RESUMEN
Aerial respiration in the pond snail Lymnaea stagnalis can be operantly conditioned. Depending on the specific training procedure used (i.e. a 0.5 h vs a 1.0 h interval between training sessions) either intermediate (ITM) or long-term memory (LTM) is formed. ITM, which persists for 2-3 h, is dependent only on de novo protein synthesis, whilst LTM persists for up to 4 weeks and is dependent on both transcription and de novo protein synthesis. We found that although the behavioural phenotype of ITM was not apparent 24 h after the last training session, a residual memory trace was present that serves as a foundation upon which a subsequent ITM-training-procedure builds on to form LTM (i.e. a "changed memory"). This residual memory trace could be perturbed by cooling, the behavioural process of context-specific extinction and by increasing the interval between the training procedures. Furthermore in preparations where the somata of RPeD1 (one of three interneurons in the central pattern generator required for aerial respiratory behavior) had been ablated before training, LTM could not be observed following a second bout of ITM-training. These data support the concept that a molecular memory trace is established as a consequence of ITM-training, which serves as a "permissive substrate", when the ITM memory is made active, sufficient to permit the necessary transcription and translation processes that are causal for LTM formation.