Muscle force depends on the amount of transversal muscle loading

Skeletal muscles are embedded in an environment of other muscles, connective tissue, and bones, which may transfer transversal forces to the muscle tissue, thereby compressing it. In a recent study we demonstrated that transversal loading of a muscle with 1.3 N cm⁻² reduces maximum isometric force (F_im) and rate of force development by approximately 5% and 25%, respectively. The aim of the present study was to examine the influence of increasing transversal muscle loading on contraction dynamics.     

A hill-type muscle model expansion accounting for effects of varying transverse muscle load

Recent studies demonstrated that uniaxial transverse loading (F_G) of a rat gastrocnemius medialis muscle resulted in a considerable reduction of maximum isometric muscle force (ΔF_im). A hill-type muscle model assuming an identical gearing G between both ΔF_im and F_G as well as lifting height of the load (Δh) and longitudinal muscle shortening (Δl_CC) reproduced experimental data for a single load.Here we tested if this model is able to reproduce experimental changes in ΔF_im and Δh for increasing transverse loads (0.64 N, 1.13 N, 1.62 N, 2.11 N, 2.60 N). Three different gearing ratios were tested: (I) constant G_c representing the idea of a muscle specific gearing parameter (e.g. predefined by the muscle geometry), (II) G_exp determined in experiments with varying transverse load, and (III) G_f that reproduced experimental ΔF_im for each transverse load.Simulations using G_c overestimated ΔF_im (up to 59%) and Δh (up to 136%) for increasing load. Although the model assumption (equal G for forces and length changes) held for the three lower loads using G_exp and G_f, simulations resulted in underestimation of ΔF_im by 38% and overestimation of Δh by 58% for the largest load, respectively. To simultaneously reproduce experimental ΔF_im and Δh for the two larger loads, it was necessary to reduce F_im by 1.9% and 4.6%, respectively. The model seems applicable to account for effects of muscle deformation within a range of transverse loading when using a linear load-dependent function for G.     

Mechanomyogram amplitude vs. isometric ankle plantarflexion torque of human medial gastrocnemius muscle at different ankle joint angles

The purpose of this study was to investigate the influence of changes in ankle joint angle on the mechanomyogram (MMG) amplitude of the human medial gastrocnemius (MG) muscle during voluntary isometric plantarflexion contractions. Ten healthy individuals were asked to perform voluntary isometric contractions at six different contraction intensities (from 10% to 100%) and at three different ankle joint angles (plantarflexion of 26°; plantarflexion of 10°; dorsiflexion of 3°). MMG signals were recorded from the surface over the MG muscle, using a 3-axis accelerometer. The relations between root mean square (RMS) MMG and isometric plantarflexion torque at different ankle joint angles were characterized to evaluate the effects of altered muscle mechanical properties on RMS MMG.We found that the relation between RMS MMG and plantarflexion torque is changed at different ankle joint angles: RMS MMG increases monotonically with increasing the plantarflexion torque but decreases as the ankle joint became dorsiflexed. Moreover, RMS MMG shows a negative correlation with muscle length, with passive torque, and with maximum voluntary torque, which were all changed significantly at different ankle joint angles.Our findings demonstrate the potential effects of changing muscle mechanical properties on muscle vibration amplitude. Future studies are required to explore the major sources of this muscle vibration from the perspective of muscle mechanics and muscle activation level, attributable to changes in the neural command.     

Passive and dynamic muscle architecture during transverse loading for gastrocnemius medialis in man

External forces from our environment impose transverse loads on our muscles. Studies in rats have shown that transverse loads result in a decrease in the longitudinal muscle force. Changes in muscle architecture during contraction may contribute to the observed force decrease. The aim of this study was to quantify changes in pennation angle, fascicle dimensions, and muscle thickness during contraction under external transverse load.Electrical stimuli were elicited to evoke maximal force twitches in the right calf muscles of humans. Trials were conducted with transverse loads of 2, 4.5, and 10 kg. An ultrasound probe was placed on the medial gastrocnemius in line with the transverse load to quantify muscle characteristics during muscle twitches.Maximum twitch force decreased with increased transverse muscle loading. The 2, 4.5, and 10 kg of transverse load showed a 9, 13, and 16% decrease in longitudinal force, respectively. Within the field of view of the ultrasound images, and thus directly beneath the external load, loading of the muscle resulted in a decrease in the muscle thickness and pennation angle, with higher loads causing greater decreases. During twitches the muscle transiently increased in thickness and pennation angle, as did fascicle thickness. Higher transverse loads showed a reduced increase in muscle thickness. Smaller increases in pennation angle and fascicle thickness strain also occurred with higher transverse loads.This study shows that increased transverse loading caused a decrease in ankle moment, muscle thickness, and pennation angle, as well as transverse deformation of the fascicles.     

Effect of insulin and contraction up on glucose transport in skeletal muscle

The major glucose transporter protein expressed in skeletal muscle is GLUT4. Both muscle contraction and insulin induce translocation of GLUT4 from the intracellular pool to the plasma membrane. The intracellular pathways that lead to contraction- and insulin-stimulated GLUT4 translocation seem to be different, allowing the attainment of a maximal effect when acting together. Insulin utilizes a phosphatidylinositol 3-kinase-dependent mechanism, whereas the exercise signal may be initiated by calcium release from the sarcoplasmic reticulum or from autocrine- or paracrine-mediated activation of glucose transport. During exercise skeletal muscle utilizes more glucose than when at rest. However, endurance training leads to decreased glucose utilization during sub-maximal exercise, in spite of a large increase in the total GLUT4 content associated with training. The mechanisms involved in this reduction have not been totally elucidated, but appear to cause the decrease of the amount of GLUT4 translocated to the plasma membrane by altering the exercise-induced enhancement of glucose transport capacity. On the other hand, the effect of resistance training is controversial. Recent studies, however, demonstrated the improvement in insulin sensitivity correlated with increasing muscle mass. New studies should be designed to define the molecular basis for these important adaptations to skeletal muscle. Since during exercise the muscle may utilize insulin-independent mechanisms to increase glucose uptake, the mechanisms involved should provide important knowledge to the understanding and managing peripheral insulin resistance.     

Effects of temperature on electromyogram and muscle function

The effects of 30 min of cooling (15 degrees C water) and warming (40 degrees C water) on arm muscle function were measured. A reference condition (24 degrees C air) was included. Of nine young male subjects the maximal grip force (Fmax), the time to reach 66% of Fmax (rate of force buildup) and the maximal rhythmic grip frequency were determined, together with surface electromyographic activity (EMG) of a forearm muscle (flexor digitorum superficialis). The results showed that in contrast to warming, cooling resulted in a significant decrease of 20% in the Fmax and a significant 50% decrease in force build-up time and the maximal rhythmic grip frequency. The relationship between the root mean square value (rms) of the EMG and the static grip force did not change due to temperature changes. The median power frequency (MPF) in the power spectrum of the EMG signal decreased by 50% due to cooling but remained unchanged with heating. During a sustained contraction at 15% of Fmax (Fmax depending on the temperature) the increase in the rms value with contraction time was 90% larger in the warm condition and 80% smaller in the cold condition compared to the increase in the reference condition. The MPF value remained constant during the warm and reference conditions but in the cold it started at a 50% lower value and increased with contraction time.(ABSTRACT TRUNCATED AT 250 WORDS)     

Viscosity as an inseparable partner of muscle contraction

A simple model is presented where, by an iterative procedure, the forces delivered by the power strokes are summed up to overcome the load. The system is moderated by the viscous hindrance. The model reproduces the features of muscle contraction as defined by the data of He et al. [1997. ATPase kinetics on activation of permeabilized isometric fibres from rabbit and frog muscle: a real time phosphate assay. J. Physiol. 501, 125-148] on rabbit psoas muscle fibres. According to the model power strokes are random. Energy summation take place if the subsequent power stroke occurs before the energy delivered by the previous power stroke is completely used. In order the sarcomere to be competent to contract initial driving force must reach a threshold whose value increases with the load. The step size of the power stroke decreases with the increase of the load. The viscous regime is simulated by the equation, where 1/k measures the viscous hindrance of the system. The relationship between water activity, viscosity and stiffness is discussed. It is concluded that the three parameters vary cyclically and that when water activity decreases (sarcomere shortening, cross-bridge attachment) viscosity and stiffness increase.     

Attachment of muscle filaments to the outer membrane of the nuclear envelope

Summary Myofilaments of striated muscles can be recognized in the electron microscope to be in structural continuity with the outer membrane of the nuclear envelope. The very site of insertion of these myofilaments at the membrane surface frequently appears characterized by a dense basal knob of 85–135 Å. It is hypothesized that this attachment of myofilaments to the nuclear membrane plays a role in mechanically transmitting the contraction of the fiber to the nucleus, thus bringing about the harmonica-like folded appearance of the nucleus which is known for the contracted states of striated, smooth and cardiac muscles.The work was supported in part by the Deutsche Forschungsgemeinschaft.The author is indebted to Miss Sigrid Krien and Miss Marianne Winter for careful technical assistance as well as to Drs. Heinz Falk and U. Scheer for valuable discussions.     

Surface mechanomyogram reflects the changes in the mechanical properties of muscle at fatigue

The contractile properties of muscle are usually investigated by analysing the force signal recorded during electrically elicited contractions. The electrically stimulated muscle shows surface oscillations that can be detected by an accelerometer; the acceleration signal is termed the surface mechanomyogram (MMG). In the study described here we compared, in the human tibialis anterior muscle, changes in the MMG and force signal characteristics before, and immediately after fatigue, as well as during 6 min of recovery, when changes in the contractile properties of muscle occur. Fatigue was induced by sustained electrical stimulation. The final aim was to evaluate the reliability of the MMG as a tool to follow the changes in the mechanical properties of muscle caused by fatigue. Because of fatigue, the parameters of the force peak, the peak rate of force production and the peak of the acceleration of force production (d2F/dt2) decreased, while the contraction time and the half-relaxation time (1/2-RT) increased. The MMG peak-to-peak (p-p) also decreased. The attenuation rate of the force oscillation amplitude and MMG p-p at increasing stimulation frequency was greater after fatigue. With the exception of 1/2-RT, all of the force and MMG parameters were restored within 2 min of recovery. A high correlation was found between MMG and d2F/dt2 in un-fatigued muscle and during recovery. In conclusion, the MMG reflects specific aspects of muscle mechanics and can be used to follow the changes in the contractile properties of muscle caused by localised muscle fatigue.     

The influence of agonist premotor silence and the stretch-shortening cycle on contractile rate in active skeletal muscle

Agonist premotor silence (PMS), a brief period of relative quiescence in active skeletal muscle prior to phasic activation, was investigated in subjects performing maximal contractions. The frequency of occurrence and potential function of the silent period were examined for elbow flexions and extensions. PMS was evident for movements in both directions, indicating that the mechanism is not primarily limited to extensors as previously hypothesized. Flexions demonstrating PMS exhibited increased velocity and acceleration; however, kinematic facilitation was only evident on trials exhibiting the muscular stretch-shortening cycle (SSC). The SSC was present on trials lacking PMS, demonstrating that biceps and triceps silence are not the sole determinants of preparatory agonist lengthening for elbow flexions and extensions, respectively. Taken together, the data indicate that agonist PMS is a mechanism under apparent central control that acts concomitantly with mechanical factors to potentiate elbow flexor contractions.     

Contractile properties of human skeletal muscle in childhood and adolescence

Using a combination of single maximal stimuli and maximum voluntary contractions, a comparison has been made of muscle properties in pre- and post-pubertal male subjects. In the dorsiflexor and plantarflexor muscles of the ankle, the twitch and maximum voluntary torques were approximately twice as large in the older subjects; the mean height and mean weight increased by factors of 1.20 and 1.86 respectively. The only other muscle parameter that changed, as a function of age, was the contraction time of the ankle dorsiflexors; the mean value was significantly longer in the older subjects. In the younger subjects, there were already clear differences between the dorsiflexor and plantarflexor muscles, the former developing smaller torques and having shorter contraction and half-relaxation times, greater post-activation potentiation and more susceptibility to fatigue. Even in the youngest subject, motor unit activation was complete in the ankle dorsiflexors; although this was not always true of the plantarflexors, the difference between the two subject groups was not significant.     

Normalization of electromyogram in the neck-shoulder region

Linear and curvilinear electromyogram (EMG) normalization methods were compared among ten healthy men during a simulated work cycle demanding attention and static holding of the arm (Solitaire test). Maximal voluntary contractions (MVC) and gradually increasing contractions up to 70% of MVC were used for normalization in different arm postures. The test contractions studied included inward and outward rotations, abduction, shoulder elevation, and flexion in different arm positions. The shoulder load moment was calculated for the flexion tests using a simple two-dimensional model. The effect of arm posture on the EMG versus shoulder load moment relationship was studied on the following muscles: supraspinatus, infraspinatus, trapezius (three parts), deltoid (two parts) and pectoralis major. All muscles participated in the MVC tests performed, and its was not possible to suggest a single recommended test for each muscle. Differences in normalized EMG median values ranging up to 30% of MVC were found between linear and curvilinear normalization methods. Short-term repeatability of normalization based on a contraction with gradually increasing force was good. Arm posture affected the relationships between shoulder load moment and EMG activity of all muscles studied. Arm posture did not, however, have a significant effect on the estimated amplitude probability distribution functions during the simulated work task. Therefore, at least for the tasks studied, the principle of normalizing in the middle position of the range of movement was deemed acceptable.     

The effect of prolonged isometric contractions on muscle fluid balance

Ultrasound scanning was performed at three sites above the fossa supraspinata on nine healthy subjects and five patients with myofascial shoulder pain. This method produced a well-defined depiction of the soft tissue layers above the fossa supraspinata and reproducible muscle thickness measurements. In the healthy subjects the average distance from the skin surface to the trapezius muscle was 7.7 mm and the average thickness of the trapezius muscle was 5.3 mm, and the average thickness of supraspinatus muscle was 20.0 mm. The supraspinatus muscle was thinner at the medial measuring site than at the other two sites. In contrast, a tendency towards a larger distance was seen from the skin to trapezius muscle at the medial measuring site than at the other two sites. No statistical differences were found between the two groups of subjects either at rest or during brief shoulder abductions. All the subjects performed a 30° unilateral isometric shoulder abduction test to exhaustion. The median endurance time was 33 min for the healthy subjects and only 5 min for the patients. The ratings of perceived exertion (RPE) were in line with this, since the increment in RPE with time was larger for the patients than for the healthy group. The reduced shoulder abduction endurance time in the patient group may have been related to impaired muscle function and/or pain development. During the 33-min shoulder abduction in the healthy subjects, the thickness of supraspinatus muscle increased by 14%, indicating muscle swelling, whereas the thickness of trapezius muscle remained constant. The fluid imbalance in the supraspinatus muscle compartment may well play a role in the development of muscle fatigue and the disorders found in industry resulting from prolonged work with arms elevated.     

Effects of contraction duration on low-frequency fatigue in voluntary and electrically induced exercise of quadriceps muscle in humans

The aims of this study were to investigate if low-frequency fatigue (LFF) dependent on the duration of repeated muscle contractions and to compare LFF in voluntary and electrically induced exercise. Male subjects performed three 9-min periods of repeated isometric knee extensions at 40% maximal voluntary contraction with contraction plus relaxation periods of 30 plus 60 s, 15 plus 30 s and 5 plus 10 s in protocols 1, 2 and 3, respectively. The same exercise protocols were repeated using feedback-controlled electrical stimulation at 40% maximal tetanic torque. Before and 15 min after each exercise period, knee extension torque at 1, 7, 10, 15, 20, 50 and 100 Hz was assessed. During voluntary exercise, electromyogram root mean square (EMG_rms) of the vastus lateralis muscle was evaluated. The 20-Hz torque:100-Hz torque (20:100 Hz torque) ratio was reduced more after electrically induced than after voluntary exercise (P < 0.05). During electrically induced exercise, the decrease in 20:100 Hz torque ratio was gradually (P < 0.05) reduced as the individual contractions shortened. During voluntary exercise, the decrease in 20:100 Hz torque ratio and the increase in EMG_rms were greater in protocol 1 (P < 0.01) than in protocols 2 and 3, which did not differ from each other. In conclusion, our results showed that LFF is dependent on the duration of individual muscle contractions during repetitive isometric exercise and that the electrically induced exercise produced a more pronounced LFF compared to voluntary exercise of submaximal intensity. It is suggested that compensatory recruitment of faster-contracting motor units is an additional factor affecting the severity of LFF during voluntary exercise. Accepted: 5 November 1997     

Calcium efflux during the cold-induced contraction of mammalian striated muscle fibres

The efflux of ⁴⁵Ca from mammalian slow twitch muscle fibres has been studied to provide a measure of the concentration of free Ca²⁺ in the sarcoplasm. The kinetically complex early phases of washout of the isotope are succeeded by a prolonged slower phase which exhibits first-order kinetics. This later phase is accelerated by caffeine, by preventing oxidative phosphorylation and also during an isometric contraction, whether this contraction is produced by lowering the temperature or by electrical stimulation. The local anaesthetic tetracaine abolishes the contraction caused by cold and in this case the rate constant for efflux is progressively lowered as the temperature is reduced (Q₁₀ value of 2.3). The removal of external Na⁺ and Ca²⁺ reduces the efflux rate constant. Caffeine, sodium removal and the inhibition of oxidative phosphorylation, all potentiate the cold contraction and the associated extra ⁴⁵Ca efflux. Ca removal causes the cold contraction to become phasic. It appears that caffeine, sodium removal, the inhibition of oxidative phosphorylation and a decrease in temperature to below 10°C are all treatments which, like electrical stimulation, increase the sarcoplasmic free calcium concentration to varying degrees.     

Reliability of muscle fibre conduction velocity distribution estimation from surface EMG

Most of the neuromuscular diseases induce changes in muscle fibre characteristics. For example, Duchenne dystrophy is characterized by a specific loss of fast fibres, and an increase in small diameter fibres. These morphological changes may lead to large modifications in the distribution of fibre diameters, possibly producing bimodal distributions. It has already been shown that it is possible to reveal these morphological modifications through the distribution of muscle fibre conduction velocity (MFCV) as estimated from needle electromyography (EMG) recordings. In this paper, we investigate whether such changes can be extracted from surface EMG signals.

Simulation allows generation of surface EMG signals in which features are well described especially at a morphological level. Therefore, we generated a database of simulated signals both in voluntary and electrically elicited contraction conditions using a bimodal distribution of muscle fibre diameters. MFCV distributions were computed using two short-term methods based on cross-correlation and peak-to-peak techniques for voluntary contraction signals, and using a deconvolution method in time domain for electrically elicited signals. MFCV distributions were compared with true ones, as generated from modelling.

This work reveals that estimating MFCV distribution through these methods does not appear yet as precise and robust enough to accurately characterize changes in redistribution of various muscle fibre diameters. However, it opens to new experimental protocols that can be explored in order to improve the robustness of MFCV distribution estimation for the follow-up of patients suffering from neuromuscular disorders.     

Effect of phospholipase A on actions of cobra venom cardiotoxins on erythrocytes and skeletal muscle

The actions of two phospholipase-free cardiotoxins from the venom of the cobra Naja naja siamensis were compared to phospholipase-contaminated cardiotoxins in terms of their ability to lyse human erythrocytes and to depolarize and contract skeletal muscle. The presence of 3–5% (w/w) phospholipase caused a 20–30-fold increase in the haemolytic activity of the two cardiotoxins, the pure cardiotoxins being virtually without haemolytic activity at 10^?7-10^?6 M. Phospholipase contamination did not enhance the ability of the cardiotoxins to cause contracture of chick biventer cervicis muscles and it caused less than a 2-fold increase in the depolarizing activity of the cardiotoxins on cultured skeletal muscle. Phospholipase-free cardiotoxins were about 10–20-times more active on cultured skeletal muscle fibres than on erythrocytes. These results support the hypothesis that some cardiotoxins have more affinity for the membranes of excitable cells than for those of other cells such as erythrocytes.     

Effect of deuterium oxide on contraction characteristics and ATPase activity in glycerinated single rabbit skeletal muscle fibers

We studied the effect of deuterium oxide (D₂O) on contraction characteristics and ATPase activity of single glycerinated muscle fibers of rabbit psoas. D₂O increased the maximum isometric force P₀ by about 20%, while the force versus stiffness relation did not change appreciably. The maximum shortening velocity under zero load V_max did not change appreciably in D₂O, so that the force-velocity (P-V) curve was scaled depending on the value of P₀. The Mg-ATPase activity of the fibers during generation of steady isometric force P₀ was reduced by about 50% in D₂O. Based on the Huxley contraction model, these results can be accounted for in terms of D₂O-induced changes in the rate constants f₁ and g₁ for making and breaking actin-myosin linkages in the isometric condition, in such a way that f₁/(f₁+g₁) increases by about 20%, while (f₁+g₁) remains unchanged. The D₂O effect at the molecular level is discussed in connection with biochemical studies on actomyosin ATPase.     

Measurement of muscle and tendon stiffness in man

Human first dorsal interosseous muscle was stimulated tetanically using several levels of percutaneous electrical current which produced forces in the muscle-tendon complex of between 30% and 100% of maximum. During the tetanus the muscle was subjected to a small fast stretch. The ratio of the force response to the displacement of the muscle-tendon complex gave a measure of the stiffness of the total complex. An adaptation of the method of Morgan (1977) allowed the stiffness to be separated into two components the stiffness of the muscle fibres and the stiffness of the tendon. The results showed that at full activation the stiffness of the muscle fibres and the tendon are approximately the same. The normalised stiffness values obtained in the experiments compared well with animal data.     

Electric dipole theory and thermodynamics of actomyosin molecular motor in muscle contraction

Movements in muscles are generated by the myosins which interact with the actin filaments. In this paper we present an electric theory to describe how the chemical energy is first stored in electrostatic form in the myosin system and how it is then released and transformed into work. Due to the longitudinal polarized molecular structure with the negative phosphate group tail, the ATP molecule possesses a large electric dipole moment (p(0)), which makes it an ideal energy source for the electric dipole motor of the actomyosin system. The myosin head contains a large number of strongly restrained water molecules, which makes the ATP-driven electric dipole motor possible. The strongly restrained water molecules can store the chemical energy released by ATP binding and hydrolysis processes in the electric form due to their myosin structure fixed electric dipole moments (p(i)). The decrease in the electric energy is transformed into mechanical work by the rotational movement of the myosin head, which follows from the interaction of the dipoles p(i) with the potential field V(0) of ATP and with the potential field Psi of the actin. The electrical meaning of the hydrolysis reaction is to reduce the dipole moment p(0)-the remaining dipole moment of the adenosine diphosphate (ADP) is appropriately smaller to return the low negative value of the electric energy nearly back to its initial value, enabling the removal of ADP from the myosin head so that the cycling process can be repeated. We derive for the electric energy of the myosin system a general equation, which contains the potential field V(0) with the dipole moment p(0), the dipole moments p(i) and the potential field psi. Using the previously published experimental data for the electric dipole of ATP (p(0) congruent with 230 debye) and for the amount of strongly restrained water molecules (N congruent with 720) in the myosin subfragment (S1), we show that the Gibbs free energy changes of the ATP binding and hydrolysis reaction steps can be converted into the form of electric energy. The mechanical action between myosin and actin is investigated by the principle of virtual work. An electric torque always appears, i.e. a moment of electric forces between dipoles p(0) and p(i)(/M/ > or = 16 pN nm) that causes the myosin head to function like a scissors-shaped electric dipole motor. The theory as a whole is illustrated by several numerical examples and the results are compared with experimental results.