Nonlinear summation of contractions in cat muscles. I. Early depression

Nerves to fast- and slow-twitch cat muscles were stimulated with various numbers of supramaximal pulses under isometric conditions. By subtracting the force produced by j - 1 pulses from that produced by j pulses, the contribution of the j th pulse could be compared with the response to one pulse (twitch response). A less-than-linear summation (depression) was observed during the rising phase of the twitch. This depression became increasingly prominent and longer in duration with repetitive stimulation. A more-than-linear summation (facilitation) was observed during the falling phase of the twitch, which became increasingly delayed and smaller in amplitude with repetitive stimulation. The early depression could be abolished for the first few pulses by Dantrolene [1-(5-p-nitrophenyl) furfurilidene amino hydantoin sodium hydrate], which reduced Ca++ release from the sarcoplasmic reticulum. The depression was less prominent at short muscle lengths or with stimulation of single motor units. A first-order, saturable reaction such as Ca++ binding to troponin or actin binding to myosin can quantitatively account for the early depression.     

Slow changes and wiener analysis of nonlinear summation in contractions of cat muscles

With regular trains of stimuli at a high frequency, the contribution of each stimulus to the force generated over time declines from the second to about the tenth stimulus, but then begins to increase again. This late increase is referred to as tetanic potentiation in analogy with the post-tetanic potentiation of the twitch after such a period of stimulation. With regular trains of stimuli at a low frequency, a progressive decrease in the essentially unfused twitches (negative staircase) is observed in the slow soleus muscle of the cat, while a progressive increase (positive staircase) is observed for the fast plantaris muscle. The time constant for the approximately exponential changes observed is on the order of 10 s. Random trains of stimuli were applied at intermediate frequencies and analyzed in terms of general methods of analysis for nonlinear systems. Systematic decreases in the magnitude and increases in the time course of the average tension per stimulus were observed with increasing mean rates of stimulation. Similar changes were observed for short intervals between stimuli within a given random train at a constant mean rate. These changes can be described in terms of an early depression and a later facilitation described in the previous papers in this series.     

Effects of elastic loads on the contractions of cat muscles

The nerves to plantaris and soleus muscles in the cat were stimulated with maximal single shocks and with random stimulus trains which produced partially fused contractions. In order to obtain information on the mechanism of muscular contraction, the effects of allowing the muscles to shorten against various elastic loads were studied in the time domain and in the frequency domain. When springs of increasing stiffness were placed in series with the muscle, the twitch tension increased greatly. The gain of the frequency response curve was also much greater with stiffer springs. The shape of the frequency response curve for plantaris muscle could usually be described by that expected for a second-order system with two real time constants or rate constants. The rate constants changed in qualitatively similar ways in response to increased stiffness of an elastic load, increased muscle length and increased mean rate of nerve stimulation. These results are in agreement with the hypothesis that the linear responses of muscles working against elastic loads are determined by the values of two rate constants. Thus, of the many processes associated with contraction, only two are rate-limiting: one associated with the viscoelastic properties of muscle and the second associated with the reuptake of Ca into the sarcoplasmic reticulum. Non-linear aspects of muscular contraction are also discussed. These are more prominent in soleus muscle than in plantaris muscle.Graduate student of the Medical Research Council of Canada.Formerly a Post-doctoral Fellow of the Muscular Dystrophy Association of Canada.     

Nonlinear summation of force in cat tibialis anterior: a muscle with intrafascicularly terminating fibers.

The complex connective tissue structure of muscle and tendon suggests that forces from two parts of a muscle may not summate linearly, particularly in muscles with intrafasciculary terminating fibers, such as cat tibialis anterior (TA). In four anesthetized cats, the TA was attached to a servomechanism to control muscle length and record force. The ventral roots were divided into two bundles, each innervating about half the TA, so the two parts could be stimulated alone or together. Nonlinear summation of force (F(nl)) was measured during isometric contractions. F(nl) was small and negative, indicating less than linear summation of the parts, which is consistent with the predicted F(nl) of muscle fibers connected in series. F(nl) was more significant when smaller parts of the muscle were tested (21.8 vs. 8% for whole muscle). These data were fit to a model where both parts of the muscle were assumed to stretch a common elasticity. Compensatory movements of the servomechanism showed the common elasticity is very stiff, and the model cannot account for F(nl) in cat TA.     

Nonlinear summation of force in cat soleus muscle results primarily from stretch of the common-elastic elements.

The complex connective tissue structure of muscle and tendon suggests that forces from two parts of a muscle may not summate linearly. This study measured the nonlinear summation of force (F(nl)) in whole cat soleus during isometric and ramp movements. In six anesthetized cats, the soleus was attached to a servomechanism to control muscle length and record force. The ventral roots were divided into two bundles, each innervating about half the soleus; thus the two parts could be stimulated alone or together. In all experiments, F(nl) was small (<6% of maximum tetanic tension). Peak F(nl) occurred during changes in muscle force, either as a result of imposed muscle movement or the onset or offset of a stimulus train. The data were fit to a model in which both parts of the muscle were assumed to stretch to a common elasticity. The servomechanism was programmed to compensate for reduced stretch of the common elasticity during partial compared with whole muscle activation. These compensatory movements showed how the model could account for most, but not all, of F(nl).     

Bombesin-induced changes in membrane potential-dependent phasic contractions of cat gastric muscle

Electrical and contractile activities of smooth muscle strips isolated from the circular muscle layer of cat gastric antrum were studied using the sucrose gap technique. Bombesin (10(-8) mol/l) depolarized the gastric muscle; this was accompanied by an increase in the strip tone, in the plateau action potential frequency and in both the frequency and the amplitude of the spike potentials as well as by a shortening of the plateau action potential duration. Both the frequency and the amplitude of the phasic contractions increased thereafter. The changes in the frequency of the plateau action potentials and contractions were not influenced either by antagonists of cholinergic and adrenergic receptors or by TTX. In the presence of the Ca antagonists D600 (10(-6) mol/l) and nifedipine (10(-7) mol/l) or in Ca-free medium containing EGTA the effect of bombesin on the frequency of the plateau action potentials and phasic contractions remained unchanged; however, spike potentials were not observed and no increase in the amplitude of phasic contractions occurred. UV-light inactivation of nifedipine restored the typical bombesin effect on the electrical and contractile activities of the gastric smooth muscle. The present data suggest that the effect of bombesin on the frequency of both plateau action potentials and phasic contractions is not linked with Ca2+ influx.     

Myoelectric changes in the triceps surae muscles under sustained contractions. Evidence for synergism

Synergistic behaviour of triceps surae muscles (medial gastrocnemius-MG, lateral gastrocnemius-LG, soleus-SOL) during sustained submaximal plantarflexions was investigated in this study. Six male subjects were asked to sustain an isometric plantar flexor effort to exhaustion at two different knee angles. Exhaustion was defined as the point when they could no longer maintain the required tension. The loads sustained at 0 and 120 degrees of knee flexion represented 50% and 36% of their maximum voluntary contraction (MVC) respectively. MVC was measured at 0 degree knee flexion. During the contractions, electromyograms (EMG) from the surface of the triceps surae muscles were recorded. Changes in the synergistic behaviour of the triceps surae were assessed via partial correlations of the average EMG (AEMG) between three muscle combinations; MG/LG, MG/SOL, LG/SOL, and correlation between SOL/MG + LG and MG/SOL + LG. The latter combinations were based on either common fibre type or innervation properties. Two types of synergisms were identified: trade-off and coactivation. Trade-off and coactivation synergies were defined by significant (p less than 0.05) positive and negative correlations respectively. Coactivation synergism was found to occur predominantly under conditions of high load or reduced length of the triceps surae, and increased with the duration of the contraction. Trade-off synergism was evident when the muscles were at their optimum length and the loads sustained were submaximum.(ABSTRACT TRUNCATED AT 250 WORDS)     

Postactivation potentiation influences differently the nonlinear summation of contractions in young and elderly adults.

The force enhancement of a twitch after a maximal conditioning muscle contraction [i.e., postactivation potentiation (PAP)] is reduced with aging, but its influence on the summation of force in response to repetitive stimulation at different frequencies is not known. The purpose of this work was to compare the electrically evoked mechanical responses of the tibialis anterior muscle between young and elderly adults after a 6-s maximal voluntary contraction (MVC). The results showed that, immediately after the conditioning MVC, twitch torque and its maximal rate of development and relaxation were significantly enhanced in both groups, but the magnitude of potentiation was greater in young (148.0 +/- 14.2, 123.7 +/- 16.5, and 185.4 +/- 36.5%, respectively) compared with elderly adults (87.4 +/- 15.2, 63.8 +/- 9.9, and 62.9 +/- 11.0%, respectively). This age-related difference in potentiation of the twitch disappeared completely 1 min after the conditioning MVC. The potentiation of torque and speed-related parameters in response to two- and three-pulse trains, delivered at a constant interval of 10 ms (100 Hz), was less than for a single pulse for both groups. In young adults, the magnitude of PAP on the successive individual mechanical contributions within a train of stimuli declined progressively such that the third contribution did not differ significantly from the same contribution before the conditioning MVC. In contrast, the second and third contributions did not potentiate (P > 0.05) in elderly adults. Although these contributions did potentiate significantly at a lower frequency of stimulation (20 Hz) in the two groups, the difference in PAP between young and elderly adults still persisted. This overall attenuation of potentiation with aging, however, appears to have a moderate influence on the decrement of the muscular performance.     

Microvibration: capable of inducing spontaneous contractions in smooth muscles.

Spontaneous contractions were elicited by the vibration of small amplitude (microvibration) imposed upon the quiescent strips prepared from the ureter and portal vein of dogs. An acceleration of the existing rhythm in the spontaneously contracting strips was also brought about by microvibration. Frequencies and amplitudes of microvibration ranged from 1 to 90 Hz and from 50 to 150 mum, respectively. Imposed cyclic strains were less than 1% at their maximum. Up to the amplitude of 150 mum, the greater the amplitude, the faster the induced rhythm. Stepwise increase of the frequencies up to 30 Hz brought about a gradual acceleration of rhythmicity. At frequencies of above 30 Hz, no appreciable frequency-dependent differences were observable among the responses induced. Local adrenergic mechanism proved to have nothing to do with these effects.     

Prediction of summation in incompletely fused tetanic contractions of rat muscle

Summation is the accumulating contractile force resulting from sequential activations applied to a muscle without sufficient interval to permit complete relaxation. The purpose of this study was to evaluate summation in the rat medial gastrocnemius muscle, and to determine if the contractile responses during summation could be predicted from the relationship between force and activation pattern. In the first part of this study, the consistency of summation in the rat gastrocnemius muscle was assessed and prediction equations were derived. The second part compared predicted summation with actual contractions obtained in a new set experiments. Summation was assessed by calculation of the contractile response, per stimulation, for up to five stimulating pulses at these frequencies: 20, 40, 60 and 80Hz. This was done by subtraction of the force transient for j-1 pulses of stimulation (where j=1-5 pulses) from the force response with j pulses of stimulation. Each of these force differences was evaluated for peak rate of force development, contraction time and half-relaxation time. Contraction and half-relaxation times changed by only a small magnitude from values obtained for the twitch. Peak rate of force development was proportional to the active force for all force transients obtained by subtraction. The force per activation increased from the first to the fifth stimulus, and was dependent on interpulse delay. In the second series of experiments, the predicted force was related to the actual force for brief tetanic contractions at 40, 50 and 60Hz (r(2)=0.875). These experiments demonstrate that the force response to sequential activations is consistent and predictable. Summation can be predicted, knowing only the amplitude of the twitch contraction and the relationship between delay and force for each activating stimulus.     

Modeling of the H-reflex facilitation during ramp and hold contractions

Healthy subjects were asked to make a voluntary ramp and hold contraction. The duration of the ramp stage was 500 ms, and the torque increment in this period was set to 15 Nm. The contraction was made from a relaxed and from a 5 Nm background torque situation. Hoffmann (H-) reflexes were elicited during the voluntary contraction, mostly with 100 ms intervals. These experiments showed an increase (facilitation) in the H-reflex before the torque or the EMG started to increase. This facilitation of the H-reflex remained during all the stages of the voluntary movement and declined to normal levels again only at the very end of the hold phase, which lasted for one second. This specific pattern of facilitation during a voluntary contraction was modeled using a modeling language, that is specifically designed to calculate neuronal systems with a high degree of reality (Ekeberg et al., 1991). Our model consisted of a motoneuron pool with 200 neurons connected to an EMG-model of the human soleus muscle and an extra group of higher-level neurons for controlling the amount of decrease of presynaptic inhibition. The model was used to simulate the observed modulation of the H-reflex with both a presynaptic and a postsynaptic mechanism. Simulations showed that a continuous change in the descending control signals is needed to make the model based on postsynaptic mechanism fit with the experimental data, whereas no extra control from the CNS over the excitatory drive to the motoneuron pool is needed when the decrease of presynaptic inhibition mechanism is applied.     

Nonlinear temporal summation of EPSP in mollusk neurons

Temporal summation of monosynaptic EPSP produced by paired afferent stimuli was investigated during experiments on identified neurons of the molluskPlanorbis corneus. Summation follows a nonlinear pattern in these neurons and falls into three categories — facilitatory, depressive, and mixed. Summation functions (ratio between test and conditioning EPSP amplitudes expressed as a function of length of interstimulus interval) displayed some local maximum in most instances. Length of the interstimulus interval corresponding to this maximum correlates with the parameters of single EPSP. The possible location of mechanisms underlying the observed characteristics of EPSP summation is discussed.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 21, No. 2, pp. 204–210, March–April, 1989.     

Histochemical fiber typing and staining intensity in cat and rat muscles.

In the gastrocnemius muscle of cat and rat, staining for oxidative enzymes differentiated three fiber types (A,B,C) and staining for adenosine triphosphate at pH 9.4 differentiated two fiber types (I, II) with a reliability of 90% and 98%, respectively. In cat 96% and in rat 90% of the fibers were typed identically after staining for nicotinamide adenine dinucleotidelinked lactic dehydrogenase (LDH) and succinic dehydrogenase (SDH). When differentiated by staining for LDH, A and B fibers were of type I. IN RAT, 80-90% OF ALL FIBERS WERE OF TYPE 22, COMPPRISING A, B and C fibers. Type I fibers stained for LDH intensely as did C fibers of type II, but stained intermediately for SDH. The degree of staining was measured by photometry. When fibers were stained for LDH, histograms of density showed three peaks corresponding to A, B and C fibers in cat, but only two peaks corresponding to A and C fibers in rat, In cat and rat, the densities of A, B and C fibers belonged to different populations. In soleus muscle of cat and rat stained for LDH, menadione-linked alpha-glycerophosphate dehydrogenase and adenosine triphosphatase at pH 9.4, the degree of staining differed from thatin any type of fiber in gastrocnemius muscle     

Local muscle blood flow and sustained contractions of human arm and back muscles.

The endurance during sustained contraction of elbow, flexors, elbow extensors, and back extensors was tested in 3 human subjects. The force level used was varied between ca. 15 and ca. 75% of maximal isometric strength (IS). The clearance of 133Xe from contracting muscles was registered during and after the endurance test. In this way it was possible to determine whether muscle blood flow (MBF) was increased or had stopped during the contraction. Experiments with artificial ischaemia of the upper arm together with MBF measurements showed that MBF was of no importance for continuing sustained contractions above a certain force level, which was 50,25, and 40% of IS for elbow flexors, elbow extensors and back extensors, respectively. However, the level, where longer lasting ( greater than 15 min) sustained contraction is possible is directly related to MBF. These levels were 22, 15, and 20% IS for elbow flexors, elbow extensors, and back extensors, respectively.     

Reproducibility of surface EMG variables in isometric sub-maximal contractions of jaw elevator muscles.

The aims of this study were: (1) to develop and assess reproducibility of a new method for measuring masticatory force in the intercuspal position; (2) to test the reproducibility of surface EMG signal amplitude and spectral variables in constant force contractions of jaw elevator muscles and its dependency on inter-electrode distance. The study was performed on the masseter and temporalis anterior muscles of both sides of nine healthy volunteers. An intraoral compressive-force sensor was used to measure maximal voluntary contraction forces in the intercuspal position and to provide a visual feedback on sub-maximal forces to the subject. Three experimental sessions were performed in three days. In each session, three isometric contractions at 80% of the maximal force were sustained by the subjects for 30s. The intra-class correlation coefficient (ICC) of the maximal force measure was 71.9%. ICC of average rectified value and mean power spectral frequency of the EMG signal increased with inter-electrode distance, with values larger than 70% with 30 mm inter-electrode distance. It was concluded that surface EMG variables measured in isometric contractions of the jaw elevator muscles with the proposed force recording system show good reproducibility for clinical applications when a 30 mm inter-electrode distance is considered.     

Fatiguing contractions of tongue protrudor and retractor muscles: influence of systemic hypoxia.

The influence of systemic hypoxia on the endurance performance of tongue protrudor and retractor muscles was examined in anesthetized, ventilated rats. Tongue protrudor (genioglossus) or retractor (hyoglossus and styloglossus) muscles were activated via medial or lateral XII nerve branch stimulation (0.1-ms pulse; 40 Hz; 330-ms trains; 1 train/s). Maximal evoked potentials (M waves) of genioglossus and hyoglossus were monitored with electromyography. Fatigue tests were performed under normoxic and hypoxic (arterial PO(2) = 50 +/- 1 Torr) conditions in separate animals. The fatigue index (FI; %initial force) after 5 min of normoxic stimulation was 85 +/- 6 and 79 +/- 7% for tongue protrudor and retractor muscles, respectively; these values were significantly lower during hypoxia (protrudor FI = 52 +/- 10, retractor FI = 18 +/- 6%; P < 0.05). Protrudor and retractor muscle M-wave amplitude declined over the course of the hypoxic fatigue test but did not change during normoxia (P < 0.05). We conclude that hypoxia attenuates tongue protrudor and retractor muscle endurance performance; potential mechanisms include neuromuscular transmission failure and/or diminished sarcolemmal excitability.     

Mechanochemical melting of collagen fibers. II. Diffusion-controlled contractions

Collagen fibers were contracted “chemically,” i.e., by transferring them from water into KCNS solutions either isometrically or isotonically. Both changes in force and fiber length and in salt and water contents were measured as functions of time. The mechanical changes were found to follow the diffusional processes. The diffusion of water exhibited a plasmolytie effect. The role of water in the melting process is discussed.