Strategies that simplify the control of quadrupedal stance. II. Electromyographic activity

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Abstract
1. This study tested the hypothesis that muscle synergies underlie the invariance in the direction of corrective forces observed following stance perturbations in the horizontal plane. Electromyographic activity was recorded from selected forelimb and hindlimb muscles of cats subjected to horizontal translations of the supporting surface in 16 different directions. The responses of muscles were quantified for each perturbation, and tuning curves were constructed that related the amplitude of muscle response to the direction of platform movement. 2. Muscle tuning curves tended to group into one of two regions, corresponding to the two directions of force vectors. A few muscles showed clearly different recruitment patterns. The same direction of correction force vector was produced by different patterns of muscle activity, and the particular EMG pattern depended on the direction of platform movement. Therefore a simple muscle synergy organization could not account for the invariance in force vector generation. 3. It is concluded that there is a hierarchy of control in the maintenance of stance in which the vector of force exerted against the ground is a high level, task-dependent controlled variable and the selection of muscles to activate in order to produce the vector is controlled at a lower level. It is proposed that muscles are controlled using a modified synergy strategy. In this scheme, a synergy is not simply a fixed group of muscles, constrained to act as a unit. Rather, muscles are organized as a task-dependent synergy that is tuned or modified as needed by the addition or subtraction of other muscles.