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• The brain spontaneously tracks the hierarchical structure in spoken and written languages. • Anterior and posterior scalp regions are respectively more engaged in the spoken and written syllable tracking. • Word forms are mainly represented in the anterior region in a modality-independent manner. The mental parsing of linguistic hierarchy is crucial for language comprehension, and while there is growing interest in the cortical tracking of auditory speech, the neurophysiological substrates for tracking written language are still unclear. We recorded electroencephalographic (EEG) responses from participants exposed to auditory and visual streams of either random syllables or tri-syllabic real words. Using a frequency-tagging approach, we analyzed the neural representations of physically presented (i.e., syllables) and mentally constructed (i.e., words) linguistic units and compared them between the two sensory modalities. We found that tracking syllables is partially modality dependent, with anterior and posterior scalp regions more involved in the tracking of spoken and written syllables, respectively. The cortical tracking of spoken and written words instead was found to involve a shared anterior region to a similar degree, suggesting a modality-independent process for word tracking. Our study suggests that basic linguistic features are represented in a sensory modality-specific manner, while more abstract ones are modality-unspecific during the online processing of continuous language input. The current methodology may be utilized in future research to examine the development of reading skills, especially the deficiencies in fluent reading among those with dyslexia. [ABSTRACT FROM AUTHOR]

A major scientific objective of cognitive neuroscience is to define cortico-cortical functional connections supporting cognitive functions. Here, we use an original approach combining frequency-tagging and direct electrical stimulation (DES) to test for bidirectional and cross-hemispheric category-specific modulations within the human cortical face network. A unique patient bilaterally implanted with depth electrodes in multiple face-selective cortical regions of the ventral occipito-temporal cortex (VOTC) was shown 70 s sequences of variable natural object images at a 6 Hz rate, objectively identifying deviant face-selective neural activity at 1.2 Hz (i.e., every five images). Concurrent electrical stimulation was separately applied for 10 seconds on four independently defined face-selective sites in the right and left VOTC. Upon stimulation, we observed reduced or even abolished face-selective neural activity locally and, most interestingly, at distant VOTC recording sites. Remote DES effects were found up to the anterior temporal lobe (ATL) in both forward and backward directions along the VOTC, as well as across the two hemispheres. This reduction was specific to face-selective neural activity, with the general 6 Hz visual response being mostly unaffected. Overall, these results shed light on the functional connectivity of the cortical face-selective network, supporting its non-hierarchical organization as well as bidirectional effective category-selective connections between posterior 'core' regions and the ATL. They also pave the way for widespread and systematic development of this approach to better understand the functional and effective connectivity of human brain networks. [ABSTRACT FROM AUTHOR]

Statistical language-learning, the capacity to extract regularities from a continuous speech stream, arguably involves the ability to segment the stream before the discrete constituents can be stored in memory. According to recent accounts, the segmentation process is reflected in the alignment of neural activity to the statistical structure embedded in the input. However, the degree to which it can predict the subsequent leaning outcome is currently unclear. As this is a relatively new avenue of research on statistical learning, a scoping review approach was adopted to identify and explore the current body of evidence on the use of neural phase entrainment as a measure of online neural statistical language-learning and its relation to the learning outcome, as well as the design characteristics of these studies. All included studies (11) observed entrainment to the underlying statistical pattern with exposure to the structured speech stream. A significant association between entrainment and learning outcome was observed in six of the studies. We discuss these findings in light of what neural entrainment in statistical word-learning experiments might represent, and speculate that it might reflect a general auditory processing mechanism, rather than segmentation of the speech stream per se. Lastly, as we find the current selection of studies to provide inconclusive evidence for neural entrainment's role in statistical learning, future research avenues are proposed. [ABSTRACT FROM AUTHOR]

The goal of measuring conceptual processing in numerical cognition is distanced by the possibility that neural responses to symbolic numerals are influenced by physical stimulus confounds. Here, we targeted conceptual responses to parity (even versus odd), using electroencephalogram (EEG) frequency-tagging with a symmetry/asymmetry design. Arabic numerals (2–9) were presented at 7.5 Hz in 50 s sequences; odd and even numbers were alternated to target differential, 'asymmetry' responses to parity at 3.75 Hz (7.5 Hz/2). Parity responses were probed with four different stimulus sets, increasing in intra-numeral stimulus variability, and with two control conditions composed of non-conceptual numeral alternations. Significant asymmetry responses were found over the occipitotemporal cortex to all conditions, even for the arbitrary controls. The large physical-differences control condition elicited the largest response in the stimulus set with the lowest variability (one font). Only in the stimulus set with the highest variability (20 drawn, coloured exemplars/numeral) did the response to parity surpass both control conditions. These findings show that physical differences across small sets of Arabic numerals can strongly influence, and even account for, automatic brain responses. However, carefully designed control conditions and highly variable stimulus sets may be used towards identifying truly conceptual neural responses. [ABSTRACT FROM AUTHOR]

• Concurrent recordings of brain function measured with frequency-tagging iEEG and DES. • Different faces presented at 6Hz perceived as the same during DES to right FFA. • DES to right FFA reduces concurrent face-selective neural activity at remote sites. • Frequency-tagging provides artifact-free measures during concurrent DES. • Cortical sites affected by DES effects are functionally connected. [ABSTRACT FROM AUTHOR]

Categorization is an essential cognitive and perceptual process, which happens spontaneously. However, earlier research often neglected the spontaneous nature of this process by mainly adopting explicit tasks in behavioral or neuroimaging paradigms. Here, we use frequency-tagging (FT) during electroencephalography (EEG) in 22 healthy human participants (both male and female) as a direct approach to pinpoint spontaneous visual categorical processing. Starting from schematic natural visual stimuli, we created morph sequences comprising 11 equal steps. Mirroring a behavioral categorical perception discrimination paradigm, we administered a FT-EEG oddball paradigm, assessing neural sensitivity for equally sized differences within and between stimulus categories. Likewise, mirroring a behavioral category classification paradigm, we administered a sweep FT-EEG oddball paradigm, sweeping from one end of the morph sequence to the other, thereby allowing us to objectively pinpoint the neural category boundary. We found that FT-EEG can implicitly measure categorical processing and discrimination. More specifically, we could derive an objective neural index of the required level to differentiate between the two categories, and this neural index showed the typical marker of categorical perception (i.e., stronger discrimination across as compared with within categories). The neural findings of the implicit paradigms were also validated using an explicit behavioral task. These results provide evidence that FT-EEG can be used as an objective tool to measure discrimination and categorization and that the human brain inherently and spontaneously (without any conscious or decisional processes) uses higher-level meaningful categorization information to interpret ambiguous (morph) shapes. [ABSTRACT FROM AUTHOR]

Research on attentional selection of stimulus features has yielded seemingly contradictory results. On the one hand, many experiments in humans and animals have observed a "global" facilitation of attended features across the entire visual field, even when spatial attention is focused on a single location. On the other hand, several event-related potential studies in humans reported that attended features are enhanced at the attended location only. The present experiment demonstrates that these conflicting results can be explained by differences in the timing of attentional allocation inside and outside the spatial focus of attention. Participants attended to fields of either red or blue randomly moving dots on either the left or right side of fixation with the task of detecting brief coherent motion targets. Recordings of steady-state visual evoked potentials elicited by the flickering stimuli allowed concurrent measurement of the time course of feature-selective attention in visual cortex on both the attended and the unattended sides. The onset of feature-selective attentional modulation on the attended side occurred around 150 ms earlier than on the unattended side. This finding that feature-selective attention is not spatially global from the outset but extends to unattended locations after a temporal delay resolves previous contradictions between studies finding global versus hierarchical selection of features and provides insight into the fundamental relationship between feature-based and location-based (spatial) attention mechanisms. [ABSTRACT FROM AUTHOR]

A promising stream of investigations is targeting ongoing neural oscillations and whether their modulation could be related to the perception of pain. Using an electroencephalography (EEG) frequency-tagging approach, sustained periodic thermonociceptive stimuli perceived as painful have been shown to modulate ongoing oscillations in the theta, alpha and beta bands at the frequency of stimulation. Nonetheless, it remains uncertain whether these modulations are indeed linked to pain perception. To test this relationship, we modulated pain perception using a cue-based expectation modulation paradigm and investigated whether ongoing oscillations in different frequency bands mirror the changes in stimulus perception. Forty healthy participants were instructed that a visual cue can precede either a high- or low-intensity stimulation. These cues were paired with three different levels of sustained periodic thermonociceptive stimuli (low, medium and high). Despite a strong effect of expectation on perceived stimulus intensity, this effect was not reflected in the modulation of the ongoing oscillations, suggesting a potential dissociation of pain perception and these oscillatory activities. Rather, it seems that the intensity of stimulation is the primary generator of the frequency-tagged EEG responses. Importantly, these results need to be confirmed by further investigations that could allow the detection of smaller effects than originally estimated.

The fluent processing of faces can be challenging for autistic individuals. Here, we assessed the neural sensitivity to rapid changes in subtle facial cues in 23 autistic men and 23 age and IQ matched non‐autistic (NA) controls using frequency‐tagging electroencephalography (EEG). In oddball paradigms examining the automatic and implicit discrimination of facial identity and facial expression, base rate images were presented at 6 Hz, periodically interleaved every fifth image with an oddball image (i.e. 1.2 Hz oddball frequency). These distinctive frequency tags for base rate and oddball stimuli allowed direct and objective quantification of the neural discrimination responses. We found no large differences in the neural sensitivity of participants in both groups, not for facial identity discrimination, nor for facial expression discrimination. Both groups also showed a clear face‐inversion effect, with reduced brain responses for inverted versus upright faces. Furthermore, sad faces generally elicited significantly lower neural amplitudes than angry, fearful and happy faces. The only minor group difference is the larger involvement of high‐level right‐hemisphere visual areas in NA men for facial expression processing. These findings are discussed from a developmental perspective, as they strikingly contrast with robust face processing deficits observed in autistic children using identical EEG paradigms. Lay Summary: We investigated the neural sensitivity of autistic and non‐autistic men to rapid changes in facial identity and facial expression. We found that brains of participants in both groups are capable to rapidly and implicitly detect changes in these facial cues, and that presenting faces upside‐down hampers identity discrimination to a similar extent in both groups. When processing facial expressions, autistic men apply a more low‐level visual processing strategy in comparison to controls. [ABSTRACT FROM AUTHOR]

Music listening often entails spontaneous perception and body movement to a periodic pulse‐like meter. There is increasing evidence that this cross‐cultural ability relates to neural processes that selectively enhance metric periodicities, even when these periodicities are not prominent in the acoustic stimulus. However, whether these neural processes emerge early in development remains largely unknown. Here, we recorded the electroencephalogram (EEG) of 20 healthy 5‐ to 6‐month‐old infants, while they were exposed to two rhythms known to induce the perception of meter consistently across Western adults. One rhythm contained prominent acoustic periodicities corresponding to the meter, whereas the other rhythm did not. Infants showed significantly enhanced representations of meter periodicities in their EEG responses to both rhythms. This effect is unlikely to reflect the tracking of salient acoustic features in the stimulus, as it was observed irrespective of the prominence of meter periodicities in the audio signals. Moreover, as previously observed in adults, the neural enhancement of meter was greater when the rhythm was delivered by low‐pitched sounds. Together, these findings indicate that the endogenous enhancement of metric periodicities beyond low‐level acoustic features is a neural property that is already present soon after birth. These high‐level neural processes could set the stage for internal representations of musical meter that are critical for human movement coordination during rhythmic musical behavior. Research Highlights: 5‐ to 6‐month‐old infants were presented with auditory rhythms that induce the perception of a periodic pulse‐like meter in adults.Infants showed selective enhancement of EEG activity at meter‐related frequencies irrespective of the prominence of these frequencies in the stimulus.Responses at meter‐related frequencies were boosted when the rhythm was conveyed by bass sounds.High‐level neural processes that transform rhythmic auditory stimuli into internal meter templates emerge early after birth. [ABSTRACT FROM AUTHOR]

Electroencephalography and magnetoencephalography recordings are non-invasive and temporally precise, making them invaluable tools in the investigation of neural responses in humans. However, these recordings are noisy, both because the neuronal electrodynamics involved produces a muffled signal and because the neuronal processes of interest compete with numerous other processes, from blinking to day-dreaming. One fruitful response to this noisiness has been to use stimuli with a specific frequency and to look for the signal of interest in the response at that frequency. Typically this signal involves measuring the coherence of response phase: here, a Bayesian approach to measuring phase coherence is described. This Bayesian approach is illustrated using two examples from neurolinguistics and its properties are explored using simulated data. We suggest that the Bayesian approach is more descriptive than traditional statistical approaches because it provides an explicit, interpretable generative model of how the data arises. It is also more data-efficient: it detects stimulus-related differences for smaller participant numbers than the standard approach.

The linguistic counting system of deaf signers consists of a manual counting format that uses specific structures for number words. Interestingly, the number signs from 1 to 4 in the Belgian sign languages correspond to the finger-montring habits of hearing individuals. These hand configurations could therefore be considered as signs (i.e., part of a language system) for deaf, while they would simply be number gestures (not linguistic) for hearing controls. A Fast Periodic Visual Stimulation design was used with electroencephalography recordings to examine whether these finger-number configurations are differently processed by the brain when they are signs (in deaf signers) as compared to when they are gestures (in hearing controls). Results showed that deaf signers show stronger discrimination responses to canonical fingermontring configurations compared to hearing controls. A second control experiment furthermore demonstrated that this finding was not merely due to the experience deaf signers have with the processing of hand configurations, as brain responses did not differ between groups for finger-counting configurations. Number configurations are therefore processed differently by deaf signers, but only when these configurations are part of their language system. [ABSTRACT FROM AUTHOR]

Non-human primate (NHP) neuroimaging can provide essential insights into the neural basis of human cognitive functions. While functional magnetic resonance imaging (fMRI) localizers can play an essential role in reaching this objective (Russ et al., 2021), they often differ substantially across species in terms of paradigms, measured signals, and data analysis, biasing the comparisons. Here we introduce a functional frequency-tagging face localizer for NHP imaging, successfully developed in humans and outperforming standard face localizers (Gao et al., 2018). FMRI recordings were performed in two awake macaques. Within a rapid 6 Hz stream of natural non-face objects images, human or monkey face stimuli were presented in bursts every 9 s. We also included control conditions with phase-scrambled versions of all images. As in humans, face-selective activity was objectively identified and quantified at the peak of the face-stimulation frequency (0.111 Hz) and its second harmonic (0.222 Hz) in the Fourier domain. Focal activations with a high signal-to-noise ratio were observed in regions previously described as face-selective, mainly in the STS (clusters PL, ML, MF; also, AL, AF), both for human and monkey faces. Robust face-selective activations were also found in the prefrontal cortex of one monkey (PVL and PO clusters). Face-selective neural activity was highly reliable and excluded all contributions from low-level visual cues contained in the amplitude spectrum of the stimuli. These observations indicate that fMRI frequency-tagging provides a highly valuable approach to objectively compare human and monkey visual recognition systems within the same framework.

Successfully engaging in social communication requires efficient processing of subtle socio-communicative cues. Voices convey a wealth of social information, such as gender, identity, and the emotional state of the speaker. We tested whether our brain can systematically and automatically differentiate and track a periodic stream of emotional utterances among a series of neutral vocal utterances. We recorded frequency-tagged EEG responses of 20 neurotypical male adults while presenting streams of neutral utterances at a 4 Hz base rate, interleaved with emotional utterances every third stimulus, hence at a 1.333 Hz oddball frequency. Four emotions (happy, sad, angry, and fear) were presented as different conditions in different streams. To control the impact of low-level acoustic cues, we maximized variability among the stimuli and included a control condition with scrambled utterances. This scrambling preserves low-level acoustic characteristics but ensures that the emotional character is no longer recognizable. Results revealed significant oddball EEG responses for all conditions, indicating that every emotion category can be discriminated from the neutral stimuli, and every emotional oddball response was significantly higher than the response for the scrambled utterances. These findings demonstrate that emotion discrimination is fast, automatic, and is not merely driven by low-level perceptual features. Eventually, here, we present a new database for vocal emotion research with short emotional utterances (EVID) together with an innovative frequency-tagging EEG paradigm for implicit vocal emotion discrimination. [ABSTRACT FROM AUTHOR]

Experiencing music often entails the perception of a periodic beat. Despite being a widespread phenomenon across cultures, the nature and neural underpinnings of beat perception remain largely unknown. In the last decade, there has been a growing interest in developing methods to probe these processes, particularly to measure the extent to which beat-related information is contained in behavioral and neural responses. Here, we propose a theoretical framework and practical implementation of an analytic approach to capture beat-related periodicity in empirical signals using frequency-tagging. We highlight its sensitivity in measuring the extent to which the periodicity of a perceived beat is represented in a range of continuous time-varying signals with minimal assumptions. We also discuss a limitation of this approach with respect to its specificity when restricted to measuring beat-related periodicity only from the magnitude spectrum of a signal and introduce a novel extension of the approach based on autocorrelation to overcome this issue. We test the new autocorrelation-based method using simulated signals and by re-analyzing previously published data and show how it can be used to process measurements of brain activity as captured with surface EEG in adults and infants in response to rhythmic inputs. Taken together, the theoretical framework and related methodological advances confirm and elaborate the frequency-tagging approach as a promising window into the processes underlying beat perception and, more generally, temporally coordinated behaviors., (© 2025 The Author(s). European Journal of Neuroscience published by Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

Previous intermodulation (IM) studies have employed two (or more) temporal modulations of a stimulus, with different local elements of the stimulus being modulated by different frequencies. Brain activities of IM obtained mainly from electroencephalograms (EEG) have been analyzed in the frequency domain. As a powerful tool, IM, which can provide a direct and objective physiological measure of neural interaction, has emerged as a promising method to decipher neural interactions in visual perception, and reveal the underlying different perceptual processing levels. In this review, we summarize the recent applications of IM in visual perception, detail the protocols and types of IM, and extend its utility and potential applications to the multisensory domain. We propose that using IM could prevail in partially revealing the potential hierarchical processing of multisensory information and contribute to a deeper understanding of the underlying brain dynamics. [ABSTRACT FROM AUTHOR]

In vivo intracranial recordings of neural activity offer a unique opportunity to understand human brain function. Intracranial electrophysiological (iEEG) activity related to sensory, cognitive or motor events manifests mostly in two types of signals: event-related local field potentials in lower frequency bands (30 Hz, High frequency, HF). While most current studies rely exclusively on HF, thought to be more focal and closely related to spiking activity, the relationship between HF and LF signals is unclear, especially in human associative cortex. Here, we provide a large-scale in-depth investigation of the spatial and functional relationship between these 2 signals based on intracranial recordings from 121 individual brains (8000 recording sites). We measure category-selective responses to complex ecologically salient visual stimuli – human faces – across a wide cortical territory in the ventral occipito-temporal cortex (VOTC), with a frequency-tagging method providing high signal-to-noise ratio (SNR) and the same objective quantification of signal and noise for the two frequency ranges. While LF face-selective activity has higher SNR across the VOTC, leading to a larger number of significant electrode contacts especially in the anterior temporal lobe, LF and HF display highly similar spatial, functional, and timing properties. Specifically, and contrary to a widespread assumption, our results point to nearly identical spatial distribution and local spatial extent of LF and HF activity at equal SNR. These observations go a long way towards clarifying the relationship between the two main iEEG signals and reestablish the informative value of LF iEEG to understand human brain function.

Stimulus-driven brain oscillations in the alpha range: Entrainment of intrinsic rhythms or frequency-following response? However, very few developmental studies to date applied rhythmic visual stimulation with the explicit goal to entrain endogenous rhythms (e.g., Köster et al., [32]), an approach that can help to uncover the functionality of brain oscillations in early cognitive and perception processes. What is important, this effect is dependent on the phase synchrony between the external stimulus and the brain response (Clouter et al., [9]; Wang et al., [49]), as well as the power of the response (Köster et al., [35]), which further suggests that the behavioral effects of rhythmic perceptual stimulation are caused by neural entrainment. Finally, Köster et al. ([35]) successfully entrained memory functions by stimulating participants' brain activity at their individual theta vs. alpha frequencies. [Extracted from the article]

Seeing a face in the real world provokes a host of automatic categorisations related to sex, emotion, identity, and more. Such individual facets of human face recognition have been extensively examined using overt categorisation judgements, yet their relative informational dependencies during the same face encounter are comparatively unknown. Here we used EEG to assess how increasing access to sensory input governs two ecologically relevant brain functions elicited by seeing a face: Distinguishing faces and nonfaces, and recognising people we know. Observers viewed a large set of natural images that progressively increased in either image duration (experiment 1) or spatial frequency content (experiment 2). We show that in the absence of an explicit categorisation task, the human brain requires less sensory input to categorise a stimulus as a face than it does to recognise whether that face is familiar. Moreover, where sensory thresholds for distinguishing faces/nonfaces were remarkably consistent across observers, there was high inter-individual variability in the lower informational bound for familiar face recognition, underscoring the neurofunctional distinction between these categorisation functions. By i) indexing a form of face recognition that goes beyond simple low-level differences between categories, and ii) tapping multiple recognition functions elicited by the same face encounters, the information minima we report bear high relevance to real-world face encounters, where the same stimulus is categorised along multiple dimensions at once. Thus, our finding of lower informational requirements for generic vs. familiar face recognition constitutes some of the strongest evidence to date for the intuitive notion that sensory input demands should be lower for recognising face category than face identity.

When learning a new language, one must segment words from continuous speech and associate them with meanings. These complex processes can be boosted by attentional mechanisms triggered by multi-sensory information. Previous electrophysiological studies suggest that brain oscillations are sensitive to different hierarchical complexity levels of the input, making them a plausible neural substrate for speech parsing. Here, we investigated the functional role of brain oscillations during concurrent speech segmentation and meaning acquisition in sixty 9-year-old children. We collected EEG data during an audio-visual statistical learning task during which children were exposed to a learning condition with consistent word-picture associations and a random condition with inconsistent word-picture associations before being tested on their ability to recall words and word-picture associations. We capitalized on the brain dynamics to align neural activity to the same rate as an external rhythmic stimulus to explore modulations of neural synchronization and phase synchronization between electrodes during multi-sensory word learning. Results showed enhanced power at both word- and syllabic-rate and increased EEG phase synchronization between frontal and occipital regions in the learning compared to the random condition. These findings suggest that multi-sensory cueing and attentional mechanisms play an essential role in children's successful word learning.

Humans can readily and effortlessly learn new faces encountered in the social environment. As a face transitions from unfamiliar to familiar, the ability to generalize across different images of the same person increases substantially. Fast periodic visual stimulation and EEG (FPVS-EEG) was used to isolate identity-specific responses that generalize across different images of the same person from low-level visual processing and face-general processes that aren't identity-specific. We observed these signals emerge and increase in magnitude as a group of strangers became lab mates (N=9). The neural response to an unfamiliar identity that remained unfamiliar did not change. Comparison of the response to the newly familiarised face to a highly overlearned face (the own-face) showed that this identity-specific signal was modulated by level of familiarity. The study presents the first examination of identity-specific processing changes as they occur in situ from normal, everyday face experience. [ABSTRACT FROM AUTHOR]

The extent to which faces and written words share neural circuitry in the human brain is actively debated. Here, we compare face-selective and word-selective responses in a large group of patients (N = 37) implanted with intracerebral electrodes in the ventral occipito-temporal cortex (VOTC). Both face-selective (i.e., significantly different responses to faces vs. non-face visual objects) and word-selective (i.e., significantly different responses to words vs. pseudofonts) neural activity is isolated with frequency-tagging. Critically, this sensitive approach allows to objectively quantify category-selective neural responses and disentangle them from general visual responses. About 70% of significant electrode contacts show either face-selectivity or word-selectivity only, with the expected right and left hemispheric dominance, respectively. Spatial dissociations are also found within core regions of face and word processing, with a medio-lateral dissociation in the fusiform gyrus (FG) and surrounding sulci, respectively. In the 30% of overlapping face- and word-selective contacts across the VOTC or in the FG and surrounding sulci, between-category-selective amplitudes (faces vs. words) show no-to-weak correlations, despite strong correlations in both the within-category-selective amplitudes (face–face, word–word) and the general visual responses to words and faces. Overall, these observations support the view that category-selective circuitry for faces and written words is largely dissociated in the human adult VOTC. [ABSTRACT FROM AUTHOR]

Humans perceive and spontaneously move to one or several levels of periodic pulses (a meter, for short) when listening to musical rhythm, even when the sensory input does not provide prominent periodic cues to their temporal location. Here, we review a multi-levelled framework to understanding how external rhythmic inputs are mapped onto internally represented metric pulses. This mapping is studied using an approach to quantify and directly compare representations of metric pulses in signals corresponding to sensory inputs, neural activity and behaviour (typically body movement). Based on this approach, recent empirical evidence can be drawn together into a conceptual framework that unpacks the phenomenon of meter into four levels. Each level highlights specific functional processes that critically enable and shape the mapping from sensory input to internal meter. We discuss the nature, constraints and neural substrates of these processes, starting with fundamental mechanisms investigated in macaque monkeys that enable basic forms of mapping between simple rhythmic stimuli and internally represented metric pulse. We propose that human evolution has gradually built a robust and flexible system upon these fundamental processes, allowing more complex levels of mapping to emerge in musical behaviours. This approach opens promising avenues to understand the many facets of rhythmic behaviours across individuals and species. This article is part of the theme issue 'Synchrony and rhythm interaction: from the brain to behavioural ecology'. [ABSTRACT FROM AUTHOR]

The auditory steady-state response (ASSR) is an oscillatory brain response generated by periodic auditory stimuli and originates mainly from the temporal auditory cortices. Recent data show that while the auditory cortices are indeed strongly activated by the stimulus when it is present (ON ASSR), the anatomical distribution of ASSR sources involves also parietal and frontal cortices, indicating that the ASSR is a more complex phenomenon than previously believed. Furthermore, while the ASSR typically continues to oscillate even after the stimulus has stopped (OFF ASSR), very little is known about the characteristics of the OFF ASSR and how it compares to the ON ASSR. Here, we assessed whether the OFF and ON ASSR powers are modulated by the stimulus properties (i.e. volume and pitch), selective attention, as well as individual musical sophistication. We also investigated the cortical source distribution of the OFF ASSR using a melody tracking task, in which attention was directed between uniquely amplitude-modulated melody streams that differed in pitch. The ON and OFF ASSRs were recorded with magnetoencephalography (MEG) on a group of participants varying from low to high degree of musical sophistication. Our results show that the OFF ASSR is different from the ON ASSR in nearly every aspect. While the ON ASSR was modulated by the stimulus properties and selective attention, the OFF ASSR was not influenced by any of these factors. Furthermore, while the ON ASSR was generated primarily from temporal sources, the OFF ASSR originated mainly from the frontal cortex. These findings challenge the notion that the OFF ASSR is merely a continuation of the ON ASSR. Rather, they suggest that the OFF ASSR is an internally-driven signal that develops from an initial sensory processing state (ON ASSR), with both types of ASSRs clearly differing in cortical representation and character. Furthermore, our results show that the ON ASSR power was enhanced by selective attention at cortical sources within each of the bilateral frontal, temporal, parietal and insular lobes. Finally, the ON ASSR proved sensitive to musicality, demonstrating positive correlations between musical sophistication and ASSR power, as well as with the degree of attentional ASSR modulation at the left and right parietal cortices. Taken together, these results show new aspects of the ASSR response, and demonstrate its usefulness as an effective tool for analysing how selective attention interacts with individual abilities in music perception.

Humans and other animal species are endowed with the ability to sense, represent, and mentally manipulate the number of items in a set without needing to count them. One central hypothesis is that this ability relies on an automated functional system dedicated to numerosity, the perception of the discrete numerical magnitude of a set of items. This system has classically been associated with intraparietal regions, however accumulating evidence in favor of an early visual number sense calls into question the functional role of parietal regions in numerosity processing. Targeting specifically numerosity among other visual features in the earliest stages of processing requires high temporal and spatial resolution. We used frequency-tagged magnetoencephalography to investigate the early automatic processing of numerical magnitudes and measured the steady-state brain responses specifically evoked by numerical and other visual changes in the visual scene. The neuromagnetic responses showed implicit discrimination of numerosity, total occupied area, and convex hull. The source reconstruction corresponding to the implicit discrimination responses showed common and separate sources along the ventral and dorsal visual pathways. Occipital sources attested the perceptual salience of numerosity similarly to both other implicitly discriminable visual features. Crucially, we found parietal responses uniquely associated with numerosity discrimination, showing automatic processing of numerosity in the parietal cortex, even when not relevant to the task. Taken together, these results provide further insights into the functional roles of parietal and occipital regions in numerosity encoding along the visual hierarchy. [ABSTRACT FROM AUTHOR]

• The ability to discriminate social hierarchy is critical for social interaction. • Fast periodic visual stimulation can be used to measure face perception. • The neural response indicated that subjects can discriminate dominant faces. • The results showed social hierarchical information can be detected automatically. Hierarchy is a pervasive feature of social organization. The ability to rapidly discriminate hierarchical information is critical for social interaction. Here, we took advantage of a special technique in electroencephalography (EEG) known as fast periodic visual stimulation (FPVS). We used this technique, which captures the automatic perception of faces, to explore the neural signature of social dominance discrimination. A stream of computer-generated faces was presented at 6 Hz, i.e. six faces/second. In the experimental condition, faces alternated from high to low social dominance within a sequence, bringing about a frequency of interest of 3 Hz (6 Hz/2), i.e. three high/low dominance faces appeared in one second. In two control conditions, we presented faces which came exclusively from one of two hierarchical ranks (either lower or higher). Participants were asked to respond to information unrelated to this hierarchical information, namely pressing the spacebar when the fixation changes color. Results revealed a significant 3 Hz response for the experimental condition only. This response was located bilaterally in the occipito-temporal region, indicating discrimination of differences in social dominance. Through the use of FPVS, we provide electrophysiological evidence to show that social hierarchical information can be detected automatically. [ABSTRACT FROM AUTHOR]

In humans, face-processing relies on a network of brain regions predominantly in the right occipito-temporal cortex. We tested congenitally deaf (CD) signers and matched hearing controls (HC) to investigate the experience dependence of the cortical organization of face processing. Specifically, we used EEG frequency-tagging to evaluate: (1) Face-Object Categorization, (2) Emotional Facial-Expression Discrimination and (3) Individual Face Discrimination. The EEG was recorded to visual stimuli presented at a rate of 6 Hz, with oddball stimuli at a rate of 1.2 Hz. In all three experiments and in both groups, significant face discriminative responses were found. Face-Object categorization was associated to a relative increased involvement of the left hemisphere in CD individuals compared to HC individuals. A similar trend was observed for Emotional Facial-Expression discrimination but not for Individual Face Discrimination. Source reconstruction suggested a greater activation of the auditory cortices in the CD group for Individual Face Discrimination. These findings suggest that the experience dependence of the relative contribution of the two hemispheres as well as crossmodal plasticity vary with different aspects of face processing.

Research on the neural basis of conscious perception has almost exclusively shown that becoming aware of a stimulus leads to increased neural responses. By designing a novel form of perceptual filling-in (PFI) overlaid with a dynamic texture display, we frequency-tagged multiple disappearing targets as well as their surroundings. We show that in a PFI paradigm, the disappearance of a stimulus and subjective invisibility is associated with increases in neural activity, as measured with steady-state visually evoked potentials (SSVEPs), in electroencephalography (EEG). We also find that this increase correlates with alpha-band activity, a well-established neural measure of attention. These findings cast doubt on the direct relationship previously reported between the strength of neural activity and conscious perception, at least when measured with current tools, such as the SSVEP. Instead, we conclude that SSVEP strength more closely measures changes in attention.

Selective auditory attention allows us to focus on relevant sounds within noisy or complex auditory environments, and is essential for the processing of speech and music. The auditory steady-state response (ASSR) has been proposed as a neural measure for tracking selective auditory attention, even within continuous and complex soundscapes. However, the current literature is inconsistent on how the ASSR is influenced by selective attention, with findings based primarily on attention being directed to either ear rather than to sound content. In this experiment, a mixture of melody streams was presented to both ears identically (diotically) as we examined if selective auditory attention to sound content influences the ASSR. Using magnetoencephalography (MEG), we assessed the stream-specific ASSRs from three frequency-tagged melody streams when attention was directed between each melody stream, based on their respective pitch and timing. Our main results showed that selective attention enhances the ASSR power of an attended melody stream by 14% at a general sensor level. This ability to readily capture attentional changes in a stimuli-precise manner makes the ASSR a useful tool for studying selective auditory attention, especially in complex auditory environments. As a secondary aim, we explored the distribution of cortical ASSR sources and their respective attentional modulation using a distributed source model of the ASSR activity. Notably, we uncovered the existence of ASSR attentional modulation outside the temporal cortices. Across-subject averages of the attentional enhancement over the cortical surface suggest that frontal regions show up to ~80% enhancement, while temporal and parietal cortices were enhanced by 20–25%. Importantly, this work advocates a novel ‘beyond the temporal cortex’ perspective on ASSR modulation and also serves as a template for future studies to precisely pin-point which cortical sites are more susceptible to ASSR attentional modulation.

To investigate face individuation (FI), a critical brain function in the human species, an oddball fast periodic visual stimulation (FPVS) approach was recently introduced (Liu‐Shuang et al., Neuropsychologia, 2014, 52, 57). In this paradigm, an image of an unfamiliar "base" facial identity is repeated at a rapid rate F (e.g., 6 Hz) and different unfamiliar "oddball" facial identities are inserted every nth item, at a F/n rate (e.g., every 5th item, 1.2 Hz). This stimulation elicits FI responses at F/n and its harmonics (2F/n, 3F/n, etc.), reflecting neural discrimination between oddball versus base facial identities, which is quantified in the frequency domain of the electroencephalogram (EEG). This paradigm, used in 20 published studies, demonstrates substantial advantages for measuring FI in terms of validity, objectivity, reliability, and sensitivity. Human intracerebral recordings suggest that this FI response originates from neural populations in the lateral inferior occipital and fusiform gyri, with a right hemispheric dominance consistent with the localization of brain lesions specifically affecting facial identity recognition (prosopagnosia). Here, we summarize the contributions of the oddball FPVS framework toward understanding FI, including its (a)typical development, with early studies supporting the application of this technique to clinical testing (e.g., autism spectrum disorder). This review also includes an in‐depth analysis of the paradigm's methodology, with guidelines for designing future studies. A large‐scale group analysis compiling data across 130 observers provides insights into the oddball FPVS FI response properties. Overall, we recommend the oddball FPVS paradigm as an alternative approach to behavioral or traditional event‐related potential EEG measures of face individuation. [ABSTRACT FROM AUTHOR]

Highly variable natural images of the same familiar face celebrity interleaved periodically in a rapid (6 images/second) train of unfamiliar faces automatically elicit an objective electroencephalographic (EEG) response over the occipito-temporal cortex of neurotypical human adults within a few minutes. However, the extent to which this frequency-tagged response goes beyond the association of common physical features of the periodically repeated face identity remains unknown. Here we compare participants who know or do not know the very same periodically repeated face celebrity and show that long-term familiarity accounts for about 80% of the neural face identity recognition response. This familiarity advantage disappears with upside-down images. Variability in response amplitude between face identities is preserved for inverted faces and in unfamiliar participants, suggesting a contribution of within-person physical face variability and distinctiveness to about 20% of the face identity response. These observations provide the strongest difference to date in human brain response between the same famous face identities perceived as familiar or unfamiliar in an implicit task. The frequency-tagged neural response largely reflects the strengthening effect of long-term memory in the human occipito-temporal cortex, and may serve to index automatic familiar face identity recognition in individual observers. [ABSTRACT FROM AUTHOR]

It is debated whether perceptual expertise of nonface objects, such as visual words, is indicated by holistic processing, which is regarded as a marker of perceptual expertise of faces. We address this question by frequency-tagged electroencephalography. Different parts of real or pseudo Chinese characters are presented at distinctive frequencies (6 or 7.2 Hz), which induce frequency-tagged steady-state visual-evoked potentials at occipital brain areas. The intermodulation response (e.g., 6 + 7.2 = 13.2 Hz) would emerge when holistic integration takes place. Our results suggest that the intermodulation response to the real characters is left lateralized, which is contralateral to previous findings with faces. Furthermore, at the left occipital area, the intermodulation response to real characters is more prominent than pseudo characters, suggesting that holistic integration is enhanced for real characters than for pseudo ones. Taken together, our findings suggest that holistic integration is potentially a general expertise marker for both faces and non-face objects. [ABSTRACT FROM AUTHOR]

Humans coordinate their movements with one another in a range of everyday activities and skill domains. Optimal joint performance requires the continuous anticipation of and adaptation to each other’s movements, especially when actions are spontaneous rather than pre-planned. Here we employ dual-EEG and frequency-tagging techniques to investigate how the neural tracking of self- and other-generated movements supports interpersonal coordination during improvised motion. LEDs flickering at 5.7 and 7.7 Hz were attached to participants’ index fingers in 28 dyads as they produced novel patterns of synchronous horizontal forearm movements. EEG responses at these frequencies revealed enhanced neural tracking of self-generated movement when leading and of other-generated movements when following. A marker of self-other integration at 13.4 Hz (inter-modulation frequency of 5.7 and 7.7 Hz) peaked when no leader was designated, and mutual adaptation and movement synchrony were maximal. Furthermore, the amplitude of EEG responses reflected differences in the capacity of dyads to synchronize their movements, offering a neurophysiologically grounded perspective for understanding perceptual-motor mechanisms underlying joint action.

To successfully interact with a rich and ambiguous visual environment, the human brain learns to differentiate visual stimuli and to produce the same response to subsets of these stimuli despite their physical difference. Although this visual categorization function is traditionally investigated from a unisensory perspective, its early development is inherently constrained by multisensory inputs. In particular, an early‐maturing sensory system such as olfaction is ideally suited to support the immature visual system in infancy by providing stability and familiarity to a rapidly changing visual environment. Here, we test the hypothesis that rapid visual categorization of salient visual signals for the young infant brain, human faces, is shaped by another highly relevant human‐related input from the olfactory system, the mother's body odor. We observe that a right‐hemispheric neural signature of single‐glance face categorization from natural images is significantly enhanced in the maternal versus a control odor context in individual 4‐month‐old infant brains. A lack of difference between odor conditions for the common brain response elicited by both face and non‐face images rules out a mere enhancement of arousal or visual attention in the maternal odor context. These observations show that face‐selective neural activity in infancy is mediated by the presence of a (maternal) body odor, providing strong support for multisensory inputs driving category acquisition in the developing human brain and having important implications for our understanding of human perceptual development. [ABSTRACT FROM AUTHOR]

When listening to music, people often perceive and move along with a periodic meter. However, the dynamics of mapping between meter perception and the acoustic cues to meter periodicities in the sensory input remain largely unknown. To capture these dynamics, we recorded the electroencephalography while nonmusician and musician participants listened to nonrepeating rhythmic sequences, where acoustic cues to meter frequencies either gradually decreased (from regular to degraded) or increased (from degraded to regular). The results revealed greater neural activity selectively elicited at meter frequencies when the sequence gradually changed from regular to degraded compared with the opposite. Importantly, this effect was unlikely to arise from overall gain, or low-level auditory processing, as revealed by physiological modeling. Moreover, the context effect was more pronounced in nonmusicians, who also demonstrated facilitated sensory-motor synchronization with the meter for sequences that started as regular. In contrast, musicians showed weaker effects of recent context in their neural responses and robust ability to move along with the meter irrespective of stimulus degradation. Together, our results demonstrate that brain activity elicited by rhythm does not only reflect passive tracking of stimulus features, but represents continuous integration of sensory input with recent context. [ABSTRACT FROM AUTHOR]

Automatic responses to brief expression changes from a neutral face have been recently isolated in the human brain using fast periodic visual stimulation (FPVS) coupled with scalp electroencephalography (EEG). Based on these observations, here we isolate specific neural signatures for the rapid categorization of each of 5 basic expressions, i.e., when they are directly discriminated from all other facial expressions. Scalp EEG was recorded in 15 participants presented with pictures alternating at a rapid 6 Hz rate (i.e., one fixation/face, backward- and forward-masked). In different stimulation sequences, an expressive (angry, disgusted, happy, fearful, or sad) or a neutral face arose every 5 pictures (i.e., at 6/5 = 1.2 Hz), among pictures of the same individual expressing the other emotions randomly. Frequency-domain analysis indicated a robust (i.e., recorded in every individual participant) and objective (i.e., at the predefined 1.2 Hz frequency and its harmonics) expression-specific brain response over occipito-temporal sites for each emotion and neutrality. In this context of variable expressions, while neural responses to the different expressions (Anger, Disgust, Happiness, Sadness) were dissimilar qualitatively, a much larger specific signature for neutral faces as compared to facial expressions was found. Interestingly, Fear also elicited a strong contrasted response with other facial expressions, associated with a specific neural signature over ventral occipito-temporal sites. Collectively, these findings reveal that specific EEG signatures for different facial expressions can be isolated in the human brain, pointing to partially different neural substrates. In addition, they provide support for a strong and highly selective neural response to fear at the system-level, in line with the importance of this emotional expression for biological survival. • We provide a neural marker for direct discrimination between facial expressions. • Specific brain signatures are isolated for each basic facial expression. • Fear and neutrality elicit larger and qualitatively distinct brain responses. • Neutral and fearful faces may have a particular status in the human brain. [ABSTRACT FROM AUTHOR]

Voices are the most relevant social sounds for humans and therefore have crucial adaptive value in development. Neuroimaging studies in adults have demonstrated the existence of regions in the superior temporal sulcus that respond preferentially to voices. Yet, whether voices represent a functionally specific category in the young infant's mind is largely unknown. We developed a highly sensitive paradigm relying on fast periodic auditory stimulation (FPAS) combined with scalp electroencephalography (EEG) to demonstrate that the infant brain implements a reliable preferential response to voices early in life. Twenty-three 4-month-old infants listened to sequences containing non-vocal sounds from different categories presented at 3.33 Hz, with highly heterogeneous vocal sounds appearing every third stimulus (1.11 Hz). We were able to isolate a voice-selective response over temporal regions, and individual voice-selective responses were found in most infants within only a few minutes of stimulation. This selective response was significantly reduced for the same frequency-scrambled sounds, indicating that voice selectivity is not simply driven by the envelope and the spectral content of the sounds. Such a robust selective response to voices as early as 4 months of age suggests that the infant brain is endowed with the ability to rapidly develop a functional selectivity to this socially relevant category of sounds. • Selective brain responses to voices emerge as early as 4 months of age • Voice selectivity in infants is partially independent of acoustic features • The topography of voice selectivity in infants resembles the adult one • The infant brain is endowed with selectivity for socially relevant sounds Calce et al. rely on a highly sensitive paradigm combining fast periodic auditory stimulation (FPAS) with scalp electroencephalography (EEG) to demonstrate that the infant brain produces a reliable preferential response to voices early in life. [ABSTRACT FROM AUTHOR]