Cognitive and sensory expectations independently shape musical expectancy and pleasure.

Expectation is crucial for our enjoyment of music, yet the underlying generative mechanisms remain unclear. While sensory models derive predictions based on local acoustic information in the auditory signal, cognitive models assume abstract knowledge of music structure acquired over the long term. To evaluate these two contrasting mechanisms, we compared simulations from four computational models of musical expectancy against subjective expectancy and pleasantness ratings of over 1000 chords sampled from 739 US Billboard pop songs. Bayesian model comparison revealed that listeners' expectancy and pleasantness ratings were predicted by the independent, non-overlapping, contributions of cognitive and sensory expectations. Furthermore, cognitive expectations explained over twice the variance in listeners' perceived surprise compared to sensory expectations, suggesting a larger relative importance of long-term representations of music structure over short-term sensory-acoustic information in musical expectancy. Our results thus emphasize the distinct, albeit complementary, roles of cognitive and sensory expectations in shaping musical pleasure, and suggest that this expectancy-driven mechanism depends on musical information represented at different levels of abstraction along the neural hierarchy. This article is part of the theme issue 'Art, aesthetics and predictive processing: theoretical and empirical perspectives'.

Sensorimotor synchronization to music reduces pain.

Pain-reducing effects of music listening are well-established, but the effects are small and their clinical relevance questionable. Recent theoretical advances, however, have proposed that synchronizing to music, such as clapping, tapping or dancing, has evolutionarily important social effects that are associated with activation of the endogenous opioid system (which supports both analgesia and social bonding). Thus, active sensorimotor synchronization to music could have stronger analgesic effects than simply listening to music. In this study, we show that sensorimotor synchronization to music significantly amplifies the pain-reducing effects of music listening. Using pressure algometry to the fingernails, pain stimuli were delivered to n = 59 healthy adults either during music listening or silence, while either performing an active tapping task or a passive control task. Compared to silence without tapping, music with tapping (but not simply listening to music) reduced pain with a large, clinically significant, effect size (d = 0.93). Simply tapping without music did not elicit such an effect. Our analyses indicate that both attentional and emotional mechanisms drive the pain-reducing effects of sensorimotor synchronization to music, and that tapping to music in addition to merely listening to music may enhance pain-reducing effects in both clinical contexts and everyday life. The study was registered as a clinical trial at ClinicalTrials.gov (registration number NCT05267795), and the trial was first posted on 04/03/2022.

Neural correlates of statistical learning in developmental dyslexia: An electroencephalography study.

The human brain extracts statistical regularities from the surrounding environment in a process called statistical learning. Behavioural evidence suggests that developmental dyslexia affects statistical learning. However, surprisingly few studies have assessed how developmental dyslexia affects the neural processing underlying this type of learning. We used electroencephalography to explore the neural correlates of an important aspect of statistical learning - sensitivity to transitional probabilities - in individuals with developmental dyslexia. Adults diagnosed with developmental dyslexia (n = 17) and controls (n = 19) were exposed to a continuous stream of sound triplets. Every so often, a triplet ending had a low transitional probability given the triplet's first two sounds ("statistical deviants"). Furthermore, every so often a triplet ending was presented from a deviant location ("acoustic deviants"). We examined mismatch negativity elicited by statistical deviants (sMMN), and MMN elicited by location deviants (i.e., acoustic changes). Acoustic deviants elicited a MMN which was larger in the control group than in the developmental dyslexia group. Statistical deviants elicited a small, yet significant, sMMN in the control group, but not in the developmental dyslexia group. However, the difference between the groups was not significant. Our findings indicate that the neural mechanisms underlying pre-attentive acoustic change detection and implicit statistical auditory learning are both affected in developmental dyslexia.

The promise of music therapy for Alzheimer's disease: A review.

Alzheimer's disease (AD) is a progressive neurodegenerative disease associated with cognitive decline. Memory problems are typically among the first signs of cognitive impairment in AD, and they worsen considerably as the disease progresses. However, musical memory is partially spared in patients with AD, despite severe deficits in episodic (and partly semantic) memory. AD patients can learn new songs, encode novel verbal information, and react emotionally to music. These effects of music have encouraged the use and development of music therapy (MT) for AD management. MT is easy to implement and well-tolerated by most patients and their caregivers. Effects of MT in patients with AD include improved mood, reduced depressive scores and trait anxiety, enhanced autobiographical recall, verbal fluency, and cognition. Here, we review musical memory in AD, therapeutic effects of studies using MT on AD, and potential mechanisms underlying those therapeutic effects. We argue that, because AD begins decades before the presentation of clinical symptoms, music interventions might be a promising means to delay and decelerate the neurodegeneration in individuals at risk for AD, such as individuals with genetic risk or subjective cognitive decline.

Study protocol for the Alzheimer and music therapy study: An RCT to compare the efficacy of music therapy and physical activity on brain plasticity, depressive symptoms, and cognitive decline, in a population with and at risk for Alzheimer's disease.

BACKGROUND: There is anecdotal evidence for beneficial effects of music therapy in patients with Alzheimer's Disease (AD). However, there is a lack of rigorous research investigating this issue. The aim of this study is to evaluate the effects of music therapy and physical activity on brain plasticity, mood, and cognition in a population with AD and at risk for AD. METHODS: One-hundred and thirty-five participants with memory complaints will be recruited for a parallel, three-arm Randomized Controlled Trial (RCT). Inclusion criteria are a diagnosis of mild (early) AD or mild cognitive impairment (MCI), or memory complaints without other neuropsychiatric pathology. Participants are randomised into either a music therapy intervention (singing lessons), an active control group (physical activity) or a passive control group (no intervention) for 12 months. The primary outcomes are the brain age gap, measured via magnetic resonance imaging (MRI), and depressive symptoms. Secondary outcomes include cognitive performance, activities of daily living, brain structure (voxel-based morphometry and diffusion tensor imaging), and brain function (resting-state functional MRI). TRIAL STATUS: Screening of participants began in April 2018. A total of 84 participants have been recruited and started intervention, out of which 48 participants have completed 12 months of intervention and post-intervention assessment. DISCUSSION: Addressing the need for rigorous longitudinal data for the effectiveness of music therapy in people with and at risk for developing AD, this trial aims to enhance knowledge regarding cost-effective interventions with potentially high clinical applicability. TRIAL REGISTRATION: ClinicalTrials.gov identifier: NCT03444181, registered on February 23, 2018.

Unpredictability of the "when" influences prediction error processing of the "what" and "where".

The capability to establish accurate predictions is an integral part of learning. Whether predictions about different dimensions of a stimulus interact with each other, and whether such an interaction affects learning, has remained elusive. We conducted a statistical learning study with EEG (electroencephalography), where a stream of consecutive sound triplets was presented with deviants that were either: (a) statistical, depending on the triplet ending probability, (b) physical, due to a change in sound location or (c) double deviants, i.e. a combination of the two. We manipulated the predictability of stimulus-onset by using random stimulus-onset asynchronies. Temporal unpredictability due to random onsets reduced the neurophysiological responses to statistical and location deviants, as indexed by the statistical mismatch negativity (sMMN) and the location MMN. Our results demonstrate that the predictability of one stimulus attribute influences the processing of prediction error signals of other stimulus attributes, and thus also learning of those attributes.

Neocortical substrates of feelings evoked with music in the ACC, insula, and somatosensory cortex.

Neurobiological models of emotion focus traditionally on limbic/paralimbic regions as neural substrates of emotion generation, and insular cortex (in conjunction with isocortical anterior cingulate cortex, ACC) as the neural substrate of feelings. An emerging view, however, highlights the importance of isocortical regions beyond insula and ACC for the subjective feeling of emotions. We used music to evoke feelings of joy and fear, and multivariate pattern analysis (MVPA) to decode representations of feeling states in functional magnetic resonance (fMRI) data of n = 24 participants. Most of the brain regions providing information about feeling representations were neocortical regions. These included, in addition to granular insula and cingulate cortex, primary and secondary somatosensory cortex, premotor cortex, frontal operculum, and auditory cortex. The multivoxel activity patterns corresponding to feeling representations emerged within a few seconds, gained in strength with increasing stimulus duration, and replicated results of a hypothesis-generating decoding analysis from an independent experiment. Our results indicate that several neocortical regions (including insula, cingulate, somatosensory and premotor cortices) are important for the generation and modulation of feeling states. We propose that secondary somatosensory cortex, which covers the parietal operculum and encroaches on the posterior insula, is of particular importance for the encoding of emotion percepts, i.e., preverbal representations of subjective feeling.

Trait Empathy Shapes Neural Responses Toward Sad Music.

Individuals with a predisposition to empathize engage with sad music in a compelling way, experiencing overall more pleasurable emotions. However, the neural mechanisms underlying these music-related experiences in empathic individuals are unknown. The present study tested whether dispositional empathy modulates neural responses to sad compared with happy music. Twenty-four participants underwent fMRI while listening to 4-min blocks of music evoking sadness or happiness. Using voxel-wise regression, we found a positive correlation between trait empathy (with scores assessed by the Interpersonal Reactivity Index) and eigenvector centrality values in the ventromedial prefrontal cortex (vmPFC), including the medial orbitofrontal cortex (mOFC). We then performed a functional connectivity (FC) analysis to detect network nodes showing stronger FC with the vmPFC/mOFC during the presentation of sad versus happy music. By doing so, we identified a "music-empathy" network (vmPFC/mOFC, dorsomedial prefrontal cortex, primary visual cortex, bilateral claustrum and putamen, and cerebellum) that is spontaneously recruited while listening to sad music and includes brain regions that support the coding of compassion, mentalizing, and visual mental imagery. Importantly, our findings extend the current understanding of empathic behaviors to the musical domain and pinpoint sad music as an effective stimulus to be employed in social neuroscience research.

When the statistical MMN meets the physical MMN.

How do listeners respond to prediction errors within patterned sequence of sounds? To answer this question we carried out a statistical learning study using electroencephalography (EEG). In a continuous auditory stream of sound triplets the deviations were either (a) statistical, in terms of transitional probability, (b) physical, due to a change in sound location (left or right speaker) or (c) a double deviants, i.e. a combination of the two. Statistical and physical deviants elicited a statistical mismatch negativity and a physical MMN respectively. Most importantly, we found that effects of statistical and physical deviants interacted (the statistical MMN was smaller when co-occurring with a physical deviant). Results show, for the first time, that processing of prediction errors due to statistical learning is affected by prediction errors due to physical deviance. Our findings thus show that the statistical MMN interacts with the physical MMN, implying that prediction error processing due to physical sound attributes suppresses processing of learned statistical properties of sounds.

Investigating the Neural Encoding of Emotion with Music.

Does our understanding of the human brain remain incomplete without a proper understanding of how the brain processes music? Here, the author makes a passionate plea for the use of music in the investigation of human emotion and its brain correlates, arguing that music can change activity in all brain structures associated with emotions, which has important implications on how we understand human emotions and their disorders and how we can make better use of beneficial effects of music in therapy.

Neural Encoding of Auditory Features during Music Perception and Imagery.

Despite many behavioral and neuroimaging investigations, it remains unclear how the human cortex represents spectrotemporal sound features during auditory imagery, and how this representation compares to auditory perception. To assess this, we recorded electrocorticographic signals from an epileptic patient with proficient music ability in 2 conditions. First, the participant played 2 piano pieces on an electronic piano with the sound volume of the digital keyboard on. Second, the participant replayed the same piano pieces, but without auditory feedback, and the participant was asked to imagine hearing the music in his mind. In both conditions, the sound output of the keyboard was recorded, thus allowing precise time-locking between the neural activity and the spectrotemporal content of the music imagery. This novel task design provided a unique opportunity to apply receptive field modeling techniques to quantitatively study neural encoding during auditory mental imagery. In both conditions, we built encoding models to predict high gamma neural activity (70-150 Hz) from the spectrogram representation of the recorded sound. We found robust spectrotemporal receptive fields during auditory imagery with substantial, but not complete overlap in frequency tuning and cortical location compared to receptive fields measured during auditory perception.

Language and music phrase boundary processing in Autism Spectrum Disorder: An ERP study.

Autism spectrum disorder (ASD) is frequently associated with communicative impairment, regardless of intelligence level or mental age. Impairment of prosodic processing in particular is a common feature of ASD. Despite extensive overlap in neural resources involved in prosody and music processing, music perception seems to be spared in this population. The present study is the first to investigate prosodic phrasing in ASD in both language and music, combining event-related brain potential (ERP) and behavioral methods. We tested phrase boundary processing in language and music in neuro-typical adults and high-functioning individuals with ASD. We targeted an ERP response associated with phrase boundary processing in both language and music - i.e., the Closure Positive Shift (CPS). While a language-CPS was observed in the neuro-typical group, for ASD participants a smaller response failed to reach statistical significance. In music, we found a boundary-onset music-CPS for both groups during pauses between musical phrases. Our results support the view of preserved processing of musical cues in ASD individuals, with a corresponding prosodic impairment. This suggests that, despite the existence of a domain-general processing mechanism (the CPS), key differences in the integration of features of language and music may lead to the prosodic impairment in ASD.

Effects of music therapy and music-based interventions in the treatment of substance use disorders: A systematic review.

Music therapy (MT) and music-based interventions (MBIs) are increasingly used for the treatment of substance use disorders (SUD). Previous reviews on the efficacy of MT emphasized the dearth of research evidence for this topic, although various positive effects were identified. Therefore, we conducted a systematic search on published articles examining effects of music, MT and MBIs and found 34 quantitative and six qualitative studies. There was a clear increase in the number of randomized controlled trials (RCTs) during the past few years. We had planned for a meta-analysis, but due to the diversity of the quantitative studies, effect sizes were not computed. Beneficial effects of MT/ MBI on emotional and motivational outcomes, participation, locus of control, and perceived helpfulness were reported, but results were inconsistent across studies. Furthermore, many RCTs focused on effects of single sessions. No published longitudinal trials could be found. The analysis of the qualitative studies revealed four themes: emotional expression, group interaction, development of skills, and improvement of quality of life. Considering these issues for quantitative research, there is a need to examine social and health variables in future studies. In conclusion, due to the heterogeneity of the studies, the efficacy of MT/ MBI in SUD treatment still remains unclear.

Effects of veridical expectations on syntax processing in music: Event-related potential evidence.

Numerous past studies have investigated neurophysiological correlates of music-syntactic processing. However, only little is known about how prior knowledge about an upcoming syntactically irregular event modulates brain correlates of music-syntactic processing. Two versions of a short chord sequence were presented repeatedly to non-musicians (n = 20) and musicians (n = 20). One sequence version ended on a syntactically regular chord, and the other one ended on a syntactically irregular chord. Participants were either informed (cued condition), or not informed (non-cued condition) about whether the sequence would end on the regular or the irregular chord. Results indicate that in the cued condition (compared to the non-cued condition) the peak latency of the early right anterior negativity (ERAN), elicited by irregular chords, was earlier in both non-musicians and musicians. However, the expectations due to the knowledge about the upcoming event (veridical expectations) did not influence the amplitude of the ERAN. These results suggest that veridical expectations modulate only the speed, but not the principle mechanisms, of music-syntactic processing.

Music and the heart.

Music can powerfully evoke and modulate emotions and moods, along with changes in heart activity, blood pressure (BP), and breathing. Although there is great heterogeneity in methods and quality among previous studies on effects of music on the heart, the following findings emerge from the literature: Heart rate (HR) and respiratory rate (RR) are higher in response to exciting music compared with tranquilizing music. During musical frissons (involving shivers and piloerection), both HR and RR increase. Moreover, HR and RR tend to increase in response to music compared with silence, and HR appears to decrease in response to unpleasant music compared with pleasant music. We found no studies that would provide evidence for entrainment of HR to musical beats. Corresponding to the increase in HR, listening to exciting music (compared with tranquilizing music) is associated with a reduction of heart rate variability (HRV), including reductions of both low-frequency and high-frequency power of the HRV. Recent findings also suggest effects of music-evoked emotions on regional activity of the heart, as reflected in electrocardiogram amplitude patterns. In patients with heart disease (similar to other patient groups), music can reduce pain and anxiety, associated with lower HR and lower BP. In general, effects of music on the heart are small, and there is great inhomogeneity among studies with regard to methods, findings, and quality. Therefore, there is urgent need for systematic high-quality research on the effects of music on the heart, and on the beneficial effects of music in clinical settings.

Music-evoked emotions: principles, brain correlates, and implications for therapy.

This paper describes principles underlying the evocation of emotion with music: evaluation, resonance, memory, expectancy/tension, imagination, understanding, and social functions. Each of these principles includes several subprinciples, and the framework on music-evoked emotions emerging from these principles and subprinciples is supposed to provide a starting point for a systematic, coherent, and comprehensive theory on music-evoked emotions that considers both reception and production of music, as well as the relevance of emotion-evoking principles for music therapy.

The effects of supervised learning on event-related potential correlates of music-syntactic processing.

Humans process music even without conscious effort according to implicit knowledge about syntactic regularities. Whether such automatic and implicit processing is modulated by veridical knowledge has remained unknown in previous neurophysiological studies. This study investigates this issue by testing whether the acquisition of veridical knowledge of a music-syntactic irregularity (acquired through supervised learning) modulates early, partly automatic, music-syntactic processes (as reflected in the early right anterior negativity, ERAN), and/or late controlled processes (as reflected in the late positive component, LPC). Excerpts of piano sonatas with syntactically regular and less regular chords were presented repeatedly (10 times) to non-musicians and amateur musicians. Participants were informed by a cue as to whether the following excerpt contained a regular or less regular chord. Results showed that the repeated exposure to several presentations of regular and less regular excerpts did not influence the ERAN elicited by less regular chords. By contrast, amplitudes of the LPC (as well as of the P3a evoked by less regular chords) decreased systematically across learning trials. These results reveal that late controlled, but not early (partly automatic), neural mechanisms of music-syntactic processing are modulated by repeated exposure to a musical piece. This article is part of a Special Issue entitled SI: Prediction and Attention.

Functional centrality of amygdala, striatum and hypothalamus in a "small-world" network underlying joy: an fMRI study with music.

Current knowledge about small-world networks underlying emotions is sparse, and confined to functional magnetic resonance imaging (fMRI) studies using resting-state paradigms. This fMRI study applied Eigenvector Centrality Mapping (ECM) and functional connectivity analysis to reveal neural small-world networks underlying joy and fear. Joy and fear were evoked using music, presented in 4-min blocks. Results show that the superficial amygdala (SF), laterobasal amygdala (LB), striatum, and hypothalamus function as computational hubs during joy. Out of these computational hubs, the amygdala nuclei showed the highest centrality values. The SF showed functional connectivity during joy with the mediodorsal thalamus (MD) and nucleus accumbens (Nac), suggesting that SF, MD, and Nac modulate approach behavior in response to positive social signals such as joyful music. The striatum was functionally connected during joy with the LB, as well as with premotor cortex, areas 1 and 7a, hippocampus, insula and cingulate cortex, showing that sensorimotor, attentional, and emotional processes converge in the striatum during music perception. The hypothalamus showed functional connectivity during joy with hippocampus and MD, suggesting that hypothalamic endocrine activity is modulated by hippocampal and thalamic activity during sustained periods of music-evoked emotion. Our study indicates high centrality of the amygdala nuclei groups within a functional network underlying joy, suggesting that these nuclei play a central role for the modulation of emotion-specific activity within this network.

Brain correlates of music-evoked emotions.

Music is a universal feature of human societies, partly owing to its power to evoke strong emotions and influence moods. During the past decade, the investigation of the neural correlates of music-evoked emotions has been invaluable for the understanding of human emotion. Functional neuroimaging studies on music and emotion show that music can modulate activity in brain structures that are known to be crucially involved in emotion, such as the amygdala, nucleus accumbens, hypothalamus, hippocampus, insula, cingulate cortex and orbitofrontal cortex. The potential of music to modulate activity in these structures has important implications for the use of music in the treatment of psychiatric and neurological disorders.

How emotional abilities modulate the influence of early life stress on hippocampal functioning.

Early life stress (ELS) is known to have considerable influence on brain development, mental health and affective functioning. Previous investigations have shown that alexithymia, a prevalent personality trait associated with difficulties experiencing and verbalizing emotions, is particularly related to ELS. The aim of the present study was to investigate how neural correlates of emotional experiences in alexithymia are altered in the presence and absence of ELS. Therefore, 50 healthy individuals with different levels of alexithymia were matched regarding ELS and investigated with respect to neural correlates of audio-visually induced emotional experiences via functional magnetic resonance imaging. The main finding was that ELS modulated hippocampal responses to pleasant (>neutral) stimuli in high-alexithymic individuals, whereas there was no such modulation in low-alexithymic individuals matched for ELS. Behavioral and psychophysiological results followed a similar pattern. When considered independent of ELS, alexithymia was associated with decreased responses in insula (pleasant > neutral) and temporal pole (unpleasant > neutral). Our results show that the influence of ELS on emotional brain responses seems to be modulated by an individual's degree of alexithymia. Potentially, protective and adverse effects of emotional abilities on brain responses to emotional experiences are discussed.