Behavioural evidence for a sleep-like quiescent state in a pulmonate mollusc, Lymnaea stagnalis (Linnaeus)

The objective of this study was to determine whether the great pond snail, Lymnaea stagnalis, expresses a sleep-like behavioural state. We found that snails spontaneously enter a relatively brief (22±1 min) quiescent state characterized by postural relaxation of the foot, mantle and tentacles, and cessation of radula rasping. Quiescence was reversed ('aroused') by appetitive (sucrose solution) and aversive (tactile) stimuli. Responsiveness to both stimuli was significantly lower in quiescent snails than in active snails. However, tactile stimuli evoked a more sustained defensive response in quiescent snails. Quiescence bouts were consolidated into 'clusters' over an infradian timescale and were only weakly affected by time of day. Clusters contained 7±0.5 bouts, lasted 13±1 h and were separated by long (37±4 h) intervals of almost continuous activity. Analysis of Kaplan-Meier survival curves revealed that the quiescent bout duration was described by an exponential probability distribution (time constant 15±1 min). Active bout duration was described by a bi-exponential probability distribution (time constants 62±4 and 592±48 min). We found no evidence for a 'sleep rebound' mechanism and quiescence expression appeared to be regulated through stochastic processes causing state transitions to resemble a Markovian random walk. We conclude that Lymnaea is a potentially valuable model system for studies of cellular function in sleep.