Effects of sea hare ink secretion and its escapin-generated components on a variety of predatory fishes

Sea hares, Aplysia californica, have a diversity of anti-predatory defenses. One is an actively released chemical defense: an ink secretion that is a mixture of two glandular products--ink from the ink gland and opaline from the opaline gland. The mechanisms of action of ink secretion and its components have recently been examined in detail against several predatory invertebrates. Our goal is to extend this mechanistic analysis to predatory vertebrates. Toward this end, the current study details the effects of ink, opaline, and one set of its components--the products of the reaction of escapin, an l-amino acid oxidase, with its natural substrates, L-lysine and L-arginine--on the palatability of food for five species of fishes: bluehead wrasses Thalassoma bifasciatum, señorita wrasses Oxyjulis californica, pinfish Lagodon rhomboides, mummichogs Fundulus heteroclitus, and bonnethead sharks Sphyrna tiburo. These fishes have different feeding styles, ranging from large fishes able to engulf sea hares to smaller fishes able to attack sea hares by pecking at them; and they live in a variety of habitats, including those that sea hares typically inhabit. We show that ink but not opaline significantly decreases the palatability of food for all five species, and that escapin products are mildly unpalatable to the two species of wrasses but not to the other species. These results, together with others, show that sea hare ink affects a diversity of predatory fishes, setting the stage for mechanistic studies using electrophysiological analysis of their chemosensory systems.