Muscle spindle feedback differs between the soleus and gastrocnemius in humans

The Hoffmann (H) reflex and motor (M) response were studied in soleus and gastrocnemius during voluntary contraction in eight male volunteers. Aims: To determine if the strength of spindle input to the muscles is the same. To assess if the M response size changes during contraction. Results: The size of the maximum M response (M max) changed during contraction in each subject. Hence, all H reflex measurements were normalized to the M max at each level of contraction for each subject. The largest H/M max was bigger in soleus than gastrocnemius at every contraction level. The overall largest H/M max for soleus (97%) and gastrocnemius (55%) were achieved at 40 and 100% maximum voluntary contraction (MVC), respectively. Conclusion: Soleus receives greater spindle feedback than the gastrocnemius both at rest and during voluntary contraction.     

Contraction level-related modulation of corticomuscular coherence differs between the tibialis anterior and soleus muscles in humans

The sensorimotor cortex activity measured by scalp EEG shows coherence with electromyogram (EMG) activity within the 15- to 35-Hz frequency band (β-band) during weak to moderate intensity of isometric voluntary contraction. This coupling is known to change its frequency band to the 35- to 60-Hz band (γ-band) during strong contraction. This study aimed to examine whether such contraction level-related modulation of corticomuscular coupling differs between muscles with different muscle compositions and functions. In 11 healthy young adults, we quantified the coherence between EEG over the sensorimotor cortex and rectified EMG during tonic isometric voluntary contraction at 10-70% of maximal voluntary contraction of the tibialis anterior (TA) and soleus (SOL) muscles, respectively. In the TA, the EEG-EMG coherence shifted from the β-band to the γ-band with increasing contraction level. Indeed, the magnitude of β-band EEG-EMG coherence was significantly decreased, whereas that of γ-band coherence was significantly increased, when the contraction level was above 60% of maximal voluntary contraction. In contrast to the TA, the SOL showed no such frequency changes of EEG-EMG coherence with alterations in the contraction levels. In other words, the maximal peak of EEG-EMG coherence in the SOL existed within the β-band, irrespective of the contraction levels. These findings suggest that the central nervous system regulates the frequency of corticomuscular coupling to exert the desired levels of muscle force and, notably, that the applicable rhythmicity of the coupling for performing strong contractions differs between muscles, depending on the physiological muscle compositions and functions of the contracting muscle.     

Biomechanical characterization and clinical implications of artificially induced toe-walking: differences between pure soleus, pure gastrocnemius and combination of soleus and gastrocnemius contractures

The purpose of this study was to characterize biomechanically three different toe-walking gait patterns, artificially induced in six neurologically intact subjects and to compare them to selected cases of pathological toe-walking. The subjects, equipped with lightweight mechanical exoskeleton with elastic ropes attached to the left leg's heel on one end and on shank and thigh on the other end in a similar anatomical locations where soleus and gastrocnemius muscles attach to skeleton, walked at speed of approximately 1m/s along the walkway under four experimental conditions: normal walking (NW), soleus contracture emulation (SOL), gastrocnemius contracture emulation (GAS) and emulation of both soleus and gastrocnemius contractures (SOLGAS). Reflective markers and force platform data were collected and ankle, knee and hip joint angles, moments and powers were calculated using inverse dynamic model for both legs. Characteristic peaks of averaged kinematic and kinetic patterns were compared among all four experimental conditions in one-way ANOVA. In the left leg SOL contracture mainly influenced the ankle angle trajectory, while GAS and SOLGAS contractures influenced the ankle and knee angle trajectories. GAS and SOLGAS contractures significantly increased ankle moment during midstance as compared to SOL contracture and NW. All three toe-walking experimental conditions exhibited significant power absorption in the ankle during loading response, which was absent in the NW condition, while during preswing significant decrease in power absorption as compared to NW was seen. In the knee joint SOL contracture diminished, GAS contracture increased while SOLGAS contracture approximately halved knee extensor moment during midstance as compared to NW. All three toe-walking experimental conditions decreased hip range of motion, hip flexor moment and power requirements during stance phase. Main difference in the right leg kinematic and kinetic patterns was seen in the knee moment trajectory, where significant increase in the knee extensor moment took place in terminal stance for GAS and SOLGAS experimental conditions as compared to SOL and NW. The kinetic trajectories under SOL and GAS experimental conditions were qualitatively compared to two selected clinical cases showing considerable similarity. This implies that distinct differences in kinetics between SOL, GAS and SOLGAS experimental conditions, as described in this paper, may be clinically relevant in determining the relative contribution of soleus and gastrocnemius muscles contractures to toe-walking in particular pathological gait.     

Myofascial force transmission between the human soleus and gastrocnemius muscles during passive knee motion

The plantarflexors of the lower limb are often assumed to act as independent actuators, but the validity of this assumption is the subject of considerable debate. This study aims to determine the degree to which passive changes in gastrocnemius muscle length, induced by knee motion, affect the tension in the adjacent soleus muscle. A second aim is to quantify the magnitude of myofascial passive force transmission between gastrocnemius and adjacent soleus. Fifteen healthy volunteers participated. Simultaneous ultrasound images of the gastrocnemius and soleus muscles were obtained during passive knee flexion (0-90°), while keeping the ankle angle fixed at either 70° or 115°. Image correlation analysis was used to quantify muscle fascicle lengths in both muscles. The data show that the soleus muscle fascicles elongate significantly during gastrocnemius shortening. The approximate change in passive soleus force as a result of the observed change in fascicle length was estimated and appears to be <5 N, but this estimate is sensitive to the assumed slack length of soleus.     

Passive knee movement-induced modulation of the soleus H-reflex and alteration in the fascicle length of the medial gastrocnemius muscle in humans

In humans, an inhibitory via Ia afferent pathway from the medial gastrocnemius (MG) to the soleus (SOL) motoneuron pool has been suggested. Herein, we examined the relation between MG fascicle length changes and the SOL H-reflex modulation during passive knee movement. Twelve subjects performed static and passive (5° s^?1) knee movement tasks with the ankle immobilized using an isokinetic dynamometer in sitting posture. The maximal H- and M-waves were measured at four target angles (20°, 40°, 60°, and 80° flexion from full knee extension). The MG fascicles length and velocity were measured using a B-mode ultrasonic apparatus. Results demonstrated that the SOL Hmax/Mmax; i.e., ratio of the maximal H- to M-waves, was attenuated with increasing MG fascicle length in static tasks. The SOL Hmax/Mmax at 20° was significantly attenuated compared with 60° and 80° with increasing MG fascicle length and lengthening velocity in passive knee extension. However, no significant differences in the SOL Hmax/Mmax were found across the target angles in the passive knee flexion task. In conclusion, as muscle spindles increase their discharge with lengthening fascicle velocity, but keep silent when fascicles shorten, our data suggest that lengthening the MG facilitates an inhibitory Ia pathway from MG to SOL, and modulates SOL motoneuron activity during movements.     

Empirical evaluation of gastrocnemius and soleus function during walking

Distinguishing gastrocnemius and soleus muscle function is relevant for treating gait disorders in which abnormal plantarflexor activity may contribute to pathological movement patterns. Our objective was to use experimental and computational analysis to determine the influence of gastrocnemius and soleus activity on lower limb movement, and determine if anatomical variability of the gastrocnemius affected its function. Our hypothesis was that these muscles exhibit distinct functions, with the gastrocnemius inducing limb flexion and the soleus inducing limb extension. To test this hypothesis, the gastrocnemius or soleus of 20 healthy participants was electrically stimulated for brief periods (90 ms) during mid- or terminal stance of a random gait cycle. Muscle function was characterized by the induced change in sagittal pelvis, hip, knee, and ankle angles occurring during the 200 ms after stimulation onset. Results were corroborated with computational forward dynamic gait models, by perturbing gastrocnemius or soleus activity during similar portions of the gait cycle. Mid- and terminal stance gastrocnemius stimulation induced posterior pelvic tilt, hip flexion and knee flexion. Mid-stance gastrocnemius stimulation also induced ankle dorsiflexion. In contrast mid-stance soleus stimulation induced anterior pelvic tilt, knee extension and plantarflexion, while late-stance soleus stimulation induced relatively little change in motion. Model predictions of induced hip, knee, and ankle motion were generally in the same direction as those of the experiments, though the gastrocnemius? results were shown to be quite sensitive to its knee-to-ankle moment arm ratio.     

A comparison of the contractile properties of the human gastrocnemius and soleus muscles

Twitch speeds and potentiating capacities have been determined for human medial and lateral gastrocnemius and soleus muscles. The experiments involved and application of submaximal stimuli to the respective muscle bellies, with monitoring of the evoked compound action potentials (M-waves) during repetitive stimulation. Contrary to an earlier report, the lateral gastrocnemius was found to have a significantly shorter mean contraction time (100.0 +/- 10.8 ms) than the soleus (156.5 +/- 14.7 ms) and this value was also significantly different from that of the medial gastrocnemius (113.7 +/- 19.6 ms). The mean half-relaxation time for each muscle also differed significantly from those for the other two muscles. A further contrast between the muscles was that potentiation of the twitch, following a 3-s tetanus at 50 Hz, was significantly greater in the lateral gastrocnemius than in soleus (mean values 60.4 +/- 43.1% and 2.6 +/- 3.3% respectively.     

Differential structural features in soleus and gastrocnemius of carnitine-treated cancer cachectic rats

Muscle wasting is associated with chronic diseases and cancer. Elucidation of the biological mechanism involved in the process of muscle mass loss and cachexia may help identify therapeutic targets. We hypothesized that l -carnitine treatment may differentially revert muscle fiber atrophy and other structural alterations in slow- and fast-twitch limb muscles of rats bearing the Yoshida ascites hepatoma. In soleus and gastrocnemius of tumor-bearing rats (10⁸ AH-130 Yoshida ascites hepatoma cells inoculated intraperitoneally) with and without treatment with l -carnitine (1 g/kg body weight for 7 days, intragastric), food intake, body and muscle weights, fiber typing and morphometry, morphological features, redox balance, autophagy and proteolytic, and signaling markers were explored. Levels of carnitine palmitoyl transferase were also measured in all the study muscles. l - Carnitine treatment ameliorated the atrophy of both slow- and fast-twitch fibers (gastrocnemius particularly), muscle structural alterations (both muscles), and attenuated oxidative stress, proteolytic and signaling markers (gastrocnemius). Despite that carnitine palmitoyl transferase-1 levels increased in both muscle types in a similar fashion, l -carnitine ameliorated muscle atrophy and proteolysis in a muscle-specific manner in cancer-induced cachexia. These data reveal the need to study muscles of different fiber type composition and function to better understand whereby l -carnitine exerts its beneficial effects on the myofibers in muscle wasting processes. These findings also have potential clinical implications, since combinations of various exercise and muscle training modalities with l -carnitine should be specifically targeted for the muscle groups to be trained.     

Effect of vertical jumping on the medial gastrocnemius and soleus muscles of rats

Ten rats were trained to perform approximately 30 jumps/day, 5 days/wk for at least 8 wk, from a force platform that enabled the number and height of jumps to be quantified. There was considerable variation in height jumped during an activity session both within and between rats. The two highest-jumping rats attained a displacement of center of mass of approximately 30 cm, estimated to be approximately 67% of the maximum attainable. The two lowest-jumping rats jumped to approximately 30% of the estimated maximum. The activity was described as "habitual activity" rather than "training" because there were no significant increases in the height of jumping by any rat over the period of activity. The isometric properties of medial gastrocnemius (MG) and soleus muscles were studied in terminal experiments on anesthetised rats. Five significant effects on MG were evoked by this pattern of exercise ("habituation"): 1) a 15-18% increase in force at frequencies of stimulation between 60 and 150 Hz and a 15% increase in maximum tetanic tension to 14.9 N, 2) a 3% increase in the maximum rate of rise of tetanic force to 3.4% of maximum tetanic tension per millisecond, 3) an increase in fatigability expressed as a smaller fatigue index in active rats (33%) than in controls (58%), 4) a decrease of 4% in the percentage of type IIa muscle fibers, and 5) an increase of 6% in the percentage of type II fibers that could not be classified with certainty as IIa or IIb.(ABSTRACT TRUNCATED AT 250 WORDS)     

Selective activation of human soleus and medial gastrocnemius muscles during walking in water

During walking in water (WW) the vertical component of ground reaction forces decreases, while the greater propulsive force is required to move forward against the greater resistance of water. In such reduced gravity environment, Hutchison et al. (1989) have demonstrated that the relative activation of rat medial gastrocnemius (MGAS) increased compared to that of the soleus (SOL) during swimming, suggesting different effects of peripheral information on motoneuron excitability of these muscles. It is conceivable that both buoyancy and resistance of water have different effects on the activation patterns of triceps surae muscles during WW, since the reduced weight in water might decrease the peripheral inflow relating load information while greater volitional command might be needed to propel a body forward against the water resistance. The present study was designed to assess each peripheral inflow and efferent input by adjusting the load and walking speed voluntarily during WW. The aim of this study is to investigate the dissociative activation pattern between the SOL and the MGAS during WW.     

Hindlimb unweighting decreases endothelium-dependent dilation and eNOS expression in soleus not gastrocnemius.

We tested the hypothesis that hindlimb unweighting (HLU) decreases endothelium-dependent vasodilation and expression of endothelial nitric oxide synthase (eNOS) and superoxide dismutase-1 (SOD-1) in arteries of skeletal muscle with reduced blood flow during HLU. Sprague-Dawley rats (300-350 g) were exposed to HLU (n = 15) or control (n = 15) conditions for 14 days. ACh-induced dilation was assessed in muscle with reduced [soleus (Sol)] or unchanged [gastrocnemius (Gast)] blood flow during HLU. eNOS and SOD-1 expression were measured in feed arteries (FA) and in first-order (1A), second-order (2A), and third-order (3A) arterioles. Dilation to infusion of ACh in vivo was blunted in Sol but not Gast. In arteries of Sol muscle, HLU decreased eNOS mRNA and protein content. eNOS mRNA content was significantly less in Sol FA (35%), 1A arterioles (25%) and 2A arterioles (18%). eNOS protein content was less in Sol FA (64%) and 1A arterioles (65%) from HLU rats. In arteries of Gast, HLU did not decrease eNOS mRNA or protein. SOD-1 mRNA expression was less in Sol 2A arterioles (31%) and 3A arterioles (29%) of HLU rats. SOD-1 protein content was less in Sol FA (67%) but not arterioles. SOD-1 mRNA and protein content were not decreased in arteries from Gast. These data indicate that HLU decreases endothelium-dependent vasodilation, eNOS expression, and SOD-1 expression primarily in arteries of Sol muscle where blood flow is reduced during HLU.     

Recruitment of the Rhesus soleus and medial gastrocnemius before, during and after spaceflight

Electromyograms were recorded from the soleus and medial gastrocnemius muscles and tendon force from the medial gastrocnemius muscle of 2 juvenile Rhesus monkeys before, during and after Cosmos flight 2229 and of ground control animals. Recording sessions were made while the Rhesus were performing a foot pedal motor task. Preflight testing indicated normal patterns of recruitment between the soleus and medial gastrocnemius, i.e. a higher level of recruitment of the soleus compared to the medial gastrocnemius during the task. Recording began two days into the spaceflight and showed that the media gastrocnemius was recruited preferentially over the soleus. This observation persisted throughout the flight and for the 2 week period of postflight testing. These data indicate a significant change in the relative recruitment of slow and fast extensor muscles under microgravity conditions. The appearance of clonic-like activity in one muscle of each Rhesus during flight further suggests a reorganization in the neuromotor system in a microgravity environment.     

Enzymic and metabolic adaptations in the gastrocnemius, plantaris and soleus muscles of hypocaloric rats.

1. The effect of hypocaloric feeding (25% of normal food intake for 21 days) of rats on the enzymic and metabolic adaptations in the gastrocnemius, plantaris and soleus muscles was studied. 2. In control and hypocaloric rats the muscle relaxation rates at 100 Hz were 35.76 and 11.38% force loss/10 ms respectively. Control rats exhibited enhanced force of muscle contraction as the frequency of stimulation increased from 10 to 100 Hz, with maximum force being at 100 Hz. Hypocaloric rats exhibited a decrease in the increment of force being exerted at high frequencies, with maintenance of force at lower stimulatory frequencies. 3. In muscles of hypocaloric rats, there were significant decreases in the maximal activities of hexokinase (17.6-37.0%), 6-phosphofructokinase (22.7-34.2%), pyruvate kinase (21.2-36.0%), citrate synthase (34.1-41.5%), oxoglutarate dehydrogenase (29.4-52.4%) and 3-hydroxyacyl-CoA dehydrogenase (26.7-32.1%), whereas the activities of glycogen phosphorylase increased (23.8-43.4%) compared with control values. 4. In soleus-muscle strip preparations of hypocaloric rats, there were significant decreases in the rates of lactate production (28.1%) and glucose oxidation (32.6%) compared with control preparations. 5. Mitochondrial preparations from muscles of hypocaloric rats incubated with various substrates exhibited decreased rates of oxygen uptake compared with control preparations. 6. In muscles of hypocaloric rats (gastrocnemius and soleus), there were significant decreases in the concentrations of glycogen (P less than 0.001) and phosphocreatine (P less than 0.001) and increases in those of pyruvate (P less than 0.001), lactate (P less than 0.001) and ADP (P less than 0.001), whereas those of ATP and AMP remained unchanged. 7. Calculated [lactate]/[pyruvate] and [ATP]/[ADP] ratios exhibited significant increases (P less than 0.05) and decreases (P less than 0.05) in muscles of hypocaloric rats respectively. 8. The results are discussed in relation to the genesis of muscle dysfunction caused by malnutrition.     

Contractile responses of the rat gastrocnemius and soleus muscles to isotonic resistance exercise.

Male rats were divided into control and weight-trained (WT) groups. WT rats performed squat-type exercises twice daily, 5 days/wk, for 14 wk. They averaged 36 lifts/day, with an average weight of 555 g. Muscle-to-body weight ratio (mg/g) of the soleus (Sol) was not different from control, but it increased 11 and 6% in the gastrocnemius (Gast) and plantaris, respectively (P < 0.05). The normalized twitch tension of the in situ Sol was elevated by 21%, whereas single-skinned type I fibers from the Sol showed an increased rate constant of tension redevelopment (K(tr)) but no other contractile adaptations to WT. In contrast, the Gast type I fibers showed an increase (P < 0.05) in maximal velocity of shortening (25%), peak power (15%), K(tr) (18%), and normalized tension (7%). The K(tr) and normalized tension of the Gast type IIa fibers increased by 24% (P < 0.05) and 12% (P < 0.05), respectively, whereas velocity and power showed a tendency to increase. Fiber size, determined by myosin ATPase histochemistry, was not different for any fiber type from the Gast or Sol. These results indicate that isotonic resistance exercise of the calf targets the Gast (type I and type IIa fibers) and has little effect on the Sol.     

Myosin heavy chains in fibers of TTX-paralyzed rat soleus and medial gastrocnemius muscles.

The expression of five myosin heavy chain (MHC) isoforms was analyzed in the rat soleus (Sol) and the deep and superficial medial gastrocnemius (dGM, sGM) muscle after 2 and 4 wk of TTX paralysis by using immunohistochemical techniques. In Sol, after 4 wk of paralysis, fibers containing type I MHC were either pure type I (14%) or also contained developmental (D; 76%), IIa (26%), or IIx (18%) MHC. Values for corresponding fibers in dGM were 8.5, 65, 38, and 22%. Also, by 4 wk an increase was seen in the proportions of fibers expressing IIa MHC in Sol (from 16 to 38%) and dGM (from 24 to 74%). In a region of sGM in control muscles containing pure IIb fibers, a major proportion (86%) remained pure after 4 wk of paralysis, with the remainder coexpressing IIb and IIx. The results indicate that TTX-induced muscle paralysis results in an increase in fibers containing multiple MHC isoforms and that the D isoform appears in a major proportion of these hybrid fibers.     

Gastrocnemius and soleus lengths in cerebral palsy equinus gait--differences between children with and without static contracture and effects of gastrocnemius recession

Equinus gait is one of the most common abnormalities in children with cerebral palsy. Although it is generally assumed that the calf muscles are abnormally short in equinus gait, no studies have been done to confirm that the muscles are short and that this shortness contributes to the equinus. This study used musculoskeletal modeling combined with computerized gait analysis to examine medial gastrocnemius (MGAS), lateral gastrocnemius (LGAS), and soleus (SOL) musculotendinous lengths during equinus gait in children with cerebral palsy. All three muscles were abnormally short during equinus gait whether or not the children had equinus contractures (P < or = 0.005). Children with static contractures had shorter maximum static MGAS and LGAS lengths than children with dynamic equinus (P < or = 0.002). The children with static contractures had ratios of peak dynamic length to maximum static length close to 1.0 for MGAS and LGAS (1.005 +/- 0.015) but lower ratios for SOL (0.984 +/- 0.024). For the children with static contracture, these ratios did not change significantly after gastrocnemius recession (P > or = 0.14) because both static and dynamic lengths increased postoperatively (P < or = 0.04). These results support the current clinical understanding of the role of calf "tightness" in equinus gait, including the appropriateness and effectiveness of gastrocnemius recession for children with equinus contracture.     

Muscle architecture and intramuscular distribution of nerves in the human soleus muscle

On the deep anterior surface of the human soleus muscle a bipenniform part is found, the portio anterior, which is innervated by the ramus anterior. This part and the innervation pattern are morphological characteristics peculiar to man. In six human soleus muscles plus three anomalies, the intramuscular distribution of the nerves supplying these muscles was macroscopically examined. In the normal soleus muscles, the R. anterior supplied not only the portio anterior but also the portio posterior through several anastomoses with twigs of the R. posterior. In the soleus without the portio anterior, the R. anterior entered the muscle at the anterior surface and joined directly with the branches of the R. posterior. In the soleus with two bipenniform parts, one bipenniform part was supplied by the R. anterior, and the other by the R. posterior. The cross-sectional area of the total nerves supplying the muscle was 0.72 +/- 0.036 mm2 (n = 8). The areas of the R. posterior and the R. anterior were in inverse relation to each other. These findings suggest that the R. anterior became separated from the R. posterior. However, the endoperineurium fiber composition of the R. anterior appeared to consist of a few nerve fiber components different from those of the R. posterior.