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SUMMARYDuring running in a natural environment, humans must routinely negotiate varied and unpredictable changes in ground level. To prevent a fall, changes in ground level, especially those that are invisible, require a quick response of the movement system within a short time. For 11 subjects we investigated two consecutive contacts during running across visible (drop of 0, 5 and 10cm) and camouflaged (drop of 0 and 10cm) changes in ground level. For both situations, we found significant variances in their leg parameters and ground reaction forces (GRFs) during the perturbed second contact but also one step ahead, in the unperturbed first contact. At visible first contact, humans linearly adapt their GRF to lower and smooth their centre of mass. During the camouflaged situation, the GRF also decreased, but it seems that the runners anticipate a drop of approximately 5-10cm. The GRF increased with drop height during the visible perturbed second contact. At the camouflaged second contact, GRFs differed noticeably from the observed reaction when crossing a similar visible drop, whereas the contact time decreased and the initial impact peak increased. This increased impact can be interpreted as a purely mechanical contribution to cope with the event. Furthermore, we observed an increased angle of attack and leg length with drop height for both situations. This is in accordance with results observed in birds running over a track with an unexpected drop, and suggests that adaptations in swing leg retraction form part of the strategy for running across uneven ground.

Pronograde trunk orientation in small birds causes prominent intra-limb asymmetries in the leg function. As yet, it is not clear whether these asymmetries induced by the trunk reflect general constraints on the leg function regardless of the specific leg architecture or size of the species. To address this, we instructed 12 human volunteers to walk at a self-selected velocity with four postures: regular erect, or with 30 deg, 50 deg and maximal trunk flexion. In addition, we simulated the axial leg force (along the line connecting hip and centre of pressure) using two simple models: spring and damper in series, and parallel spring and damper. As trunk flexion increases, lower limb joints become more flexed during stance. Similar to birds, the associated posterior shift of the hip relative to the centre of mass leads to a shorter leg at toe-off than at touchdown, and to a flatter angle of attack and a steeper leg angle at toe-off. Furthermore, walking with maximal trunk flexion induces right-skewed vertical and horizontal ground reaction force profiles comparable to those in birds. Interestingly, the spring and damper in series model provides a superior prediction of the axial leg force across trunk-flexed gaits compared with the parallel spring and damper model; in regular erect gait, the damper does not substantially improve the reproduction of the human axial leg force. In conclusion, mimicking the pronograde locomotion of birds by bending the trunk forward in humans causes a leg function similar to that of birds despite the different morphology of the segmented legs.

In locomotion, humans have to deal with changes in ground level like pavement or stairs.When they encounter uneven ground with changes in terrain height, they reduce their angle of attack and leg stiffness on a step. This strategy was found for the single step upward movement. However, are these adjustments the result of a general strategy? In our study we focused on leg adjustments while running up and down, implying permanent adaptation to a new track level. To investigate this, we measured ten healthy participants as they ran along a runway with 10 cm increased and 10 cm lowered steps. We found that ground reaction force, leg length, leg stiffness, and angle of attack were adjusted to the direction of the vertical disturbance (up or down) but also to its length. When running upwards, leg stiffness decreased by about 20.4% on the single step and by about 9.3% on the permanently elevated track step. In addition to that -when running downwards -leg stiffness decreased in preparation for the downward step by about 18.8%. We also observed that the angle of attack diminished on elevated contact from 61° to 59°, and increased on lowered contact from 61° to 65°. The adjustment of leg stiffness seemed to be actively achieved, whereas the angle of attack appeared to be passively adjusted, consistent with a running model that includes leg retraction in late swing phase.

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While running on uneven ground, humans are able to negotiate visible but also camouflaged changes in ground level. Previous studies have shown that the leg kinematics before touch down change with ground level. The present study experimentally investigated the contributions of visual perception (visual feedback), proprioceptive feedback and feed-forward patterns to the muscle activity responsible for these adaptations. The activity of three bilateral lower limb muscles (m. gastrocnemius medialis, m. tibialis anterior and m. vastus medialis) of nine healthy subjects was recorded during running across visible (drop of 0, −5 and −10 cm) and camouflaged changes in ground level (drop of 0 and −10 cm). The results reveal that at touchdown with longer flight time, m. tibialis anterior activation decreases and m. vastus medialis activation increases purely by feed-forward driven (flight time-dependent) muscle activation patterns, while m. gastrocnemius medialis activation increase is additionally influenced by visual feedback. Thus, feed-forward driven muscle activation patterns are sufficient to explain the experimentally observed adjustments of the leg at touchdown.