Changes in fascicle length from rest to maximal voluntary contraction affect the assessment of voluntary activation

The purpose of this study was to investigate the effect of the differences between the actual fascicle length during a voluntary contraction and the fascicle length at rest of the triceps surae muscle on the determination of the voluntary activation (VA) by using the interpolated twitch technique. Twelve participants performed isometric voluntary maximal (MVC) and submaximal (20%, 40%, 60% and 80% MVC) contractions at two different ankle angles (75 degrees and 90 degrees ) under application of the interpolated twitch technique. Two ultrasound probes were used to determine the fascicle length of soleus, gastrocnemius medialis and gastrocnemius lateralis muscles. Further, the MVCs and the twitches were repeated for six more ankle angles (85 degrees , 95 degrees , 100 degrees , 105 degrees , 110 degrees and 115 degrees ). The VA of the triceps surae muscle were calculated (a) using the rest twitch force (RTF) measured during the same trial as the interpolated twitch force (ITF; traditional method) and (b) using the RTF at an ankle angle where the fascicle length showed similar values between ITF and RTF (fascicle length consideration method). The continuous changes in fascicle length from rest to MVC affect the accuracy of the assessment of the VA. The traditional method overestimates the assessment of the VA on average 4% to 12%, especially at 90 degrees ankle angle (i.e. short muscle length). The reason for this influence is the unequal force-length potential of the muscle at twitch application by the measure of ITF and RTF. These findings provide evidence that the fascicle length consideration method permits a more precise prediction (an improvement of 4-12%) of the voluntary contraction compared to the traditional method.     

Twitch interpolation technique in testing of maximal muscle strength: Influence of potentiation, force level, stimulus intensity and preload

The aim was to study the methodological aspects of the muscle twitch interpolation technique in estimating the maximal force of contraction in the quadriceps muscle utilizing commercial muscle testing equipment. Six healthy subjects participated in seven sets of experiments testing the effects on twitch size of potentiation, time lag after potentiation, magnitude of voluntary force, stimulus amplitude, stimulus duration, angle of the knee, and angle of the hip. In addition, the consequences of submaximal potentiation on the estimation of maximal force from twitch sizes were studied in five healthy subjects. We found an increase in twitch size with increasing levels of potentiation and twitch size decreased exponentially following potentiation. We found a curvilinear relationship between twitch size and voluntary force, and these properties were more obvious when the stimulation intensity of the preload was reduced. The relationship between twitch size and force was only linear, for force levels greater than 25% of maximum. It was concluded that to achieve an accurate estimate of true maximal force of muscle contraction, it would be necessary for the subject to be able to perform at least 75% of the true maximal force.     

Effects of series compliance on twitches superimposed on voluntary contractions.

The activation of skeletal muscle during voluntary isometric contraction has been assessed by measuring the increase in force caused by a superimposed maximal shock to the motor nerve (the twitch-interpolation technique). When the muscle is held isometric, the increase in force with stimulation (superimposed twitch force) decreases with increasing voluntary force, and a line fit through the data can be extrapolated to maximal voluntary force at the zero twitch force axis. In a previous paper we questioned the applicability of this technique in situations where a high series compliance allows the muscle to shorten during the superimposed twitch. To explore effects of series compliance, we measured force of the adductor pollicis during voluntary isometric contractions with noncompliant and compliant loading devices. With the compliant loading device, superimposed twitch force was systematically less than with the noncompliant device, and the plot of superimposed twitch force vs. voluntary force was often concave upward, preventing easy extrapolation to maximal voluntary force. These findings are consistent with force-velocity characteristics of muscle and suggest that twitch-interpolation data must be interpreted with caution when the muscle is not held isometric during the superimposed twitch.     

Juxtaposition of the changes in intracellular calcium and force during staircase potentiation at 30 and 37°C

Ca²⁺ entry during the action potential stimulates muscle contraction. During repetitive low frequency stimulation, skeletal muscle undergoes staircase potentiation (SP), a progressive increase in the peak twitch force induced by each successive stimulus. Multiple mechanisms, including myosin regulatory light chain phosphorylation, likely contribute to SP, a temperature-dependent process. Here, we used the Ca²⁺-sensitive fluorescence indicators acetoxymethyl (AM)-furaptra and AM-fura-2 to examine the intracellular Ca²⁺ transient (ICT) and the baseline Ca²⁺ level at the onset of each ICT during SP at 30 and 37°C in mouse lumbrical muscle. The stimulation protocol, 8 Hz for 8 s, resulted in a 27 ± 3% increase in twitch force at 37°C and a 7 ± 2% decrease in twitch force at 30°C (P < 0.05). Regardless of temperature, the peak rate of force production (+df/dt) was higher in all twitches relative to the first twitch (P < 0.05). Consistent with the differential effects of stimulation on twitch force at the two temperatures, raw ICT amplitude decreased during repetitive stimulation at 30°C (P < 0.05) but not at 37°C. Cytosolic Ca²⁺ accumulated during SP such that baseline Ca²⁺ at the onset of ICTs occurring late in the train was higher (P < 0.05) than that of those occurring early in the train. ICT duration increased progressively at both temperatures. This effect was not entirely proportional to the changes in twitch duration, as twitch duration characteristically decreased before increasing late in the protocol. This is the first study identifying a changing ICT as an important, and temperature-sensitive, modulator of muscle force during repetitive stimulation. Moreover, we extend previous observations by demonstrating that contraction-induced increases in baseline Ca²⁺ coincide with greater +df/dt but not necessarily with higher twitch force.     

Stiffness, force, and sarcomere shortening during a twitch in frog semitendinosus muscle bundles

The time course of force and stiffness during a twitch was determined at 6 and 26 degrees C in frog semitendinosus muscle bundles using the transmission time technique of Schoenberg, M., J.B. Wells, and R.J. Podolsky, 1974, J. Gen. Physiol. 64:623-642. Sarcomere shortening due to series compliance was also measured using a laser light diffraction technique. Following stimulation, stiffness developed more rapidly than force, but had a slower time course than published Ca2+ transients determined from light signals using Ca2+ sensitive dyes (Baylor, S.M., W.K. Chandler, and M.W. Marshall, 1982, J. Physiol. (Lond.). 331:139-177). Stiffness (S) did not reach its tetanic value during a twitch at 6 or 26 degrees C, although at 6 degrees C, it approached close to this value with S-twitch/S-tetanus = 0.82 +/- 0.07 (+/- SEM). During relaxation, force fell more rapidly than stiffness both for a twitch and also a tetanus. Also in this paper, several of the assumptions inherent in using the transmission time technique for the measurement of stiffness are considered in detail.     

Reproducible measurement of voluntary activation of human elbow flexors with motor cortical stimulation.

Voluntary activation of muscle is commonly quantified by comparison of the extra force added by motor nerve stimulation during a contraction [superimposed twitch (SIT)] with that produced at rest by the same stimulus (resting twitch). An inability to achieve 100% voluntary activation implies that failure to produce maximal force output from the muscle must have occurred at a site at or above the level of the motoneurons. We have used cortical stimulation to quantify voluntary activation. Here, incomplete activation implies a failure at or above the level of motor cortical output. With cortical stimulation, it is inappropriate to compare extra force evoked during a contraction with the twitch evoked in resting muscle because motor cortical and spinal cord excitability both increase with activity. However, an appropriate "resting twitch" can be estimated. We previously estimated its amplitude by extrapolation of the linear relation between SIT amplitude and voluntary torque calculated from 35 contractions of >50% maximum (Todd G, Taylor JL, and Gandevia SC. J Physiol 551: 661-671, 2003). In this study, we improved the utility of this method to enable evaluation of voluntary activation when it may be changing over time, such as during the development of fatigue, or in patients who may be unable to perform large numbers of contractions. We have reduced the number of contractions required to only three. Estimation of the resting twitch from three contractions was reliable over time with low variability. Furthermore, its reliability and variability were similar to the resting twitch estimated from 30 contractions and to that evoked by conventional motor nerve stimulation.     

Effect of twitch interval duration on the contractile function of subsequent twitches in isolated rat, rabbit, and dog myocardium under physiological conditions

Many studies have shown that a change in stimulation frequency leads to altered contractility of the myocardium. However, it remains unclear what changes occur directly after a change in frequency and which ones are a result of the slow processes that lead to the altered homeostasis, which develops after a change in stimulation frequency. To distinguish the immediate from the slow responses, we assessed contractile function in two species that have distinctively different calcium (Ca(2+))-handling properties using a recently developed, randomized pacing protocol. In isolated dog and rat right ventricular trabeculae, twitch contractions at five different cycle lengths within the physiologic range of each species were randomized around a steady-state frequency. We found, in both species, that the duration of the cycle length just prior to the analyzed twitch (primary) positively correlated with the increased force of the analyzed twitch. In sharp contrast, the cycle lengths, one and two more removed from the analyzed twitch ("secondary" and "tertiary"), displayed a negative correlation with force of the analyzed twitch. In additional experiments, assessment of intracellular Ca(2+) transients in rabbit trabeculae revealed that diastolic Ca(2+) levels were closely correlated to contractile function outcome. The relative contribution of the primary cycle length was different between dog (51%) and rat (71%), whereas in neither species was a significant effect on relaxation time observed. With the use of randomized cycle lengths, we have distinguished the intrinsic response from the signaling-mediated effects of frequency-dependent activation on myofilament properties and Ca(2+) handling.     

Relationship between back muscle endurance and voluntary activation

There is some evidence that the Biering-Sorensen endurance test can discriminate low back pain sufferers from healthy individuals and can predict future back pain. This test relies on the subject's ability to voluntarily drive the back muscles. This neural drive, termed voluntary activation (VA) can be measured using the twitch interpolation technique. The aim of the current study was to investigate the relationship between back muscle endurance and VA. Twenty-one healthy volunteers (10 males) participated. Bilateral electromyographic recordings were obtained from erector spinae and rectus abdominis. Back extensor torque was recorded using a dynamometer. The protocol consisted of measurement of VA (using magnetic stimulation of the brain and assessment of the sizes of the evoked twitches) and measurement of endurance. There was a linear correlation (r(2)=1, P<0.01) between voluntary torque and VA. The mean (SEM) endurance time was 174.9 (12.8)s. There was no correlation between endurance and VA at either 100% MVC (r(2)=0.01, P=0.72) or at 50% MVC (r(2)=0.11, P=0.16). These findings indicate that the endurance of the back muscles, as assessed using this widely utilised test does not appear to be related to a subject's ability to drive their back muscles voluntarily either maximally or submaximally.     

Variability in the interpolated twitch torque for maximal and submaximal voluntary contractions.

The superimposed twitch technique is frequently used to study the degree of motor unit activation during voluntary effort. This technique is one of the preferred methods to determine the activation deficit (AD) in normal, athletic, and patient populations. One of the limitations of the superimposed twitch technique is its variability under given contractile conditions. The objective of this research was to determine the source(s) of variability in the superimposed twitch force (STF) for repeat measurements. We hypothesized that the variability in the AD measurements may be caused by the timing of the twitch force relative to the onset of muscle activation, by force transients during the twitch application, by small variations in the actual force from the nominal target force, and by variations in the resting twitch force. Twenty-eight healthy subjects participated in this study. Sixteen of these subjects participated in a protocol involving contractions at 50% of their maximal voluntary contraction (MVC) effort, whereas the remaining 12 participated in a protocol involving contractions at 100% of their MVC. Doublet-twitch stimuli were superimposed onto the 50 and 100% effort knee extensor muscle contractions, and the resting twitch forces, voluntary knee extensor forces, and STFs were then measured. The mean resting twitch forces obtained before and after 8 s of 50% of MVC were the same. Similarly, the mean STFs determined at 1, 3, 5, and 7 s into the 50% MVC were the same. The variations in twitch force were significantly smaller after accounting for the actual force at twitch application than those calculated from the prescribed forces during the 50% MVC protocol (P < 0.05). Furthermore, the AD and the actual force showed statistically significant negative correlations for the 50% MVC tests. The interpolated twitch torque determined for the maximal effort contractions ranged from 1 to 70%. In contrast to the protocol at 50% of MVC, negative correlations were only observed in 5 of the 12 subjects during the 100% effort contractions. These results suggest that small variations in the actual force from the target force can account for the majority of the variations in the STFs for submaximal but not maximal effort contractions. For the maximal effort contractions, large variations in the STF exist due to undetermined causes.     

Assessment of Neuromuscular Function Using Percutaneous Electrical Nerve Stimulation

Percutaneous electrical nerve stimulation is a non-invasive method commonly used to evaluate neuromuscular function from brain to muscle (supra-spinal, spinal and peripheral levels). The present protocol describes how this method can be used to stimulate the posterior tibial nerve that activates plantar flexor muscles. Percutaneous electrical nerve stimulation consists of inducing an electrical stimulus to a motor nerve to evoke a muscular response. Direct (M-wave) and/or indirect (H-reflex) electrophysiological responses can be recorded at rest using surface electromyography. Mechanical (twitch torque) responses can be quantified with a force/torque ergometer. M-wave and twitch torque reflect neuromuscular transmission and excitation-contraction coupling, whereas H-reflex provides an index of spinal excitability. EMG activity and mechanical (superimposed twitch) responses can also be recorded during maximal voluntary contractions to evaluate voluntary activation level. Percutaneous nerve stimulation provides an assessment of neuromuscular function in humans, and is highly beneficial especially for studies evaluating neuromuscular plasticity following acute (fatigue) or chronic (training/detraining) exercise.     

Differences in the log-transformed electromyographic–force relationships of the plantar flexors between high- and moderate-activated subjects

The present study examined the log-transformed electromyographic amplitude (EMG) versus force relationships for the medial gastrocnemius (MG) and soleus (SOL) in high- and moderate-activated subjects. Twenty-five (age = 21 ± 2 year; mass = 62 ± 12 kg) participants performed six submaximal contractions (30–90% maximal voluntary contraction [MVC]) with the interpolated twitch technique (ITT) performed at 90% MVC to calculate percent voluntary activation (% VA). Sixteen participants with > 90% VA at 90% MVC were categorized high-activated group; the remaining nine were the moderate-activated group. Linear regression models were fit to the log-transformed EMG–force relationships. The slope (b value) and the antilog of the Y-intercept (a value) were calculated. The b values from the MG EMG–force relationships were higher (P < 0.05) for the high-activated group (1.27 ± 0.13) than the moderate-activated group (0.88 ± 0.06). The a values and p–p M-wave amplitude values (collapsed across twitches [superimposed and potentiated]) were larger (P < 0.05) for the MG (1.17 ± 0.40 and 8.98 ± 0.46 mV) than the SOL (0.24 ± 0.07 and 4.48 ± 0.20 mV) when collapsed across groups. The b value from the log-transformed EMG–force relationships is an attractive model to determine if a subject has the ability to achieve high activation of their MG without muscle or nerve stimulation.     

Effect of nifedipine on the contractile function of the rat diaphragm in vitro

The isometric contractile response of the directly-stimulated rat diaphragm was studied before and following addition of the calcium channel blocker, nifedipine. Nifedipine (10 micrograms/ml and 30 micrograms/ml bath concentrations) significantly increased isometric force output during twitch and unfused tetanic stimulation. Force potentiation during unfused tetanic stimulation was equivalent during either high or low voltage stimulation. Nifedipine had no effect on the time to peak force, half relaxation time, or relaxation time during twitch stimulation; thus, both activation and relaxation rates were increased. The force potentiating actions of nifedipine persisted in a calcium-free bathing solution and were enhanced by d-tubocurarine. In contrast to the force enhancing effects found with twitch and unfused tetanic stimulation, nifedipine caused a small but significant reduction in isometric force during maximal fused tetanic stimulation. It is concluded that the force potentiating effects of nifedipine on rat diaphragm are not due to fiber recruitment, enhancement of neuromuscular excitation, or altered inward trans-sarcolemmal calcium flux, but may result from a direct effect of the drug on the rate of activation of the contractile apparatus.     

Peripheral and central fatigue after muscle-damaging exercise is muscle length dependent and inversely related

Healthy untrained men performed 10 series of 12 knee eccentric extension repetitions (EE) at 160°/s. The maximal voluntary isometric contraction force of the quadriceps muscle, the maximal rate of electrically induced torque development (RTD) and relaxation (RTR), isokinetic concentric torque at 30°/s, the electrostimulation-induced torque at 20 and 100 Hz frequencies were established before and after EE at shorter and longer muscle lengths. Besides, voluntary activation (VA) index and central activation ratio (CAR) were tested. There was more peripheral fatigue than central after EE. We established more central fatigue as well as low frequency fatigue at a shorter muscle length compared to the longer muscle length. Relative RTD as well as relative RTR, improved after EE and did not depend on the muscle length. Finally, central fatigue is inversely significantly related with the eccentric torque reduction during eccentric exercise and with the changes in muscle torque induced by low frequency stimulation.     

Methodological issues with the interpolated twitch technique.

A number of methodological issues in the use of the interpolated twitch technique were investigated for their effect on true maximum force (TMF) and activation (ACT): timing of control (pre- vs post-contraction) and superimposed twitches (first vs second); type of twitch stimulus (primarily magnitude); and the type of extrapolation utilised. On three occasions subjects performed a series of maximal and sub-maximal contractions of the knee extensors, with electrically evoked twitches delivered before, during and after each contraction. The twitch-voluntary force relationship was concave for all types of twitch stimuli, and extrapolation using this relationship typically calculated TMF 39N (7%) higher, and ACT 7% lower than linear extrapolation. The timing of the control (2-4%) and superimposed twitches (approximately 4%) both influenced TMF and ACT. Despite the different twitch stimuli being a range of magnitudes (13-32% maximum voluntary force) they did not affect TMF and ACT. A novel finding was that prior potentiation changed the shape of the twitch-voluntary force relationship. For precise measurement of TMF and ACT it is recommended that: extrapolation is based on the twitch-voluntary force relationship of the experimental model; and post-contraction potentiated twitches be used, as the superimposed twitch on a high level contraction appears to be potentiated.     

Is maximum stimulation intensity required in the assessment of muscle activation capacity?

Voluntary activation assessment using the interpolation twitch technique (ITT) has almost invariably been done using maximal stimulation intensity, i.e., an intensity beyond which no additional joint moment or external force is produced by increasing further the intensity of stimulation. The aim of the study was to identify the minimum stimulation intensity at which percutaneous ITT yields valid results. Maximal stimulation intensity and the force produced at that intensity were identified for the quadriceps muscle using percutaneous electrodes in eight active men. The stimulation intensities producing 10–90% (in 10% increments) of that force were determined and subsequently applied during isometric contractions at 90% of maximum voluntary contraction (MVC) via twitch doublets. Muscle activation was calculated with the ITT and pain scores were obtained for each stimulation intensity and compared to the respective values at maximum stimulation intensity. Muscle activation at maximal stimulation intensity was 91.6 (2.5)%. The lowest stimulation intensity yielding comparable muscle activation results to maximal stimulation was 50% (88.8 (3.9)%, p < 0.05). Pain score at maximal stimulation intensity was 6.6 (1.5) cm and it was significantly reduced at 60% stimulation intensity (3.7 (1.5) cm, p < 0.05) compared to maximal stimulation intensity. Submaximal stimulation can produce valid ITT results while reducing the discomfort obtained by the subjects, widening the assessment of ITT to situations where discomfort may otherwise impede maximal electrostimulation.     

Different effects of verapamil and low calcium on repetitive contractile activity of frog fatigue-resistant and easily-fatigued muscle fibres.

The effects of low calcium and verapamil on contractility of two muscle fibre types (m. iliofibularis, Rana temporaria) upon different stimulation protocols were been compared. Verapamil (0.02 mmol/l) induced temporal excitation-contraction coupling failure during single tetanic stimulation and enhanced the decline of tetanic force during 30 s repetitive tetanic stimulation in both fatigue-resistant fibres and easily-fatigued fibres. In contrast to verapamil, low extracellular calcium (0.02 mmol/l) only enhanced the decline of tetanic force in fatigue-resistant during repetitive tetanic stimulation but had no effect on easily-fatigued fibres. The effect of verapamil on the decline of tetanic force in fatigue-resistant fibres was more profound in low calcium conditions. Both verapamil and low calcium eliminated twitch facilitation that appeared after prolonged contractile activity in fatigue-resistant fibres. 4mmol/l Ni+2, used as calcium channel antagonist, had effects similar to low calcium medium. It could be concluded that (i) extracellular Ca2+-requirements for excitation-contraction coupling are different in fatigue-resistant and easily-fatigued fibres; (ii) the effects of verapamil on force performance are not entirely dependent upon calcium channel blockade.     

Extracellular determinants of cardiac contractility in the cold anoxic turtle

Painted turtles (Chrysemys picta) survive months of anoxic submergence, which is associated with large changes in the extracellular milieu where pH falls by 1, while extracellular K+, Ca++, and adrenaline levels all increase massively. While the effect of each of these changes in the extracellular environment on the heart has been previously characterized in isolation, little is known about their interactions and combined effects. Here we examine the isolated and combined effects of hyperkalemia, acidosis, hypercalcemia, high adrenergic stimulation, and anoxia on twitch force during isometric contractions in isolated ventricular strip preparations from turtles. Experiments were performed on turtles that had been previously acclimated to warm (25 degrees C), cold (5 degrees C), or cold anoxia (submerged in anoxic water at 5 degrees C). The differences between acclimation groups suggest that cold acclimation, but not anoxic acclimation per se, results in a downregulation of processes in the excitation-contraction coupling. Hyperkalemia (10 mmol L(-1) K+) exerted a strong negative inotropic effect and caused irregular contractions; the effect was most pronounced at low temperature (57%-97% reductions in twitch force). Anoxia reduced twitch force at both temperatures (14%-38%), while acidosis reduced force only at 5 degrees C (15%-50%). Adrenergic stimulation (10 micromol L(-1)) increased twitch force by 5%-19%, but increasing extracellular [Ca++] from 2 to 6 mmol L(-1) had only small effects. When all treatments were combined with anoxia, twitch force was higher at 5 degrees C than at 25 degrees C, whereas in normoxia twitch force was higher at 25 degrees C. We propose that hyperkalemia may account for a large part of the depressed cardiac contractility during long-term anoxic submergence.     

Acidosis counteracts the negative inotropic effect of K+ on ventricular muscle strips from the toad Bufo marinus

Strenuous activity is associated with acidosis, increased extracellular potassium concentration ([K+]o), and elevated levels of circulating catecholamines. Acidosis and elevated [K+]o are normally considered harmful to cardiac function, and a high sympathetic tone on the heart may lead to arrhythmia. During activity, however, the heart must be able to increase rate and strength of contraction. While the individual effects of [K+]o, acidosis, and adrenaline on contractile properties of cardiac muscle have been characterized for some ectothermic species, less information is available on their interactions. Here we examine the isolated and combined effects of [K+]o, acidosis, and adrenaline on ventricular muscle strips from the toad Bufo marinus. This study showed that increased [K+]o significantly reduced twitch force, while lactic acid significantly increased twitch force and more than counteracted the negative inotropic effects of elevated [K+]o. There was no inotropic effect of Na-lactate (neutralized lactic acid), which suggests that lactic acid stimulated twitch force through reduced pH and not by serving as a substrate. Adrenaline had a positive effect on twitch force in all preparations. Irrespective of treatment, twitch force decreased as stimulation rate increased. During high [K+]o, there was a severe reduction in maximal frequency of toad ventricular strips that was not alleviated by lactic acidosis and/or adrenaline, which suggests that high [K+]o influences twitch force and maximal rate by different mechanisms. In vivo levels of lactic acid, [K+]o, adrenaline, and heart rate previously observed during forced activity in Bufo did not significantly affect the contractile properties of heart muscle strips in vitro. Thus, although [K+]o significantly decreased twitch force, this detrimental effect was more than counteracted by the positive inotropic effect of lactic acid and adrenaline.     

In vivo magnetic and electric recordings from nerve bundles and single motor units in mammalian skeletal muscle. Correlations with muscle force

Recent advances in the technology of recording magnetic fields associated with electric current flow in biological tissues have provided a means of examining action currents that is more direct and possibly more accurate than conventional electrical recording. Magnetic recordings are relatively insensitive to muscle movement, and, because the recording probes are not directly connected to the tissue, distortions of the data due to changes in the electrochemical interface between the probes and the tissue are eliminated. In vivo magnetic recordings of action currents of rat common peroneal nerve and extensor digitorum longus (EDL) muscle were obtained by a new magnetic probe and amplifier system that operates within the physiological temperature range. The magnetically recorded waveforms were compared with those obtained simultaneously by conventional, extracellular recording techniques. We used the amplitude of EDL twitch force (an index of stimulus strength) generated in response to graded stimulation of the common peroneal nerve to enable us to compare the amplitudes of magnetically recorded nerve and muscle compound action currents (NCACs and MCACs, respectively) with the amplitudes of electrically recorded nerve compound action potentials (NCAPs). High, positive correlations to stimulus strength were found for NCACs (r = 0.998), MCACs (r = 0.974), and NCAPs (r = 0.998). We also computed the correlations of EDL single motor unit twitch force with magnetically recorded single motor unit compound action currents (SMUCACs) and electrically recorded single motor unit compound action potentials (SMUCAPs) obtained with both a ring electrode and a straight wire serving as a point electrode. Only the SMUCACs had a relatively strong positive correlation (r = 0.768) with EDL twitch force. Correlations for ring and wire electrode-recorded SMUCAPs were 0.565 and -0.366, respectively. This study adds a relatively direct examination of action currents to the characterization of the normal biophysical properties of peripheral nerve, muscle, and muscle single motor units.     

K+-induced twitch potentiation is not due to longer action potential

The objective of this study was to determine whether an increased duration of the action potential contributes to the K+-induced twitch potentiation at 37 degrees C. Twitch contractions were elicited by field stimulation, and action potentials were measured with conventional microelectrodes. For mouse extensor digitorum longus (EDL) muscle, twitch force was greater at 7-13 mM K+ than at 4.7 mM (control). For soleus muscle, twitch force potentiation was observed between 7 and 11 mM K+. Time to peak and half-relaxation time were not affected by the increase in extracellular K+ concentration in EDL muscle, whereas both parameters became significantly longer in soleus muscle. Decrease in overshoot and prolongation of the action potential duration observed at 9 and 11 mM K+ were mimicked when muscles were respectively exposed to 25 and 50 nM tetrodotoxin (TTX; used to partially block Na+ channels). Despite similar action potentials, twitch force was not potentiated by TTX. It is therefore suggested that the K+-induced potentiation of the twitch in EDL muscle is not due to a prolongation of the action potential and contraction time, whereas a longer contraction, especially the relaxation phase, may contribute to the potentiation in soleus muscle.