Rapid eye movement sleep

Electrical activity in the brain record of REM Sleep. EEG highlighted by red box. Eye movement highlighted by red line. 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 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. 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. 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 amygdala may also regulate cardiac function in lieu of the less active insular cortex. Two other neurotransmitters, orexin and gamma-Aminobutyric acid (GABA), seem to promote wakefulness, diminish during deep sleep, and inhibit paradoxical sleep. Unlike the abrupt transitions in electrical patterns, the chemical changes in the brain show continuous periodic oscillation. Models of REM regulation According to the activation-synthesis hypothesis proposed by Robert McCarley and Allan Hobson in 1975–1977, control over REM sleep involves pathways of "REM-on" and "REM-off" neurons in the brain stem. REM-on neurons are primarily cholinergic (i.e., involve acetylcholine); REM-off neurons activate serotonin and noradrenaline, which among other functions suppress the REM-on neurons. McCarley and Hobson suggested that the REM-on neurons actually stimulate REM-off neurons, thereby serving as the mechanism for the cycling between REM and non-REM sleep. They used Lotka–Volterra equations to describe this cyclical inverse relationship. The withdrawal of orexin and GABA may cause the absence of the other excitatory neurotransmitters; researchers in recent years increasingly include GABA regulation in their models. Eye movements Most of the eye movements in "rapid eye movement" sleep are in fact less rapid than those normally exhibited by waking humans. They are also shorter in duration and more likely to loop back to their starting point. About seven such loops take place over one minute of REM sleep. In slow-wave sleep, the eyes can drift apart; however, the eyes of the paradoxical sleeper move in tandem. but a direct relationship remains to be clearly established. Congenitally blind people, who do not typically have visual imagery in their dreams, still move their eyes in REM sleep. Circulation, respiration, and thermoregulation Generally speaking, the body suspends homeostasis during paradoxical sleep. Heart rate, cardiac pressure, cardiac output, arterial pressure, and breathing rate quickly become irregular when the body moves into REM sleep. If a male has erectile dysfunction (ED) while awake, but has NPT episodes during REM, it would suggest that the ED is from a psychological rather than a physiological cause. In females, erection of the clitoris (nocturnal clitoral tumescence or NCT) causes enlargement, with accompanying vaginal blood flow and transudation (i.e. lubrication). During a normal night of sleep, the penis and clitoris may be erect for a total time of from one hour to as long as three and a half hours during REM. Body temperature is not well regulated during REM sleep, and thus organisms become more sensitive to temperatures outside their thermoneutral zone. Cats and other small furry mammals will shiver and breathe faster to regulate temperature during NREMS—but not during REMS. their already-negative membrane potential decreases by another 2–10 millivolts, thereby raising the threshold which a stimulus must overcome to excite them. Muscle inhibition may result from unavailability of monoamine neurotransmitters (restraining the abundance of acetylcholine in the brainstem) and perhaps from mechanisms used in waking muscle inhibition. Some localized twitching and reflexes can still occur. Lack of REM atonia causes REM behavior disorder, where those affected physically act out their dreams, or conversely "dream out their acts", under an alternative theory on the relationship between muscle impulses during REM and associated mental imagery (which would also apply to people without the condition, except that commands to their muscles are suppressed). Lesions of the pons to prevent atonia have induced functional "REM behavior disorder" in animals. == Psychology ==

Dreaming Rapid eye movement sleep (REM) has since its discovery been closely associated with dreaming. Waking up sleepers during a REM phase is a common experimental method for obtaining dream reports; 80% of people can give some kind of dream report under these circumstances. (In fact these could be considered a hybrid state combining essential elements of REM sleep and waking consciousness.) Because of non-REM dreaming, some sleep researchers have strenuously contested the importance of connecting dreaming to the REM sleep phase. The prospect that well-known neurological aspects of REM do not themselves cause dreaming suggests the need to re-examine the neurobiology of dreaming per se. A study at Harvard Medical School in 2000 tested the effects of paroxetine and fluvoxamine on healthy young adult males and females for 31 days: a drug-free baseline week, 19 days on either paroxetine or fluvoxamine with morning and evening doses, and 5 days of absolute discontinuation. Results showed that SSRI treatment decreased the average amount of dream recall frequency in comparison to baseline measurements as a result of serotonergic REM suppression. Sleep aids the process by which creativity forms associative elements into new combinations that are useful or meet some requirement. This occurs in REM sleep rather than in NREM sleep. Rather than being due to memory processes, this has been attributed to changes during REM sleep in cholinergic and noradrenergic neuromodulation. This is in contrast to waking consciousness, where higher levels of norepinephrine and acetylcholine inhibit recurrent connections in the neocortex. REM sleep through this process adds creativity by allowing "neocortical structures to reorganise associative hierarchies, in which information from the hippocampus would be reinterpreted in relation to previous semantic representations or nodes." == Timing ==

(electroencephalogram of sleep) showing sleep cycles characterized by increasing paradoxical (REM) sleep In the ultradian sleep cycle, an organism alternates between deep sleep (slow, large, synchronized brain waves) and paradoxical sleep (faster, desynchronized waves). Sleep happens in the context of the larger circadian rhythm, which influences sleepiness and physiological factors based on timekeepers within the body. Sleep can be distributed throughout the day or clustered during one part of the rhythm: in nocturnal animals, during the day, and in diurnal animals, at night. REM sleep typically occupies 20–25% of total sleep in adult humans: about 90–120 minutes of a night's sleep. The first REM episode occurs about 70 minutes after falling asleep. Cycles of about 90 minutes each follow, with each cycle including a larger proportion of REM sleep. Infants spend more time in REM sleep than adults. The proportion of REM sleep then decreases significantly in childhood. Older people tend to sleep less overall, but sleep in REM for about the same absolute time (and therefore spend a greater proportion of sleep in REM). Tonic REM is characterized by theta rhythms in the brain; phasic REM is characterized by PGO waves and actual "rapid" eye movements. Processing of external stimuli is heavily inhibited during phasic REM, and recent evidence suggests that sleepers are more difficult to arouse from phasic REM than in slow-wave sleep. == Deprivation effects ==

Selective REMS deprivation causes a significant increase in the number of attempts to go into REM stage while asleep. On recovery nights, an individual will usually move to stage 3 and REM sleep more quickly and experience a REM rebound, which refers to an increase in the time spent in REM stage over normal levels. These findings are consistent with the idea that REM sleep is biologically necessary. However, the "rebound" REM sleep usually does not last fully as long as the estimated length of the missed REM periods. Higher norepinepherine is a possible cause of these results. It has been suggested that acute REM sleep deprivation can improve certain types of depression—when depression appears to be related to an imbalance of certain neurotransmitters. Although sleep deprivation in general annoys most of the population, it has repeatedly been shown to alleviate depression, albeit temporarily. More than half the individuals who experience this relief report it to be rendered ineffective after sleeping the following night. Thus, researchers have devised methods such as altering the sleep schedule for a span of days following a REM deprivation period and combining sleep-schedule alterations with pharmacotherapy to prolong this effect. Antidepressants (including selective serotonin reuptake inhibitors, tricyclics, and monoamine oxidase inhibitors) and stimulants (such as amphetamine, methylphenidate and cocaine) interfere with REM sleep by stimulating the monoamine neurotransmitters which must be suppressed for REM sleep to occur. Administered at therapeutic doses, these drugs may stop REM sleep entirely for weeks or months. Withdrawal causes a REM rebound. ==In other animals==

es sleeping, with REM and slow-wave sleep phases Although it manifests differently in different animals, REM sleep or something like it occurs in all land mammals—as well as in birds. The primary criteria used to identify REM are the change in electrical activity, measured by EEG, and loss of muscle tone, interspersed with bouts of twitching in phasic REM. The amount of REM sleep and cycling varies among animals; predators experience more REM sleep than prey. Observations of jumping spiders in their nocturnal resting position also suggest a REM sleep-like state characterized by bouts of twitching and retinal movements and hints of muscle atonia (legs curling up as a result of pressure loss caused by muscle atonia in the prosoma). Sleep deprivation experiments on non-human animals can be set up differently than those on humans. The "flower pot" method involves placing a laboratory animal above water on a platform so small that it falls off upon losing muscle tone. The naturally rude awakening which results may elicit changes in the organism which necessarily exceed the simple absence of a sleep phase. ==Possible functions==

Some researchers argue that the perpetuation of a complex brain process such as REM sleep indicates that it serves an important function for the survival of mammalian and avian species. It fulfills important physiological needs vital for survival to the extent that prolonged REM sleep deprivation leads to death in experimental animals. In both humans and experimental animals, REM sleep loss leads to several behavioral and physiological abnormalities. Loss of REM sleep has been noticed during various natural and experimental infections. Survivability of the experimental animals decreases when REM sleep is totally attenuated during infection; this leads to the possibility that the quality and quantity of REM sleep is generally essential for normal body physiology. Further, the existence of a "REM rebound" effect suggests the possibility of a biological need for REM sleep. While the precise function of REM sleep is not well understood, several theories have been proposed. Memory Sleep in general aids memory. REM sleep may favor the preservation of certain types of memories: specifically, procedural memory, spatial memory, and emotional memory. In rats, REM sleep increases following intensive learning, especially several hours after, and sometimes for multiple nights. Experimental REM sleep deprivation has sometimes inhibited memory consolidation, especially regarding complex processes (e.g., how to escape from an elaborate maze). Tucker et al. demonstrated that a daytime nap containing solely non-REM sleep enhances declarative memory—but not procedural memory. According to the sequential hypothesis, the two types of sleep work together to consolidate memory. Graeme Mitchison and Francis Crick proposed in 1983 that by virtue of its inherent spontaneous activity, the function of REM sleep "is to remove certain undesirable modes of interaction in networks of cells in the cerebral cortex"—a process they characterize as "unlearning". As a result, those memories which are relevant (whose underlying neuronal substrate is strong enough to withstand such spontaneous, chaotic activation) are further strengthened, whilst weaker, transient, "noise" memory traces disintegrate. Memory consolidation during paradoxical sleep is specifically correlated with the periods of rapid eye movement, which do not occur continuously. One explanation for this correlation is that the PGO electrical waves, which precede the eye movements, also influence memory. Sleep deprivation studies have shown that deprivation early in life can result in behavioral problems, permanent sleep disruption, and decreased brain mass. Shift of gaze According to "scanning hypothesis", the directional properties of REM sleep are related to a shift of gaze in dream imagery. Against this hypothesis is that such eye movements occur in those born blind and in fetuses in spite of lack of vision. Also, binocular REMs are non-conjugated (i.e., the two eyes do not point in the same direction at a time) and so lack a fixation point. In support of this theory, research finds that in goal-oriented dreams, eye gaze is directed towards the dream action, determined from correlations in the eye and body movements of REM sleep behavior disorder patients who enact their dreams. Oxygen supply to cornea Dr. David M. Maurice, an eye specialist and former adjunct professor at Columbia University, proposed that REM sleep was associated with oxygen supply to the cornea, and that aqueous humor, the liquid between cornea and iris, was stagnant if not stirred. Among the supportive evidence, he calculated that if aqueous humor was stagnant, oxygen from the iris had to reach the cornea by diffusion through aqueous humor, which was not sufficient. According to the theory, when the organism is awake, eye movement (or cool environmental temperature) enables the aqueous humor to circulate. When the organism is sleeping, REM provides the much needed stir to aqueous humor. This theory is consistent with the observation that fetuses, as well as eye-sealed newborn animals, spend much time in REM sleep, and that during a normal sleep, a person's REM sleep episodes become progressively longer deeper into the night. However, owls experience REM sleep, but do not move their head more than in non-REM sleep and it is well known that owls' eyes are nearly immobile. Other theories Another theory suggests that monoamine shutdown is required so that the monoamine receptors in the brain can recover to regain full sensitivity. The sentinel hypothesis of REM sleep was put forward by Frederick Snyder in 1966. It is based upon the observation that REM sleep in several mammals (the rat, the hedgehog, the rabbit, and the rhesus monkey) is followed by a brief awakening. This does not occur for either cats or humans, although humans are more likely to wake from REM sleep than from NREM sleep. Snyder hypothesized that REM sleep activates an animal periodically, to scan the environment for possible predators. This hypothesis does not explain the muscle paralysis of REM sleep; however, a logical analysis might suggest that the muscle paralysis exists to prevent the animal from fully waking up unnecessarily, and allowing it to return easily to deeper sleep. Jim Horne, a sleep researcher at Loughborough University, has suggested that REM in modern humans compensates for the reduced need for wakeful food foraging. Furthermore, eye movements are also theorized to play a role in certain psychotherapies such as eye movement desensitization and reprocessing (EMDR). == See also ==