Architectural and functional features of human triceps surae muscles during contraction

Architecturalproperties of the triceps surae muscles were determined in vivo for sixmen. The ankle was positioned at 15° dorsiflexion (15°)and 0, 15, and 30° plantar flexion, with the knee set at 0, 45, and90°. At each position, longitudinal ultrasonic images of the medial(MG) and lateral (LG) gastrocnemius and soleus (Sol) muscles wereobtained while the subject was relaxed (passive) and performed maximalisometric plantar flexion (active), from which fascicle lengths andangles with respect to the aponeuroses were determined. In the passivecondition, fascicle lengths changed from 59, 65, and 43 mm (knee,0°; ankle, 15°) to 32, 41, and 30 mm (knee, 90°ankle, 30°) for MG, LG, and Sol, respectively. Fascicle shorteningby contraction was more pronounced at longer fascicle lengths. MG hadgreatest fascicle angles, ranging from 22 to 67°, and was in a verydisadvantageous condition when the knee was flexed at 90°,irrespective of ankle positions. Different lengths and angles offascicles, and their changes by contraction, might be related todifferences in force-producing capabilities of the muscles and elasticcharacteristics of tendons and aponeuroses.

     

Sway-dependent modulation of the triceps surae H-reflex during standing.

Previous research has shown that changes in spinal excitability occur during the postural sway of quiet standing. In the present study, it was of interest to examine the independent effects of sway position and sway direction on the efficacy of the triceps surae Ia pathway, as reflected by the Hoffman (H)-reflex amplitude, during standing. Eighteen participants, tested under two different experimental protocols, stood quietly on a force platform. Percutaneous electrical stimulation was applied to the posterior tibial nerve when the position and direction of anteroposterior (A-P) center of pressure (COP) signal satisfied the criteria for the various experimental conditions. It was found that, regardless of sway position, a larger amplitude of the triceps surae H-reflex (difference of 9-14%; P = 0.005) occurred when subjects were swaying in the forward compared with the backward direction. The effects of sway position, independent of the sway direction, on spinal excitability exhibited a trend (P = 0.075), with an 8.9 +/- 3.7% increase in the H-reflex amplitude occurring when subjects were in a more forward position. The observed changes to the efficacy of the Ia pathway cannot be attributed to changes in stimulus intensity, as indicated by a constant M-wave amplitude, or to the small changes in the level of background electromyographic activity. One explanation for the changes in reflex excitability with respect to the postural sway of standing is that the neural modulation may be related to the small lengthening and shortening contractions occurring in the muscles of the triceps surae.     

Spontaneous activity of passive muscle spindles of the cat triceps surae muscle

The hypothesis that one possible cause of the spontaneous discharge of muscle spindles in the totally relaxed and de-efferented triceps surae muscle, with its tendon divided, is the mechanical factor due to extrafusal-intrafusal interaction was tested in experiments on anesthetized cats. Responses of spontaneously active units were similar with respect to many indices to responses of silent receptors capable of generating a long discharge in response to an increase in static length of the muscle. Isotonic contraction of the relaxed muscle during direct or indirect stimulation was accompanied by a pause in spontaneous activity. The prolonged increase in discharge frequency sometimes arising as a result of pressure on the muscle in the region of the receptor also was abolished by weak isotonic contraction of the muscle. If a long posttetanic sensory discharge was induced in a spontaneously active receptor by intensive tetanization of the nerve to the muscle, and it was then abolished by short stretching of the muscle, the initial spontaneous discharge frequency was restored. The dynamic thresholds of spontaneously active receptors were lower than for silent receptors. In some spontaneously active receptors an initial slowing of the discharge was observed during linear or stepwise stretching of the muscle. It is suggested that the ability of sensory endings to generate a long spontaneous discharge is due to initial stretching of the spindle inside the relaxed muscle.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 10, No. 3, pp. 287–294, May–June, 1978.     

Mechanomyographic and electromyographic responses of the triceps surae during maximal voluntary contractions.

The objective of this study was to examine the effect of joint angle on the electromyogram (EMG) and mechanomyogram (MMG) during maximal voluntary contraction (MVC). Eight subjects performed maximal isometric plantar flexor torque productions at varying knee and/or ankle angles. Maximal voluntary torque, EMG, and MMG from the soleus (Sol), medial (MG) and lateral gastrocnemius (LG) muscles were measured at different joint angles. At varying knee angles, the root mean squared (rms) MMG amplitude of the MG and LG increased with knee joint extension from 60 degrees to 180 degrees (full extension) in steps of 30 degrees, whereas that of the Sol was constant. At varying ankle angles, the rms-MMG of all muscles (Sol, MG, and LG) decreased with torque as ankle joint extending from 80 degrees (10 degrees dorsiflexion position) to 120 degrees (30 degrees plantar flexion position) in steps of 10 degrees. In each case, changes in the rms-MMG of the three muscles were almost parallel to those in torque. In contrast, there were no significant differences in the rms-EMG of all muscles among all joint angles. Our data suggest that the MMG amplitudes recorded from individual muscles during MVCs can represent relative torque-angle relationships that cannot be represented by the EMG signals.     

The membrane systems in different fibre types of the triceps surae muscle of cat

Summary The volume and surface area of mitochondria and sarcoplasmic reticulum in fast and slow twitch fibres of the cat triceps surae muscle were determined from thin sections. The width of the Z-line and the array of glycogen granules identified fast and slow twitch fibres.The relative volume occupied by mitochondria was largest in slow twitch gastrocnemius fibres. Fast twitch fibres showed the greatest scatter of mitochondrial content. This corresponds with the fact that motor units of the fast twitch type differ most with respect to resistance to fatigue.The relative volume of the sarcoplasmic reticulum was twice as large in fast as in slow twitch fibres. The volume fraction occupied by longitudinal tubules of the reticulum was the same in fast and slow twitch gastrocnemius fibres but was only half as large in the slow twitch soleus fibres. This difference may be related to post-tetanic potentiation: this property is present in all gastrocnemius fibres but is absent in soleus fibres.The specific tetanic force is 3 to 5 times smaller in slow twitch gastrocnemius than in slow twitch soleus fibres or fast gastrocnemius fibres. There was, however, no detectable morphological difference that might be related to this difference in force.Freeze fractures demonstrated directly that, in soleus fibres, terminal cisternae and longitudinal tubules of the reticulum were scarce as compared to gastrocnemius fibres. The plasma membranes of some gastrocnemius fibres displayed square arrays of 60-nm particles; these arrays were absent in other gastrocnemius fibres and in all soleus fibres. They probably characterize plasma membranes of fast twitch fibres.This study was supported by grants from the Danish Medical Research Council. I wish to thank Mrs. M. Bjærg for valuable technical help     

Fatigue responses of human triceps surae muscles during repetitive maximal isometric contractions.

Nine healthy men (22-45 yr) completed 100 repetitive maximal isometric contractions of the ankle plantar flexor muscles in two knee positions of full extension (K0) and flexion at 90 degrees (K90), positions that varied the contribution of the gastrocnemii. Electromyographic activity was recorded from the medial and lateral gastrocnemii and soleus muscles by using surface electrodes. Plantar flexion torque in K0 was greater and decreased more rapidly than in K90. The electromyographic amplitude decreased over time, and there were no significant differences between muscles and knee joint positions. The level of voluntary effort, assessed by a supramaximal electrical stimulation during every 10th contraction, decreased from 96 to 70% (P < 0.05) with no difference between K0 and K90. It was suggested that a decrease in plantar flexion torque was attributable to both central and peripheral fatigue and that greater fatigability in K0 than in K90 would result from a greater contribution and hence more pronounced fatigue of the gastrocnemius muscle. Further support for this possibility was provided from changes in twitch torque.     

Effect of muscle contraction levels on the force-length relationship of the human Achilles tendon during lengthening of the triceps surae muscle-tendon unit

Findings from animal experiments are sometimes contradictory to the idea that the tendon structure is a simple elastic spring in series with muscle fibers, and suggest influence of muscle contraction on the tendon mechanical properties. The purpose of the present study was to investigate the influence of muscle contraction levels on the force-length relationship of the human Achilles tendon during lengthening of the triceps surae muscle-tendon unit. For seven subjects, ankle dorsiflexion was performed without (passive condition) and with contraction of plantar flexor muscles (eccentric conditions, at 3 contraction levels) on an isokinetic dynamometer. Deformation of the Achilles tendon during each trial was measured using ultrasonography. The Achilles tendon force corresponding to the tendon elongation of 10mm in the passive condition was significantly smaller than those in the eccentric conditions (p<0.05 or p<0.01). Within the eccentric conditions, the Achilles tendon force corresponding to the tendon elongation of 10mm was significantly greater in the maximal contraction level than those in submaximal eccentric conditions (p<0.05 or p<0.01). In addition, the tendon stiffness was greater in higher contraction levels (p<0.05 or p<0.01). Present results suggest that the human tendon structure is not a simple elastic spring in series with muscle fibers.     

Passive triceps surae stretch inhibits vasoconstriction in the nonexercised limb during posthandgrip muscle ischemia.

We investigated whether selective muscle mechanoreceptor activation in the lower limb opposes arm muscle metaboreceptor activation-mediated limb vasoconstriction. Seven subjects completed two trials: one control trial and one stretch trial. Both trials included 2 min of handgrip and 2 min of posthandgrip exercise muscle ischemia (PEMI). In the stretch trial, a 2-min sustained triceps surae stretch, by brief passive dorsiflexion of the right foot, was performed simultaneously during PEMI. Mean arterial pressure, heart rate, and forearm blood flow (FBF) in the nonexercised arm and forearm vascular conductance (FVC) in the nonexercised arm were measured. During PEMI in the control trial, mean arterial pressure was significantly greater and FBF and FVC were significantly lower than baseline values (P < 0.05 for each). In contrast, FBF and FVC during PEMI in the stretch trial exhibited different responses than in the control trial. FBF and FVC were significantly greater in the stretch trial than in the control trial (FBF, 5.5 +/- 0.4 vs. 3.8 +/- 0.4 ml x 100 ml(-1) x min(-1); FVC, 0.048 +/- 0.004 vs. 0.033 +/- 0.003 unit, respectively; P < 0.05). These results indicate that passive triceps surae stretch can inhibit vasoconstriction in the nonexercised forearm mediated via muscle metaboreceptor activation in the exercised arm.     

Voluntary activation of the triceps surae in prepubertal children

The activation capacities and neuromuscular efficiency (NME) of the triceps surae (TS) of prepubescent children (7–11 years) and adults were evaluated during submaximal and maximal (MVC) isometric plantarflexion to determine whether they varied with age. TS-EMG were obtained by summing-up the rectified electromyograms of the soleus and gastrocnemii muscles; these data were quantified using a sliding average method and normalized with reference to the TS maximal compound action potential (TS-M-wave).

The maximal EMG increased significantly with age in the children, but less than MVC, what led to a significant increase in NME_Max (MVC/TS-EMG_max ratio). The EMG–torque relationship indicated an age-related overactivation of TS at low torque, what led to a lower NME_Sub-max (inverse of the slope of the EMG–torque relationship) for the youngest children. The overactivation of TS was accompanied by contraction of the TA, which decreased with age. The youngest children were also less able to maintain a target torque and muscle activation. Finally, the twitch interpolated method revealed an age-dependant activation deficit. We conclude that central mechanisms are the main cause of the lower torques developed by children and they appear to vary with age in prepubertal children.     

A comparison of the triceps surae and residual muscle moments at the ankle during cycling

The rigid linked system model and principles of inverse dynamics have been widely used to calculate residual muscle moments during various activities. EMG driven models and optimization algorithms have also been presented in the literature in efforts to estimate skeletal muscle forces and evaluate their possible contribution to the residual muscle moment. Additionally, skeletal muscle-tendon forces have been measured, directly, in both animals and humans. The purpose of this investigation was to calculate the moment produced by the triceps surae muscles and compare it to the residual muscle moment at the ankle during cycling at three power outputs (90, 180 and 270 W). Inferences were made regarding the potential contribution made by each triceps surae component to the tendon force using EMG and muscle-tendon length changes. A buckle-type transducer was surgically implanted on the right Achilles tendon of one male subject. Achilles tendon forces measured in vivo were multiplied by their corresponding moment arms to yield the triceps surae moment during the three working conditions. Moment arm lengths were obtained in a separate experiment using magnetic resonance imaging (MRI). Pedal reaction forces, body segment accelerations (determined from high speed film), and appropriate mass parameters served as input to the inverse solution. The triceps surae moment was temporally in phase with and consistently represented approximately 65% of the residual muscle moment at the ankle. These data demonstrate the feasibility of using implanted transducers in human subjects and provide a greater understanding of musculoskeletal mechanics during normal human movements.     

Muscle activation and its distribution within human triceps surae muscles.

The purposes of this study were 1) to quantify the volume of activated parts within a whole muscle and 2) to examine activated area distributions along the length of muscle. Seven male subjects performed five sets of 10 repetitions of a single-leg calf-raise exercise with the knee fully extended. Transverse relaxation time (T2)-weighted spin echo images were acquired before and immediately after the exercise. A range of pixels with a T2 greater than the mean +1 SD of the region of interest (ROI) from the preexercise image and pixels with a T2 lower than the mean + SD of the ROI from the postexercise image were defined as "active" muscle. The active muscle images were three dimensionally reconstructed, from which the volume of the activated muscle was determined for individual triceps surae (TS) muscles. Our data indicate that approximately 46% of the medial gastrocnemius (MG) muscle was activated during the exercise, with activation of the lateral gastrocnemius (LG) and soleus (Sol) muscles being approximately 35%. In the MG, distal portions had a greater percentage area of activated muscle than the proximal portions (P < 0.05), which was consistent with the results regarding electromyogram activity. In contrast, regional activation differences were not observed in the LG and Sol. These findings suggest that the amounts of activated muscle and its distribution would be different among TS muscles.     

Contractile adaptations in the human triceps surae after isometric exercise

Ultrastructural and twitch contractile characteristics of the human triceps surae were determined in seven healthy but very sedentary subjects before and after 16 wk of unilateral isometric training at 100% maximal voluntary contraction. After training, twitch contraction time decreased by approximately 20%. One-half relaxation time, peak twitch torque, and percent fiber type in any of the muscles of the triceps surae complex were not changed by training. Type I and type II fiber areas increased in the soleus by approximately 30%, but only type II fibers showed an increased in area in the lateral gastrocnemius (40%). Despite such changes in fiber area, the volume density of the sarcoplasmic reticulum-transverse tubular (SR) network averaged 3.2 +/- 0.6 and 5.9 +/- 0.9% in type I and type II fibers, respectively, before and after training in the two heads of the gastrocnemius. Type I SR fraction increased to 3.5 +/- 1.2% after training in the soleus; however, correlations were not significant between the change in the volume density of SR and the change in twitch contraction time (R = 0.46, P = 0.45) or the change in one-half relaxation time (R = -0.68, P = 0.08). The results demonstrate that isometric training at 100% maximal voluntary contraction induced changes in twitch contraction time that were not directly related to changes in the volume density of SR in fibers of the triceps surae.     

A comparison of electrical activity in the triceps surae at maximum isometric contraction with the knee and ankle at various angles

The purpose of this study was to test the endurance of the soleus muscle, and to examine the joint position at which it is most active, while simultaneously suppressing the activity of the gastrocnemius. Ten young males performed maximum isometric contraction of the triceps surae for 100 s, and the endurance and plantar flexion torque of this muscle were measured at various angles of the knee and ankle joints. The electromyogram was measured simultaneously and subsequently converted into integrated electromyogram (IEMG) values. With the knee flexed at 130 degrees, the rate of change in IEMG values for the soleus (0.454% x s(-1)) with the ankle in a neutral position was significantly higher than that for the medial and lateral gastrocnemius. Both with the ankle dorsiflexed at 10 degrees and in the neutral position, the rate of change in IEMG for the soleus was significantly higher with the knee flexed at 90 degrees and 130 degrees than with the knee fully extended. With the knee flexed at 90 degrees and 130 degrees, the IEMG activity of the soleus during the initial (5-10 s) and final 5 s tended to be higher than those for the medial and lateral gastrocnemius, regardless of the ankle joint position. We conclude that the position in which the soleus acts most selectively during a sustained maximum isometric contraction of the triceps surae is with the ankle in a neutral position and the knee flexed at 130 degrees.     

Electrically evoked contractions of the triceps surae during and following 21 days of voluntary leg immobilization

The effects of 21 days voluntary leg (plaster) immobilization on the mechanical properties of the triceps surae have been studied in 11 young female subjects, mean age 19.4 years. The results show that during the period of immobilization the mean time to peak tension (TPT) and half relaxation time (1/2RT) and tension (Pt) of the maximal twitch increased significantly (p less than 0.001) but the effects were short lived. Maximal tension and contraction times of the twitch recovered within 2-14 days following the removal of the plaster cast. The electrically evoked tetanic tensions at 10 Hz and 20 Hz did not change significantly (P greater than 0.1) during immobilization, but the 50 Hz tetanic tension (Po50) and maximal voluntary contraction (MVC) were reduced (p less than 0.05). The fall in Po50 and MVC was associated with 10% decrease in the estimated muscle (plus bone) cross-sectional area. The relative (%) change in Po50 and MVC following immobilization was related to the initial physiological status (as indicated by the response of the triceps surae to a standard fatigue test prior to immobilization) of the muscle. The rate of rise and recovery fall of the tetanus were slightly but significantly (p less than 0.01) reduced on day 7 of immobilization, but thereafter remained constant. The isokinetic properties of the triceps surae as reflected in the measured torque/velocity relation of the muscle in 4 subjects did not change significantly if account was taken of the slight degree of atrophy present following immobilization.(ABSTRACT TRUNCATED AT 250 WORDS)     

The influence of seat height on the mechanical function of the triceps surae muscles during steady-rate cycling

The purpose of this study was to manipulate bicycle seat height in order to perturbate muscle length, contraction velocity and excitation of soleus and medial gastrocnemius muscles. One group of female riders (n = 13) rode a stationary ergometer at 200 W and a cadence of 80 rpm. Individuals rode at a self-selected seat height, a 10% lowered and 5% raised seat position. It was hypothesized that because the two muscles would operate at decreased contraction velocities at the low seat, the integrated EMG would be less for the lowest seat position. The soleus and medial gastrocnemius muscles showed a significant decrease in integrated EMG value with decreased seat height (soleus F_2,24 = 5.4, p < 0.01, gastrocnemius F_2,24 = 51.6, p < 0.0001). The combined effect of the movement at the ankle and knee joints resulted in increased length of gastrocnemius rather than shortening at the lowered seat-height position as anticipated. This suggested that there was a greater role of knee-joint angle in determining the muscle excitation for medial gastrocnemius. The original hypothesis was accepted, confirming the importance of setting proper seat height.     

Peak triceps surae muscle activity is not specific to knee flexion angles during MVIC

There is limited research on peak activity of the separate triceps surae muscles in select knee flexion (KF) positions during a maximum voluntary isometric contraction (MVIC) used to normalize EMG signals. The aim of this study was to determine how frequent peak activity occurred during an MVIC for soleus (SOL), gastrocnemius medialis (GM), and gastrocnemius lateralis (GL) in select KF positions, and if these peaks were recorded in similar KF positions. Forty-eight healthy individuals performed unilateral plantar-flexion MVIC in standing with 0°KF and 45°KF, and in sitting with 90°KF. Surface EMG of SOL, GM, and GL were collected and processed in 250 ms epochs to determine peak root-mean-square amplitude. Peak activity was most frequently captured in standing and rarely in sitting, with no position selective to SOL, GM or GL activity. Peak GM and GL activity was more frequent in 0°KF than 45°KF, and more often in similar KF positions than not. Peak SOL activity was just as likely in 45°KF as 0°KF, and more in positions similar to GM, but not GL. The EMG amplitudes were at least 20% greater in positions that captured peak activity over those that did not. The overall findings support performing an MVIC in more than one KF position to normalize triceps surae EMG. It is emphasized that no KF position is selective to SOL, GM, or GL alone.     

Mechanical and morphological properties of the triceps surae muscle-tendon unit in old and young adults and their interaction with a submaximal fatiguing contraction.

The purposes of this study were to examine (a) whether the morphological properties of the muscle gastrocnemius medialis (GM) contribute to the known enhanced muscle fatigue resistance during submaximal sustained isometric plantar flexion contraction of old compared to young adults and (b) whether a submaximal fatiguing contraction differently affects the mechanical properties of the GM tendon and aponeurosis of old and young adults. Fourteen old and 12 young male subjects performed maximal voluntary isometric plantar flexions (MVC) on a dynamometer before and after a submaximal fatiguing task (40% MVC). Moments and EMG signals from the gastrocnemius medialis and lateralis, soleus and tibialis anterior muscles were measured. The elongation of the GM tendon and aponeurosis and the morphological properties of its contractile element were examined by means of ultrasonography. The old adults showed lower maximal ankle joint moment, stiffness and fascicle length in both tested conditions. The submaximal fatiguing contraction did not affect the force-strain relationship of the GM tendon and aponeurosis of either young or old adults. The time to task failure was longer for the old adults and was strongly correlated with the fascicle length (r(2)=0.50, P<0.001). This provides evidence on that the lower ratio of the active muscle volume to muscle force for the old adults might be an additional mechanism contributing to the known age related increase in muscle fatigue resistance.     

Muscle activation and the isokinetic torque-velocity relationship of the human triceps surae

The influence of muscle activation and the time allowed for torque generation on the angle-specific torque-velocity relationship of the triceps surae was studied during plantar flexion using supramaximal electrical stimulation and a release technique on six male subjects [mean (SD) age 25 (4) years]. Torque-velocity data were obtained under different levels of constant muscle activation by varying the stimulus frequency and the time allowed for isometric torque generation prior to release and isokinetic shortening. To eliminate the effects of the frequency response on absolute torque the isokinetic data were normalized to the maximum isometric torque values at 0.44 rad. There were no significant differences in the normalized torques generated at any angular velocity using stimulus frequencies of 20, 50 or 80 Hz. When the muscle was stimulated at 50 Hz the torques obtained after a 400 ms and 1 s pre-release isometric contraction did not differ significantly. However, with no pre-release contraction significantly less torque was generated at all angular velocities beyond 1.05 rad · s^–1 when compared with either the 200, 400 ms or 1 s condition. With a 200 ms pre-release contraction significantly less torque was generated at angular velocities beyond 1.05 rad · s^–1 when compared with the 400 ms or 1 s conditions. It would seem that the major factor governing the shape of the torque-velocity curve at a constant level of muscle activation is the time allowed for torque generation.     

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)