Alpha fluctuations regulate the accrual of visual information to awareness.

Endogenous brain processes play a paramount role in shaping up perceptual phenomenology. This is illustrated by the alternations experienced by humans (and other animals) when watching perceptually ambiguous, static images. We hypothesised that endogenous alpha fluctuations in the visual cortex pace the accumulation of sensory information leading to perceptual outcomes. Here, we addressed this hypothesis using binocular rivalry combined with visual entrainment and electroencephalography in humans (64 female, 53 male). The results revealed a correlation between the individual frequency of alpha oscillations in the occipital cortex and perceptual alternation rates experienced during binocular rivalry. In subsequent experiments we show that regulating endogenous brain activity via rhythmic entrainment produced corresponding changes in perceptual alternation rate. These changes were observed only in the alpha range but not at lower entrainment frequencies, and were much reduced when using arrhythmic stimulation. Additionally, entraining at frequencies above the alpha range did not result in speeding up perceptual alternation rates. Overall, these findings support the notion that visual information is accumulated via alpha cycles to promote the emergence of conscious perceptual representations. We suggest that models of binocular rivalry incorporating posterior alpha as a pacemaker can provide an important advance in the comprehension of the dynamics of visual awareness.

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.

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.

Flexibility in reaction time analysis: many roads to a false positive?

In the present article, we explore the influence of undisclosed flexibility in the analysis of reaction times (RTs). RTs entail some degrees of freedom of their own, due to their skewed distribution, the potential presence of outliers and the availability of different methods to deal with these issues. Moreover, these degrees of freedom are usually not considered part of the analysis itself, but preprocessing steps that are contingent on data. We analysed the impact of these degrees of freedom on the false-positive rate using simulations over real and simulated data. When several preprocessing methods are used in combination, the false-positive rate can easily rise to 17%. This figure becomes more concerning if we consider that more degrees of freedom are awaiting down the analysis pipeline, potentially making the final false-positive rate much higher.

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.

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.

Theta oscillations reflect conflict processing in the perception of the McGurk illusion.

The McGurk illusion is one of the most famous illustrations of cross-modal integration in human perception. It has been often used as a proxy of audiovisual (AV) integration and to infer the properties of the integration process in natural (congruent) AV conditions. Nonetheless, a blatant difference between McGurk stimuli and natural, congruent, AV speech is the conflict between the auditory and the visual information in the former. Here, we hypothesized that McGurk stimuli (and any AV incongruency) engage brain responses similar to those found in more general cases of perceptual conflict (e.g., Stroop), and propose that the McGurk illusion arises as a result of the resolution of such conflict. We used electroencephalography to measure variations in the power of theta, a well-known marker of the brain response to conflict. The results showed that perception of AV McGurk stimuli, just like AV incongruence in general, induces an increase in activity in the theta band. This response was similar to that evoked by Stroop stimuli, as measured in the same participants. This finding suggests that the McGurk illusion is mediated by general-purpose conflict mechanisms, and calls for caution in generalizing findings obtained using the McGurk illusion, to the general case of multisensory integration.

Audiovisual integration as conflict resolution: The conflict of the McGurk illusion.

There are two main behavioral expressions of multisensory integration (MSI) in speech; the perceptual enhancement produced by the sight of the congruent lip movements of the speaker, and the illusory sound perceived when a speech syllable is dubbed with incongruent lip movements, in the McGurk effect. These two models have been used very often to study MSI. Here, we contend that, unlike congruent audiovisually (AV) speech, the McGurk effect involves brain areas related to conflict detection and resolution. To test this hypothesis, we used fMRI to measure blood oxygen level dependent responses to AV speech syllables. We analyzed brain activity as a function of the nature of the stimuli-McGurk or non-McGurk-and the perceptual outcome regarding MSI-integrated or not integrated response-in a 2 × 2 factorial design. The results showed that, regardless of perceptual outcome, AV mismatch activated general-purpose conflict areas (e.g., anterior cingulate cortex) as well as specific AV speech conflict areas (e.g., inferior frontal gyrus), compared with AV matching stimuli. Moreover, these conflict areas showed stronger activation on trials where the McGurk illusion was perceived compared with non-illusory trials, despite the stimuli where physically identical. We conclude that the AV incongruence in McGurk stimuli triggers the activation of conflict processing areas and that the process of resolving the cross-modal conflict is critical for the McGurk illusion to arise. Hum Brain Mapp 38:5691-5705, 2017. © 2017 Wiley Periodicals, Inc.

Hand gestures as visual prosody: BOLD responses to audio-visual alignment are modulated by the communicative nature of the stimuli.

During public addresses, speakers accompany their discourse with spontaneous hand gestures (beats) that are tightly synchronized with the prosodic contour of the discourse. It has been proposed that speech and beat gestures originate from a common underlying linguistic process whereby both speech prosody and beats serve to emphasize relevant information. We hypothesized that breaking the consistency between beats and prosody by temporal desynchronization, would modulate activity of brain areas sensitive to speech-gesture integration. To this aim, we measured BOLD responses as participants watched a natural discourse where the speaker used beat gestures. In order to identify brain areas specifically involved in processing hand gestures with communicative intention, beat synchrony was evaluated against arbitrary visual cues bearing equivalent rhythmic and spatial properties as the gestures. Our results revealed that left MTG and IFG were specifically sensitive to speech synchronized with beats, compared to the arbitrary vision-speech pairing. Our results suggest that listeners confer beats a function of visual prosody, complementary to the prosodic structure of speech. We conclude that the emphasizing function of beat gestures in speech perception is instantiated through a specialized brain network sensitive to the communicative intent conveyed by a speaker with his/her hands.

Top-down attention regulates the neural expression of audiovisual integration.

The interplay between attention and multisensory integration has proven to be a difficult question to tackle. There are almost as many studies showing that multisensory integration occurs independently from the focus of attention as studies implying that attention has a profound effect on integration. Addressing the neural expression of multisensory integration for attended vs. unattended stimuli can help disentangle this apparent contradiction. In the present study, we examine if selective attention to sound pitch influences the expression of audiovisual integration in both behavior and neural activity. Participants were asked to attend to one of two auditory speech streams while watching a pair of talking lips that could be congruent or incongruent with the attended speech stream. We measured behavioral and neural responses (fMRI) to multisensory stimuli under attended and unattended conditions while physical stimulation was kept constant. Our results indicate that participants recognized words more accurately from an auditory stream that was both attended and audiovisually (AV) congruent, thus reflecting a benefit due to AV integration. On the other hand, no enhancement was found for AV congruency when it was unattended. Furthermore, the fMRI results indicated that activity in the superior temporal sulcus (an area known to be related to multisensory integration) was contingent on attention as well as on audiovisual congruency. This attentional modulation extended beyond heteromodal areas to affect processing in areas classically recognized as unisensory, such as the superior temporal gyrus or the extrastriate cortex, and to non-sensory areas such as the motor cortex. Interestingly, attention to audiovisual incongruence triggered responses in brain areas related to conflict processing (i.e., the anterior cingulate cortex and the anterior insula). Based on these results, we hypothesize that AV speech integration can take place automatically only when both modalities are sufficiently processed, and that if a mismatch is detected between the AV modalities, feedback from conflict areas minimizes the influence of this mismatch by reducing the processing of the least informative modality.