High-fidelity transmission of auditory symbolic material is associated with reduced right-left neuroanatomical asymmetry between primary auditory regions.

The intergenerational stability of auditory symbolic systems, such as music, is thought to rely on brain processes that allow the faithful transmission of complex sounds. Little is known about the functional and structural aspects of the human brain which support this ability, with a few studies pointing to the bilateral organization of auditory networks as a putative neural substrate. Here, we further tested this hypothesis by examining the role of left-right neuroanatomical asymmetries between auditory cortices. We collected neuroanatomical images from a large sample of participants (nonmusicians) and analyzed them with Freesurfer's surface-based morphometry method. Weeks after scanning, the same individuals participated in a laboratory experiment that simulated music transmission: the signaling games. We found that high accuracy in the intergenerational transmission of an artificial tone system was associated with reduced rightward asymmetry of cortical thickness in Heschl's sulcus. Our study suggests that the high-fidelity copying of melodic material may rely on the extent to which computational neuronal resources are distributed across hemispheres. Our data further support the role of interhemispheric brain organization in the cultural transmission and evolution of auditory symbolic systems.

The early adolescent brain on music: Analysis of functional dynamics reveals engagement of orbitofrontal cortex reward system.

Music listening plays a pivotal role for children and adolescents, yet it remains unclear how music modulates brain activity at the level of functional networks in this young population. Analysing the dynamics of brain networks occurring and dissolving over time in response to music can provide a better understanding of the neural underpinning of music listening. We collected functional magnetic resonance imaging (fMRI) data from 17 preadolescents aged 10-11 years while listening to two similar music pieces separated by periods without music. We subsequently tracked the occurrence of functional brain networks over the recording time using a recent method that detects recurrent patterns of phase-locking in the fMRI signals: the leading eigenvector dynamics analysis (LEiDA). The probabilities of occurrence and switching profiles of different functional networks were compared between periods of music and no music. Our results showed significantly increased occurrence of a specific functional network during the two music pieces compared to no music, involving the medial orbitofrontal and ventromedial prefrontal cortices-a brain subsystem associated to reward processing. Moreover, the higher the musical reward sensitivity of the preadolescents, the more this network was preceded by a pattern involving the insula. Our findings highlight the involvement of a brain subsystem associated with hedonic and emotional processing during music listening in the early adolescent brain. These results offer novel insight into the neural underpinnings of musical reward in early adolescence, improving our understanding of the important role and the potential benefits of music at this delicate age.

Enhanced mismatch negativity in harmonic compared with inharmonic sounds.

Many natural sounds have frequency spectra composed of integer multiples of a fundamental frequency. This property, known as harmonicity, plays an important role in auditory information processing. However, the extent to which harmonicity influences the processing of sound features beyond pitch is still unclear. This is interesting because harmonic sounds have lower information entropy than inharmonic sounds. According to predictive processing accounts of perception, this property could produce more salient neural responses due to the brain's weighting of sensory signals according to their uncertainty. In the present study, we used electroencephalography to investigate brain responses to harmonic and inharmonic sounds commonly occurring in music: Piano tones and hi-hat cymbal sounds. In a multifeature oddball paradigm, we measured mismatch negativity (MMN) and P3a responses to timbre, intensity, and location deviants in listeners with and without congenital amusia-an impairment of pitch processing. As hypothesized, we observed larger amplitudes and earlier latencies (for both MMN and P3a) in harmonic compared with inharmonic sounds. These harmonicity effects were modulated by sound feature. Moreover, the difference in P3a latency between harmonic and inharmonic sounds was larger for controls than amusics. We propose an explanation of these results based on predictive coding and discuss the relationship between harmonicity, information entropy, and precision weighting of prediction errors.

Musicianship and melodic predictability enhance neural gain in auditory cortex during pitch deviance detection.

When listening to music, pitch deviations are more salient and elicit stronger prediction error responses when the melodic context is predictable and when the listener is a musician. Yet, the neuronal dynamics and changes in connectivity underlying such effects remain unclear. Here, we employed dynamic causal modeling (DCM) to investigate whether the magnetic mismatch negativity response (MMNm)-and its modulation by context predictability and musical expertise-are associated with enhanced neural gain of auditory areas, as a plausible mechanism for encoding precision-weighted prediction errors. Using Bayesian model comparison, we asked whether models with intrinsic connections within primary auditory cortex (A1) and superior temporal gyrus (STG)-typically related to gain control-or extrinsic connections between A1 and STG-typically related to propagation of prediction and error signals-better explained magnetoencephalography responses. We found that, compared to regular sounds, out-of-tune pitch deviations were associated with lower intrinsic (inhibitory) connectivity in A1 and STG, and lower backward (inhibitory) connectivity from STG to A1, consistent with disinhibition and enhanced neural gain in these auditory areas. More predictable melodies were associated with disinhibition in right A1, while musicianship was associated with disinhibition in left A1 and reduced connectivity from STG to left A1. These results indicate that musicianship and melodic predictability, as well as pitch deviations themselves, enhance neural gain in auditory cortex during deviance detection. Our findings are consistent with predictive processing theories suggesting that precise and informative error signals are selected by the brain for subsequent hierarchical processing.

Signaling games and music as a credible signal.

The argument by Mehr et al. that music emerged and evolved culturally as a credible signal is convincing, but it lacks one essential ingredient: a model of signaling behavior that supports the main hypothesis theoretically and empirically. We argue that signaling games can help us explain how musical structures emerge as population-level phenomena, through sender-receiver signaling interactions.

The chronnectome of musical beat.

Keeping time is fundamental for our everyday existence. Various isochronous activities, such as locomotion, require us to use internal timekeeping. This phenomenon comes into play also in other human pursuits such as dance and music. When listening to music, we spontaneously perceive and predict its beat. The process of beat perception comprises both beat inference and beat maintenance, their relative importance depending on the salience of beat in the music. To study functional connectivity associated with these processes in a naturalistic situation, we used functional magnetic resonance imaging to measure brain responses of participants while they were listening to a piece of music containing strong contrasts in beat salience. Subsequently, we utilized dynamic graph analysis and psychophysiological interactions (PPI) analysis in connection with computational modelling of beat salience to investigate how functional connectivity manifests these processes. As the main effect, correlation analyses between the obtained dynamic graph measures and the beat salience measure revealed increased centrality in auditory-motor cortices, cerebellum, and extrastriate visual areas during low beat salience, whereas regions of the default mode- and central executive networks displayed high centrality during high beat salience. PPI analyses revealed partial dissociation of functional networks belonging to this pathway indicating complementary neural mechanisms crucial in beat inference and maintenance, processes pivotal for extracting and predicting temporal regularities in our environment.

Musical prediction error responses similarly reduced by predictive uncertainty in musicians and non-musicians.

Auditory prediction error responses elicited by surprising sounds can be reliably recorded with musical stimuli that are more complex and realistic than those typically employed in EEG or MEG oddball paradigms. However, these responses are reduced as the predictive uncertainty of the stimuli increases. In this study, we investigate whether this effect is modulated by musical expertise. Magnetic mismatch negativity (MMNm) responses were recorded from 26 musicians and 24 non-musicians while they listened to low- and high-uncertainty melodic sequences in a musical multi-feature paradigm that included pitch, slide, intensity and timbre deviants. When compared to non-musicians, musically trained participants had significantly larger pitch and slide MMNm responses. However, both groups showed comparable reductions in pitch and slide MMNm amplitudes in the high-uncertainty condition compared with the low-uncertainty condition. In a separate, behavioural deviance detection experiment, musicians were more accurate and confident about their responses than non-musicians, but deviance detection in both groups was similarly affected by the uncertainty of the melodies. In both experiments, the interaction between uncertainty and expertise was not significant, suggesting that the effect is comparable in both groups. Consequently, our results replicate the modulatory effect of predictive uncertainty on prediction error; show that it is present across different types of listeners; and suggest that expertise-related and stimulus-driven modulations of predictive precision are dissociable and independent.

Weighting of neural prediction error by rhythmic complexity: A predictive coding account using mismatch negativity.

The human brain's ability to extract and encode temporal regularities and to predict the timing of upcoming events is critical for music and speech perception. This work addresses how these mechanisms deal with different levels of temporal complexity, here the number of distinct durations in rhythmic patterns. We use electroencephalography (EEG) to relate the mismatch negativity (MMN), a proxy of neural prediction error, to a measure of information content of rhythmic sequences, the Shannon entropy. Within each of three conditions, participants listened to repeatedly presented standard rhythms of five tones (four inter-onset intervals) and of a given level of entropy: zero (isochronous), medium entropy (two distinct interval durations), or high entropy (four distinct interval durations). Occasionally, the fourth tone was moved forward in time that is it occurred 100 ms (small deviation) or 300 ms early (large deviation). According to the predictive coding framework, high-entropy stimuli are more difficult to model for the brain, resulting in less confident predictions and yielding smaller prediction errors for deviant sounds. Our results support this hypothesis, showing a gradual decrease in MMN amplitude as a function of entropy, but only for small timing deviants. For large timing deviants, in contrast, a modulation of activity in the opposite direction was observed for the earlier N1 component, known to also be sensitive to sudden changes in directed attention. Our results suggest the existence of a fine-grained neural mechanism that weights neural prediction error to the complexity of rhythms and that mostly manifests in the absence of directed attention.

Neuroanatomical substrate of noise sensitivity.

Recent functional studies suggest that noise sensitivity, a trait describing attitudes towards noise and predicting noise annoyance, is associated with altered processing in the central auditory system. In the present work, we examined whether noise sensitivity could be related to the structural anatomy of auditory and limbic brain areas. Anatomical MR brain images of 80 subjects were parcellated with FreeSurfer to measure grey matter volume, cortical thickness, cortical area and folding index of anatomical structures in the temporal lobe and insular cortex. The grey matter volume of amygdala and hippocampus was measured as well. According to our findings, noise sensitivity is associated with the grey matter volume in the selected structures. Among those, we propose and discuss particular areas, previously linked to auditory perceptual, emotional and interoceptive processing, in which larger grey matter volume seems to be related to higher noise sensitivity.

Connectivity patterns during music listening: Evidence for action-based processing in musicians.

Musical expertise is visible both in the morphology and functionality of the brain. Recent research indicates that functional integration between multi-sensory, somato-motor, default-mode (DMN), and salience (SN) networks of the brain differentiates musicians from non-musicians during resting state. Here, we aimed at determining whether brain networks differentially exchange information in musicians as opposed to non-musicians during naturalistic music listening. Whole-brain graph-theory analyses were performed on participants' fMRI responses. Group-level differences revealed that musicians' primary hubs comprised cerebral and cerebellar sensorimotor regions whereas non-musicians' dominant hubs encompassed DMN-related regions. Community structure analyses of the key hubs revealed greater integration of motor and somatosensory homunculi representing the upper limbs and torso in musicians. Furthermore, musicians who started training at an earlier age exhibited greater centrality in the auditory cortex, and areas related to top-down processes, attention, emotion, somatosensory processing, and non-verbal processing of speech. We here reveal how brain networks organize themselves in a naturalistic music listening situation wherein musicians automatically engage neural networks that are action-based while non-musicians use those that are perception-based to process an incoming auditory stream. Hum Brain Mapp 38:2955-2970, 2017. © 2017 Wiley Periodicals, Inc.

The urge to judge: Why the judgmental attitude has anything to do with the aesthetic enjoyment of negative emotions.

Based on arguments from both philosophical and empirical aesthetics, we hereby propose that the enjoyment of negative emotions in art and fiction is distinct from the immediate pleasure deriving from sensory features, because it requires a conscious, intentional attitude toward the object. This attitude is linked with the compelling goal of providing a judgment of liking, beauty, perfection, or similar.

The reliability of continuous brain responses during naturalistic listening to music.

Low-level (timbral) and high-level (tonal and rhythmical) musical features during continuous listening to music, studied by functional magnetic resonance imaging (fMRI), have been shown to elicit large-scale responses in cognitive, motor, and limbic brain networks. Using a similar methodological approach and a similar group of participants, we aimed to study the replicability of previous findings. Participants' fMRI responses during continuous listening of a tango Nuevo piece were correlated voxelwise against the time series of a set of perceptually validated musical features computationally extracted from the music. The replicability of previous results and the present study was assessed by two approaches: (a) correlating the respective activation maps, and (b) computing the overlap of active voxels between datasets at variable levels of ranked significance. Activity elicited by timbral features was better replicable than activity elicited by tonal and rhythmical ones. These results indicate more reliable processing mechanisms for low-level musical features as compared to more high-level features. The processing of such high-level features is probably more sensitive to the state and traits of the listeners, as well as of their background in music.

The association of noise sensitivity with music listening, training, and aptitude.

After intensive, long-term musical training, the auditory system of a musician is specifically tuned to perceive musical sounds. We wished to find out whether a musician's auditory system also develops increased sensitivity to any sound of everyday life, experiencing them as noise. For this purpose, an online survey, including questionnaires on noise sensitivity, musical background, and listening tests for assessing musical aptitude, was administered to 197 participants in Finland and Italy. Subjective noise sensitivity (assessed with the Weinstein's Noise Sensitivity Scale) was analyzed for associations with musicianship, musical aptitude, weekly time spent listening to music, and the importance of music in each person's life (or music importance). Subjects were divided into three groups according to their musical expertise: Nonmusicians (N = 103), amateur musicians (N = 44), and professional musicians (N = 50). The results showed that noise sensitivity did not depend on musical expertise or performance on musicality tests or the amount of active (attentive) listening to music. In contrast, it was associated with daily passive listening to music, so that individuals with higher noise sensitivity spent less time in passive (background) listening to music than those with lower sensitivity to noise. Furthermore, noise-sensitive respondents rated music as less important in their life than did individuals with lower sensitivity to noise. The results demonstrate that the special sensitivity of the auditory system derived from musical training does not lead to increased irritability from unwanted sounds. However, the disposition to tolerate contingent musical backgrounds in everyday life depends on the individual's noise sensitivity.

Capturing the musical brain with Lasso: Dynamic decoding of musical features from fMRI data.

We investigated neural correlates of musical feature processing with a decoding approach. To this end, we used a method that combines computational extraction of musical features with regularized multiple regression (LASSO). Optimal model parameters were determined by maximizing the decoding accuracy using a leave-one-out cross-validation scheme. The method was applied to functional magnetic resonance imaging (fMRI) data that were collected using a naturalistic paradigm, in which participants' brain responses were recorded while they were continuously listening to pieces of real music. The dependent variables comprised musical feature time series that were computationally extracted from the stimulus. We expected timbral features to obtain a higher prediction accuracy than rhythmic and tonal ones. Moreover, we expected the areas significantly contributing to the decoding models to be consistent with areas of significant activation observed in previous research using a naturalistic paradigm with fMRI. Of the six musical features considered, five could be significantly predicted for the majority of participants. The areas significantly contributing to the optimal decoding models agreed to a great extent with results obtained in previous studies. In particular, areas in the superior temporal gyrus, Heschl's gyrus, Rolandic operculum, and cerebellum contributed to the decoding of timbral features. For the decoding of the rhythmic feature, we found the bilateral superior temporal gyrus, right Heschl's gyrus, and hippocampus to contribute most. The tonal feature, however, could not be significantly predicted, suggesting a higher inter-participant variability in its neural processing. A subsequent classification experiment revealed that segments of the stimulus could be classified from the fMRI data with significant accuracy. The present findings provide compelling evidence for the involvement of the auditory cortex, the cerebellum and the hippocampus in the processing of musical features during continuous listening to music.

Making sense of music: Insights from neurophysiology and connectivity analyses in naturalistic listening conditions.

According to the latest frameworks, auditory perception and memory involve the constant prediction of future sound events by the brain, based on the continuous extraction of feature regularities from the environment. The neural hierarchical mechanisms for predictive processes in perception and memory for sounds are typically studied in relation to simple acoustic features in isolated sounds or sound patterns inserted in highly certain contexts. Such studies have identified reliable prediction formation and error signals, e.g., the N100 or the mismatch negativity (MMN) evoked responses. In real life, though, individuals often face situations in which uncertainty prevails and where making sense of sounds becomes a hard challenge. In music, not only deviations from predictions are masterly set up by composers to induce emotions but sometimes the sheer uncertainty of sound scenes is exploited for aesthetic purposes, especially in compositional styles such as Western atonal classical music. In very recent magnetoencephalography (MEG) and electroencephalography (EEG) studies, experimental and technical advances in stimulation paradigms and analysis approaches have permitted the identification of prediction-error responses from highly uncertain, atonal contexts and the extraction of prediction-related responses from real, continuous music. Moreover, functional connectivity analyses revealed the emergence of cortico-hippocampal interactions during the formation of auditory memories for more predictable vs. less predictable patterns. These findings contribute to understanding the general brain mechanisms that enable us to predict even highly uncertain sound environments and to possibly make sense of and appreciate even atonal music.

Aesthetics and predictive processing: grounds and prospects of a fruitful encounter.

In the last few years, a remarkable convergence of interests and results has emerged between scholars interested in the arts and aesthetics from a variety of perspectives and cognitive scientists studying the mind and brain within the predictive processing (PP) framework. This convergence has so far proven fruitful for both sides: while PP is increasingly adopted as a framework for understanding aesthetic phenomena, the arts and aesthetics, examined under the lens of PP, are starting to be seen as important windows into our mental functioning. The result is a vast and fast-growing research programme that promises to deliver important insights into our aesthetic encounters as well as a wide range of psychological phenomena of general interest. Here, we present this developing research programme, describing its grounds and highlighting its prospects. We start by clarifying how the study of the arts and aesthetics encounters the PP picture of mental functioning (§1). We then go on to outline the prospects of this encounter for the fields involved: philosophy and history of art (§2), psychology of aesthetics and neuroaesthetics (§3) and psychology and neuroscience more generally (§4). The upshot is an ambitious but well-defined framework within which aesthetics and cognitive science can partner up to illuminate crucial aspects of the human mind. This article is part of the theme issue 'Art, aesthetics and predictive processing: theoretical and empirical perspectives'.

The impact of crossmodal predictions on the neural processing of aesthetic stimuli.

Neuroaesthetic research has focused on neural predictive processes involved in the encounter with art stimuli or the related evaluative judgements, and it has been mainly conducted unimodally. Here, with electroencephalography, magnetoencephalography and an affective priming protocol, we investigated whether and how the neural responses to non-representational aesthetic stimuli are top-down modulated by affective representational (i.e. semantically meaningful) predictions between audition and vision. Also, the neural chronometry of affect processing of these aesthetic stimuli was investigated. We hypothesized that the early affective components of crossmodal aesthetic responses are dependent on the affective and representational predictions formed in another sensory modality resulting in differentiated brain responses, and that audition and vision indicate different processing latencies for affect. The target stimuli were aesthetic visual patterns and musical chords, and they were preceded by a prime from the opposing sensory modality. We found that early auditory-cortex responses to chords were more affected by valence than the corresponding visual-cortex ones. Furthermore, the assessments of visual targets were more facilitated by affective congruency of crossmodal primes than the acoustic targets. These results indicate, first, that the brain uses early affective information for predictively guiding aesthetic responses; second, that an affective transfer of information takes place crossmodally, mainly from audition to vision, impacting the aesthetic assessment. This article is part of the theme issue 'Art, aesthetics and predictive processing: theoretical and empirical perspectives'.

Musical and electrical stimulation as intervention in disorder of consciousness (DOC) patients: A randomised cross-over trial.

BACKGROUND: Disorders of consciousness (DOC), i.e., unresponsive wakefulness syndrome (UWS) or vegetative state (VS) and minimally conscious state (MCS), are conditions that can arise from severe brain injury, inducing widespread functional changes. Given the damaging implications resulting from these conditions, there is an increasing need for rehabilitation treatments aimed at enhancing the level of consciousness, the quality of life, and creating new recovery perspectives for the patients. Music may represent an additional rehabilitative tool in contexts where cognition and language are severely compromised, such as among DOC patients. A further type of rehabilitation strategies for DOC patients consists of Non-Invasive Brain Stimulation techniques (NIBS), including transcranial electrical stimulation (tES), affecting neural excitability and promoting brain plasticity. OBJECTIVE: We here propose a novel rehabilitation protocol for DOC patients that combines music-based intervention and NIBS in neurological patients. The main objectives are (i) to assess the residual neuroplastic processes in DOC patients exposed to music, (ii) to determine the putative neural modulation and the clinical outcome in DOC patients of non-pharmacological strategies, i.e., tES(control condition), and music stimulation, and (iii) to evaluate the putative positive impact of this intervention on caregiver's burden and psychological distress. METHODS: This is a randomised cross-over trial in which a total of 30 participants will be randomly allocated to one of three different combinations of conditions: (i) Music only, (ii) tES only (control condition), (iii) Music + tES. The music intervention will consist of listening to an individually tailored playlist including familiar and self-relevant music together with fixed songs; concerning NIBS, tES will be applied for 20 minutes every day, 5 times a week, for two weeks. After these stimulations two weeks of placebo treatments will follow, with sham stimulation combined with noise for other two weeks. The primary outcomes will be clinical, i.e., based on the differences in the scores obtained on the neuropsychological tests, such as Coma Recovery Scale-Revised, and neurophysiological measures as EEG, collected pre-intervention, post-intervention and post-placebo. DISCUSSION: This study proposes a novel rehabilitation protocol for patients with DOC including a combined intervention of music and NIBS. Considering the need for rigorous longitudinal randomised controlled trials for people with severe brain injury disease, the results of this study will be highly informative for highlighting and implementing the putative beneficial role of music and NIBS in rehabilitation treatments. TRIAL REGISTRATION: ClinicalTrials.gov identifier: NCT05706831, registered on January 30, 2023.

From Vivaldi to Beatles and back: predicting lateralized brain responses to music.

We aimed at predicting the temporal evolution of brain activity in naturalistic music listening conditions using a combination of neuroimaging and acoustic feature extraction. Participants were scanned using functional Magnetic Resonance Imaging (fMRI) while listening to two musical medleys, including pieces from various genres with and without lyrics. Regression models were built to predict voxel-wise brain activations which were then tested in a cross-validation setting in order to evaluate the robustness of the hence created models across stimuli. To further assess the generalizability of the models we extended the cross-validation procedure by including another dataset, which comprised continuous fMRI responses of musically trained participants to an Argentinean tango. Individual models for the two musical medleys revealed that activations in several areas in the brain belonging to the auditory, limbic, and motor regions could be predicted. Notably, activations in the medial orbitofrontal region and the anterior cingulate cortex, relevant for self-referential appraisal and aesthetic judgments, could be predicted successfully. Cross-validation across musical stimuli and participant pools helped identify a region of the right superior temporal gyrus, encompassing the planum polare and the Heschl's gyrus, as the core structure that processed complex acoustic features of musical pieces from various genres, with or without lyrics. Models based on purely instrumental music were able to predict activation in the bilateral auditory cortices, parietal, somatosensory, and left hemispheric primary and supplementary motor areas. The presence of lyrics on the other hand weakened the prediction of activations in the left superior temporal gyrus. Our results suggest spontaneous emotion-related processing during naturalistic listening to music and provide supportive evidence for the hemispheric specialization for categorical sounds with realistic stimuli. We herewith introduce a powerful means to predict brain responses to music, speech, or soundscapes across a large variety of contexts.

Age differences in central auditory system responses to naturalistic music.

Aging influences the central auditory system leading to difficulties in the decoding and understanding of overlapping sound signals, such as speech in noise or polyphonic music. Studies on central auditory system evoked responses (ERs) have found in older compared to young listeners increased amplitudes (less inhibition) of the P1 and N1 and decreased amplitudes of the P2, mismatch negativity (MMN), and P3a responses. While preceding research has focused on simplified auditory stimuli, we here tested whether the previously observed age-related differences could be replicated with sounds embedded in medium and highly naturalistic musical contexts. Older (age 55-77 years) and younger adults (age 21-31 years) listened to medium naturalistic (synthesized melody) and highly naturalistic (studio recording of a music piece) stimuli. For the medium naturalistic music, the age group differences on the P1, N1, P2, MMN, and P3a amplitudes were all replicated. The age group differences, however, appeared reduced with the highly compared to the medium naturalistic music. The finding of lower P2 amplitude in older than young was replicated for slow event rates (0.3-2.9 Hz) in the highly naturalistic music. Moreover, the ER latencies suggested a gradual slowing of the auditory processing time course for highly compared to medium naturalistic stimuli irrespective of age. These results support that age-related differences on ERs can partly be observed with naturalistic stimuli. This opens new avenues for including naturalistic stimuli in the investigation of age-related central auditory system disorders.