Mechanisms of force and power production in unsteady ricochetal brachiation

Brachiators travel by swinging beneath handholds, and it is not obvious how these animals manage to accelerate and decelerate in a horizontal direction, especially when moving rapidly. Most previous analyses focused on brachiation in highly constrained laboratory conditions that induced steady-state locomotion. Emerging understanding of brachiation suggests that much of gibbon locomotory behavior and morphology must be considered within the context of the complexities of the natural environment: the forest canopy is three-dimensional, with high variation in handhold availability and properties. The goal of this paper is to quantify the active mechanisms by which gibbons can dynamically control their velocity. Force production and kinematics were analyzed from a white-handed gibbon Hylabates lar during ricochetal brachiation. Both the mechanisms of force production and power input may be inferred for accelerating and decelerating brachiation by combining force data with kinematics. Examples of steady-state, accelerating, and decelerating ricochetal brachiation are highlighted. Gibbons are able to produce net horizontal impulses by releasing early (resulting in a loss of potential energy, but an accelerating horizontal impulse) or delaying release (associated with an increase in potential energy, and a decelerating horizontal impulse). Torque about the shoulder, leg-lifting (or dropping), and elbow flexing (or straightening) are discussed as potential mechanisms for controlling energy within the brachiating system. Of these possibilities, leg-lifting and arm-flexing were observed as mechanisms of adding mechanical energy. Net energy loss, and substantial torques about the shoulder, were not observed.