An Enduring Role for Hippocampal Pattern Completion in Addition to an Emergent Nonhippocampal Contribution to Holistic Episodic Retrieval after a 24†h Delay.

Episodic memory retrieval is associated with the holistic neocortical reinstatement of all event information, an effect driven by hippocampal pattern completion. However, whether holistic reinstatement occurs, and whether hippocampal pattern completion continues to drive reinstatement, after a period of consolidation is unclear. Theories of systems consolidation predict either a time-variant or time-invariant role of the hippocampus in the holistic retrieval of episodic events. Here, we assessed whether episodic events continue to be reinstated holistically and whether hippocampal pattern completion continues to facilitate holistic reinstatement following a period of consolidation. Female and male human participants learned "events" that comprised multiple overlapping pairs of event elements (e.g., person-location, object-location, location-person). Importantly, encoding occurred either immediately before or 24 h before retrieval. Using fMRI during the retrieval of events, we show evidence for holistic reinstatement, as well as a correlation between reinstatement and hippocampal pattern completion, regardless of whether retrieval occurred immediately or 24 h after encoding. Thus, hippocampal pattern completion continues to contribute to holistic reinstatement after a delay. However, our results also revealed that some holistic reinstatement can occur without evidence for a corresponding signature of hippocampal pattern completion after a delay (but not immediately after encoding). We therefore show that hippocampal pattern completion, in addition to a nonhippocampal process, has a role in holistic reinstatement following a period of consolidation. Our results point to a consolidation process where the hippocampus and neocortex may work in an additive, rather than compensatory, manner to support episodic memory retrieval.

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.

Sleep loss disrupts the neural signature of successful learning.

Sleep supports memory consolidation as well as next-day learning. The influential "Active Systems" account of offline consolidation suggests that sleep-associated memory processing paves the way for new learning, but empirical evidence in support of this idea is scarce. Using a within-subjects (n = 30), crossover design, we assessed behavioral and electrophysiological indices of episodic encoding after a night of sleep or total sleep deprivation in healthy adults (aged 18-25 years) and investigated whether behavioral performance was predicted by the overnight consolidation of episodic associations from the previous day. Sleep supported memory consolidation and next-day learning as compared to sleep deprivation. However, the magnitude of this sleep-associated consolidation benefit did not significantly predict the ability to form novel memories after sleep. Interestingly, sleep deprivation prompted a qualitative change in the neural signature of encoding: Whereas 12-20 Hz beta desynchronization-an established marker of successful encoding-was observed after sleep, sleep deprivation disrupted beta desynchrony during successful learning. Taken together, these findings suggest that effective learning depends on sleep but not necessarily on sleep-associated consolidation.

Targeted memory reactivation during sleep can induce forgetting of overlapping memories.

Memory reactivation during sleep can shape new memories into a long-term form. Reactivation of memories can be induced via the delivery of auditory cues during sleep. Although this targeted memory reactivation (TMR) approach can strengthen newly acquired memories, research has tended to focus on single associative memories. It is less clear how TMR affects retention for overlapping associative memories. This is critical, given that repeated retrieval of overlapping associations during wake can lead to forgetting, a phenomenon known as retrieval-induced forgetting (RIF). We asked whether a similar pattern of forgetting occurs when TMR is used to cue reactivation of overlapping pairwise associations during sleep. Participants learned overlapping pairs-learned separately, interleaved with other unrelated pairs. During sleep, we cued a subset of overlapping pairs using TMR. While TMR increased retention for the first encoded pairs, memory decreased for the second encoded pairs. This pattern of retention was only present for pairs not tested prior to sleep. The results suggest that TMR can lead to forgetting, an effect similar to RIF during wake. However, this effect did not extend to memories that had been strengthened via retrieval prior to sleep. We therefore provide evidence for a reactivation-induced forgetting effect during sleep.

Nap effects on preschool children's learning of letter-sound mappings.

This study explored whether a daytime nap aids children's acquisition of letter-sound knowledge, which is a fundamental component for learning to read. Thirty-two preschool children in Sydney, Australia (Mage  = 4 years;3 months) were taught letter-sound mappings in two sessions: one followed by a nap and the other by a wakeful period. Learning was assessed by explicit letter-sound mappings ("Which sound does this letter make?") and knowledge generalization tasks ("Here's Tav and Cav, which one is /kav/?"). Results from the knowledge generalization task showed better performance after a nap than after wake. However, no nap benefit was found for explicit letter-sound knowledge. This study provides initial evidence that naps could be beneficial for preschool children's learning of letter-sound mappings.

Does the maturation of early sleep patterns predict language ability at school entry? A Born in Bradford study.

Children's vocabulary ability at school entry is highly variable and predictive of later language and literacy outcomes. Sleep is potentially useful in understanding and explaining that variability, with sleep patterns being predictive of global trajectories of language acquisition. Here, we looked to replicate and extend these findings. Data from 354 children (without English as an additional language) in the Born in Bradford study were analysed, describing the mean intercepts and linear trends in parent-reported day-time and night-time sleep duration over five time points between 6 and 36 months-of-age. The mean difference between night-time and day-time sleep was predictive of receptive vocabulary at age five, with more night-time sleep relative to day-time sleep predicting better language. An exploratory analysis suggested that socioeconomic status was predictive of vocabulary outcomes, with sleep patterns partially mediating this relationship. We suggest that the consolidation of sleep patterns acts as a driver of early language development.

Sleep-dependent consolidation in children with comprehension and vocabulary weaknesses: it'll be alright on the night?

BACKGROUND: Vocabulary is crucial for an array of life outcomes and is frequently impaired in developmental disorders. Notably, 'poor comprehenders' (children with reading comprehension deficits but intact word reading) often have vocabulary deficits, but underlying mechanisms remain unclear. Prior research suggests intact encoding but difficulties consolidating new word knowledge. We test the hypothesis that poor comprehenders' sleep-associated vocabulary consolidation is compromised by their impoverished lexical-semantic knowledge. METHODS: Memory for new words was tracked across wake and sleep to assess encoding and consolidation in 8-to-12-year-old good and poor comprehenders. Each child participated in two sets of sessions, one beginning in the morning (AM-encoding) and the other in the evening (PM-encoding). In each case, they were taught 12 words and were trained on a spatial memory task. Memory was assessed immediately, 12- and 24-hr later via stem-completion, picture-naming, and definition tasks to probe different aspects of word knowledge. Long-term retention was assessed 1-2 months later. RESULTS: Recall of word-forms improved over sleep and postsleep wake, as measured in both stem-completion and picture-naming tasks. Counter to hypotheses, deficits for poor comprehenders were not observed in consolidation but instead were seen across measures and throughout testing, suggesting a deficit from encoding. Variability in vocabulary knowledge across the whole sample predicted sleep-associated consolidation, but only when words were learned early in the day and not when sleep followed soon after learning. CONCLUSIONS: Poor comprehenders showed weaker memory for new words than good comprehenders, but sleep-associated consolidation benefits were comparable between groups. Sleeping soon after learning had long-lasting benefits for memory and may be especially beneficial for children with weaker vocabulary. These results provide new insights into the breadth of poor comprehenders' vocabulary weaknesses, and ways in which learning might be better timed to remediate vocabulary difficulties.

Atypicalities in sleep and semantic consolidation in autism.

Sleep is known to support the neocortical consolidation of declarative memory, including the acquisition of new language. Autism spectrum disorder (ASD) is often characterized by both sleep and language learning difficulties, but few studies have explored a potential connection between the two. Here, 54 children with and without ASD (matched on age, nonverbal ability and vocabulary) were taught nine rare animal names (e.g., pipa). Memory was assessed via definitions, naming and speeded semantic decision tasks immediately after learning (pre-sleep), the next day (post-sleep, with a night of polysomnography between pre- and post-sleep tests) and roughly 1 month later (follow-up). Both groups showed comparable performance at pre-test and similar levels of overnight change on all tasks; but at follow-up children with ASD showed significantly greater forgetting of the unique features of the new animals (e.g., pipa is a flat frog). Children with ASD had significantly lower central non-rapid eye movement (NREM) sigma power. Associations between spindle properties and overnight changes in speeded semantic decisions differed by group. For the TD group, spindle duration predicted overnight changes in responses to novel animals but not familiar animals, reinforcing a role for sleep in the stabilization of new semantic knowledge. For the ASD group, sigma power and spindle duration were associated with improvements in responses to novel and particularly familiar animals, perhaps reflecting more general sleep-associated improvements in task performance. Plausibly, microstructural sleep atypicalities in children with ASD and differences in how information is prioritized for consolidation may lead to cumulative consolidation difficulties, compromising the quality of newly formed semantic representations in long-term memory.

The nature of delayed dream incorporation ('dream-lag effect'): Personally significant events persist, but not major daily activities or concerns.

Incorporation of details from waking life events into rapid eye movement (REM) sleep dreams has been found to be highest on the 2 nights after, and then 5-7 nights after, the event. These are termed, respectively, the day-residue and dream-lag effects. This study is the first to categorize types of waking life experiences and compare their incorporation into dreams across multiple successive nights. Thirty-eight participants completed a daily diary each evening and a dream diary each morning for 14 days. In the daily diary, three categories of experiences were reported: major daily activities (MDAs), personally significant events (PSEs) and major concerns (MCs). After the 14-day period each participant identified the correspondence between items in their daily diaries and subsequent dream reports. The day-residue and dream-lag effects were found for the incorporation of PSEs into dreams (effect sizes of .33 and .27, respectively), but only for participants (n = 19) who had a below-median total number of correspondences between daily diary items and dream reports (termed "low-incorporators" as opposed to "high-incorporators"). Neither the day-residue or dream-lag effects were found for MDAs or MCs. This U-shaped timescale of incorporation of events from daily life into dreams has been proposed to reflect REM sleep-dependent memory consolidation, possibly related to emotional memory processing. This study had a larger sample size of dreams than any dream-lag study hitherto with trained participants. Coupled with previous successful replications, there is thus substantial evidence supporting the dream-lag effect and further explorations of its mechanism, including its neural underpinnings, are warranted.

Offline consolidation supersedes prior knowledge benefits in children's (but not adults') word learning.

Prior linguistic knowledge is proposed to support the acquisition and consolidation of new words. Adults typically have larger vocabularies to support word learning than children, but the developing brain shows enhanced neural processes that are associated with offline memory consolidation. This study investigated contributions of prior knowledge to initial word acquisition and consolidation at different points in development, by teaching children and adults novel words (e.g., ballow) that varied in the number of English word-form "neighbours" (e.g., wallow, bellow). Memory for the novel word-forms was tested immediately after training, the next day and 1 week later, to assess the time-course of prior knowledge contributions. Children aged 7-9 years (Experiments 1, 3) and adults (Experiment 2) recalled words with neighbours better than words without neighbours when tested immediately after training. However, a period of offline consolidation improved overall recall and reduced the influence of word-form neighbours on longer term memory. These offline consolidation benefits were larger in children than adults, supporting theories that children have a greater propensity for consolidating phonologically distinctive language information. Local knowledge of just a single word-form neighbour was enough to enhance learning, and this led to the individual differences in word recall that were related to adults' global vocabulary ability. The results support the proposal that the relative contributions of different learning mechanisms change across the lifespan, and highlight the importance of testing theoretical models of word learning in the context of development.

No effect of targeted memory reactivation during slow-wave sleep on emotional recognition memory.

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.

Consolidation of vocabulary is associated with sleep in typically developing children, but not in children with dyslexia.

Sleep is known to play an active role in consolidating new vocabulary in adults; however, the mechanisms by which sleep promotes vocabulary consolidation in childhood are less well understood. Furthermore, there has been no investigation into whether previously reported differences in sleep architecture might account for variability in vocabulary consolidation in children with dyslexia. Twenty-three children with dyslexia and 29 age-matched typically developing peers were exposed to 16 novel spoken words. Typically developing children showed overnight improvements in novel word recall; the size of the improvement correlated positively with slow wave activity, similar to previous findings with adults. Children with dyslexia showed poorer recall of the novel words overall, but nevertheless showed overnight improvements similar to age-matched peers. However, comparisons with younger children matched on initial levels of novel word recall pointed to reduced consolidation in dyslexics after 1 week. Crucially, there were no significant correlations between overnight consolidation and sleep parameters in the dyslexic group. This suggests a reduced role of sleep in vocabulary consolidation in dyslexia, possibly as a consequence of lower levels of learning prior to sleep, and highlights how models of sleep-associated memory consolidation can be usefully informed by data from typical and atypical development.

Accent modulates access to word meaning: Evidence for a speaker-model account of spoken word recognition.

Speech carries accent information relevant to determining the speaker's linguistic and social background. A series of web-based experiments demonstrate that accent cues can modulate access to word meaning. In Experiments 1-3, British participants were more likely to retrieve the American dominant meaning (e.g., hat meaning of "bonnet") in a word association task if they heard the words in an American than a British accent. In addition, results from a speeded semantic decision task (Experiment 4) and sentence comprehension task (Experiment 5) confirm that accent modulates on-line meaning retrieval such that comprehension of ambiguous words is easier when the relevant word meaning is dominant in the speaker's dialect. Critically, neutral-accent speech items, created by morphing British- and American-accented recordings, were interpreted in a similar way to accented words when embedded in a context of accented words (Experiment 2). This finding indicates that listeners do not use accent to guide meaning retrieval on a word-by-word basis; instead they use accent information to determine the dialectic identity of a speaker and then use their experience of that dialect to guide meaning access for all words spoken by that person. These results motivate a speaker-model account of spoken word recognition in which comprehenders determine key characteristics of their interlocutor and use this knowledge to guide word meaning access.

Sleep underpins the plasticity of language production.

The constraints that govern acceptable phoneme combinations in speech perception and production have considerable plasticity. We addressed whether sleep influences the acquisition of new constraints and their integration into the speech-production system. Participants repeated sequences of syllables in which two phonemes were artificially restricted to syllable onset or syllable coda, depending on the vowel in that sequence. After 48 sequences, participants either had a 90-min nap or remained awake. Participants then repeated 96 sequences so implicit constraint learning could be examined, and then were tested for constraint generalization in a forced-choice task. The sleep group, but not the wake group, produced speech errors at test that were consistent with restrictions on the placement of phonemes in training. Furthermore, only the sleep group generalized their learning to new materials. Polysomnography data showed that implicit constraint learning was associated with slow-wave sleep. These results show that sleep facilitates the integration of new linguistic knowledge with existing production constraints. These data have relevance for systems-consolidation models of sleep.

The selective role of premotor cortex in speech perception: a contribution to phoneme judgements but not speech comprehension.

Several accounts of speech perception propose that the areas involved in producing language are also involved in perceiving it. In line with this view, neuroimaging studies show activation of premotor cortex (PMC) during phoneme judgment tasks; however, there is debate about whether speech perception necessarily involves motor processes, across all task contexts, or whether the contribution of PMC is restricted to tasks requiring explicit phoneme awareness. Some aspects of speech processing, such as mapping sounds onto meaning, may proceed without the involvement of motor speech areas if PMC specifically contributes to the manipulation and categorical perception of phonemes. We applied TMS to three sites-PMC, posterior superior temporal gyrus, and occipital pole-and for the first time within the TMS literature, directly contrasted two speech perception tasks that required explicit phoneme decisions and mapping of speech sounds onto semantic categories, respectively. TMS to PMC disrupted explicit phonological judgments but not access to meaning for the same speech stimuli. TMS to two further sites confirmed that this pattern was site specific and did not reflect a generic difference in the susceptibility of our experimental tasks to TMS: stimulation of pSTG, a site involved in auditory processing, disrupted performance in both language tasks, whereas stimulation of occipital pole had no effect on performance in either task. These findings demonstrate that, although PMC is important for explicit phonological judgments, crucially, PMC is not necessary for mapping speech onto meanings.

A registered report testing the effect of sleep on Deese-Roediger-McDermott false memory: greater lure and veridical recall but fewer intrusions after sleep.

Human memory is known to be supported by sleep. However, less is known about the effect of sleep on false memory, where people incorrectly remember events that never occurred. In the laboratory, false memories are often induced via the Deese-Roediger-McDermott (DRM) paradigm where participants are presented with wordlists comprising semantically related words such as nurse, hospital and sick (studied words). Subsequently, participants are likely to falsely remember that a related lure word such as doctor was presented. Multiple studies have examined whether these false memories are influenced by sleep, with contradictory results. A recent meta-analysis suggests that sleep may increase DRM false memory when short lists are used. We tested this in a registered report (N = 488) with a 2 (Interval: Immediate versus 12 h delay) × 2 (Test Time: 9:00 versus 21:00) between-participant DRM experiment, using short DRM lists (N = 8 words/list) and free recall as the memory test. We found an unexpected time-of-day effect such that completing free recall in the evening led to more intrusions (neither studied nor lure words). Above and beyond this time-of-day effect, the Sleep participants produced fewer intrusions than their Wake counterparts. When this was statistically controlled for, the Sleep participants falsely produced more critical lures. They also correctly recalled more studied words (regardless of intrusions). Exploratory analysis showed that these findings cannot be attributed to differences in output bias, as indexed by the number of total responses. Our overall results cannot be fully captured by existing sleep-specific theories of false memory, but help to define the role of sleep in two more general theories (Fuzzy-Trace and Activation/Monitoring theories) and suggest that sleep may benefit gist abstraction/spreading activation on one hand and memory suppression/source monitoring on the other.

Episodic memory and sleep are involved in the maintenance of context-specific lexical information.

Familiar words come with a wealth of associated knowledge about their variety of usage, accumulated over a lifetime. How do we track and adjust this knowledge as new instances of a word are encountered? A recent study (Cognition) found that, for homonyms (e.g., bank), sleep-associated consolidation facilitates the updating of meaning dominance. Here, we tested the generality of this finding by exposing participants to (Experiment 1; N = 125) nonhomonyms (e.g., bathtub) in sentences that biased their meanings toward a specific interpretation (e.g., bathtub-slip vs. bathtub-relax), and (Experiment 2; N = 128) word-class ambiguous words (e.g., loan) in sentences where the words were used in their dispreferred word class (e.g., "He will loan me money"). Both experiments showed that such sentential experience influenced later interpretation and usage of the words more after a night's sleep than a day awake. We interpret these results as evidence for a general role of episodic memory in language comprehension such that new episodic memories are formed every time a sentence is comprehended, and these memories contribute to lexical processing next time the word is encountered, as well as potentially to the fine-tuning of long-term lexical knowledge. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Learning new meanings for old words: effects of semantic relatedness.

Changes to our everyday activities mean that adult language users need to learn new meanings for previously unambiguous words. For example, we need to learn that a "tweet" is not only the sound a bird makes, but also a short message on a social networking site. In these experiments, adult participants learned new fictional meanings for words with a single dominant meaning (e.g., "ant") by reading paragraphs that described these novel meanings. Explicit recall of these meanings was significantly better when there was a strong semantic relationship between the novel meaning and the existing meaning. This relatedness effect emerged after relatively brief exposure to the meanings (experiment 1), but it persisted when training was extended across 7 days (experiment 2) and when semantically demanding tasks were used during this extended training (experiment 3). A lexical decision task was used to assess the impact of learning on online recognition. In Experiment 3, participants responded more quickly to words whose new meaning was semantically related than to those with an unrelated meaning. This result is consistent with earlier studies showing an effect of meaning relatedness on lexical decision, and it indicates that these newly acquired meanings become integrated with participants' preexisting knowledge about the meanings of words.

Schematic information influences memory and generalisation behaviour for schema-relevant and -irrelevant information.

Schemas modulate memory performance for schema-congruent and -incongruent information. However, it is assumed they do not influence behaviour for information irrelevant to themselves. We assessed memory and generalisation behaviour for information related to an underlying pattern, where a schema could be extracted (schema-relevant), and information that was unrelated and therefore irrelevant to the extracted schema (schema-irrelevant). Using precision measures of long-term memory, where participants learnt associations between words and locations around a circle, we assessed memory and generalisation for schema-relevant and -irrelevant information. Words belonged to two semantic categories: human-made and natural. For one category, word-locations were clustered around one point on the circle (clustered condition), while the other category had word-locations randomly distributed (non-clustered condition). The presence of an underlying pattern in the clustered condition allows for the extraction of a schema that can support both memory and generalisation. At test, participants were presented with old (memory) and new (generalisation) words, requiring them to identify a remembered location or make a best guess. The presence of the clustered pattern modulated memory and generalisation. In the clustered condition, participants placed old and new words in locations consistent with the underlying pattern. In contrast, for the non-clustered condition, participants were less likely to place old and new non-clustered words in locations consistent with the clustered condition. Therefore, we provide evidence that the presence of schematic information modulates memory and generalisation for schema-relevant and -irrelevant information. Our results highlight the need to carefully construct appropriate schema-irrelevant control conditions such that behaviour in these conditions is not modulated by the presence of a schema. Theoretically, models of schema processing need to account for how the presence of schematic information can have consequences for information that is irrelevant to itself.

UK children's sleep and anxiety during the COVID-19 pandemic.

BACKGROUND: Sleep and mental wellbeing are intimately linked. This relationship is particularly important to understand as it emerges over childhood. Here we take the opportunity that the COVID-19 pandemic, and resulting lockdown in the UK, presented to study sleep-related behaviour and anxiety in school-aged children. METHODS: Parents and children were asked to complete questionnaires towards the start of the UK lockdown in April-to-May of 2020, then again in August of that year (when many restrictions had been lifted). We explored children's emotional responses to the pandemic and sleep patterns at both time points, from the perspectives of parents and children themselves. RESULTS: Children's bedtime anxiety increased at the start of the lockdown as compared to a typical week; however, by August, bedtime anxiety had ameliorated along with children's COVID-19 related anxiety. Bedtime anxiety predicted how long it took children to fall asleep at night at both the start and the end of the lockdown. Bedtime and wake-up time shifted at the start of lockdown, but interestingly total sleep time was resilient (likely owing to an absence of early school start times) and was not predicted by child anxiety. CONCLUSIONS: These findings further support calls for sleep quality (in particular, time taken to fall asleep) to be taken as a key indicator of mental health in children, particularly under usual circumstances when schools are open and sleep duration may be less resilient.