Defensive behaviour in the pond snail, Lymnaea stagnalis: the whole body withdrawal associated with exsanguination

The freshwater snail L. stagnalis is known to be able to respond to a strong, noxious stimulus with a full retraction of the foot and head into the shell accompanied with expelling the blood through the hemal pore. We have found that this behavioural response, besides graded local retractions, can be triggered by mild tactile stimulation provided that the stimulus is repeated several times. Only a complete exsanguination could be obtained, indicating that it is an all-or-none defensive behaviour. In an electrophysiological investigation of isolated brain, a number of similarities were found between this all-or-none behaviour and the so-called input 3 to central neurons, as described by Benjamin and Winlow. These include ability to be selectively activated by high calcium solutions, and blocked by keeping the snails in a spoiled water. Injection of snails with naloxone (0.5-2.0/micrograms/g) or ergotamine (0.4/microgram/g) blocked selectively the whole body withdrawal induced by tactile stimulation, but not that induced by injection of a high calcium saline or acetylcholine solution, indicating that enkephalinergic and/or dopaminergic mechanosensory neurons might be involved. The consideration of available data has led to a working hypothesis that the activity of input 3 might be the neurophysiological correlate of the high threshold all-or-none whole body withdrawal associated with exsanguination.