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Energetics and mechanics of terrestrial locomotion. IV. Total mechanical energy changes as a function of speed and body size in birds and mammals
N. C. Heglund, M. A. Fedak, C. R. Taylor, G. A. Cavagna


This is the final paper in or series examining the link between the energetics and mechanics of terrestrial locomotion. In this paper the kinetic energy of the limbs and body relative to the centre of mass (EKE, tot of paper two) is combined with the potential plus kinetic energy of the centre of mass (ECM, tot of paper three) to obtain the total mechanical energy (excluding elastic energy) of an animal during constant average-speed locomotion. The minimum mass-specific power required of the muscles and tendons to maintain the observed oscillations in total energy, Etot/Mb, can be described by one equation: Etot/Mb = 0.478. vg 1.53 + 0.685. vg + 0.072 where Etot/Mb is in W kg-1 and vg is in m s-1. This equation is independent of body size, applying equally as well to a chipmunk or a quail as to a horse or an ostrich. In marked contrast, the metabolic energy consumed by each gram of an animal as it moves along the ground at a constant speed increases linearly with speed and is proportional to Mb-0.3. Thus, we have found that each gram of tissue of a 30 g quail or chipmunk running at 3 m s-1 consumes metabolic energy at a rate about 15 times that of a 100 kg ostrich, horse or human running at the same speed while their muscles are performing work at the same rate. Our measurements demonstrate the importance of storage and recovery of elastic energy in larger animals, but they cannot confirm or exclude the possibility of elastic storage of energy in small animals. It seems clear that the rate at which animals consume energy during locomotion cannot be explained by assuming a constant efficiency between the energy consumed and the mechanical work performed by the muscles. It is suggested that the intrinsic velocity of shortening of the active muscle motor units (which is related to the rate of cycling of the cross bridges between actin and myosin) and the rate at which the muscles are turned on and off are the most important factors in determining the metabolic cost of constant-speed locomotion. Faster motor units are recruited as animals increase speed, and equivalent muscles of small animals have faster fibres than those of larger animals. Also, the muscles are turned on and off more quickly as an animal increases speed, and at the same speed a small animal will be turning muscles on and off at a much higher rate. These suggestions are testable, and future studies should determine if they are correct.