Control of ground reaction forces by hindlimb muscles during cat locomotion

It has been proposed that biarticular muscles are primarily responsible for the control of the direction of external forces, as their activation is closely related and highly sensitive to the direction of external forces. This functional role for biarticular muscles has been supported qualitatively by experimental evidence, but has never been tested quantitatively for lack of a mathematical/mechanical formulation of this theory and the difficulty of measuring individual muscle forces during voluntary movements. The purposes of this study were: (1) to define rules for muscular coordination based on the control of external forces; (2) to develop a model of the cat hindlimb that allows for the calculation of the magnitude and direction of the ground reaction forces (GRFs) produced by individual hindlimb muscles; and (3) to test if the coordination of mono- and biarticular cat hindlimb muscles is related to the control of the resultant GRF. We measured the GRF, hindlimb kinematics, selected muscle forces and activations during cat locomotion. Then, the measured muscle forces were used as input to the hindlimb model to compute the muscle-induced GRF. We assume that if activation (and possibly force) increased as the muscle-induced component of GRF approximated the resultant GRF, then that muscle was used by the central nervous system (CNS) to help control the direction of the external GRF. During cat walking, medial gastrocnemius (MG) and plantaris (PL) forces increased with increasing proximity to the GRF, while soleus (SOL) forces and vastus lateralis (VL) activations did not. SOL and VL activation were most strongly related to the vertical and parallel (braking/accelerating) component of the GRF, respectively. We concluded from these results that MG and PL are primarily responsible for the control of the direction of the GRF, while SOL primarily functions as an anti-gravity muscle, and VL as an acceleration/deceleration muscle.