Development of learning and memory in Aplysia. II. Habituation and dishabituation

The defensive withdrawal reflex of the mantle organs of Aplysia californica exhibits a variety of forms of both nonassociative and associative learning, which can exist in both short- and long-term forms. In addition, the reflex can be readily elicited and quantified as soon as the effector organs (siphon and gill) emerge at their respective developmental stages. Thus, this reflex system provides a useful preparation in which to study the development of learning and memory. In the present series of experiments we investigated the development of 2 forms of nonassociative learning, habituation and dishabituation, in the siphon withdrawal component of the reflex. This reflex response could be examined throughout the juvenile life of the animal (stages 9-12) since the reflex is functionally intact as soon as the siphon emerges in stage 9 (Rankin et al., 1987). We studied the development of habituation in stages 9-12 using tactile stimuli to the siphon delivered at interstimulus intervals (ISIs) of 1, 5, 10, and 30 sec. Habituation of siphon withdrawal was evident as early as juvenile stage 9. However, it existed in an immature form: Significant habituation was produced only with a very short (1 sec) ISI. No significant habituation occurred in response to 5 or 10 sec ISIs. Approximately 4 d later, in stage 10, significant habituation occurred to both 1 and 5 sec ISIs but not to a 10 sec ISI. Finally, approximately 1-2 weeks later, in stage 11, significant habituation occurred to 1, 5, and 10 sec ISIs but not to a 30 sec ISI, whereas stage 12 juveniles and adults (stage 13) readily habituated to a 30 sec ISI. Thus, there was a systematic developmental trend in the ability of the animals to habituate: Progressively older animals were capable of habituation to stimuli presented at progressively longer intervals. The systematic development of habituation was also evident by examining the amount of habituation exhibited to comparable ISIs by animals at different developmental stages. For all 4 ISIs examined, older animals showed significantly greater habituation than younger animals. Thus, our results show that habituation is present as soon as the siphon response system emerges and that it develops progressively throughout the juvenile life of the animal. Whereas habituation was present in the earliest developmental stage we examined (stage 9), dishabituation (in response to tail shock) did not emerge until 4-7 d later, in stage 10.(ABSTRACT TRUNCATED AT 400 WORDS)