Nonassociative and associative modification of head-waving produced by aversive tentacular stimuli in Aplysia.

Head-waving, a spontaneously occurring exploratory and appetitive behavior of the marine mollusc Aplysia, provides an opportunity to examine mechanisms of learning expressed in a nonreflexive behavior. The present study explores nonassociative and associative forms of learned modification of head-waving produced using an aversive stimulus as reinforcement. Experiments on intact, freely behaving animals demonstrate that training with electric shock as an aversive unconditioned stimulus, delivered unilaterally to the anterior tentacles, produces a learned shift in head-waving behavior away from the side on which shock was applied. This behavioral change is a novel learned behavioral response that is influenced by the topographic location of an aversive stimulus. Furthermore, training with application of tentacle shock reinforcement, contingent upon the animal's head position, produces operant conditioning of head-waving. Thus, anterior tentacle shock is effective as an aversive reinforcer for both nonassociative and operant learning expressed in the head-waving behavior of Aplysia.