Cursorial ground birds are paragons of bipedal running that span a 500-fold mass range from quail to ostrich. Here we investigate the task-level control priorities of cursorial birds by analysing how they negotiate single-step obstacles that create a conflict between body stability (attenuating deviations in body motion) and consistent leg force–length dynamics (for economy and leg safety). We also test the hypothesis that control priorities shift between body stability and leg safety with increasing body size, reflecting use of active control to overcome size-related challenges. Weight-support demands lead to a shift towards straighter legs and stiffer steady gait with increasing body size, but it remains unknown whether non-steady locomotor priorities diverge with size. We found that all measured species used a consistent obstacle negotiation strategy, involving unsteady body dynamics to minimise fluctuations in leg posture and loading across multiple steps, not directly prioritising body stability. Peak leg forces remained remarkably consistent across obstacle terrain, within 0.35 body weights of level running for obstacle heights from 0.1 to 0.5 times leg length. All species used similar stance leg actuation patterns, involving asymmetric force–length trajectories and posture-dependent actuation to add or remove energy depending on landing conditions. We present a simple stance leg model that explains key features of avian bipedal locomotion, and suggests economy as a key priority on both level and uneven terrain. We suggest that running ground birds target the closely coupled priorities of economy and leg safety as the direct imperatives of control, with adequate stability achieved through appropriately tuned intrinsic dynamics.
Rapid unloading and a peak in power output of the ankle joint have been widely observed during push-off in human walking. Modelbased studies hypothesize that this push-off causes redirection of the body center of mass just before touch-down of the leading leg. Other research suggests that work done by the ankle extensors provides kinetic energy for the initiation of swing. Also, muscle work is suggested to power a catapult-like action in late stance of human walking. However, there is a lack of knowledge about the biomechanical process leading to this widely observed high power output of the ankle extensors. In our study, we use kinematic and dynamic data of human walking collected at speeds between 0.5 and 2.5 m s −1 for a comprehensive analysis of push-off mechanics. We identify two distinct phases, which divide the push-off: first, starting with positive ankle power output, an alleviation phase, where the trailing leg is alleviated from supporting the body mass, and second, a launching phase, where stored energy in the ankle joint is released. Our results show a release of just a small part of the energy stored in the ankle joint during the alleviation phase. A larger impulse for the trailing leg than for the remaining body is observed during the launching phase. Here, the buckling knee joint inhibits transfer of power from the ankle to the remaining body. It appears that swing initiation profits from an impulsive ankle push-off resulting from a catapult without escapement.
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