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In addition to a vestibular system, birds uniquely have a balance-sensing organ within the pelvis, called the lumbosacral organ (LSO). The LSO is well developed in terrestrial birds, possibly to facilitate balance control in perching and terrestrial locomotion. No previous studies have quantified the functional benefits of the LSO for balance. We suggest two main benefits of hip-localized balance sense: reduced sensorimotor delay and improved estimation of foot-ground acceleration. We used system identification to test the hypothesis that hip-localized balance sense improves estimates of foot acceleration compared to a head-localized sense, due to closer proximity to the feet. We built a physical model of a standing guinea fowl perched on a platform, and used 3D accelerometers at the hip and head to replicate balance sense by the LSO and vestibular systems. The horizontal platform was attached to the end effector of a 6 DOF robotic arm, allowing us to apply perturbations to the platform analogous to motions of a compliant branch. We also compared state estimation between models with low and high neck stiffness. Cross-correlations revealed that foot-to-hip sensing delays were shorter than foot-to-head, as expected. We used multi-variable output error state-space (MOESP) system identification to estimate foot-ground acceleration as a function of hip- and head-localized sensing, individually and combined. Hip-localized sensors alone provided the best state estimates, which were not improved when fused with head-localized sensors. However, estimates from head-localized sensors improved with higher neck stiffness. Our findings support the hypothesis that hip-localized balance sense improves the speed and accuracy of foot state estimation compared to head-localized sense. The findings also suggest a role of neck muscles for active sensing for balance control: increased neck stiffness through muscle co-contraction can improve the utility of vestibular signals. Our engineering approach provides, to our knowledge, the first quantitative evidence for functional benefits of the LSO balance sense in birds. The findings support notions of control modularity in birds, with preferential vestibular sense for head stability and gaze, and LSO for body balance control,respectively. The findings also suggest advantages for distributed and active sensing for agile locomotion in compliant bipedal robots.

A Physical Model Suggests That Hip-Localized Balance Sense in Birds Improves State Estimation in Perching: Implications for Bipedal Robots