Neurophysiology of the States of Sleep
Michel Jouvet
Physiological Reviews 47 (2) pp : 117-177 (1967)


Definitions and Abbreviations

State of Sleep Characterized by Slow Cortical Activity Slow Sleep

Behavioral aspect

Electrophysiological aspect

Structures and mechanisms responsible for slow sleep

State of Sleep Characterized by Fast Cortical Activity-Paradoxical Sleep

Behavioral aspects

Electrophysiological aspects

Structures and mechanisms responsible for paradoxical sleep

A synthesis of paradoxical sleep mechanisms

Relationship with oneiric activity in man

Phylogenesis of the States of Sleep

Ontogenesis of the States of Sleep

Relationship Between Slow Sleep and Paradoxical Sleep Unicity or Duality of Sleep Mechanisms

A Possible Monoaminergic Theory of Sleep

Figure 1

Figure 2


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1 Introduction

As long as we do not know how and why sleep forces on us a necessary and recurrent change in the process of our relations with our environment, it is impossible to give a definition of sleep that would satisfy everybody. Indeed the causes and mechanisms of sleep remain unknown despite a great amount of work. Kleitman's huge book (277) itself includes 4,377 references, and yet the latest achievements in the neurophysiology of sleep have only limited coverage. Since it is thus out of the question to undertake a general survey of the physiology of sleep, this paper is limited to a review of some particular aspects of the neurophysiology of sleep in the light of results achieved in the last 5 or 6 years. On the one hand, we must assume that our brain, like our kidneys and heart but unlike our muscular system, does not rest during sleep. On the contrary, it undergoes an active reorganization rather than a real inhibition, and so sleep seems to be an active phenomenon. On the other hand, it appears that behavioral sleep does not proceed from a single process but is the manifestation of two different states of nervous activity, though these are closely interconnected. The electrical brain activity of a sleeping mammal has a recurring evolution proceeding from two opposite modes. The first mode, which is the earliest known (and which is called slow sleep), manifests itself in the presence of a synchronized cortical activity of spindles and/or of high-voltage slow waves. The other mode reveals itself by a low-voltage fast cortical activity similar to arousal activity [activated sleep (122) or paradoxical or rhombencephalic sleep tPS) (254)]. Though it has not yet been proved that these two electric aspects of sleep are the manifestation of a single hypnogenic mechanism or the manifestation of two fundamentally different states, I shall set forth their behavioral and electric aspects successively and then discuss the mechanism of their appearance the last chapter is devoted to their respective interrelations in the light of their phylo- and ontogenetic evolution. I shall not deal in detail with the problems of vegetative, respiratory, digestive, blood, or metabolic (267) variations during sleep for they are clearly stated by Kleitman (277); this study is also limited to the nervous theories of sleep state, though humoral theories are again of topical interest (279, 319, 320).

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