Challenging human locomotion: stability and modular organisation in unsteady conditions.

The need to move over uneven terrain is a daily challenge. In order to face unexpected perturbations due to changes in the morphology of the terrain, the central nervous system must flexibly modify its control strategies. We analysed the local dynamic stability and the modular organisation of muscle activation (muscle synergies) during walking and running on an even- and an uneven-surface treadmill. We hypothesized a reduced stability during uneven-surface locomotion and a reorganisation of the modular control. We found a decreased stability when switching from even- to uneven-surface locomotion (p < 0.001 in walking, p = 0.001 in running). Moreover, we observed a substantial modification of the time-dependent muscle activation patterns (motor primitives) despite a general conservation of the time-independent coefficients (motor modules). The motor primitives were considerably wider in the uneven-surface condition. Specifically, the widening was significant in both the early (+40.5%, p < 0.001) and late swing (+7.7%, p = 0.040) phase in walking and in the weight acceptance (+13.6%, p = 0.006) and propulsion (+6.0%, p = 0.041) phase in running. This widening highlighted an increased motor output's robustness (i.e. ability to cope with errors) when dealing with the unexpected perturbations. Our results confirmed the hypothesis that humans adjust their motor control strategies' timing to deal with unsteady locomotion.