The dynamics of hylobatid bipedalism: evidence for an energy-saving mechanism?

Abstract

When gibbons travel through the forest canopy, brachiation is alternated with short bipedal bouts over horizontal boughs. We know, from previous research, that brachiation is a very efficient locomotor mode that makes use of a pendulum-like exchange of energy, but to date, nothing is known about the dynamics of hylobatid bipedalism. We wondered if gibbons also make use of an efficient gait mechanism during bipedal locomotion. To investigate this, we calculated oscillations of the centre of mass (COM), energy fluctuations, recovery rates and power outputs from the 3D ground reaction forces. These ground reaction forces were collected during spontaneous bipedal locomotion of four untrained white-handed gibbons (Hylobates lar) over an instrumented walkway (with an AMTI force plate). Excursions of the COM are relatively large during hylobatid bipedalism and the fluctuations of potential and kinetic energy are largely in-phase. Together with the low inverted pendulum recovery rates, this points to a spring-mass mechanism during bipedal locomotion. Although the well-developed Achilles tendon of gibbons seems to be a good candidate for the storage and recoil of elastic energy, this is not supported by kinematical data of the ankle joint. Instead, we suggest that the knee extensor muscle tendon unit functions as an energy-saving mechanism during hylobatid bipedalism, but detailed anatomical data is needed to confirm this suggestion. At low speeds gibbons use either pendular or spring mechanics, but a clear gait transition as seen in most quadrupedal mammals is absent. At moderate to high velocities, gibbons use a bouncing gait, generally without aerial phases. This supports the view that aerial phases are not a prerequisite for spring-mass mechanics and reinforces the claim that duty factor alone should not be used to distinguish between a walk and run.