Predicting path from undulations for C. elegans using linear and nonlinear resistive force theory

A basic issue in the physics of behaviour is the mechanical relationship between an animal and its surroundings. The model nematode C. elegans provides an excellent platform to explore this relationship due to its anatomical simplicity. Nonetheless, the physics of nematode crawling, in which the worm undulates its body to move on a wet surface, is not completely understood and the mathematical models often used to describe this phenomenon are empirical. We confirm that linear resistive force theory, one such empirical model, is effective at predicting a worm's path from its sequence of body postures for forward crawling, reversing, and turning and for a broad range of different behavioural phenotypes observed in mutant worms. Worms recently isolated from the wild have a higher effective drag anisotropy than the laboratory-adapted strain N2 and most mutant strains. This means the wild isolates crawl with less surface slip, perhaps reflecting more efficient gaits. The drag anisotropies required to fit the observed locomotion data (70  ±  28 for the wild isolates) are significantly larger than the values measured by directly dragging worms along agar surfaces (3-10 in Rabets et al (2014 Biophys. J. 107 1980-7)). A proposed nonlinear extension of the resistive force theory model also provides accurate predictions, but does not resolve the discrepancy between the parameters required to achieve good path prediction and the experimentally measured parameters. We confirm that linear resistive force theory provides a good effective model of worm crawling that can be used in applications such as whole-animal simulations and advanced tracking algorithms, but that the nature of the physical interaction between worms and their most commonly studied laboratory substrate remains unresolved.