Slow oscillations during slow-wave sleep (SWS) may facilitate memory consolidation by regulating interactions between hippocampal and cortical networks. Slow oscillations appear as high-amplitude, synchronized EEG activity, corresponding to upstates of neuronal depolarization and downstates of hyperpolarization. Memory reactivations occur spontaneously during SWS, and can also be induced by presenting learning-related cues associated with a prior learning episode during sleep. This technique, targeted memory reactivation (TMR), selectively enhances memory consolidation. Given that memory reactivation is thought to occur preferentially during the slow-oscillation upstate, we hypothesized that TMR stimulation effects would depend on the phase of the slow oscillation. Participants learned arbitrary spatial locations for objects that were each paired with a characteristic sound (eg, cat-meow). Then, during SWS periods of an afternoon nap, one-half of the sounds were presented at low intensity. When object location memory was subsequently tested, recall accuracy was significantly better for those objects cued during sleep. We report here for the first time that this memory benefit was predicted by slow-wave phase at the time of stimulation. For cued objects, location memories were categorized according to amount of forgetting from pre- to post-nap. Conditions of high versus low forgetting corresponded to stimulation timing at different slow-oscillation phases, suggesting that learning-related stimuli were more likely to be processed and trigger memory reactivation when they occurred at the optimal phase of a slow oscillation. These findings provide insight into mechanisms of memory reactivation during sleep, supporting the idea that reactivation is most likely during cortical upstates. Slow-wave sleep (SWS) is characterized by synchronized neural activity alternating between active upstates and quiet downstates. The slow-oscillation upstates are thought to provide a window of opportunity for memory consolidation, particularly conducive to cortical plasticity. Recent evidence shows that sensory cues associated with previous learning can be delivered subtly during SWS to selectively enhance memory consolidation. Our results demonstrate that this behavioral benefit is predicted by slow-oscillation phase at stimulus presentation time. Cues associated with high versus low forgetting based on analysis of subsequent recall performance were delivered at opposite slow-oscillation phases. These results provide evidence of an optimal slow-oscillation phase for memory consolidation during sleep, supporting the idea that memory processing occurs preferentially during cortical upstates. Copyright © 2016 the authors 0270-6474/16/361401-09$15.00/0.

Over the past two decades, research has accumulated compelling evidence that sleep supports the formation of long-term memory. The standard two-stage memory model that has been originally elaborated for declarative memory assumes that new memories are transiently encoded into a temporary store (represented by the hippocampus in the declarative memory system) before they are gradually transferred into a long-term store (mainly represented by the neocortex), or are forgotten. Based on this model, we propose that sleep, as an offline mode of brain processing, serves the 'active system consolidation' of memory, i.e. the process in which newly encoded memory representations become redistributed to other neuron networks serving as long-term store. System consolidation takes place during slow-wave sleep (SWS) rather than rapid eye movement (REM) sleep. The concept of active system consolidation during sleep implicates that (a) memories are reactivated during sleep to be consolidated, (b) the consolidation process during sleep is selective inasmuch as it does not enhance every memory, and (c) memories, when transferred to the long-term store undergo qualitative changes. Experimental evidence for these three central implications is provided: It has been shown that reactivation of memories during SWS plays a causal role for consolidation, that sleep and specifically SWS consolidates preferentially memories with relevance for future plans, and that sleep produces qualitative changes in memory representations such that the extraction of explicit and conscious knowledge from implicitly learned materials is facilitated.

Slow wave sleep (SWS) is known to favour episodic memory consolidation. Given that ageing is associated with a reduction in SWS and episodic memory impairment, our aim was to investigate whether memory continues to benefit from sleep in older adults. Episodic memory consolidation was tested in 20 young (22.1 ± 1.7 years) and 20 older volunteers (68.9 ± 5.3 years) who performed a visuospatial two-dimensional object-location task. Retention capacities were evaluated after 12 h of wakefulness or 12 h of sleep. Performances before and after the interval allowed us to calculate a forgetting rate. Sleep architecture was measured by polysomnography (older adults = 410 min; young adults: 467 min). Our results showed that the beneficial effect of sleep on memory consolidation was reduced in older adults compared to young adults. In older adults, sleep did not enhance memory consolidation significantly compared to wakefulness. Sleep prevented young adults from forgetting (-0.10% ± 2.1), while the forgetting rate in older adults was still important after a period of sleep (16.60% ± 4.2; P = 0.05). The sleep architecture of older adults was characterized by a decrease in sleep efficiency (-12%; P < 0.05), in total cycle time (-137 min; P < 0.05), in percentage of total cycle time (-21%; P < 0.05) and in rapid eye movement time (-41 min; P < 0.05) compared to young adults. However, no difference in slow wave sleep was observed (-1%; not significant) and no correlation was found with performance. Age-related changes in sleep parameters may have a negative impact on memory consolidation in older adults. © 2013 European Sleep Research Society.

Human creativity has been claimed to rely on the neurotransmitter dopamine, but evidence is still sparse. We studied whether individual performance (N=117) in divergent thinking (alternative uses task) and convergent thinking (remote association task) can be predicted by the individual spontaneous eye blink rate (EBR), a clinical marker of dopaminergic functioning. EBR predicted flexibility in divergent thinking and convergent thinking, but in different ways. The relationship with flexibility was independent of intelligence and followed an inverted U-shape function with medium EBR being associated with greatest flexibility. Convergent thinking was positively correlated with intelligence but negatively correlated with EBR, suggesting that higher dopamine levels impair convergent thinking. These findings support the claim that creativity and dopamine are related, but they also call for more conceptual differentiation with respect to the processes involved in creative performance.

Memories are gradually consolidated after initial encoding, and this can sometimes lead to a transition from implicit to explicit knowledge. The exact physiological processes underlying this reorganization remain unclear. Here, we used a serial reaction time task to determine whether targeted memory reactivation (TMR) of specific memory traces during slow-wave sleep promotes the emergence of explicit knowledge. Human participants learned two 12-item sequences of button presses (A and B). These differed in both cue order and in the auditory tones associated with each of the four fingers (one sequence had four higher-pitched tones). Subsequent overnight sleep was monitored, and the tones associated with one learned sequence were replayed during slow-wave sleep. After waking, participants demonstrated greater explicit knowledge (p = 0.005) and more improved procedural skill (p = 0.04) for the cued sequence relative to the uncued sequence. Furthermore, fast spindles (13.5-15 Hz) at task-related motor regions predicted overnight enhancement in procedural skill (r = 0.71, p = 0.01). Auditory cues had no effect on post-sleep memory performance in a control group who received TMR before sleep. These findings suggest that TMR during sleep can alter memory representations and promote the emergence of explicit knowledge, supporting the notion that reactivation during sleep is a key mechanism in this process. Copyright © 2014 Cousins et al.

Sleep has been shown to foster the process of insight generation in young adults during problem solving activities. Aging is characterized by substantial changes in sleep architecture altering memory consolidation. Whether sleep might promote the occurrence of insight in older adults as well has not yet been tested experimentally. To address this issue, we tested healthy young and old volunteers on an insight problem solving task, involving both explicit and implicit features, before and after a night of sleep or a comparable wakefulness period. Data showed that insight emerged significantly less frequently after a night of sleep in older adults compared to young. Moreover, there was no difference in the magnitude of insight occurrence following sleep and daytime -consolidation in aged participants. We further found that acquisition of implicit knowledge in the task before sleep potentiated the gain of insight in young participants, but this effect was not observed in aged participants. Overall, present findings demonstrate that a period of sleep does not significantly promote insight in problem solving in older adults. Copyright © 2017 Elsevier Inc. All rights reserved.

Extracting regularities from a sequence of events is essential for understanding our environment. However, there is no consensus regarding the extent to which such regularities can be generalised beyond the modality of learning. One reason for this could be the variation in consolidation intervals used in different paradigms, also including an opportunity to sleep. Using a novel statistical learning paradigm in which structured information is acquired in the auditory domain and tested in the visual domain over either 30 min or 24 h consolidation intervals, we show that cross-modal transfer can occur, but this transfer is only seen in the 24 h group. Importantly, the extent of cross-modal transfer is predicted by the amount of slow wave sleep (SWS) obtained. Additionally, cross-modal transfer is associated with the same pattern of decreasing medial temporal lobe and increasing striatal involvement which has previously been observed to occur across 24 h in unimodal statistical learning. We also observed enhanced functional connectivity after 24 h in a network of areas which have been implicated in cross-modal integration including the precuneus and the middle occipital gyrus. Finally, functional connectivity between the striatum and the precuneus was also enhanced, and this strengthening was predicted by SWS. These results demonstrate that statistical learning can generalise to some extent beyond the modality of acquisition, and together with our previously published unimodal results, support the notion that statistical learning is both domain-general and domain-specific. Copyright © 2016 Elsevier Ltd. All rights reserved.

Sleep after learning promotes the quantitative strengthening of new memories. Less is known about the impact of sleep on the qualitative reorganization of memory content. This study tested the hypothesis that sleep facilitates both memory strengthening and reorganization as indexed by a verbal creativity task. Sixty healthy university students (30 female, 30 male, 20-30 years) were investigated in a randomized, controlled parallel-group study with three experimental groups (sleep, sleep deprivation, daytime wakefulness). At baseline, 60 items of the Compound Remote Associate (CRA) task were presented. At retest after the experimental conditions, the same items were presented again together with 20 new control items to disentangle off-line incubation from online performance effects. Sleep significantly strengthened formerly encoded memories in comparison to both wake conditions (improvement in speed of correctly resolved items). Offline reorganization was not enhanced following sleep, but was enhanced following sleep-deprivation in comparison to sleep and daytime wakefulness (solution time of previously incubated, newly solved items). Online performance did not differ between the groups (solution time of new control items). The results support the notion that sleep promotes the strengthening, but not the reorganization, of newly encoded memory traces in a verbal creativity task. Future studies are needed to further determine the impact of sleep on different types of memory reorganization, such as associative thinking, creativity and emotional memory processing, and potential clinical translations, such as the augmentation of psychotherapy through sleep interventions. © 2016 Associated Professional Sleep Societies, LLC.

Sleep after learning promotes the quantitative strengthening of new memories. Less is known about the impact of sleep on the qualitative reorganisation of memory, which is the focus of this review. Studies have shown that, in the declarative system, sleep facilitates the abstraction of rules (schema formation), the integration of knowledge into existing schemas (schema integration) and creativity that requires the disbandment of existing patterns (schema disintegration). Schema formation and integration might primarily benefit from slow wave sleep, whereas the disintegration of a schema might be facilitated by rapid eye movement sleep. In the procedural system, sleep fosters the reorganisation of motor memory. The neural mechanisms of these processes remain to be determined. Notably, emotions have been shown to modulate the sleep-related reorganisation of memories. In the final section of this review, we propose that the sleep-related reorganisation of memories might be particularly relevant for mental disorders. Thus, sleep disruptions might contribute to disturbed memory reorganisation and to the development of mental disorders. Therefore, sleep-related interventions might modulate the reorganisation of memories and provide new inroads into treatment. Copyright © 2014 Elsevier Ltd. All rights reserved.

Sleep facilitates declarative memory processing. However, we know little about whether sleep plays a role in the processing of a fundamental feature of declarative memory, relational memory - the flexible representation of items not directly learned prior to sleep. Thirty-one healthy participants first learned at 12 pm two sets of face-object photograph pairs (direct associative memory), in which the objects in each pair were common to both lists, but paired with two different faces. Participants either were given approximately 90 min to have a NREM-only daytime nap (n=14) or an equivalent waking period (n=17). At 4:30 pm, participants who napped demonstrated significantly better retention of direct associative memory, as well as better performance on a surprise task assessing their relational memory, in which participants had to associate the two faces previously paired with the same object during learning. Particularly noteworthy, relational memory performance was correlated with the amount of NREM sleep during the nap, with only slow-wave sleep predicting relational memory performance. Sleep stage data did not correlate with direct associative memory retention. These results suggest an active role for sleep in facilitating multiple processes that are not limited to the mere strengthening of rote memories, but also the binding of items that were not directly learned together, reorganizing them for flexible use at a later time. Copyright 2010 Elsevier Inc. All rights reserved.

This article argues both rapid eye movement (REM) and non-rapid eye movement (NREM) sleep contribute to overnight episodic memory processes but their roles differ. Episodic memory may have evolved from memory for spatial navigation in animals and humans. Equally, mnemonic navigation in world and mental space may rely on fundamentally equivalent processes. Consequently, the basic spatial network characteristics of pathways which meet at omnidirectional nodes or junctions may be conserved in episodic brain networks. A pathway is formally identified with the unidirectional, sequential phases of an episodic memory. In contrast, the function of omnidirectional junctions is not well understood. In evolutionary terms, both animals and early humans undertook tours to a series of landmark junctions, to take advantage of resources (food, water and shelter), whilst trying to avoid predators. Such tours required memory for emotionally significant landmark resource-place-danger associations and the spatial relationships amongst these landmarks. In consequence, these tours may have driven the evolution of both spatial and episodic memory. The environment is dynamic. Resource-place associations are liable to shift and new resource-rich landmarks may be discovered, these changes may require re-wiring in neural networks. To realise these changes, REM may perform an associative, emotional encoding function between memory networks, engendering an omnidirectional landmark junction which is instantiated in the cortex during NREM Stage 2. In sum, REM may preplay associated elements of past episodes (rather than replay individual episodes), to engender an unconscious representation which can be used by the animal on approach to a landmark junction in wake. Copyright © 2015 Elsevier Inc. All rights reserved.

This study was conceived by the idea that there exist different kinds of cognitive style-based, task competencies that have implications for task motivation and cognitive performance on creative problem-solving tasks/insight. Specifically, the relationships among the Assimilator Explorer styles (Kaufmann, 1979), experimentally manipulated task competence for each style, and performance on insight tasks was examined. A total of 264 participants with a mean age of 17.4 years completed a cognitive style test, two measures of task motivation, and three practical construction-type insight tasks. Explorers with experimentally increased competence beliefs were hypothesized to perform less well on typical insight problems than Explorers with experimentally decreased competence beliefs, while Assimilators with increased levels of competence beliefs were expected to perform better than Assimilators with decreased competence beliefs. A randomized experiment with written instructions was conducted to test these hypotheses. The results supported the main hypothesis, yet an additional three-way interaction hypothesis among styles, experimentally manipulated task competence, and task structure was not supported. Limitations are discussed.

Damage to the hippocampal system disrupts recent memory but leaves remote memory intact. The account presented here suggests that memories are first stored via synaptic changes in the hippocampal system, that these changes support reinstatement of recent memories in the neocortex, that neocortical synapses change a little on each reinstatement, and that remote memory is based on accumulated neocortical changes. Models that learn via changes to connections help explain this organization. These models discover the structure in ensembles of items if learning of each item is gradual and interleaved with learning about other items. This suggests that the neocortex learns slowly to discover the structure in ensembles of experiences. The hippocampal system permits rapid learning of new items without disrupting this structure, and reinstatement of new memories interleaves them with others to integrate them into structured neocortical memory systems.

The nine-dot problem is often used to demonstrate and explain mental impasse, creativity, and out of the box thinking. The present study investigated the interplay of a restricted initial search space, the likelihood of invoking a representational change, and the subsequent constraining of an unrestricted search space. In three experimental conditions, participants worked on different versions of the nine-dot problem that hinted at removing particular sources of difficulty from the standard problem. The hints were incremental such that the first suggested a possible route for a solution attempt; the second additionally indicated the dot at which lines meet on the solution path; and the final condition also provided non-dot locations that appear in the solution path. The results showed that in the experimental conditions, representational change is encountered more quickly and problems are solved more often than for the control group. We propose a cognitive model that focuses on general problem-solving heuristics and representational change to explain problem difficulty.

Sleep deprivation is common among university students, and has been associated with poor academic performance and physical dysfunction. However, current literature has a narrow focus in regard to domains tested, this study aimed to investigate the effects of a night of sleep deprivation on cognitive and physical performance in students. A randomized controlled crossover study was carried out with 64 participants [58% male ( = 37); 22 ± 4 years old (mean ± SD)]. Participants were randomized into two conditions: normal sleep or one night sleep deprivation. Sleep deprivation was monitored using an online time-stamped questionnaire at 45 min intervals, completed in the participants' homes. The outcomes were cognitive: working memory (Simon game© derivative), executive function (Stroop test); and physical: reaction time (ruler drop testing), lung function (spirometry), rate of perceived exertion, heart rate, and blood pressure during submaximal cardiopulmonary exercise testing. Data were analysed using paired two-tailed tests and MANOVA. Reaction time and systolic blood pressure post-exercise were significantly increased following sleep deprivation (mean ± SD change: reaction time: 0.15 ± 0.04 s,  = 0.003; systolic BP: 6 ± 17 mmHg,  = 0.012). No significant differences were found in other variables. Reaction time and vascular response to exercise were significantly affected by sleep deprivation in university students, whilst other cognitive and cardiopulmonary measures showed no significant changes. These findings indicate that acute sleep deprivation can have an impact on physical but not cognitive ability in young healthy university students. Further research is needed to identify mechanisms of change and the impact of longer term sleep deprivation in this population.

Anecdotal reports recount of individuals obtaining insights during sleep. For instance, various acclaimed scientists have attributed some of their greatest insights to sleep-related mentation. To date, this phenomenon has not been systematically investigated. The current study explored the occurrence and characteristics of Sleep-Related Insights (SRIs) in a large population sample, using a questionnaire approach. We found that a large majority of participants (~80%) experienced SRIs at some point in their lives and about 40% obtained SRIs regularly. Most of these subjects could link SRIs to remembered sleep mentation. SRIs were reported to occur in both sleep and half-sleep states, and at any point of the sleep period. Furthermore, SRIs regarded emotional preoccupations about twice as often as theoretical problems. Finally, SRIs were not robustly related to subjective sleep-quality, but small positive correlations with insomnia and narcolepsy-like symptoms were observed. In conclusion, SRIs are much more common than might have been expected, manifest in several forms and appear to be part of normal, healthy sleep. Importantly, the strong link of SRIs with sleep mentation suggests they result from some form of higher-order information processing during sleep, rather than being (fully) secondary to general restorative effects of sleep. Finally, our findings show that a large portion of the sampled population is aware of sleep's benefits for real life problem solving and experiences such benefits on a regular basis.

During creative problem solving, initial solution attempts often fail because of self-imposed constraints that prevent us from thinking out of the box. In order to solve a problem successfully, the problem representation has to be restructured by combining elements of available knowledge in novel and creative ways. It has been suggested that sleep supports the reorganization of memory representations, ultimately aiding problem solving. In this study, we systematically tested the effect of sleep and time on problem solving, using classical insight tasks and magic tricks. Solving these tasks explicitly requires a restructuring of the problem representation and may be accompanied by a subjective feeling of insight. In two sessions, 77 participants had to solve classical insight problems and magic tricks. The two sessions either occurred consecutively or were spaced 3 h apart, with the time in between spent either sleeping or awake. We found that sleep affected neither general solution rates nor the number of solutions accompanied by sudden subjective insight. Our study thus adds to accumulating evidence that sleep does not provide an environment that facilitates the qualitative restructuring of memory representations and enables problem solving.

Schreiner, Thomas; Lehmann, Mick; Rasch, Bjoern Univ Zurich, Dept Psychol, CH-8050 Zurich, Switzerland. Schreiner, Thomas; Rasch, Bjoern Univ Fribourg, Dept Psychol, CH-1701 Fribourg, Switzerland. Lehmann, Mick Psychiat Univ Hosp Zurich, Clin Affect Disorders & Gen Psychiat, CH-8032 Zurich, Switzerland. Rasch, Bjoern Zurich Ctr Interdisciplinary Sleep Res ZiS, CH-8091 Zurich, Switzerland.

Memory reactivation appears to be a fundamental process in memory consolidation. In this study we tested the influence of memory reactivation during rapid eye movement (REM) sleep on memory performance and brain responses at retrieval in healthy human participants. Fifty-six healthy subjects (28 women and 28 men, age [mean ± standard deviation]: 21.6 ± 2.2 y) participated in this functional magnetic resonance imaging (fMRI) study. Auditory cues were associated with pictures of faces during their encoding. These memory cues delivered during REM sleep enhanced subsequent accurate recollections but also false recognitions. These results suggest that reactivated memories interacted with semantically related representations, and induced new creative associations, which subsequently reduced the distinction between new and previously encoded exemplars. Cues had no effect if presented during stage 2 sleep, or if they were not associated with faces during encoding. Functional magnetic resonance imaging revealed that following exposure to conditioned cues during REM sleep, responses to faces during retrieval were enhanced both in a visual area and in a cortical region of multisensory (auditory-visual) convergence. These results show that reactivating memories during REM sleep enhances cortical responses during retrieval, suggesting the integration of recent memories within cortical circuits, favoring the generalization and schematization of the information.

Sleep spindle activity has been associated with improvements in procedural and declarative memory. Here, for the first time, we looked at the role of spindles in the integration of newly learned information with existing knowledge, contrasting this with explicit recall of the new information. Two groups of participants learned novel spoken words (e.g., cathedruke) that overlapped phonologically with familiar words (e.g., cathedral). The sleep group was exposed to the novel words in the evening, followed by an initial test, a polysomnographically monitored night of sleep, and a second test in the morning. The wake group was exposed and initially tested in the morning and spent a retention interval of similar duration awake. Finally, both groups were tested a week later at the same circadian time to control for possible circadian effects. In the sleep group, participants recalled more words and recognized them faster after sleep, whereas in the wake group such changes were not observed until the final test 1 week later. Following acquisition of the novel words, recognition of the familiar words was slowed in both groups, but only after the retention interval, indicating that the novel words had been integrated into the mental lexicon following consolidation. Importantly, spindle activity was associated with overnight lexical integration in the sleep group, but not with gains in recall rate or recognition speed of the novel words themselves. Spindle activity appears to be particularly important for overnight integration of new memories with existing neocortical knowledge.

In recent years, vibrant research has developed on "consolidation" during sleep: To what extent are newly experienced impressions reprocessed or even restructured during sleep? We used the number reduction task (NRT) to study if and how sleep does not only reiterate new experiences but may even lead to new insights. In the NRT, covert regularities may speed responses. This implicit acquisition of regularities may become explicitly conscious at some point, leading to a qualitative change in behavior which reflects this insight. By applying the NRT at two consecutive sessions separated by an interval, we investigated the role of sleep in this interval for attaining insight at the second session. In the first study, a night of sleep was shown to triple the number of participants attaining insight above the base rate of about 20%. In the second study, this hard core of 20% discoverers differed from other participants in their task-related EEG potentials from the very beginning already. In the third study, the additional role of sleep was specified as an effect of the deep-sleep phase of slow-wave sleep on participants who had implicitly acquired the covert regularity before sleep. It was in these participants that a specific increase of EEG during slow-wave sleep in the 10-12 Hz band was obtained. These results support the view that neuronal memory reprocessing during slow-wave sleep restructures task-related representations in the brain, and that such restructuring promotes the gain of explicit knowledge.