Mechanics of soft-body rolling motion without external torque

The Drosophila larva, a soft-body animal, can bend its body and roll efficiently to escape danger. However, contrary to common belief, this rolling motion is not driven by the imbalance of gravity and ground reaction forces. Through functional imaging and ablation experiments, we demonstrate that the sequential actuation of axial muscles within an appropriate range of angles is critical for generating rolling. We model the interplay between muscle contraction, hydrostatic skeleton deformation, and body-environment interactions, and systematically explain how sequential muscle actuation generates the rolling motion. Additionally, we constructed a pneumatic soft robot to mimic the larval rolling strategy, successfully validating our model. This mechanics model of soft-body rolling motion not only advances the study of related neural circuits, but also holds potential for applications in soft robotics.