Delineating memory reactivation in sleep with verbal and non-verbal retrieval cues.

Sleep supports memory consolidation via the reactivation of newly formed memory traces. One way to investigate memory reactivation in sleep is by exposing the sleeping brain to auditory retrieval cues; a paradigm known as targeted memory reactivation. To what extent the acoustic properties of memory cues influence the effectiveness of targeted memory reactivation, however, has received limited attention. We addressed this question by exploring how verbal and non-verbal memory cues affect oscillatory activity linked to memory reactivation in sleep. Fifty-one healthy male adults learned to associate visual stimuli with spoken words (verbal cues) and environmental sounds (non-verbal cues). Subsets of the verbal and non-verbal memory cues were then replayed during sleep. The voice of the verbal cues was either matched or mismatched to learning. Memory cues (relative to unheard control cues) prompted an increase in theta/alpha and spindle power, which have been heavily implicated in sleep-associated memory processing. Moreover, verbal memory cues were associated with a stronger increase in spindle power than non-verbal memory cues. There were no significant differences between the matched and mismatched verbal cues. Our findings suggest that verbal memory cues may be most effective for triggering memory reactivation in sleep, as indicated by an amplified spindle response.

Targeted Memory Reactivation during Nonrapid Eye Movement Sleep Enhances Neutral, But Not Negative, Components of Memory.

Emotionally salient components of memory are preferentially remembered at the expense of accompanying neutral information. This emotional memory trade-off is enhanced over time, and possibly sleep, through a process of memory consolidation. Sleep is believed to benefit memory through a process of reactivation during nonrapid eye movement sleep (NREM). Here, targeted memory reactivation (TMR) was used to manipulate the reactivation of negative and neutral memories during NREM sleep. Thirty-one male and female participants encoded composite scenes containing either a negative or neutral object superimposed on an always neutral background. During NREM sleep, sounds associated with the scene object were replayed, and memory for object and background components was tested the following morning. We found that TMR during NREM sleep improved memory for neutral, but not negative scene objects. This effect was associated with sleep spindle activity, with a larger spindle response following TMR cues predicting TMR effectiveness for neutral items only. These findings therefore do not suggest a role of NREM memory reactivation in enhancing the emotional memory trade-off across a 12 h period but do align with growing evidence of spindle-mediated memory reactivation in service of neutral declarative memory.

Age-related positivity effect in emotional memory consolidation from middle age to late adulthood.

Background: While younger adults are more likely to attend to, process, and remember negative relative to positive information, healthy older adults show the opposite pattern. The current study evaluates when, exactly, this positivity shift begins, and how it influences memory performance for positive, negative, and neutral information. Methods: A total of 274 healthy early middle-aged (35-47), late middle-aged (48-59), and older adults (>59) viewed scenes consisting of a negative, positive, or a neutral object placed on a plausible neutral background, and rated each scene for its valence and arousal. After 12 h spanning a night of sleep (n = 137) or a day of wakefulness (n = 137), participants completed an unexpected memory test during which they were shown objects and backgrounds separately and indicated whether the scene component was the "same," "similar," or "new" to what they viewed during the study session. Results and conclusions: We found that both late middle-aged and older adults rated positive and neutral scenes more positively compared to early middle-aged adults. However, only older adults showed better memory for positive objects relative to negative objects, and a greater positive memory trade-off magnitude (i.e., remembering positive objects at the cost of their associated neutral backgrounds) than negative memory trade-off magnitude (i.e., remembering negative objects at the cost of their associated neutral backgrounds). Our findings suggest that while the positivity bias may not emerge in memory until older adulthood, a shift toward positivity in terms of processing may begin in middle age.

Neural reactivation during human sleep.

Sleep promotes memory consolidation: the process by which newly acquired memories are stabilised, strengthened, and integrated into long-term storage. Pioneering research in rodents has revealed that memory reactivation in sleep is a primary mechanism underpinning sleep's beneficial effect on memory. In this review, we consider evidence for memory reactivation processes occurring in human sleep. Converging lines of research support the view that memory reactivation occurs during human sleep, and is functionally relevant for consolidation. Electrophysiology studies have shown that memory reactivation is tightly coupled to the cardinal neural oscillations of non-rapid eye movement sleep, namely slow oscillation-spindle events. In addition, functional imaging studies have found that brain regions recruited during learning become reactivated during post-learning sleep. In sum, the current evidence paints a strong case for a mechanistic role of neural reactivation in promoting memory consolidation during human sleep.

NREM sleep oscillations and their relations with sleep-dependent memory consolidation in early course psychosis and first-degree relatives.

Sleep spindles are believed to mediate sleep-dependent memory consolidation, particularly when coupled to neocortical slow oscillations. Schizophrenia is characterized by a deficit in sleep spindles that correlates with reduced overnight memory consolidation. Here, we examined sleep spindle activity, slow oscillation-spindle coupling, and both motor procedural and verbal declarative memory consolidation in early course, minimally medicated psychosis patients and non-psychotic first-degree relatives. Using a four-night experimental procedure, we observed significant deficits in spindle density and amplitude in patients relative to controls that were driven by individuals with schizophrenia. Schizophrenia patients also showed reduced sleep-dependent consolidation of motor procedural memory, which correlated with spindle density. Contrary to expectations, there were no group differences in the consolidation of declarative memory on a word pairs task. Nor did the relatives of patients differ in spindle activity or memory consolidation compared with controls, however increased consistency in the timing of SO-spindle coupling were seen in both patient and relatives. Our results extend prior work by demonstrating correlated deficits in sleep spindles and sleep-dependent motor procedural memory consolidation in early course, minimally medicated patients with schizophrenia, but not in first-degree relatives. This is consistent with other work in suggesting that impaired sleep-dependent memory consolidation has some specificity for schizophrenia and is a core feature rather than reflecting the effects of medication or chronicity.

The influence of encoding strategy on associative memory consolidation across wake and sleep.

Sleep benefits memory consolidation. However, factors present at initial encoding may moderate this effect. Here, we examined the role that encoding strategy plays in subsequent memory consolidation during sleep. Eighty-nine participants encoded pairs of words using two different strategies. Each participant encoded half of the word pairs using an integrative visualization technique, where the two items were imagined in an integrated scene. The other half were encoded nonintegratively, with each word pair item visualized separately. Memory was tested before and after a period of nocturnal sleep (N = 47) or daytime wake (N = 42) via cued recall tests. Immediate memory performance was significantly better for word pairs encoded using the integrative strategy compared with the nonintegrative strategy (P < 0.001). When looking at the change in recall across the delay, there was significantly less forgetting of integrated word pairs across a night of sleep compared with a day spent awake (P < 0.001), with no significant difference in the nonintegrated pairs (P = 0.19). This finding was driven by more forgetting of integrated compared with not-integrated pairs across the wake delay (P < 0.001), whereas forgetting was equivalent across the sleep delay (P = 0.26). Together, these results show that the strategy engaged in during encoding impacts both the immediate retention of memories and their subsequent consolidation across sleep and wake intervals.

Beta spectral power during sleep is associated with impaired recall of extinguished fear.

The failure to retain memory for extinguished fear plays a major role in the maintenance of posttraumatic stress disorder (PTSD), with successful extinction recall necessary for symptom reduction. Disturbed sleep, a hallmark symptom of PTSD, impairs fear extinction recall. However, our understanding of the electrophysiological mechanisms underpinning sleep's role in extinction retention remains underdetermined. We examined the relationship between the microarchitecture of sleep and extinction recall in healthy humans (n = 71, both male and females included) and a pilot study in individuals with PTSD (n = 12). Participants underwent a fear conditioning and extinction protocol over 2 days, with sleep recording occurring between conditioning and extinction. Twenty-four hours after extinction learning, participants underwent extinction recall. Power spectral density (PSD) was computed for pre- and post-extinction learning sleep. Increased beta-band PSD (~17-26 Hz) during pre-extinction learning sleep was associated with worse extinction recall in healthy participants (r = 0.41, p = .004). Beta PSD was highly stable across three nights of sleep (intraclass correlation coefficients > 0.92). Results suggest beta-band PSD is specifically implicated in difficulties recalling extinguished fear.

Ears Wide Shut: A Complete Failure to Recognize Relevant Acoustic Stimuli.

The filtering out of apparently extraneous and redundant stimuli is critical for the effective processing of novel and relevant sensory information. But brain mechanisms that evolved to perform this function are necessarily less than perfect, in some cases failing to filter out irrelevant stimuli and in others filtering out important information. We report here on a stimulus from everyday life-the sound made by an arriving elevator, which contains information indicating the car's direction of movement-that not one of over 1,100 study participants was aware of, despite encountering this information repeatedly throughout their lives. Evidence of implicit knowledge of this information was also absent, suggesting that this valuable information is filtered out at an early stage of sensory processing.

The effects of cognitive reappraisal and sleep on emotional memory formation.

Emotion regulation (i.e. either up- or down-regulating affective responses to emotional stimuli) has been shown to modulate long-term emotional memory formation. Further, research has demonstrated that the emotional aspects of scenes are preferentially remembered relative to neutral aspects (known as the emotional memory trade-off effect). This trade-off is often enhanced when sleep follows learning, compared to an equivalent period of time spent awake. However, the interactive effects of sleep and emotion regulation on emotional memory are poorly understood. We presented 87 participants with pictures of neutral or negative objects on neutral backgrounds paired with instructions to either increase or decrease their emotional response by altering personal relevance, or to passively view the stimuli. Following a 12 h period of sleep or wakefulness, participants were tested for their memory of objects and backgrounds separately. Although we replicated the emotional memory trade-off effect, no differences in the magnitude of the trade-off effect were observed between regulation conditions. Sleep improved all aspects of memory, but it did not preferentially benefit memory for emotional components of scenes. Irrespective of a period of sleep or wake following encoding, findings suggest emotion regulation during encoding did not influence memory for emotional items at a 12-hour delay.

Associations between sleep variables and ostensibly paranormal experiences and paranormal beliefs: A scoping review.

Night-time is a period of great significance for many people who report paranormal experiences. However, there is limited understanding of the associations between sleep variables and seemingly paranormal experiences and/or beliefs. The aim of this review is to improve our understanding of these associations while unifying a currently fragmented literature-base into a structured, practical review. In this pre-registered scoping review, we searched for relevant studies in MEDLINE (PubMed), PsycINFO (EBSCO), Web of Science and EMBASE using terms related to sleep and ostensibly paranormal experiences and beliefs. Forty-four studies met all inclusion criteria. All were cross-sectional and most investigated sleep paralysis and/or lucid dreaming in relation to ostensibly paranormal experiences and paranormal beliefs. Overall, there were positive associations between many sleep variables (including sleep paralysis, lucid dreams, nightmares, and hypnagogic hallucinations) and ostensibly paranormal experiences and paranormal beliefs (including those of ghosts, spirits, and near-death experiences). The findings of this review have potential clinical implications such as reducing misdiagnosis and treatment development and provide foundations for further research. Our findings also highlight the importance of understanding why so many people report 'things that go bump in the night'.

Emotional memories are enhanced when reactivated in slow wave sleep, but impaired when reactivated in REM.

Sleep supports memory consolidation. However, it is not completely clear how different sleep stages contribute to this process. While rapid eye movement sleep (REM) has traditionally been implicated in the processing of emotionally charged material, recent studies indicate a role for slow wave sleep (SWS) in strengthening emotional memories. Here, to directly examine which sleep stage is primarily involved in emotional memory consolidation, we used targeted memory reactivation (TMR) in REM and SWS during a daytime nap. Contrary to our hypothesis, reactivation of emotional stimuli during REM led to impaired memory. Consistent with this, REM% was correlated with worse recall in the group that took a nap without TMR. Meanwhile, cueing benefit in SWS was strongly correlated with the product of times spent in REM and SWS (SWS-REM product), and reactivation significantly enhanced memory in those with high SWS-REM product. Surprisingly, SWS-REM product was associated with better memory for reactivated items and poorer memory for non-reactivated items, suggesting that sleep both preserved and eliminated emotional memories, depending on whether they were reactivated. Notably, the emotional valence of cued items modulated both sleep spindles and delta/theta power. Finally, we found that emotional memories benefited from TMR more than did neutral ones. Our results suggest that emotional memories decay during REM, unless they are reactivated during prior SWS. Furthermore, we show that active forgetting complements memory consolidation, and both take place across SWS and REM. In addition, our findings expand upon recent evidence indicating a link between sleep spindles and emotional processing.

Isolated sleep paralysis: Clinical features, perception of aetiology, prevention and disruption strategies in a large international sample.

STUDY OBJECTIVES: Isolated sleep paralysis is a benign but frightening condition characterised by a temporary inability to move at sleep onset or upon awakening. Despite the prevalence of this condition, little is known concerning its clinical features, associated demographic characteristics, and prevention as well as disruption strategies. METHODS: An online cross-sectional study was conducted. The sample comprised 3523 participants who had reported at least one lifetime episode of ISP and 3288 participants without a lifetime episode. Participants answered a survey including questions about sleep quality, sleep paralysis, and sleep paralysis prevention/disruption techniques. RESULTS: A total of 6811 participants were investigated (mean age = 46.9, SD = 15.4, age range = 18-89, 66.1% female). Those who reported experiencing ISP at least once during their lives reported longer sleep onset latencies, shorter sleep duration, and greater insomnia symptoms. Females (vs. male) and younger (vs. older) participants were more likely to experience ISP. Significant fear during episodes was reported by 76.0% of the participants. Most people (63.3%) who experienced ISP believed it to be caused by 'something in the brain'. A minority endorsed supernatural causes (7.1%). Five prevention strategies (e.g., changing sleep position, adjusting sleep patterns) with at least 60.0% effectiveness, and five disruption strategies (e.g., physical/bodily action, making noise) with varying degrees of effectiveness (ranging from 29.5 to 61.8) were identified through open-ended responses. CONCLUSIONS: ISP is associated with shorter sleep duration, longer sleep onset latency, and greater insomnia symptoms. The multiple prevention and disruption techniques identified in this study support existing treatment approaches and may inform subsequent treatment development. Implications for current diagnostic criteria are discussed.

The associations between paranormal beliefs and sleep variables.

Previous studies have found significant associations between paranormal beliefs and sleep variables. However, these have been conducted on a small scale and are limited in the number of sleep variables investigated. This study aims to fill a gap in the literature by investigating paranormal beliefs in relation to a wide range of sleep variables in a large sample. Participants (N = 8853) completed a survey initiated by the BBC Focus Magazine. They reported on their demographics, sleep disturbances and paranormal beliefs. Poorer subjective sleep quality (lower sleep efficiency, longer sleep latency, shorter sleep duration and increased insomnia symptoms) was associated with greater endorsement of belief in: (1) the soul living on after death; (2) the existence of ghosts; (3) demons; (4) an ability for some people to communicate with the dead; (5) near-death experiences are evidence for life after death; and (6) aliens have visited earth. In addition, episodes of exploding head syndrome and isolated sleep paralysis were associated with the belief that aliens have visited earth. Isolated sleep paralysis was also associated with the belief that near-death experiences are evidence for life after death. Findings obtained here indicate that there are associations between beliefs in the paranormal and various sleep variables. This information could potentially better equip us to support sleep via psychoeducation. Mechanisms underlying these associations are likely complex, and need to be further explored to fully understand why people sometimes report "things that go bump in the night".

Sleep preferentially consolidates negative aspects of human memory: Well-powered evidence from two large online experiments.

Research suggests that sleep benefits memory. Moreover, it is often claimed that sleep selectively benefits memory for emotionally salient information over neutral information. However, not all scientists are convinced by this relationship [e.g., J. M. Siegel. Curr. Sleep Med. Rep., 7, 15-18 (2021)]. One criticism of the overall sleep and memory literature-like other literature-is that many studies are underpowered and lacking in generalizability [M. J. Cordi, B. Rasch. Curr. Opin. Neurobiol., 67, 1-7 (2021)], thus leaving the evidence mixed and confusing to interpret. Because large replication studies are sorely needed, we recruited over 250 participants spanning various age ranges and backgrounds in an effort to confirm sleep's preferential emotional memory consolidation benefit using a well-established task. We found that sleep selectively benefits memory for negative emotional objects at the expense of their paired neutral backgrounds, confirming our prior work and clearly demonstrating a role for sleep in emotional memory formation. In a second experiment also using a large sample, we examined whether this effect generalized to positive emotional memory. We found that while participants demonstrated better memory for positive objects compared to their neutral backgrounds, sleep did not modulate this effect. This research provides strong support for a sleep-specific benefit on memory consolidation for specifically negative information and more broadly affirms the benefit of sleep for cognition.

How the 2020 US Presidential election impacted sleep and its relationship to public mood and alcohol consumption.

OBJECTIVES: Major sociopolitical events can influence the general public's affective state and other affect-related processes, such as sleep. Here, we investigated the extent that the 2020 US presidential election impacted sleep, public mood, and alcohol consumption. We also explored the relationship between affect and sleep changes during the peak period of election stress. PARTICIPANTS: US-residing (n = 437) and non-US-residing (n = 106) participants were recruited online for participation in the study. METHODS: A non-representative, convenience sample responded to daily assessments of their affect, sleep, and alcohol consumption during a baseline period (October 1-13, 2020) and in the days surrounding the 2020 US Election (October 30-November 12, 2020). RESULTS: Analyses determined changes within and between US and non-US participants. Election Day evoked significantly reduced sleep amount and efficiency, coupled with heightened stress, negative affect, and increased alcohol use. While US participants were significantly more impacted in a number of domains, non-US participants also reported reduced sleep and greater stress compared to baseline. Across participants, disrupted sleep on Election Night correlated with changes in emotional well-being and alcohol consumption on Election Day. CONCLUSION: These results suggest that major sociopolitical events can have global impacts on sleep that may interact with significant fluctuations in public mood and well-being. Further, while the largest impact is on the local population, these results suggest that the effects can extend beyond borders. These findings highlight the potential impact of future sociopolitical events on public well-being.

Slow oscillation-spindle coupling is negatively associated with emotional memory formation following stress.

Both stress and sleep enhance emotional memory. They also interact, with the largest effect of sleep on emotional memory being seen when stress occurs shortly before or after encoding. Slow wave sleep (SWS) is critical for long-term episodic memory, facilitated by the temporal coupling of slow oscillations and sleep spindles. Prior work in humans has shown these associations for neutral information in non-stressed participants. Whether coupling interacts with stress to facilitate emotional memory formation is unknown. Here, we addressed this question by reanalyzing an existing dataset of 64 individuals. Participants underwent a psychosocial stressor (32) or comparable control (32) prior to the encoding of 150-line drawings of neutral, positive, and negative images. All participants slept overnight with polysomnography, before being given a surprise memory test the following day. In the stress group, time spent in SWS was positively correlated with memory for images of all valences. Results were driven by those who showed a high cortisol response to the stressor, compared to low responders. The amount of slow oscillation-spindle coupling during SWS was negatively associated with neutral and emotional memory in the stress group only. The association with emotional memory was significantly stronger than for neutral memory within the stress group. These results suggest that stress around the time of initial memory formation impacts the relationship between slow wave sleep and memory.

Changes in Sleep Regularity and Perceived Life Stress across the COVID-19 Pandemic: A Longitudinal Analysis of a Predominately Female United States Convenience Sample.

The Coronavirus Disease 2019 (COVID-19) pandemic had a profound impact on sleep and psychological well-being for individuals worldwide. This pre-registered investigation extends our prior study by tracking self-reported social jetlag (SJL), social sleep restriction (SSR), and perceived life stress from May 2020 through October 2021. Using web-based surveys, we collected self-reported sleep information with the Ultrashort Munich Chronotype Questionnaire at three additional timepoints (September 2020, February 2021 and October 2021). Further, we measured perceived life stress with the Perceived Stress Scale at two additional timepoints (February 2021 and October 2021). In a subsample of 181, predominantly female (87%), United States adults aged 19-89 years, we expanded our prior findings by showing that the precipitous drop in SJL during the pandemic first wave (May 2020), compared to pre-pandemic (February, 2020), rapidly rose with loosening social restrictions (September 2020), though never returned to pre-pandemic levels. This effect was greatest in young adults, but not associated with self-reported chronotype. Further, perceived life stress decreased across the pandemic, but was unrelated to SJL or SSR. These findings suggest that sleep schedules were sensitive to pandemic-related changes in social restrictions, especially in younger participants. We posit several possible mechanisms supporting these findings.

Sleep Power Spectral Density and Spindles in PTSD and Their Relationship to Symptom Severity.

Sleep disturbances are common in post-traumatic stress disorder (PTSD), although which sleep microarchitectural characteristics reliably classify those with and without PTSD remains equivocal. Here, we investigated sleep microarchitectural differences (i.e., spectral power, spindle activity) in trauma-exposed individuals that met (n = 45) or did not meet (n = 52) criteria for PTSD and how these differences relate to post-traumatic and related psychopathological symptoms. Using ecologically-relevant home sleep polysomnography recordings, we show that individuals with PTSD exhibit decreased beta spectral power during NREM sleep and increased fast sleep spindle peak frequencies. Contrary to prior reports, spectral power in the beta frequency range (20.31-29.88 Hz) was associated with reduced PTSD symptoms, reduced depression, anxiety and stress and greater subjective ability to regulate emotions. Increased fast frequency spindle activity was not associated with individual differences in psychopathology. Our findings may suggest an adaptive role for beta power during sleep in individuals exposed to a trauma, potentially conferring resilience. Further, we add to a growing body of evidence that spindle activity may be an important biomarker for studying PTSD pathophysiology.

Sleep spectral power correlates of prospective memory maintenance.

Prospective memory involves setting an intention to act that is maintained over time and executed when appropriate. Slow wave sleep (SWS) has been implicated in maintaining prospective memories, although which SWS oscillations most benefit this memory type remains unclear. Here, we investigated SWS spectral power correlates of prospective memory. Healthy young adult participants completed three ongoing tasks in the morning or evening. They were then given the prospective memory instruction to remember to press "Q" when viewing the words "horse" or "table" when repeating the ongoing task after a 12-h delay including overnight, polysomnographically recorded sleep or continued daytime wakefulness. Spectral power analysis was performed on recorded sleep EEG. Two additional groups were tested in the morning or evening only, serving as time-of-day controls. Participants who slept demonstrated superior prospective memory compared with those who remained awake, an effect not attributable to time-of-day of testing. Contrary to prior work, prospective memory was negatively associated with SWS. Furthermore, significant increases in spectral power in the delta-theta frequency range (1.56 Hz-6.84 Hz) during SWS was observed in participants who failed to execute the prospective memory instructions. Although sleep benefits prospective memory maintenance, this benefit may be compromised if SWS is enriched with delta-theta activity.

Using EEG microstates to examine post-encoding quiet rest and subsequent word-pair memory.

Evidence suggests that the brain preferentially consolidates memories during "offline" periods, in which an individual is not performing a task and their attention is otherwise undirected, including spans of quiet, resting wakefulness. Moreover, research has demonstrated that factors such as the initial encoding strength of information influence which memories receive the greatest benefit. Recent studies have begun to investigate these periods of post-learning quiet rest using EEG microstate analysis to observe the electrical dynamics of the brain during these stretches of memory consolidation, specifically finding an increase in the amount of the canonical microstate D during a post-encoding rest period. Here, we implement an exploratory analysis to probe the activity of EEG microstates during a post-encoding session of quiet rest in order to scrutinize the impact of learning on microstate dynamics and to further understand the role these microstates play in the consolidation of memories. We examined 54 subjects (41 female) as they completed a word-pair memory task designed to use repetition to vary the initial encoding strength of the word-pair memories. In this study, we were able to replicate previous research in which there was a significant increase (p < .05) in the amount of microstate D (often associated with the dorsal attention network) during post-encoding rest. This change was accompanied by a significant decrease (p < .05) in the amount of microstate C (often associated with the default mode network). We also found preliminary evidence indicating a positive relationship between the amount of microstate D and improved memory for weakly encoded memories, which merits further exploration.