Rapid eye movement (REM) sleep is a peculiar neural state that occupies 20-25% of nighttime sleep in healthy human adults and seems to play critical roles in a variety of functions spanning from basic physiological mechanisms to complex cognitive processes. REM sleep exhibits a plethora of transient neurophysiological features, such as eye movements, muscle twitches, and changes in autonomic activity, however, despite its heterogeneous nature, it is usually conceptualized as a homogeneous sleep state. We propose here that differentiating and exploring the fine microstructure of REM sleep, especially its phasic and tonic constituents would provide a novel framework to examine the mechanisms and putative functions of REM sleep. In this review, we show that phasic and tonic REM periods are remarkably different neural states with respect to environmental alertness, spontaneous and evoked cortical activity, information processing, and seem to contribute differently to the dysfunctions of REM sleep in several neurological and psychiatric disorders. We highlight that a distinctive view on phasic and tonic REM microstates would facilitate the understanding of the mechanisms and functions of REM sleep in healthy and pathological conditions.

A great body of research indicates that eveningness is associated with negative psychological outcomes, including depressive and anxiety symptoms, behavioral dyscontrol and different health impairing behaviors. Impaired subjective sleep quality, increased circadian misalignment and daytime sleepiness were also reported in evening-type individuals in comparison with morning-types. Although sleep problems were consistently reported to be associated with poor psychological functioning, the effects of sleep disruption on the relationship between eveningness preference and negative emotionality have scarcely been investigated. Here, based on questionnaire data of 756 individuals (25.5% males, age range = 18-43 years, mean = 25.3 ± 5.8 years), as well as of the evening-type (N = 211) and morning-type (N = 189) subgroups, we examined the relationship among sleep problems, eveningness and negative emotionality. Subjects completed the Hungarian Version of the Horne and Östberg Morningness-Eveningness Questionnaire (MEQ-14), The Athen Insomnia Scale (AIS) and the Epworth Sleepiness Scale (ESS). Moreover, a composite score of Negative Emotionality (NE) was computed based on the scores of the Short Beck Depression Inventory (BDI-9), the Perceived Stress Scale (PSS-4) and the General Health Questionnaire (GHQ-12). Morning and evening circadian misalignment was calculated based on the difference between preferred and real wake- and bedtimes. Two possible models were tested, hypothesizing that sleep problems (circadian misalignment, insomniac symptoms and daytime sleepiness) moderate or mediate the association between eveningness and negative emotionality. Eveningness preference was correlated with increased NE and increased AIS, ESS and circadian misalignment scores. Our results indicate that eveningness-preference is an independent risk factor for higher negative emotionality regardless of the effects of age, gender, circadian misalignment and sleep complaints. Nevertheless, while chronotype explained ∼6%, sleep problems (AIS and ESS) accounted for a much larger proportion (∼28%) of the variance of NE. We did not find a significant effect of interaction (moderation) between chronotype and sleep problems. In contrast, insomniac symptoms (AIS) emerged as a partial mediator between chronotype and NE. These findings argue against the assumption that indicators of mental health problems in evening-type individuals can be explained exclusively on the basis of disturbed sleep. Nevertheless, negative psychological outcomes seem to be partially attributable to increased severity of insomniac complaints in evening-types.

Sleep spindles are thalamocortical oscillations in nonrapid eye movement sleep, which play an important role in sleep-related neuroplasticity and offline information processing. Sleep spindle features are stable within and vary between individuals, with, for example, females having a higher number of spindles and higher spindle density than males. Sleep spindles have been associated with learning potential and intelligence; however, the details of this relationship have not been fully clarified yet. In a sample of 160 adult human subjects with a broad IQ range, we investigated the relationship between sleep spindle parameters and intelligence. In females, we found a positive age-corrected association between intelligence and fast sleep spindle amplitude in central and frontal derivations and a positive association between intelligence and slow sleep spindle duration in all except one derivation. In males, a negative association between intelligence and fast spindle density in posterior regions was found. Effects were continuous over the entire IQ range. Our results demonstrate that, although there is an association between sleep spindle parameters and intellectual performance, these effects are more modest than previously reported and mainly present in females. This supports the view that intelligence does not rely on a single neural framework, and stronger neural connectivity manifesting in increased thalamocortical oscillations in sleep is one particular mechanism typical for females but not males.

Procedural learning is a fundamental cognitive function that facilitates efficient processing of and automatic responses to complex environmental stimuli. Here, we examined training-dependent and off-line changes of two sub-processes of procedural learning: namely, sequence learning and statistical learning. Whereas sequence learning requires the acquisition of order-based relationships between the elements of a sequence, statistical learning is based on the acquisition of probabilistic associations between elements. Seventy-eight healthy young adults (58 females and 20 males) completed the modified version of the Alternating Serial Reaction Time task that was designed to measure Sequence and Statistical Learning simultaneously. After training, participants were randomly assigned to one of three conditions: active wakefulness, quiet rest, or daytime sleep. We examined off-line changes in Sequence and Statistical Learning as well as further improvements after extended practice. Performance in Sequence Learning increased during training, while Statistical Learning plateaued relatively rapidly. After the off-line period, both the acquired sequence and statistical knowledge was preserved, irrespective of the vigilance state (awake, quiet rest or sleep). Sequence Learning further improved during extended practice, while Statistical Learning did not. Moreover, within the sleep group, cortical oscillations and sleep spindle parameters showed differential associations with Sequence and Statistical Learning. Our findings can contribute to a deeper understanding of the dynamic changes of multiple parallel learning and consolidation processes that occur during procedural memory formation.