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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III. State of sleep characterized by fast cortical activity paradoxical sleep

E. Relationship With Oneiric Activity in Man

The assimilation of the paradoxical phase of sleep in the cat with the oneiric activity phase (REM phase) in man has raised many arguments. Today, if we compare the paradoxical phase of sleep in the cat and the REM phase in normal man (24, 121, 126, 127, 168, 233, 249, 257, 377) or in patients suffering from various cerebral lesions (255, 353), there is no doubt about the similarity of the phenomena (241, 249, 408) (Table 2) and that there is "a need" (first elicited in man) "for oneiric activity" appearing during selective deprivation experiments (123, 125). The demonstration of the relationship between oneiric activity and REM sleep in man allows us to speculate that mechanisms and structures, progressively elicited in the cat during PS, will enable us to understand eventually the causes and functions of the oneiric state. Moreover, subjective data given by subjects awakened when dreaming, immediately after REM, seem to reveal a close relationship between the direction of ocular movements and the oneiric scenery ( 124, 377) . It then appears possible that postsynaptic geniculate and occipItal. events originating in the pons (described in the cat) may be responsible for oneiric imagery in man. Numerous questions are still to be settled before explaining the relationship of REM to this imagery. As a matter of fact, as in the cat, rapid eye movements in man during sleep have patterns and speeds different from those of visual observation, but resembling those of ocular movements during attempts to remember events (228); on the other hand, PS ocular movements persist in subjects suffering from a decortication syndrome, unable to have ocular movements during wakefulness (249, 255). Moreover, REM occur during sleep in newborn infants (376) and in blind-born adults (43, 345), who cannot have visual imagery during dreaming. If phasic electrical and oculomotor phenomena both appear to be triggered from the pontine area, the precise determination of the temporal relationship existing between ponto-occipItal. spikes and REM is essential in order to understand the integrating mechanisms responsible for oneiric imagery.

TABLE 2. Similarities between paradoxical sleep in the cat and REM sleep in humans

EEG Cat (adult) Man (Adult)
Cortical activity Fast, low voltage (similar to arousal) Scalp recording: low-voltage, 6-8 cycles/sec in occipItal. areas
Evoked responses by visual or auditory stimuli Decreased Average response decreased

Physiological behavior Cat (adult) Man (adult)
EMG Activity EMG Activity of the neck totally decreased Subhyoidian muscle activity totally decreased
Monosynaptic reflexes Decreased or abolished Decreased or abolished (H-reflexes)
Ocular movements Present Present
Pupillary diameter Myosis  
Heart rate Mostly decreased, irregularity Mostly increased, irregularity
Blood pressure Decreased Increased but different method
Respiration Irregularity Irregularity
GSR Decreased Decreased

  Cat (adult) Man (adult)
Functionnal aspects - -
Arousal threshold - -
Auditory stimulation Increased Superior or equal to stage III equal to stage IV
Periodicity 20-28 % of total sleep time, always follows slow sleep 20-25 % of total sleep time, always follows slow sleep
Result of selective deprivation Increased during recovery Increased during recovery
Structural aspects Persist after decortication Suppressed by pontine lesions Persist in case of decortication Suppressed by brain-stem lesion involving the pons
Ontogenesis Present immediately after birth, may directly follow waking Present immediately after birth, may directly follow waking
Subjective experience Unknown Dreaming

From Dement, W. C., and M. Jouvet. General discussion. In Aspects anatomo-fonctionnels de la physiologie du sommeil, edited by M. Jouvet. Paris Centre Natl. Rech. Sci., 1965.

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