Abstract
Limbless crawling is a fundamental form of biological locomotion adopted by a wide variety of species, including the amoeba, earthworm and snake. An interesting question from a biomechanics perspective is how limbless crawlers control their flexible bodies in order to realize directional migration. In this paper, we discuss the simple but instructive problem of peristalsis-like locomotion driven by elongation-contraction waves that propagate along the body axis, a process frequently observed in slender species such as the earthworm. We show that the basic equation describing this type of locomotion is a linear, one-dimensional diffusion equation with a time-space-dependent diffusion coefficient and a source term, both of which express the biological action that drives the locomotion. A perturbation analysis of the equation reveals that adequate control of friction with the substrate on which locomotion occurs is indispensable in order to translate the internal motion (propagation of the elongation-contraction wave) into directional migration. Both the locomotion speed and its direction (relative to the wave propagation) can be changed by the control of friction. The biological relevance of this mechanism is discussed.