In the present study we investigated in vivo length changes in the fascicles and tendon of the human gastrocnemius medialis (GM) muscle during walking. The experimental protocol involved real-time ultrasound scanning of the GM muscle, recording of the electrical activity of the muscle, measurement of knee-and ankle-joint rotations, and measurement of ground reaction forces in six men during walking at 3 km h À1 on a treadmill. Fascicular lengths were measured from the sonographs recorded. Musculotendon complex length changes were estimated from anatomical and joint kinematic data. Tendon length changes were obtained combining the musculotendon complex and fascicular length-change data. The fascicles followed a di¡erent length-change pattern from those of the musculotendon complex and tendon throughout the step cycle. Two important features emerged: (i) the muscle contracted near-isometrically in the stance phase, with the fascicles operating at ca. 50 mm; and (ii) the tendon stretched by ca. 7 mm during single support, and recoiled in push-o¡. The behaviour of the muscle in our experiment indicates consumption of minimal metabolic energy for eliciting the contractile forces required to support and displace the body. On the other hand, the spring-like behaviour of the tendon indicates storage and release of elastic-strain energy. Either of the two mechanisms would favour locomotor economy.

Architectural properties of the triceps surae muscles were determined in vivo for six men. The ankle was positioned at 15 degrees dorsiflexion (-15 degrees) and 0, 15, and 30 degrees plantar flexion, with the knee set at 0, 45, and 90 degrees. At each position, longitudinal ultrasonic images of the medial (MG) and lateral (LG) gastrocnemius and soleus (Sol) muscles were obtained while the subject was relaxed (passive) and performed maximal isometric plantar flexion (active), from which fascicle lengths and angles with respect to the aponeuroses were determined. In the passive condition, fascicle lengths changed from 59, 65, and 43 mm (knee, 0 degrees; ankle, -15 degrees) to 32, 41, and 30 mm (knee, 90 degrees ankle, 30 degrees) for MG, LG, and Sol, respectively. Fascicle shortening by contraction was more pronounced at longer fascicle lengths. MG had greatest fascicle angles, ranging from 22 to 67 degrees, and was in a very disadvantageous condition when the knee was flexed at 90 degrees, irrespective of ankle positions. Different lengths and angles of fascicles, and their changes by contraction, might be related to differences in force-producing capabilities of the muscles and elastic characteristics of tendons and aponeuroses.

Muscle force (MF) is linearly related to physiological cross-sectional area (CSA), which is obtained from muscle volume (MV) divided by fibre length. Taking into account the fact that joint torque (TQ) is determined by MF multiplied by the moment arm, the maximal TQ would be a function of MV. This proposition was tested in the present study by investigating the relationship between MV and TQ for elbow flexor (EF) and extensor (EE) muscles of 26 males. The MVs of EF and EE were determined from a series of muscle CSA by magnetic resonance imaging (MRI), and pennation angle (theta) and FL by ultrasonography (US). Maximal isometric TQ was measured at right angle of elbow joint for EF and EE. There was a highly significant correlation between MV and TQ both for EF and EE (r=0.95 and 0.96 respectively) compared with that between muscle CSA and TQ, suggesting the dependence of TQ on MV. Furthermore, prediction equations for MV (MVULT) from muscle thickness (MT) measured by US was developed with reference to MVMRI by the MRI on 26 subjects, and the equations were applied to estimate MV of healthy university students (CON; 160 males) and sports athletes (ATH; 99 males). There were significant linear relationships between MVULT and TQ both for EF (r=0.783) and EE (r=0.695) for all subjects (n=259). The MVULT was significantly higher in ATH (by 32% for EF and 33% for EE, respectively) than in CON. Similarly, significantly greater TQ was observed in ATH (by 35% for EF, 37% for EE, respectively). The theta for EE showed no difference between both groups (17.8 degrees for CON and 17.5 degrees for ATH). On the other hand, the TQ to MV ratio were identical for CON and ATH. The results reveal that the muscle volume of the upper arm is a major determinant of joint torque (TQ), regardless of athletic training.

Muscle-fiber pennation angles were measured in vivo with the use of ultrasonography to investigate the relationship between fiber pennation and muscle size for 32 male subjects (from untrained subjects to highly trained bodybuilders). From the image of a B-mode ultrasonogram, fiber pennation angles and thickness of triceps brachii were determined, the former as angles between echoes from the interspaces of fascicles and from the aponeurosis of long and medial heads of triceps and the latter as the distance between the fat-muscle and muscle-bone interfaces. The pennation angles were in the range of 15 and 53 degrees for the long head and 9 to 26 degrees for the medial head, which were similar to or greater than the published and the present data on human cadavers. Significant differences were observed between normal subjects and bodybuilders in muscle thickness and pennation angles (P < 0.01), and there were significant correlations between muscle thickness and pennation angles for both long (r = 0.884) and medial (r = 0.833) heads of triceps, suggesting that muscle hypertrophy involves an increase in fiber pennation angles.

We have developed a technique to determine fascicle length in human vastus lateralis muscle in vivo by using ultrasonography. When the subjects had the knee fully extended passively from a position of 110 degree flexion (relaxed condition), the fascicle length decreased from 133 to 97 mm on average. During static contractions at 10% of maximal voluntary contraction strength (tensed condition), fascicle shortening was more pronounced (from 126 to 67 mm), especially when the knee was closer to full extension. Similarly, as the knee was extended, the angle of pennation (fascicle angle, defined as the angle between fascicles and aponeurosis) increased (relaxed, from 14 to 18 degrees; tensed, from 14 to 21 degrees), and a greater increase in the pennation angle was observed in the tensed than in the relaxed condition when the knee was close to extension (< 40 degrees). We conclude that there are differences in fascicle lengths and pennation angles when the muscle is in a relaxed and isometrically tensed conditions and that the differences are affected by joint angles, at least at the submaximal contraction level.

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.

The purpose of this study was to quantify the elastic properties of tendon structures in vivo and to investigate the influence of the tendon properties on jump performance with and without countermovement. Elongation of the tendon and aponeurosis of vastus lateralis muscle (dL) was directly measured by ultrasonography while subjects (n = 31) performed ramp isometric knee extension up to the voluntary maximum (MVC). The relationship between muscle force and dL was fitted to a linear regression above 50% MVC, the slope of which was defined as stiffness of the tendon structures. Statistical analysis revealed no significant difference between duplicated measurements of stiffness, with an interday reliability of r = 0.88 and a coefficient of variance of 6.1%. Although the stiffness was not significantly related to absolute jump height in either vertical jump, it was inversely correlated with the difference in jump height between the vertical jumps performed with and without countermovement. The results suggested that the stiffness of tendon structures has a favorable effect on stretch-shortening cycle exercise, possibly due to adequate storage and recoil of elastic energy.

Five men underwent unilateral resistance training of elbow extensor (triceps brachii) muscles for 16 weeks. Before and after training, muscle layer thickness and fascicle angles of the long head of the triceps muscle were measured in vivo using B-mode ultrasound, and fascicle lengths were estimated. Series anatomical cross-sectional areas (ACSA) of the triceps brachii muscle were measured by magnetic resonance imaging, from which muscle volume (Vm) was determined and physiological cross-sectional area (PCSA) was calculated. Elbow extension strength (isometric; concentric and eccentric at 30, 90 and 180 degrees.s-1) was measured using an isokinetic dynamometer to determine specific tension. Muscle volumes, ACSA, PCSA, muscle layer thickness and fascicle angles increased after training and their relative changes were similar, while muscle and fascicle length did not change. Muscle strength increased at all velocities; however, specific tension decreased after training. Increase in fascicle angles, which would be the result of increased Vm and PCSA, would seem to imply the occurrence of changes in muscle architecture. This might have given a negative effect on the force-generating properties of the muscles.