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1Concord Field Station, Department of Organismic and Evolutionary Biology, Harvard University; Bedford, Massachusetts 01730; 2Department of Integrative Physiology, University of Colorado, Boulder, Colorado 80309; 3Department of Zoology, Oregon State University, Corvallis, Oregon 97331; and 4Department of Organismal Biology and Anatomy, University of Chicago, Chicago, Illinois 60637
Submitted 5 January 2004 ; accepted in final form 9 July 2004
Muscular forces generated during locomotion depend on an animal's speed, gait, and size and underlie the energy demand to power locomotion. Changes in limb posture affect muscle forces by altering the mechanical advantage of the ground reaction force (R) and therefore the effective mechanical advantage (EMA = r/R, where r is the muscle mechanical advantage) for muscle force production. We used inverse dynamics based on force plate and kinematic recordings of humans as they walked and ran at steady speeds to examine how changes in muscle EMA affect muscle force-generating requirements at these gaits. We found a 68% decrease in knee extensor EMA when humans changed gait from a walk to a run compared with an 18% increase in hip extensor EMA and a 23% increase in ankle extensor EMA. Whereas the knee joint was extended (154–176°) during much of the support phase of walking, its flexed position (134–164°) during running resulted in a 5.2-fold increase in quadriceps impulse (time-integrated force during stance) needed to support body weight on the ground. This increase was associated with a 4.9-fold increase in the ground reaction force moment about the knee. In contrast, extensor impulse decreased 37% (P < 0.05) at the hip and did not change at the ankle when subjects switched from a walk to a run. We conclude that the decrease in limb mechanical advantage (mean limb extensor EMA) and increase in knee extensor impulse during running likely contribute to the higher metabolic cost of transport in running than in walking. The low mechanical advantage in running humans may also explain previous observations of a greater metabolic cost of transport for running humans compared with trotting and galloping quadrupeds of similar size.
gait; muscle mechanical advantage; muscle force; metabolic cost
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