Max Planck Institute for Empirical Aesthetics

The human brain has evolved for group living [1Dunbar I.R. The social brain hypothesis.in: Cacioppo J.T. Foundations in Social Neuroscience. MIT Press, 2002: 69-88Google Scholar]. Yet we know so little about how it supports dynamic group interactions that the study of real-world social exchanges has been dubbed the “dark matter of social neuroscience” [2Schilbach L. Timmermans B. Reddy V. Costall A. Bente G. Schlicht T. Vogeley K. Toward a second-person neuroscience.Behav. Brain Sci. 2013; 36: 393-414Crossref PubMed Scopus (890) Google Scholar]. Recently, various studies have begun to approach this question by comparing brain responses of multiple individuals during a variety of (semi-naturalistic) tasks [3Babiloni F. Astolfi L. Social neuroscience and hyperscanning techniques: past, present and future.Neurosci. Biobehav. Rev. 2014; 44: 76-93Crossref PubMed Scopus (289) Google Scholar, 4Dmochowski J.P. Bezdek M.A. Abelson B.P. Johnson J.S. Schumacher E.H. Parra L.C. Audience preferences are predicted by temporal reliability of neural processing.Nat. Commun. 2014; 5: 4567Crossref PubMed Scopus (164) Google Scholar, 5Dumas G. Nadel J. Soussignan R. Martinerie J. Garnero L. Inter-brain synchronization during social interaction.PLoS ONE. 2010; 5: e12166Crossref PubMed Scopus (495) Google Scholar, 6Hasson U. Ghazanfar A.A. Galantucci B. Garrod S. Keysers C. Brain-to-brain coupling: a mechanism for creating and sharing a social world.Trends Cogn. Sci. 2012; 16: 114-121Abstract Full Text Full Text PDF PubMed Scopus (545) Google Scholar, 7Hari R. Himberg T. Nummenmaa L. Hämäläinen M. Parkkonen L. Synchrony of brains and bodies during implicit interpersonal interaction.Trends Cogn. Sci. 2013; 17: 105-106Abstract Full Text Full Text PDF PubMed Scopus (67) Google Scholar, 8Jiang J. Chen C. Dai B. Shi G. Ding G. Liu L. Lu C. Leader emergence through interpersonal neural synchronization.Proc. Natl. Acad. Sci. USA. 2015; 112: 4274-4279Crossref PubMed Scopus (168) Google Scholar, 9Pfeiffer U.J. Timmermans B. Vogeley K. Frith C.D. Schilbach L. Towards a neuroscience of social interaction.Front. Hum. Neurosci. 2013; 7: 22Crossref PubMed Scopus (30) Google Scholar, 10Stephens G.J. Silbert L.J. Hasson U. Speaker-listener neural coupling underlies successful communication.Proc. Natl. Acad. Sci. USA. 2010; 107: 14425-14430Crossref PubMed Scopus (546) Google Scholar, 11Babiloni C. Buffo P. Vecchio F. Marzano N. Del Percio C. Spada D. Rossi S. Bruni I. Rossini P.M. Perani D. Brains “in concert”: frontal oscillatory alpha rhythms and empathy in professional musicians.Neuroimage. 2012; 60: 105-116Crossref PubMed Scopus (84) Google Scholar, 12Duan L. Dai R.N. Xiao X. Sun P.P. Li Z. Zhu C.Z. Cluster imaging of multi-brain networks (CIMBN): a general framework for hyperscanning and modeling a group of interacting brains.Front. Neurosci. 2015; 9: 267Crossref PubMed Scopus (28) Google Scholar, 13Hasson U. Nir Y. Levy I. Fuhrmann G. Malach R. Intersubject synchronization of cortical activity during natural vision.Science. 2004; 303: 1634-1640Crossref PubMed Scopus (1063) Google Scholar, 14Nummenmaa L. Glerean E. Viinikainen M. Jääskeläinen I.P. Hari R. Sams M. Emotions promote social interaction by synchronizing brain activity across individuals.Proc. Natl. Acad. Sci. USA. 2012; 109: 9599-9604Crossref PubMed Scopus (287) Google Scholar, 15Dikker S. Silbert L.J. Hasson U. Zevin J.D. On the same wavelength: predictable language enhances speaker-listener brain-to-brain synchrony in posterior superior temporal gyrus.J. Neurosci. 2014; 34: 6267-6272Crossref PubMed Scopus (71) Google Scholar]. These experiments reveal how stimulus properties [13Hasson U. Nir Y. Levy I. Fuhrmann G. Malach R. Intersubject synchronization of cortical activity during natural vision.Science. 2004; 303: 1634-1640Crossref PubMed Scopus (1063) Google Scholar], individual differences [14Nummenmaa L. Glerean E. Viinikainen M. Jääskeläinen I.P. Hari R. Sams M. Emotions promote social interaction by synchronizing brain activity across individuals.Proc. Natl. Acad. Sci. USA. 2012; 109: 9599-9604Crossref PubMed Scopus (287) Google Scholar], and contextual factors [15Dikker S. Silbert L.J. Hasson U. Zevin J.D. On the same wavelength: predictable language enhances speaker-listener brain-to-brain synchrony in posterior superior temporal gyrus.J. Neurosci. 2014; 34: 6267-6272Crossref PubMed Scopus (71) Google Scholar] may underpin similarities and differences in neural activity across people. However, most studies to date suffer from various limitations: they often lack direct face-to-face interaction between participants, are typically limited to dyads, do not investigate social dynamics across time, and, crucially, they rarely study social behavior under naturalistic circumstances. Here we extend such experimentation drastically, beyond dyads and beyond laboratory walls, to identify neural markers of group engagement during dynamic real-world group interactions. We used portable electroencephalogram (EEG) to simultaneously record brain activity from a class of 12 high school students over the course of a semester (11 classes) during regular classroom activities (Figures 1A–1C; Supplemental Experimental Procedures, section S1). A novel analysis technique to assess group-based neural coherence demonstrates that the extent to which brain activity is synchronized across students predicts both student class engagement and social dynamics. This suggests that brain-to-brain synchrony is a possible neural marker for dynamic social interactions, likely driven by shared attention mechanisms. This study validates a promising new method to investigate the neuroscience of group interactions in ecologically natural settings.

Noninvasive brain stimulation techniques are used in experimental and clinical fields for their potential effects on brain network dynamics and behavior. Transcranial electrical stimulation (TES), including transcranial direct current stimulation (tDCS) and transcranial alternating current stimulation (tACS), has gained popularity because of its convenience and potential as a chronic therapy. However, a mechanistic understanding of TES has lagged behind its widespread adoption. Here, we review data and modelling on the immediate neurophysiological effects of TES in vitro as well as in vivo in both humans and other animals. While it remains unclear how typical TES protocols affect neural activity, we propose that validated models of current flow should inform study design and artifacts should be carefully excluded during signal recording and analysis. Potential indirect effects of TES (e.g., peripheral stimulation) should be investigated in more detail and further explored in experimental designs. We also consider how novel technologies may stimulate the next generation of TES experiments and devices, thus enhancing validity, specificity, and reproducibility.

What is universal about music, and what varies? We built a corpus of ethnographic text on musical behavior from a representative sample of the world's societies, as well as a discography of audio recordings. The ethnographic corpus reveals that music (including songs with words) appears in every society observed; that music varies along three dimensions (formality, arousal, religiosity), more within societies than across them; and that music is associated with certain behavioral contexts such as infant care, healing, dance, and love. The discography-analyzed through machine summaries, amateur and expert listener ratings, and manual transcriptions-reveals that acoustic features of songs predict their primary behavioral context; that tonality is widespread, perhaps universal; that music varies in rhythmic and melodic complexity; and that elements of melodies and rhythms found worldwide follow power laws.

Departing from traditional linguistic models, advances in deep learning have resulted in a new type of predictive (autoregressive) deep language models (DLMs). Using a self-supervised next-word prediction task, these models generate appropriate linguistic responses in a given context. In the current study, nine participants listened to a 30-min podcast while their brain responses were recorded using electrocorticography (ECoG). We provide empirical evidence that the human brain and autoregressive DLMs share three fundamental computational principles as they process the same natural narrative: (1) both are engaged in continuous next-word prediction before word onset; (2) both match their pre-onset predictions to the incoming word to calculate post-onset surprise; (3) both rely on contextual embeddings to represent words in natural contexts. Together, our findings suggest that autoregressive DLMs provide a new and biologically feasible computational framework for studying the neural basis of language.

Why are negative emotions so central in art reception far beyond tragedy? Revisiting classical aesthetics in the light of recent psychological research, we present a novel model to explain this much discussed (apparent) paradox. We argue that negative emotions are an important resource for the arts in general, rather than a special license for exceptional art forms only. The underlying rationale is that negative emotions have been shown to be particularly powerful in securing attention, intense emotional involvement, and high memorability, and hence is precisely what artworks strive for. Two groups of processing mechanisms are identified that conjointly adopt the particular powers of negative emotions for art's purposes. The first group consists of psychological distancing mechanisms that are activated along with the cognitive schemata of art, representation, and fiction. These schemata imply personal safety and control over continuing or discontinuing exposure to artworks, thereby preventing negative emotions from becoming outright incompatible with expectations of enjoyment. This distancing sets the stage for a second group of processing components that allow art recipients to positively embrace the experiencing of negative emotions, thereby rendering art reception more intense, more interesting, more emotionally moving, more profound, and occasionally even more beautiful. These components include compositional interplays of positive and negative emotions, the effects of aesthetic virtues of using the media of (re)presentation (musical sound, words/language, color, shapes) on emotion perception, and meaning-making efforts. Moreover, our Distancing-Embracing model proposes that concomitant mixed emotions often help integrate negative emotions into altogether pleasurable trajectories.

BACKGROUND: Time-limited, early-life exposures to institutional deprivation are associated with disorders in childhood, but it is unknown whether effects persist into adulthood. We used data from the English and Romanian Adoptees study to assess whether deprivation-associated adverse neurodevelopmental and mental health outcomes persist into young adulthood. METHODS: The English and Romanian Adoptees study is a longitudinal, natural experiment investigation into the long-term outcomes of individuals who spent from soon after birth to up to 43 months in severe deprivation in Romanian institutions before being adopted into the UK. We used developmentally appropriate standard questionnaires, interviews completed by parents and adoptees, and direct measures of IQ to measure symptoms of autism spectrum disorder, inattention and overactivity, disinhibited social engagement, conduct or emotional problems, and cognitive impairment (IQ score <80) during childhood (ages 6, 11, and 15 years) and in young adulthood (22-25 years). For analysis, Romanian adoptees were split into those who spent less than 6 months in an institution and those who spent more than 6 months in an institution. We used a comparison group of UK adoptees who did not experience deprivation. We used mixed-effects regression models for ordered-categorical outcome variables to compare symptom levels and trends between groups. FINDINGS: Romanian adoptees who experienced less than 6 months in an institution (n=67 at ages 6 years; n=50 at young adulthood) and UK controls (n=52 at age 6 years; n=39 at young adulthood) had similarly low levels of symptoms across most ages and outcomes. By contrast, Romanian adoptees exposed to more than 6 months in an institution (n=98 at ages 6 years; n=72 at young adulthood) had persistently higher rates than UK controls of symptoms of autism spectrum disorder, disinhibited social engagement, and inattention and overactivity through to young adulthood (pooled p<0·0001 for all). Cognitive impairment in the group who spent more than 6 months in an institution remitted from markedly higher rates at ages 6 years (p=0·0001) and 11 years (p=0·0016) compared with UK controls, to normal rates at young adulthood (p=0·76). By contrast, self-rated emotional symptoms showed a late-onset pattern with minimal differences versus UK controls at ages 11 years (p=0·0449) and 15 years (p=0·17), and then marked increases by young adulthood (p=0·0005), with similar effects seen for parent ratings. The high deprivation group also had a higher proportion of people with low educational achievement (p=0·0195), unemployment (p=0·0124), and mental health service use (p=0·0120, p=0·0032, and p=0·0003 for use when aged <11 years, 11-14 years, and 15-23 years, respectively) than the UK control group. A fifth (n=15) of individuals who spent more than 6 months in an institution were problem-free at all assessments. INTERPRETATION: Notwithstanding the resilience shown by some adoptees and the adult remission of cognitive impairment, extended early deprivation was associated with long-term deleterious effects on wellbeing that seem insusceptible to years of nurturance and support in adoptive families. FUNDING: Economic and Social Research Council, Medical Research Council, Department of Health, Jacobs Foundation, Nuffield Foundation.

Aesthetic perception and judgement are not merely cognitive processes, but also involve feelings. Therefore, the empirical study of these experiences requires conceptualization and measurement of aesthetic emotions. Despite the long-standing interest in such emotions, we still lack an assessment tool to capture the broad range of emotions that occur in response to the perceived aesthetic appeal of stimuli. Elicitors of aesthetic emotions are not limited to the arts in the strict sense, but extend to design, built environments, and nature. In this article, we describe the development of a questionnaire that is applicable across many of these domains: the Aesthetic Emotions Scale (Aesthemos). Drawing on theoretical accounts of aesthetic emotions and an extensive review of extant measures of aesthetic emotions within specific domains such as music, literature, film, painting, advertisements, design, and architecture, we propose a framework for studying aesthetic emotions. The Aesthemos, which is based on this framework, contains 21 subscales with two items each, that are designed to assess the emotional signature of responses to stimuli's perceived aesthetic appeal in a highly differentiated manner. These scales cover prototypical aesthetic emotions (e.g., the feeling of beauty, being moved, fascination, and awe), epistemic emotions (e.g., interest and insight), and emotions indicative of amusement (humor and joy). In addition, the Aesthemos subscales capture both the activating (energy and vitality) and the calming (relaxation) effects of aesthetic experiences, as well as negative emotions that may contribute to aesthetic displeasure (e.g., the feeling of ugliness, boredom, and confusion).

Recent studies establish that cortical oscillations track naturalistic speech in a remarkably faithful way. Here, we test whether such neural activity, particularly low-frequency (<8 Hz; delta-theta) oscillations, similarly entrain to music and whether experience modifies such a cortical phenomenon. Music of varying tempi was used to test entrainment at different rates. In three magnetoencephalography experiments, we recorded from nonmusicians, as well as musicians with varying years of experience. Recordings from nonmusicians demonstrate cortical entrainment that tracks musical stimuli over a typical range of tempi, but not at tempi below 1 note per second. Importantly, the observed entrainment correlates with performance on a concurrent pitch-related behavioral task. In contrast, the data from musicians show that entrainment is enhanced by years of musical training, at all presented tempi. This suggests a bidirectional relationship between behavior and cortical entrainment, a phenomenon that has not previously been reported. Additional analyses focus on responses in the beta range (∼15-30 Hz)-often linked to delta activity in the context of temporal predictions. Our findings provide evidence that the role of beta in temporal predictions scales to the complex hierarchical rhythms in natural music and enhances processing of musical content. This study builds on important findings on brainstem plasticity and represents a compelling demonstration that cortical neural entrainment is tightly coupled to both musical training and task performance, further supporting a role for cortical oscillatory activity in music perception and cognition.

This is the first comprehensive theoretical article on aesthetic emotions. Following Kant's definition, we propose that it is the first and foremost characteristic of aesthetic emotions to make a direct contribution to aesthetic evaluation/appreciation. Each aesthetic emotion is tuned to a special type of perceived aesthetic appeal and is predictive of the subjectively felt pleasure or displeasure and the liking or disliking associated with this type of appeal. Contrary to the negativity bias of classical emotion catalogues, emotion terms used for aesthetic evaluation purposes include far more positive than negative emotions. At the same time, many overall positive aesthetic emotions encompass negative or mixed emotional ingredients. Appraisals of intrinsic pleasantness, familiarity, and novelty are preeminently important for aesthetic emotions. Appraisals of goal relevance/conduciveness and coping potential are largely irrelevant from a pragmatic perspective, but in some cases highly relevant for cognitive and affective coping. Aesthetic emotions are typically sought and savored for their own sake, with subjectively felt intensity and/or emotional arousal being rewards in their own right. The expression component of aesthetic emotions includes laughter, tears, and facial and bodily movements, along with applause or booing and words of praise or blame. Aesthetic emotions entail motivational approach and avoidance tendencies, specifically, tendencies toward prolonged, repeated, or interrupted exposure and wanting to possess aesthetically pleasing objects. They are experienced across a broad range of experiential domains and not coextensive with art-elicited emotions. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

The emotional state of being moved, though frequently referred to in both classical rhetoric and current language use, is far from established as a well-defined psychological construct. In a series of three studies, we investigated eliciting scenarios, emotional ingredients, appraisal patterns, feeling qualities, and the affective signature of being moved and related emotional states. The great majority of the eliciting scenarios can be assigned to significant relationship and critical life events (especially death, birth, marriage, separation, and reunion). Sadness and joy turned out to be the two preeminent emotions involved in episodes of being moved. Both the sad and the joyful variants of being moved showed a coactivation of positive and negative affect and can thus be ranked among the mixed emotions. Moreover, being moved, while featuring only low-to-mid arousal levels, was experienced as an emotional state of high intensity; this applied to responses to fictional artworks no less than to own-life and other real, but media-represented, events. The most distinctive findings regarding cognitive appraisal dimensions were very low ratings for causation of the event by oneself and for having the power to change its outcome, along with very high ratings for appraisals of compatibility with social norms and self-ideals. Putting together the characteristics identified and discussed throughout the three studies, the paper ends with a sketch of a psychological construct of being moved.

How does the human brain support real-world learning? We used wireless electroencephalography to collect neurophysiological data from a group of 12 senior high school students and their teacher during regular biology lessons. Six scheduled classes over the course of the semester were organized such that class materials were presented using different teaching styles (videos and lectures), and students completed a multiple-choice quiz after each class to measure their retention of that lesson's content. Both students' brain-to-brain synchrony and their content retention were higher for videos than lectures across the six classes. Brain-to-brain synchrony between the teacher and students varied as a function of student engagement as well as teacher likeability: Students who reported greater social closeness to the teacher showed higher brain-to-brain synchrony with the teacher, but this was only the case for lectures-that is, when the teacher is an integral part of the content presentation. Furthermore, students' retention of the class content correlated with student-teacher closeness, but not with brain-to-brain synchrony. These findings expand on existing social neuroscience research by showing that social factors such as perceived closeness are reflected in brain-to-brain synchrony in real-world group settings and can predict cognitive outcomes such as students' academic performance.

It is a common experience-and well established experimentally-that music can engage us emotionally in a compelling manner. The mechanisms underlying these experiences are receiving increasing scrutiny. However, the extent to which other domains of aesthetic experience can similarly elicit strong emotions is unknown. Using psychophysiology, neuroimaging and behavioral responses, we show that recited poetry can act as a powerful stimulus for eliciting peak emotional responses, including chills and objectively measurable goosebumps that engage the primary reward circuitry. Importantly, while these responses to poetry are largely analogous to those found for music, their neural underpinnings show important differences, specifically with regard to the crucial role of the nucleus accumbens. We also go beyond replicating previous music-related studies by showing that peak aesthetic pleasure can co-occur with physiological markers of negative affect. Finally, the distribution of chills across the trajectory of poems provides insight into compositional principles of poetry.

The hippocampus is crucial for episodic memory, but it is also involved in online prediction. Evidence suggests that a unitary hippocampal code underlies both episodic memory and predictive processing, yet within a predictive coding framework the hippocampal-neocortical interactions that accompany these two phenomena are distinct and opposing. Namely, during episodic recall, the hippocampus is thought to exert an excitatory influence on the neocortex, to reinstate activity patterns across cortical circuits. This contrasts with empirical and theoretical work on predictive processing, where descending predictions suppress prediction errors to 'explain away' ascending inputs via cortical inhibition. In this hypothesis piece, we attempt to dissolve this previously overlooked dialectic. We consider how the hippocampus may facilitate both prediction and memory, respectively, by inhibiting neocortical prediction errors or increasing their gain. We propose that these distinct processing modes depend upon the neuromodulatory gain (or precision) ascribed to prediction error units. Within this framework, memory recall is cast as arising from fictive prediction errors that furnish training signals to optimise generative models of the world, in the absence of sensory data.

Since the second-half of the twentieth century, intracranial electroencephalography (iEEG), including both electrocorticography (ECoG) and stereo-electroencephalography (sEEG), has provided an intimate view into the human brain. At the interface between fundamental research and the clinic, iEEG provides both high temporal resolution and high spatial specificity but comes with constraints, such as the individual's tailored sparsity of electrode sampling. Over the years, researchers in neuroscience developed their practices to make the most of the iEEG approach. Here we offer a critical review of iEEG research practices in a didactic framework for newcomers, as well addressing issues encountered by proficient researchers. The scope is threefold: (i) review common practices in iEEG research, (ii) suggest potential guidelines for working with iEEG data and answer frequently asked questions based on the most widespread practices, and (iii) based on current neurophysiological knowledge and methodologies, pave the way to good practice standards in iEEG research. The organization of this paper follows the steps of iEEG data processing. The first section contextualizes iEEG data collection. The second section focuses on localization of intracranial electrodes. The third section highlights the main pre-processing steps. The fourth section presents iEEG signal analysis methods. The fifth section discusses statistical approaches. The sixth section draws some unique perspectives on iEEG research. Finally, to ensure a consistent nomenclature throughout the manuscript and to align with other guidelines, e.g., Brain Imaging Data Structure (BIDS) and the OHBM Committee on Best Practices in Data Analysis and Sharing (COBIDAS), we provide a glossary to disambiguate terms related to iEEG research.

Preprocessing of functional MRI (fMRI) involves numerous steps to clean and standardize data before statistical analysis. Generally, researchers create ad hoc preprocessing workflows for each new dataset, building upon a large inventory of tools available for each step. The complexity of these workflows has snowballed with rapid advances in MR data acquisition and image processing techniques. We introduce fMRIPrep , an analysis-agnostic tool that addresses the challenge of robust and reproducible preprocessing for task-based and resting fMRI data. FMRIPrep automatically adapts a best-in-breed workflow to the idiosyncrasies of virtually any dataset, ensuring high-quality preprocessing with no manual intervention. By introducing visual assessment checkpoints into an iterative integration framework for software-testing, we show that fMRIPrep robustly produces high-quality results on a diverse fMRI data collection comprising participants from 54 different studies in the OpenfMRI repository. We review the distinctive features of fMRIPrep in a qualitative comparison to other preprocessing workflows. We demonstrate that fMRIPrep achieves higher spatial accuracy as it introduces less uncontrolled spatial smoothness than commonly used preprocessing tools. FMRIPrep has the potential to transform fMRI research by equipping neuroscientists with a high-quality, robust, easy-to-use and transparent preprocessing workflow which can help ensure the validity of inference and the interpretability of their results.

Using the method or time-delayed embedding, a signal can be embedded into higher-dimensional space in order to study its dynamics. This requires knowledge of two parameters: The delay parameter $\tau$, and the embedding dimension parameter $D$. Two standard methods to estimate these parameters in one-dimensional time-series involve the inspection of the Average Mutual Information (AMI) function and the False Nearest Neighbor (FNN) function. In some contexts, however, such as phase-space reconstruction for Multidimensional Recurrence Quantification Analysis (MdRQA), the empirical time series that need to be embedded already possess a dimensionality higher than one. In the current article, we present extensions of the AMI and FNN functions for higher dimensional time-series and their application to data from the Lorenz system coded in Matlab.

Significance Previous work in humans has found rhythmic cortical activity while listening to rhythmic sounds such as speech or music. Whether this activity reflects oscillatory dynamics of a neural circuit or instead evoked responses to the rhythmic stimulus has been difficult to determine. Here, we devised a metric to tease apart the two hypotheses by analyzing phase lag across many stimulation rates. We test this phase concentration metric using numerical simulations and generate quantitative predictions to compare against recorded magnetoencephalography data. Both previously recorded and new data were better predicted by a model of oscillatory dynamics than evoked responses. This work, therefore, provides definitive evidence for the presence of an oscillatory dynamic in auditory cortex during processing of rhythmic stimuli.

Transcranial electrical stimulation has widespread clinical and research applications, yet its effect on ongoing neural activity in humans is not well established. Previous reports argue that transcranial alternating current stimulation (tACS) can entrain and enhance neural rhythms related to memory, but the evidence from non-invasive recordings has remained inconclusive. Here, we measure endogenous spindle and theta activity intracranially in humans during low-frequency tACS and find no stable entrainment of spindle power during non-REM sleep, nor of theta power during resting wakefulness. As positive controls, we find robust entrainment of spindle activity to endogenous slow-wave activity in 66% of electrodes as well as entrainment to rhythmic noise-burst acoustic stimulation in 14% of electrodes. We conclude that low-frequency tACS at common stimulation intensities neither acutely modulates spindle activity during sleep nor theta activity during waking rest, likely because of the attenuated electrical fields reaching the cortical surface.

Championing open science, an adversarial collaboration aims to unravel the footprints of consciousness

Most humans have a near-automatic inclination to tap, clap, or move to the beat of music. The capacity to extract a periodic beat from a complex musical segment is remarkable, as it requires abstraction from the temporal structure of the stimulus. It has been suggested that nonlinear interactions in neural networks result in cortical oscillations at the beat frequency, and that such entrained oscillations give rise to the percept of a beat or a pulse. Here we tested this neural resonance theory using MEG recordings as female and male individuals listened to 30 s sequences of complex syncopated drumbeats designed so that they contain no net energy at the pulse frequency when measured using linear analysis. We analyzed the spectrum of the neural activity while listening and compared it to the modulation spectrum of the stimuli. We found enhanced neural response in the auditory cortex at the pulse frequency. We also showed phase locking at the times of the missing pulse, even though the pulse was absent from the stimulus itself. Moreover, the strength of this pulse response correlated with individuals' speed in finding the pulse of these stimuli, as tested in a follow-up session. These findings demonstrate that neural activity at the pulse frequency in the auditory cortex is internally generated rather than stimulus-driven. The current results are both consistent with neural resonance theory and with models based on nonlinear response of the brain to rhythmic stimuli. The results thus help narrow the search for valid models of beat perception.