Hydrodynamics of larval settlement from a larva's point of view

Many benthic marine invertebrate animals release larvae that are dispersed by ocean currents. These larvae swim and can respond to environmental factors such as chemical cues. However, larvae are so small (generally 0.01-1 mm) that they are often assumed to be passive particles whose trajectories are determined by the motion of the water in which they are riding. Therefore, marine larvae are useful model organisms to study the more general question of how the locomotion of very small animals in complex, variable natural habitats is affected by the motion of the fluid (water or air) around them. Studying larval locomotion under conditions of water flow encountered in nature is challenging because measuring the behavior of an individual microscopic organism requires high magnification imaging that is difficult to do in the field. The purpose of this article is to synthesize in one place the various approaches that we have been using to address the technical challenges of studying the locomotion of microscopic larvae in realistic ambient flow. The steps in our process include: (1) measuring water flow in the field; (2) mimicking realistic water movement in laboratory flumes to measure larval scale fluctuations in velocity of flow and concentration of chemical cues; (3) mimicking fine scale temporal patterns of larval encounters with a dissolved chemical cue to record larval responses; (4) using individual-based models to put larvae back into the larger scale environmental flow to determine trajectories; and (5) mimicking fine scale spatial and temporal patterns of larval encounters with water velocities and shear to determine the instantaneous forces on larvae. We illustrate these techniques using examples from our ongoing research on the settlement of larvae onto fouling communities and from our published work on settlement of larvae onto coral reefs. These examples show that water velocities and concentrations of chemical cues encountered by microscopic organisms can fluctuate in fractions of a second and vary over scales of less than a millimeter.