Catchlike-inducing train activation of human muscle during isotonic contractions: burst modulation.

Stimulation trains that exploit the catchlike property [catchlike-inducing trains (CITs)] produce greater forces and rates of rise of force than do constant-frequency trains (CFTs) during isometric contractions and isovelocity movements. This study examined the effect of CITs during isotonic contractions in healthy subjects. Knee extension was electrically elicited against a load of 10% of maximum voluntary isometric contraction. The stimulation intensity was set to produce 20% of maximum voluntary isometric contraction. The muscle was tested before and after fatigue with a 6-pulse CFT and 6-pulse CITs that contained an initial doublet, triplet, or quadruplet. For prefatigue responses, the greatest isotonic performance was produced by CITs with initial doublets. When the muscles were fatigued, triplet CITs were best. CITs produce greater excursion, work, peak power, and average power than do CFTs, because CITs produced more rapid rates of rise of force. Faster rates of rise of force enabled the preload on the muscle to be exceeded earlier during the stimulation train.     

Variable-frequency train stimulation of canine latissimus dorsi muscle during shortening contractions

George, David T., Stuart A. Binder-Macleod, Thomas N. Delosso, and William P. Santamore. Variable-frequency trainstimulation of canine latissimus dorsi muscle during shorteningcontractions. J. Appl. Physiol. 83(3):994-1001, 1997.In cardiomyoplasty, the latissimus dorsi muscle(LDM) is wrapped around the heart ventricles and electrically activatedwith a constant-frequency train (CFT). This study tested the hypothesesthat increased mechanical performance from the LDM could be achieved byactivating the muscle with variable-frequency trains (VFTs) of shorterduration or containing fewer stimulus pulses than the CFT now used. Themechanical performance of the canine LDM (n = 7) during shortening contractionswas measured while the muscle was stimulated with 5- and 6-pulse CFTs(of duration 132 and 165 ms, respectively) and 5- and 6-pulse VFTs (ofduration 104 and 143 ms, respectively) that were designed to takeadvantage of the catchlike property of skeletal muscle. Measurementswere made from fresh and fatigued muscles. For the fresh muscles, the VFTs elicited significantly greater peak power than did the 6-pulse CFT. When the muscles were fatigued, VFT stimulation significantly improved both the peak and mean power produced compared withstimulation by CFTs. These results show that stimulation of the LDMwith shorter duration VFTs is potentially useful for application incardiomyoplasty.

     

Voluntary activation of human quadriceps during and after isokinetic exercise.

The extent of voluntary activation in fresh and fatigued quadriceps muscles was investigated during isometric and isokinetic voluntary contractions at 20 and 150 degrees/s in 23 normal human subjects. The muscles were fatigued by a total of 4 min of maximal knee extension at an angular velocity of 85 degrees/s. Voluntary activation was determined by the superimposition of tetanic electrical stimulation at 100 Hz for 250 ms, initiated at a constant knee angle. The relationship between voluntary and stimulated force was similar to that found with the established twitch superimposition technique used on isometric contractions. In fresh muscle all the subjects showed full voluntary activation during isometric contractions. Some activation failure was seen in five subjects at 20 degrees/s [2.0 +/- 0.9 degrees (SE)] and in two subjects at 150 degrees/s (0.7 +/- 0.5). After fatigue all subjects showed some activation failure at 0 and 20 degrees/s (36.4 +/- 3.1 and 28.8 +/- 4.1 degrees, respectively), but only two showed any at 150 degrees/s (1.4 +/- 5.7). We conclude that brief high-intensity dynamic exercise can cause a considerable failure of voluntary activation. This failure was most marked during isometric and the lower-velocity isokinetic contractions. Thus a failure of voluntary activation may have greater functional significance than previous studies of isometric contractions have indicated.     

Changes of the force-frequency relationship in human tibialis anterior at fatigue.

This work estimates the influence of the single twitch (ST) parameters changes on specific regions of the force-frequency relationship (FFR) in fatigued human tibialis anterior (TA). In 20 subjects (age 20-40) the TA underwent three stimulation phases: (a) five STs at 1 Hz followed by 5 s stimulation with increasing rate (1-50 Hz, to obtain FFR); (b) fatiguing stimulation (35 Hz for 40 s); (c) same as in "a". By the average STs (mean of the five responses) of a and c phases, the peak twitch (Pt) was calculated. Moreover, after ST normalization to Pt, the maximum contraction rate (MCR) and the maximum relaxation rate (MRR) were computed. By the FFR, normalized to the 50 Hz force, we first defined the threshold frequency (TF) when the force oscillation presented the same value in (a) and (c), and then the areas below the FFR in the 1 Hz-TF and in the TF-50 Hz ranges. RESULTS: In unfatigued and fatigued muscle Pt, and MRR changed from 6.12 +/- 3.08 to 3.27 +/- 1.16 N and from 0.87 +/- 0.13 to 0.65 +/- 0.09% Pt/ms, respectively. MCR did not change significantly. The 1 Hz-TF area ratio (c/a) was > 1 for muscles having fatigued Pt > 60% of its basal value. The TF-50 Hz area ratio (c/a) was mostly below 1. CONCLUSIONS: At fatigue, MRR reduction, leading to a better fusion of muscle mechanical output, is able to compensate, in the 1 Hz-TF frequency range, up to 40% Pt loss; beyond TF, the changes of FFR are related to the degree of force loss indicated by the fatigued Pt.     

Sensitivity of the motor unit unfused tetanus to changes in the pattern of motoneuronal firing

The effects of decreasing as well as increasing the interpulse intervals on the tension produced by motor units in the rat medial gastrocnemius muscle were investigated. The increase and decrease in tension, resulting from these changes in interpulse intervals were observed and compared for each motor unit in tetani fused to a various degree. It was found that amplitudes of both changes in tension were the same when the tetanic fusion index was approximately 0.70 and the tension corresponded to 40-50% of the maximal tetanic tension. This observation concerned the all three types of motor units. We also studied the effects of introducing a short interpulse interval ("doublet") at the beginning of the stimulation. The doublet produced an increased tetanic tension with a more fused profile, however the tension was similarly sensitive to an increase as well as a decrease in the interpulse interval when the tetanic fusion index was also about 0.70. Therefore, the extent of tetanic fusion determines the sensitivity of motor unit tetani to changes in a pattern of stimulation. The tetanus of the fusion index about 0.70 can be considered as the most sensitive to changes in the pattern of motoneuronal firing.     

Sound production in the whitemouth croaker and relationship between fish size and disturbance call characteristics

The whitemouth croaker produces two different sounds using extrinsic sonic muscles: (1) male advertisement calls during the spawning season and (2) disturbance calls, produced by both sexes. The advertisement call, related to courtship, was recorded in the field and from two marked spawning males of 28 and 30.5 cm L_T in the laboratory. It consists of individual pulses with average durations of 19.7 ms and 17.8 ms for the two males respectively, interpulse intervals of 496 ms and 718 ms, and dominant frequencies of 280 Hz and 316 Hz. Pulses are emitted in bouts of one to three min duration. Disturbance calls consist of a burst of pulses produced at short intervals, and the pulse duration, interpulse interval and dominant frequency of the pulses average 19.8 ms, 17.1 ms, and 363 Hz, respectively. Dominant frequency and interpulse interval decrease and pulse duration increases with fish size. Sound characteristics change markedly in young of the year individuals (lower than 25 cm L_T) after which they appear to stabilize. Higher dominant frequency in the advertisement than in the disturbance call and the relationship of dominant frequency to pulse duration suggest that dominant frequency is determined as a forced response to muscle contraction parameters rather than by the natural frequency of the swimbladder.     

Effects of peripheral inputs from hindlimb on the monosynaptic reflex of motoneurons innervating tail muscles.

The effects of group II muscle (PBSt, GS) and cutaneous afferent (Sur, SPc, Tib) inputs from the hindlimb on the monosynaptic reflexes of motoneurons innervating tail muscles were studied in lower spinalized cats. Stimulation of the cutaneous nerves at the conditioning-test stimulus interval of about 10-20 ms facilitated and inhibited the monosynaptic reflexes of ipsilateral and contralateral tail muscles, respectively. The effects of the muscle nerve stimulation were not so prominent as those elicited by cutaneous nerve stimulation. The monosynaptic reflex was also inhibited by muscle nerve stimulation at 10-50 ms intervals. The effects of conditioning stimulation of the hindlimb peripheral nerves at short intervals were depressed or blocked by section of the ipsilateral lateral funiculus at S1 spinal segment. These findings show that the neuronal pathway from hindlimb afferents to tail muscle motoneurons passed the lateral funiculus of the spinal cord and modulates the motoneuronal activity of tail muscles.     

Activation of human quadriceps femoris muscle during dynamic contractions: effects of load on fatigue.

Muscle fatigue is both multifactorial and task dependent. Electrical stimulation may assist individuals with paralysis to perform functional activities [functional electrical stimulation (FES), e.g., standing or walking], but muscle fatigue is a limiting factor. One method of optimizing force is to use stimulation patterns that exploit the catchlike property of skeletal muscle [catchlike-inducing trains (CITs)]. Although nonisometric (dynamic) contractions are important parts of both normal physiological activation of skeletal muscles and FES, no previous studies have attempted to identify the effect that the load being lifted by a muscle has on the fatigue produced. This study examined the effects of load on fatigue during dynamic contractions and the augmentation produced by CITs as a function of load. Knee extension in healthy subjects was electrically elicited against three different loads. The highest load produced the least excursion, work, and average power, but it produced the greatest fatigue. CIT augmentation was greatest at the highest load and increased with fatigue. Because CITs were effective during shortening contractions for a variety of loads, they may be of benefit during FES applications.     

Effect of caffeine on skeletal muscle function before and after fatigue

We studied the effect of caffeine on voluntary and electrically stimulated contractions of the adductor pollicis muscle in five adult volunteers. Caffeine (500 mg) was administered orally in a double-blind fashion. Electrical stimulation of the ulnar nerve was performed at 10, 20, 30, 50, and 100 Hz before and after a sustained voluntary contraction held at 50% of the maximal voluntary contraction (MVC). A brief tetanus at 30 Hz was also performed to calculate relaxation rate in the fresh muscle. Contractile properties, relaxation rate, and endurance were then assessed after caffeine and placebo, as well as the response of the fatigued muscle to different frequencies of stimulation. There was no difference in the maximal tension obtained with electrical stimulation (T100) or in the MVC between placebo and caffeine. The tensions developed with electrical stimulation at lower frequencies increased significantly with caffeine ingestion, shifting the frequency-force curve to the left, both before and after fatigue. Mean plasma caffeine concentration associated with these responses was 12.2 +/- 4.9 mg/l. We conclude that caffeine has a direct effect on skeletal muscle contractile properties both before and after fatigue as demonstrated by electrical stimulation.     

Differential effects of temperature on contractile behavior in isolated frog skeletal muscle.

1. The thermal dependence of contractile behavior at different stimulation frequencies was investigated in isolated frog sartorius muscles. 2. Increasing incubation temperature (10-30 degrees C) produced decreases in Pt (43.7%) and P15 (70.3%), and an increase in Po (26.0%). 3. Thermal ratios (R10) calculated for Pt, P15 and Po indicated high thermal dependence at lower temperatures (10-20 degrees C; 0.60, 0.44 and 1.38, respectively) but relative thermal independence at higher temperatures (20-30 degrees C; 0.95, 0.75 and 0.95, respectively). 4. Contractile ratios (Pt/Po and P15/Po) decreased with increased temperature (10-30 degrees C; 56.3% and 76.0%, respectively). 5. Thermal ratios (R20) calculated for peak tension at different stimulation frequencies demonstrated high thermal dependence at the lower frequencies (10-30 pps, 0.46-0.48) and decreasing dependence at higher frequencies (40-50 pps, 0.69-0.82).     

Catchlike property of rat diaphragm: subsequent train frequency effects in variable-train stimulation.

A high-frequency burst of pulses at the onset of a subtetanic train of stimulation allows skeletal muscle to hold force at a higher level than expected from the extra pulses alone because of the catchlike property of muscle. The present study tested the hypothesis that the presence and degree of force increase induced by a high-frequency burst are strongly modulated by the subsequent train frequency. Rat diaphragm muscle strips (studied in vitro at 37 degrees C) underwent two-, three-, or four-pulse bursts [interpulse interval (IPI) of 5 or 10 ms] at the onset of 10- to 50-Hz subtetanic trains. Force was quantified during the train with respect to its peak value (F(peak)), mean value (F(mean)), and force-time integral (F(area)), and it was compared with that produced during subtetanic trains of an equal number of pulses without preceding pulse bursts (Diff-F(peak), Diff-F(mean), Diff-F(area)). F(peak) and F(mean) increased with two-, three-, and four-pulse bursts, and Diff-F(peak) and Diff-F(mean) increased progressively with decreasing frequency of the subtetanic train. F(area), the best reflection of catchlike force augmentation, was increased mainly by the four-pulse bursts with an IPI of 10 ms, and Diff-F(area) was maximal at subsequent train frequencies of 15-25 Hz. The use of incorrect patterns of burst stimulation could also precipitate F(area) decreases, which were observed with the four-pulse, 5-ms IPI paradigm. The time required to reach 80% of maximal force (T(80%)) became shorter for each of the pulse burst stimulation patterns, with maximal reduction of Diff-T(80%) occurring at a subsequent train frequency of 20 Hz in all cases. These data indicate that extra-pulse burst stimulation paradigms need to incorporate the optimal combinations of extra-pulse number, IPI, and the frequency of the subsequent subtetanic train to take greatest advantage of the catchlike property of muscle.     

Effects of modulators of sarcoplasmic Ca2+ release on the development of skeletal muscle fatigue.

The reduced release of Ca2+ from sarcoplasmic reticulum (SR) is considered a major determinant of muscle fatigue. In the present study, we investigated whether the presence of dantrolene, an established inhibitor of SR Ca2+ release, or caffeine, a drug facilitating SR Ca2+ release, modifies muscle fatigue development. Accordingly, the effects of Ca2+ release modulators were analyzed in vitro in mouse fast-twitch [extensor digitorum longus (EDL)] and slow-twitch (soleus) muscles, fatigued by repeated short tetani (40 Hz for 300 ms, 0.5 s(-1) in soleus and 60 Hz for 300 ms, 0.3 s(-1) in EDL, for 6 min). Caffeine produced a substantial increase of tetanic tension of both EDL and soleus muscles, whereas dantrolene decreased tetanic tension only in EDL muscle. In both EDL and soleus muscles, 5 microM dantrolene did not affect fatigue development, whereas 20 microM dantrolene produced a positive staircase during the first 3 min of stimulation in EDL muscle and a slowing of fatigue development in soleus muscle. The development of the positive staircase was abolished by the addition of 15 microM ML-7, a selective inhibitor of myosin light chain kinase. On the other hand, caffeine caused a larger and faster loss of tension in both EDL and soleus muscles. The results seem to indicate that the changes in fatigue profile induced by caffeine or dantrolene are mainly due to the changes in the initial tetanic tension caused by the drugs, with the resulting changes in the level of contraction-dependent factors of fatigue, rather than to changes in the SR Ca2+ release during fatigue development.     

Post-Activation Potentiation--A Clue for Simplifying Models of Muscle Dynamics

SYNOPSIS. Post-activation potentiation is a phenomena that occursonly in fasttwitch muscle fibers. Its main effect is to enhancemuscle force at sub-maximal activation levels for a short durationof time following previous muscle activation. We characterizedthis phenomenon in feline caudofemoralis (CF) muscle (composedof 100% fast-twitch muscle fibers) to understand its importanceduring physiological patterns of activation. During such patterns(e.g., 43 pps, 8 pulse trains delivered at 1 sec intervals)CF potentiated rapidly and apparently maximally. When CF wasallowed to relax, potentiation decayed slowly with a time constant20–40 x slower than the rise-time. The level of potentiationreached during the potentiating paradigm was stable in responseto a wide range of stimuli, including various stimulation rates(15–120 pps) and various inter-train intervals (up to10 sec). The shape of the twitch force-length curve for potentiatedCF was similar to that of the tetanic force-length curve ineither the potentiated or unpotentiated state. In contrast,the shape of the twitch force-length curve for unpotentiatedCF was shifted markedly to the right accompanied by a narrowingof the curve's peak. We conclude from our observations thatfast-twitch muscle fibers operate and should be modeled in astate of full potentiation, and that modeling the potentiatedstate may actuaUy be simpler than modeling the unpotentiatedstate.     

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.     

A method of bite force measurement in primates

A bite force transducer consisting of two differential strain beams with four strain gages in a full bridge configuration was modified for measuring occlusal forces in rhesus monkeys. A procedure of muscle stimulation (20-50 V, 60 Hz, and 0.8 ms duration) produced maximal unilateral masticatory muscle contraction when stimulating electrodes were placed in the masseter muscle. Tests of this procedure revealed reproducible results and a potential for use in studies of the force of isometric contraction of the masticatory muscles in normal and experimentally altered macaques and other primates.     

Fatigue in mammalian skeletal muscle stimulated under computer control

Functional electrical stimulation (FES) is used to provideparalyzed human subjects with postural support and a limited range ofmovements. Problems encountered with FES include jerky movements fromtension oscillations during stimulation and rapid muscle fatigue. Inthis paper, we report experiments on anesthetized cats that test a new,computer-controlled method of stimulation of the muscle nerve supply,distributed across several inputs, which reduces these problems. After5 min of continuous, distributed stimulation of the medialgastrocnemius muscle at 6 pulses per second (pps) across 6 channels,tension fell to 55.9 ± 3.9% (SE) of its original value. Incomparison, after 5 min of synchronous stimulation of one muscleportion at 36 pps, tension fell to 11 ± 3.7%. At higherstimulation rates, 10 pps per channel (distributed) and 60 pps(synchronous), the differences in fatigue were even greater. Similarresults were obtained when an intermittent, rather than a continuous,stimulation protocol was used. These findings indicate that distributedstimulation has important advantages over other methods forapplications such as FES.

     

Characterization of the synaptic properties of olfactory bulb projections

The olfactory bulb directly projects to several diverse telencephalic structures, but, to date, few studies have investigated the physiological characteristics of most of these areas. As an initial step towards understanding the odor processing functions of these secondary olfactory structures, we recorded evoked field potentials in response to lateral olfactory tract stimulation in vivo in urethane-anesthetized Sprague-Dawley rats in the following brain structures: anterior olfactory nucleus, ventral and dorsal tenia tecta, olfactory tubercle, anterior and posterior piriform cortex, the anterior cortical nucleus of the amygdala, and lateral entorhinal cortex. Using paired-pulse stimulation with interpulse intervals of 25-1000 ms, we observed facilitation of the response to the second pulse in every structure examined, although the degree of facilitation varied among the target structures. Additionally, pulse train stimulation at three different frequencies (40, 10 and 2 Hz) produced facilitation of evoked field potentials that also varied among target structures. We discuss the potential utility of such short-term facilitation in olfactory processing.     

Neurophysiological characteristics of human leg muscle action potentials evoked by transcutaneous magnetic stimulation of the spine

The objectives of this study were to establish the neurophysiological properties of the compound muscle action potentials (CMAPs) evoked by transcutaneous magnetic stimulation of the spine (tsMSS) and the effects of tsMSS on the soleus H‐reflex. In semi‐prone seated subjects with trunk semi‐flexed, the epicenter of a figure‐of‐eight magnetic coil was placed at Thoracic 10 with the handle on the midline of the vertebral column. The magnetic stimulator was triggered by monophasic single pulses of 1 ms, and the intensity ranged from 90% to 100% of the stimulator output across subjects. CMAPs were recorded bilaterally from ankle and knee muscles at the interstimulus intervals of 1, 3, 5, 8, and 10 s. The CMAPs evoked were also conditioned by posterior tibial and common peroneal nerve stimulation at a conditioning‐test (C‐T) interval of 50 ms. The soleus H‐reflex was conditioned by tsMSS at the C‐T intervals of 50, 20, ?20, and ?50 ms. The amplitude of the CMAPs was not decreased when evoked at low stimulation frequencies, excitation of group I afferents from mixed peripheral nerves in the leg affected the CMAPs in a non‐somatotopical neural organization pattern, and tsMSS depressed soleus H‐reflex excitability. These CMAPs are likely due to orthodromic excitation of nerve motor fibers and antidromic depolarization of different types of afferents. The latency of these CMAPs may be utilized to establish the spine‐to‐muscle conduction time in central and peripheral nervous system disorders in humans. tsMSS may constitute a non‐invasive modality to decrease spinal reflex hyperexcitability and treat hypertonia in neurological disorders. Bioelectromagnetics 34:200–210, 2013. © 2012 Wiley Periodicals, Inc.     

Rapid-Rate Paired Associative Stimulation over the Primary Somatosensory Cortex

Rapid-rate paired associative stimulation (rPAS) involves repeat pairing of peripheral nerve stimulation and Transcranial magnetic stimulation (TMS) pulses at a 5 Hz frequency. RPAS over primary motor cortex (M1) operates with spike-timing dependent plasticity such that increases in corticospinal excitability occur when the nerve and TMS pulse temporally coincide in cortex. The present study investigates the effects of rPAS over primary somatosensory cortex (SI) which has not been performed to date. In a series of experiments, rPAS was delivered over SI and M1 at varying timing intervals between the nerve and TMS pulse based on the latency of the N20 somatosensory evoked potential (SEP) component within each participant (intervals for SI-rPAS: N20, N20-2.5 ms, N20 + 2.5 ms, intervals for M1-rPAS: N20, N20+5 ms). Changes in SI physiology were measured via SEPs (N20, P25, N20-P25) and SEP paired-pulse inhibition, and changes in M1 physiology were measured with motor evoked potentials and short-latency afferent inhibition. Measures were obtained before rPAS and at 5, 25 and 45 minutes following stimulation. Results indicate that paired-pulse inhibition and short-latency afferent inhibition were reduced only when the SI-rPAS nerve-TMS timing interval was set to N20-2.5 ms. SI-rPAS over SI also led to remote effects on motor physiology over a wider range of nerve-TMS intervals (N20-2.5 ms – N20+2.5 ms) during which motor evoked potentials were increased. M1-rPAS increased motor evoked potentials and reduced short-latency afferent inhibition as previously reported. These data provide evidence that, similar to M1, rPAS over SI is spike-timing dependent and is capable of exerting changes in SI and M1 physiology.