Whereas respiratory rhythm, heart rate, and blood pressure tend to be basal in NREM sleep, they display greater activity and greater variability during REM phases (3-5, 11, 12, 31). The fine muscles of the face and extremities contract frequently, though there are few gross body displacements (14). In the absence of movement, however, muscle tone measured from the head and neck virtually disappears in REM periods (20, 32). The extrinsic ocular muscles are an exception to this rule. Before and during shifts of eyeball position, tone may be sustained in uninvolved and antagonist muscles (33). Penile erections are specific to the REM periods, detumescence occurring as NREM sleep ensues (34). Basal skin resistance, which should fall with heightened arousal, has been reported to rise in REM sleep by some investigators (35) but not by others (11, 30). In view of the eye activity and oneiric phenomena during REM sleep, it is intriguing that the REM sleep EEG is remarkably similar to that of a subject awake under circumstances of visual imaging or stimulation, when alpha activity is blocked. Furthermore, cortical responses evoked during the waking state are extremely similar to those evoked in REM sleep (36).An unexpected finding has been that motor-response and arousal thresholds are no higher in deep NREM sleep than in REM sleep (5, 37). This seems to fit with the finding in cats that during REM sleep there is a high response threshold in the mesencephalic reticular formation to auditory (38) as well as to direct stimulation (39, 40). In spite of the lowered responsiveness to stimulation, however, there is greater spontaneous activity in the reticular formation during REM sleep (39). (It is precisely this aspect of brain functioning in REM sleep which renders "depth of sleep" so difficult to designate.) Huttenlocher has speculated that in REM sleep evoked responses may be occluded because of this high level of spontaneous activity. Recently, however, Adey et al. (41) were not able to demonstrate higher response thresholds in the mesencephalic reticular formations of chimpanzees during REM sleep.
Studies in AnimalsA dual neurophysiological organization of sleep is not specific to human beings. Every species of mammal so 29 APRIL 1966 far studied exhibits rhythmically alternating periods of REM and NREM activity which are marked by vegetative alterations similar in most respects to those that are found in man (12, 41-52).Moreover, animal experimentation has greatly extended our knowledge of the active processes occurring in the central nervous system during the REM state, such as: increase in blood flow to the cortex (47); rise in brain temperature (52); elevation in frequency of spontaneous neuronal firing in the MRF (39), medial and descending vestibular nuclei (53), pyramidal tract (54, 55), and occipital cortex (56); development of monophasic wave aggregates in the pons, lateral geniculate body, and other subcortical areas; continuous theta activity in the hippocampus (even more regular than during arousal) (9, 51...