11Coordinated rhythmic movements are ubiquitous in animal behavior. In many organisms, chains 12 of neural oscillators underlie the generation of these rhythms. In C. elegans, locomotor wave 13 generation has been poorly understood; in particular, it is unclear where in the circuit rhythms 14 are generated, and whether there exists more than one such generator. We used optogenetic 15and ablation experiments to probe the nature of rhythm generation in the locomotor circuit. We 16 found that multiple sections of forward locomotor circuitry are capable of independently 17 generating rhythms. By perturbing different components of the motor circuit, we localize the 18 source of secondary rhythms to cholinergic motor neurons in the midbody. Using rhythmic 19 optogenetic perturbation we demonstrate bidirectional entrainment of oscillations between 20 different body regions. These results show that, as in many other vertebrates and invertebrates, 21 the C. elegans motor circuit contains multiple oscillators that coordinate activity to generate 22
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