Limb and trunk mechanisms for balance control during locomotion in quadrupeds

In quadrupeds, the most critical aspect of postural control during locomotion is lateral stability. However, neural mechanisms underlying lateral stability are poorly understood. Here, we studied lateral stability in decerebrate cats walking on a treadmill with their hindlimbs. Two destabilizing factors were used: a brief lateral push of the cat and a sustained lateral tilt of the treadmill. It was found that the push caused considerable trunk bending and twisting, as well as changes in the stepping pattern, but did not lead to falling. Due to postural reactions, locomotion with normal body configuration was restored in a few steps. It was also found that the decerebrate cat could keep balance during locomotion on the laterally tilted treadmill. This postural adaptation was based on the transformation of the symmetrical locomotor pattern into an asymmetrical one, with different functional lengths of the right and left limbs. Then, we analyzed limb and trunk neural mechanisms contributing to postural control during locomotion. It was found that one of the limb mechanisms operates in the transfer phase and secures a standard (relative to the trunk) position for limb landing. Two other limb mechanisms operate in the stance phase; they counteract distortions of the locomotor pattern by regulating the limb stiffness. The trunk configuration mechanism controls the body shape on the basis of sensory information coming from trunk afferents. We suggest that postural reactions generated by these four mechanisms are integrated, thus forming a response of the whole system to perturbation of balance during locomotion.