Vanderwolf C H, Kramis R, Robinson T E
Ciba Found Symp. 1977(58):199-226. doi: 10.1002/9780470720394.ch10.
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.
节律性慢活动（RSA）在包括清醒行为、主动睡眠和外科麻醉在内的多种情况下，会在海马体中出现。在所有这些情况下，RSA显然是由CA1和齿状回发生器的耦合运作产生的。两个上行脑干系统似乎能够启动这些耦合发生器的活动。一个系统经斜角带和内侧隔核上行，可能含有胆碱能突触，因为它会被阿托品阻断并被毒扁豆碱刺激。这个系统产生的RSA通常频率为4 - 7赫兹，可在清醒状态下的完全不动、主动睡眠或麻醉期间出现。第二个上行系统产生较高频率（通常为7 - 12赫兹）的RSA，且仅在诸如行走等运动发生时的清醒状态下活跃。在主动睡眠期间，这个系统仅在阶段性肌肉抽搐时活跃。麻醉剂（乙醚、乌拉坦）和吗啡会消除第二个系统的活动，但它对阿托品和烟碱类药物有抗性。苯丙胺刺激、而主要的镇静剂抑制抗阿托品系统，但这些药物不会消除其与行为的正常关系。新皮层活动似乎受两个上行系统控制，这两个系统与上行至海马体的系统密切平行。
