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Activation Pattern in Lower Level in the Neo-,Paleo-,And Archicortices

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Activation Pattern in Lower Level in the Neo-,Paleo-,And Archicortices ACTIVATION PATTERN IN LOWER LEVEL IN THE NEO-,PALEO-,AND ARCHICORTICES Goro IMAMURA* AND Hiroshi KAWAMURA** Laboratory of Neurophysiology,Institute of Brain-Research, School of Medicine,University of Tokyo In the classical works of brain physiology,solely the neocortical electrical activity was recorded in the most cases,and two different patterns;low volt- age fast waves and slow waves with spindle bursts were classified as an indi- cator of the functional level of the brain.These two patterns,namely arousal pattern and sleep pattern of the neocortex respectively,were utilized to study the effects of various kinds of physiological procedures and the effects of drugs upon the brain. After the administration of some kinds of central depressants,especially narcotics(dial-urethane,pentobarbital etc.)often occurred a peculiar anesthe- tized pattern in the neocortex,and high voltage"arousal"pattern appeared by stimulation.For example,in the works of MURPHY & GELLHORN(1945)1), electrical stimulation of the posterior hypothalamus did not produce ordinary low voltage fast waves in the neocortex under dial-urethane anesthesia,and instead,4-8 cps high amplitude sinusoidal waves were elicited.SLOAN et al. (1950)2)also showed that electrical stimulation of the anterior limbic lobe elicited in some cases high voltage"arousal"pattern in the neocortical area. Similar pattern was also observed in the paleo-cortex(pyriform lobe)and its subcortical nuclei(amygdala),by MACLEAN et al.(1952)3)and described as olfactory-like responses.FEINDEL & GLOOR(1954)"also described 12-16 cps high amplitude waves as"arousal"pattern in the amygdala under pento- barbital anesthesia. In the present paper,the observation of this peculiar pattern in the neo-, paleo-,and archicortices was made and named as the activation pattern in lower level,and the conditions for appearance of this pattern were discussed. Received for publication April 22,1962. 今 村 護 郎,川 村 浩 *Present Address:Department of Psychology,Faculty of Letters,University of Tokyo **Present Address:2nd Department of Physiology,Yokohama University School of Medicine. 494 LOWER LEVEL ACTIVATION PATTERN OF EEG 495 METHODS Experiments were carried out on cats.The animals were prepared under ether anesthesia,tracheal and venal canulae were inserted.After the animal was fixed on a stereotaxic apparatus(Todai-Noken type),ether was removed and d-tubocurarine chloride was injected,and respiration was maintained artificially.Then trepanation of the skull was performed.Concentric needle electrodes,distance of the tips,0.5mm apart,were inserted stereotaxically in the amygdala,hippocampus,midbrain reticular formation,preoptic region and posterior part of the hypothalamus and some other subcortical nuclei.EEG from the neocortex was led by the silver-ball electrodes,ball diameter 1mm and polar separation being 2mm.Square wave pulses for stimulation were delivered from an electronic stimulator(Nihon-Koden MSE 3 type)and for re- cording of EEG a 12-channel ink-writing electroencephalograph(San'ei model EG 126) was used. At the end of each experiment,the brain was fixed and the positions of electrode tips were subsequently determined by histological examination.To provoke the acti- vation pattern in lower level in the electrical activity of the brain,various kinds of central depressants(narcotics:pentobarbital,hexobarbital etc.;tranquillizers:chlor- promazine,meprobamate)were used.In some cases,to elevate the activity level of the brain,central excitants(caffein,megimide,and pentylene-tetrazol)were also used. RESULTS Activation pattern in lower level in the neocortex.High voltage 10-15cps(in some cases down to 4-8cps or up to 20cps)activation pattern in lower level in the neocortex was easily elicited after the administration of a small dose of barbiturates.FIG.1 shows the effect of intravenous injection of pento- barbital(8mg/kg)upon the electrical activity of the paleocortex and neocortex. Before the injection of the drug,high frequency electrical stimulation(100 cps, 1msec,1.6volt)of the midbrain reticular formation elicited low voltage fast activity both in the anterior sigmoid gyrus and in the pyriform lobe.Olfactory bulb remained almost unchanged.Ten minutes after intravenous injection of pentobarbital,reticular stimulation with the same parameter showed the dif- ferent effrect in the EEG patterns.Thus,the pyriform lobe showed high amplitude 12cps waves during reticular stimulation,and the neocortex also occurred 13-14cps rhythmic waves of high amplitude. Similar effect on the electrical activity was seen after the administration of hexobarbital(5mg/kg,i.p.).By this dose,the activation pattern in lower level was obtained only in the neocortex,while other portions of the brain showed rather ordinary low voltage fast waves.Slow waves were reduced by pain stimulation in the amygdala,and this tendency was also observed in the caudate nucleus.The hippocampus showed regular slow arousal waves with superimposed enhanced fast waves(FIG.2). The activation pattern of high voltage 10-15cps waves was observed in the spontaneous recovery process from deep anesthesia or by addition of cen- 496 G.IMAMURA AND H.KAWAMURA BEFORE AFTER PENTOBARBITAL 8MG/KG I.V. FIG.1.•@ Activation pattern in lower level in the neocortex and paleocortex evoked by high frequency electrical stimulation of the midbrain reticular forma- tion(100 cps,1msec,1.6 volt).Upper traces:control records.Lower traces:after intravenous injection of 8mg/kg pentobarbital.High amplitude rhythmic waves were produced in the pyriform lobe(12 cps)and anterior sigmoid gyrus(13-14 cps)instead of low voltage fast activity . Abbreviations;PYR;pyriform lobe,OB;olfactory bulb,ANT .SIG;anterior sigmoid gyrus,AMY;amygdala,HIP;hipnocampus.NC:caudate nucleus .ANT. CIN;anterior cingulate gyrus,POST.CIN;posterior cingulate gvrus .ECT.SY; ectosylvian gyrus,LAT;lateral gyrus,SEP;septal nuclei . FIG.2.•@ Activation pattern in lower level in the neocortex.Noxious stimu- lation of pinching paw of the hind leg produced high voltage activation pattern solely in the neocortex(anterior sigmoid gyrus),after the intravenous administ- ration of hexobarbital(5mg/kg).Note the mixed pattern in the hippocampus with regular slow waves and intermingled fast activity.Caudate nucleus and amygdala showed low voltage fast pattern. LOWER LEVEL ACTIVATION PATTERN OF EEG 497 tral excitants(caffein or megimide)to the deeply anesthetized animal.FIG.3 shows the activation pattern in lower level produced in all areas of the brain by reticular stimulation(100 cps,1 msec.2 volts)after addition of caffein 10 mg/kg intravenously,to a cat previously administered a large dose of mepro- bamate(1g/kg,i.p.). Among tranquillizers,meprobamate easily produced this activation pattern especially in the neocortex.Frequency of this high amplitude waves decreased with lowering of the activity level of the animal(by hypothermia,administ- ration of larger dose of central depressants). Activation pattern in lower level in the archicortex.Similar 10-15 cps activation pattern of high amplitude was also observed in the hippocampus.FIG.4 shows 14 cps high amplitude rhythmic waves in the hippocampus during electrical stimulation of the midbrain reticular formation.In this case,no central depressant drug was used.This pattern more easily appeared in the hippocampus than in the neocortex and paleocortex without any drug admin- istration. FIG.5 shows the different effects of reticular and hypothalamic stimulation. upon EEG pattern of the hippocampus and neocortex.Stimulation of the mid- brain reticular formation suppressed slow waves,and produced fast activity in the neocortex,while posterior hypothalamic stimulation elicited fast activity in the hippocampus and almost no effect on the electrical activity of the neo- cortex. As was reported in the previous papers5)6),stimulation of the preoptic region desynchronized the hippocampal regular slow waves and stimulation of the posterior hypothalamus produced marked hippocampal arousal waves.After the administration of a moderate dose of barbiturate,however,both preoptic and posterior hypothalamic stimulation produced the activation pattern in lower level in the hippocampus. In the state of light anesthesia,posterior hypothalamic stimulation produced regular slow waves intermingled with fast waves,which seemed to suggest the different origin of regular slow waves and fast waves in the hippocampus. Activation pattern in lower level in the paleocortex.As was seen in FIG.1,the administration of 10mg/kg barbiturate elicited the activation pattern of high amplitude in the paleocortex and amygdala as well as in the neocortex.These waves also appeared during spontaneous awakening from deep barbiturate anesthesia or by addition of the central excitant drugs(caffein or megimide) to the deeply anesthetized animal.FIG.6 shows the marked 12 cps high am- plitude activation pattern in the paleocortex and amygdala elicited by noxious stimulation 2hours after the intraperitoneal injection of 35mg/kg pentobarbital. In this case,the neocortical area(lower three traces)showed almost no change in their pattern of electrical activity and the hippocampus showed less change than the amygdala and pyriform lobe. 498 G.IMAMURA AND H.KAW AMURA AFTER PMEPROBAMATE a CAFFEIN FIG.3.•@ High voltage activation pattern in the whole cortical and subcortical areas.Four hours after intraperitoneal injection of 1g/kg meprobamate,10mg/ kg caffein was added intravenously.Under this condition,reticular stimulation (100 cps,1 msec,2
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