Multiple mechanisms contribute to the generation, propagation, and coordination of 8 rhythmic patterns necessary for locomotion in Caenorhabditis elegans. Current experiments have 9 focused on two possibilities: pacemaker neurons and stretch-receptor feedback. Here, we focus on 10 whether locomotion behavior can be produced by a chain of network oscillators in the ventral 11 nerve cord. We use a simulation model to demonstrate that a repeating neural circuit identified in 12 the worm's connectome can be chained together to drive forward locomotion on agar in a 13 neuromechanical model of the nematode, in the absence of pacemaker neurons or 14 stretch-receptor feedback. Systematic exploration of the space of possible solutions reveals that 15 there are multiple configurations that result in locomotion that match the kinematics of the worm 16 on agar. Analysis of the best solutions reveals that gap junctions between different classes of 17 motoneurons are likely to play key roles in coordinating oscillations along the ventral nerve cord. 18 19 40 1 of 24 Manuscript submitted to eLife that modulate C. elegans locomotion (Tavernarakis et al., 1997), as well as evidence of a direct 41 relationship between body curvature and neural activity (Wen et al., 2012). However, coordinated 42 rhythmic patterns can also be produced internally, while remaining open to modulation through 43 external contributions. Central pattern generators (CPGs) are known to be involved in a wide variety 44 of behaviors in a number of different organisms, including insect flight, swimming in molluscs, gut 45 movements in crustaceans, and swimming and respiration in vertebrates (Marder and Bucher, 2001; 46 Goulding, 2009; Katz, 2016; Arshavsky et al., 2016; Dasen, 2018; Minassian et al., 2017). In a CPG, 47 the rhythmic pattern can be generated through the intrinsic oscillatory properties of pacemaker 48 neurons or it can emerge from the interaction of networks of non-oscillatory neurons (Goulding, 49 2009). Recent experiments have provided support for the role of intrinsic oscillations in C. elegans 50 locomotion (Gao et al., 2018; Fouad et al., 2018; Xu et al., 2018). Although the work attributes the 51 source of these rhythm generators to pacemaker neurons, the evidence provided does not discard 52 the possibility of network oscillators (Wen et al., 2018). 53 It is increasingly acknowledged that simulation models play an important role in elucidating 54 how brain-body-environment systems produce behavior (Ijspeert, 2008; Abbott, 2008; Izquierdo, 55 2018). In C. elegans, there has been an surge of theoretical work focused on understanding the 56 neuromechanical basis of locomotion. Several computational models have demonstrated that 57 proprioception alone can be used to generate rhythmic patterns and propagate them along the 58 body (Niebur and Erdös, 1991; Karbowski et al., 2008; Boyle, 2009; Mailler et al., 2010; Wen et al., 59 2012; Izquierdo and Beer, 2018; Fieseler et al., 2018; Gleeson et al., 2018). There have also been a 60 number of m...