Dispersion Relations for Active Undulators in Overdamped Environments.

Organisms that locomote by propagating waves of body bending can maintain performance across heterogeneous environments by modifying their gait frequency $\omega$ or wavenumber $k$. We identify a unifying relationship between these parameters for overdamped undulatory swimmers (including nematodes, spermatozoa, and mm-scale fish) moving in diverse environmental rheologies, in the form of an active `dispersion relation' $\omega\propto k^{\pm2}$. A model treating the organisms as actively driven viscoelastic beams reproduces the experimentally observed scaling. The relative strength of rate-dependent dissipation in the body and the environment determines whether $k^2$ or $k^{-2}$ scaling is observed. The existence of these scaling regimes reflects the $k$ and $\omega$ dependence of the various underlying force terms and how their relative importance changes with the external environment and the neuronally commanded gait.