Considerations towards a neurobiologically-informed EEG measurement of sleepiness.

Sleep is a daily experience across humans and other species, yet our understanding of how and why we sleep is presently incomplete. This is particularly prevalent in research examining the neurophysiological measurement of sleepiness in humans, where several electroencephalogram (EEG) phenomena have been linked with prolonged wakefulness. This leaves researchers without a solid basis for the measurement of homeostatic sleep need and complicates our understanding of the nature of sleep. Recent theoretical and technical advances may allow for a greater understanding of the neurobiological basis of homeostatic sleep need: this may result from increases in neuronal excitability and shifts in excitation/inhibition balance in neuronal circuits and can potentially be directly measured via the aperiodic component of the EEG. Here, we review the literature on EEG-derived markers of sleepiness in humans and argue that changes in these electrophysiological markers may actually result from neuronal activity represented by changes in aperiodic markers. We argue for the use of aperiodic markers derived from the EEG in predicting sleepiness and suggest areas for future research based on these.

Prior Context and Individual Alpha Frequency Influence Predictive Processing during Language Comprehension.

The extent to which the brain predicts upcoming information during language processing remains controversial. To shed light on this debate, the present study reanalyzed Nieuwland and colleagues' (2018) [Nieuwland, M. S., Politzer-Ahles, S., Heyselaar, E., Segaert, K., Darley, E., Kazanina, N., et al. Large-scale replication study reveals a limit on probabilistic prediction in language comprehension. eLife, 7, e33468, 2018] replication of DeLong and colleagues (2015) [DeLong, K. A., Urbach, T. P., & Kutas, M. Probabilistic word pre-activation during language comprehension inferred from electrical brain activity. Nature Neuroscience, 8, 1117-1121, 2005]. Participants (n = 356) viewed sentences containing articles and nouns of varying predictability, while their EEG was recorded. We measured ERPs preceding the critical words (namely, the semantic prediction potential), in conjunction with postword N400 patterns and individual neural metrics. ERP activity was compared with two measures of word predictability: cloze probability and lexical surprisal. In contrast to prior literature, semantic prediction potential amplitudes did not increase as cloze probability increased, suggesting that the component may not reflect prediction during natural language processing. Initial N400 results at the article provided evidence against phonological prediction in language, in line with Nieuwland and colleagues' findings. Strikingly, however, when the surprisal of the prior words in the sentence was included in the analysis, increases in article surprisal were associated with increased N400 amplitudes, consistent with prediction accounts. This relationship between surprisal and N400 amplitude was not observed when the surprisal of the two prior words was low, suggesting that expectation violations at the article may be overlooked under highly predictable conditions. Individual alpha frequency also modulated the relationship between article surprisal and the N400, emphasizing the importance of individual neural factors for prediction. The present study extends upon existing neurocognitive models of language and prediction more generally, by illuminating the flexible and subject-specific nature of predictive processing.

Augmenting complex and dynamic performance through mindfulness-based cognitive training: An evaluation of training adherence, trait mindfulness, personality and resting-state EEG.

Human performance applications of mindfulness-based training have demonstrated its utility in enhancing cognitive functioning. Previous studies have illustrated how these interventions can improve performance on traditional cognitive tests, however, little investigation has explored the extent to which mindfulness-based training can optimise performance in more dynamic and complex contexts. Further, from a neuroscientific perspective, the underlying mechanisms responsible for performance enhancements remain largely undescribed. With this in mind, the following study aimed to investigate how a short-term mindfulness intervention (one week) augments performance on a dynamic and complex task (target motion analyst task; TMA) in young, healthy adults (n = 40, age range = 18-38). Linear mixed effect modelling revealed that increased adherence to the web-based mindfulness-based training regime (ranging from 0-21 sessions) was associated with improved performance in the second testing session of the TMA task, controlling for baseline performance. Analyses of resting-state electroencephalographic (EEG) metrics demonstrated no change across testing sessions. Investigations of additional individual factors demonstrated that enhancements associated with training adherence remained relatively consistent across varying levels of participants' resting-state EEG metrics, personality measures (i.e., trait mindfulness, neuroticism, conscientiousness), self-reported enjoyment and timing of intervention adherence. Our results thus indicate that mindfulness-based cognitive training leads to performance enhancements in distantly related tasks, irrespective of several individual differences. We also revealed nuances in the magnitude of cognitive enhancements contingent on the timing of adherence, regardless of total volume of training. Overall, our findings suggest that mindfulness-based training could be used in a myriad of settings to elicit transferable performance enhancements.

Awareness of errors is reduced by sleep loss.

The ability to detect and subsequently correct errors is important in preventing the detrimental consequences of sleep loss. The Error Related Negativity (ERN), and the error positivity (Pe) are established neural correlates of error processing. Previous work has shown sleep loss reduces ERN and Pe, indicating sleep loss impairs error-monitoring processes. However, no previous work has examined behavioral error awareness, in conjunction with EEG measures, under sleep loss conditions, and studies of sleep restriction are lacking. Using combined behavioral and EEG measures, we report two studies investigating the impact of total sleep deprivation (TSD) and sleep restriction (SR) on error awareness. Fourteen healthy participants completed the Error Awareness Task under conditions of TSD and 27 completed the same task under conditions of SR. It was found that TSD did not influence behavioral error awareness or ERN or Pe amplitude, however, SR reduced behavioral error awareness, increased the time taken to detect errors, and reduced Pe amplitude. Findings indicate individuals who are chronically sleep restricted are at risk for reduced recognition of errors. Reduced error awareness may be one factor contributing to the increased accidents and injuries seen in contexts where sleep loss is prevalent.

Individual differences in information processing during sleep and wake predict sleep-based memory consolidation of complex rules.

Memory is critical for many cognitive functions, from remembering facts, to learning complex environmental rules. While memory encoding occurs during wake, memory consolidation is associated with sleep-related neural activity. Further, research suggests that individual differences in alpha frequency during wake (∼7 - 13 Hz) modulate memory processes, with higher individual alpha frequency (IAF) associated with greater memory performance. However, the relationship between wake-related EEG individual differences, such as IAF, and sleep-related neural correlates of memory consolidation has been largely unexplored, particularly in a complex rule-based memory context. Here, we aimed to investigate whether wake-derived IAF and sleep neurophysiology interact to influence rule learning in a sample of 35 healthy adults (16 males; mean age = 25.4, range: 18 - 40). Participants learned rules of a modified miniature language prior to either 8hrs of sleep or wake, after which they were tested on their knowledge of the rules in a grammaticality judgement task. Results indicate that sleep neurophysiology and wake-derived IAF do not interact but modulate memory for complex linguistic rules separately. Phase-amplitude coupling between slow oscillations and spindles during non-rapid eye-movement (NREM) sleep also promoted memory for rules that were analogous to the canonical English word order. As an exploratory analysis, we found that rapid eye-movement (REM) sleep theta power at posterior regions interacts with IAF to predict rule learning and proportion of time in REM sleep predicts rule learning differentially depending on grammatical rule type. Taken together, the current study provides behavioural and electrophysiological evidence for a complex role of NREM and REM sleep neurophysiology and wake-derived IAF in the consolidation of rule-based information.

Neural and cognitive correlates of performance in dynamic multi-modal settings.

The endeavour to understand human cognition has largely relied upon investigation of task-related brain activity. However, resting-state brain activity can also offer insights into individual information processing and performance capabilities. Previous research has identified electroencephalographic resting-state characteristics (most prominently: the individual alpha frequency; IAF) that predict cognitive function. However, it has largely overlooked a second component of electrophysiological signals: aperiodic 1/ƒ activity. The current study examined how both oscillatory and aperiodic resting-state EEG measures, alongside traditional cognitive tests, can predict performance in a dynamic and complex, semi-naturalistic cognitive task. Participants' resting-state EEG was recorded prior to engaging in a Target Motion Analysis (TMA) task in a simulated submarine control room environment (CRUSE), which required participants to integrate dynamically changing information over time. We demonstrated that the relationship between IAF and cognitive performance extends from simple cognitive tasks (e.g., digit span) to complex, dynamic measures of information processing. Further, our results showed that individual 1/ƒ parameters (slope and intercept) differentially predicted performance across practice and testing sessions, whereby flatter slopes and higher intercepts were associated with improved performance during learning. In addition to the EEG predictors, we demonstrate a link between cognitive skills most closely related to the TMA task (i.e., spatial imagery) and subsequent performance. Overall, the current study highlights (1) how resting-state metrics - both oscillatory and aperiodic - have the potential to index higher-order cognitive capacity, while (2) emphasising the importance of examining these electrophysiological components within more dynamic settings and over time.

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.

Task-related, intrinsic oscillatory and aperiodic neural activity predict performance in naturalistic team-based training scenarios.

Effective teams are essential for optimally functioning societies. However, little is known regarding the neural basis of two or more individuals engaging cooperatively in real-world tasks, such as in operational training environments. In this exploratory study, we recruited forty individuals paired as twenty dyads and recorded dual-EEG at rest and during realistic training scenarios of increasing complexity using virtual simulation systems. We estimated markers of intrinsic brain activity (i.e., individual alpha frequency and aperiodic activity), as well as task-related theta and alpha oscillations. Using nonlinear modelling and a logistic regression machine learning model, we found that resting-state EEG predicts performance and can also reliably differentiate between members within a dyad. Task-related theta and alpha activity during easy training tasks predicted later performance on complex training to a greater extent than prior behaviour. These findings complement laboratory-based research on both oscillatory and aperiodic activity in higher-order cognition and provide evidence that theta and alpha activity play a critical role in complex task performance in team environments.

Oscillatory and Aperiodic Neural Activity Jointly Predict Language Learning.

Memory formation involves the synchronous firing of neurons in task-relevant networks, with recent models postulating that a decrease in low-frequency oscillatory activity underlies successful memory encoding and retrieval. However, to date, this relationship has been investigated primarily with face and image stimuli; considerably less is known about the oscillatory correlates of complex rule learning, as in language. Furthermore, recent work has shown that nonoscillatory (1/ƒ) activity is functionally relevant to cognition, yet its interaction with oscillatory activity during complex rule learning remains unknown. Using spectral decomposition and power-law exponent estimation of human EEG data (17 women, 18 men), we show for the first time that 1/ƒ and oscillatory activity jointly influence the learning of word order rules of a miniature artificial language system. Flexible word-order rules were associated with a steeper 1/ƒ slope, whereas fixed word-order rules were associated with a shallower slope. We also show that increased theta and alpha power predicts fixed relative to flexible word-order rule learning and behavioral performance. Together, these results suggest that 1/ƒ activity plays an important role in higher-order cognition, including language processing, and that grammar learning is modulated by different word-order permutations, which manifest in distinct oscillatory profiles.

Theta Activity During Encoding Interacts With NREM Sleep Oscillations to Predict Memory Generalization.

Relatively little is known regarding the interaction between encoding-related neural activity and sleep-based memory consolidation. One suggestion is that a function of encoding-related theta power may be to "tag" memories for subsequent processing during sleep. This study aimed to extend previous work on the relationships between sleep spindles, slow oscillation-spindle coupling, and task-related theta activity with a combined Deese-Roediger-McDermott (DRM) and nap paradigm. This allowed us to examine the influence of task- and sleep-related oscillatory activity on the recognition of both encoded list words and associative theme words. Thirty-three participants (29 females, mean age = 23.2 years) learned and recognised DRM lists separated by either a 2 h wake or sleep period. Mixed-effects modelling revealed the sleep condition endorsed more associative theme words and fewer list words in comparison to the wake group. Encoding-related theta power was also found to influence sleep spindle density, and this interaction was predictive of memory outcomes. The influence of encoding-related theta was specific to sleep spindle density, and did not appear to influence the strength of slow oscillation-spindle coupling as it relates to memory outcomes. The finding of interactions between wakeful and sleep oscillatory-related activity in promoting memory and learning has important implications for theoretical models of sleep-based memory consolidation.

Gender bias in academia: A lifetime problem that needs solutions.

Despite increased awareness of the lack of gender equity in academia and a growing number of initiatives to address issues of diversity, change is slow, and inequalities remain. A major source of inequity is gender bias, which has a substantial negative impact on the careers, work-life balance, and mental health of underrepresented groups in science. Here, we argue that gender bias is not a single problem but manifests as a collection of distinct issues that impact researchers' lives. We disentangle these facets and propose concrete solutions that can be adopted by individuals, academic institutions, and society.

Resting-state aperiodic neural dynamics predict individual differences in visuomotor performance and learning.

An emerging body of work has demonstrated that resting-state non-oscillatory, or aperiodic, 1/f neural activity is a functional and behaviorally relevant marker of cognitive function capacity. In the motor domain, previous work has only applied 1/f analyses to investigations of motor coordination and performance measures. The value of aperiodic resting-state neural dynamics as a marker of individual visuomotor performance capacity remains unknown. Accordingly, the aim of this work was to investigate if individual 1/f intercept and slope parameters of aperiodic resting-state neural activity predict reaction time and perceptual sensitivity in an immersive virtual reality marksmanship task. The marksmanship task required speeded selection of target stimuli and avoidance of selecting non-target stimuli. Motor and perceptual demands were incrementally increased across task blocks and participants performed the task across three training sessions spanning one week. When motor demands were high, steeper individual 1/f slope predicted shorter reaction time. This relationship did not change with practice. Increased 1/f intercept and a steeper 1/f slope were associated with higher perceptual sensitivity, measured as d'. However, this association was only observed under the highest levels of perceptual demand and only in the initial exposure to these conditions. Individuals with a lower 1/f intercept and a shallower 1/f slope demonstrated the greatest gains in perceptual sensitivity from task practice. These findings demonstrate that individual differences in motor and perceptual performance can be accounted for with resting-state aperiodic neural dynamics. The 1/f aperiodic parameters are most informative in predicting visuomotor performance under complex and demanding task conditions. In addition to predicting capacity for high visuomotor performance with a novel task, 1/f aperiodic parameters might also be useful in predicting which individuals might derive the most improvements from practice.

Mini Pinyin: A modified miniature language for studying language learning and incremental sentence processing.

Artificial grammar learning (AGL) paradigms are used extensively to characterise (neuro)cognitive bases of language learning. However, despite their effectiveness in characterising the capacity to learn complex structured sequences, AGL paradigms lack ecological validity and typically do not account for cross-linguistic differences in sentence comprehension. Here, we describe a new modified miniature language paradigm - Mini Pinyin - that mimics natural language as it is based on an existing language (Mandarin Chinese) and includes both structure and meaning. Mini Pinyin contains a number of cross-linguistic elements, including varying word orders and classifier-noun rules. To evaluate the effectiveness of Mini Pinyin, 76 (mean age = 24.9; 26 female) monolingual native English speakers completed a learning phase followed by a sentence acceptability judgement task. Generalised mixed effects modelling revealed that participants attained a moderate degree of accuracy on the judgement task, with performance scores ranging from 25% to 100% accuracy depending on the word order of the sentence. Further, sentences compatible with the canonical English word order were learned more efficiently than non-canonical word orders. We controlled for inter-individual differences in statistical learning ability, which accounted for ~20% of the variance in performance on the sentence judgement task. We provide stimuli and statistical analysis scripts as open-source resources and discuss how future research can utilise this paradigm to study the neurobiological basis of language learning. Mini Pinyin affords a convenient tool for improving the future of language learning research by building on the parameters of traditional AGL or existing miniature language paradigms.

Individual alpha frequency modulates sleep-related emotional memory consolidation.

Alpha-band oscillatory activity is involved in modulating memory and attention. However, few studies have investigated individual differences in oscillatory activity during the encoding of emotional memory, particularly in sleep paradigms where sleep is thought to play an active role in memory consolidation. The current study aimed to address the question of whether individual alpha frequency (IAF) modulates the consolidation of declarative memory across periods of sleep and wake. 22 participants aged 18-41 years (mean age = 25.77) viewed 120 emotionally valenced images (positive, negative, neutral) and completed a baseline memory task before a 2hr afternoon sleep opportunity and an equivalent period of wake. Following the sleep and wake conditions, participants were required to distinguish between 120 learned (target) images and 120 new (distractor) images. This method allowed us to delineate the role of different oscillatory components of sleep and wake states in the emotional modulation of memory. Linear mixed-effects models revealed interactions between IAF, rapid eye movement sleep theta power, and slow-wave sleep slow oscillatory density on memory outcomes. These results highlight the importance of individual factors in the EEG in modulating oscillatory-related memory consolidation and subsequent behavioural outcomes and test predictions proposed by models of sleep-based memory consolidation.

Sleep-Dependent Memory Consolidation and Incremental Sentence Comprehension: Computational Dependencies during Language Learning as Revealed by Neuronal Oscillations.

We hypothesize a beneficial influence of sleep on the consolidation of the combinatorial mechanisms underlying incremental sentence comprehension. These predictions are grounded in recent work examining the effect of sleep on the consolidation of linguistic information, which demonstrate that sleep-dependent neurophysiological activity consolidates the meaning of novel words and simple grammatical rules. However, the sleep-dependent consolidation of sentence-level combinatorics has not been studied to date. Here, we propose that dissociable aspects of sleep neurophysiology consolidate two different types of combinatory mechanisms in human language: sequence-based (order-sensitive) and dependency-based (order-insensitive) combinatorics. The distinction between the two types of combinatorics is motivated both by cross-linguistic considerations and the neurobiological underpinnings of human language. Unifying this perspective with principles of sleep-dependent memory consolidation, we posit that a function of sleep is to optimize the consolidation of sequence-based knowledge (the when) and the establishment of semantic schemas of unordered items (the what) that underpin cross-linguistic variations in sentence comprehension. This hypothesis builds on the proposal that sleep is involved in the construction of predictive codes, a unified principle of brain function that supports incremental sentence comprehension. Finally, we discuss neurophysiological measures (EEG/MEG) that could be used to test these claims, such as the quantification of neuronal oscillations, which reflect basic mechanisms of information processing in the brain.