Flexible muscle modes and synergies in challenging whole-body tasks

We used the idea of hierarchical control to study multi-muscle synergies during a whole-body sway task performed by a standing person. Within this view, at the lower level of the hierarchy, muscles are united into groups (M-modes). At the higher level, gains at the M-modes are co-varied by the controller in a task-specific way to ensure low variability of important physical variables. In particular, we hypothesized that (1) the composition of M-modes could adjust and (2) an index of M-mode co-variation would become weaker in more challenging conditions. Subjects were required to perform a whole-body sway at 0.5 Hz paced by a metronome. They performed the task with eyes open and closed, while standing on both feet or on one foot only, with and without vibration applied to the Achilles tendons. Integrated indices of muscle activation were subjected to principal component analysis to identify M-modes. An increase in the task complexity led to an increase in the number of principal components that contained significantly loaded indices of muscle activation from 3 to 5. Hence, in more challenging tasks, the controller manipulated a larger number of variables. Multiple regression analysis was used to define the Jacobian of the system mapping small changes in M-mode gains onto shifts of the center of pressure (COP) in the anterior-posterior direction. Further, the variance in the M-mode space across sway cycles was partitioned into two components, one that did not affect an average across cycles COP coordinate and the other that did (good and bad variance, respectively). Under all conditions, the subjects showed substantially more good variance than bad variance interpreted as a multi-M-mode synergy stabilizing the COP trajectory. An index of the strength of the synergy was comparable across all conditions, and there was no modulation of this index over the sway cycle. Hence, our first hypothesis that the composition of M-modes could adjust under challenging conditions has been confirmed while the second hypothesis stating that the index of M-mode co-variation would become weaker in more challenging conditions has been falsified. We interpret the observations as suggesting that adjustments at the lower level of the hierarchy-in the M-mode composition-allowed the subjects to maintain a comparable level of stabilization of the COP trajectory in more challenging tasks. The findings support the (at least) two-level hierarchical control scheme of whole-body movements.