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Fruit flies or Drosophila larvae exhibit a diverse range of locomotion gaits enabled by their soft, segmented bodies and intricate muscle arrangements. Their bodies, composed of multiple segments, are synchronously activated to propel forward through a combination of muscle elongation and contraction. Soft robotic systems, inspired by such biological marvels, face significant challenges in replicating these complex behaviors due to the intricate interplay between muscle activation, soft body dynamics, and frictional forces. To address these challenges, we propose a reduced-order model that captures the essential features of larval crawling. By modeling segments as a combination of prismatic and revolute joints, we can simulate the nonlinear motion resulting from muscle activation and body deformation. Our model demonstrates the potential of this approach to accurately describe larval movement, as validated by comparisons with actual larval trajectories. This research offers valuable insights into the design and control of soft robots and provides a framework for biologists to investigate the complex mechanisms of neuromuscular coordination in larvae. 

When threatened by dangerous or harmful stimuli, animals engage in diverse forms of rapid escape behaviors. InDrosophilalarvae, one type of escape response involves C-shaped bending and lateral rolling followed by rapid forward crawling. The sensory circuitry that promotes larval escape has been extensively characterized; however, the motor programs underlying rolling are unknown. Here, we characterize the neuromuscular basis of rolling escape behavior. We used high-speed, volumetric, Swept Confocally Aligned Planar Excitation (SCAPE) microscopy to image muscle activity during larval rolling. Unlike sequential peristaltic muscle contractions that progress from segment to segment during forward and backward crawling, muscle activity progresses circumferentially during bending and rolling escape behavior. We propose that progression of muscular contraction around the larva’s circumference results in a transient misalignment between weight and the ground support forces, which generates a torque that induces stabilizing body rotation. Therefore, successive cycles of slight misalignment followed by reactive aligning rotation lead to continuous rolling motion. Supporting our biomechanical model, we found that disrupting the activity of muscle groups undergoing circumferential contraction progression leads to rolling defects. We use EM connectome data to identify premotor to motor connectivity patterns that could drive rolling behavior and perform neural silencing approaches to demonstrate the crucial role of a group of glutamatergic premotor neurons in rolling. Our data reveal body-wide muscle activity patterns and putative premotor circuit organization for execution of the rolling escape response.