Interactions between benthic predators and zooplanktonic prey are affected by turbulent waves.

Predators capture prey in complex and variable environments. In the ocean, bottom-dwelling (benthic) organisms are subjected to water currents, waves, and turbulent eddies. For benthic predators that feed on small animals carried in the water (zooplankton), flow not only delivers prey, but can also shape predator-prey interactions. Benthic passive suspension feeders collect prey delivered by movement of ambient water onto capture-surfaces, whereas motile benthic predators, such as burrow-dwelling fish, dart out to catch passing zooplankton. How does the flow of ambient water affect these contrasting modes of predation by benthic zooplanktivores? We studied the effects of turbulent, wavy flow on the encounter, capture, and retention of motile zooplanktonic prey (copepods, Acartia spp.) by passive benthic suspension feeders (sea anemones, Anthopleura elegantissima). Predator-prey interactions were video-recorded in a wave-generating flume under two regimes of oscillating flow with different peak wave velocities and levels of turbulent kinetic energy ("weak" and "strong" waves). Rates of encounter (number of prey passing through a sea anemone's capture zone per time), capture (prey contacting and sticking to tentacles per time), and retention (prey retained on tentacles, without struggling free or washing off, per time) were measured at both strengths of waves. Strong waves enhanced encounter rates both for dead copepods and for actively swimming copepods, but there was so much variability in the behavior of the live prey that the effect of wave strength on encounter rates was not significant. Trapping efficiency (number of prey retained per number encountered) was the same in both flow regimes because, although fewer prey executed maneuvers to escape capture in strong waves, more of the captured prey was washed off the predators' tentacles. Although peak water velocities and turbulence of waves did not affect feeding rates of passive suspension-feeding sea anemones, increases in these aspects of flow have been shown to enhance feeding rates and efficiency of motile benthic fish that lunge out of their burrows to catch zooplankton. Faster, more turbulent flow interferes with the ability of prey to detect predators and execute escape maneuvers, and thus enhances capture rates both for passive suspension-feeding predators and for actively swimming predators. However, prey captured in the mouths of fish are not washed away by ambient flow, whereas prey captured on the tentacles of suspension feeders can be swept off before they are ingested. Therefore, the effects of flowing water on predation on zooplankton by benthic animals depend on the feeding mode of the predator.

Chemical Induction of Larval Settlement Behavior in Flow.

The ability of dissolved chemical cues to induce larval settlement from the water column has long been debated. Through computer-assisted video motion analysis, we quantified the movements of individual oyster (Crassostrea virginica) larvae in a small racetrack flume at free-stream flow speeds of 2.8, 6.2, and 10.4 cm/s. In response to waterborne chemical cues, but not to seawater (control), oyster larvae moved downward in the water column and swam in slow curved paths before attaching to the flume bottom. Effective stimuli were adult-oyster-conditioned seawater (OCW) and a synthetic peptide analog (glycyl-glycyl-L-arginine) for the natural cue. The chemically mediated behavioral responses of oyster larvae in flow were essentially identical to those responses previously reported in still water. Our experimental results therefore demonstrate the capacity of waterborne cues to evoke settlement behavior in oyster pediveligers under varying hydrodynamic conditions.

Odor Plumes and Animal Navigation in Turbulent Water Flow: A Field Study.

Turbulence causes chemical stimuli to be highly variable in time and space; hence the study of animal orientation in odor plumes presents a formidable challenge. Through combined chemical and physical measurements, we characterized the transport of attractant released by clam prey in a turbulent aquatic environment. Concurrently, we quantified the locomotory responses of predatory crabs successfully searching for sources of clam attractant. Our results demonstrate that both rheotaxis and chemotaxis are necessary for successful orientation. Perception of chemical cues causes crabs to move in the upstream direction, but feedback from attractant distributions directly regulates movement across-stream in the plume. Orientation mechanisms used by crabs difler from those employed by flying insects, the only other system in which navigation relative to odor plumes has been coupled with fluid dynamics. Insects respond to odors by moving upstream, but they do not use chemical distributions to determine across-stream direction, whereas crabs do. Turbulent eddy diffusivities in crab habitats are 100 to 1000 times lower than those of terrestrial grasslands and forests occupied by insects. Insects must respond to plumes consisting of highly dispersed, tiny filaments or parcels of odor. Crabs rely more heavily on spatial aspects of chemical stimulus distributions because their fluid dynamic environment creates a more stable plume structure, thus permitting chemotaxis.