On the speech envelope in the cortical tracking of speech.

The synchronization between the speech envelope and neural activity in auditory regions, referred to as cortical tracking of speech (CTS), plays a key role in speech processing. The method selected for extracting the envelope is a crucial step in CTS measurement, and the absence of a consensus on best practices among the various methods can influence analysis outcomes and interpretation. Here, we systematically compare five standard envelope extraction methods the absolute value of Hilbert transform (absHilbert), gammatone filterbanks, heuristic approach, Bark scale, and vocalic energy), analyzing their impact on the CTS. We present performance metrics for each method based on the recording of brain activity from participants listening to speech in clear and noisy conditions, utilizing intracranial EEG, MEG and EEG data. As expected, we observed significant CTS in temporal brain regions below 10 Hz across all datasets, regardless of the extraction methods. In general, the gammatone filterbanks approach consistently demonstrated superior performance compared to other methods. Results from our study can guide scientists in the field to make informed decisions about the optimal analysis to extract the CTS, contributing to advancing the understanding of the neuronal mechanisms implicated in CTS.

Integrating when and what information in the left parietal lobe allows language rule generalization.

A crucial aspect when learning a language is discovering the rules that govern how words are combined in order to convey meanings. Because rules are characterized by sequential co-occurrences between elements (e.g., "These cupcakes are unbelievable"), tracking the statistical relationships between these elements is fundamental. However, purely bottom-up statistical learning alone cannot fully account for the ability to create abstract rule representations that can be generalized, a paramount requirement of linguistic rules. Here, we provide evidence that, after the statistical relations between words have been extracted, the engagement of goal-directed attention is key to enable rule generalization. Incidental learning performance during a rule-learning task on an artificial language revealed a progressive shift from statistical learning to goal-directed attention. In addition, and consistent with the recruitment of attention, functional MRI (fMRI) analyses of late learning stages showed left parietal activity within a broad bilateral dorsal frontoparietal network. Critically, repetitive transcranial magnetic stimulation (rTMS) on participants' peak of activation within the left parietal cortex impaired their ability to generalize learned rules to a structurally analogous new language. No stimulation or rTMS on a nonrelevant brain region did not have the same interfering effect on generalization. Performance on an additional attentional task showed that this rTMS on the parietal site hindered participants' ability to integrate "what" (stimulus identity) and "when" (stimulus timing) information about an expected target. The present findings suggest that learning rules from speech is a two-stage process: following statistical learning, goal-directed attention-involving left parietal regions-integrates "what" and "when" stimulus information to facilitate rapid rule generalization.

Conflict monitoring and attentional adjustment during binocular rivalry.

To make sense of ambiguous and, at times, fragmentary sensory input, the brain must rely on a process of active interpretation. At any given moment, only one of several possible perceptual representations prevails in our conscious experience. Our hypothesis is that the competition between alternative representations induces a pattern of neural activation resembling cognitive conflict, eventually leading to fluctuations between different perceptual outcomes in the case of steep competition. To test this hypothesis, we probed changes in perceptual awareness between competing images using binocular rivalry. We drew our predictions from the conflict monitoring theory, which holds that cognitive control is invoked by the detection of conflict during information processing. Our results show that fronto-medial theta oscillations (5-7 Hz), an established electroencephalography (EEG) marker of conflict, increases right before perceptual alternations and decreases thereafter, suggesting that conflict monitoring occurs during perceptual competition. Furthermore, to investigate conflict resolution via attentional engagement, we looked for a neural marker of perceptual switches as by parieto-occipital alpha oscillations (8-12 Hz). The power of parieto-occipital alpha displayed an inverse pattern to that of fronto-medial theta, reflecting periods of high interocular inhibition during stable perception, and low inhibition around moments of perceptual change. Our findings aim to elucidate the relationship between conflict monitoring mechanisms and perceptual awareness.

Can the occipital alpha-phase speed up visual detection through a real-time EEG-based brain-computer interface (BCI)?

Electrical brain oscillations reflect fluctuations in neural excitability. Fluctuations in the alpha band (α, 8-12 Hz) in the occipito-parietal cortex are thought to regulate sensory responses, leading to cyclic variations in visual perception. Inspired by this theory, some past and recent studies have addressed the relationship between α-phase from extra-cranial EEG and behavioural responses to visual stimuli in humans. The latest studies have used offline approaches to confirm α-gated cyclic patterns. However, a particularly relevant implication is the possibility to use this principle online, whereby stimuli are time-locked to specific α-phases leading to predictable outcomes in performance. Here, we aimed at providing a proof of concept for such real-time neurotechnology. Participants performed a speeded response task to visual targets that were presented upon a real-time estimation of the α-phase via an EEG closed-loop brain-computer interface (BCI). According to the theory, we predicted a modulation of reaction times (RTs) along the α-cycle. Our BCI system achieved reliable trial-to-trial phase locking of stimuli to the phase of individual occipito-parietal α-oscillations. Yet, the behavioural results did not support a consistent relation between RTs and the phase of the α-cycle neither at group nor at single participant levels. We must conclude that although the α-phase might play a role in perceptual decisions from a theoretical perspective, its impact on EEG-based BCI application appears negligible.

The phase of pre-stimulus brain oscillations correlates with cross-modal synchrony perception.

In everyday life multisensory events, such as a glass crashing on the floor, the different sensory inputs are often experienced as simultaneous, despite the sensory processing of sound and sight within the brain are temporally misaligned. This lack of cross-modal synchrony is the unavoidable consequence of different light and sound speeds, and their different neural transmission times in the corresponding sensory pathways. Hence, cross-modal synchrony must be reconstructed during perception. It has been suggested that spontaneous fluctuations in neural excitability might be involved in the temporal organisation of sensory events during perception and account for variability in behavioural performance. Here, we addressed the relationship between ongoing brain oscillations and the perception of cross-modal simultaneity. Participants performed an audio-visual simultaneity judgement task while their EEG was recorded. We focused on pre-stimulu activity, and found that the phase of neural oscillations at 13 ± 2 Hz 200 ms prior to the stimulus correlated with subjective simultaneity of otherwise identical sound-flash events. Remarkably, the correlation between EEG phase and behavioural report occurred in the absence of concomitant changes in EEG amplitude. The probability of simultaneity perception fluctuated significantly as a function of pre-stimulus phase, with the largest perceptual variation being accounted for phase angles nearly 180º apart. This pattern was strongly reliable for sound-flash pairs but not for flash-sound pairs. Overall, these findings suggest that the phase of ongoing brain activity might underlie internal states of the observer that influence cross-modal temporal organisation between the senses and, in turn, subjective synchrony.

Perceptual and Physiological Consequences of Dark Adaptation: A TMS-EEG Study.

Existing literature on sensory deprivation suggests that short-lasting periods of dark adaptation (DA) can cause changes in visual cortex excitability. DA cortical effects have previously been assessed through phosphene perception, i.e., the ability to report visual sensations when a transcranial magnetic stimulation (TMS) pulse is delivered over the visual cortex. However, phosphenes represent an indirect measure of visual cortical excitability which relies on a subjective report. Here, we aimed at overcoming this limitation by assessing visual cortical excitability by combining subjective (i.e., TMS-induced phosphenes) and objective (i.e., TMS-evoked potentials - TEPs) measurements in a TMS-EEG protocol after 30 min of DA. DA effects were compared to a control condition, entailing 30 min of controlled light exposure. TMS was applied at 11 intensities in order to estimate the psychometric function of phosphene report and explore the relationship between TEPs and TMS intensity. Compared to light adaptation, after DA the slope of the psychometric function was significantly steeper, and the amplitude of a TEP component (P60) was lower, only for high TMS intensities. The perceptual threshold was not affected by DA. These results support the idea that DA leads to a change in the excitability of the visual cortex, accompanied by a behavioral modification of visual perception. Furthermore, this study provides a first valuable description of the relationship between TMS intensity and visual TEPs.

The breakdown of the Simon effect in cross-modal contexts: EEG evidence.

In everyday life, we often must coordinate information across spatial locations and different senses for action. It is well known, for example, that reactions are faster when an imperative stimulus and its required response are congruent than when they are not, even if stimulus location itself is completely irrelevant for the task (the so-called Simon effect). However, because these effects have been frequently investigated in single-modality scenarios, the consequences of spatial congruence when more than one sensory modality is at play are less well known. Interestingly, at a behavioral level, the visual Simon effect vanishes in mixed (visual and tactile) modality scenarios, suggesting that irrelevant spatial information ceases to exert influence on vision. To shed some light on this surprising result, here we address the expression of irrelevant spatial information in EEG markers typical of the visual Simon effect (P300, theta power modulation, LRP) in mixed-modality contexts. Our results show no evidence for the visual-spatial information to affect performance at behavioral and neurophysiological levels. The absence of evidence of the neural markers of visual S-R conflict in the mixed-modality scenario implies that some aspects of spatial representations that are strongly expressed in single-modality scenarios might be bypassed.

Confounders in the detection of minimal hepatic encephalopathy: a neuropsychological and quantified EEG study.

BACKGROUND & AIMS: Chronic alcohol misuse, HCV infection and cirrhosis may cause cognitive alterations. The aim of the present study was to assess the influence of alcohol misuse, HCV infection and cirrhosis per se on the neuropsychological and electroencephalogram (EEG) profile and to evaluate the role of alcohol misuse and HCV infections as potential confounding factors in the detection of minimal hepatic encephalopathy. METHODS: A comprehensive neuropsychological profile and EEG spectral parameters were obtained in six age-matched groups of 30 subjects each: (i) HCV-related hepatitis without cirrhosis, (ii) chronic alcohol abusers, (iii) patients with HCV-related cirrhosis, (iv) alcohol-related cirrhosis, (v) cirrhosis not related to alcohol or HCV and (vi) healthy subjects. Cirrhotic patients were matched for MELD score. RESULTS: The factor 'cirrhosis' was associated with low Phonemic Verbal Fluency (PVF) and Difference between Trail Making Test B and A (TMT) (B-A) (P < 0.001). Chronic alcohol misuse was associated with low PVF, TMT (B-A), Memory with Interference Task at 10 (ITM 10) and 30 s (ITM 30) (all P < 0.05). An interaction was found between the factors 'cirrhosis', 'alcohol misuse' and tests (P < 0.01). HCV hepatitis reduced ITM 10 (P < 0.05), but no interaction was found between 'cirrhosis', 'HCV infection' and tests (P = 0.14). The EEG parameters were mainly influenced by 'cirrhosis' (P < 0.05), and EEG alterations were more pronounced in patients with alcoholic cirrhosis (P = 0.04). CONCLUSIONS: Cirrhosis per se, chronic alcohol misuse and HCV infection were found to be associated with cognitive dysfunction. In patients with cirrhosis, the interaction with alcohol misuse further impinged on brain dysfunction.

Long-Range α-Synchronization as Control Signal for BCI: A Feasibility Study.

Shifts in spatial attention are associated with variations in α band (α, 8-14 Hz) activity, specifically in interhemispheric imbalance. The underlying mechanism is attributed to local α-synchronization, which regulates local inhibition of neural excitability, and frontoparietal synchronization reflecting long-range communication. The direction-specific nature of this neural correlate brings forward its potential as a control signal in brain-computer interfaces (BCIs). In the present study, we explored whether long-range α-synchronization presents lateralized patterns dependent on voluntary attention orienting and whether these neural patterns can be picked up at a single-trial level to provide a control signal for active BCI. We collected electroencephalography (EEG) data from a cohort of healthy adults (n = 10) while performing a covert visuospatial attention (CVSA) task. The data show a lateralized pattern of α-band phase coupling between frontal and parieto-occipital regions after target presentation, replicating previous findings. This pattern, however, was not evident during the cue-to-target orienting interval, the ideal time window for BCI. Furthermore, decoding the direction of attention trial-by-trial from cue-locked synchronization with support vector machines (SVMs) was at chance level. The present findings suggest EEG may not be capable of detecting long-range α-synchronization in attentional orienting on a single-trial basis and, thus, highlight the limitations of this metric as a reliable signal for BCI control.

Developmental changes in individual alpha frequency: Recording EEG data during public engagement events.

Statistical power in cognitive neuroimaging experiments is often very low. Low sample size can reduce the likelihood of detecting real effects (false negatives) and increase the risk of detecting non-existing effects by chance (false positives). Here, we document our experience of leveraging a relatively unexplored method of collecting a large sample size for simple electroencephalography (EEG) studies: by recording EEG in the community during public engagement and outreach events. We collected data from 346 participants (189 females, age range 6-76 years) over 6 days, totalling 29 hours, at local science festivals. Alpha activity (6-15 Hz) was filtered from 30 seconds of signal, recorded from a single electrode placed between the occipital midline (Oz) and inion (Iz) while the participants rested with their eyes closed. A total of 289 good-quality datasets were obtained. Using this community-based approach, we were able to replicate controlled, lab-based findings: individual alpha frequency (IAF) increased during childhood, reaching a peak frequency of 10.28 Hz at 28.1 years old, and slowed again in middle and older age. Total alpha power decreased linearly, but the aperiodic-adjusted alpha power did not change over the lifespan. Aperiodic slopes and intercepts were highest in the youngest participants. There were no associations between these EEG indexes and self-reported fatigue, measured by the Multidimensional Fatigue Inventory. Finally, we present a set of important considerations for researchers who wish to collect EEG data within public engagement and outreach environments.

The effect of TMS on visual motion sensitivity: an increase in neural noise or a decrease in signal strength?

The underlying mechanisms of action of transcranial magnetic stimulation (TMS) are still a matter of debate. TMS may impair a subject's performance by increasing neural noise, suppressing the neural signal, or both. Here, we delivered a single pulse of TMS (spTMS) to V5/MT during a motion direction discrimination task while concurrently manipulating the level of noise in the motion stimulus. Our results indicate that spTMS essentially acts by suppressing the strength of the relevant visual signal. We suggest that TMS may induce a pattern of neural activity that complements the ongoing activation elicited by the sensory signal in a manner that partially impoverishes that signal.

The phase of Theta oscillations modulates successful memory formation at encoding.

Several studies have shown that attention and perception can depend upon the phase of ongoing neural oscillations at stimulus onset. Here, we extend this idea to the memory domain. We tested the hypothesis that ongoing fluctuations in neural activity impact memory encoding in two experiments using a picture paired-associates task in order to gauge episodic memory performance. Experiment 1 was behavioural only and capitalized on the principle of phase resetting. We tested if subsequent memory performance fluctuates rhythmically, time-locked to a resetting cue presented before the to-be-remembered pairs at different time intervals. We found an indication that behavioural performance was periodically and selectively modulated at Theta frequency (~4 Hz). In Experiment 2, we focused on pre-stimulus ongoing activity using scalp EEG while participants performed a paired-associates task. The pre-registered analysis, using large electrode clusters and generic Theta and Alpha spectral ranges, returned null results of the pre-stimulus phase-behaviour correlation. However, as expected from prior literature, we found that variations in stimulus-related Theta-power predicted subsequent memory performance. Therefore, we used this post-stimulus effect in Theta power to guide a post-hoc pre-stimulus phase analysis in terms of scalp and frequency of interest. This analysis returned a correlation between the pre-stimulus Theta phase and subsequent memory. Altogether, these results suggest that pre-stimulus Theta activity at encoding may impact later memory performance.

#EEGManyLabs: Investigating the replicability of influential EEG experiments.

There is growing awareness across the neuroscience community that the replicability of findings about the relationship between brain activity and cognitive phenomena can be improved by conducting studies with high statistical power that adhere to well-defined and standardised analysis pipelines. Inspired by recent efforts from the psychological sciences, and with the desire to examine some of the foundational findings using electroencephalography (EEG), we have launched #EEGManyLabs, a large-scale international collaborative replication effort. Since its discovery in the early 20th century, EEG has had a profound influence on our understanding of human cognition, but there is limited evidence on the replicability of some of the most highly cited discoveries. After a systematic search and selection process, we have identified 27 of the most influential and continually cited studies in the field. We plan to directly test the replicability of key findings from 20 of these studies in teams of at least three independent laboratories. The design and protocol of each replication effort will be submitted as a Registered Report and peer-reviewed prior to data collection. Prediction markets, open to all EEG researchers, will be used as a forecasting tool to examine which findings the community expects to replicate. This project will update our confidence in some of the most influential EEG findings and generate a large open access database that can be used to inform future research practices. Finally, through this international effort, we hope to create a cultural shift towards inclusive, high-powered multi-laboratory collaborations.

The neural mechanisms of the effects of transcranial magnetic stimulation on perception.

Transcranial magnetic stimulation (TMS) is a technique used to study perceptual, motor, and cognitive functions in the human brain. Its effects have been likened to a "virtual brain lesion," but a direct test of this assumption is lacking. To verify this hypothesis, we measured psychophysically the interaction between the neural activity induced by a visual motion-direction discrimination task and that induced by TMS. The visual stimulus featured two elements: a visual signal (dots that moved coherently in one direction) and visual noise (dots that moved randomly in many directions). Three hypotheses were tested to explain the impairment in performance as a result of TMS: 1) a decrease in signal strength; 2) an induction of randomly distributed neural noise with an accompanying decrement in system sensitivity; and 3) a suppression of relevant information processing and addition of neural noise. We provide evidence in favor of the second hypothesis by showing that TMS basically acts by adding neural noise to the perceptual process.

From cognitive control to visual incongruity: Conflict detection in surrealistic images.

This study explored brain responses to images that exploit incongruity as a creative technique, often used in advertising (i.e., surrealistic images). We hypothesized that these images would reveal responses akin to cognitive conflict resulting from incongruent trials in typical laboratory tasks (i.e., Stroop Task). Indeed, in many surrealistic images, common visual elements are juxtaposed to create un-ordinary associations with semantically conflicting representations. We expected that these images engage the conflict processing network that has been described in cognitive neuroscience theories. We addressed this hypothesis by measuring the power of mid-frontal Theta oscillations using EEG while participants watched images through a social media-like interface. Incongruent images, compared to controls, produced a significant Theta power increase, as predicted from the cognitive conflict theory. We also found increased memory for incongruent images one week after exposure, compared to the controls. These findings provide evidence for the incongruent images to effectively engage the viewer's cognitive control and boost memorability. The results of this study provide validation of cognitive theories in real-life scenarios (i.e., surrealistic ads or art) and offer insights regarding the use of neural correlates as effectiveness metrics in advertising.

The relevance of alpha phase in human perception.

Neural oscillations in the low frequencies, roughly in the alpha band (α, 5-15 Hz), have been suggested to act as a gateway from sensation to perception. This hypothesis assumes discrete perception. In particular, the amplitude and the phase of the low frequency rhythm entails a cortical mechanism which paces the access of sensory information into the cognitive system. Evidence supporting this idea includes correlations between the phase of neural oscillations and behavioral performance in perception, spatial attention and working memory. Despite a widespread confidence in the theory, these findings have been mostly based on a varied range of exploratory approaches and inferential group statistics. Here, we aimed at validating the involvement of low frequency cortical rhythm in perception and at providing a clear-cut EEG analysis pipeline. Such an analytical pipeline should support the adoption of a hypothesis-driven framework for future replications and applications. The design, the analyses and the statistical power of the present experiment were based on prior studies in which phase opposition was successfully found. However, our results provide evidence for the involvement of pre-stimulus oscillatory alpha amplitude but not phase in perception. We discuss the null findings from the present study within the existing literature.

Modality-switching in the Simon task: The clash of reference frames.

The representation of spatial information related to an event can influence behavior even when location is task-irrelevant, as in the case of Stimulus-Response (S-R) compatibility effects on the Simon task. However, unlike single-modality situations, which are often used to study the Simon effect, in real-life scenarios various sensory modalities provide spatial information coded in different coordinate systems. Here, we address the expression of S-R compatibility effects in mixed-modality contexts, where events can occur in 1 of various sensory modalities (i.e., vision, touch or audition). The results confirm that, in single-modality cases, Simon effects in vision are expressed in an external spatial frame of reference, while touch information is coded anatomically. Remarkably, when mixing visual and tactile trials in an unpredictable way, the Simon effect disappeared in vision whereas tactile Simon effects remained expressed in their own (anatomical) frame of reference. Mixing visual and auditory stimuli did not obliterate the visual Simon effect and S-R compatibility effects in an external reference frame were evident for both modalities. The extinction of visual Simon effects as a result of mixing visual and tactile modalities can be interpreted as a consequence of the dynamic reorganization of the weights associated to the different sources of spatial information at play. (PsycINFO Database Record

Reliability of TMS phosphene threshold estimation: Toward a standardized protocol.

BACKGROUND: Phosphenes induced by transcranial magnetic stimulation (TMS) are a subjectively described visual phenomenon employed in basic and clinical research as index of the excitability of retinotopically organized areas in the brain. OBJECTIVE: Phosphene threshold estimation is a preliminary step in many TMS experiments in visual cognition for setting the appropriate level of TMS doses; however, the lack of a direct comparison of the available methods for phosphene threshold estimation leaves unsolved the reliability of those methods in setting TMS doses. The present work aims at fulfilling this gap. METHODS: We compared the most common methods for phosphene threshold calculation, namely the Method of Constant Stimuli (MOCS), the Modified Binary Search (MOBS) and the Rapid Estimation of Phosphene Threshold (REPT). In two experiments we tested the reliability of PT estimation under each of the three methods, considering the day of administration, participants' expertise in phosphene perception and the sensitivity of each method to the initial values used for the threshold calculation. RESULTS: We found that MOCS and REPT have comparable reliability when estimating phosphene thresholds, while MOBS estimations appear less stable. CONCLUSIONS: Based on our results, researchers and clinicians can estimate phosphene threshold according to MOCS or REPT equally reliably, depending on their specific investigation goals. We suggest several important factors for consideration when calculating phosphene thresholds and describe strategies to adopt in experimental procedures.

The mismatch negativity as an index of cognitive decline for the early detection of Alzheimer's disease.

Evidence suggests that Alzheimer's disease (AD) is part of a continuum, characterized by long preclinical phases before the onset of clinical symptoms. In several cases, this continuum starts with a syndrome, defined as mild cognitive impairment (MCI), in which daily activities are preserved despite the presence of cognitive decline. The possibility of having a reliable and sensitive neurophysiological marker that can be used for early detection of AD is extremely valuable because of the incidence of this type of dementia. In this study, we aimed to investigate the reliability of auditory mismatch negativity (aMMN) as a marker of cognitive decline from normal ageing progressing from MCI to AD. We compared aMMN elicited in the frontal and temporal locations by duration deviant sounds in short (400 ms) and long (4000 ms) inter-trial intervals (ITI) in three groups. We found that at a short ITI, MCI showed only the temporal component of aMMN and AD the frontal component compared to healthy elderly who presented both. At a longer ITI, aMMN was elicited only in normal ageing subjects at the temporal locations. Our study provides empirical evidence for the possibility to adopt aMMN as an index for assessing cognitive decline in pathological ageing.