Neuronal correlates of sleep, wakefulness and arousal in a diurnal insect

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Abstract
The discovery that various states of sleep, rest, wakefulness and arousal in man can be correlated with specific forms of the electroencephalogram1 has led to intensive studies of these states, mostly in mammals2-5. Today it is generally accepted that circadian sleep-wakefulness cycles occur in mammals and birds2,3,6. Behavioural observations on sleep in moths have also been published7; many other invertebrates demonstrate rest/activity cycles8. Circadian sensitivity fluctuations in both central9 and peripheral10-15 components of the visual system of various nocturnal arthropod species have been demonstrated. We now report that long-term, extracellular, single-unit recordings from optomotor interneurones in the optic lobes of forager honey bees reveal an oscillation in their sensitivity to moving visual stimuli16, 17. The oscillation displays properties typical of a circadian rhythm6, 18. The sensitivity of the neurons is higher during the subjective day than during the subjective night. The locomotor activity of individual, fixed walking forager bees shows a similar circadian oscillation and is also higher during the subjective day. Visual and mechanical stimuli can act directly on the interneurones and restore their sensitivity during times of reduced neuronal responsiveness. A comparison with results available for mammals makes it likely that the neuronal phenomena presented here are correlates of the bee's circadian sleep-wakefulness rhythm.