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First published online March 14, 2005
Journal of Experimental Biology 208, 1175-1190 (2005)
Published by The Company of Biologists 2005
doi: 10.1242/jeb.01486
In vivo muscle function vs speed I. Muscle strain in relation to length change of the muscle-tendon unit
1 Biological Sciences Department, California State Polytechnic University,
Pomona, CA 91768-4032, USA
2 Equine Research Center, California State Polytechnic University, Pomona,
CA 91768-4032, USA
3 Concord Field Station, Harvard University, Bedford, MA 01730,
USA
* Author for correspondence (e-mail: dfhoyt{at}csupomona.edu)
Accepted 22 December 2004
The activity of muscles can be concentric (shortening), eccentric (lengthening) or isometric (constant length). When studying muscle function it is important to know what the muscle fascicles are actually doing because the performance of muscle is strongly influenced by the type of activity: force decreases as a function of shortening velocity during concentric contractions; force produced during eccentric contractions can be stronger than maximum isometric force, and force production is enhanced if a concentric contraction follows an eccentric phase. It is well known that length changes of muscle fascicles may be different from length changes of the overall muscle-tendon unit because of the compliance of the series elasticity. Consequently, fascicles of joint extensor muscles may not undergo eccentric activity even when the joint flexes, but the extent to which this occurs may vary with the compliance of the series elasticity and may differ between species: the vastus lateralis, a knee extensor, shortens when active during trotting in dogs and lengthens in rats. Previous studies of kinematics of trotting in horses have shown that during stance, the elbow extends nearly continuously with a brief period of flexion near mid-stance and the knee exhibits two phases of flexion followed by extension. The lateral triceps (an elbow extensor) has no external tendon but the vastus lateralis has a relatively long external tendon and the fascicles insert on an aponeurosis. Thus, one might expect the relation between fascicle strain and overall length change of the muscle-tendon units to be quite different in these two muscles. In the present study in horses, fascicle length changes of the lateral triceps and vastus lateralis were measured with sonomicrometry and length changes of the muscle-tendon units were estimated from muscle architecture and joint kinematics for four horses trotting on a treadmill at nine speeds. Because the focus of this study was the relation between length changes of the muscle-tendon unit (estimated from kinematics) and length changes in the muscle fascicles, we divided the stance-phase sonomicrometry records into phases that corresponded to the alternating flexion and extension of the joint as indicated by the kinematic records. During its one eccentric phase, the triceps shortened by 0.7±0.4% despite a predicted lengthening of 1%. Similarly, the vastus shortened by 3.7±1.9% when kinematics predicted 3.2% lengthening. During their concentric phases the triceps shortened by 10.6% and the vastus shortened by 8.1%. Strain in the triceps did not change with speed but it did in the vastus. Strain rate increased with speed in both muscles as did the integrated EMG, indicating an increase in the volume of muscle recruited. Thus, despite differences in their architecture and the kinematic patterns of the associated joints, these two joint extensors exhibited similar activity.
Key words: Locomotion, quadruped, sonomicrometry, muscle
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