Shell damage leads to enhanced memory formation in Lymnaea

Ecologically relevant stressors alter the ability of the pond snail, Lymnaea stagnalis, to form long-term memory (LTM). Here, we show that an environmentally relevant stressor, shell damage, has a dramatic effect on the enhancement of LTM formation. Damage in the form of a shell clip 24 h before operant conditioning training resulted in long-term memory (LTM) formation following a single 0.5 h training session (TS). Typically, in these snails, two 0.5 h TSs with a 1 h interval between the sessions are required to cause LTM formation. We show here that even with a 72 h interval between shell clip and training, memory enhancement still occurred. The stress associated with shell clip could be mitigated by an ongoing high-Ca2+ pond water environment, an injection of propranolol and a DNA methylation blocker. However, use of an anaesthetic (MgCl2) during the clip or intermittent exposure to the high-Ca2+ pond water environment did not mitigate the stress associated with the shell clip. Shell clip was also sufficient to cause juvenile snails, which neither learn nor form memory, to gain the capacity to form LTM. Together, the experiments demonstrate that shell clipping is an environmentally relevant stressor that can cause enhancement of LTM formation.