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Sleep consolidates declarative memory after deep but not shallow incidental encoding, but little is known about this form of consolidation. One unexplored area is the extent to which the amount of exposure to incidentally encoded information affects consolidation processes. In two experiments, we manipulated the number of times information was presented. In Experiment 1, participants encoded words either one or three times in a deep or shallow incidental encoding task and completed a surprise recognition test after sleep or wake. Sleep consolidated information after deep encoding after one and three exposures, but not after shallow encoding. In Experiment 2, we explored the relationship between sleep architecture and memory after deep encoding. There was a trend for accuracy to be negatively related to N1 sleep, and reaction time to be negatively related to slow-wave sleep for words encoded once; however, the correlations did not survive corrections for multiple comparisons. These results are discussed with respect to active and passive consolidation processes., (© 2024 European Sleep Research Society.)

Sleep improves the consolidation and long-term stability of newly formed memories and associations. Most research on human declarative memory and its consolidation during sleep uses word-pair associations requiring exhaustive learning. In the present study, we present the visual paired association learning (vPAL) paradigm, in which participants learn new associations between images of celebrities and animals. The vPAL is based on a one-shot exposure that resembles learning in natural conditions. We tested if vPAL can reveal a role for sleep in memory consolidation by assessing the specificity of memory recognition, and the cued recall performance, before and after sleep. We found that a daytime nap improved the stability of recognition memory and discrimination abilities compared to identical intervals of wakefulness. By contrast, cued recall of associations did not exhibit significant sleep-dependent effects. High-density electroencephalography during naps further revealed an association between sleep spindle density and stability of recognition memory. Thus, the vPAL paradigm opens new avenues for future research on sleep and memory consolidation across ages and heterogeneous populations in health and disease., (© 2024 The Author(s). Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Experimental studies suggest that short or disrupted sleep impairs memory consolidation, mood, and perception of emotional stimuli. However, studies have chiefly relied on laboratory-based study designs and small sample sizes. The aim of this fully online and pre-registered study was to investigate the association between sleep and overnight memory consolidation, emotion perception, and affect in a large, self-selected UK sample. A total of 1646 participants (473 completed) took part in an online study, where they completed a declarative (word-pairs) memory task, emotion perception task (valence ratings of images), and rated their affect within 2 h of bed-time. The following morning, participants reported on their state affect, sleep for the previous night, completed a cued recall task for the previously presented word-pairs, rated the valence of previously viewed images, and completed a surprise recognition task. Demographic data and habitual sleep quality and duration (sleep traits) were also recorded. Habitual sleep traits were associated with immediate recall for the word-pairs task, while self-reported sleep parameters for the specific night were not associated with overnight memory consolidation. Neither habitual sleep traits, nor nightly sleep parameters were associated with unpleasantness ratings to negative stimuli or overnight habituation. Habitual poor sleep was associated with less positive and more negative affect, and morning affect was predicted by the specific night's sleep. This study suggests that overnight emotional processing and declarative memory may not be associated with self-reported sleep across individuals. More work is needed to understand how findings from laboratory-based studies extrapolate to real-world samples and contexts., (© 2023 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Infants face the constant challenge of selecting information for encoding and storage from a continuous incoming stream of data. Sleep might help in this process by selectively consolidating new memory traces that are likely to be of future relevance. Using a deferred imitation paradigm and an experimental design, we asked whether 15- and 24-month-old infants (N = 105) who slept soon after encoding a televised demonstration of target actions would show higher imitation scores (retention) after a 24-h delay than same-aged infants who stayed awake for ≥4 h after encoding. In light of infants' well-known difficulties in learning and remembering information from screens, we tested if increasing the relevance of the televised content via standardised caregiver verbalisations might yield the highest imitation scores in the sleep condition. Regardless of sleep condition, 24-month-olds exhibited retention of target actions while 15-month-olds consistently failed to do so. For 24-month-olds, temporal recall was facilitated by sleep, but not by parental verbalisations. Correlational analyses revealed that more time asleep within 4 h after encoding was associated with better retention of the target actions and their temporal order in 24-months-olds. These results suggest that sleep facilitates memory consolidation of screen-based content in late infancy and that this effect might not hinge on caregivers' verbal engagement during viewing., (© 2023 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Motor adaptation reflects the ability of the brain's sensorimotor system to flexibly deal with environmental changes to generate effective motor behaviour. Whether sleep contributes to the consolidation of motor adaptation remains controversial. In this study, we investigated the impact of sleep on motor adaptation and its neurophysiological correlates in a novel motor adaptation task that leverages a highly automatised motor skill, that is, typing. We hypothesised that sleep-associated memory consolidation would benefit motor adaptation and induce modulations in task-related beta band (13-30 Hz) activity during adaptation. Healthy young male experts in typing on the regular computer keyboard were trained to type on a vertically mirrored keyboard while brain activity was recorded using electroencephalography. Typing performance was assessed either after a full night of sleep with polysomnography or a similar period of daytime wakefulness. Results showed improved motor adaptation performance after nocturnal sleep but not after daytime wakefulness, and decreased beta power: (a) during mirrored typing as compared with regular typing; and (b) in the post-sleep versus the pre-sleep mirrored typing sessions. Furthermore, the slope of the electroencephalography signal, a measure of aperiodic brain activity, decreased during mirrored as compared with regular typing. Changes in the electroencephalography spectral slope from pre- to post-sleep mirrored typing sessions were correlated with changes in task performance. Finally, increased fast sleep spindle density (13-15 Hz) during the night following motor adaptation training was predictive of successful motor adaptation. These findings suggest that post-training sleep modulates neural activity supporting adaptive motor functions., (© 2023 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Acute exercise has been shown to affect long-term memory and sleep. However, it is unclear whether exercise-induced changes in sleep architecture are associated with enhanced memory. Recently, it has been shown that exercise followed by a nap improved declarative memory. Whether these effects transfer to night sleep and other memory domains has not yet been studied. Here, we investigate the influence of exercise on nocturnal sleep architecture and associations with sleep-dependent procedural and declarative memory consolidation. Nineteen subjects (23.68 ± 3.97 years) were tested in a balanced cross-over design. In two evening sessions, participants either exercised (high-intensity interval training) or rested immediately after encoding two memory tasks: (1) a finger tapping task and (2) a paired-associate learning task. Subsequent nocturnal sleep was recorded by polysomnography. Retrieval was conducted the following morning. High-intensity interval training lead to an increased declarative memory retention (p = 0.047, d = 0.40) along with a decrease in REM sleep (p = 0.012, d = 0.75). Neither procedural memory nor NREM sleep were significantly affected. Exercise-induced changes in N2 showed a positive correlation with procedural memory retention which did not withstand multiple comparison correction. Exploratory analyses on sleep spindles and slow wave activity did not reveal significant effects. The present findings suggest an exercise-induced enhancement of declarative memory which aligns with changes in nocturnal sleep architecture. This gives additional support for the idea of a potential link between exercise-induced sleep modifications and memory formation which requires further investigation in larger scaled studies., (© 2023 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Targeted memory reactivation (TMR) during sleep enhances memory consolidation in young adults by modulating electrophysiological markers of neuroplasticity. Interestingly, older adults exhibit deficits in motor memory consolidation, an impairment that has been linked to age-related degradations in the same sleep features sensitive to TMR. We hypothesised that TMR would enhance consolidation in older adults via the modulation of these markers. A total of 17 older participants were trained on a motor task involving two auditory-cued sequences. During a post-learning nap, two auditory cues were played: one associated to a learned (i.e., reactivated) sequence and one control. Performance during two delayed re-tests did not differ between reactivated and non-reactivated sequences. Moreover, both associated and control sounds modulated brain responses, yet there were no consistent differences between the auditory cue types. Our results collectively demonstrate that older adults do not benefit from specific reactivation of a motor memory trace by an associated auditory cue during post-learning sleep. Based on previous research, it is possible that auditory stimulation during post-learning sleep could have boosted motor memory consolidation in a non-specific manner., (© 2023 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Sleep is able to contribute not only to memory consolidation, but also to post-sleep learning. The notion exists that either synaptic downscaling or another process during sleep increase post-sleep learning capacity. A correlation between augmentation of the sleep slow oscillation and hippocampal activation at encoding support the contribution of sleep to encoding of declarative memories. In the present study, the effect of closed-loop acoustic stimulation during an afternoon nap on post-sleep encoding of two verbal (word pairs, verbal learning and memory test) and non-verbal (figural pairs) tasks and on electroencephalogram during sleep and learning were investigated in young healthy adults (N = 16). Closed-loop acoustic stimulation enhanced slow oscillatory and spindle activity, but did not affect encoding at the group level. Subgroup analyses and comparisons with similar studies lead us to the tentative conclusion that further parameters such as time of day and subjects' cognitive ability influenced responses to closed-loop acoustic stimulation., (© 2022 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Hippocampal ripple oscillations have been associated with memory reactivations during wake and sleep. These reactivations should contribute to working memory and memory consolidation respectively. In the past decade studies have moved from being observational to actively disrupting ripple-related activity in closed-loop approaches to enable causal investigations into their function. All together these studies have been able to provide evidence that wake, task-related ripple activity is important for working memory and planning but less important for stabilisation of spatial representations. Rest and sleep-related ripple activity, in contrast, is important for long-term memory performance and thus memory consolidation. In this review, we summarise results from different closed-loop approaches in rodents. Further, we highlight differences in detection and stimulation methods as well as controls and discuss how these differences could influence outcomes., (© 2021 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

The beneficial effects of sleep for memory consolidation are assumed to rely on the reactivation of memories in conjunction with the coordinated interplay of sleep rhythms like slow oscillations and spindles. Specifically, slow oscillations are assumed to provide the temporal frame for spindles to occur in the slow oscillations up-states, enabling a redistribution of reactivated information within hippocampal-neocortical networks for long-term storage. Memory reactivation can also be triggered externally by presenting learning-associated cues (like odours or sounds) during sleep, but it is presently unclear whether there is an optimal time-window for the presentation of such cues in relation to the phase of the slow oscillations. In the present within-subject comparison, participants (n = 16) learnt word-pairs visually presented with auditory cues of the first syllable. These syllables were subsequently used for real-time cueing either in the up- or down-state of endogenous slow oscillations. Contrary to our hypothesis, we found differences in memory performance neither between up- and down-state cueing, nor between word-pairs that were cued versus uncued. In the up-state cueing condition, higher amounts of rapid eye movement sleep were associated with better memory for cued contents, whereas higher amounts of slow-wave sleep were associated with better memory for uncued contents. Evoked response analyses revealed signs of cue processing in both conditions. Interestingly, both up- and down-state cueing evoked a similar spindle response with the induced slow oscillations up-state at ~1000 ms post-cue. We speculate that our cueing procedure triggered generalised reactivation processes that facilitated the consolidation of both cued and uncued memories irrespective of the slow oscillation phase., (© 2022 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

There have been numerous attempts over the decades to introduce closed-loop feedback to induce sleep oscillations. Recently, our group also introduced closed-loop acoustic feedback to the sleep spindle and reported improved procedural memory consolidation during a nap with spindle-targeted pink noise stimulation. In this study, we replicated our previous work with a control condition in an attempt to investigate the effect of closed-loop feedback on procedural memory. The results demonstrated a significant improvement in the subjects' procedural learning and reduced wake time during the nap with closed-loop acoustic stimulation compared with the control condition. Further, we found that randomized acoustic stimuli lead to more frequent spindle activity and a faster decrement in slow oscillation power compared with the sham condition. There were strong correlations between slow oscillation and measures related to sleep efficiency as well. Interestingly, we found a marginal enhancement in procedural learning during the nap with the closed-loop acoustic stimulation compared with the sham nap. We also found a marginal decrement in theta power during the nap with closed-loop feedback compared with the sham nap, and a negative correlation between slow oscillation and theta power. We speculate that the marginal improvement in procedural learning may be related to closed-loop acoustic feedback's stabilization of non-rapid eye movement sleep. Taken together, this study shows that the closed-loop feedback method has the potential to stabilize sleep and improve procedural memory., (© 2022 European Sleep Research Society.)

Sleep-dependent memory consolidation is disturbed in patients with schizophrenia, who furthermore show reductions in sleep spindles and probably also in delta power during sleep. The memory dysfunction in these patients is one of the strongest markers for worse long-term functional outcome. However, therapeutic interventions to normalise memory functions, e.g., with medication, still do not exist. Against this backdrop, we investigated to what extent a non-invasive approach enhancing sleep with real-time auditory stimulation in-phase with slow oscillations might affect overnight memory consolidation in patients with schizophrenia. To this end, we examined 18 patients with stably medicated schizophrenia in a double-blinded sham-controlled design. Memory performance was assessed by a verbal (word list) and a non-verbal (complex figure) declarative memory task. In comparison to a sham condition without auditory stimuli, we found that in patients with schizophrenia, auditory stimulation evokes an electrophysiological response similar to that in healthy participants leading to an increase in slow wave and temporally coupled sleep spindle activity during stimulation. Despite this finding, patients did not show any beneficial effect on the overnight change in memory performance by stimulation. Although the stimulation in our study did not improve the patient's memory, the electrophysiological response gives hope that auditory stimulation could enable us to provide better treatment for sleep-related detriments in these patients in the future., (© 2022 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

A widely accepted view in memory research is that previously acquired information can be reactivated during sleep, leading to persistent memory storage. Targeted memory reactivation (TMR) was developed as a technique whereby specific memories can be reactivated during sleep using a sensory stimulus linked to prior learning. As a research tool, TMR can improve memory, raising the possibility that it may be useful for cognitive enhancement and clinical therapy. A major challenge for the expanded use of TMR is that a skilled operator must manually control stimulation, which is impractical in many settings. To address this limitation, we developed the SleepStim system for automated TMR in the home. SleepStim includes a smartwatch to collect movement and heart-rate data, plus a smartphone to emit auditory cues. A machine-learning model identifies periods of deep sleep and triggers TMR sounds within these periods. We tested whether this system could replicate the spatial-memory benefit of in-laboratory TMR. Participants learned locations of objects on a grid, and then half of the object locations were reactivated during sleep over 3 nights. Recall was tested each morning. In an experiment with 61 participants, the TMR effect was not significant but varied systematically with stimulus intensity; low-intensity but not high-intensity stimuli produced memory benefits. In a second experiment with 24 participants, we limited stimulus intensity and found that TMR reliably improved spatial memory, consistent with effects observed in laboratory studies. We conclude that SleepStim can effectively accomplish automated TMR, and that avoiding sleep disruption is critical for TMR benefits., (© 2022 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Increasingly studied in a systematic manner since the 1970s, the cognitive processes of the brain taking place during sleeping periods remain an important object of scrutiny in the scientific community. In particular, sleep has been demonstrated to play a significant role for learning and memory consolidation processes, and sleep scientists have started unravelling its underlying neurophysiological mechanisms. However, sleep remains a multidimensional phenomenon, and many questions remain left open for future research. In this selective review article, we address recent advances in particular domains in which sleep research has further progressed in the past decade. We highlight the developmental trajectory of sleep-dependent learning and memory consolidation processes, from their development in childhood to their potential impairments in ageing, and the nature and extent of our capabilities for information processing, learning, and memory reinforcement during sleep., (© 2022 European Sleep Research Society.)

We summarize here the progress in identifying the neuronal network as well as the function of paradoxical sleep and the gaps of knowledge that should be filled in priority. The core system generating paradoxical sleep localized in the brainstem is now well identified, and the next step is to clarify the role of the forebrain in particular that of the hypothalamus including the melanin-concentrating hormone neurons and of the basolateral amygdala. We discuss these two options, and also the discovery that cortical activation during paradoxical sleep is restricted to a few limbic cortices activated by the lateral supramammillary nucleus and the claustrum. Such activation nicely supports the findings recently obtained showing that neuronal reactivation occurs during paradoxical sleep in these structures, and induces both memory consolidation of important memory and forgetting of less relevant ones. The question that still remains to be answered is whether paradoxical sleep is playing more crucial roles in processing emotional and procedural than other types of memories. One attractive hypothesis is that paradoxical sleep is responsible for erasing negative emotional memories, and that this function is not properly functioning in depressed patients. On the other hand, the presence of a muscle atonia during paradoxical sleep is in favour of a role in procedural memory as new types of motor behaviours can be tried without harm during the state. In a way, it also fits with the proposed role of paradoxical sleep in setting up the sensorimotor system during development., (© 2022 European Sleep Research Society.)

The relationship between sleep and cognition has long been recognized, with slow-wave sleep thought to play a critical role in long-term memory consolidation. Recent research has presented the possibility that non-invasive acoustic stimulation during sleep could enhance memory consolidation. Herein, we report a random-effects model meta-analysis examining the impact of this intervention on memory and sleep architecture in healthy adults. Sixteen studies were identified through a systematic search. We found a medium significant effect of acoustic stimulation on memory task performance (g = 0.68, p = .031) in young adults <35 years of age, but no statistically significant effect in adults >35 years of age (g = -0.83, p = .223). In young adults, there was a large statistically significant effect for declarative memory tasks (g = 0.87, p = .014) but no effect for non-declarative tasks (g = -0.25, p = .357). There were no statistically significant differences in polysomnography-derived sleep architecture values between sham and stimulation conditions in either young or older adults. Based on these results, it appears that acoustic stimulation during sleep may only be an effective intervention for declarative memory consolidation in young adults. However, the small number of studies in this area, their small sample sizes, the short-term nature of most investigations and the high between-studies heterogeneity highlight a need for high-powered and long-term experiments to better elucidate, and subsequently maximise, any potential benefits of this novel approach., (© 2021 European Sleep Research Society.)

The general consensus is that sleep promotes neuronal recovery and plasticity, whereas sleep deprivation (SD) impairs brain function, including cognitive processes. Indeed, a wealth of data has shown a negative impact of SD on learning and memory processes, particularly those that involve the hippocampus. The mechanisms underlying these negative effects of sleep loss are only partly understood, but a reoccurring question is whether they are in part caused by stress hormones that may be released during SD. The purpose of the present study is therefore to examine the role of glucocorticoid stress hormones in SD-induced memory impairment. Male C57BL/6J mice were trained in an object-location memory paradigm, followed by 6 hr of SD by mild stimulation. At the beginning of the SD mice were injected with the corticosterone synthesis inhibitor metyrapone. Memory was tested 24 hr after training. Blood samples taken in a separate group of mice showed that SD resulted in a mild but significant increase in plasma corticosterone levels, which was prevented by metyrapone. However, the SD-induced impairment in object-location memory was not prevented by metyrapone treatment. This indicates that glucocorticoids play no role in causing the memory impairments seen after a short period of SD., (© 2019 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

The current study was designed to further clarify the influence of brain morphology, sleep oscillatory activity and age on memory consolidation. Specifically, we hypothesized, that a smaller volume of hippocampus, parahippocampal and medial prefrontal cortex negatively impacts declarative, but not procedural, memory consolidation. Explorative analyses were conducted to demonstrate whether a decrease in slow-wave activity negatively impacts declarative memory consolidation, and whether these factors mediate age effects on memory consolidation. Thirty-eight healthy participants underwent an acquisition session in the evening and a retrieval session in the morning after night-time sleep with polysomnographic monitoring. Declarative memory was assessed with the paired-associate word list task, while procedural memory was tested using the mirror-tracing task. All participants underwent high-resolution magnetic resonance imaging. Participants with smaller hippocampal, parahippocampal and medial prefrontal cortex volumes displayed a reduced overnight declarative, but not procedural memory consolidation. Mediation analyses showed significant age effects on overnight declarative memory consolidation, but no significant mediation effects of brain morphology on this association. Further mediation analyses showed that the effects of age and brain morphology on overnight declarative memory consolidation were not mediated by polysomnographic variables or sleep electroencephalogram spectral power variables. Thus, the results suggest that the association between age, specific brain area volume and overnight memory consolidation is highly relevant, but does not necessarily depend on slow-wave sleep as previously conceptualized., (© 2020 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research.)

Sleep and memory studies often focus on overnight rather than long-term memory changes, traditionally associating overnight memory change (OMC) with sleep architecture and sleep patterns such as spindles. In addition, (para-)sympathetic innervation has been associated with OMC after a daytime nap using heart rate variability (HRV). In this study we investigated overnight and long-term performance changes for procedural memory and evaluated associations with sleep architecture, spindle activity (SpA) and HRV measures (R-R interval [RRI], standard deviation of R-R intervals [SDNN], as well as spectral power for low [LF] and high frequencies [HF]). All participants (N = 20, M age  = 23.40 ± 2.78 years) were trained on a mirror-tracing task and completed a control (normal vision) and learning (mirrored vision) condition. Performance was evaluated after training (R1), after a full-night sleep (R2) and 7 days thereafter (R3). Overnight changes (R2-R1) indicated significantly higher accuracy after sleep, whereas a significant long-term (R3-R2) improvement was only observed for tracing speed. Sleep architecture measures were not associated with OMC after correcting for multiple comparisons. However, individual SpA change from the control to the learning night indicated that only "SpA enhancers" exhibited overnight improvements for accuracy and long-term improvements for speed. HRV analyses revealed that lower SDNN and LF power was associated with better OMC for the procedural speed measure. Altogether, this study indicates that overnight improvement for procedural memory is specific for spindle enhancers, and is associated with HRV during sleep following procedural learning., (© 2019 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Individuals diagnosed with posttraumatic stress disorder (PTSD) experience disruption at both slow-wave sleep (SWS) and rapid-eye movement (REM) sleep stages and demonstrate marked memory impairment. A small group of studies suggests that, within the disorder, there is a mechanistic relation between these sleep and memory impairments. This study sought to extend that literature by examining whether, in PTSD-diagnosed individuals, memory-retention deficits are present after a sleep-filled (but not after a wake-filled) delay (i.e., whether memory deficits can be traced to interruptions of sleep-dependent memory consolidation). Moreover, we investigated whether SWS- or REM-based disturbances, or both, contribute to retention deficits. We recruited participants into three groups: PTSD (n = 21), trauma-exposed non-PTSD (TE; n = 19) and healthy control (HC; n = 20). Using a crossover design, we assessed memory recall before and after an 8-hr period of polysomnography-monitored sleep and an 8-hr period of regular waking activity. PTSD-diagnosed participants retained less information than controls over the sleep-filled (but not wake-filled) delay. Furthermore, increased REM fragmentation predicted postsleep memory retention in PTSD-diagnosed individuals only. No SWS parameter was associated with or predictive of the amount of information retained postsleep. We conclude that specific REM-related changes in PTSD-diagnosed individuals affected sleep-dependent neutral declarative memory consolidation. Generally, these findings extend the literature suggesting that the co-occurrence of sleep and memory difficulties in PTSD is not accidental, but that these two symptom clusters are meaningfully related. Specifically, the study illustrates that subtle REM-related disruptions contribute most strongly to memory impairment in PTSD., (© 2019 European Sleep Research Society.)

Initially independent lines of research suggest that sleep-specific brain activity patterns, observed as electroencephalographic slow oscillatory and sleep spindle activity, promote memory consolidation and underlying synaptic refinements. Here, we further tested the emerging concept that specifically the coordinated interplay of slow oscillations and spindle activity (phase-amplitude coupling) support memory consolidation. Particularly, we associated indices of the interplay between slow oscillatory (0.16-1.25 Hz) and spindle activity (12-16 Hz) during non-rapid eye movement sleep (strength [modulation index] and phase degree of coupling) in 20 healthy adults with parameters of overnight declarative (word-list task) and procedural (mirror-tracing task) memory consolidation. The pattern of results supports the notion that the interplay between oscillations facilitates memory consolidation. The coincidence of the spindle amplitude maximum with the up-state of the slow oscillation (phase degree) was significantly associated with declarative memory consolidation (r = .65, p = .013), whereas the overall strength of coupling (modulation index) correlated with procedural memory consolidation (r = .45, p = .04). Future studies are needed to test for potential causal effects of the observed association between neural oscillations during sleep and memory consolidation, and to elucidate ways of modulating these processes, for instance through non-invasive brain-stimulation techniques., (© 2019 European Sleep Research Society.)

Insomnia involves disruption of sleep initiation, maintenance and/or overall quality, and may interfere with cognition. Here, we evaluated memory impairment produced by rodent mild (acute) insomnia models. Insomnia models consisted of either single or repeated exposure to cages previously occupied (dirtied) by an unfamiliar rat for 5-7 days. Rats were trained in the Morris water maze to remember the platform location (acquisition), and were then exposed to: (a) 6 hr of undisturbed baseline; (b) dirty cage change-induced insomnia (animal placed into a cage dirtied by another rat for 6 hr); or (c) double-dirty cage change-induced insomnia (animal placed into a cage dirtied by another rat for 3 hr, and then another dirty cage 3 hr later). The animal's memory for the platform location was then evaluated in a probe trial. Double-dirty cage change-induced insomnia significantly disrupted sleep, although the effects of dirty cage change-induced insomnia were overall not significant. In the fourth hour of double-dirty cage change-induced insomnia (following the second cage change), sleep episode number and duration alterations indicated sleep fragmentation. Furthermore, power spectral analysis revealed diminished wake and, to a lesser extent, rapid eye movement theta power (indicated by trend difference) in the last 3 hr of exposure. Significant deficits were noted for measures of water maze performance following double-dirty cage change-induced insomnia, indicating impaired memory. In summary, one variant of the rodent insomnia model, double-dirty cage change-induced insomnia, disrupted sleep and attenuated memory consolidation, indicating this paradigm may be useful to evaluate the effects of hypnotics on memory consolidation., (© 2018 European Sleep Research Society.)

Sleep following learning benefits memory. One model attributes this effect to the iterative "reactivation" of memory traces in the sleeping brain, demonstrated in animal models. Although technical limitations prohibit using the same methods to observe memory reactivation in the human brain, the study of mental activity during sleep provides an alternative method of observing memory activation during sleep. In fact, the content of dream experience may reflect the process of memory reactivation and consolidation in the sleeping brain. In line with this hypothesis, we previously reported that dreaming about a spatial learning task during a nap strongly predicts subsequent performance improvements. Here, we replicate this observation in an overnight sleep study, for the first time demonstrating that pre-sleep training on a virtual maze navigation task is reflected in dreams reported from all phases of sleep, with unambiguous representation of the task in dream content associated with improved next-morning performance. These observations are consistent with reactivation-based models of memory consolidation in sleep, confirming our earlier finding that the cognitive-level activation of recent experience during sleep is associated with subsequent performance gains., (© 2018 European Sleep Research Society.)

Collecting dream reports typically requires waking subjects up from their sleep-a method that has been used to study the relationship between dreams and memory consolidation. However, it is unclear whether these awakenings influence sleep-associated memory consolidation processes. Furthermore, it is unclear how the incorporation of the learning task into dreams is related to memory consolidation. In this study we compared memory performance in a word-picture association learning task after a night with and without awakenings in 22 young and healthy participants. We then examined if the stimuli from the learning task are successfully incorporated into dreams, and if this incorporation is related to the task performance the next morning. We show that while the awakenings impaired both subjective and objective sleep quality, they did not affect sleep-associated memory consolidation. When dreams were collected during the night by awakenings, memories of the learning task were successfully incorporated into dreams. When dreams were collected in the morning, no incorporations were detected. Task incorporation into non-rapid eye movement sleep dreams, but not rapid eye movement sleep dreams positively predicted memory performance the next morning. We conclude that the method of awakenings to collect dream reports is suitable and necessary for dream and memory studies. Furthermore, our study suggests that dreams in non-rapid eye movement rather than rapid eye movement sleep might be related to processes of memory consolidation during sleep., (© 2018 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Recent work has suggested that the benefits of sleep for memory consolidation are enhanced for highly salient (versus non-salient) memories. Using a technique known as targeted memory reactivation, it is possible to selectively strengthen newly learned memories by re-exposing the sleeping brain to auditory cues. The aim of the current study was to examine whether emotionally salient memories are also more responsive to targeted memory reactivation in slow-wave sleep than neutral memories. In an initial training phase, participants memorised emotionally negative and neutral pictures, which were each paired with a semantically related sound. Recognition for the pictures was assessed before and after a 90-min nap opportunity, during which half the sounds were re-presented during slow-wave sleep (as assessed via online polysomnographic sleep monitoring). We observed no effect of targeted memory reactivation on the recognition of emotionally negative or neutral memories. Our results highlight the importance of the memory paradigm used to assess targeted memory reactivation, and suggest that the robust and durable nature of recognition memory may make it an insensitive measure of behavioural targeted memory reactivation benefits. To fully assess the impacts of targeted memory reactivation on emotional memory processing in sleep, future studies should adopt experimental paradigms that maximise the salience of emotional stimuli while also providing a sensitive index of memory accuracy., (© 2017 European Sleep Research Society.)

Face recognition is a highly specialized capability that has implicit and explicit memory components. Studies show that learning tasks with facial components are dependent on rapid eye movement and non-rapid eye movement sleep features, including rapid eye movement sleep density and fast sleep spindles. This study aimed to investigate the relationship between sleep-dependent consolidation of memory for faces and partial rapid eye movement sleep deprivation, rapid eye movement density, and fast and slow non-rapid eye movement sleep spindles. Fourteen healthy participants spent 1 night each in the laboratory. Prior to bed they completed a virtual reality task in which they interacted with computer-generated characters. Half of the participants (REMD group) underwent a partial rapid eye movement sleep deprivation protocol and half (CTL group) had a normal amount of rapid eye movement sleep. Upon awakening, they completed a face recognition task that contained a mixture of previously encountered faces from the task and new faces. Rapid eye movement density and fast and slow sleep spindles were detected using in-house software. The REMD group performed worse than the CTL group on the face recognition task; however, rapid eye movement duration and rapid eye movement density were not related to task performance. Fast and slow sleep spindles showed differential relationships to task performance, with fast spindles being positively and slow spindles negatively correlated with face recognition. The results support the notion that rapid eye movement and non-rapid eye movement sleep characteristics play complementary roles in face memory consolidation. This study also raises the possibility that fast and slow spindles contribute in opposite ways to sleep-dependent memory consolidation., (© 2017 European Sleep Research Society.)

Although many types of newly encoded information can be consolidated during sleep, an enhanced effect has been found for memories tagged as relevant to the future, such as through knowledge of future testing or payment for successful recall. In the current study, participants (n = 80) learned Welsh and Breton translations of English words, and intrinsic relevance of learned material was operationalized as perceived value of the Welsh and Breton languages. Participants were non-Welsh native English speakers who had recently arrived in Wales. Memory for the words was tested immediately and 12 h later, after either a period of wake or a period of sleep. An increase in recall for both languages was found after sleep, but not after wake. Importantly, for the sleep condition, overnight improvement in Welsh word recall was associated with participants' level of valuing the Welsh language. This association was not found for the wake period condition. These findings support previous indications of an active role of sleep in the consolidation of memories relevant for the future, and demonstrate that this effect may be modulated by individual differences in perceived value of the learned material. It remains to be established whether this association is mediated by an emotional attachment to the language or a cognitive facility with it, or both., (© 2016 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Effectiveness of memory consolidation is determined by multiple factors, including sleep after learning, emotional valence, arousal and novelty. Few studies investigated how the effect of sleep compares with (and interacts with) these other factors, of which virtually none are in children. The present study did so by repeated assessment of declarative memory in 386 children (45% boys) aged 9-11 years through an online word-pair task. Children were randomly assigned to either a morning or evening learning session of 30 unrelated word-pairs with positive, neutral or negative valenced cues and neutral targets. After immediately assessing baseline recognition, delayed recognition was recorded either 12 or 24 h later, resulting in four different assessment schedules. One week later, the procedure was repeated with exactly the same word-pairs to evaluate whether effects differed for relearning versus original novel learning. Mixed-effect logistic regression models were used to evaluate how the probability of correct recognition was affected by sleep, valence, arousal, novelty and their interactions. Both immediate and delayed recognition were worse for pairs with negatively valenced or less arousing cue words. Relearning improved immediate and delayed word-pair recognition. In contrast to these effects, sleep did not affect recognition, nor did sleep moderate the effects of arousal, valence and novelty. The findings suggest a robust inclination of children to specifically forget the pairing of words to negatively valenced cue words. In agreement with a recent meta-analysis, children seem to depend less on sleep for the consolidation of information than has been reported for adults, irrespective of the emotional valence, arousal and novelty of word-pairs., (© 2017 European Sleep Research Society.)

Retrieving false information can have serious consequences. Sleep is important for memory, but voluntary sleep curtailment is becoming more rampant. Here, the misinformation paradigm was used to investigate false memory formation after 1 night of total sleep deprivation in healthy young adults (N = 58, mean age ± SD = 22.10 ± 1.60 years; 29 males), and 7 nights of partial sleep deprivation (5 h sleep opportunity) in these young adults and healthy adolescents (N = 54, mean age ± SD = 16.67 ± 1.03 years; 25 males). In both age groups, sleep-deprived individuals were more likely than well-rested persons to incorporate misleading post-event information into their responses during memory retrieval (P < 0.050). These findings reiterate the importance of adequate sleep in optimal cognitive functioning, reveal the vulnerability of adolescents' memory during sleep curtailment, and suggest the need to assess eyewitnesses' sleep history after encountering misleading information., (© 2016 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

As chronic sleep restriction is a widespread problem among adolescents, the present study investigated the effects of a 1-week sleep restriction (SR) versus control period on the consolidation of long-term memory for prose passages. We also determined whether the benefit of prioritization on memory is modulated by adequate sleep occurring during consolidation. Fifty-six healthy adolescents (25 male, aged 15-19 years) were instructed to remember a prose passage in which half of the content was highlighted (prioritized), and were told that they would receive an additional bonus for remembering highlighted content. Following an initial free recall test, participants underwent a 7-night period in which they received either a 5-h (SR) or 9-h (control) nightly sleep opportunity, monitored by polysomnography on selected nights. Free recall of the passage was tested at the end of the sleep manipulation period (1 week after encoding), and again 6 weeks after encoding. Recall of highlighted content was superior to that of non-highlighted content at all three time-points (initial, 1 week, 6 weeks). This beneficial effect of prioritization on memory was stronger 1 week relative to a few minutes after encoding for the control, but not the SR group. N3 duration was similar in the control and SR groups. Overall, the present study shows that the benefits of prioritization on memory are enhanced over time, requiring time and sleep to unfold fully. Partial sleep deprivation (i.e. 5-h nocturnal sleep opportunity) may attenuate such benefits, but this may be offset by preservation of N3 sleep duration., (© 2016 The Authors. Journal of Sleep Research published by John Wiley & Sons Ltd on behalf of European Sleep Research Society.)

Both sleep and future relevance influence memory consolidation; however, limited research has investigated their role in memory reconsolidation. We manipulated the future relevance of both stable and labile memories in need of reconsolidation. Two groups learned two blocks of syllable pairs on one evening and were told they would be tested on one of the blocks later. On the second evening, one group (Labile) received reminders designed to return their memories of syllable pairs to a labile state, while a second group (Stable) received reminders designed to leave these memories in a stable state. No significant differences in memory retention were found between blocks or groups the following morning. Frontal delta (0.5-4 Hz) electroencephalographic power during Stage 2 sleep correlated positively with retention of future-relevant material exclusively in the Labile group. Central theta (4-8 Hz) electroencephalographic power during rapid eye movement (REM) sleep correlated positively with the extent to which the Labile group selectively retained future-relevant material. These relationships suggest that sleep-dependent processes are involved in an economical reprocessing of memories beyond the initial stages of consolidation., (© 2016 European Sleep Research Society.)

Although sleep facilitates learning and memory, the roles of dreaming and habitual levels of recalling dreams remain unknown. This study examined if performance and overnight improvement on a rapid eye movement sleep-sensitive visuomotor task is associated differentially with habitually high or low dream recall frequency. As a relation between dream production and visuospatial skills has been demonstrated previously, one possibility is that frequency of dream recall will be linked to performance on visuomotor tasks such as the Mirror Tracing Task. We expected that habitually low dream recallers would perform more poorly on the Mirror Tracing Task than would high recallers and would show less task improvement following a night of sleep. Fifteen low and 20 high dream recallers slept one night each in the laboratory and performed the Mirror Tracing Task before and after sleep. Low recallers had overall worse baseline performance but a greater evening-to-morning improvement than did high recallers. Greater improvements in completion time in low recallers were associated with Stage 2 rather than rapid eye movement sleep. Findings support the separate notions that dreaming is related to visuomotor processes and that different levels of visuomotor skill engage different sleep- and dream-related consolidation mechanisms., (© 2015 European Sleep Research Society.)