Side effects of monaural beat stimulation during sustained mental work on mind wandering and performance measures.

Excessive mind wandering (MW) contributes to the development and maintenance of psychiatric disorders. Previous studies have suggested that auditory beat stimulation may represent a method enabling a reduction of MW. However, little is known about how different auditory stimulation conditions are subjectively perceived and whether this perception is in turn related to changes in subjective states, behavioral measures of attention and MW. In the present study, we therefore investigated MW under auditory beat stimulation and control conditions using experience sampling during a sustained attention to response task (SART). The subjective perception of the stimulation conditions, as well as changes in anxiety, stress and negative mood after versus before stimulation were assessed via visual-analog scales. Results showed that any auditory stimulation applied during the SART was perceived as more distracting, disturbing, uncomfortable and tiring than silence and was related to more pronounced increases of stress and negative mood. Importantly, the perception of the auditory conditions as disturbing was directly correlated with MW propensity. Additionally, distracting, disturbing and uncomfortable perceptions predicted negative mood. In turn, negative mood was inversely correlated with response accuracy for target stimuli, a behavioral indicator of MW. In summary, our data show that MW and attentional performance are affected by the adverse perception of auditory stimulation, and that this influence may be mediated by changes in mood.

Insights into the time course of mind wandering during task execution.

Our minds tend to wander, sometimes with little control. Despite this phenomenon, that can affect our ability to perform everyday tasks gaining much interest, relatively little is understood about the actual time course of MW across an experimental task. With this in mind, we collated data from two previously reported studies investigating the effect of auditory beat stimulation on MW. Taking experience sampling probes intermittently dispersed throughout a sustained-attention-to-response task (SART), we re-evaluated responses to theta monaural beat stimulation, as well as to two control conditions (silence (headphones only) and a sine wave control tone). The experience sampling probes were binned into shorter intervals of approximately five minutes duration, chronologically as they appeared within the paradigm. Experience sampling probes assayed whether MW had occurred, with or without meta-awareness, and lastly in which temporal orientation (past/present/future). By applying this somewhat temporally better resolved approach, we were able to examine the time course of attentional fluctuations related to MW during the execution of the SART, as well as interactions arising from the auditory beat stimulation. As anticipated, MW increased during task execution, most prominently at the beginning of the experiment. We also observed that levels of meta-awareness declined over time. Moreover, the temporal evolution of meta-awareness and past-orientation appeared to depend on the stimulation condition. These data demonstrate that time-on-task is a crucial factor in measuring MW, during the performance of an attentional task.

Mind wandering in anxiety disorders: A status report.

Many investigations have targeted the subject of worry in anxiety disorders. Worry can be regarded as a subtype of mind wandering (MW), which is undeliberate, perseverative, negatively-valenced, and mainly future-oriented. Nevertheless, until now only a few studies have explored the role of overall MW in the origin and course of anxiety disorders. To foster progress in this field, we briefly describe and discuss relevant studies addressing MW in subjects with anxiety disorders or symptoms or disorders associated with anxiety symptoms. Provisional synthesis suggests that: a) the overall amount of MW is positively correlated with anxiety symptoms; b) MW characteristics reflecting worry and rumination appear to be relevant in anxiety; c) comorbid depressive and ADHD symptoms may contribute to excessive MW in anxiety; d) MW-related therapeutic interventions may be useful as complementary treatments in anxiety disorders. However, more studies related to MW in anxiety disorders or symptoms are necessary to corroborate and extend these initial findings. Such investigations should ideally combine experience sampling with self-rating assessments of both MW and worry/rumination.

A key role of the hippocampal P3 in the attentional blink.

The attentional blink (AB) refers to an impaired identification of target stimuli (T2), which are presented shortly after a prior target (T1) within a rapid serial visual presentation (RSVP) stream. It has been suggested that the AB is related to a failed transfer of T2 into working memory and that hippocampus (HC) and entorhinal cortex (EC) are regions crucial for this transfer. Since the event-related P3 component has been linked to inhibitory processes, we hypothesized that the hippocampal P3 elicited by T1 may impact on T2 processing within HC and EC. To test this hypothesis, we reanalyzed microwire data from 21 patients, who performed an RSVP task, during intracranial recordings for epilepsy surgery assessment (Reber et al., 2017). We identified T1-related hippocampal P3 components in the local field potentials (LFPs) and determined the temporal onset of T2 processing in HC/EC based on single-unit response onset activity. In accordance with our hypothesis, T1-related single-trial P3 amplitudes at the onset of T2 processing were clearly larger for unseen compared to seen T2-stimuli. Moreover, increased T1-related single-trial P3 peak latencies were found for T2[unseen] versus T2[seen] trials in case of lags 1 to 3, which was in line with our predictions. In conclusion, our findings support inhibition models of the AB and indicate that the hippocampal P3 elicited by T1 plays a central role in the AB.

What is mind blanking: A conceptual clarification.

The investigation of so-called mind blanking has been recently established in the context of mind wandering research. Unfortunately, the description of the term 'mind blanking' and its experimental assessment are often ambiguous. More specifically, two distinct phenomena have been conceptually blended and both characterized using the term 'mind blanking' in experimental studies: the absence of task-focus and thought, versus complete lack of conscious experiences. At least in part, this confusion can be traced back to the writings of William James, who referred to 'thought' as a superordinate term to address different conscious experiences. Applying the technique of so-called experience sampling, experimental studies, up to now, probably assessed the absence of thought, but not a complete lack of conscious experiences. There is no clear evidence yet for instances of the latter phenomenon occurring during non-pathological waking state. Possibly, such evidence could be revealed in the future using a conceptually more refined experience sampling.

Modulation of Mind Wandering Using Monaural Beat Stimulation in Subjects With High Trait-Level Mind Wandering.

Mind wandering (MW) refers to a state when attention shifts from the task at hand or current situation toward thoughts, feelings, and imaginations. This state is often accompanied by a decline in mood, and patients suffering from major depression exhibit more perseverative MW. Hence, although the directionality of the relationship between mood and MW is still under investigation, it may be useful to explore possible avenues to reduce MW. In an earlier pilot study, we investigated MW during auditory beat stimulation in healthy subjects using thought-probes during a sustained attention to response task (SART). We found evidence for reduced MW during monaural 5 Hz beats compared to silence, sine tones, and binaural 5 Hz beats. Moreover, the data tentatively suggested that this reduction was particularly pronounced in subjects with high levels of MW during silence. In the current study, we therefore asked whether MW can be reduced by monaural theta beats in subjects with high trait-levels of MW, as indicated by an online MW questionnaire. Preselected subjects performed a SART task with thought-probes assessing the propensity to mind wander, meta-awareness, and the temporal orientation of MW. Stimulation conditions comprised monaural theta beats, as well as silence (headphones on), and sine tones as control conditions. Our main hypothesis stating that the propensity to mind wander during monaural theta beats is reduced compared to both control conditions was only partly confirmed. Indeed, MW was significantly diminished during exposure to the theta beats compared to sine tones. However, reduced MW during theta beats versus silence was only observed in a subgroup using stricter inclusion criteria. Considering possible reasons for this outcome, our data suggest that the preselection procedure was suboptimal and that beat effects are modulated by the individual responses to auditory stimulation in general.

On the relationship between mind wandering and mindfulness.

Mind wandering (MW) and mindfulness have both been reported to be vital moderators of psychological wellbeing. Here, we aim to examine how closely associated these phenomena are and evaluate the psychometrics of measures often used to quantify them. We investigated two samples, one consisting of German-speaking unpaid participants (GUP, n [Formula: see text] 313) and one of English-speaking paid participants (EPP, n [Formula: see text] 228) recruited through MTurk.com. In an online experiment, we collected data using the Mindful Attention Awareness Scale (MAAS) and the sustained attention to response task (SART) during which self-reports of MW and meta-awareness of MW were recorded using experience sampling (ES) probes. Internal consistency of the MAAS was high (Cronbachs [Formula: see text] of 0.96 in EPP and 0.88 in GUP). Split-half reliability for SART measures and self-reported MW was overall good with the exception of SART measures focusing on Nogo trials, and those restricted to SART trials preceding ES in a 10 s time window. We found a moderate negative association between trait mindfulness and MW as measured with ES probes in GUP, but not in EPP. Our results suggest that MW and mindfulness are on opposite sides of a spectrum of how attention is focused on the present moment and the task at hand.

Neuropsychological features of mind wandering in left-, right- and extra temporal lobe epilepsy.

PURPOSE: Mind wandering, i.e. mental time-travelling and imagery unrelated to the current situation has recently been related to mesial temporal lobe (memory) function. In this regard we evaluated as to whether parameters of mind wandering are related to material specific memory in patients with a left-, right-, or extra- temporal lobe epilepsy. METHODS: In this prospective controlled study we analyzed mind wandering, material specific memory, and executive functions in 29 right-handed patients with right-, left-, or extra-temporal lobe epilepsies. Mind wandering was assessed with a sustained attention to response task containing embedded inquiries on mind wandering. In addition, verbal list learning and memory (VLMT), design list learning (DCS-R), and executive function (EpiTrack) were assessed. RESULTS: In patients with right temporal lobe epilepsy, the propensity to mind wander was positively related to verbal memory performance, while in left temporal lobe epilepsy, the propensity and future related mind wandering were positively correlated to the performance in visual/figural memory. Generally, the propensity of MW was related to executive function as well. CONCLUSION: The results suggest that mind wandering in lateralized temporal lobe epilepsy appears to be non-specifically driven by executive function and specifically by the mode and functionality of the memory system of the non-epileptic hemisphere. Repeated assessments would be required to discern as to how much such patterns depend on lesions versus epileptic dysfunction and whether they change with successful medical or surgical treatment.

Mind wandering and depression: A status report.

While many clinical studies and overviews on the contribution of rumination to depression exist, relatively little information regarding the role of mind wandering (MW) in general is available. Therefore, it remains an open question whether patterns of MW are altered in depression and, if so, how these alterations are related to rumination. Here, we review and discuss studies investigating MW in cohorts, showing either a clinically significant depression or with clinically significant disorders accompanied by depressive symptoms. These studies yield first tentative insights into major issues. However, further investigations are required, specifically studies which: i) compare patients with a primary diagnosis of major depression with healthy and appropriately matched controls, ii) implement measures of both MW and rumination, iii) are based on experience sampling (in combination with other key approaches), iv) compare experience sampling during daily life, resting state and attentional tasks, v) explore possible biases in the assessment of MW, vi) acquire data not only related to the propensity and contents of MW, but also regarding meta-awareness and intentionality.

Neural activity in the human anterior thalamus during natural vision.

In natural vision humans and other primates explore environment by active sensing, using saccadic eye movements to relocate the fovea and sample different bits of information multiple times per second. Saccades induce a phase reset of ongoing neuronal oscillations in primary and higher-order visual cortices and in the medial temporal lobe. As a result, neuron ensembles are shifted to a common state at the time visual input propagates through the system (i.e., just after fixation). The extent of the brain's circuitry that is modulated by saccades is not yet known. Here, we evaluate the possibility that saccadic phase reset impacts the anterior nuclei of the thalamus (ANT). Using recordings in the human thalamus of three surgical patients during natural vision, we found that saccades and visual stimulus onset both modulate neural activity, but with distinct field potential morphologies. Specifically, we found that fixation-locked field potentials had a component that preceded saccade onset. It was followed by an early negativity around 50 ms after fixation onset which is significantly faster than any response to visual stimulus presentation. The timing of these events suggests that the ANT is predictively modulated before the saccadic eye movement. We also found oscillatory phase concentration, peaking at 3-4 Hz, coincident with suppression of Broadband High-frequency Activity (BHA; 80-180 Hz), both locked to fixation onset supporting the idea that neural oscillations in these nuclei are reorganized to a low excitability state right after fixation onset. These findings show that during real-world natural visual exploration neural dynamics in the human ANT is influenced by visual and oculomotor events, which supports the idea that ANT, apart from their contribution to episodic memory, also play a role in natural vision.

Auditory Beat Stimulation Modulates Memory-Related Single-Neuron Activity in the Human Medial Temporal Lobe.

Auditory beats are amplitude-modulated signals (monaural beats) or signals that subjectively cause the perception of an amplitude modulation (binaural beats). We investigated the effects of monaural and binaural 5 Hz beat stimulation on neural activity and memory performance in neurosurgical patients performing an associative recognition task. Previously, we had reported that these beat stimulation conditions modulated memory performance in opposite directions. Here, we analyzed data from a patient subgroup, in which microwires were implanted in the amygdala, hippocampus, entorhinal cortex and parahippocampal cortex. We identified neurons responding with firing rate changes to binaural versus monaural 5 Hz beat stimulation. In these neurons, we correlated the differences in firing rates for binaural versus monaural beats to the memory-related differences for remembered versus forgotten items and associations. In the left hemisphere, we detected statistically significant negative correlations between firing rate differences for binaural versus monaural beats and remembered versus forgotten items/associations. Importantly, such negative correlations were also observed between beat stimulation-related firing rate differences in the pre-stimulus window and memory-related firing rate differences in the post-stimulus windows. In line with concepts of homeostatic plasticity, our findings suggest that beat stimulation is linked to memory performance via shifting baseline firing levels.

Sharp Wave-Ripples in Human Amygdala and Their Coordination with Hippocampus during NREM Sleep.

Cooperative interactions between the amygdala and hippocampus are widely regarded as critical for overnight emotional processing of waking experiences, but direct support from the human brain for such a dialog is absent. Using overnight intracranial recordings in 4 presurgical epilepsy patients (3 female), we discovered ripples within human amygdala during nonrapid eye movement (NREM) sleep, a brain state known to contribute to affective processing. Like hippocampal ripples, amygdala ripples are associated with sharp waves, linked to sleep spindles, and tend to co-occur with their hippocampal counterparts. Moreover, sharp waves and ripples are temporally linked across the 2 brain structures, with amygdala ripples occurring during hippocampal sharp waves and vice versa. Combined with further evidence of interregional sharp-wave and spindle synchronization, these findings offer a potential physiological substrate for the NREM-sleep-dependent consolidation and regulation of emotional experiences.

Analyzing human sleep EEG: A methodological primer with code implementation.

Recent years have witnessed a surge in human sleep electroencephalography (EEG) studies, employing increasingly sophisticated analysis strategies to relate electrophysiological activity to cognition and disease. However, properly calculating and interpreting metrics used in contemporary sleep EEG requires attention to numerous theoretical and practical signal-processing details that are not always obvious. Moreover, the vast number of outcome measures that can be derived from a single dataset inflates the risk of false positives and threatens replicability. We review several methodological issues related to 1) spectral analysis, 2) montage choice, 3) extraction of phase and amplitude information, 4) surrogate construction, and 5) minimizing false positives, illustrating both the impact of methodological choices on downstream results, and the importance of checking processing steps through visualization and simplified examples. By presenting these issues in non-mathematical form, with sleep-specific examples, and with code implementation, this paper aims to instill a deeper appreciation of methodological considerations in novice and non-technical audiences, and thereby help improve the quality of future sleep EEG studies.

Sleep spindles mediate hippocampal-neocortical coupling during long-duration ripples.

Sleep is pivotal for memory consolidation. According to two-stage accounts, memory traces are gradually translocated from hippocampus to neocortex during non-rapid-eye-movement (NREM) sleep. Mechanistically, this information transfer is thought to rely on interactions between thalamocortical spindles and hippocampal ripples. To test this hypothesis, we analyzed intracranial and scalp Electroencephalography sleep recordings from pre-surgical epilepsy patients. We first observed a concurrent spindle power increase in hippocampus (HIPP) and neocortex (NC) time-locked to individual hippocampal ripple events. Coherence analysis confirmed elevated levels of hippocampal-neocortical spindle coupling around ripples, with directionality analyses indicating an influence from NC to HIPP. Importantly, these hippocampal-neocortical dynamics were particularly pronounced during long-duration compared to short-duration ripples. Together, our findings reveal a potential mechanism underlying active consolidation, comprising a neocortical-hippocampal-neocortical reactivation loop initiated by the neocortex. This hippocampal-cortical dialogue is mediated by sleep spindles and is enhanced during long-duration hippocampal ripples.

Reduced past-oriented mind wandering in left compared to right medial temporal lobe epilepsy.

Mind wandering refers to a shift of attention away from a task at hand to task-unrelated thoughts. Several groups have shown increased activation of the left medial temporal lobe (MTL) before and during spontaneous thoughts suggesting that the left MTL may play a crucial role in mind wandering. Due to its relevance for long-term memory, we further hypothesized that the left MTL is particularly involved in mind wandering towards the past. Accordingly, we predicted a reduced propensity to mind wander and less past-oriented mind wandering in patients with left MTL epilepsies. To this end, we experimentally investigated mind wandering in 89 in-patients undergoing diagnostic evaluation of their putative epileptic disorder. Patients performed a sustained attention to response task with embedded experience sampling probes aiming to assess occurrence, meta-awareness and temporal orientation (past/present/future) of mind-wandering episodes. We did not find significant differences in the propensity to mind wander between patient subgroups. However, the left MTL epilepsy subgroup showed significantly reduced past-oriented mind wandering compared to right MTL epilepsies, as well as a trend towards diminished past-oriented mind wandering compared to idiopathic epilepsies. Possibly due to compensatory mechanisms, the right MTL epilepsy subgroup showed significantly increased past-oriented mind wandering compared to extratemporal epilepsies and patients with syncopes. These behavioural findings point to a rejection of the hypothesis that the amount of time engaged in mind wandering crucially depends on the left MTL. However, our data do support the idea that the left MTL is particularly involved in mind wandering towards the past.

Phase-based coordination of hippocampal and neocortical oscillations during human sleep.

During sleep, new memories undergo a gradual transfer from hippocampal (HPC) to neocortical (NC) sites. Precisely timed neural oscillations are thought to mediate this sleep-dependent memory consolidation, but exactly how sleep oscillations instantiate the HPC-NC dialog remains elusive. Employing overnight invasive electroencephalography in ten neurosurgical patients, we identified three broad classes of phase-based communication between HPC and lateral temporal NC. First, we observed interregional phase synchrony for non-rapid eye movement (NREM) spindles, and N2 and rapid eye movement (REM) theta activity. Second, we found asymmetrical N3 cross-frequency phase-amplitude coupling between HPC slow oscillations (SOs) and NC activity spanning the delta to high-gamma/ripple bands, but not in the opposite direction. Lastly, N2 theta and NREM spindle synchrony were themselves modulated by HPC SOs. These forms of interregional communication emphasize the role of HPC SOs in the HPC-NC dialog, and may offer a physiological basis for the sleep-dependent reorganization of mnemonic content.

No evidence for spontaneous cross-frequency phase-phase coupling in the human hippocampus.

Cross-frequency phase-phase coupling (PPC) has been suggested to play a role in cognitive processing and, in particular, in memory consolidation during sleep. Controversial results have been reported regarding the existence of spontaneous phase-phase coupling in the hippocampus. Here, we investigated this phenomenon in intracranial EEG recordings from the human hippocampus acquired during waking state and different sleep stages. We estimated the strength of interactions between different pairs of frequency bands and evaluated the statistical significance of findings using surrogates that build on different null hypotheses. Indications for spontaneous phase-phase coupling were only observed when testing with less rigorous surrogates. When requiring that all four surrogate tests be passed, however, there were no significant indications for phase-phase coupling. In conclusion, we did not detect evidence for spontaneous cross-frequency phase-phase coupling in the human hippocampus.

Heterogeneous profiles of coupled sleep oscillations in human hippocampus.

Cross-frequency coupling of sleep oscillations is thought to mediate memory consolidation. While the hippocampus is deemed central to this process, detailed knowledge of which oscillatory rhythms interact in the sleeping human hippocampus is lacking. Combining intracranial hippocampal and non-invasive electroencephalography from twelve neurosurgical patients, we characterized spectral power and coupling during non-rapid eye movement (NREM) and rapid eye movement (REM) sleep. Hippocampal coupling was extensive, with the majority of channels expressing spectral interactions. NREM consistently showed delta-ripple coupling, but ripples were also modulated by slow oscillations (SOs) and sleep spindles. SO-delta and SO-theta coupling, as well as interactions between delta/theta and spindle/beta frequencies also occurred. During REM, limited interactions between delta/theta and beta frequencies emerged. Moreover, oscillatory organization differed substantially between i) hippocampus and scalp, ii) sites along the anterior-posterior hippocampal axis, and iii) individuals. Overall, these results extend and refine our understanding of hippocampal sleep oscillations.