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.

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.

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.

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.

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.

The Temporal Pole Top-Down Modulates the Ventral Visual Stream During Social Cognition.

The temporal pole (TP) has been associated with diverse functions of social cognition and emotion processing. Although the underlying mechanism remains elusive, one possibility is that TP acts as domain-general hub integrating socioemotional information. To test this, 26 participants were presented with 60 empathy-evoking film clips during fMRI scanning. The film clips were preceded by a linguistic sad or neutral context and half of the clips were accompanied by sad music. In line with its hypothesized role, TP was involved in the processing of sad context and furthermore tracked participants' empathic concern. To examine the neuromodulatory impact of TP, we applied nonlinear dynamic causal modeling to a multisensory integration network from previous work consisting of superior temporal gyrus (STG), fusiform gyrus (FG), and amygdala, which was extended by an additional node in the TP. Bayesian model comparison revealed a gating of STG and TP on fusiform-amygdalar coupling and an increase of TP to FG connectivity during the integration of contextual information. Moreover, these backward projections were strengthened by emotional music. The findings indicate that during social cognition, TP integrates information from different modalities and top-down modulates lower-level perceptual areas in the ventral visual stream as a function of integration demands.

The Distancing-Embracing model of the enjoyment of negative emotions in art reception.

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

Negative emotions in art reception: Refining theoretical assumptions and adding variables to the Distancing-Embracing model.

While covering all commentaries, our response specifically focuses on the following issues: How can the hypothesis of emotional distancing (qua art framing) be compatible with stipulating high levels of felt negative emotions in art reception? Which concept of altogether pleasurable mixed emotions does our model involve? Can mechanisms of predictive coding, social sharing, and immersion enhance the power of our model?

Processing of hierarchical syntactic structure in music.

Hierarchical structure with nested nonlocal dependencies is a key feature of human language and can be identified theoretically in most pieces of tonal music. However, previous studies have argued against the perception of such structures in music. Here, we show processing of nonlocal dependencies in music. We presented chorales by J. S. Bach and modified versions in which the hierarchical structure was rendered irregular whereas the local structure was kept intact. Brain electric responses differed between regular and irregular hierarchical structures, in both musicians and nonmusicians. This finding indicates that, when listening to music, humans apply cognitive processes that are capable of dealing with long-distance dependencies resulting from hierarchically organized syntactic structures. Our results reveal that a brain mechanism fundamental for syntactic processing is engaged during the perception of music, indicating that processing of hierarchical structure with nested nonlocal dependencies is not just a key component of human language, but a multidomain capacity of human cognition.

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.

Superficial amygdala and hippocampal activity during affective music listening observed at 3 T but not 1.5 T fMRI.

The purpose of this study was to compare 3 T and 1.5 T fMRI results during emotional music listening. Stimuli comprised of psychoacoustically balanced instrumental musical pieces, with three different affective expressions (fear, neutral, joy). Participants (N=32) were split into two groups, one subjected to fMRI scanning using 3 T and another group scanned using 1.5 T. Whole brain t-tests (corrected for multiple comparisons) compared joy and fear in each of the two groups. The 3 T group showed significant activity differences between joy and fear localized in bilateral superficial amygdala, bilateral hippocampus and bilateral auditory cortex. The 1.5 T group showed significant activity differences between joy and fear localized in bilateral auditory cortex and cuneus. This is the first study to compare results obtained under different field strengths with regard to affective processes elicited by means of auditory/musical stimulation. The findings raise concern over false negatives in the superficial amygdala and hippocampus in affective studies conducted under 1.5 T and caution that imaging improvements due to increasing magnetic field strength can be influenced by region-specific characteristics.

Differential effects of early life stress on hippocampus and amygdala volume as a function of emotional abilities.

Early life stress (ELS) is known to have considerable influence on brain development and affective functioning. Previous studies in clinical populations have shown that hippocampus and amygdala, two central structures of limbic emotion processing circuits, are predominantly affected by early stress exposure. Given the inconsistent findings on ELS-related effects in healthy populations and the associations of ELS and affective functioning, the question arises which additional emotion-relevant variables need to be considered to better understand the effects of ELS. We, therefore, investigated the volume of hippocampus and amygdala in 25 high alexithymic (h-ALEX) and 25 low alexithymic (l-ALEX) individuals, which were matched with regard to ELS, but significantly differed in their degree of emotional functioning. Volumetric analyses were performed using FSL-FIRST, a method to automatically segment subcortical structures on T1-weighted magnetic resonance images. Alexithymia was assessed using the Toronto Alexithymia Scale and Bermond-Vorst Alexithymia Questionnaire. ELS was assessed by Childhood Trauma Questionnaire (CTQ) and Early Trauma Inventory. Our data showed that ELS was negatively associated with right hippocampus volume in h-ALEX individuals, while there was no such association in the l-ALEX group. Furthermore, ELS was positively associated with left amygdala volume in l-ALEX individuals, but not in individuals with high levels of alexithymia. The present study emphasizes a substantial relationship between intrapersonal factors, such as alexithymia and neural alterations related to the experience of ELS. Longitudinal study designs are necessary to pursue the question of how emotional abilities interact with individual adaptations to early stress exposure on the neural level.

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.

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'.

The roles of superficial amygdala and auditory cortex in music-evoked fear and joy.

This study investigates neural correlates of music-evoked fear and joy with fMRI. Studies on neural correlates of music-evoked fear are scant, and there are only a few studies on neural correlates of joy in general. Eighteen individuals listened to excerpts of fear-evoking, joy-evoking, as well as neutral music and rated their own emotional state in terms of valence, arousal, fear, and joy. Results show that BOLD signal intensity increased during joy, and decreased during fear (compared to the neutral condition) in bilateral auditory cortex (AC) and bilateral superficial amygdala (SF). In the right primary somatosensory cortex (area 3b) BOLD signals increased during exposure to fear-evoking music. While emotion-specific activity in AC increased with increasing duration of each trial, SF responded phasically in the beginning of the stimulus, and then SF activity declined. Psychophysiological Interaction (PPI) analysis revealed extensive emotion-specific functional connectivity of AC with insula, cingulate cortex, as well as with visual, and parietal attentional structures. These findings show that the auditory cortex functions as a central hub of an affective-attentional network that is more extensive than previously believed. PPI analyses also showed functional connectivity of SF with AC during the joy condition, taken to reflect that SF is sensitive to social signals with positive valence. During fear music, SF showed functional connectivity with visual cortex and area 7 of the superior parietal lobule, taken to reflect increased visual alertness and an involuntary shift of attention during the perception of auditory signals of danger.

Co-localizing linguistic and musical syntax with intracranial EEG.

Despite general agreement on shared syntactic resources in music and language, the neuroanatomical underpinnings of this overlap remain largely unexplored. While previous studies mainly considered frontal areas as supramodal grammar processors, the domain-general syntactic role of temporal areas has been so far neglected. Here we capitalized on the excellent spatial and temporal resolution of subdural EEG recordings to co-localize low-level syntactic processes in music and language in the temporal lobe in a within-subject design. We used Brain Surface Current Density mapping to localize and compare neural generators of the early negativities evoked by violations of phrase structure grammar in both music and spoken language. The results show that the processing of syntactic violations relies in both domains on bilateral temporo-fronto-parietal neural networks. We found considerable overlap of these networks in the superior temporal lobe, but also differences in the hemispheric timing and relative weighting of their fronto-temporal constituents. While alluding to the dissimilarity in how shared neural resources may be configured depending on the musical or linguistic nature of the perceived stimulus, the combined data lend support for a co-localization of early musical and linguistic syntax processing in the temporal lobe.

Auditory stroop and absolute pitch: an fMRI study.

To date, the underlying cognitive and neural mechanisms of absolute pitch (AP) have remained elusive. In the present fMRI study, we investigated verbal and tonal perception and working memory in musicians with and without absolute pitch. Stimuli were sine wave tones and syllables (names of the scale tones) presented simultaneously. Participants listened to sequences of five stimuli, and then rehearsed internally either the syllables or the tones. Finally participants indicated whether a test stimulus had been presented during the sequence. For an auditory stroop task, half of the tonal sequences were congruent (frequencies of tones corresponded to syllables which were the names of the scale tones) and half were incongruent (frequencies of tones did not correspond to syllables). Results indicate that first, verbal and tonal perception overlap strongly in the left superior temporal gyrus/sulcus (STG/STS) in AP musicians only. Second, AP is associated with the categorical perception of tones. Third, the left STG/STS is activated in AP musicians only for the detection of verbal-tonal incongruencies in the auditory stroop task. Finally, verbal labelling of tones in AP musicians seems to be automatic. Overall, a unique feature of AP appears to be the similarity between verbal and tonal perception.

Functional specializations for music processing in the human newborn brain.

In adults, specific neural systems with right-hemispheric weighting are necessary to process pitch, melody, and harmony as well as structure and meaning emerging from musical sequences. It is not known to what extent the specialization of these systems results from long-term exposure to music or from neurobiological constraints. One way to address this question is to examine how these systems function at birth, when auditory experience is minimal. We used functional MRI to measure brain activity in 1- to 3-day-old newborns while they heard excerpts of Western tonal music and altered versions of the same excerpts. Altered versions either included changes of the tonal key or were permanently dissonant. Music evoked predominantly right-hemispheric activations in primary and higher order auditory cortex. During presentation of the altered excerpts, hemodynamic responses were significantly reduced in the right auditory cortex, and activations emerged in the left inferior frontal cortex and limbic structures. These results demonstrate that the infant brain shows a hemispheric specialization in processing music as early as the first postnatal hours. Results also indicate that the neural architecture underlying music processing in newborns is sensitive to changes in tonal key as well as to differences in consonance and dissonance.

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.

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.