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

(b) Results of deafferentations

The possibility that a reflex mechanism with a peripheral point of departure is hlvolved in the production of a P.S. reflex should not be excluded a priori, since P.S. can be obtained by proprioceptive stimulation in pontile animals. Furthermore, the part played by the carotid sinus has often been referred to since the work of Koch (1932). Finally, changes in blood pressure (Candia et al., 1962) and in cardiac and respiratory rates allow for the hypothesis that P.S. is induced during slow sleep by means of nervous afferents with a vascular point of departure. In order therefore to rule out the possibility of exclusive triggering of P.S. by some extracerebral nervous mechanism we systematically eliminated the majority of nervous afferents (Fig. 17).

The following operations were carried out alone or successively in 10 normal cats: total section in two stages of the sino-aortic nerve, bilateral stellectomy verified by the appearance of a bilateral Claude Bernard-Horner's syndrome, bilateral vagotomy in the neck, and intradural section of the posterior nerve roots from C1 to C6.

The results, which were in the main negative, need not be reported in detail. None of the operations carried out alone produced appreciable EEG or behavioural changes during slow sleep or P.S. The average proportion of P.S. was not significantly different postoperatively. In 2 animals, in addition, total section of the sino-aortic nerve was successively followed by bilateral stellectomy and bilateral vagotomy, carried out in several stages. No great differences in slow sleep or P.S. were observed here either. Finally, it is noteworthy that suppression of nuchal EMG activity during P.S. was absolute in the 2 animals that had undergone section of the posterior cervical roots.

Six dogs exhibited completely normal slow sleep and P.S. on the criteria of EEG, behaviour, and the duration, after the fol]owing operations: total section of the spinal cord at C6, total medullary destruction from D5 to S2 together with bilateral section of the brachial plexus, abdominal sympathectomy, bilateral splanchnicectomy together with medullo-adrenal curettage, and total thyroidectomy.

These results are represented diagrammatically in Fig. 17. As will be seen, exclusive involvement in the inception of P.S. can be ruled out for the spinal cord below C6, the cervical sympathetic chain, the sinus and aortic nerves, the vagus, and, lastly, the posterior roots from C1 to C6. Participation of medullo-adrenal and thyroid hormones can also be excluded.

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