Comparison of elasticity of human tendon and aponeurosis in knee extensors and ankle plantar flexors in vivo

The purposes of this study were to compare the elasticity of tendon and aponeurosis in human knee extensors and ankle plantar flexors in vivo and to examine whether the maximal strain of tendon was correlated to that of aponeurosis. The elongation of tendon and aponeurosis during isometric knee extension (n = 23) and ankle plantar flexion (n = 22), respectively, were determined using a real-time ultrasonic apparatus, while the participants performed ramp isometric contractions up to voluntary maximum. To calculate the strain values from the measured elongation, we measured the respective length of tendon and aponeurosis. For the knee extensors, the maximal strain of aponeurosis (12.1 +/- 2.8 %) was significantly greater than that of the patella tendon (8.3 +/- 2.4 %), p < 0.001. On the contrary, the maximal strain of Achilles tendon (5.9 +/- 1.4 %) was significantly greater than that of aponeurosis in ankle plantar flexors (2.7 +/- 1.4 %), p < 0.001. Furthermore, for both knee extensors and ankle plantar flexors there was no significant correlation between maximal strain of tendon and aponeurosis. These results would be important for understanding the different roles of tendon and aponeurosis during human movements and for more accurate muscle modeling.     

Superficial aponeurosis of human gastrocnemius is elongated during contraction: implications for modeling muscle-tendon unit.

Two questions were addressed in this study: (1) how much strain of the superficial aponeurosis of the human medial gastrocnemius muscle (MG) was obtained during voluntary isometric contractions in vivo, (2) whether there existed inhomogeneity of the strain along the superficial aponeurosis. Seven male subjects, whose knees were extended and ankles were flexed at right angle, performed isometric plantar flexion while elongation of superficial aponeurosis of MG was determined from the movements of the intersections made by the superficial aponeurosis and fascicles using ultrasonography. The strain of the superficial aponeurosis at the maximum voluntary contraction, estimated from the elongation and length data, was 5.6+/-1.2%. There was no significant difference in strain between the proximal and distal parts of the superficial aponeurosis. Based on the present result and that of our previous study for the same subjects (J. Appl. Physiol 90 (2001) 1671), a model was formulated for a contracting uni-pennate muscle-tendon unit. This model, which could be applied to isometric contractions at other angles and therefore of wide use, showed that similar strain between superficial and deep aponeuroses of MG contributed to homogeneous fascicle length change within MG during contractions. These findings would contribute to clarifying the functions of the superficial aponeurosis and the effects of the superficial aponeurosis elongation on the whole muscle behavior.     

Elastic properties of human Achilles tendon are correlated to muscle strength.

The purpose of this study was to investigate whether the mechanical properties of the Achilles tendon were correlated to muscle strength in the triceps surae in humans. Twenty-four men and twelve women exerted maximal voluntary isometric plantar flexion (MVIP) torque. The elongation (DeltaX) and strain of the Achilles tendon (epsilon), the proximal part of which is the composite of the gastrocnemius tendon and the soleus aponeurosis, at MVIP were determined from the displacement of the distal myotendinous junction of the medial gastrocnemius using ultrasonography. The Achilles tendon force at MVIP (F) was calculated from the MVIP torque and the Achilles tendon moment arm. There were no significant differences in either the F-DeltaX or F-epsilon relationships between men and women. DeltaX and epsilon were 9.8 +/- 2.6 mm and 5.3 +/- 1.6%, respectively, and were positively correlated to F (r = 0.39, P < 0.05; r = 0.39, P < 0.05), which meant that subjects with greater muscle strength could store more elastic energy in the tendon. The regression y-intercepts for the F-DeltaX (P < 0.01) and F-epsilon (P < 0.05) relationship were significantly positive. These results might indicate that the Achilles tendon was stiffer in subjects with greater muscle strength, which may play a role in reducing the probability of tendon strain injuries. It was suggested that the Achilles tendon of subjects with greater muscle strength did not impair the potential for storing elastic energy in tendons and may be able to deliver the greater force supplied from a stronger muscle more efficiently. Furthermore, the difference in the Achilles tendon mechanical properties between men and women seemed to be correlated to the difference in muscle strength rather than gender.     

Influence of static stretching on viscoelastic properties of human tendon structures in vivo.

The purpose of this study was to investigate the influences of static stretching on the viscoelastic properties of human tendon structures in vivo. Seven male subjects performed static stretching in which the ankle was passively flexed to 35 degrees of dorsiflexion and remained stationary for 10 min. Before and after the stretching, the elongation of the tendon and aponeurosis of medial gastrocnemius muscle (MG) was directly measured by ultrasonography while the subjects performed ramp isometric plantar flexion up to the maximum voluntary contraction (MVC), followed by a ramp relaxation. The relationship between the estimated muscle force (Fm) of MG and tendon elongation (L) during the ascending phase was fitted to a linear regression, the slope of which was defined as stiffness of the tendon structures. The percentage of the area within the Fm-L loop to the area beneath the curve during the ascending phase was calculated as an index representing hysteresis. Stretching produced no significant change in MVC but significantly decreased stiffness and hysteresis from 22.9 +/- 5.8 to 20.6 +/- 4.6 N/mm and from 20.6 +/- 8.8 to 13.5 +/- 7.6%, respectively. The present results suggest that stretching decreased the viscosity of tendon structures but increased the elasticity.     

Soleus aponeurosis strain distribution following chronic unloading in humans: an in vivo MR phase-contrast study.

The in vivo strain properties of human skeletal muscle-tendon complexes are poorly understood, particularly following chronic periods of reduced load bearing. We studied eight healthy volunteers who underwent 4 wk of unilateral lower limb suspension (ULLS) to induce chronic unloading. Before and after the ULLS, maximum isometric ankle plantar flexion torque was determined by using a magnetic resonance (MR)-compatible dynamometry. Volumes of the triceps surae muscles and strain distribution of the soleus aponeurosis and the Achilles tendon at a constant submaximal plantar flexion (20% pre-maximal voluntary contraction) were measured by using MRI and velocity-encoded, phase-contrast MRI techniques. Following ULLS, volumes of the soleus and the medial gastrocnemius and the maximum isometric ankle plantar flexion (maximum voluntary contraction) decreased by 5.5+/-1.9, 7.5+/-2.7, and 48.1+/-6.1%, respectively. The strain of the aponeurosis along the length of the muscle before the ULLS was 0.3+/-0.3%, ranging from -1.5 to 2.7% in different locations of the aponeurosis. Following ULLS, the mean strain was -6.4+/-0.3%, ranging from -1.6 to 1.3%. The strain distribution of the midregion of the aponeurosis was significantly influenced by the ULLS, whereas the more distal component showed no consistent changes. Achilles tendon strain was not affected by the ULLS. These results raise the issue as to whether these changes in strain distribution affect the functional properties of the triceps surae and whether the probability of strain injuries within the triceps surae increases following chronic unloading in those regions of this muscle complex in which unusual strains occur.     

Differential displacement of the human soleus and medial gastrocnemius aponeuroses during isometric plantar flexor contractions in vivo.

The human triceps surae muscle-tendon complex is a unique structure with three separate muscle compartments that merge via their aponeuroses into the Achilles tendon. The mechanical function and properties of these structures during muscular contraction are not well understood. The purpose of the study was to investigate the extent to which differential displacement occurs between the aponeuroses of the medial gastrocnemius (MG) and soleus (Sol) muscles during plantar flexion. Eight subjects (mean +/- SD; age 30 +/- 7 yr, body mass 76.8 +/- 5.5 kg, height 1.83 +/- 0.06 m) performed maximal isometric ramp contractions with the plantar flexor muscles. The experiment was performed in two positions: position 1, in which the knee joint was maximally extended, and position 2, in which the knee joint was maximally flexed (125 degrees ). Plantarflexion moment was assessed with a strain gauge load cell, and the corresponding displacement of the MG and Sol aponeuroses was measured by ultrasonography. Differential shear displacement of the aponeurosis was quantified by subtracting displacement of Sol from that of MG. Maximal plantar flexion moment was 36% greater in position 1 than in position 2 (132 +/- 20 vs. 97 +/- 11 N.m). In position 1, the displacement of the MG aponeurosis at maximal force exceeded that of the Sol (12.6 +/- 1.7 vs. 8.9 +/- 1.5 mm), whereas in position 2 displacement of the Sol was greater than displacement of the MG (9.6 +/- 1.0 vs. 7.9 +/- 1.2 mm). The amount and "direction" of shear between the aponeuroses differed significantly between the two positions across the entire range of contraction, indicating that the Achilles tendon may be exposed to intratendinous shear and stress gradients during human locomotion.     

In vivo human tendinous tissue stretch upon maximum muscle force generation

In the present study, we examined the hypothesis that stretch of tendinous tissue in the human tibialis anterior (TA) muscle-tendon unit upon isometric dorsiflexion maximum voluntary contraction (MVC) varies along the entire tendinous component length. Ultrasound-based measurements of the excursions of the TA tendon origin and proximal end of the TA central aponeurosis were taken in the transition from rest to MVC in six men. Subtracting the TA tendon origin excursion from the excursion of the aponeurosis proximal end, the aponeurosis excursion was estimated. Estimation of the aponeurosis proximal region excursion was obtained subtracting the excursion of the insertion point of a central region fascicle on the aponeurosis from the whole aponeurosis excursion. Subtracting tendon excursion from the excursion of the central fascicle insertion point, the aponeurosis distal region excursion was estimated. Strain values were calculated dividing the excursions obtained by the original resting lengths. All excursions and lengths were measured in the mid-longitudinal axis of the TA muscle-tendon unit at the neutral anatomical ankle position. Tendon excursion and strain were 0.5+/-0. 08 cm (mean+/-SE) and 3.1+/-0.2%, respectively. Aponeurosis excursion and strain were 1.1+/-0.15 cm and 6.5+/-0.6%, respectively. Aponeurosis distal region excursion and strain were 0.3+/-0.05 cm and 3.5+/-0.3%, respectively. Aponeurosis proximal region excursion and strain were 0.8+/-0.12 cm and 9.2+/-1%, respectively. Aponeurosis excursion and strain were larger by 110-120% (P<0.05) compared with tendon. Aponeurosis proximal region excursion and strain were larger by 165-170% (P<0.05) compared with aponeurosis distal region. These findings are in line with results from in vitro animal material testing and have important implications for theoretical models of muscle function.     

Effect of joint rotation correction when measuring elongation of the gastrocnemius medialis tendon and aponeurosis

It is well known that during maximal plantar flexion contractions the ankle joint rotation overestimates the actual elongation of the tendon and aponeurosis. The aim of this study was to examine the influence of the curve length changes of the Achilles tendon on the joint rotation corrected elongation and strain of the gastrocnemius medialis (GM) tendon and aponeurosis. Nine subjects (age: 29.4 ± 5.7 years, body mass: 78.8 ± 6.8 kg, body height: 178 ± 4 cm) participated in the study. The subjects performed maximal voluntary isometric plantarflexion contractions in the prone position on a Biodex-dynamometer. Ultrasonography (Aloka SSD 4000) was used to visualize the muscle belly of the GM muscle-tendon unit. To calculate the curve length changes of the Achilles tendon its surface contour was reconstructed using a series of small reflective skin markers having a diameter of 2.5 mm. The elongation of the GM tendon and aponeurosis was calculated (a) as the difference of the measured and the passive (due to joint rotation) displacement of the tendon and aponeurosis and (b) as the difference of the measured displacement and the length changes of the reconstructed Achilles tendon surface contour. The absolute difference between the elongation obtained by both methods were 1.2 ± 0.4 mm. These differences were due to the higher changes in length obtained by the reconstruction of the tendon curved surface contour as compared to the changes observed in the passive displacement of the digitised point at the aponeurosis. Without correcting for angle joint rotation, the measured elongation clearly overestimates the actual elongation of the GM tendon and aponeurosis. After the passive displacement correction the calculated elongation still overestimates the actual elongation of the GM tendon and aponeurosis. However, this overestimation has a negligible effect on the examined in vivo strain (0.3%) of the tendon and aponeurosis.     

Nonuniform strain of human soleus aponeurosis-tendon complex during submaximal voluntary contractions in vivo.

The distribution of strain along the soleus aponeurosis tendon was examined during voluntary contractions in vivo. Eight subjects performed cyclic isometric contractions (20 and 40% of maximal voluntary contraction). Displacement and strain in the apparent Achilles tendon and in the aponeurosis were calculated from cine phase-contrast magnetic resonance images acquired with a field of view of 32 cm. The apparent Achilles tendon lengthened 2.8 and 4.7% in 20 and 40% maximal voluntary contraction, respectively. The midregion of the aponeurosis, below the gastrocnemius insertion, lengthened 1.2 and 2.2%, but the distal aponeurosis shortened 2.1 and 2.5%, respectively. There was considerable variation in the three-dimensional anatomy of the aponeurosis and muscle-tendon junction. We suggest that the nonuniformity in aponeurosis strain within an individual was due to the presence of active and passive motor units along the length of the muscle, causing variable force along the measurement site. Force transmission along intrasoleus connective tissue may also be a significant source of nonuniform strain in the aponeurosis.     

Strain and elongation of the human gastrocnemius tendon and aponeurosis during maximal plantarflexion effort

Regarding the strain and elongation distribution along the tendon and aponeurosis the literature is reporting different findings. Therefore, the purpose of this study was to examine in vivo the elongation and the strain of the human gastrocnemius medialis tendon and aponeurosis simultaneously at the same trial during maximal voluntary plantarflexion efforts. Twelve subjects participated in the study. The subjects performed isometric maximal voluntary contractions of their left leg on a Biodex-dynamometer. The kinematics of the leg were recorded using the Vicon 624 system with 8 cameras operating at 120 Hz. Two ultrasound probes were used to visualise the tendon (myotendinous junction region) and the distal aponeurosis of the gastrocnemius medialis respectively. The main findings were: (a) the absolute elongation of the gastrocnemius medialis tendon was different to that of the aponeurosis, (b) the strain of the gastrocnemius medialis tendon did not differ from the strain of the aponeurosis, (c) during the "isometric" plantarflexion the ankle angle exhibited significant changes, and (d) the non-rigidity of the dynamometer arm-foot system and the coactivity of the tibialis anterior both have a significant influence on the moment exerted at the ankle joint. Thus the strain of the human gastrocnemius medialis tendon and aponeurosis estimated in vivo using two-dimensional ultrasonography is uniform. To calculate the elongation of the whole tendon it is necessary to multiply the strain calculated for the examined part of the tendon by the total length of the tendon.     

Low-intensity tensile loading increases intratendinous glucose uptake in the Achilles tendon.

The metabolic activity of tendinous tissues has traditionally been considered to be of limited magnitude. However, recent studies have suggested that glucose uptake increases in the force-transmitting tissues as a response to contractile loading, which in turn indicates an elevated tissue metabolism. The purpose of the present study was to investigate whether such a mechanism could be observed for the human Achilles tendon following tensile loading. Six subjects participated in the study. Unilateral Achilles tendon loading was applied by 25-min intermittent voluntary plantar flexor contractions. A radioactive tracer ([18F]-2-fluoro-2-deoxy-D-glucose) was administered during muscle action, and glucose uptake was measured by use of PET. Regions of interest were defined on the PET images corresponding to the cross section of Achilles tendon at two longitudinally separated sites (insertion and free tendon). Glucose uptake index was determined within respective regions of interest for the active and resting leg. Tendon force during voluntary contractions was approximately 13% of maximal voluntary contraction force. Tendon loading induced an elevated glucose uptake index compared with that of the contralateral resting tendon in the region of tendon insertion (0.13 +/- 0.05 vs. 0.09 +/- 0.02; P < 0.05) and at the free tendon (0.12 +/- 0.01 vs. 0.08 +/- 0.02; P < 0.05). The present data suggest that tissue metabolism is elevated in the human Achilles tendon in response to low-intensity loading.     

Dynamic viscoelastic behavior of lower extremity tendons during simulated running.

The aim of this project was to see whether the tendon would show creep during long-term dynamic loading (here referred to as dynamic creep). Pig tendons were loaded by a material-testing machine with a human Achilles tendon force profile (1.37 Hz, 3% strain, 1,600 cycles), which was obtained in an earlier in vivo experiment during running. All the pig tendons showed some dynamic creep during cyclic loading (between 0.23 +/- 0.15 and 0.42 +/- 0.21%, means +/- SD). The pig tendon data were used as an input of a model to predict dynamic creep in the human Achilles tendon during running of a marathon and to evaluate whether there might consequently be an influence on group Ia afferent-mediated length and velocity feedback from muscle spindles. The predicted dynamic creep in the Achilles tendon was considered to be too small to have a significant influence on the length and velocity feedback from soleus during running. In spite of the characteristic nonlinear viscoelastic behavior of tendons, our results demonstrate that these properties have a minor effect on the ability of tendons to act as predictable, stable, and elastic force transmitters during long-term cyclic loading.     

Effect of muscle fatigue on the compliance of the gastrocnemius medialis tendon and aponeurosis

The aim of the present study was to examine whether or not the compliance of the gastrocnemius medialis (GM) tendon and aponeurosis is influenced by submaximal fatiguing efforts. Fourteen elderly male subjects performed isometric maximal voluntary plantarflexion contractions (MVC) on a dynamometer before and after two fatiguing protocols. The protocols consisted of: (1) submaximal concentric isokinetic contractions (70% isokinetic MVC) at 60 degrees /s and (2) a sustained isometric contraction (40% isometric MVC) until failure to hold the defined moment. Ultrasonography was used to determine the elongation and strain of the GM tendon and aponeurosis. To account for the axis misalignment between ankle and dynamometer, the kinematics of the leg were captured at 120 Hz. The maximum moment decreased from 85.9+/-17.9 Nm prior fatigue to 79.2+/-19 Nm after isokinetic fatigue and to 69.9+/-16.4 Nm after isometric fatigue. The maximal strain of the GM tendon and aponeurosis before fatigue, after isokinetic and after isometric fatigue were 4.9+/-1.1%, 4.4+/-1.1% and 4.3+/-1.1% respectively. Neither the strain nor the elongation showed significant differences before and after each fatiguing task at any 100 N step of the calculated tendon force. This implies that the compliance was not altered after either the isokinetic or the isometric fatiguing task. Therefore it was concluded that the strains during the performed submaximal fatiguing tasks, were too small to provoke any structural changes in tendon and aponeurosis.     

Effect of stretching training on the viscoelastic properties of human tendon structures in vivo.

The purpose of this study was to examine whether stretching training altered the viscoelastic properties of human tendon structures in vivo. Eight men performed the stretching training for 3 wk. Before and after the stretching training, the elongation of the tendon and aponeurosis of medial gastrocnemius muscle was directly measured by ultrasonography while the subjects performed ramp isometric plantar flexion up to the voluntary maximum, followed by a ramp relaxation. The relationship between the estimated muscle force (Fm) and tendon elongation (L) during the ascending phase was fitted to a linear regression, the slope of which was defined as stiffness of tendon structures. The percentage of the area within the Fm-L loop to the area beneath the curve during ascending phase was calculated as an index representing hysteresis. To assess the flexibility, the passive torque of the plantar flexor muscles was measured during the passive stretch from 0 degrees (anatomic position) to 25 degrees of dorsiflexion with a constant velocity of 5 degrees/s. The slope of the linear portion of the passive torque-angle curve during stretching was defined as flexibility index. Flexibility index decreased significantly after stretching training (-13.4 +/- 4.6%). On the other hand, the stretching training produced no significant change in stiffness but significantly decreased hysteresis from 19.9 +/- 11.7 to 12.5 +/- 9.5%. The present results suggested that stretching training affected the viscosity of tendon structures but not the elasticity.     

Muscle-tendon interaction and elastic energy usage in human walking.

The present study was designed to explore how the interaction between the fascicles and tendinous tissues is involved in storage and utilization of elastic energy during human walking. Eight male subjects walked with a natural cadence (1.4 +/- 0.1 m/s) on a 10-m-long force plate system. In vivo techniques were employed to record the Achilles tendon force and to scan real-time fascicle lengths for two muscles (medial gastrocnemius and soleus). The results showed that tendinous tissues of both medial gastrocnemius and soleus muscles lengthened slowly throughout the single-stance phase and then recoiled rapidly close to the end of the ground contact. However, the fascicle length changes demonstrated different patterns and amplitudes between two muscles. The medial gastrocnemius fascicles were stretched during the early single-stance phase and then remained isometrically during the late-stance phase. In contrast, the soleus fascicles were lengthened until the end of the single-stance phase. These findings suggest that the elastic recoil takes place not as a spring-like bouncing but as a catapult action in natural human walking. The interaction between the muscle fascicles and tendinous tissues plays an important role in the process of release of elastic energy, although the leg muscles, which are commonly accepted as synergists, do not have similar mechanical behavior of fascicles in this catapult action.     

Biceps femoris and semitendinosus tendon/aponeurosis strain during passive and active (isometric) conditions

The purpose of this study was to quantify strain and elongation of the long head of the biceps femoris (BFlh) and the semitendinosus (ST) tendon/aponeurosis. Forty participants performed passive knee extension trials from 90° of knee flexion to full extension (0°) followed by ramp isometric contractions of the knee flexors at 0°, 45° and 90° of knee flexion. Two ultrasound probes were used to visualize the displacement of BFlh and ST tendon/aponeurosis. Three-way analysis of variance designs indicated that: (a) Tendon/aponeurosis (passive) elongation and strain were higher for the BFlh than the ST as the knee was passively extended (p < 0.05), (b) contraction at each angular position was accompanied by a smaller BFlh tendon/aponeurosis (active) strain and elongation than the ST at higher levels of effort (p < 0.05) and (c) combined (passive and active) strain was significantly higher for the BFlh than ST during ramp contraction at 0° but the opposite was observed for the 45° and 90° flexion angle tests (p < 0.05). Passive elongation of tendon/aponeurosis has an important effect on the tendon/aponeurosis behavior of the hamstrings and may contribute to a different loading of muscle fibers and tendinous tissue between BFlh and ST.     

Influence of structure on the tissue dynamics of the human soleus muscle observed in MRI studies during isometric contractions

This article investigates how the internal structure of muscle and its relationship with tendon and even skeletal structures influence the translation of muscle fiber contractions into movement of a limb. Reconstructions of the anatomy of the human soleus muscle from the Visible Human Dataset (available from the National Library of Medicine), magnetic resonance images (MRI), and cadaver studies revealed a complex 3D connective tissue structure populated with pennate muscle fibers. The posterior aponeurosis and the median septum of the soleus form the insertion of the muscle and are continuous with the Achilles tendon. The distal extremities of the pennate muscle fibers attach to these structures. The anterior aponeurosis is located intramuscularly, between the posterior aponeurosis and the median septum. It forms the origin of the muscle and contacts the proximal extremities of the soleus muscle fibers. MRI measurements of in vivo tissue velocities during isometric contractions (20% and 40% maximum voluntary contractions) revealed a similarly complex 3D distribution of tissue movements. The distribution of velocities was similar to the distribution of major connective tissue structures within the muscle. During an isometric contraction, muscle fiber contractions move the median septum and posterior aponeurosis proximally, relative to the anterior aponeurosis. The pennate arrangement of muscle fibers probably amplifies muscle fiber length changes but not sufficiently to account for the twofold difference in muscle fiber length changes relative to excursion of the calcaneus. The discrepancy may be accounted for by an additional gain mechanism operating directly on the Achilles tendon by constraining the posterior movement of the tendon, which would otherwise occur due to the increasingly posterior location of the calcaneus in plantarflexeion.     

In vivo behavior of muscle fascicles and tendinous tissues of human gastrocnemius and soleus muscles during twitch contraction.

The present study investigated the differences between the human medial gastrocnemius (MG) and soleus (SOL) muscles in length changes of muscle fascicles and tendinous tissues during twitch contraction induced by an electrical nerve stimulus. Also, the time-course characteristics of twitch torque were related with changes in the length of muscle fascicles and tendinous tissues. No significant difference was observed between MG and SOL in contraction and half relaxation times of the changes in lengths and velocities of both muscle fascicles and tendinous tissues. The time-course of changes in twitch torque was nearly identical to that of the length of muscle fascicles and tendinous tissues. It was suggested that the behavior of MG and SOL during twitch contraction is practically similar in spite of their known physiological and architectural differences, and that the time-course of twitch torque is greatly influenced by the changes in the length of muscle fascicles and tendinous tissues.     

Behavior of fascicles and tendinous structures of human gastrocnemius during vertical jumping.

Behavior of fascicles and tendinous structures of human gastrocnemius medialis (MG) was determined by use of ultrasonography in vivo during jumping. Eight male subjects jumped vertically without countermovement (squat jump, SQJ). Simultaneously, kinematics, kinetics, and electromyography from lower leg muscles were recorded during SQJ. During phase I (-350 to -100 ms before toe-off), muscle-tendon complex (MTC) length was almost constant. Fascicles, however, shortened by 26%, and tendinous structures were stretched by 6%, storing elastic energy of 4.9 J during phase I. During phase II (-100 ms to toe-off), although fascicles generated force quasi-isometrically, MTC shortened rapidly by 5.3%, releasing prestored elastic energy with a higher peak positive power than that of fascicles. Also, the compliance of tendinous structures in vivo was somewhat higher than that of external tendon used in the simulation studies. The results demonstrate that the compliance of tendinous structures, together with no yielding of muscle fibers, allows MTC to effectively generate relatively large power at a high joint angular velocity region during the last part of push-off.     

In vivo fascicle behavior of synergistic muscles in concentric and eccentric plantar flexions in humans.

Ultrasonography was used to directly measure in vivo fascicle behavior of the medial gastrocnemius (MG) and soleus (SOL) muscles while the subjects (n=6 men) performed maximal voluntary concentric and eccentric plantar flexions at 60, 120, 180 and 240 deg/s. Fascicle shortening and lengthening velocities of MG, obtained from fascicle length changes over time, were significantly higher than those of SOL at +/-120, +/-180 and +240 deg/s, possibly reflecting physiological and mechanical differences between these muscles. On the other hand, the effective fascicle shortening and lengthening velocities, defined as the velocities in the longitudinal direction of muscle belly, were not significantly different between MG and SOL. This could be due to difference in fascicle architecture and/or the existence of mechanical linkages between these muscles. Moreover, when the contribution of tendinous tissues to muscle-tendon complex length change was determined from fascicle length, pennation angle, moment arm and joint angle, it accounted for approximately 50% in both concentric and eccentric trials, but showed considerable intra-subject variations. This result quantifiably demonstrates the importance of tendinous tissues in isokinetically controlled joint movements.