was tested by the superimposed electrical stimulation method during maximal voluntary contraction and by computing the ratio between voluntary average EMG and compound muscle action potential (M wave) induced by electrical stimulation (average EMG/M wave). Contractile properties of the dorsiflexor muscles were investigated by recording the mechanical responses to single and paired maximal stimuli. The results showed that the age-related deficit in force (collapsed across genders and velocities) was greater for Iso (20.5%; P < 0.05) and Con (38.6%; P < 0.001) contractions compared with Ecc contractions (6.5%; P > 0.05). When the torque produced during Con and Ecc contractions was expressed relative to the maximal Iso torque, it was significantly reduced in Con contractions and increased in Ecc contractions with aging, with the latter effect being more pronounced for women. In both genders, voluntary activation was not significantly impaired in elderly adults and did not differ from young subjects. Similarly, coactivation was not changed with aging. In contrast, the mechanical responses to single and paired stimuli showed a general slowing of the muscle contractile kinetics with a slightly greater effect in women. It is concluded that the force deficit during Con and Iso contractions of the ankle dorsiflexors in advanced age cannot be explained by impaired voluntary activation or changes in coactivation. Instead, this age-related adaptation and the mechanisms that preserve force in Ecc contractions appeared to be located at the muscular level.     

Discovery of TBC1D1 as an insulin-, AICAR-, and contraction-stimulated signaling nexus in mouse skeletal muscle

The Akt substrate of 160 kDa (AS160) is phosphorylated on Akt substrate (PAS) motifs in response to insulin and contraction in skeletal muscle, regulating glucose uptake. Here we discovered a dissociation between AS160 protein expression and apparent AS160 PAS phosphorylation among soleus, tibialis anterior, and extensor digitorum longus muscles. Immunodepletion of AS160 in tibialis anterior muscle lysates resulted in minimal depletion of the PAS band at 160 kDa, suggesting the presence of an additional PAS immunoreactive protein. By immunoprecipitation and mass spectrometry, we identified this protein as the AS160 paralog TBC1D1, an obesity candidate gene regulating GLUT4 translocation in adipocytes. TBC1D1 expression was severalfold higher in skeletal muscles compared with all other tissues and was the dominant protein detected by the anti-PAS antibody at 160 kDa in tibialis anterior and extensor digitorum longus but not soleus muscles. In vivo stimulation by insulin, contraction, and the AMP-activated protein kinase (AMPK) activator AICAR increased TBC1D1 PAS phosphorylation. Using mass spectrometry on TBC1D1 from mouse skeletal muscle, we identified several novel phosphorylation sites on TBC1D1 and found the majority were consensus or near consensus sites for AMPK. Semiquantitative analysis of spectra suggested that AICAR caused greater overall phosphorylation of TBC1D1 sites compared with insulin. Purified Akt and AMPK phosphorylated TBC1D1 in vitro, and AMPK, but not Akt, reduced TBC1D1 electrophoretic mobility. TBC1D1 is a major PAS immunoreactive protein in skeletal muscle that is phosphorylated in vivo by insulin, AICAR, and contraction. Both Akt and AMPK phosphorylate TBC1D1, but AMPK may be the more robust regulator.     

Functional electrical stimulation using microstimulators to correct foot drop: a case study

This paper presents a case study that tested the feasibility and efficacy of using injectable microstimulators (BIONs) in a functional electrical stimulation (FES) device to correct foot drop. Compared with surface stimulation of the common peroneal nerve, stimulation with BIONs provides more selective activation of specific muscles. For example, stimulation of the tibialis anterior (TA) and extensor digitorum longus (EDL) muscles with BIONs produces ankle flexion without excessive inversion or eversion of the foot (i.e., balanced flexion). Efficacy was assessed using a 3-dimensional motion analysis of the ankle and foot trajectories during walking with and without stimulation. Without stimulation, the toe on the affected leg drags across the ground. BION stimulation of the TA muscle and deep peroneal nerve (which innervates TA and EDL) elevates the foot such that the toe clears the ground by 3 cm, which is equivalent to the toe clearance in the less affected leg. The physiological cost index (PCI) measured effort during walking. The PCI equals the change in heart rate (from rest to activity) divided by the walking speed; units are beats per metre. The PCI is high without stimulation (2.29 +/- 0.37, mean +/- SD) and greatly reduced with surface (1.29 +/- 0.10) and BIONic stimulation (1.46 +/- 0.24). Also, walking speed increased from 9.4 +/- 0.4 m/min without stimulation to 19.6 +/- 2.0 m/min with surface and 17.8 +/- 0.7 m/min with BIONic stimulation. These results suggest that FES delivered by a BION is an alternative to surface stimulation and provides selective control of muscle activation.     

Comparison between EMG to force processing and kinetic analysis for the calf muscle moment in walking and stepping

In EMG to force processing the force of a muscle or the corresponding muscle moment with respect to the joint is determined from EMG and muscle length by means of a model of the muscle's contractile and elastic properties. In kinetic analysis the joint moment is calculated from the ground reaction force and the positions and accelerations of the body segments by means of Newtonian equations, based on a rigid body segment model. These two fundamentally different methods have been compared by determining simultaneously the calf muscle moment and the total ankle moment in walking at slow, moderate and fast speed and in stepping up and down a low bench. As long as there is no activity of other muscles, the moments obtained by either method should be identical. The analysis was restricted to such periods, as assessed from the EMG of tibialis anterior. In each experimental condition three steps were analyzed in five subjects of 21-22 yr. In comparison with the results of kinetic analysis, EMG processing shows differences between 7 and 54 Nm r.m.s., with an average of 22 Nm r.m.s. Normalized with respect to the r.m.s. value of the moment itself, the differences amount to 0.13-0.51, with an average of 0.22. In the majority of cases there was no systematic deviation between EMG processing and kinetic analysis and the r.m.s. difference was of the same order of magnitude as determined previously in ergometer experiments.     

Effect of physical activity on MRI-measured blood oxygen level-dependent transients in skeletal muscle after brief contractions.

The signal intensity (SI) in gradient-echo, echo-planar magnetic resonance images (repetition time/echo time = 1,000/40) of anterior tibialis muscle in active [estimated energy expenditure 42.4 +/- 3.7 (SD), n = 8] vs. sedentary (32.3 +/- 0.6 kcal.kg(-1).day(-1), n = 8) young adult (18-34 yr old) human subjects was measured after single, 1-s-duration maximum voluntary ankle dorsiflexion contractions. There was no difference between groups in anterior tibial muscle cross-sectional area or peak force. In both groups there was a transient increase in anterior tibialis muscle SI, which peaked 5-7 s after the end of each contraction. The magnitude of the SI transient was over threefold greater [5.5 +/- 1.0 (SE) vs. 1.5 +/- 0.4%] and persisted twice as long (half-recovery time 5.4 +/- 0.4 vs. 2.7 +/- 0.3 s) in the active subjects. In the same subjects, blood flow in popliteal, anterior tibial, and posterior tibial arteries was measured by cardiac-gated CINE magnetic resonance angiography before and after 2 min of dynamic, repetitive ankle dorsiflexion exercise. There was no difference between groups in resting or postexercise flow in anterior tibial artery, although popliteal and posterior tibial artery flow after exercise tended to be greater in the active group. The results indicate that transient hyperemia and oxygenation in muscle after single contractions are enhanced by chronic physical activity to a greater extent than peak muscle blood flow.     

Inference of motor unit recruitment order in voluntary and electrically elicited contractions

The relationship between surface myoelectric signal parameters and the level of voluntary or electrically elicited contractions was studied in 32 experiments on the tibialis anterior muscle of 22 healthy human subjects. Contractions were performed at 20 and 80% of the maximum voluntary contraction torque. Two levels of stimulation current were used, yielding, respectively, a maximum M wave and an M wave approximately 30% of the maximum. A four-bar electrode probe was used to detect single- and double-differential signals from which mean and median frequency of the power spectrum and average muscle fiber conduction velocity were estimated. Measurements obtained from voluntary contractions showed a positive correlation between contraction levels and both conduction velocity and spectral parameters. Conduction velocity increased by 21.2 +/- 10.9% when voluntary contraction level increased from 20 to 80% of the maximal value. Spectral parameters increased by similar amounts. Tetanic electrical stimulation was applied to a muscle motor point for 20 s via surface electrodes. Rectangular current pulses with 0.1-ms width and frequencies of 20, 25, 30, 35, and 40 Hz were used. Four types of behavior were observed with increasing stimulation level: 1) the two spectral parameters and conduction velocity both increased with stimulation in 15 experiments, 2) the two spectral parameters decreased and conduction velocity increased in 8 experiments, 3) the two spectral parameters and conduction velocity both decreased in 6 experiments, and 4) the two spectral parameters increased and conduction velocity decreased in 3 experiments. Conduction velocity increased with increasing stimulation current in 72% of the experiments, indicating a recruitment order similar to that of voluntary contractions, although it decreased in the other 28% of the cases, indicating a reverse order of recruitment. Contrary to what is observed in direct stimulation of nerves, motor units are not in general recruited in reverse order of size during electrical stimulation of a muscle motor point. This discrepancy may be the result of geometric factors or a lack of correlation between axonal branch diameter and the diameter of the parent motoneuron axon. Changes of conduction velocity and spectral parameters in opposite directions may be the result of the combined effect of the motor unit recruitment order and of the different tissue filtering function associated with the geometric location of the recruited motor units within the muscle.     

In vivo specific tension of human skeletal muscle.

In this study, we estimated the specific tensions of soleus (Sol) and tibialis anterior (TA) muscles in six men. Joint moments were measured during maximum voluntary contraction (MVC) and during electrical stimulation. Moment arm lengths and muscle volumes were measured using magnetic resonance imaging, and pennation angles and fascicular lengths were measured using ultrasonography. Tendon and muscle forces were modeled. Two approaches were followed to estimate specific tension. First, muscle moments during electrical stimulation and moment arm lengths, fascicular lengths, and pennation angles during MVC were used (data set A). Then, MVC moments, moment arm lengths at rest, and cadaveric fascicular lengths and pennation angles were used (data set B). The use of data set B yielded the unrealistic specific tension estimates of 104 kN/m(2) in Sol and 658 kN/m(2) in TA. The use of data set A, however, yielded values of 150 and 155 kN/m(2) in Sol and TA, respectively, which agree with in vitro results from fiber type I-predominant muscles. In fact, both Sol and TA are such muscles. Our study demonstrates the feasibility of accurate in vivo estimates of human muscle intrinsic strength.     

Isometric muscle length-tension curves do not predict angle-torque curves of human wrist in continuous active movements

In this study we tested the hypothesis that during steady contractions of human wrist extensors or flexors, the torque-angle relationship during movements imposed about the wrist is predicted by the classical isometric muscle length-tension curve, with ascending, descending and ascending limbs. Angle-torque relationships were measured during steady muscle activation (10% of maximal voluntary contraction: MVC), elicited either by electrical stimulation or voluntary regulation of the electromyogram (EMG). Flexion-extension movements of constant speed (+/-10 degrees /s) were imposed on the subjects' hands with a servo actuator, either through the full physiological range of motion +/-50 degrees, or through +/-10 degrees. During extensor contractions, angle-torque curves in +/-50 degrees movements had ascending, descending and ascending limbs, as in isometric contractions. However, in +/-10 degrees movements, torque always increased with increasing muscle length and decreased with decreasing length, even over angles corresponding to the descending limb of isometric curves. For flexor activation, angle-torque curves had similar properties, though descending limbs were less obvious or absent. During imposed movements, hysteresis was observed in the angle-torque curves. This was attributed to non-linearities of the active muscles. Hysteresis reached a maximum at intermediate wrist angles and declined at maximal muscle length, contradicting the recent hypothesis that sarcomere non-uniformity is responsible for the hysteresis. We conclude that the classical isometric length-tension curve, with its prominent descending limb, does not predict angle-torque curves of human wrist muscles in continuous movements. A more appropriate model is one in which stiffness about the wrist is always positive and hysteresis is a significant factor.     

Real-Time Changes in Corticospinal Excitability during Voluntary Contraction with Concurrent Electrical Stimulation

While previous studies have assessed changes in corticospinal excitability following voluntary contraction coupled with electrical stimulation (ES), we sought to examine, for the first time in the field, real-time changes in corticospinal excitability. We monitored motor evoked potentials (MEPs) elicited by transcranial magnetic stimulation and recorded the MEPs using a mechanomyogram, which is less susceptible to electrical artifacts. We assessed the MEPs at each level of muscle contraction of wrist flexion (0%, 5%, or 20% of maximum voluntary contraction) during voluntary wrist flexion (flexor carpi radialis (FCR) voluntary contraction), either with or without simultaneous low-frequency (10 Hz) ES of the median nerve that innervates the FCR. The stimulus intensity corresponded to 1.2× perception threshold. In the FCR, voluntary contraction with median nerve stimulation significantly increased corticospinal excitability compared with FCR voluntary contraction without median nerve stimulation (p<0.01). In addition, corticospinal excitability was significantly modulated by the level of FCR voluntary contraction. In contrast, in the extensor carpi radialis (ECR), FCR voluntary contraction with median nerve stimulation significantly decreased corticospinal excitability compared with FCR voluntary contraction without median nerve stimulation (p<0.05). Thus, median nerve stimulation during FCR voluntary contraction induces reciprocal changes in cortical excitability in agonist and antagonist muscles. Finally we also showed that even mental imagery of FCR voluntary contraction with median nerve stimulation induced the same reciprocal changes in cortical excitability in agonist and antagonist muscles. Our results support the use of voluntary contraction coupled with ES in neurorehabilitation therapy for patients.     

Induction of neuronal type nitric oxide synthase in skeletal muscle by chronic electrical stimulation in vivo

Reiser, Peter J., William O. Kline, and Pal L. Vaghy.Induction of neuronal type nitric oxide synthase in skeletal muscle by chronic electrical stimulation in vivo. J. Appl. Physiol. 82(4): 1250-1255, 1997.Fast-twitch skeletal muscles contain more neuronal-type nitricoxide synthase (nNOS) than slow-twitch muscles because nNOS is presentonly in fast (type II) muscle fibers. Chronic in vivo electricalstimulation of tibialis anterior and extensor digitorum longus musclesof rabbits was used as a method of inducing fast-to-slow fiber typetransformation. We have studied whether an increase in musclecontractile activity induced by electrical stimulation alters nNOSexpression, and if so, whether the nNOS expression decreases to thelevels present in slow muscles. Changes in the expression of myosinheavy chain isoforms and maximum velocity of shortening of skinnedfibers indicated characteristic fast-to-slow fiber type transformationafter 3 wk of stimulation. At the same time, activity of NOS doubled inthe stimulated muscles, and this correlated with an increase in theexpression of nNOS shown by immunoblot analysis. These data suggestthat nNOS expression in skeletal muscle is regulated by muscle activityand that this regulation does not necessarily follow the fast-twitchand slow-twitch pattern during the dynamic phase of phenotypetransformation.

     

Muscle damage and repair in voluntarily running mice: strain and muscle differences

Summary Soleus, extensor digitorum longus and tibialis anterior muscles of mice voluntarily running in wheels for periods of 5 to 120 days were studied in spaced serial and serial cross-sections. Shortly after the onset of running and during the next 2 weeks, degeneration, necrosis, phagocytosis and regeneration of muscle fibers, satellite cell proliferation and cellular infiltration were found in soleus muscles of mice from all strains investigated (CBA/J, NMRI, C57b, NIH, SWS and Balb/c). Tibialis anterior but not extensor digitorum longus muscles were also damaged. Predominantly high-oxidative fibers were affected (both slow-oxidative and fast oxidative glycolytic in soleus, fast-oxidative glycolytic in tibialis anterior). Denervated soleus muscles that had been passively stretched during running were not damaged. Evidence was found that, during the early period of running, split fibers form by myogenesis within (regeneration) or outside (satellite cell proliferation) necrotic muscle fiber segments. Split fibers persisted in solei of long-term (2 to 3 months) exercised CBA/J but not NMRI mice. In 6 out of 20 solei of CBA/J runners exercised for 2 months or longer, fiber-type grouping was observed in the areas where extensive damage usually occurred in the early periods. The results show that different muscles are damaged and repaired to varying degrees and that marked interstrain and inter-individual differences are present. It appears that acute muscle injury occurring upon onset of voluntary running is a usual event in the adaptation of muscles to altered use.     

Age-related decline in rate of torque development is accompanied by lower maximal motor unit discharge frequency during fast contractions

The aim of this study was to investigate the association between the rate of torque development and maximal motor unit discharge frequency in young and elderly adults as they performed rapid submaximal contractions with the ankle dorsiflexors. Recordings were obtained of the torque exerted by the dorsiflexors during the isometric contractions and the surface and intramuscular electromyograms (EMGs) from the tibialis anterior. The maximal rate of torque development and integrated EMG (percentage of total EMG burst) at peak rate of torque development during fast contractions were lower in elderly than young adults by 48% (P < 0.05) and 16.5% (P < 0.05), respectively. The young adults, but not the elderly adults, exhibited a positive association (r2 = 0.33; P < 0.01) between the integrated EMG computed up to the peak rate of torque development and the maximal rate of torque development achieved during the fast contractions. These age-related changes during fast voluntary contractions were accompanied by a decline (P < 0.001) in motor unit discharge frequency (19, 28, and 34% for first 3 interspike intervals, respectively) and in the percentage of units (45%; P < 0.05) that exhibited double discharges (doublets) at brief intervals (<5 ms). Because aging decreased the maximal rate of torque development of fast voluntary contractions to a greater extent ( approximately 10%) than that of an electrically evoked twitch, collectively the results indicate that the age-related decline in maximal motor unit discharge frequency likely limit, in addition to the slowing of muscle contractile properties, the performance of fast voluntary contractions.     

Effect of muscle fatigue on the compliance of the gastrocnemius medialis tendon and aponeurosis

The aim of the present study was to examine whether or not the compliance of the gastrocnemius medialis (GM) tendon and aponeurosis is influenced by submaximal fatiguing efforts. Fourteen elderly male subjects performed isometric maximal voluntary plantarflexion contractions (MVC) on a dynamometer before and after two fatiguing protocols. The protocols consisted of: (1) submaximal concentric isokinetic contractions (70% isokinetic MVC) at 60 degrees /s and (2) a sustained isometric contraction (40% isometric MVC) until failure to hold the defined moment. Ultrasonography was used to determine the elongation and strain of the GM tendon and aponeurosis. To account for the axis misalignment between ankle and dynamometer, the kinematics of the leg were captured at 120 Hz. The maximum moment decreased from 85.9+/-17.9 Nm prior fatigue to 79.2+/-19 Nm after isokinetic fatigue and to 69.9+/-16.4 Nm after isometric fatigue. The maximal strain of the GM tendon and aponeurosis before fatigue, after isokinetic and after isometric fatigue were 4.9+/-1.1%, 4.4+/-1.1% and 4.3+/-1.1% respectively. Neither the strain nor the elongation showed significant differences before and after each fatiguing task at any 100 N step of the calculated tendon force. This implies that the compliance was not altered after either the isokinetic or the isometric fatiguing task. Therefore it was concluded that the strains during the performed submaximal fatiguing tasks, were too small to provoke any structural changes in tendon and aponeurosis.     

Hyperthermia and central fatigue during prolonged exercise in humans.

The present study investigated the effects of hyperthermia on the contributions of central and peripheral factors to the development of neuromuscular fatigue. Fourteen men exercised at 60% maximal oxygen consumption on a cycle ergometer in hot (40 degrees C; hyperthermia) and thermoneutral (18 degrees C; control) environments. In hyperthermia, the core temperature increased throughout the exercise period and reached a peak value of 40.0 +/- 0.1 degrees C (mean +/- SE) at exhaustion after 50 +/- 3 min of exercise. In control, core temperature stabilized at approximately 38.0 +/- 0.1 degrees C, and exercise was maintained for 1 h without exhausting the subjects. Immediately after the cycle trials, subjects performed 2 min of sustained maximal voluntary contraction (MVC) either with the exercised legs (knee extension) or with a "nonexercised" muscle group (handgrip). The degree of voluntary activation during sustained maximal knee extensions was assessed by superimposing electrical stimulation (EL) to nervus femoralis. Voluntary knee extensor force was similar during the first 5 s of contraction in hyperthermia and control. Thereafter, force declined in both trials, but the reduction in maximal voluntary force was more pronounced in the hyperthermic trial, and, from 30 to 120 s, the force was significantly lower in hyperthermia compared with control. Calculation of the voluntary activation percentage (MVC/MVC + EL) revealed that the degree of central activation was significantly lower in hyperthermia (54 +/- 7%) compared with control (82 +/- 6%). In contrast, total force of the knee extensors (MVC + force from EL) was not different in the two trials. Force development during handgrip contraction followed the same pattern of response as was observed for the knee extensors. In conclusion, these data demonstrate that the ability to generate force during a prolonged MVC is attenuated with hyperthermia, and the impaired performance is associated with a reduction in the voluntary activation percentage.