A probabilistic model for the ultradian timing of REM sleep in mice

A salient feature of mammalian sleep is the alternation between rapid eye movement (REM) and non-REM (NREM) sleep. However, how these two sleep stages influence each other and thereby regulate the timing of REM sleep episodes is still largely unresolved. Here, we developed a statistical model that specifies the relationship between REM and subsequent NREM sleep to quantify how REM sleep affects the following NREM sleep duration and its electrophysiological features in mice. We show that a lognormal mixture model well describes how the preceding REM sleep duration influences the amount of NREM sleep till the next REM sleep episode. The model supports the existence of two different types of sleep cycles: Short cycles form closely interspaced sequences of REM sleep episodes, whereas during long cycles, REM sleep is first followed by an interval of NREM sleep during which transitions to REM sleep are extremely unlikely. This refractory period is characterized by low power in the theta and sigma range of the electroencephalogram (EEG), low spindle rate and frequent microarousals, and its duration proportionally increases with the preceding REM sleep duration. Using our model, we estimated the propensity for REM sleep at the transition from NREM to REM sleep and found that entering REM sleep with higher propensity resulted in longer REM sleep episodes with reduced EEG power. Compared with the light phase, the buildup of REM sleep propensity was slower during the dark phase. Our data-driven modeling approach uncovered basic principles underlying the timing and duration of REM sleep episodes in mice and provides a flexible framework to describe the ultradian regulation of REM sleep in health and disease.     

Some indices of the activity of the brain in "rapid" sleep preceeded or not preceeded by delta-sleep]

In order to study the functional interaction between the delta sleep and the REM sleep some psychophysiological features of REM sleep were examined in REM-onset (without any preceding delta sleep--"early REM period") and in the REM period (REMP) terminating the normal sleep cycle (with the preceding delta sleep) of 92 daytime sleep attacks in 10 narcoleptic patients. Under these conditions the significant differences exist in the characteristics of the dream reports and in subjective estimations of sleep quality and duration. Sleep was evaluated as "superficial" and underestimations of sleep duration took place after an early REMP. Correct estimations of sleep duration and evaluations of sleep as "deep" dominated after REMP enging sleep cycles. The results obtained indicate the functional interaction between the delta sleep and REM sleep existing in the sleep cycle and largely determining the psychic content of the brain activity in the REM sleep.     

Effects of essential oil inhalation on objective and subjective sleep quality in healthy university students

Aromatherapy with essential oils is one of the most popular complementary medical tools for improving sleep quality. However, only a few reports have objectively measured the effects of essential oils on sleep. Here, we used objective and subjective measures to analyze the effects of the essential oils of lavender (Lavandula angustifolia) and sweet orange (Citrus sinensis) on the sleep quality of healthy university students. The participants were monitored for 15 consecutive nights as they inhaled lavender oil and sweet orange oil, in a crossover design. Their sleep was monitored objectively by actigraphy, and total sleep time (TST), sleep efficiency, sleep latency, and wake after sleep onset (WASO) were analyzed. Their sleep was analyzed subjectively using Oguri–Shirakawa–Azumi (OSA) sleep inventory scores. Inhalation of an essential oil improved sleep measures only in participant whose sleep quality was poor in the control condition. Lavender seemed more effective than sweet orange in objective measures, especially in improving sleep latency. In the subjective sleep analysis, the essential oils improved sleep maintenance, dreaming, and sleep length in subjects who had poor sleep quality. Sweet orange seemed more effective than lavender in the subjective sleep measures. The difference between the two oils suggests that expectancy bias had little effect on the hypnotic effect of lavender on objective sleep. Although no obvious effect was observed in good sleepers, the inhalation of lavender oil could be effective for helping poor sleepers improve objective sleep quality.

     

Effects of thermal environment on sleep and circadian rhythm

ABSTRACT: The thermal environment is one of the most important factors that can affect human sleep. The stereotypical effects of heat or cold exposure are increased wakefulness and decreased rapid eye movement sleep and slow wave sleep. These effects of the thermal environment on sleep stages are strongly linked to thermoregulation, which affects the mechanism regulating sleep. The effects on sleep stages also differ depending on the use of bedding and/or clothing. In semi-nude subjects, sleep stages are more affected by cold exposure than heat exposure. In real-life situations where bedding and clothing are used, heat exposure increases wakefulness and decreases slow wave sleep and rapid eye movement sleep. Humid heat exposure further increases thermal load during sleep and affects sleep stages and thermoregulation. On the other hand, cold exposure does not affect sleep stages, though the use of beddings and clothing during sleep is critical in supporting thermoregulation and sleep in cold exposure. However, cold exposure affects cardiac autonomic response during sleep without affecting sleep stages and subjective sensations. These results indicate that the impact of cold exposure may be greater than that of heat exposure in real-life situations; thus, further studies are warranted that consider the effect of cold exposure on sleep and other physiological parameters.     

Comparing Sleep‐Loss Sleepiness and Sleep Inertia: Lapses Make the Difference

To compare the behavioral effects of sleep‐loss sleepiness (performance impairment due to sleep loss) and sleep inertia (period of impaired performance that follows awakening), mean response latencies and number of lapses from a visual simple reaction‐time task were analyzed. Three experimental conditions were designed to manipulate sleepiness and sleep‐inertia levels: uninterrupted sleep, partial sleep reduction, and total sleep deprivation. Each condition included two consecutive nights (the first always a night of uninterrupted sleep, and the second either a night of uninterrupted sleep, a night when sleep was reduced to 3 h, or a night of total sleep deprivation), as well as two days in which performance was assessed at 10 different time points (08:00, 08:30, 09:00, 09:30, 10:00, 11:00, 14:00, 17:00, 20:00, and 23:00 h). From 08:00 to 09:00 h, reaction times in the partial sleep‐reduction and total sleep‐deprivation conditions were at a similar level and were slower than those observed in the uninterrupted sleep condition. In the same time period, the frequency of lapses in the total sleep‐deprivation condition was higher than in the partial sleep‐reduction condition, while this latter condition never differed from the uninterrupted sleep condition. The results indicate that both sleep inertia and sleep‐loss sleepiness lead to an increase in response latencies, but only extreme sleepiness leads to an increase in lapse frequency. We conclude that while reaction times slow as a result of both sleep inertia and sleep‐loss sleepiness, lapses appear to be a specific feature of sleep‐loss sleepiness.     

Increased food intake after starvation enhances sleep in Drosophila melanogaster

Feeding and sleep are highly conserved,interconnected behaviors essential for survival.Starvation has been shown to potently suppress sleep across species;however,whether satiety promotes sleep is still unclear.Here we use the fruit fly,Drosophila melanogaster,as a model organism to address the interaction between feeding and sleep.We first monitored the sleep of flies that had been starved for 24 h and found that sleep amount increased in the first 4 h after flies were given food.Increased sleep after starvation was due to an increase in sleep bout number and average sleep bout length.Mutants of translin or adipokinetic hormone,which fail to suppress sleep during starvation,still exhibited a sleep increase after starvation,suggesting that sleep increase after starvation is not a consequence of sleep loss during starvation.We also found that feeding activity and food consumption were higher in the first 10-30 min after starvation.Restricting food consumption in starved flies to 30 min was sufficient to increase sleep for 1 h.Although flies ingested a comparable amount of food at differing sucrose concentrations,sleep increase after starvation on a lower sucrose concentration was undetectable.Taken together,our results suggest that increased food intake after starvation enhances sleep and reveals a novel relationship between feeding and sleep.     

Variability of Sleep Duration Is Related to Subjective Sleep Quality and Subjective Well-Being: An Actigraphy Study

While there is a large body of evidence that poor subjective sleep quality is related to lower subjective well-being, studies on the relation of objective sleep measures and subjective well-being are fewer in number and less consistent in their findings. Using data of the Survey of Mid-Life in the United States (MIDUS), we investigated whether duration and quality of sleep, assessed by actigraphy, were related to subjective well-being and whether this relationship was mediated by subjective sleep quality. Three hundred and thirteen mainly white American individuals from the general population and 128 urban-dwelling African American individuals between 35 and 85 years of age were studied cross-sectionally. Sleep duration, variability of sleep duration, sleep onset latency, and time awake after sleep onset were assessed by actigraphy over a period of 7 days. Subjective sleep quality was assessed with the Pittsburgh Sleep Quality Index, positive psychological well-being and symptoms of psychological distress were assessed with the Satisfaction with Life Scale and the Mood and Anxiety Symptom Questionnaire. In both white and African Americans high day-to-day variability in sleep duration was related to lower levels of subjective well-being controlling age, gender, educational and marital status, and BMI. By contrast, sleep duration, sleep onset latency, and time awake after sleep onset were not related to subjective well-being controlling covariates and other sleep variables. Moreover, the relationship between variability in sleep duration and well-being was partially mediated by subjective sleep quality. The findings show that great day-to-day variability in sleep duration – more than average sleep duration – is related to poor subjective sleep quality and poor subjective well-being.     

Extended nights, sleep loss, and recovery sleep in adolescents

In summary, this study of sleep in adolescents on an atypical schedule of 18-hour nights showed marked but not unanticipated differences in sleep as function of prior sleep deprivation. Unanticipated was the evidence of "recovery" sleep in adolescents who not only were not sleep deprived, but who had been on a sleep "optimizing" schedule and had been awake for only 10 hours. Extended sleep beginning about 4 hours in advance of entrained sleep onset phase was not associated with a return of SWS, a finding coinciding with predictions from studies in adults. Finally, this study provides an indication that the homeostatic sleep/wake process becomes less robust or sleep responsive during adolescent development, a phenomenon that may influence the delay of sleep common in adolescents.     

基于蓝牙4.0的便携式睡眠监测仪的研制

目的:实时监测睡眠状况,从而帮助人们特别是老人找到影响睡眠的原因。方法:设计了一个低功耗便携式睡眠监测仪,它是通过加速度传感器采集腕动信号、蓝牙4.0低功耗无线传输、Micro-SD卡存储、上位机显示等实现对睡眠状态的检测。为了验证睡眠监测仪的准确性,本文采用了视频分析方法,并且对不同人群进行监测。结果:研制的睡眠监测仪具有便携低功耗等特点,能够准确监测睡眠状态。结论:睡眠监测仪的研制对使用者特别是老人帮助很大,能够帮助使用者方便适时了解自己的睡眠状况,找到影响睡眠原因和改善睡眠质量方法。     

Selective REM sleep deprivation during daytime. II. Muscle atonia in non-REM sleep

One of the hallmarks of rapid eye movement (REM) sleep is muscle atonia. Here we report extended epochs of muscle atonia in non-REM sleep (MAN). Their extent and time course was studied in a protocol that included a baseline night, a daytime sleep episode with or without selective REM sleep deprivation, and a recovery night. The distribution of the latency to the first occurrence of MAN was bimodal with a first mode shortly after sleep onset and a second mode 40 min later. Within a non-REM sleep episode, MAN showed a U-shaped distribution with the highest values before and after REM sleep. Whereas MAN was at a constant level over consecutive 2-h intervals of nighttime sleep, MAN showed high initial values when sleep began in the morning. Selective daytime REM sleep deprivation caused an initial enhancement of MAN during recovery sleep. It is concluded that episodes of MAN may represent an REM sleep equivalent and that it may be a marker of homeostatic and circadian REM sleep regulating processes. MAN episodes may contribute to the compensation of an REM sleep deficit.     

Gender and sleep in nightworkers: a quantitative analysis of sleep in days off

Differences in sleep patterns between workdays and days off contribute to shiftwork effects on workers' health and well-being. But regardless of shift schedules, female workers face more difficulties in fulfilling their sleep need because of housework. This study analyzes gender differences concerning sleep in days off by comparing sleep patterns in male and female nightworkers, analyzing sleep as related to the presence of children and testing the association of sleep features between workdays and days off. Male (n = 16) and female (n = 30) workers at a plastic plant, working from 10 p.m. to 6 a.m., on weekdays, filled sleep logs for seven consecutive weeks. Male and female samples did not differ in length of night sleep or in total length of sleep. For both samples, sleep length/day in days off increased, but the difference was lager among females. Also important were the relations between sleep in workdays and days off, specially among women. Among female workers, the results indicated that workers with children tended to sleep less in Saturday mornings, suggesting a negative effect of motherhood on sleep not restricted to workdays. The general results indicate that sleep need on the one hand, and social factors on the other determine the actual amount of sleep.     

Phylogenetic analysis of the ecology and evolution of mammalian sleep

The amount of time asleep varies greatly in mammals, from 3 h in the donkey to 20 h in the armadillo. Previous comparative studies have suggested several functional explanations for interspecific variation in both the total time spent asleep and in rapid-eye movement (REM) or "quiet" (non-REM) sleep. In support of specific functional benefits of sleep, these studies reported correlations between time in specific sleep states (NREM or REM) and brain size, metabolic rate, and developmental variables. Here we show that estimates of sleep duration are significantly influenced by the laboratory conditions under which data are collected and that, when analyses are limited to data collected under more standardized procedures, traditional functional explanations for interspecific variation in sleep durations are no longer supported. Specifically, we find that basal metabolic rate correlates negatively rather than positively with sleep quotas, and that neither adult nor neonatal brain mass correlates positively with REM or NREM sleep times. These results contradict hypotheses that invoke energy conservation, cognition, and development as drivers of sleep variation. Instead, the negative correlations of both sleep states with basal metabolic rate and diet are consistent with trade-offs between sleep and foraging time. In terms of predation risk, both REM and NREM sleep quotas are reduced when animals sleep in more exposed sites, whereas species that sleep socially sleep less. Together with the fact that REM and NREM sleep quotas correlate strongly with each other, these results suggest that variation in sleep primarily reflects ecological constraints acting on total sleep time, rather than the independent responses of each sleep state to specific selection pressures. We propose that, within this ecological framework, interspecific variation in sleep duration might be compensated by variation in the physiological intensity of sleep.     

The evolution and diversification of sleep

The evolutionary origins of sleep and its sub-states, rapid eye movement (REM) and non-REM (NREM) sleep, found in mammals and birds, remain a mystery. Although the discovery of a single type of sleep in jellyfish suggests that sleep evolved much earlier than previously thought, it is unclear when and why sleep diversified into multiple types of sleep. Intriguingly, multiple types of sleep have recently been found in animals ranging from non-avian reptiles to arthropods to cephalopods. Although there are similarities between these states and those found in mammals and birds, notable differences also exist. The diversity in the way sleep is expressed confounds attempts to trace the evolution of sleep states, but also serves as a rich resource for exploring the functions of sleep.     

The effect of memorization processes on the structural organization of natural human sleep

The effect of sleep on learning was investigated, comparing results of memorizing and reproduction of an unknown text before and after subsequent sleep, with a detailed analysis of sleep patterns. Psychological tests excluded the possibility of emotional and stress factors. Presleep learning did not influence mean values of such sleep parameters as total sleep time or duration of different sleep stages. The main finding ot the present experiment was a redistribution of stage REM during nocturnal sleep following learning--its increase in the second sleep cycle with a corresponding decrease toward the end of night. Also, individual difficulties in learning were inversely related to REM latency. Changes in sleep patterns after learning didn't influence the total number of sleep cycles. It is suggested that the REM phase of sleep might be involved in the processing of information acquired during wakefulness.     

Age-related reduction in the maximal capacity for sleep--implications for insomnia

Sleep changes markedly across the life span and complaints about insomnia are prevalent in older people [1]. Whether age-related alterations in sleep are due to modifications in social factors, circadian physiology, homeostatic drive, or the ability to sleep remains unresolved. We assessed habitual sleep duration at home and then quantified daytime sleep propensity, sleep duration, and sleep structure in an inpatient protocol that included extended sleep opportunities covering 2/3 of the circadian cycle (12 hr at night and 4 hr in the afternoon) for 3-7 days in 18 older and 35 younger healthy men and women. At baseline, older subjects had less daytime sleep propensity than did younger subjects. Total daily sleep duration, which was initially longer than habitual sleep duration, declined during the experiment to asymptotic values that were 1.5 hr shorter in older (7.4 +/- 0.4 SEM, hour) than in younger subjects (8.9 +/- 0.4). Rapid-eye-movement sleep and non-rapid-eye-movement sleep contributed about equally to this reduction. Thus, in the absence of social and circadian constraints, both daytime sleep propensity and the maximal capacity for sleep are reduced in older people. These data have important implications for understanding age-related insomnia.     

Circadian preference and sleep-wake regularity: associations with self-report sleep parameters in daytime-working adults

The aim of this study was to explore how interindividual differences in circadian type (morningness) and sleep timing regularity might be related to subjective sleep quality and quantity. Self-report circadian phase preference, sleep timing, sleep quality, and sleep duration were assessed in a sample of 62 day-working adults (33.9% male, age 23?48 yrs). The Pittsburgh Sleep Quality Index (PSQI) measured subjective sleep quality and the Sleep Timing Questionnaire (STQ) assessed habitual sleep latency and minutes awake after sleep onset. The duration, timing, and stability of sleep were assessed using the STQ separately for work-week nights (Sunday?Thursday) and for weekend nights (Friday and Saturday). Morningness-eveningness was assessed using the Composite Scale of Morningness (CSM). Daytime sleepiness was measured using the Epworth Sleepiness Scale (ESS). A morning-type orientation was associated with longer weekly sleep duration, better subjective sleep quality, and shorter sleep-onset latency. Stable weekday rise-time correlated with better self-reported sleep quality and shorter sleep-onset latency. A more regular weekend bedtime was associated with a shorter sleep latency. A more stable weekend rise-time was related to longer weekday sleep duration and lower daytime sleepiness. Increased overall regularity in rise-time was associated with better subjective sleep quality, shorter sleep-onset latency, and higher weekday sleep efficiency. Finally, a morning orientation was related to increased regularity in both bedtimes and rise-times. In conclusion, in daytime workers, a morning-type orientation and more stable sleep timing are associated with better subjective sleep quality. (Author correspondence: asoehner@berkeley.edu ).     

Age-related changes in the circadian and homeostatic regulation of human sleep

The reduction of electroencephalographic (EEG) slow-wave activity (SWA) (EEG power density between 0.75-4.5 Hz) and spindle frequency activity, together with an increase in involuntary awakenings during sleep, represent the hallmarks of human sleep alterations with age. It has been assumed that this decrease in non-rapid eye movement (NREM) sleep consolidation reflects an age-related attenuation of the sleep homeostatic drive. To test this hypothesis, we measured sleep EEG characteristics (i.e., SWA, sleep spindles) in healthy older volunteers in response to high (sleep deprivation protocol) and low sleep pressure (nap protocol) conditions. Despite the fact that the older volunteers had impaired sleep consolidation and reduced SWA levels, their relative SWA response to both high and low sleep pressure conditions was similar to that of younger persons. Only in frontal brain regions did we find an age-related diminished SWA response to high sleep pressure. On the other hand, we have clear evidence that the circadian regulation of sleep during the 40 h nap protocol was changed such that the circadian arousal signal in the evening was weaker in the older study participants. More sleep occurred during the wake maintenance zone, and subjective sleepiness ratings in the late afternoon and evening were higher than in younger participants. In addition, we found a diminished melatonin secretion and a reduced circadian modulation of REM sleep and spindle frequency-the latter was phase-advanced relative to the circadian melatonin profile. Therefore, we favor the hypothesis that age-related changes in sleep are due to weaker circadian regulation of sleep and wakefulness. Our data suggest that manipulations of the circadian timing system, rather than the sleep homeostat, may offer a potential strategy to alleviate age-related decrements in sleep and daytime alertness levels.     

Selective REM sleep deprivation during daytime I. Time course of interventions and recovery sleep

Although repeated selective rapid eye movement (REM) sleep deprivation by awakenings during nighttime has shown that the number of sleep interruptions required to prevent REM sleep increases within and across consecutive nights, the underlying regulatory processes remained unspecified. To assess the role of circadian and homeostatic factors in REM sleep regulation, REM sleep was selectively deprived in healthy young adult males during a daytime sleep episode (7-15 h) after a night without sleep. Circadian REM sleep propensity is known to be high in the early morning. The number of interventions required to prevent REM sleep increased from the first to the third 2-h interval by a factor of two and then leveled off. Only a minor REM sleep rebound (11.6%) occurred in the following undisturbed recovery night. It is concluded that the limited rise of interventions during selective daytime REM sleep deprivation may be due to the declining circadian REM sleep propensity, which may partly offset the homeostatic drive and the sleep-dependent disinhibition of REM sleep.     

Effects of selective sleep deprivation on ventilation during recovery sleep in normal humans.

To assess the effects of selective sleep loss on ventilation during recovery sleep, we deprived 10 healthy young adult humans of rapid-eye-movement (REM) sleep for 48 h and compared ventilation measured during the recovery night with that measured during the baseline night. At a later date we repeated the study using awakenings during non-rapid-eye-movement (NREM) sleep at the same frequency as in REM sleep deprivation. Neither intervention produced significant changes in average minute ventilation during presleep wakefulness, NREM sleep, or the first REM sleep period. By contrast, both interventions resulted in an increased frequency of breaths, in which ventilation was reduced below the range for tonic REM sleep, and in an increased number of longer episodes, in which ventilation was reduced during the first REM sleep period on the recovery night. The changes after REM sleep deprivation were largely due to an increase in the duration of the REM sleep period with an increase in the total phasic activity and, to a lesser extent, to changes in the relationship between ventilatory components and phasic eye movements. The changes in ventilation after partial NREM sleep deprivation were associated with more pronounced changes in the relationship between specific ventilatory components and eye movement density, whereas no change was observed in the composition of the first REM sleep period. These findings demonstrate that sleep deprivation leads to changes in ventilation during subsequent REM sleep.