title: Sensory-Motor Systems of Copepods involved in their Escape from Suction Feeding.
abstract: Copepods escape well by detecting minute gradients in the flow field; they react quickly, and swim away strongly. As a key link in the aquatic food web, these small planktonic organisms often encounter suction-feeding fish. Studies have identified certain hydrodynamic features that are created by the approach of this visual predator and the generation of its suction flow for capturing food. Similarly, studies have identified certain hydrodynamic features that evoke the evasive response of copepods. This is a review of the copepod sensory motor system as pertains to understanding their response to suction-feeding fish. Analyses of the reaction time, threshold sensitivity, structure of sensors, and evasive behavior by this key prey of fish can be useful for evaluating the effectiveness of feeding tactics in response to suction flow. To illustrate, we present results comparing a copepod from a fishless lake (Hesperodiaptomus shoshone) to a copepod from a rich fishing ground (Calanus finmarchicus). We designed a flow mimic that produces a realistic mushroom-cap-shaped flow field and realistic accelerations of flow; the copepods treated the mimic as a threat and performed jumps directed up and away from the siphon. Calanus finmarchicus responded at an average threshold strain rate of 18.7/s, escaped at 0.46 m/s, and traveled 5.99 mm, most frequently as a single jump. Hesperodiaptomus shoshone responded at a strain rate of 15.1/s that is not significantly different, escaped more slowly at 0.22 m/s and traveled a shorter distance of 3.01 mm using a series of hops. The high variability noted in the initial angle of the body and the maximum change in body angle suggests that unpredictability in the escape maneuver is another aspect of the tactic of copepods. The speed of the escape by small copepods 2-3 mm long is overwhelmed by the speed of the attack by the much larger, faster fish; if the copepod reacts when it is within the fish's arena of capture (<1.5 mm from mouth), it will be eaten. The copepod, however, has an acutely sensitive array of mechanosensors that perceive the flow field of the fish at distances of 3-6 mm, or outside the fish's range of capture. The copepod also has a rapid and strong locomotory response, thereby increasing the odds that the copepod will survive-but speed is unlikely to be the best tactic for staying alive. Instead, the copepod accelerates from 61.3 to 96.5 m/s(2) or more than 20 times stronger than the lunge of a fish. This collection of capabilities of copepods enables them to remain one of the most abundant multicellular organisms on our planet.
authors.affiliation: *School of Biology, Georgia Institute of Technology, Atlanta, GA 30332, USA; Department of Mechanical Engineering, The Johns Hopkins University, Baltimore, MD 21218, USA; Department of Mechanical Engineering, Stanford University, Stanford, CA 94305, USA; School of Civil and Environmental Engineering, Georgia Institute of Technology, Atlanta, GA 30332, USA jeannette.yen@biology.gatech.edu.
authors.forename: Jeannette
authors.initials: J
authors.lastname: Yen
authors.affiliation: *School of Biology, Georgia Institute of Technology, Atlanta, GA 30332, USA; Department of Mechanical Engineering, The Johns Hopkins University, Baltimore, MD 21218, USA; Department of Mechanical Engineering, Stanford University, Stanford, CA 94305, USA; School of Civil and Environmental Engineering, Georgia Institute of Technology, Atlanta, GA 30332, USA.
authors.forename: David W
authors.initials: DW
authors.lastname: Murphy
authors.affiliation: *School of Biology, Georgia Institute of Technology, Atlanta, GA 30332, USA; Department of Mechanical Engineering, The Johns Hopkins University, Baltimore, MD 21218, USA; Department of Mechanical Engineering, Stanford University, Stanford, CA 94305, USA; School of Civil and Environmental Engineering, Georgia Institute of Technology, Atlanta, GA 30332, USA.
authors.forename: Lin
authors.initials: L
authors.lastname: Fan
authors.affiliation: *School of Biology, Georgia Institute of Technology, Atlanta, GA 30332, USA; Department of Mechanical Engineering, The Johns Hopkins University, Baltimore, MD 21218, USA; Department of Mechanical Engineering, Stanford University, Stanford, CA 94305, USA; School of Civil and Environmental Engineering, Georgia Institute of Technology, Atlanta, GA 30332, USA.
authors.forename: Donald R
authors.initials: DR
authors.lastname: Webster
mesh_terms: D000818:Animals; D033342:Copepoda / Q000502:physiology*; D004924:Escape Reaction; D020387:Food Chain; D057446:Hydrodynamics; D013396:Suction; D013550:Swimming