Directional control and the functional organization of defensive responses inAplysia

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Abstract
Noxious cutaneous stimulation of anterior sites onAplysia californica causes withdrawal and turning followed by escape locomotion (Figs. 1, 2). Stimulation of anterior sites causes significantly larger turning responses than does stimulation of posterior sites (Fig. 3), so that escape locomotion is always directed away from a site of ‘attack’. Later phases of escape locomotion are often the same, regardless of the site of the triggering stimulus (Fig. 4). The defensive secretions, ink and opaline, are directed along the anterior-posterior axis at the source of noxious stimulation (Figs. 5, 6). Ink and opaline ejections are directed to the front or back of the animal by characteristic responses of the siphon, mantle, and parapodia (Fig. 7). Ink and opaline are ejected by a series of coordinated pumping movements of the mantle, gill, and parapodia that closely resemble triggered ‘respiratory pumping’ or ‘Interneuron II’ episodes (Fig. 8; Kupfermann and Kandel 1969; Byrne and Koester 1978; Hening 1982). The directed ejection of secretions from the mantle cavity in response to noxious stimulation suggests a number of potential defensive functions for these secretions including aggressive retaliation, startle display, diversion, and alarm signalling (Edmunds 1975). Taken together, our results and others' suggest an integrated scheme for the functional organization of overt defensive behavior inAplysia (Fig. 9), and begin to suggest testable hypotheses about the integration of defensive responses on the cellular level in this animal.