The rich repertoire of locomotor behaviors in quadrupedal animals requires flexible inter-limb and inter-segmental coordination. Here we studied the kinematic coordination of different gaits (walk, trot, gallop, and swim) of six dogs (Canis lupus familiaris) and, in particular, the planar covariation of limb segment elevation angles. The results showed significant variations in the relative duration of rearward limb movement, amplitude of angular motion, and inter-limb coordination, with gait patterns ranging from a lateral sequence of footfalls during walking to a diagonal sequence in swimming. Despite these differences, the planar law of inter-segmental coordination was maintained across different gaits in both forelimbs and hindlimbs. Notably, phase relationships and orientation of the covariation plane were highly limb specific, consistent with the functional differences in their neural control. Factor analysis of published muscle activity data also demonstrated differences in the characteristic timing of basic activation patterns of the forelimbs and hindlimbs. Overall, the results demonstrate that the planar covariation of inter-segmental coordination has emerged for both fore- and hindlimbs and all gaits, although in a limb-specific manner.

Locomotion of tetrapods on land adapted to different environments and needs resulting in a variety of different gait styles. However, comparative analyses reveal common principles of limb movement control. Here, we report that a kinematic synergy involving the planar covariation of limb segment motion holds in 54 different animal species (10 birds and 44 mammals), despite large differences in body size, mass (ranging from 30 g to 4 tonnes), limb configuration, and amplitude of movements. This kinematic synergy lies at the interface between the neural command signals output by locomotor pattern generators, the mechanics of the body center of mass and the external environment, and it may represent one neuromechanical principle conserved in evolution to save mechanical energy.