Swimming at low Reynolds number: a beginners guide to undulatory locomotion

Undulatory locomotion is a means of self-propulsion that relies on the generation and propagation of waves along a body. As a mode of locomotion it is primitive and relatively simple, yet can be remarkably robust. No wonder then, that it is so prevalent across a range of biological scales from motile bacteria to gigantic prehistoric snakes. Key to understanding undulatory locomotion is the body's interplay with the physical environment, which the swimmer or crawler will exploit to generate propulsion, and in some cases, even to generate the underlying undulations. This review focuses by and large on undulators in the low Reynolds number regime, where the physics of the environment can be much more tractable. We review some key concepts and theoretical advances, as well as simulation tools and results applied to selected examples of biological swimmers. In particular, we extend the discussion to some simple cases of locomotion in non-Newtonian media as well as to small animals, in which the nervous system, motor control, body properties and the environment must all be considered to understand how undulations are generated and modulated. To conclude, we review recent progress in microrobotic undulators that may one day become commonplace in applications ranging from toxic waste disposal to minimally invasive surgery.