SUMMARY
Do all animals sleep? Sleep has been observed in many vertebrates, and there is a growing body of evidence for sleep-like states in arthropods and nematodes [1–5]. Here we show that sleep is also present in Cnidaria [6–8], an earlier branching metazoan lineage. Cnidaria, along with Ctenophora, are the first metazoan phyla to evolve tissue-level organization and differentiated cell types, such as neurons and muscle [9–15]. In Cnidaria, neurons are organized into a non-centralized radially symmetric nerve net [11,13,15–17] that nevertheless shares fundamental properties with the vertebrate nervous system: action potentials, synaptic transmission, neuropeptides, and neurotransmitters [15–20]. It was reported that cnidarian soft corals [21] and box jellyfish [22,23] exhibit periods of quiescence, a pre-requisite for sleep-like states, prompting us to ask if sleep is present in Cnidaria. Within Cnidaria, the upside-down jellyfish Cassiopea spp. displays a quantifiable pulsing behavior, allowing us to perform long-term behavioral tracking. Monitoring Cassiopea pulsing activity for consecutive days and nights revealed behavioral quiescence at night that is rapidly reversible, and a delayed response to stimulation in the quiescent state. When deprived of nighttime quiescence, Cassiopea exhibited decreased activity and reduced responsiveness to a sensory stimulus during the subsequent day, consistent with homeostatic regulation of the quiescent state. Together these results indicate that Cassiopea has a sleep-like state, supporting the hypothesis that sleep arose early in the metazoan lineage, prior to the emergence of a centralized nervous system.