Hippocampal electrical activity during waking behaviour and sleep: analyses using centrally acting drugs.

Rhythmical slow activity (RSA) occurs in the hippocampus under many conditions including waking behaviour, active sleep and surgical anaesthesia. Under all these conditions RSA, apparently, is produced by the coupled operation of CA1 and dentate gyrus generators. Two ascending brainstem systems appear capable of initiating activity in these coupled generators. One system, ascending via the diagonal band and medial septal nucleus, may contain cholinergic synapses since it is blocked by atropine and stimulated by eserine. The RSA produced by this system usually has a frequency of 4--7 Hz and can occur during total immobility during the waking state, active sleep or anaesthesia. A second ascending system produces RSA of higher frequency (usually 7--12 Hz) and is active during waking if, and only if, movements such as walking occur. During active sleep this system is active only during phasic muscular twitches. Anaesthetics (ether, urethane) and morphine abolish activity in this second system but it is resistant to atropinic and nicotinic drugs. Amphetamine stimulates, and major tranquillizers depress the atropine-resistant system but these drugs do not abolish its normal relation to behaviour. Neocortical activity appears to be controlled by two ascending systems which parallel closely those ascending to the hippocampus.