Search

It has become clear that sleep after learning has beneficial effects on the later retrieval of newly acquired memories. The neural mechanisms underlying these effects are becoming increasingly clear as well, particularly those of non-REM sleep. However, much is still unknown about the sleep and memory relationship: the sleep state or features of sleep physiology that associate with memory performance often vary by task or experimental design, and the nature of this variability is not entirely clear. This paper describes pertinent features of sleep physiology and provides a detailed review of the scientific literature indicating beneficial effects of post-learning sleep on memory retrieval. This paper additionally introduces a hypothesis which attributes these beneficial effects of post-learning sleep to separable processes of memory reinforcement and memory refinement whereby reinforcement supports one's ability to retrieve a given memory and refinement supports the precision of that memory retrieval in the context of competitive alternatives. It is observed that features of non-REM sleep are involved in a post-learning substantiation of memory representations that benefit memory performance; thus, memory reinforcement is primarily attributed to non-REM sleep. Memory refinement is primarily attributed to REM sleep given evidence of bidirectional synaptic plasticity in REM sleep and findings from studies of selective REM sleep deprivation., Competing Interests: Conflicts of interest The authors do not have any conflicts of interest to disclose., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Electroencephalogram (EEG) alpha (8-12 Hz) asymmetries were collected from the mid-frontal and central regions during presleep wakefulness and Stage 1, Stage 2, and rapid eye movement (REM) sleep in 11 healthy right-handed participants who were free of psychiatric, neurological, and sleep problems. The authors found significant correlations between presleep wakefulness and different stages of sleep in the frontal, but not central, EEG alpha asymmetry measure. The strongest correlation was between presleep waking and REM sleep, replicating and extending relation earlier work to a normal population. The high degree of association between presleep waking and REM sleep may be a result of high cortical activation common to these states and may reflect a predisposition to different styles of emotional reactivity.

Sleep spindles are 12-14 Hz oscillations in EEG, which are thought to inhibit or 'gate' information processing. Event-related potentials may be employed to probe the extent of information processing during sleep. Previous research indicates that event-related potentials elicited by moderate intensity stimuli show increased positivity (or further removal of negativity) when stimuli are presented concurrent with spindles. However, the effectiveness of spindles to inhibit the processing of much louder stimuli remains unknown. The purpose of the present study was to investigate the extent of this gating, by using a range of stimuli including those that are loud and intrusive. Eight good sleepers were recorded during a single night. Auditory stimuli were delivered randomly at 0, 60, 80 or 100 dB SPL. Trials were sorted off-line by sleep stage, stimulus intensity and spindle characteristic (i.e. spindle absent, spindle present). During the sleep-onset period, the often-reported changes in event-related potentials were observed - N1 decreased and P2 increased in amplitude. In Stage 2 sleep, P2 was affected by the presence of spindles, particularly when stimulus intensity was loud. Its amplitude was greatest when spindles occurred following the onset of the stimulus. Scalp-recorded spindles might, therefore, be a consequence of the prior thalamic inhibition of information processing, especially when confronted by loud, intrusive external stimuli.

During NREM sleep a very large amplitude wave-form, known as the K-complex, may be elicited upon presentation of an external stimulus. The present study compared the scalp distribution of a prominent negative wave peaking at about 550 ms and a later positive wave peaking between 900 and 1300 ms in stage 2 and slow wave sleep (SWS). Nine subjects spent a single night in the laboratory. They were presented with an 80 dB SPL 2000 Hz auditory tone pip every 15 s. The EEG was recorded from 29 electrode sites and referenced to the nose. A K-complex was elicited on 34% of trials in stage 2 and on 46% of trials in SWS. A negative wave peaking at 330 ms was larger on trials in which the K-complex was elicited than on trials in which it was not. The large amplitude N550 was readily observable on trials in which the K-complex was elicited but could not be observed on trials in which it was not. The N550 was bilaterally symmetrical and was maximum over fronto-central areas of the scalp in both stage 2 and SWS. It inverted in polarity at the mastoid and inferior parietal regions. The scalp distribution of N550 significantly differed between stage 2 and SWS. It showed a sharper decline in amplitude over parietal and posterior-inferior areas of the scalp in stage 2 compared to SWS. A later P900 was maximum over centro-frontal areas of the scalp and was also bilaterally symmetrical. It showed a significantly sharper decline in amplitude over widespread inferior areas during SWS. Because the scalp maps of the N550 and P900 are different in stage 2 and SWS, their intracranial sources must also be different.

Event-related potentials (ERPs) are often used to measure the extent of information processing during sleep. Previous studies have indicated that a late positive wave, P300, can be elicited during REM sleep if stimuli are very rare and/or very loud. The present study examined the role of stimulus intensity in eliciting a P300 during REM sleep. Eight subjects were presented with auditory tone pips with an intensity of either 0, 60, 80 or 100 dB SPL. Stimuli were delivered at random with equal probability. Trials were sorted by stage of sleep, stimulus intensity, and presence or absence of rapid eye movements in REM sleep. During the waking state, when subjects read a book, the loud 100 dB stimulus elicited short (P3a) and long latency (P300) positive waves (peaking at 293 and 373 ms respectively). In stage of 2 non-REM sleep, N1 decreased to baseline level while P2 increased in amplitude compared to the waking state. A P300 could not be observed in stage 2 sleep regardless of the level of stimulus intensity. During REM sleep, a late P300 (latency 363 ms) was elicited by the 100 dB stimulus. The earlier positive peak (i.e. P3a) was not apparent. The P300 was reduced in amplitude compared to the waking state. Its amplitude did not differ between phasic and tonic states of REM sleep. A late parietal negative slow wave (SW) was also apparent during REM. Although the SW was larger during phasic compared to tonic REM, the difference was not significant. These data suggest that stimuli which are sufficiently intrusive to elicit a P300 in the waking state continue to do so in REM sleep.

Objectives: The P300 component of the auditory event-related potential (ERP) is a large amplitude positive wave peaking at approximately 300 ms following detection of a rare significant stimulus. Although P300 is typically only evoked when the subject attends to the stimulus, it may also be elicited in an awake, inattentive subject if the stimulus is sufficiently intrusive. We therefore employed an oddball task to determine if high intensity stimuli would elicit the P300 during sleep., Methods: A loud 90 dB SPL tone pip was presented infrequently (P = 0.05) in a train of lower intensity 70 dB SPL standard stimuli. A multiple channel EEG was recorded from 8 good sleepers during a single night., Results: A large amplitude parieto-central P300, peaking at 321 ms, was apparent in REM sleep to the loud deviant stimulus. In stage two non-REM sleep, a later positive wave, peaking at 446 ms, was apparent even after K-Complexes were removed from the average. This non-REM P450 was however maximum over occipito-parietal areas of the scalp., Conclusion: The presence of a P300 in REM sleep following a loud, rare stimulus indicates that sensory discrimination capabilities remain intact during this state. This may be associated with either pre- or conscious processing of relevant stimuli.