Swimming in an Unsteady World

When animals swim in marine habitats, the water through which they move is usually flowing. Therefore, an important part of understanding the physics of how animals swim in nature is determining how they interact with the fluctuating turbulent water currents in their environment. The research systems we have been using to address this question are microscopic marine animals swimming in turbulent, wavy water flow over spatially-complex communities of organisms growing on surfaces. Field measurements of water motion were used to design realistic turbulent flow in a laboratory wave-flume over different substrata, particle-image velocimetry was used to measure fine-scale, rapidly-varying water velocity vector fields, and planar laser-induced fluorescence was used to measure concentrations of chemical cues from the substratum. We used individual-based models of small animals swimming in this unsteady flow to determine how their trajectories and contacts with substrata were affected by their locomotion through the water, rotation by local shear, response to odors, and transport by ambient flow. We found that the shears, accelerations, and odor concentrations encountered by small swimmers fluctuate rapidly, with peaks much higher than mean values lasting fractions of a second. We identified ways in which the behavior of small, weak swimmers can bias how they are transported by ambient flow (e.g. sinking during brief encounters with shear or odor enhances settlement onto substrata below, whereas constant swimming enhances contact with surfaces above or beside larvae). Although microscopic organisms swim slowly relative to ambient water flow, their locomotory behavior in response to the rapidly-fluctuating shears and odors they encounter can affect where they are transported by ambient water movement.