Rapid eye movement sleep (REM sleep or REMS) is a unique phase of sleep in all mammals (including humans) and birds and in some reptiles, characterized by random rapid movement of the eyes, accompanied by low muscle tone throughout the body, and the propensity of the sleeper to dream vividly. The core body and brain temperatures increase during REM sleep and skin temperature decreases to the lowest values. The REM phase is also known as paradoxical sleep (PS) and sometimes desynchronized sleep or dreamy sleep, because of physiological similarities to waking states including rapid, low-voltage desynchronized brain waves. Electrical and chemical activity regulating this phase seem to originate in the brain stem, and is characterized most notably by an abundance of the neurotransmitter acetylcholine, combined with a nearly complete absence of monoamine neurotransmitters histamine, serotonin and norepinephrine. Experiences of REM sleep are not transferred to permanent memory due to absence of norepinephrine. REM sleep is physiologically different from the other phases of sleep, which are collectively referred to as non-REM sleep (NREM sleep, NREMS, synchronized sleep). The absence of visual and auditory stimulation (sensory deprivation) during REM sleep can cause hallucinations. REM and non-REM sleep alternate within one sleep cycle, which lasts about 90 minutes in adult humans. As sleep cycles continue, they shift towards a higher proportion of REM sleep. The transition to REM sleep brings marked physical changes, beginning with electrical bursts called "ponto-geniculo-occipital waves" (PGO waves) originating in the brain stem. REM sleep occurs 4 times in a 7-hour sleep. Organisms in REM sleep suspend central homeostasis, allowing large fluctuations in respiration, thermoregulation and circulation which do not occur in any other modes of sleeping or waking. The body abruptly loses muscle tone, a state known as REM atonia. In 1953, Professor Nathaniel Kleitman and his student Eugene Aserinsky defined rapid eye movement and linked it to dreams. REM sleep was further described by researchers, including William Dement and Michel Jouvet. Many experiments have involved awakening test subjects whenever they begin to enter the REM phase, thereby producing a state known as REM deprivation. Subjects allowed to sleep normally again usually experience a modest REM rebound. Techniques of neurosurgery, chemical injection, electroencephalography, positron emission tomography, and reports of dreamers upon waking have all been used to study this phase of sleep.
Physiology
Electrical activity in the brain
REM sleep is called "paradoxical" because of its similarities to wakefulness. Although the body is paralyzed, the brain acts as if it is somewhat awake, with cerebral neurons firing with the same overall intensity as in wakefulness. Electroencephalography during REM sleep reveals fast, low amplitude, desynchronized neural oscillation (brainwaves) that resemble the pattern seen during wakefulness, which differ from the slow δ (delta) waves pattern of NREM deep sleep. An important element of this contrast is the 3–10 Hz theta rhythm in the hippocampus and 40–60 Hz gamma waves in the cortex; patterns of EEG activity similar to these rhythms are also observed during wakefulness. The cortical and thalamic neurons in the waking and REM sleeping brain are more depolarized (fire more readily) than in the NREM deep sleeping brain. Human theta wave activity predominates during REM sleep in both the hippocampus and the cortex. During REM sleep, electrical connectivity among different parts of the brain manifests differently than during wakefulness. Frontal and posterior areas are less coherent in most frequencies, a fact which has been cited in relation to the chaotic experience of dreaming. However, the posterior areas are more coherent with each other; as are the right and left hemispheres of the brain, especially during lucid dreams. Brain energy use in REM sleep, as measured by oxygen and glucose metabolism, equals or exceeds energy use in waking. The rate in non-REM sleep is 11–40% lower.
Brain stem Neural activity during REM sleep seems to originate in the brain stem, especially the pontine tegmentum and locus coeruleus. REM sleep is punctuated and immediately preceded by PGO (ponto-geniculo-occipital) waves, bursts of electrical activity originating in the brain stem. (PGO waves have long been measured directly in cats but not in humans because of constraints on experimentation; however, comparable effects have been observed in humans during "phasic" events which occur during REM sleep, and the existence of similar PGO waves is thus inferred.) These waves occur in clusters about every 6 seconds for 1–2 minutes during the transition from deep to paradoxical sleep. They exhibit their highest amplitude upon moving into the visual cortex and are a cause of the "rapid eye movements" in paradoxical sleep. Other muscles may also contract under the influence of these waves.
Forebrain Research in the 1990s using positron emission tomography (PET) confirmed the role of the brain stem and suggested that, within the forebrain, the limbic and paralimbic systems showed more activation than other areas. The areas activated during REM sleep are approximately inverse to those activated during non-REM sleep and display greater activity than in quiet waking. The "anterior paralimbic REM activation area" (APRA) includes areas linked with emotion, memory, fear and sex, and may thus relate to the experience of dreaming during REMS. More recent PET research has indicated that the distribution of brain activity during REM sleep varies in correspondence with the type of activity seen in the prior period of wakefulness. The superior frontal gyrus, medial frontal areas, intraparietal sulcus, and superior parietal cortex, areas involved in sophisticated mental activity, show equal activity in REM sleep as in wakefulness. The amygdala is also active during REM sleep and may participate in generating the PGO waves, and experimental suppression of the amygdala results in less REM sleep. The amygdala may also regulate cardiac function in lieu of the less active insular cortex.
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