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.