Strategies that simplify the control of quadrupedal stance. I. Forces at the ground.

1. Postural reactions were studied in six cats subjected to small, linear translations of the supporting surface in each of 16 different directions in the horizontal plane. Directions were specified in a polar coordinate system, with posterior translations being 0 degrees and leftward translations, 90 degrees. The data consisted of the forces exerted by each paw of the cat against the ground, measured in three orthogonal directions, vertical (z-axis), longitudinal (y-axis), and lateral (x-axis). 2. The force traces were analyzed by measuring the area under the curve during the postural reaction and dividing by the time of integration to give an average change in force. These values were normalized and plotted against direction of translation in polar coordinates, to give force tuning curves. The longitudinal and lateral force components were combined to generate force vectors in the horizontal plane. 3. Every cat responded to the platform translations with the same, simple strategy in which each hindlimb actively produced a correction force vector in one of only two possible directions. Participation of the forelimbs in the horizontal plane correction was not obligatory. While the direction of each hindlimb force vector was invariant, the amplitude was modulated according to the direction of platform movement. The resultant force vector, that acts through the center of mass of the animal, was in a direction opposite to the platform movement and directly opposed the perturbation. By this strategy, the cat was able to correct for destabilizing movements of the supporting surface in any direction in the horizontal plane. 4. It is concluded that the generation of forces between the paws and the ground is a high-level parameter that is controlled by the nervous system in a task-dependent manner. By using the strategy of restricting these forces to a set of two direction-invariant vectors, the problem of maintaining stance in the face of horizontal plane disturbances is greatly simplified.