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.

According to the force–length–velocity relationships, the muscle force potential is determined by the operating length and velocity, which affects the energetic cost of contraction. During running, the human soleus muscle produces mechanical work through active shortening and provides the majority of propulsion. The trade-off between work production and alterations of the force–length and force–velocity potentials (i.e. fraction of maximum force according to the force–length–velocity curves) might mediate the energetic cost of running. By mapping the operating length and velocity of the soleus fascicles onto the experimentally assessed force–length and force–velocity curves, we investigated the association between the energetic cost and the force–length–velocity potentials during running. The fascicles operated close to optimal length (0.90 ± 0.10L₀) with moderate velocity (0.118 ± 0.039V_max[maximum shortening velocity]) and, thus, with a force–length potential of 0.92 ± 0.07 and a force–velocity potential of 0.63 ± 0.09. The overall force–length–velocity potential was inversely related (r= −0.52,p= 0.02) to the energetic cost, mainly determined by a reduced shortening velocity. Lower shortening velocity was largely explained (p< 0.001,R²= 0.928) by greater tendon gearing, shorter Achilles tendon lever arm, greater muscle belly gearing and smaller ankle angle velocity. Here, we provide the first experimental evidence that lower shortening velocities of the soleus muscle improve running economy.

According to the force-length-velocity relationships, the muscle force potential during locomotion is determined by the operating fibre length and velocity. We measured fascicle and muscle-tendon unit length and velocity as well as the activity of the human vastus lateralis muscle (VL) during walking and running. Furthermore, we determined the VL force-length relationship experimentally and calculated the force-length and force-velocity potentials (i.e. fraction of maximum force according to the force-length-velocity curves) for both gaits. During the active state of the stance phase, fascicles showed significantly (p < 0.05) smaller length changes (walking: 9.2 ± 4.7% of optimal length (L₀); running: 9.0 ± 8.4%L₀) and lower velocities (0.46 ± 0.36 L₀/s; 0.03 ± 0.83 L₀/s) compared to the muscle-tendon unit (walking: 19.7 ± 5.3%L₀, −0.94 ± 0.32 L₀/s; running: 34.5 ± 5.8%L₀, −2.59 ± 0.41 L₀/s). The VL fascicles operated close to optimum length (L₀ = 9.4 ± 0.11 cm) in both walking (8.6 ± 0.14 cm) and running (10.1 ± 0.19 cm), resulting in high force-length (walking: 0.92 ± 0.08; running: 0.91 ± 0.14) and force-velocity (0.91 ± 0.08; 0.97 ± 0.13) potentials. For the first time we demonstrated that, in contrast to the current general conception, the VL fascicles operate almost isometrically and close to L₀ during the active state of the stance phase of walking and running. The findings further verify an important contribution of the series-elastic element to VL fascicle dynamics.

Is the control of movement less stable when we walk or run in challenging settings? Intuitively, one might answer that it is, given that challenging locomotion externally (e.g., rough terrain) or internally (e.g., age-related impairments) makes our movements more unstable. Here, we investigated how young and old humans synergistically activate muscles during locomotion when different perturbation levels are introduced. Of these control signals, called muscle synergies, we analyzed the local stability and the complexity (or irregularity) over time. Surprisingly, we found that perturbations force the central nervous system to produce muscle activation patterns that are less unstable and less complex. These outcomes show that robust locomotion control in challenging settings is achieved by producing less complex control signals that are more stable over time, whereas easier tasks allow for more unstable and irregular control.