Inter-Hemispheric Oscillations in Human Sleep

Sleep is generally categorized into discrete stages based on characteristic electroencephalogram (EEG) patterns. This traditional approach represents sleep architecture in a static way, but it cannot reflect variations in sleep across time and across the cortex. To investigate these dynamic aspects of sleep, we analyzed sleep recordings in 14 healthy volunteers with a novel, frequency-based EEG analysis. This approach enabled comparison of sleep patterns with low inter-individual variability. We then implemented a new probability dependent, automatic classification of sleep states that agreed closely with conventional manual scoring during consolidated sleep. Furthermore, this analysis revealed a previously unrecognized, interhemispheric oscillation during rapid eye movement (REM) sleep. This quantitative approach provides a new way of examining the dynamic aspects of sleep, shedding new light on the physiology of human sleep.     

A new mathematical model for the homeostatic effects of sleep loss on neurobehavioral performance

The two-process model of sleep regulation makes accurate predictions of sleep timing and duration for a variety of experimental sleep deprivation and nap sleep scenarios. Upon extending its application to waking neurobehavioral performance, however, the model fails to predict the effects of chronic sleep restriction. Here we show that the two-process model belongs to a broader class of models formulated in terms of coupled non-homogeneous first-order ordinary differential equations, which have a dynamic repertoire capturing waking neurobehavioral functions across a wide range of wake/sleep schedules. We examine a specific case of this new model class, and demonstrate the existence of a bifurcation: for daily amounts of wakefulness less than a critical threshold, neurobehavioral performance is predicted to converge to an asymptotically stable state of equilibrium; whereas for daily wakefulness extended beyond the critical threshold, neurobehavioral performance is predicted to diverge from an unstable state of equilibrium. Comparison of model simulations to laboratory observations of lapses of attention on a psychomotor vigilance test (PVT), in experiments on the effects of chronic sleep restriction and acute total sleep deprivation, suggests that this bifurcation is an essential feature of performance impairment due to sleep loss. We present three new predictions that may be experimentally verified to validate the model. These predictions, if confirmed, challenge conventional notions about the effects of sleep and sleep loss on neurobehavioral performance. The new model class implicates a biological system analogous to two connected compartments containing interacting compounds with time-varying concentrations as being a key mechanism for the regulation of psychomotor vigilance as a function of sleep loss. We suggest that the adenosinergic neuromodulator/receptor system may provide the underlying neurobiology.     

Sleep intensity and the evolution of human cognition

Over the past four decades, scientists have made substantial progress in understanding the evolution of sleep patterns across the Tree of Life. 1 , 2 Remarkably, the specifics of sleep along the human lineage have been slow to emerge. This is surprising, given our unique mental and behavioral capacity and the importance of sleep for individual cognitive performance. 3 - 5 One view is that our species' sleep architecture is in accord with patterns documented in other mammals. 6 We promote an alternative view, that human sleep is highly derived relative to that of other primates. Based on new and existing evidence, we specifically propose that humans are more efficient in their sleep patterns than are other primates, and that human sleep is shorter, deeper, and exhibits a higher proportion of REM than expected. Thus, we propose the sleep intensity hypothesis: Early humans experienced selective pressure to fulfill sleep needs in the shortest time possible. Several factors likely served as selective pressures for more efficient sleep, including increased predation risk in terrestrial environments, threats from intergroup conflict, and benefits arising from increased social interaction. Less sleep would enable longer active periods in which to acquire and transmit new skills and knowledge, while deeper sleep may be critical for the consolidation of those skills, leading to enhanced cognitive abilities in early humans.     

Peculiarities of human sleep in the case of double alternation of "sleep-waking" cycles

The dynamics of sleep was studied in seven healthy subjects in two schedules, differing in the magnitude of the sleep intervals and their distribution over the 24 hour period. The first series provided for the use of two four-hour sleep intervals, the second six- and two-hour intervals. The investigations showed that the transition to a new schedule was accompanied by substantial qualitative and quantitative sleep disturbances, followed by a restoration of sleep as the subjects adapted. Less pronounced sleep disorders and more rapid restoration were observed in the schedule with nonuniform distribution of the intervals; more pronounced sleep disorders and a slowed restoration of sleep were observed on the schedule with two four-hour intervals.     

The maturational trajectories of NREM and REM sleep durations differ across adolescence on both school-night and extended sleep

We recorded sleep electroencephalogram longitudinally across ages 9-18 yr in subjects sleeping at home. Recordings were made twice yearly on 4 consecutive nights: 2 nights with the subjects maintaining their ongoing school-night schedules, and 2 nights with time in bed extended to 12 h. As expected, school-night total sleep time declined with age. This decline was entirely produced by decreasing non-rapid eye movement (NREM) sleep. Rapid eye movement (REM) sleep durations increased slightly but significantly. NREM and REM sleep durations also exhibited different age trajectories when sleep was extended. Both durations exceeded those on school-night schedules. However, the elevated NREM duration did not change with age, whereas REM durations increased significantly. We interpret the adolescent decline in school-night NREM duration in relation to our hypothesis that NREM sleep reverses changes produced in plastic brain systems during waking. The "substrate" produced during waking declines across adolescence, because synaptic elimination decreases the intensity (metabolic rate) of waking brain activity. Declining substrate reduces both NREM intensity (i.e., delta power) and NREM duration. The absence of a decline in REM sleep duration on school-night sleep and its age-dependent increase in extended sleep pose new challenges to understanding its physiological role. Whatever their ultimate explanation, these robust findings demonstrate that the two physiological states of human sleep respond differently to the maturational brain changes of adolescence. Understanding these differences should shed new light on both brain development and the functions of sleep.     

The Sleeping Brain's Influence on Verbal Memory: Boosting Resistance to Interference

Memories evolve. After learning something new, the brain initiates a complex set of post-learning processing that facilitates recall (i.e., consolidation). Evidence points to sleep as one of the determinants of that change. But whenever a behavioral study of episodic memory shows a benefit of sleep, critics assert that sleep only leads to a temporary shelter from the damaging effects of interference that would otherwise accrue during wakefulness. To evaluate the potentially active role of sleep for verbal memory, we compared memory recall after sleep, with and without interference before testing. We demonstrated that recall performance for verbal memory was greater after sleep than after wakefulness. And when using interference testing, that difference was even more pronounced. By introducing interference after sleep, this study confirms an experimental paradigm that demonstrates the active role of sleep in consolidating memory, and unmasks the large magnitude of that benefit.     

Longitudinal Change in Sleep and Daytime Sleepiness in Postpartum Women

Sleep disruption strongly influences daytime functioning; resultant sleepiness is recognised as a contributing risk-factor for individuals performing critical and dangerous tasks. While the relationship between sleep and sleepiness has been heavily investigated in the vulnerable sub-populations of shift workers and patients with sleep disorders, postpartum women have been comparatively overlooked. Thirty-three healthy, postpartum women recorded every episode of sleep and wake each day during postpartum weeks 6, 12 and 18. Although repeated measures analysis revealed there was no significant difference in the amount of nocturnal sleep and frequency of night-time wakings, there was a significant reduction in sleep disruption, due to fewer minutes of wake after sleep onset. Subjective sleepiness was measured each day using the Karolinska Sleepiness Scale; at the two earlier time points this was significantly correlated with sleep quality but not to sleep quantity. Epworth Sleepiness Scores significantly reduced over time; however, during week 18 over 50% of participants were still experiencing excessive daytime sleepiness (Epworth Sleepiness Score ≥12). Results have implications for health care providers and policy makers. Health care providers designing interventions to address sleepiness in new mothers should take into account the dynamic changes to sleep and sleepiness during this initial postpartum period. Policy makers developing regulations for parental leave entitlements should take into consideration the high prevalence of excessive daytime sleepiness experienced by new mothers, ensuring enough opportunity for daytime sleepiness to diminish to a manageable level prior to reengagement in the workforce.     

The cognitive neuroscience of sleep: neuronal systems,consciousness and learning

Sleep can be addressed across the entire hierarchy of biological organization. We discuss neuronal-network and regional forebrain activity during sleep, and its consequences for consciousness and cognition. Complex interactions in thalamocortical circuits maintain the electroencephalographic oscillations of non-rapid eye movement (NREM) sleep. Functional neuroimaging affords views of the human brain in both NREM and REM sleep, and has informed new concepts of the neural basis of dreaming during REM sleep -- a state that is characterized by illogic, hallucinosis and emotionality compared with waking. Replay of waking neuronal activity during sleep in the rodent hippocampus and in functional images of human brains indicates possible roles for sleep in neuroplasticity. Different forms and stages of learning and memory might benefit from different stages of sleep and be subserved by different forebrain regions.     

The ecological relevance of sleep: the trade-off between sleep,memory and energy conservation

All animals in which sleep has been studied express signs of sleep-like behaviour, suggesting that sleep must have some fundamental functions that are sustained by natural selection. Those functions, however, are still not clear. Here, we examine the ecological relevance of sleep from the perspective of behavioural trade-offs that might affect fitness. Specifically, we highlight the advantage of using food-caching animals as a system in which a conflict might occur between engaging in sleep for memory/learning and hypothermia/torpor to conserve energy. We briefly review the evidence for the importance of sleep for memory, the importance of memory for food-caching animals and the conflicts that might occur between sleep and energy conservation in these animals. We suggest that the food-caching paradigm represents a naturalistic and experimentally practical system that provides the opportunity for a new direction in sleep research that will expand our understanding of sleep, especially within the context of ecological and evolutionary processes.     

Shared and unique features of mammalian sleep spindles – insights from new and old animal models

Sleep spindles are phasic events observed in mammalian non-rapid eye movement sleep. They are relevant today in the study of memory consolidation, sleep quality, mental health and ageing. We argue that our advanced understanding of their mechanisms has not exhausted the utility and need for animal model work. This is both because some topics, like cognitive ageing, have not yet been addressed sufficiently in comparative efforts and because the evolutionary history of this oscillation is still poorly understood. Comparisons across species often are either limited to referencing the classical cat and rodent models, or are over-inclusive, uncritically including reports of sleep spindles in rarely studied animals. In this review, we discuss the emergence of new (dog and sheep) models for sleep spindles and compare the strengths and shortcomings of new and old models based on the three validation criteria for animal models – face, predictive, and construct validity. We conclude that an emphasis on cognitive ageing might dictate the future of comparative sleep spindle studies, a development that is already becoming visible in studies on dogs. Moreover, reconstructing the evolutionary history of sleep spindles will require more stringent criteria for their identification, across more species. In particular, a stronger emphasis on construct and predictive validity can help verify if spindle-like events in other species are actual sleep spindles. Work in accordance with such stricter validation suggests that sleep spindles display more universally shared features, like defining frequency, than previously thought.     

Evaluation of a fast forward rotating shift schedule in the steel industry with a special focus on ageing and sleep

The aim of this study was to explore the effect of a change in the speed and direction of shift rotation on the sleep and wakefulness of younger and older workers. A continuous three-shift schedule was changed from a slow backward rotating (EEE-MMMNNN-----) to a fast forward rotating system (MMEENN----). Sixteen subjects (mean age 42 years) were studied before and one year after the change in schedule. Two age groups were compared: ten younger men (mean age 35 years) and six older men (mean age 53 years). The effects of the new work schedule were evaluated by a questionnaire (modified SSI), and on-site registrations with an actigraph and sleep log for one shift cycle (10-15 days) before and after the new schedule. After the change in schedule, subjective sleep problems decreased and alertness increased during the morning shifts. The change in schedule influenced sleep differently in the two age-groups. Both the subjective and objective quality of sleep improved among the older workers. The results indicate that a fast forward rotating shift schedule is more suitable for older workers than a slower backward rotating system.     

Invited review: genetic dissection of sleep.

Recent advances in genomics open up new avenues in the analysis of complex behaviors such as sleep. In this analysis, the mouse is the model species of choice because it is amenable to high throughput phenotype and genotype analysis. With the use of the mouse model, unprecedented progress in our understanding of sleep physiology and the treatment of sleep disorders is awaited. This review is intended to provide an overview of available methods and techniques for genetic dissection of sleep in mice. Limits and advantages of different approaches are discussed to highlight the necessity for combining methods to avoid erroneous interpretations. The gap between understanding mechanisms of sleep and its functions may be bridged by finding its molecular bases.     

Neurophysiological characteristics of the resting forms of primary sleep and hypobiosis in the frog Rana temporaria

Neurophysiological studies have been made on natural resting forms of the primary sleep, wakefulness and hypobiosis in the frog. New method of presentation of the distribution density of EEG signal by a frequency spectrum was used. New data were presented concerning seasonal changes in the frequency parameters of one of the resting forms of the primary sleep, SLS-2. The role of this form of rest in the origin of hypobiosis in the frog Rana temporaria is discussed. Besides, new data were obtained which confirm that the resting form, SLS-3, of the primary sleep is an evolutionary precursor of the intermediate form of the sleep in reptiles and slow wave and activated phase of the sleep in warm-blooded vertebrates.     

An automatic EEG-based sleep staging system with introducing NAoSP and NAoGP as new metrics for sleep staging systems

Different biological signals are recorded in sleep labs during sleep for the diagnosis and treatment of human sleep problems. Classification of sleep stages with electroencephalography (EEG) is preferred to other biological signals due to its advantages such as providing clinical information, cost-effectiveness, comfort, and ease of use. The evaluation of EEG signals taken during sleep by clinicians is a tiring, time-consuming, and error-prone method. Therefore, it is clinically mandatory to determine sleep stages by using software-supported systems. Like all classification problems, the accuracy rate is used to compare the performance of studies in this domain, but this metric can be accurate when the number of observations is equal in classes. However, since there is not an equal number of observations in sleep stages, this metric is insufficient in the evaluation of such systems. For this purpose, in recent years, Cohen’s kappa coefficient and even the sensitivity of NREM1 have been used for comparing the performance of these systems. Still, none of them examine the system from all dimensions. Therefore, in this study, two new metrics based on the polygon area metric, called the normalized area of sensitivity polygon and normalized area of the general polygon, are proposed for the performance evaluation of sleep staging systems. In addition, a new sleep staging system is introduced using the applications offered by the MATLAB program. The existing systems discussed in the literature were examined with the proposed metrics, and the best systems were compared with the proposed sleep staging system. According to the results, the proposed system excels in comparison with the most advanced machine learning methods. The single-channel method introduced based on the proposed metrics can be used for robust and reliable sleep stage classification from all dimensions required for real-time applications.Electronic supplementary materialThe online version of this article (10.1007/s11571-020-09641-2) contains supplementary material, which is available to authorized users.     

Polygenic impact of morningness on the overnight dynamics of sleep spindle amplitude

Sleep spindles are thalamocortical oscillations that contribute to sleep maintenance and sleep‐related brain plasticity. The current study is an explorative study of the circadian dynamics of sleep spindles in relation to a polygenic score (PGS) for circadian preference towards morningness. The participants represent the 17‐year follow‐up of a birth cohort having both genome‐wide data and an ambulatory sleep electroencephalography measurement available ( N = 154, Mean age = 16.9, SD = 0.1 years, 57% girls). Based on a recent genome‐wide association study, we calculated a PGS for circadian preference towards morningness across the whole genome, including 354 single‐nucleotide polymorphisms. Stage 2 slow (9‐12.5 Hz, N = 186 739) and fast (12.5‐16 Hz, N = 135 504) sleep spindles were detected using an automated algorithm with individual time tags and amplitudes for each spindle. There was a significant interaction of PGS for morningness and timing of sleep spindles across the night. These growth curve models showed a curvilinear trajectory of spindle amplitudes: those with a higher PGS for morningness showed higher slow spindle amplitudes in frontal derivations, and a faster dissipation of spindle amplitude in central derivations. Overall, the findings provide new evidence on how individual sleep spindle trajectories are influenced by genetic factors associated with circadian type. The finding may lead to new hypotheses on the associations previously observed between circadian types, psychiatric problems and spindle activity.     

Importance of sleep for avian vocal communication

Sleep is one of the few truly ubiquitous animal behaviours, and though many animals spend enormous periods of time asleep, we have only begun to understand the consequences of sleep disturbances. In humans, sleep is crucial for effective communication. Birds are classic models for understanding the evolution and mechanisms of human language and speech. Bird vocalizations are remarkably diverse, critical, fitness-related behaviours, and the way sleep affects vocalizations is likely similarly varied. However, research on the effects of sleep disturbances on avian vocalizations is shockingly scarce. Consequently, there is a critical gap in our understanding of the extent to which sleep disturbances disrupt communication. Here, we argue that sleep disturbances are likely to affect all birds'' vocal performance by interfering with motivation, memory consolidation and vocal maintenance. Further, we suggest that quality sleep is likely essential when learning new vocalizations and that sleep disturbances will have especially strong effects on learned vocalizations. Finally, we advocate for future research to address gaps in our understanding of how sleep influences vocal learning and performance in birds.     

注意缺陷多动障碍低觉醒模型的新证据

注意缺陷多动障碍（Attention Deficit Hyperactivity Disorder,ADHD）是儿童期常见的一种发展性的异常,其病因及发生机理至今未明。低觉醒模型是ADHD成因的一种假设。本文从睡眠障碍导致的低觉醒探讨ADHD发生机理。通过对ADHD儿童的睡眠障碍进行分析以及将ADHD外在表现与睡眠剥夺后的表现进行对比分析,得出ADHD儿童存在的低觉醒是由于外显的或内隐的睡眠障碍引起的,一方面间接证明了低觉醒模型,另一方面为ADHD的成因研究开拓了新的思路。     

Multilevel modeling of chronotype and weekdays versus weekends to predict nonrestorative sleep

Nonrestorative sleep, a form of subjective sleep disturbance that has been largely neglected in the literature, is newly accessible to researchers via the validated restorative sleep questionnaire (RSQ). The daily version of the RSQ allows for analysis of within-subjects variation in restorative sleep across repeated samplings, and such day-to-day regularity in sleep variables has been highlighted as an important new direction for research. The present study used a sophisticated statistical approach, multilevel modeling, to examine the contributions of circadian chronotype, calendar day of questionnaire completion (weekends versus weekdays), and their interaction in explaining both interindividual and intraindividual variance in restorative sleep. Analyses were conducted using an archival dataset of college undergraduates who continuously completed daily RSQs over a 14-day sampling period. In the final multilevel model, possessing an evening type predicted lower restorative sleep between subjects, while sampling on weekdays predicted lower restorative sleep within subjects. Furthermore, a cross-level interaction was observed, such that the difference in restorative sleep on weekends versus weekdays was more pronounced among those with greater evening circadian preference. All of the effects were maintained after accounting for the significant influence of gender (women had less restorative sleep than men). These results are theoretically consistent with findings that evening types display stronger disparities in sleep schedules across free and workdays (i.e., social jet lag), and attest to the usefulness of multilevel models for statistically investigating how stable traits interact with factors that vary day to day (e.g., work or school schedules) in influencing sleep outcomes.     

Sleep and dreaming: induction and mediation of REM sleep by cholinergic mechanisms

The most important recent work on the neurobiology of sleep has focused on the precise cellular and biochemical mechanisms of rapid eye movement sleep mediation. Direct and indirect evidence implicates acetylcholine-containing neurons in the peribrachial pons as critical in the triggering and maintenance of rapid eye movement sleep. Other new studies provide support for the hypothesis that the cholinergic generator system is gated during waking by serotonergic and noradrenergic influences. A growing consensus regarding the basic neurobiology has stimulated new thinking about the brain basis of consciousness during waking and dreaming.