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Abstract Aging is often associated with decline in brain processing power and neural predictive capabilities. To challenge this notion, we used magnetoencephalography (MEG) and magnetic resonance imaging (MRI) to record the whole-brain activity of 39 older adults (over 60 years old) and 37 young adults (aged 18–25 years) during recognition of previously memorised and varied musical sequences. Results reveal that when recognising memorised sequences, the brain of older compared to young adults reshapes its functional organisation. In fact, it shows increased early activity in sensory regions such as the left auditory cortex (100 ms and 250 ms after each note), and only moderate decreased activity (350 ms) in medial temporal lobe and prefrontal regions. When processing the varied sequences, older adults show a marked reduction of the fast-scale functionality (250 ms after each note) of higher-order brain regions including hippocampus, ventromedial prefrontal and inferior temporal cortices, while no differences are observed in the auditory cortex. Accordingly, young outperform older adults in the recognition of novel sequences, while no behavioural differences are observed with regards to memorised ones. Our findings show age-related neural changes in predictive and memory processes, integrating existing theories on compensatory neural mechanisms in non-pathological aging.

BackgroundDisorders 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.ObjectiveWe 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.MethodsThis 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.DiscussionThis 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 registrationClinicalTrials.gov identifier: NCT05706831, registered on January 30, 2023.

A magnetoencephalography study reveals that tonal musical sequences recruit neural activity in the hippocampus and anterior cingulate cortex, while atonal sequences activate auditory regions, suggesting the involvement of a cortico-subcortical brain network in auditory recognition.

Abstract Through long-term training, music experts acquire complex and specialized sensorimotor skills, which are paralleled by continuous neuro-anatomical and -functional adaptations. The underlying neuroplasticity mechanisms have been extensively explored in decades of research in music, cognitive, and translational neuroscience. However, the absence of a comprehensive review and quantitative meta-analysis prevents the plethora of variegated findings to ultimately converge into a unified picture of the neuroanatomy of musical expertise. Here, we performed a comprehensive neuroimaging meta-analysis of publications investigating neuro-anatomical and -functional differences between musicians (M) and non-musicians (NM). Eighty-four studies were included in the qualitative synthesis. From these, 58 publications were included in coordinate-based meta-analyses using the anatomic/activation likelihood estimation (ALE) method. This comprehensive approach delivers a coherent cortico-subcortical network encompassing sensorimotor and limbic regions bilaterally. Particularly, M exhibited higher volume/activity in auditory, sensorimotor, interoceptive, and limbic brain areas and lower volume/activity in parietal areas as opposed to NM. Notably, we reveal topographical (dis-)similarities between the identified functional and anatomical networks and characterize their link to various cognitive functions by means of meta-analytic connectivity modelling. Overall, we effectively synthesized decades of research in the field and provide a consistent and controversies-free picture of the neuroanatomy of musical expertise.

Music training, in all its forms, is known to have an impact on behavior both in childhood and even in aging. In the delicate life period of transition from childhood to adulthood, music training might have a special role for behavioral and cognitive maturation. Among the several kinds of music training programs implemented in the educational communities, we focused on instrumental training incorporated in the public middle school curriculum in Italy that includes both individual, group and collective (orchestral) lessons several times a week. At three middle schools, we tested 285 preadolescent children (aged 10–14 years) with a test and questionnaire battery including adaptive tests for visuo-spatial working memory skills (with the Jack and Jill test), fluid intelligence (with a matrix reasoning test) and music-related perceptual and memory abilities (with listening tests). Of these children, 163 belonged to a music curriculum within the school and 122 to a standard curriculum. Significant differences between students of the music and standard curricula were found in both perceptual and cognitive domains, even when controlling for pre-existing individual differences in musical sophistication. The music children attending the third and last grade of middle school had better performance and showed the largest advantage compared to the control group on both audiovisual working memory and fluid intelligence. Furthermore, some gender differences were found for several tests and across groups in favor of females. The present results indicate that learning to play a musical instrument as part of the middle school curriculum represents a resource for preadolescent education. Even though the current evidence is not sufficient to establish the causality of the found effects, it can still guide future research evaluation with longitudinal data.

Summary: Neuronal oscillations, their inter-areal synchronization, and scale-free dynamics constitute fundamental mechanisms for cognition by regulating communication in neuronal networks. These oscillatory dynamics have large inter-individual variability that is partly heritable. We hypothesized that this variability could be partially explained by genetic polymorphisms in neuromodulatory genes. We recorded resting-state magnetoencephalography (MEG) from 82 healthy participants and investigated whether oscillation dynamics were influenced by genetic polymorphisms in catechol-O-methyltransferase (COMT) Val158Met and brain-derived neurotrophic factor (BDNF) Val66Met. Both COMT and BDNF polymorphisms influenced local oscillation amplitudes and their long-range temporal correlations (LRTCs), while only BDNF polymorphism affected the strength of large-scale synchronization. Our findings demonstrate that COMT and BDNF genetic polymorphisms contribute to inter-individual variability in neuronal oscillation dynamics. Comparison of these results to computational modeling of near-critical synchronization dynamics further suggested that COMT and BDNF polymorphisms influenced local oscillations by modulating the excitation-inhibition balance according to the brain criticality framework.

Atonal music is often characterized by low predictability stemming from the absence of tonal or metrical hierarchies. In contrast, Western tonal music exhibits intrinsic predictability due to its hierarchical structure and therefore, offers a directly accessible predictive model to the listener. In consequence, a specific challenge of atonal music is that listeners must generate a variety of new predictive models. Listeners must not only refrain from applying available tonal models to the heard music, but they must also search for statistical regularities and build new rules that may be related to musical properties other than pitch, such as timbre or dynamics. In this article, we propose that the generation of such new predictive models and the aesthetic experience of atonal music are characterized by internal states related to exploration. This is a behavior well characterized in behavioral neuroscience as fulfilling an innate drive to reduce uncertainty but which has received little attention in empirical music research. We support our proposal with emerging evidence that the hedonic value is associated with the recognition of patterns in low-predictability sound sequences and that atonal music elicits distinct behavioral responses in listeners. We end by outlining new research avenues that might both deepen our understanding of the aesthetic experience of atonal music in particular, and reveal core qualities of the aesthetic experience in general.

Liking and pleasantness are common concepts in psychological emotion theories and in everyday language related to emotions. Despite obvious similarities between the terms, several empirical and theoretical notions support the idea that pleasantness and liking are cognitively different phenomena, becoming most evident in the context of emotion regulation and art enjoyment. In this study it was investigated whether liking and pleasantness indicate behaviourally measurable differences, not only in the long timespan of emotion regulation, but already within the initial affective responses to visual and auditory stimuli. A cross-modal affective priming protocol was used to assess whether there is a behavioural difference in the response time when providing an affective rating to a liking or pleasantness task. It was hypothesized that the pleasantness task would be faster as it is known to rely on rapid feature detection. Furthermore, an affective priming effect was expected to take place across the sensory modalities and the presentative and non-presentative stimuli. A linear mixed effect analysis indicated a significant priming effect as well as an interaction effect between the auditory and visual sensory modalities and the affective rating tasks of liking and pleasantness: While liking was rated fastest across modalities, it was significantly faster in vision compared to audition. No significant modality dependent differences between the pleasantness ratings were detected. The results demonstrate that liking and pleasantness rating scales refer to separate processes already within the short time scale of one to two seconds. Furthermore, the affective priming effect indicates that an affective information transfer takes place across modalities and the types of stimuli applied. Unlike hypothesized, liking rating took place faster across the modalities. This is interpreted to support emotion theoretical notions where liking and disliking are crucial properties of emotion perception and homeostatic self-referential information, possibly overriding pleasantness-related feature analysis. Conclusively, the findings provide empirical evidence for a conceptual delineation of common affective processes.

Hallucinations are a complex and multidimensional phenomenon which can differ based on the involved pathology, typology and sensory modality. Hallucinations are common in patients with neurodegenerative diseases. Recent sparse evidence from resting state functional magnetic resonance imaging (rs-fMRI) studies has identified altered functional connectivity in those patients within several brain networks, such as the default mode, attentional and sensory ones, without, however, providing an organized picture of the mechanisms involved. This systematic review, following PRISMA guidelines, aims at critically analyzing the current literature on the brain networks associated with the phenomenon of hallucinations in patients with neurodegenerative diseases. Ten rs-fMRI studies fulfilled our selection criteria. All these studies focused on synucleinopathies, and most of them focused on visual hallucinations and were characterized by a heterogeneous methodology. Thus, instead of offering a definite picture of the mechanisms underlying hallucinations in neurodegeneration, this systematic review encourages further research especially concerning tauopathies. Notwithstanding, the findings overall suggest a disruption in the top-down (associated with memory intrusion and difficulty of inhibition) and in the bottom-up processes (associated with the sensory areas involved in the hallucinations). Further investigations are needed in order to disentangle the brain mechanisms involved in hallucinations and to overcome possible limitations characterizing the current literature.

Frontotemporal dementia (FTD) is a rare neurodegenerative disease, characterized by behavioral and language impairments. Primary progressive aphasia (PPA) is the linguistic variant of this heterogeneous disorder. To date, there is a lack of consensus about which interventions are effective in these patients. However, several studies show that music-based interventions are beneficial in neurological diseases. This study aims, primarily, to establish the state of the art of music-based interventions designed for PPA due to FTD and, secondarily, to inform the planning of PPA-dedicated future interventions for Italian neurological institutions. The first aim is fulfilled by a review which critically screens the neurological studies examining the effects of music- and/or rhythm-based interventions, especially, on language rehabilitation in aphasic FTD. We found that only two papers fulfilled our criteria and concerned specifically aphasic patients due to FTD. Of those, one paper reported a study conducted in an Italian institution. Most of the reviewed studies focused, instead, on aphasia in post-stroke patients. The results of our review invite further studies to investigate the role of music as a valuable support in the therapy for neurodegenerative patients with language problems and in particular to PPA due to FTD. Moreover, based on this initial work, we can delineate new music-based interventions dedicated to PPA for Italian institutions.

The properties of functional brain networks strongly depend on how their nodes are chosen. Commonly, nodes are defined by Regions of Interest (ROIs), predetermined groupings of fMRI measurement voxels. Earlier, we demonstrated that the functional homogeneity of ROIs, captured by their spatial consistency, varies widely across ROIs in commonly used brain atlases. Here, we ask how ROIs behave as nodes of dynamic brain networks. To this end, we use two measures: spatiotemporal consistency measures changes in spatial consistency across time and network turnover quantifies the changes in the local network structure around an ROI. We find that spatial consistency varies non-uniformly in space and time, which is reflected in the variation of spatiotemporal consistency across ROIs. Furthermore, we see time-dependent changes in the network neighborhoods of the ROIs, reflected in high network turnover. Network turnover is nonuniformly distributed across ROIs: ROIs with high spatiotemporal consistency have low network turnover. Finally, we reveal that there is rich voxel-level correlation structure inside ROIs. Because the internal structure and the connectivity of ROIs vary in time, the common approach of using static node definitions may be surprisingly inaccurate. Therefore, network neuroscience would greatly benefit from node definition strategies tailored for dynamical networks. Regions of Interest (ROIs) are often used as the nodes of functional brain networks. ROIs consist of several fMRI measurement voxels that are assumed to be functionally homogeneous, that is, behave similarly. Earlier, we showed that the assumption of similar voxel dynamics is not always true: functional homogeneity varies widely across ROIs. In this paper, we demonstrate that functional homogeneity changes in time. These changes are connected to changes in local network structure around ROIs, which suggests that an ROI’s functional homogeneity may reflect its role in the network. Finally, we show that there is rich, time-dependent structure of voxel-level connectivity inside ROIs. This leads us to ask if the dynamic brain networks can be described by any set of static ROIs.

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.

Cochlear implants (CIs) allow good perception of speech while music listening is unsatisfactory, leading to reduced music enjoyment. Hence, a number of ongoing efforts aim to improve music perception with a CI. Regardless of the nature of these efforts, effect measurements must be valid and reliable. While auditory skills are typically examined by behavioral methods, recording of the mismatch negativity (MMN) response, using electroencephalography (EEG), has recently been applied successfully as a supplementary objective measure. Eleven adult CI users and 14 normally hearing (NH) controls took part in the present study. To measure their detailed discrimination of fundamental features of music we applied a new multifeature MMN-paradigm which presented four music deviants at four levels of magnitude, incorporating a novel “no-standard” approach to be tested with CI users for the first time. A supplementary test measured behavioral discrimination of the same deviants and levels. The MMN-paradigm elicited significant MMN responses to all levels of deviants in both groups. Furthermore, the CI-users’ MMN amplitudes and latencies were not significantly different from those of NH controls. Both groups showed MMN strength that was in overall alignment with the deviation magnitude. In CI users, however, discrimination of pitch levels remained undifferentiated. On average, CI users’ behavioral performance was significantly below that of the NH group, mainly due to poor pitch discrimination. Although no significant effects were found, CI users’ behavioral results tended to be in accordance with deviation magnitude, most prominently manifested in discrimination of the rhythm deviant. In summary, the study indicates that CI users may be able to discriminate subtle changes in basic musical features both in terms of automatic neural responses and of attended behavioral detection. Despite high complexity, the new CI MuMuFe paradigm and the “no-standard” approach provided reliable results, suggesting that it may serve as a relevant tool in future CI research. For clinical use, future studies should investigate the possibility of applying the paradigm with the purpose of assessing discrimination skills not only at the group level but also at the individual level.

Music producers, whether original composers or performers, vary in their ability to acquire and faithfully transmit music. This form of variation may serve as a mechanism for the emergence of new traits in musical systems. In this study, we aim to investigate whether individual differences in the social learning and transmission of music relate to intrinsic neural dynamics of auditory processing systems. We combined auditory and resting-state functional magnetic resonance imaging (fMRI) with an interactive laboratory model of cultural transmission, the signaling game, in an experiment with a large cohort of participants (N=51). We found that the degree of interhemispheric rs-FC within fronto-temporal auditory networks predicts—weeks after scanning—learning, transmission, and structural modification of an artificial tone system. Our study introduces neuroimaging in cultural transmission research and points to specific neural auditory processing mechanisms that constrain and drive variation in the cultural transmission and regularization of musical systems.

The last decades have seen a proliferation of music and brain studies, with a major focus on plastic changes as the outcome of continuous and prolonged engagement with music. Thanks to the advent of neuroaesthetics, research on music cognition has broadened its scope by considering the multifarious phenomenon of listening in all its forms, including incidental listening up to the skillful attentive listening of experts, and all its possible effects. These latter range from objective and sensorial effects directly linked to the acoustic features of the music to the subjectively affective and even transformational effects for the listener. Of special importance is the finding that neural activity in the reward circuit of the brain is a key component of a conscious listening experience. We propose that the connection between music and the reward system makes music listening a gate towards not only hedonia but also eudaimonia, namely a life well lived, full of meaning that aims at realizing one’s own “daimon” or true nature. It is argued, further, that music listening, even when conceptualized in this aesthetic and eudaimonic framework, remains a learnable skill that changes the way brain structures respond to sounds and how they interact with each other.

Brain responses are often studied under strictly experimental conditions in which electroencephalograms (EEGs) are recorded to reflect reactions to short and repetitive stimuli. However, in real life, aural stimuli are continuously mixed and cannot be found isolated, such as when listening to music. In this audio context, the acoustic features in music related to brightness, loudness, noise, and spectral flux, among others, change continuously; thus, significant values of these features can occur nearly simultaneously. Such situations are expected to give rise to increased brain reaction with respect to a case in which they would appear in isolation. In order to assert this, EEG signals recorded while listening to a tango piece were considered. The focus was on the amplitude and time of the negative deflation (N100) and positive deflation (P200) after the stimuli, which was defined on the basis of the selected music feature saliences, in order to perform a statistical analysis intended to test the initial hypothesis. Differences in brain reactions can be identified depending on the concurrence (or not) of such significant values of different features, proving that coterminous increments in several qualities of music influence and modulate the strength of brain responses.

This commentary discusses the paper by Schubert and colleagues in the context of the recent proposals in the field of empirical aesthetics and neuroaesthetics, emphasizing the need for inclusion of the time variable for accounting for the dynamic nature of a musical experience. At the same time, the efforts of the authors for systematizing the concepts in the field are praised.

Converging evidence has demonstrated that musical training is associated with improved perceptual and cognitive skills, including executive functions and general intelligence, particularly in childhood. In contrast, in adults the relationship between cognitive performance and musicianship is less clear and seems to be modulated by a number of background factors, such as personality and socio-economic status. Aiming to shed new light on this topic, we administered the Wechsler Adult Intelligence Scale III (WAIS-III), the Wechsler Memory Scale III (WMS-III), and the Stroop Test to 101 Finnish healthy adults grouped according to their musical expertise (non-musicians, amateurs, and musicians). After being matched for socio-economic status, personality traits and other demographic variables, adult musicians exhibited higher cognitive performance than non-musicians in all the mentioned measures. Moreover, linear regression models showed significant positive relationships between executive functions (working memory and attention) and the duration of musical practice, even after controlling for intelligence and background variables, such as personality traits. Hence, our study offers further support for the association between cognitive abilities and musical training, even in adulthood.HIGHLIGHTS- Musicians show higher general intelligence (FSIQ), verbal intelligence (VIQ), working memory (WMI) and attention skills than non-musicians. Amateurs score in between.- Significant positive correlations between years of musical playing and cognitive abilities support the hypothesis that long-term musical practice is associated with intelligence and executive functions.

Survey studies have shown that participating in music groups produces several benefits, such as discipline, cooperation and responsibility. Accordingly, recent longitudinal studies showed that orchestral music training has a positive impact on inhibitory control in school-age children. However, most of these studies examined long periods of training not always feasible for all families and institutions and focused on children’s measures ignoring the viewpoint of the teachers. Considering the crucial role of inhibitory control on hyperactivity, inattention and impulsivity, we wanted to explore if short orchestral music training would promote a reduction of these impulsive behaviors in children. This study involved 113 Italian children from 8 to 10 years of age. 55 of them attended 3 months of orchestral music training. The training included a 2-hour lesson per week at school and a final concert. The 58 children in the control group did not have any orchestral music training. All children were administered tests and questionnaires measuring inhibitory control and hyperactivity near the beginning and end of the 3-month training period. We also collected information regarding the levels of hyperactivity of the children as perceived by the teachers at both time points. Children in the music group showed a significant improvement in inhibitory control. Moreover, in the second measurement the control group showed an increase in self-reported hyperactivity that was not found in the group undergoing the music training program. This change was not noticed by the teachers, implying a discrepancy between self-reported and observed behavior at school. Our results suggest that even an intense and brief period of orchestral music training is sufficient to facilitate the development of inhibitory control by modulating the levels of self-reported hyperactivity. This research has implications for music pedagogy and education especially in children with high hyperactivity. Future investigations will test whether the findings can be extended to children diagnosed with ADHD.

Learning, attention and action play a crucial role in determining how stimulus predictions are formed, stored, and updated. Years-long experience with the specific repertoires of sounds of one or more musical styles is what characterizes professional musicians. Here we contrasted active experience with sounds, namely long-lasting motor practice, theoretical study and engaged listening to the acoustic features characterizing a musical style of choice in professional musicians with mainly passive experience of sounds in laypersons. We hypothesized that long-term active experience of sounds would influence the neural predictions of the stylistic features in professional musicians in a distinct way from the mainly passive experience of sounds in laypersons. Participants with different musical backgrounds were recruited: professional jazz and classical musicians, amateur musicians and non-musicians. They were presented with a musical multi-feature paradigm eliciting mismatch negativity (MMN), a prediction error signal to changes in six sound features for only 12 minutes of electroencephalography (EEG) and magnetoencephalography (MEG) recordings. We observed a generally larger MMN amplitudes-indicative of stronger automatic neural signals to violated priors-in jazz musicians (but not in classical musicians) as compared to non-musicians and amateurs. The specific MMN enhancements were found for spectral features (timbre, pitch, slide) and sound intensity. In participants who were not musicians, the higher preference for jazz music was associated with reduced MMN to pitch slide (a feature common in jazz music style). Our results suggest that long-lasting, active experience of a musical style is associated with accurate neural priors for the sound features of the preferred style, in contrast to passive listening.

In recent years, the field of neuroaesthetics has gained considerable attention with music being a favored object of study. The majority of studies concerning music have, however, focused on the experience of Western tonal music (TM), which is characterized by tonal hierarchical organization, a high degree of consonance, and a tendency to provide the listener with a tonic as a reference point during the listening experience. We argue that a narrow focus on Western TM may have led to a one-sided view regarding the qualities of the aesthetic experience of music since Western art music from the 20th and 21st century like atonal music (AM) – while lacking a tonal hierarchical structure, and while being highly dissonant and hard to predict – is nevertheless enjoyed by a group of avid listeners. We propose a research focus that investigates, in particular, the experience of AM as a novel and compelling way with which to enhance our understanding of both the aesthetic appreciation of music and the role of predictive models in the context of musical pleasure. We use music theoretical analysis and music information retrieval methods to demonstrate how AM presents the listener with a highly uncertain auditory environment. Specifically, an analysis of a corpus of 100 musical segments is used to illustrate how tonal classical music and AM differ quantitatively in terms of both key and pulse clarity values. We then examine person related, extrinsic and intrinsic factors, that point to potential mechanisms underlying the appreciation and pleasure derived from AM. We argue that personality traits like “openness to experience,” the framing of AM as art, and the mere exposure effect are key components of such mechanisms. We further argue that neural correlates of uncertainty estimation could represent a central mechanism for engaging with AM and that such contexts engender a comparatively weak predictive model in the listener. Finally we argue that in such uncertain contexts, correct predictions may be more subjectively rewarding than prediction errors since they signal to the individual that their predictive model is improving.

Motor exercise, such as sport or musical activities, helps with a plethora of diseases by modulating brain functions in neocortical and subcortical regions, resulting in behavioural changes related to mood regulation, well-being, memory, and even cognitive preservation in aging and neurodegenerative diseases. Although evidence is accumulating on the systemic neural mechanisms mediating these brain effects, the specific mechanisms by which exercise acts upon the cellular level are still under investigation. This is particularly the case for music training, a much less studied instance of motor exercise than sport. With regards to sport, consistent neurobiological research has focused on the brain-derived neurotrophic factor (BDNF), an essential player in the central nervous system. BDNF stimulates the growth and differentiation of neurons and synapses. It thrives in the hippocampus, the cortex, and the basal forebrain, which are the areas vital for memory, learning, and higher cognitive functions. Animal models and neurocognitive experiments on human athletes converge in demonstrating that physical exercise reliably boosts BDNF levels. In this review, we highlight comparable early findings obtained with animal models and elderly humans exposed to musical stimulation, showing how perceptual exposure to music might affect BDNF release, similar to what has been observed for sport. We subsequently propose a novel hypothesis that relates the neuroplastic changes in the human brains after musical training to genetically- and exercise-driven BDNF levels.

People can experience different emotions when listening to music. A growing number of studies have investigated the brain structures and neural connectivities associated with perceived emotions. However, very little is known about the effect of an explicit act of judgment on the neural processing of emotionally-valenced music. In this study, we adopted the novel consensus clustering paradigm, called binarisation of consensus partition matrices (Bi-CoPaM), to study whether and how the conscious aesthetic evaluation of the music would modulate brain connectivity networks related to emotion and reward processing. Participants listened to music under three conditions – one involving a non-evaluative judgment, one involving an explicit evaluative aesthetic judgment, and one involving no judgment at all (passive listening only). During non-evaluative attentive listening we obtained auditory-limbic connectivity whereas when participants were asked to decide explicitly whether they liked or disliked the music excerpt, only two clusters of intercommunicating brain regions were found: one including areas related to auditory processing and action observation, and the other comprising higher-order structures involved with visual processing. Results indicate that explicit evaluative judgment has an impact on the neural auditory-limbic connectivity during affective processing of music.

Capitalizing from neuroscience knowledge on how individuals are affected by the sound environment, we propose to adopt a cybernetic and ecological point of view on the musical aesthetic experience, which includes subprocesses, such as feature extraction and integration, early affective reactions and motor actions, style mastering and conceptualization, emotion and proprioception, evaluation and preference. In this perspective, the role of the listener/composer/performer is seen as that of an active “agent” coping in highly individual ways with the sounds. The findings concerning the neural adaptations in musicians, following long-term exposure to music, are then reviewed by keeping in mind the distinct subprocesses of a musical aesthetic experience. We conclude that these neural adaptations can be conceived of as the immediate and lifelong interactions with multisensorial stimuli (having a predominant auditory component), which result in lasting changes of the internal state of the “agent”. In a continuous loop, these changes affect, in turn, the subprocesses involved in a musical aesthetic experience, towards the final goal of achieving better perceptual, motor and proprioceptive responses to the immediate demands of the sounding environment. The resulting neural adaptations in musicians closely depend on the duration of the interactions, the starting age, the involvement of attention, the amount of motor practice and the musical genre played.

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.

The present literature review investigated how pleasure induced by music and visual-art has been conceptually understood in empirical research over the past 20 years. After an initial selection of abstracts from seven databases (keywords: pleasure, reward, enjoyment, and hedonic), twenty music and eleven visual-art papers were systematically compared. The following questions were addressed: (1) What is the role of the keyword in the research question? (2) Is pleasure considered a result of variation in the perceiver’s internal or external attributes? (3) What are the most commonly employed methods and main variables in empirical settings? Based on these questions, our critical integrative analysis aimed to identify which themes and processes emerged as key features for conceptualizing art-induced pleasure. The results demonstrated great variance in how pleasure has been approached: In the music studies pleasure was often a clear object of investigation, whereas in the visual-art studies the term was often embedded into the context of an aesthetic experience, or used otherwise in a descriptive, indirect sense. Music studies often targeted different emotions, their intensity or anhedonia. Biographical and background variables and personality traits of the perceiver were often measured. Next to behavioral methods, a common method was brain imaging which often targeted the reward circuitry of the brain in response to music. Visual-art pleasure was also frequently addressed using brain imaging methods, but the research focused on sensory cortices rather than the reward circuit alone. Compared with music research, visual-art research investigated more frequently pleasure in relation to conscious, cognitive processing, where the variations of stimulus features and the changing of viewing modes were regarded as explanatory factors of the derived experience. Despite valence being frequently applied in both domains, we conclude, that in empirical music research pleasure seems to be part of core affect and hedonic tone modulated by stable personality variables, whereas in visual-art research pleasure is a result of the so called conceptual act depending on a chosen strategy to approach art. We encourage an integration of music and visual-art into to a multi-modal framework to promote a more versatile understanding of pleasure in response to aesthetic artifacts.

Our sense of rhythm relies on orchestrated activity of several cerebral and cerebellar structures. Although functional connectivity studies have advanced our understanding of rhythm perception, this phenomenon has not been sufficiently studied as a function of musical training and beyond the General Linear Model (GLM) approach. Here, we studied pulse clarity processing during naturalistic music listening using a data-driven approach (independent component analysis; ICA). Participants' (18 musicians and 18 controls) functional magnetic resonance imaging (fMRI) responses were acquired while listening to music. A targeted region of interest (ROI) related to pulse clarity processing was defined, comprising auditory, somatomotor, basal ganglia, and cerebellar areas. The ICA decomposition was performed under different model orders, i.e., under a varying number of assumed independent sources, to avoid relying on prior model order assumptions. The components best predicted by a measure of the pulse clarity of the music, extracted computationally from the musical stimulus, were identified. Their corresponding spatial maps uncovered a network of auditory (perception) and motor (action) areas in an excitatory-inhibitory relationship at lower model orders, while mainly constrained to the auditory areas at higher model orders. Results revealed (a) a strengthened functional integration of action-perception networks associated with pulse clarity perception hidden from GLM analyses, and (b) group differences between musicians and non-musicians in pulse clarity processing, suggesting lifelong musical training as an important factor that may influence beat processing.

We dealt with the problem of artifacts in EEG signals in relation to the usage of lengthy trials. Specifically, we considered eye artifacts found in EEG signals, their influence in the analysis of the data and alternatives to diminish their impact on later studies of brain activity on lengthy tasks. We proposed a scheme of partial rejection of independent signal components, provided a method to extract EEG signal components with diminished influence of eye artifacts, and assess the importance of using artifact free signal excerpts to extract signal components in order to analyze brain activity in a musical context.

Listening to music is above all a human experience, which becomes an aesthetic experience when an individual immerses himself/herself in the music, dedicating attention to perceptual-cognitive-affective interpretation and evaluation. The study of these processes where the individual perceives, understands, enjoys and evaluates a set of auditory stimuli has mainly been focused on the effect of music on specific brain structures, as measured with neurophysiology and neuroimaging techniques. The very recent application of network science algorithms to brain research allows an insight into the functional connectivity between brain regions. These studies in network neuroscience have identified distinct circuits that function during goal-directed tasks and resting states. We review recent neuroimaging findings which indicate that music listening is traceable in terms of network connectivity and activations of target regions in the brain, in particular between the auditory cortex, the reward brain system and brain regions active during mind wandering.

Music information retrieval (MIR) methods offer interesting possibilities for automatically identifying time points in music recordings that relate to specific brain responses. However, how the acoustical features and the novelty of the music structure affect the brain response is not yet clear. In the present study, we tested a new method for automatically identifying time points of brain responses based on MIR analysis. We utilized an existing database including brain recordings of 48 healthy listeners measured with electroencephalography (EEG) and magnetoencephalography (MEG). While we succeeded in capturing brain responses related to acoustical changes in the modern tango piece Adios Nonino, we obtained less reliable brain responses with a metal rock piece and a modern symphony orchestra musical composition. However, brain responses might also relate to the novelty of the music structure. Hence, we added a manual musicological analysis of novelty in the musical structure to the computational acoustic analysis, obtaining strong brain responses even to the rock and modern pieces. Although no standardized method yet exists, these preliminary results suggest that analysis of novelty in music is an important aid to MIR analysis for investigating brain responses to realistic music.

Musical training leads to sensory and motor neuroplastic changes in the human brain. Motivated by findings on enlarged corpus callosum in musicians and asymmetric somatomotor representation in string players, we investigated the relationship between musical training, callosal anatomy, and interhemispheric functional symmetry during music listening. Functional symmetry was increased in musicians compared to nonmusicians, and in keyboardists compared to string players. This increased functional symmetry was prominent in visual and motor brain networks. Callosal size did not significantly differ between groups except for the posterior callosum in musicians compared to nonmusicians. We conclude that the distinctive postural and kinematic symmetry in instrument playing cross-modally shapes information processing in sensory-motor cortical areas during music listening. This cross-modal plasticity suggests that motor training affects music perception.

Musical expertise is associated with structural and functional changes in the brain that underlie facilitated auditory perception. We investigated whether the phase locking (PL) and amplitude modulations (AM) of neuronal oscillations in response to musical chords are correlated with musical expertise and whether they reflect the prototypicality of chords in Western tonal music. To this aim, we recorded magnetoencephalography (MEG) while musicians and non-musicians were presented with common prototypical major and minor chords, and with uncommon, non-prototypical dissonant and mistuned chords, while watching a silenced movie. We then analyzed the PL and AM of ongoing oscillations in the theta (4-8 Hz) alpha (8-14 Hz), beta- (14-30 Hz) and gamma- (30-80 Hz) bands to these chords. We found that musical expertise was associated with strengthened PL of ongoing oscillations to chords over a wide frequency range during the first 300 ms from stimulus onset, as opposed to increased alpha-band AM to chords over temporal MEG channels. In musicians, the gamma-band PL was strongest to non-prototypical compared to other chords, while in non-musicians PL was strongest to minor chords. In both musicians and non-musicians the long-latency (> 200 ms) gamma-band PL was also sensitive to chord identity, and particularly to the amplitude modulations (beats) of the dissonant chord. These findings suggest that musical expertise modulates oscillation PL to musical chords and that the strength of these modulations is dependent on chord prototypicality.

The ability to recognize emotions contained in facial expressions are affected by both affective traits and states and varies widely between individuals. While affective traits are stable in time, affective states can be regulated more rapidly by environmental stimuli, such as music, that indirectly modulate the brain state. Here, we tested whether a relaxing or irritating sound environment affects implicit processing of facial expressions. Moreover, we investigated whether and how individual traits of anxiety and emotional control interact with this process. 32 healthy subjects performed an implicit emotion processing task (presented to subjects as a gender discrimination task) while the sound environment was defined either by a) a therapeutic music sequence (MusiCure), b) a noise sequence or c) silence. Individual changes in mood were sampled before and after the task by a computerized questionnaire. Additionally, emotional control and trait anxiety were assessed in a separate session by paper and pencil questionnaires. Results showed a better mood after the MusiCure condition compared with the other experimental conditions and faster responses to happy faces during MusiCure compared with angry faces during Noise. Moreover, individuals with higher trait anxiety were faster in performing the implicit emotion processing task during MusiCure compared with Silence. These findings suggest that sound-induced affective states are associated with differential responses to angry and happy emotional faces at an implicit stage of processing, and that a relaxing sound environment facilitates the implicit emotional processing in anxious individuals.

Listening to music has been found to reduce acute and chronic pain. The underlying mechanisms are poorly understood; however, emotion and cognitive mechanisms have been suggested to influence the analgesic effect of music. In this study we investigated the influence of familiarity, emotional and cognitive features, and cognitive style on music-induced analgesia. Forty-eight healthy participants were divided into three groups (empathizers, systemizers and balanced) and received acute pain induced by heat while listening to different sounds. Participants listened to unfamiliar Mozart music rated with high valence and low arousal, unfamiliar environmental sounds with similar valence and arousal as the music, an active distraction task (mental arithmetic) and a control, and rated the pain. Data showed that the active distraction led to significantly less pain than did the music or sounds. Both unfamiliar music and sounds reduced pain significantly when compared to the control condition; however, music was no more effective than sound to reduce pain. Furthermore, we found correlations between pain and emotion ratings. Finally, systemizers reported less pain during the mental arithmetic compared with the other two groups. These findings suggest that familiarity may be key in the influence of the cognitive and emotional mechanisms of music-induced analgesia, and that cognitive styles may influence pain perception.

The importance of music in our daily life has given rise to an increased number of studies addressing the brain regions involved in its appreciation. Some of these studies controlled only for the familiarity of the stimuli, while others relied on pleasantness ratings, and others still on musical preferences. With a listening test and a functional magnetic resonance imaging (fMRI) experiment, we wished to clarify the role of familiarity in the brain correlates of music appreciation by controlling, in the same study, for both familiarity and musical preferences. First, we conducted a listening test, in which participants rated the familiarity and liking of song excerpts from the pop/rock repertoire, allowing us to select a personalized set of stimuli per subject. Then, we used a passive listening paradigm in fMRI to study music appreciation in a naturalistic condition with increased ecological value. Brain activation data revealed that broad emotion-related limbic and paralimbic regions as well as the reward circuitry were significantly more active for familiar relative to unfamiliar music. Smaller regions in the cingulate cortex and frontal lobe, including the motor cortex and Broca's area, were found to be more active in response to liked music when compared to disliked one. Hence, familiarity seems to be a crucial factor in making the listeners emotionally engaged with music, as revealed by fMRI data.

Musical competence may confer cognitive advantages that extend beyond processing of familiar musical sounds. Behavioural evidence indicates a general enhancement of both working memory and attention in musicians. It is possible that musicians, due to their training, are better able to maintain focus on task-relevant stimuli, a skill which is crucial to working memory. We measured the blood oxygenation-level dependent (BOLD) activation signal in musicians and non-musicians during working memory of musical sounds to determine the relation among performance, musical competence and generally enhanced cognition. All participants easily distinguished the stimuli. We tested the hypothesis that musicians nonetheless would perform better, and that differential brain activity would mainly be present in cortical areas involved in cognitive control such as the lateral prefrontal cortex. The musicians performed better as reflected in reaction times and error rates. Musicians also had larger BOLD responses than non-musicians in neuronal networks that sustain attention and cognitive control, including regions of the lateral prefrontal cortex, lateral parietal cortex, insula, and putamen in the right hemisphere, and bilaterally in the posterior dorsal prefrontal cortex and anterior cingulate gyrus. The relationship between the task performance and the magnitude of the BOLD response was more positive in musicians than in non-musicians, particularly during the most difficult working memory task. The results confirm previous findings that neural activity increases during enhanced working memory performance. The results also suggest that superior working memory task performance in musicians rely on an enhanced ability to exert sustained cognitive control. This cognitive benefit in musicians may be a consequence of focused musical training.