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.

Is musical engagement enough to keep the brain young?

Music-making and engagement in music-related activities have shown procognitive benefits for healthy and pathological populations, suggesting reductions in brain aging. A previous brain aging study, using Brain Age Gap Estimation (BrainAGE), showed that professional and amateur-musicians had younger appearing brains than non-musicians. Our study sought to replicate those findings and analyze if musical training or active musical engagement was necessary to produce an age-decelerating effect in a cohort of healthy individuals. We scanned 125 healthy controls and investigated if musician status, and if musical behaviors, namely active engagement (AE) and musical training (MT) [as measured using the Goldsmiths Musical Sophistication Index (Gold-MSI)], had effects on brain aging. Our findings suggest that musician status is not related to BrainAGE score, although involvement in current physical activity is. Although neither MT nor AE subscales of the Gold-MSI are predictive for BrainAGE scores, dispositional resilience, namely the ability to deal with challenge, is related to both musical behaviors and sensitivity to musical pleasure. While the study failed to replicate the findings in a previous brain aging study, musical training and active musical engagement are related to the resilience factor of challenge. This finding may reveal how such musical behaviors can potentially strengthen the brain's resilience to age, which may tap into a type of neurocognitive reserve.

Brain-correlates of processing local dependencies within a statistical learning paradigm.

Statistical learning refers to the implicit mechanism of extracting regularities in our environment. Numerous studies have investigated the neural basis of statistical learning. However, how the brain responds to violations of auditory regularities based on prior (implicit) learning requires further investigation. Here, we used functional magnetic resonance imaging (fMRI) to investigate the neural correlates of processing events that are irregular based on learned local dependencies. A stream of consecutive sound triplets was presented. Unbeknown to the subjects, triplets were either (a) standard, namely triplets ending with a high probability sound or, (b) statistical deviants, namely triplets ending with a low probability sound. Participants (n = 33) underwent a learning phase outside the scanner followed by an fMRI session. Processing of statistical deviants activated a set of regions encompassing the superior temporal gyrus bilaterally, the right deep frontal operculum including lateral orbitofrontal cortex, and the right premotor cortex. Our results demonstrate that the violation of local dependencies within a statistical learning paradigm does not only engage sensory processes, but is instead reminiscent of the activation pattern during the processing of local syntactic structures in music and language, reflecting the online adaptations required for predictive coding in the context of statistical learning.

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.

Tormenting thoughts: The posterior cingulate sulcus of the default mode network regulates valence of thoughts and activity in the brain's pain network during music listening.

Many individuals spend a significant amount of their time "mind-wandering". Mind-wandering often includes spontaneous, nonintentional thought, and a neural correlate of this kind of thought is the default mode network (DMN). Thoughts during mind-wandering can have positive or negative valence, but only little is known about the neural correlates of positive or negative thoughts. We used resting-state functional magnetic resonance imaging (fMRI) and music to evoke mind-wandering in n = 33 participants, with positive-sounding music eliciting thoughts with more positive valence and negative-sounding music eliciting thoughts with more negative valence. Applying purely data-driven analysis methods, we show that medial orbitofrontal cortex (mOFC, part of the ventromedial prefrontal cortex) and the posterior cingulate sulcus (likely area 23c of the posterior cingulate cortex), two sub-regions of the DMN, modulate the valence of thought-contents during mind-wandering. In addition, across two independent experiments, we observed that the posterior cingulate sulcus, a region involved in pain, shows valence-specific functional connectivity with core regions of the brain's putative pain network. Our results suggest that two DMN regions (mOFC and posterior cingulate sulcus) support the formation of negative spontaneous, nonintentional thoughts, and that the interplay between these structures with regions of the putative pain network forms a neural mechanism by which thoughts can become painful.

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.

Cortical thickness and resting-state cardiac function across the lifespan: A cross-sectional pooled mega-analysis.

Understanding the association between autonomic nervous system [ANS] function and brain morphology across the lifespan provides important insights into neurovisceral mechanisms underlying health and disease. Resting-state ANS activity, indexed by measures of heart rate [HR] and its variability [HRV] has been associated with brain morphology, particularly cortical thickness [CT]. While findings have been mixed regarding the anatomical distribution and direction of the associations, these inconsistencies may be due to sex and age differences in HR/HRV and CT. Previous studies have been limited by small sample sizes, which impede the assessment of sex differences and aging effects on the association between ANS function and CT. To overcome these limitations, 20 groups worldwide contributed data collected under similar protocols of CT assessment and HR/HRV recording to be pooled in a mega-analysis (N = 1,218 (50.5% female), mean age 36.7 years (range: 12-87)). Findings suggest a decline in HRV as well as CT with increasing age. CT, particularly in the orbitofrontal cortex, explained additional variance in HRV, beyond the effects of aging. This pattern of results may suggest that the decline in HRV with increasing age is related to a decline in orbitofrontal CT. These effects were independent of sex and specific to HRV; with no significant association between CT and HR. Greater CT across the adult lifespan may be vital for the maintenance of healthy cardiac regulation via the ANS-or greater cardiac vagal activity as indirectly reflected in HRV may slow brain atrophy. Findings reveal an important association between CT and cardiac parasympathetic activity with implications for healthy aging and longevity that should be studied further in longitudinal research.

A coordinate-based meta-analysis of music-evoked emotions.

Since the publication of the first neuroscience study investigating emotion with music about two decades ago, the number of functional neuroimaging studies published on this topic has increased each year. This research interest is in part due to the ubiquity of music across cultures, and to music's power to evoke a diverse range of intensely felt emotions. To support a better understanding of the brain correlates of music-evoked emotions this article reports a coordinate-based meta-analysis of neuroimaging studies (n = 47 studies with n = 944 subjects). The studies employed a range of diverse experimental approaches (e.g., using music to evoke joy, sadness, fear, tension, frissons, surprise, unpleasantness, or feelings of beauty). The results of an activation likelihood estimation (ALE) indicate large clusters in a range of structures, including amygdala, anterior hippocampus, auditory cortex, and numerous structures of the reward network (ventral and dorsal striatum, anterior cingulate cortex, orbitofrontal cortex, secondary somatosensory cortex). The results underline the rewarding nature of music, the role of the auditory cortex as an emotional hub, and the role of the hippocampus in attachment-related emotions and social bonding.

Aesthetic emotions are a key factor in aesthetic evaluation: Reply to Skov and Nadal (2020).

Our theoretical model (Menninghaus et al., 2019) defines aesthetic emotions by reference to their role in aesthetic evaluation, and specifically as being predictive of aesthetic liking/disliking. Skov and Nadal (2020) dismiss the construct of "aesthetic emotions" as a "dated supposition" adopted from a "speculative" tradition and assert that there are no such emotions. Accordingly, they question all pieces of empirical evidence we referred to as supporting our model. In our response, we rebut these objections point by point and defend as well as expand the empirical evidence in support of our model. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

The Effect of Emotional Valence on Ventricular Repolarization Dynamics Is Mediated by Heart Rate Variability: A Study of QT Variability and Music-Induced Emotions.

BACKGROUND: Emotions can affect cardiac activity, but their impact on ventricular repolarization variability, an important parameter providing information about cardiac risk and autonomic nervous system activity, is unknown. The beat-to-beat variability of the QT interval (QTV) from the body surface ECG is a non-invasive marker of repolarization variability, which can be decomposed into QTV related to RR variability (QTVrRRV) and QTV unrelated to RRV (QTVuRRV), with the latter thought to be a marker of intrinsic repolarization variability. AIM: To determine the effect of emotional valence (pleasant and unpleasant) on repolarization variability in healthy volunteers by means of QTV analysis. METHODS: 75 individuals (24.5 ± 3.2 years, 36 females) without a history of cardiovascular disease listened to music-excerpts that were either felt as pleasant (n = 6) or unpleasant (n = 6). Excerpts lasted about 90 s and were presented in a random order along with silent intervals (n = 6). QTV and RRV were derived from the ECG and the time-frequency spectrum of RRV, QTV, QTVuRRV and QTVrRRV as well as time-frequency coherence between QTV and RRV were estimated. Analysis was performed in low-frequency (LF), high frequency (HF) and total spectral bands. RESULTS: The heart rate-corrected QTV showed a small but significant increase from silence (median 347/interquartile range 31 ms) to listening to music felt as unpleasant (351/30 ms) and pleasant (355/32 ms). The dynamic response of QTV to emotional valence showed a transient phase lasting about 20 s after the onset of each musical excerpt. QTV and RRV were highly correlated in both HF and LF (mean coherence ranging 0.76-0.85). QTV and QTVrRRV decreased during listening to music felt as pleasant and unpleasant with respect to silence and further decreased during listening to music felt as pleasant. QTVuRRV was small and not affected by emotional valence. CONCLUSION: Emotional valence, as evoked by music, has a small but significant effect on QTV and QTVrRRV, but not on QTVuRRV. This suggests that the interaction between emotional valence and ventricular repolarization variability is mediated by cycle length dynamics and not due to intrinsic repolarization variability.

Author Correction: When the statistical MMN meets the physical MMN.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Uncertainty and Surprise Jointly Predict Musical Pleasure and Amygdala, Hippocampus, and Auditory Cortex Activity.

Listening to music often evokes intense emotions [1, 2]. Recent research suggests that musical pleasure comes from positive reward prediction errors, which arise when what is heard proves to be better than expected [3]. Central to this view is the engagement of the nucleus accumbens-a brain region that processes reward expectations-to pleasurable music and surprising musical events [4-8]. However, expectancy violations along multiple musical dimensions (e.g., harmony and melody) have failed to implicate the nucleus accumbens [9-11], and it is unknown how music reward value is assigned [12]. Whether changes in musical expectancy elicit pleasure has thus remained elusive [11]. Here, we demonstrate that pleasure varies nonlinearly as a function of the listener's uncertainty when anticipating a musical event, and the surprise it evokes when it deviates from expectations. Taking Western tonal harmony as a model of musical syntax, we used a machine-learning model [13] to mathematically quantify the uncertainty and surprise of 80,000 chords in US Billboard pop songs. Behaviorally, we found that chords elicited high pleasure ratings when they deviated substantially from what the listener had expected (low uncertainty, high surprise) or, conversely, when they conformed to expectations in an uninformative context (high uncertainty, low surprise). Neurally, we found using fMRI that activity in the amygdala, hippocampus, and auditory cortex reflected this interaction, while the nucleus accumbens only reflected uncertainty. These findings challenge current neurocognitive models of music-evoked pleasure and highlight the synergistic interplay between prospective and retrospective states of expectation in the musical experience. VIDEO ABSTRACT.

Heroic music stimulates empowering thoughts during mind-wandering.

It is generally well-known, and scientifically well established, that music affects emotions and moods. However, only little is known about the influence of music on thoughts. This scarcity is particularly surprising given the importance of the valence of thoughts for psychological health and well-being. We presented excerpts of heroic- and sad-sounding music to n = 62 individuals, and collected thought probes after each excerpt, assessing the valence and the nature of thoughts stimulated by the music. Our results show that mind-wandering emerged during listening to either type of music (heroic, sad), and that the type of music strongly influenced the thought contents during mind-wandering. Heroic-sounding music evoked more positive, exciting, constructive, and motivating thoughts, while sad-sounding music evoked more calm or demotivating thoughts. The results thus indicate that music has a strong effect on the valence of thought contents during mind-wandering, with heroic music evoking more empowering and motivating thoughts, and sad music more relaxing or depressive thoughts. These findings have important implications for the use of music in everyday life to promote health and well-being in both clinical populations and healthy individuals.

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.

What are aesthetic emotions?

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