Birds achieve high robustness in uneven terrain through active control of landing conditions

We understand little about how animals adjust locomotor behaviour to negotiate uneven terrain. The mechanical demands and constraints of such behaviours likely differ from uniform terrain locomotion. Here we investigated how common pheasants negotiate visible obstacles with heights from 10 to 50% of leg length. Our goal was to determine the neuro-mechanical strategies used to achieve robust stability, and address whether strategies vary with obstacle height. We found that control of landing conditions was crucial for minimising fluctuations in stance leg loading and work in uneven terrain. Variation in touchdown leg angle (theta(TD)) was correlated with the orientation of ground force during stance, and the angle between the leg and body velocity vector at touchdown (beta(TD)) was correlated with net limb work. Pheasants actively targeted obstacles to control body velocity and leg posture at touchdown to achieve nearly steady dynamics on the obstacle step. In the approach step to an obstacle, the birds produced net positive limb work to launch themselves upward. On the obstacle, body dynamics were similar to uniform terrain. Pheasants also increased swing leg retraction velocity during obstacle negotiation, which we suggest is an active strategy to minimise fluctuations in peak force and leg posture in uneven terrain. Thus, pheasants appear to achieve robustly stable locomotion through a combination of path planning using visual feedback and active adjustment of leg swing dynamics to control landing conditions. We suggest that strategies for robust stability are context specific, depending on the quality of sensory feedback available, especially visual input.