Effects of limb mass distribution on mechanical power outputs during quadrupedalism

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Abstract
Many researchers have suggested that cursorial mammals concentrate limb muscle mass proximally to reduce energy costs during locomotion. Although supported by experiments where mass is added to an individual's limbs, mammals with naturally occurring distally heavy limbs such as primates have similar energy costs compared with other mammals. This study presents a new hypothesis to explain how animals with distally heavy limbs maintain low energy costs. Since distal mass should increase energy costs due to higher amounts of muscular power outputs, this hypothesis is based on the divergent effects of stride frequency on internal and external power outputs (the power output to move the limbs and the body center of mass, respectively). The use of low stride frequencies reduces limb velocities and therefore decreases internal power, while associated long strides increase the vertical displacement of the body center of mass and therefore increase external power. Total power (the sum of internal and external power) may therefore not differ among mammals with different limb mass distributions. To test this hypothesis, I examined a sample of infant baboons (Papio cynocephalus) during ontogeny and compared them with more cursorial mammals. Limb mass distribution changes with age (from distal to more proximally concentrated) in baboons, and the infants used shorter strides and higher stride frequencies when limb mass was most proximally concentrated. Compared with non-primates who have more proximally concentrated limb mass, the infants used longer strides and lower stride frequencies. Relatively low internal power was associated with low stride frequencies in both the intra- and inter-specific samples. However, only in the inter-specific comparison were relatively long strides associated with high external power outputs. In both the intra-specific and the inter-specific samples, total power did not differ between groups who differed in limb mass distribution. The results of this study suggest that a trade-off mechanism is available to quadrupeds with distally heavy limbs allowing them to maintain similar total power outputs (and likely similar energy costs) compared with mammals with more proximally concentrated limb mass.