Paradoxical Sleep - A Study of its Nature and Mechanisms
Michel Jouvet
Progress In Brain Research Vol. 18 Sleep Mechanisms 1965
TABLE OF CONTENTS
Introduction
Evidence of the duality of the states of sleep

(a) EEG and behavioural findings

(b) Phylogenetic findings

(c) Ontogenetic findings

(d) Functional findings

(e) Structural findings
Mechanisms of paradoxical sleep

(a) Producing P.S. as a reflex

(b) Results of deafferentations

(c) Role of the hypothalamus and pituitary

(d) Deprivation of P.S. in the pontile animal

(e) Effects of temperature on P.S. in the pontile animal

(f) Action of gamma-butyrolactone (G.B.L.)

(g) Osmolarity of the blood and paradoxical sleep
Discussion

(a) Duality of the states of sleep

(b) Mechanisms underlying the appearance of P.S.

Summary and Conclusions

Discussion
Figures

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Mechanisms of paradoxical sleep

(a) Producing P.S. as a reflex

The immediate triggering of P.S. has been obtained during slow sleep in intact animals by stimulation of the pontile reticular formation (M. Jouvet, 1961), mid-brain reticular formation (Rossi et al.,1961), and hippocampus (Cadilhac et al., 1961). The same phenomenon has been seen following stimulation of the pontine reticular formation in pontile and mesencephalic animals and the existence of refractory phases after phases of spontaneous or provoked P.S. was also noted (M. Jouvet, 1961). Furthermore, periods of slow sleep can be induced by low-frequency stimulation of the cutaneous or muscular nerves during wakefulness (Pompeiano and Swett, 1962a,b), whilst the same stimulation during slow sleep in intact cats only exceptionally produces P.S. (Pompeiano, 1964).

In the pontile animal, on the other hand, the P.S. reflex can regularly be obtained by proprioceptive and nociceptive stimulation. This phenomenon occurs only when certain conditions are fulfilled.

Anatomical conditions: Reflex P.S. occurs only when section of the brain stem is placed behind the origin of III. If the section is at or in front of this point in the mesencephalic animal, nociceptive and proprioceptive stimuli increase tonus and rigidity. Fig. 15 shows the most posterior (A) section to prevent the P.S. reflex;

(B) and (C) indicate the anterior and posterior limits of sections compatible with the appearance of the P.S. reflex and (D) the level of the section in retropontile animals in which no P.S. reflex could be obtained. These sections thus delimit two zones: the first, anterior to (A), at the level of the mesencephalic tegmentum, the integrity of which prevents the appearance of the P.S. reflex; these cond, between (C) and (D), at the level of the pons, which is necessary for the reflex and which encompasses the posterior part of the nucleus reticularis pontis oralis and the anterior part of the N.R.P. caudalis (based on the coordinates of the atlas of Snyder and Niemer, 1961). (It should be noted that the coordinates of the reticular formation nuclei of the pons vary from atlas to atlas. Thus, the anterior limits of the nucleus reticularis pontis caudalis are situated at P2 according to Snyder and Niemer, (1961) and at P5 according to Reinoso-Suarez (1961).)

Stimuli: P.S. can be triggered off as a reflex by the following stimuli: opening the mouth, introducing a tube into the esophagus with or without fluid, pinching the ear or fore or hind paws, flexion, extension, passive rotation of the head, passive flexion or hyperextension of the limbs. Cutaneous stimuli (stroking the back, face, stomach, limbs), and auditive stimuli, on the other hand, had no effect. Nociceptive stimuli almost always produce an immediate extension reaction of very short duration (a few seconds) and very brief apnea, and it is possible that the common denominator of all these reactions is the involvement of proprioceptive afferents.

The P.S. obtained by these stimuli appears either immediately (after 1-3 sec) or after a delay of 20-30 sec. It is accompanied by the same EEG, autonomic and behavioural (eye movements) phenomena as spontaneous P.S., and the duration of both is identical. A 'refractory period' of 10-20 min follows spontaneous or evoked P.S., during which the same stimuli either are totally ineffective or produce phasic suppression or, more rarely, tonic suppression of the EMG (cataplexy), which can last for 1 or 2 min. But then no monophasic spikes occur in the pons and there are no eye movements (Fig. 16). Such cataplexic periods cannot therefore be identified with P.S. The duration of the refractory phases following nociceptive and proprioceptive stimulation was identical. It was shorter than the mean interval occurring between two phases of spontaneous P.S. Finally intravenous injection of 1-5 microg/kg adrenalin never caused a P.S. reflex in these animals.

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