Scaling of muscle fibres and locomotion

To reconcile the scaling of the mechanics and energetics of locomotion to recent data on the scaling of the mechanics of muscle fibres, I have extended the theory of Taylor and colleagues that the energetic cost of locomotion is determined by the cost of generating force by the fibres. By assuming (1) that the cost of generating force in a fibre is proportional to V(max) (maximum velocity of shortening) and (2) that, at physiologically equivalent speeds, animals of different body sizes recruit the same fibre types, this extension quantitatively predicts the scaling of the energetics of locomotion, as well as other observations, from the scaling of V(max) of the muscle fibres. First, the energetic cost of locomotion at physiologically equivalent speeds scales with Mb-0.16, where Mb is body mass, as does V(max) of a given fibre type. However, the energetic cost at absolute speeds (cost of transport) scales with Mb-0.30, because small animals must compress their recruitment order into a narrower speed range and, hence, recruit faster muscle fibre types at a given running speed. Thus, it costs more for small animals to move 1 kg of their body mass 1 km not only because a given muscle fibre type from a small animal costs more to generate force than from a large one, but also because small animals recruit faster fibre types at a given absolute running speed. In addition, this analysis provides evidence that V(max) scales similarly to 1/tc (where tc is foot contact time) and muscle shortening velocity (V), in agreement with recent models. Finally, this extension predicts that, at physiologically equivalent speeds, the weight-specific energetic cost per step is independent of body size, as has been found empirically.