Energetics of bipedal running. II. Limb design and running mechanics.

Compared with quadrupeds, bipedal runners of the same weight have longer legs, take longer steps and can presumably use slower, more economical muscle fibers. One might predict that bipedal running is less expensive, but it is not. We hypothesized that bipeds recruit a larger volume of muscle to support their weight, eliminating the potential economy of longer legs and slower steps. To test our hypothesis, we calculated the relative volume of muscle needed to support body weight over a stride in small dogs (Canis familiaris) and wild turkeys (Meleagris gallopavo) of the same weight. First, we confirmed that turkeys and dogs use approximately the same amount of energy to run at the same speed, and found that turkeys take 1. 8-fold longer steps. Higher muscle forces and/or longer muscle fibers would require a greater volume of active muscle, since muscle volume is proportional to the product of force and fascicle length. We measured both mean fascicle length and mean mechanical advantage for limb extensor muscles. Turkeys generated approximately the same total muscle force to support their weight during running and used muscle fascicles that are on average 2.1 times as long as in dogs, thus requiring a 2.5-fold greater active muscle volume. The greater volume appears to offset the economy of slower rates of force generation, supporting our hypothesis and providing a simple explanation for why it costs the same to run on two and four legs.