Walking on the Moon: Hypogravity drives the emergence of a proprioception-dependent locomotor state

Santuz, A.; Luciano, F.; Natalucci, V.; Mbaye, A.; Ma, N.; Cazzola, D.; Colyer, S.; Cowburn, J.; Albracht, K.; Braunstein, B.; Rittweger, J.; Herssens, N.; Weber, T.; Green, D. A.; de Noiij, J.; Minetti, A. E.; Pavei, G.; Zampieri, N.

doi:10.64898/2026.06.02.729513

bioRxiv · 10.64898/2026.06.02.729513

Walking on the Moon: Hypogravity drives the emergence of a proprioception-dependent locomotor state

Santuz, A.·Luciano, F.·Natalucci, V.·Mbaye, A.·Ma, N.·Cazzola, D.·Colyer, S.·Cowburn, J.·Albracht, K.·Braunstein, B.·Rittweger, J.·Herssens, N.·Weber, T.·Green, D. A.·de Noiij, J.·Minetti, A. E.·Pavei, G.·Zampieri, N.

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Abstract

Animals must adapt locomotion to changing environmental conditions, but how the nervous system flexibly controls movement remains unclear. Gravity provides a powerful perturbation because it alters body loading, limb dynamics, and proprioceptive feedback. Apollo astronauts often skipped on the Moon, adopting an asymmetric gait rarely used on Earth, yet the sensorimotor basis of this behaviour is unknown. Here, by studying hypogravity locomotion in humans and mice, we show that reduced gravity reveals a conserved proprioception-dependent mechanism for locomotor state expression. In humans, skipping in simulated lunar gravity is generated through flexible temporal reconfiguration of existing motor modules rather than construction of a new motor architecture. In mice, simulated lunar gravity elicited a skipping-like locomotor state, and genetic elimination of muscle proprioceptors abolished this response. Thus, hypogravity reveals how proprioceptive feedback enables flexible expression of locomotor states.

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Santuz, A., Luciano, F., Natalucci, V., Mbaye, A., Ma, N., Cazzola, D., Colyer, S., Cowburn, J., Albracht, K., Braunstein, B., Rittweger, J., Herssens, N., Weber, T., Green, D. A., de Noiij, J., Minetti, A. E., Pavei, G., Zampieri, N.. 2026-06-05. Walking on the Moon: Hypogravity drives the emergence of a proprioception-dependent locomotor state. https://doi.org/10.64898/2026.06.02.729513

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