State- or trait-like individual differences in dream recall. Preliminary findings from a within-subjects study of multiple nap REM sleep awakenings

We examined the question whether the role of EEG oscillations in predicting presence/absence of dream recall (DR) is explained by "state-" or "trait-like" factors. Six healthy subjects were awakened from REM sleep in a within-subjects design with multiple naps, until a recall and a non-recall condition were obtained. Naps were scheduled in the early afternoon and were separated by 1 week. Topographical EEG data of the 5-min of REM sleep preceding each awakening were analyzed by power spectral analysis [Fast Fourier Transform (FFT)] and by a method to detect oscillatory activity [Better OSCillations (BOSC)]. Both analyses show that REC is associated to higher frontal theta activity (5-7 Hz) and theta oscillations (6.06 Hz) compared to NREC condition, but only the second comparison reached significance. Our pilot study provides support to the notion that sleep and wakefulness share similar EEG correlates of encoding in episodic memories, and supports the "state-like hypothesis": DR may depend on the physiological state related to the sleep stage from which the subject is awakened rather than on a stable individual EEG pattern

Oscillatory EEG activity during REM sleep in elderly people predicts subsequent dream recall after awakenings

Several findings underlined that the electrophysiological (EEG) background of the last segment of sleep before awakenings may predict the presence/absence of dream recall (DR) in young subjects. However, little is known about the EEG correlates of DR in elderly people. Only an investigation found differences between recall and non-recall conditions during NREM sleep EEG in older adults, while—surprisingly—no EEG predictor of DR was found for what concerns REMsleep. Considering REMsleep as a privileged scenario to produce mental sleep activity related to cognitive processes, our study aimed to investigate whether specific EEG topography and frequency changes during REM sleep in elderly people may predict a subsequent recall of mental sleep activity. Twenty-one healthy older volunteers (mean age 69.2 ± 6.07 SD) and 20 young adults (mean age 23.4 ± 2.76 SD) were recorded for one night from19 scalp derivations. Dreams were collected upon morning awakenings from REM sleep. EEG signals of the last 5min were analyzed by the Better OSCillation algorithm to detect the peaks of oscillatory activity in both groups. Statistical comparisons revealed that older as well as young individuals recall their dream experience when the last segment of REM sleep is characterized by frontal theta oscillations. No Recall (Recall vs. Non-Recall) × Age (Young vs. Older) interaction was found. This result replicated the previous evidence in healthy young subjects, as shown in within- and between-subjects design. The findings are completely original for older individuals, demonstrating that theta oscillations are crucial for the retrieval of dreaming also in this population. Furthermore, our results did not confirm a greater presence of the theta activity in healthy aging. Conversely, we found a greater amount of rhythmic theta and alpha activity in young than older participants. It is worth noting that the theta oscillations detected are related to cognitive functioning. We emphasize the notion that the oscillatory theta activity should be distinguished from the non-rhythmic theta activity identified in relation to other phenomena such as (a) sleepiness and hypoarousal conditions during the waking state and (b) cortical slowing, considered as an EEG alteration in clinical samples

The benefits of targeted memory reactivation for consolidation in sleep are contingent on memory accuracy and direct cue-memory associations

Objectives: To investigate how the effects of targeted memory reactivation (TMR) are influenced by memory accuracy prior to sleep and the presence or absence of direct cue-memory associations. Methods: 30 participants associated each of 50 pictures with an unrelated word and then with a screen location in two separate tasks. During picture-location training, each picture was also presented with a semantically related sound. The sounds were therefore directly associated with the picture locations but indirectly associated with the words. During a subsequent nap, half of the sounds were replayed in slow wave sleep (SWS) (TMR). The effect of TMR on memory for the picture locations (direct cue-memory associations) and picture-word pairs (indirect cue-memory associations) was then examined. Results: TMR reduced overall memory decay for recall of picture locations. Further analyses revealed a benefit of TMR for picture locations recalled with a low degree of accuracy prior to sleep, but not those recalled with a high degree of accuracy. The benefit of TMR for low accuracy memories was predicted by time spent in SWS. There was no benefit of TMR for memory of the picture-word pairs, irrespective of memory accuracy prior to sleep. Conclusions: TMR provides the greatest benefit to memories recalled with a low degree of accuracy prior to sleep. The memory benefits of TMR may also be contingent on direct cue-memory associations

Cross-linguistic differences in case marking shape neural power dynamics and gaze behavior during sentence planning

Languages differ in how they mark the dependencies between verbs and arguments, e.g., by case. An eye tracking and EEG picture description study examined the influence of case marking on the time course of sentence planning in Basque and Swiss German. While German assigns an unmarked (nominative) case to subjects, Basque specifically marks agent arguments through ergative case. Fixations to agents and event-related synchronization (ERS) in the theta and alpha frequency bands, as well as desynchronization (ERD) in the alpha and beta bands revealed multiple effects of case marking on the time course of early sentence planning. Speakers decided on case marking under planning early when preparing sentences with ergative-marked agents in Basque, whereas sentences with unmarked agents allowed delaying structural commitment across languages. These findings support hierarchically incremental accounts of sentence planning and highlight how cross-linguistic differences shape the neural dynamics underpinning language use.This work was funded by Swiss National Science Foundation Grant Nr. 100015_160011 (B.B. and M.M.), the NCCR Evolving Language, Swiss National Science Foundation Agreement Nr. #51NF40_180888 (B.B. and M. M.), and the PhD Program in Linguistics and the Graduate Research Campus of the University of Zurich (A.E.). DEB is supported by a grant from the Harvard Data Science Initiative and the Branco Weiss Foundation. I.B.-S. is supported by an Australian Research Council Future Fellowship (FT160100437). I.L. is supported by grants from the Spanish Ministry of Economy and Competitiveness (Grant No. FFI2015-64183-P) and the Basque Government (IT1169-19). The authors thank Anne-Lise Giraud for the suggestion to include beta-band analyses, Vitória Piai for advice on EEG data processing, Giuachin Kreiliger for statistical consultation, Andrina Balsofiore and Edurne Petrirena for help recording the lead-in fragments, Nathalie Rieser and Debora Beuret for help with data collection and processing, and the Phonogram Archives of the University of Zurich for technical support. The authors also thank two anonymous reviewers for their helpful comments on an earlier version of the manuscript

Interactions between the hippocampus and prefrontal cortex in context-dependent overlapping memory retrieval

Activation in the hippocampus (HC) and prefrontal cortex (PFC) is critical to accurately retrieve overlapping sequences. Experiments 1 and 2 tested the hypotheses that activation in and interaction between HC and PFC increases as overlap between sequences increases in a non-spatial task. Experiment 3 tested the hypothesis that theta oscillations are involved in orchestrating interactions between HC and PFC in a spatial task with overlapping elements. In the first two studies, 17 participants (aged 18-34; 11 female) learned sequences consisting of a picture frame, face, and scene. Conditions varied by degree of overlap. Using fMRI, Experiment 1 tested how degree of overlap affected HC and PFC activation. In overlapping sequences, middle and posterior HC were active when predictability of the correct response increased, dorsolateral PFC was active when participants were able to ascertain the correct set of sequences, and ventrolateral PFC was active when inhibition of interfering associations was required. Experiment 2 examined functional connectivity of HC and PFC during disambiguation. Low- and high-overlap conditions were associated with increased connectivity in separate regions at different times indicating that retrieval under the two conditions used different neural networks and strategies. Low-overlap trials were associated with increased connectivity between HC and prefrontal and parietal regions. High-overlap trials showed increased connectivity between lateral PFC and visual areas, indicating that imagery may be necessary for accurate performance. Using EEG recording, Experiment 3 examined theta activity during retrieval of well-learned, overlapping and non-overlapping mazes in 17 participants (aged 18-34, 11 female). Theta activity increased in overlapping mazes during the first of four hallways, suggesting participants were looking ahead to upcoming turns in the maze. Theta activity increased at the beginning and choice point of the third overlapping hallway, possibly in response to interference from the paired, overlapping maze. These studies provide evidence that (1) overlapping associations in non-spatial sequences elicit interactions between hippocampus and lateral prefrontal cortex, (2) increasing the degree of overlap changes the neural processes required to perform the task, and (3) theta power increases in response to increased cognitive demand and maintenance of sequence information needed to differentiate between overlapping spatial routes

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 (thewhen) and the establishment of semantic schemas of unordered items (thewhat) 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

Neural oscillations and a nascent corticohippocampal theory of reference

The ability to use words to refer to the world is vital to the communicative power of human language. In particular, the anaphoric use of words to refer to previously mentioned concepts (antecedents) allows dialogue to be coherent and meaningful. Psycholinguistic theory posits that anaphor comprehension involves reactivating a memory representation of the antecedent. Whereas this implies the involvement of recognition memory, or the mnemonic sub-routines by which people distinguish old from new, the neural processes for reference resolution are largely unknown. Here, we report time-frequency analysis of four EEG experiments to reveal the increased coupling of functional neural systems associated with referentially coherent expressions compared to referentially problematic expressions. Despite varying in modality, language, and type of referential expression, all experiments showed larger gamma-band power for referentially coherent expressions compared to referentially problematic expressions. Beamformer analysis in high-density Experiment 4 localised the gamma-band increase to posterior parietal cortex around 400-600 ms after anaphor-onset and to frontaltemporal cortex around 500-1000 ms. We argue that the observed gamma-band power increases reflect successful referential binding and resolution, which links incoming information to antecedents through an interaction between the brain’s recognition memory networks and frontal-temporal language network. We integrate these findings with previous results from patient and neuroimaging studies, and we outline a nascent cortico-hippocampal theory of reference

The spatiotemporal pattern of the human electroencephalogram at sleep onset after a period of prolonged wakefulness

During the sleep onset (SO) process, the human electroencephalogram (EEG) is characterized by an orchestrated pattern of spatiotemporal changes. Sleep deprivation (SD) strongly affects both wake and sleep EEG, but a description of the topographical EEG power spectra and oscillatory activity during the wake-sleep transition after a period of prolonged wakefulness is still missing. The increased homeostatic sleep pressure should induce an earlier onset of sleep-related EEG oscillations. The aim of the present study was to assess the spatiotemporal EEG pattern at SO following SD. A dataset of a previous study was analyzed. We assessed the spatiotemporal EEG changes (19 cortical derivations) during the SO (5 min before vs. 5 min after the first epoch of Stage 2) of a recovery night after 40 h of SD in 39 healthy subjects, analyzing the EEG power spectra (fast Fourier transform) and the oscillatory activity [better oscillation (BOSC) detection method]. The spatiotemporal pattern of the EEG power spectra mostly confirmed the changes previously observed during the wake-sleep transition at baseline. The comparison between baseline and recovery showed a wide increase of the post- vs. pre-SO ratio during the recovery night in the frequency bins 10 Hz. We found a predominant alpha oscillatory rhythm in the pre-SO period, while after SO the theta oscillatory activity was prevalent. The oscillatory peaks showed a generalized increase in all frequency bands from delta to sigma with different predominance, while beta activity increased only in the fronto-central midline derivations. Overall, the analysis of the EEG power replicated the topographical pattern observed during a baseline night of sleep but with a stronger intensity of the SO-induced changes in the frequencies 10 Hz, and the detection of the rhythmic activity showed the rise of several oscillations at SO after SD that was not observed during the wake-sleep transition at baseline (e.g., alpha and frontal theta in correspondence of their frequency peaks). Beyond confirming the local nature of the EEG pattern at SO, our results show that SD has an impact on the spatiotemporal modulation of cortical activity during the falling-asleep process, inducing the earlier emergence of sleep-related EEG oscillations