The neuro-oscillatory profiles of static and dynamic music-induced visual imagery.

Visual imagery, i.e., seeing in the absence of the corresponding retinal input, has been linked to visual and motor processing areas of the brain. Music listening provides an ideal vehicle for exploring the neural correlates of visual imagery because it has been shown to reliably induce a broad variety of content, ranging from abstract shapes to dynamic scenes. Forty-two participants listened with closed eyes to twenty-four excerpts of music, while a 15-channel EEG was recorded, and, after each excerpt, rated the extent to which they experienced static and dynamic visual imagery. Our results show both static and dynamic imagery to be associated with posterior alpha suppression (especially in lower alpha) early in the onset of music listening, while static imagery was associated with an additional alpha enhancement later in the listening experience. With regard to the beta band, our results demonstrate beta enhancement to static imagery, but first beta suppression before enhancement in response to dynamic imagery. We also observed a positive association, early in the listening experience, between gamma power and dynamic imagery ratings that was not present for static imagery ratings. Finally, we offer evidence that musical training may selectively drive effects found with respect to static and dynamic imagery and alpha, beta, and gamma band oscillations. Taken together, our results show the promise of using music listening as an effective stimulus for examining the neural correlates of visual imagery and its contents. Our study also highlights the relevance of future work seeking to study the temporal dynamics of music-induced visual imagery.

A model of time-varying music engagement.

The current paper offers a model of time-varying music engagement, defined as changes in curiosity, attention and positive valence, as music unfolds over time. First, we present research (including new data) showing that listeners tend to allocate attention to music in a manner that is guided by both features of the music and listeners' individual differences. Next, we review relevant predictive processing literature before using this body of work to inform our model. In brief, we propose that music engagement, over the course of an extended listening episode, may constitute several cycles of curiosity, attention and positive valence that are interspersed with moments of mind-wandering. Further, we suggest that refocusing on music after an episode of mind-wandering can be due to triggers in the music or, conversely, mental action that occurs when the listener realizes they are mind-wandering. Finally, we argue that factors that modulate both overall levels of music engagement and how it changes over time include music complexity, listener background and the listening context. Our paper highlights how music can be used to provide insights into the temporal dynamics of attention and into how curiosity might emerge in everyday contexts. This article is part of the theme issue 'Art, aesthetics and predictive processing: theoretical and empirical perspectives'.

Music listening evokes story-like visual imagery with both idiosyncratic and shared content.

There is growing evidence that music can induce a wide range of visual imagery. To date, however, there have been few thorough investigations into the specific content of music-induced visual imagery, and whether listeners exhibit consistency within themselves and with one another regarding their visual imagery content. We recruited an online sample (N = 353) who listened to three orchestral film music excerpts representing happy, tender, and fearful emotions. For each excerpt, listeners rated how much visual imagery they were experiencing and how vivid it was, their liking of and felt emotional intensity in response to the excerpt, and, finally, described the content of any visual imagery they may have been experiencing. Further, they completed items assessing a number of individual differences including musical training and general visual imagery ability. Of the initial sample, 254 respondents completed the survey again three weeks later. A thematic analysis of the content descriptions revealed three higher-order themes of prominent visual imagery experiences: Storytelling (imagined locations, characters, actions, etc.), Associations (emotional experiences, abstract thoughts, and memories), and References (origins of the visual imagery, e.g., film and TV). Although listeners demonstrated relatively low visual imagery consistency with each other, levels were higher when considering visual imagery content within individuals across timepoints. Our findings corroborate past literature regarding music's capacity to encourage narrative engagement. It, however, extends it (a) to show that such engagement is highly visual and contains other types of imagery to a lesser extent, (b) to indicate the idiosyncratic tendencies of listeners' imagery consistency, and (c) to reveal key factors influencing consistency levels (e.g., vividness of visual imagery and emotional intensity ratings in response to music). Further implications are discussed in relation to visual imagery's purported involvement in music-induced emotions and aesthetic appeal.

Investigating the interaction of pleasantness and arousal and the role of aesthetic emotions on episodic memory using a musical what-where-when paradigm.

Recent evidence suggests that the presence of music experienced as pleasant positively influences episodic memory (EM) encoding. However, it is unclear whether this impact of pleasant music holds regardless of how arousing the music is, and what influence, if any, music-induced aesthetic emotions have. Furthermore, most music EM studies have used verbal or facial memory tasks limiting the generalisability of these findings to everyday EMs with spatiotemporal richness. The current study used an online what-where-when paradigm to assess music's influence on EM encoding in a rich spatiotemporal environment. 105 participants carried out the what-where-when task in the presence of either silence or one of four musical stimuli falling into the four corners of the 2-D circumplex emotion model. We observed an interaction effect between the pleasantness and arousingness of music stimuli, whereby for pleasant stimuli, the low arousing excerpt was associated with better recall performance compared to the high arousing excerpt. We also observed that, across all excerpts, experience of negative aesthetic emotions was associated with compromised recall performance. Together, our results confirm the deleterious influence arousing stimuli can have on memory and support the notion that aesthetic-emotional experience of music can influence how memories are encoded in everyday life.

Hierarchically nested networks optimize the analysis of audiovisual speech.

In conversational settings, seeing the speaker's face elicits internal predictions about the upcoming acoustic utterance. Understanding how the listener's cortical dynamics tune to the temporal statistics of audiovisual (AV) speech is thus essential. Using magnetoencephalography, we explored how large-scale frequency-specific dynamics of human brain activity adapt to AV speech delays. First, we show that the amplitude of phase-locked responses parametrically decreases with natural AV speech synchrony, a pattern that is consistent with predictive coding. Second, we show that the temporal statistics of AV speech affect large-scale oscillatory networks at multiple spatial and temporal resolutions. We demonstrate a spatial nestedness of oscillatory networks during the processing of AV speech: these oscillatory hierarchies are such that high-frequency activity (beta, gamma) is contingent on the phase response of low-frequency (delta, theta) networks. Our findings suggest that the endogenous temporal multiplexing of speech processing confers adaptability within the temporal regimes that are essential for speech comprehension.

In touch: Cardiac and respiratory patterns synchronize during ensemble singing with physical contact.

Musical ensemble performances provide an ideal environment to gain knowledge about complex human interactions. Network structures of synchronization can reflect specific roles of individual performers on the one hand and a higher level of organization of all performers as a superordinate system on the other. This study builds on research on joint singing, using hyperscanning of respiration and heart rate variability (HRV) from eight professional singers. Singers performed polyphonic music, distributing their breathing within the same voice and singing without and with physical contact: that is touching each other's shoulder or waist. The idea of singing with touch was motivated by historical depictions of ensemble performances that showed singers touching each other. It raises the question of the potential benefit of touch for group performances. From a psycho-physiological point of view, physical contact should increase the synchronization of singing coordination. The results confirm previous findings on synchronization of respiration and HRV during choir singing and extend those findings to a non-homophonic musical repertoire while also revealing an increase in synchronization in respiration during physical contact. These effects were significant across different frequency ranges. The effect of physical contact was stronger when all singers were singing in comparison to the partial ensemble. Importantly, the synchronization could not be fully explained by the singing action (i.e., singing the same voice, or singing vs. listening) or by the standing position or touch. This finding suggests a higher level of organization of all singers, forming a superordinate system.

Atonal Music as a Model for Investigating Exploratory Behavior.

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.

Prediction under uncertainty: Dissociating sensory from cognitive expectations in highly uncertain musical contexts.

Predictive models in the brain rely on the continuous extraction of regularities from the environment. These models are thought to be updated by novel information, as reflected in prediction error responses such as the mismatch negativity (MMN). However, although in real life individuals often face situations in which uncertainty prevails, it remains unclear whether and how predictive models emerge in high-uncertainty contexts. Recent research suggests that uncertainty affects the magnitude of MMN responses in the context of music listening. However, musical predictions are typically studied with MMN stimulation paradigms based on Western tonal music, which are characterized by relatively high predictability. Hence, we developed an MMN paradigm to investigate how the high uncertainty of atonal music modulates predictive processes as indexed by the MMN and behavior. Using MEG in a group of 20 subjects without musical training, we demonstrate that the magnetic MMN in response to pitch, intensity, timbre, and location deviants is evoked in both tonal and atonal melodies, with no significant differences between conditions. In contrast, in a separate behavioral experiment involving 39 non-musicians, participants detected pitch deviants more accurately and rated confidence higher in the tonal than in the atonal musical context. These results indicate that contextual tonal uncertainty modulates processing stages in which conscious awareness is involved, although deviants robustly elicit low-level pre-attentive responses such as the MMN. The achievement of robust MMN responses, despite high tonal uncertainty, is relevant for future studies comparing groups of listeners' MMN responses to increasingly ecological music stimuli.

Locus of emotion influences psychophysiological reactions to music.

It is now widely accepted that the perception of emotional expression in music can be vastly different from the feelings evoked by it. However, less understood is how the locus of emotion affects the experience of music, that is how the act of perceiving the emotion in music compares with the act of assessing the emotion induced in the listener by the music. In the current study, we compared these two emotion loci based on the psychophysiological response of 40 participants listening to 32 musical excerpts taken from movie soundtracks. Facial electromyography, skin conductance, respiration and heart rate were continuously measured while participants were required to assess either the emotion expressed by, or the emotion they felt in response to the music. Using linear mixed effects models, we found a higher mean response in psychophysiological measures for the "perceived" than the "felt" task. This result suggested that the focus on one's self distracts from the music, leading to weaker bodily reactions during the "felt" task. In contrast, paying attention to the expression of the music and consequently to changes in timbre, loudness and harmonic progression enhances bodily reactions. This study has methodological implications for emotion induction research using psychophysiology and the conceptualization of emotion loci. Firstly, different tasks can elicit different psychophysiological responses to the same stimulus and secondly, both tasks elicit bodily responses to music. The latter finding questions the possibility of a listener taking on a purely cognitive mode when evaluating emotion expression.

Pupil responses to pitch deviants reflect predictability of melodic sequences.

Humans automatically detect events that, in deviating from their expectations, may signal prediction failure and a need to reorient behaviour. The pupil dilation response (PDR) to violations has been associated with subcortical signals of arousal and prediction resetting. However, it is unclear how the context in which a deviant occurs affects the size of the PDR. Using ecological musical stimuli that we characterised using a computational model, we showed that the PDR to pitch deviants is sensitive to contextual uncertainty (quantified as entropy), whereby the PDR was greater in low than high entropy contexts. The PDR was also positively correlated with the unexpectedness of notes. No effects of music expertise were found, suggesting a ceiling effect due to enculturation. These results show that the same sudden environmental change can lead to differing arousal levels depending on contextual factors, providing evidence for a sensitivity of the PDR to long-term context.

Intracranial Recordings and Computational Modeling of Music Reveal the Time Course of Prediction Error Signaling in Frontal and Temporal Cortices.

Prediction is held to be a fundamental process underpinning perception, action, and cognition. To examine the time course of prediction error signaling, we recorded intracranial EEG activity from nine presurgical epileptic patients while they listened to melodies whose information theoretical predictability had been characterized using a computational model. We examined oscillatory activity in the superior temporal gyrus (STG), the middle temporal gyrus (MTG), and the pars orbitalis of the inferior frontal gyrus, lateral cortical areas previously implicated in auditory predictive processing. We also examined activity in anterior cingulate gyrus (ACG), insula, and amygdala to determine whether signatures of prediction error signaling may also be observable in these subcortical areas. Our results demonstrate that the information content (a measure of unexpectedness) of musical notes modulates the amplitude of low-frequency oscillatory activity (theta to beta power) in bilateral STG and right MTG from within 100 and 200 msec of note onset, respectively. Our results also show this cortical activity to be accompanied by low-frequency oscillatory modulation in ACG and insula-areas previously associated with mediating physiological arousal. Finally, we showed that modulation of low-frequency activity is followed by that of high-frequency (gamma) power from approximately 200 msec in the STG, between 300 and 400 msec in the left insula, and between 400 and 500 msec in the ACG. We discuss these results with respect to models of neural processing that emphasize gamma activity as an index of prediction error signaling and highlight the usefulness of musical stimuli in revealing the wide-reaching neural consequences of predictive processing.

Atonal Music: Can Uncertainty Lead to Pleasure?

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.

Personalized translational epilepsy research - Novel approaches and future perspectives: Part II: Experimental and translational approaches.

Despite the availability of more than 15 new "antiepileptic drugs", the proportion of patients with pharmacoresistant epilepsy has remained constant at about 20-30%. Furthermore, no disease-modifying treatments shown to prevent the development of epilepsy following an initial precipitating brain injury or to reverse established epilepsy have been identified to date. This is likely in part due to the polyetiologic nature of epilepsy, which in turn requires personalized medicine approaches. Recent advances in imaging, pathology, genetics, and epigenetics have led to new pathophysiological concepts and the identification of monogenic causes of epilepsy. In the context of these advances, the First International Symposium on Personalized Translational Epilepsy Research (1st ISymPTER) was held in Frankfurt on September 8, 2016, to discuss novel approaches and future perspectives for personalized translational research. These included new developments and ideas in a range of experimental and clinical areas such as deep phenotyping, quantitative brain imaging, EEG/MEG-based analysis of network dysfunction, tissue-based translational studies, innate immunity mechanisms, microRNA as treatment targets, functional characterization of genetic variants in human cell models and rodent organotypic slice cultures, personalized treatment approaches for monogenic epilepsies, blood-brain barrier dysfunction, therapeutic focal tissue modification, computational modeling for target and biomarker identification, and cost analysis in (monogenic) disease and its treatment. This report on the meeting proceedings is aimed at stimulating much needed investments of time and resources in personalized translational epilepsy research. This Part II includes the experimental and translational approaches and a discussion of the future perspectives, while the diagnostic methods, EEG network analysis, biomarkers, and personalized treatment approaches were addressed in Part I [1].

Personalized translational epilepsy research - Novel approaches and future perspectives: Part I: Clinical and network analysis approaches.

Despite the availability of more than 15 new "antiepileptic drugs", the proportion of patients with pharmacoresistant epilepsy has remained constant at about 20-30%. Furthermore, no disease-modifying treatments shown to prevent the development of epilepsy following an initial precipitating brain injury or to reverse established epilepsy have been identified to date. This is likely in part due to the polyetiologic nature of epilepsy, which in turn requires personalized medicine approaches. Recent advances in imaging, pathology, genetics and epigenetics have led to new pathophysiological concepts and the identification of monogenic causes of epilepsy. In the context of these advances, the First International Symposium on Personalized Translational Epilepsy Research (1st ISymPTER) was held in Frankfurt on September 8, 2016, to discuss novel approaches and future perspectives for personalized translational research. These included new developments and ideas in a range of experimental and clinical areas such as deep phenotyping, quantitative brain imaging, EEG/MEG-based analysis of network dysfunction, tissue-based translational studies, innate immunity mechanisms, microRNA as treatment targets, functional characterization of genetic variants in human cell models and rodent organotypic slice cultures, personalized treatment approaches for monogenic epilepsies, blood-brain barrier dysfunction, therapeutic focal tissue modification, computational modeling for target and biomarker identification, and cost analysis in (monogenic) disease and its treatment. This report on the meeting proceedings is aimed at stimulating much needed investments of time and resources in personalized translational epilepsy research. Part I includes the clinical phenotyping and diagnostic methods, EEG network-analysis, biomarkers, and personalized treatment approaches. In Part II, experimental and translational approaches will be discussed (Bauer et al., 2017) [1].

Basic, specific, mechanistic? Conceptualizing musical emotions in the brain.

The number of studies investigating music processing in the human brain continues to increase, with a large proportion of them focussing on the correlates of so-called musical emotions. The current Review highlights the recent development whereby such studies are no longer concerned only with basic emotions such as happiness and sadness but also with so-called music-specific or "aesthetic" ones such as nostalgia and wonder. It also highlights how mechanisms such as expectancy and empathy, which are seen as inducing musical emotions, are enjoying ever-increasing investigation and substantiation with physiological and neuroimaging methods. It is proposed that a combination of these approaches, namely, investigation of the precise mechanisms through which so-called music-specific or aesthetic emotions may arise, will provide the most important advances for our understanding of the unique nature of musical experience.

Music and literature: are there shared empathy and predictive mechanisms underlying their affective impact?

It has been suggested that music and language had a shared evolutionary precursor before becoming mainly responsible for the communication of emotive and referential meaning respectively. However, emphasis on potential differences between music and language may discourage a consideration of the commonalities that music and literature share. Indeed, one possibility is that common mechanisms underlie their affective impact, and the current paper carefully reviews relevant neuroscientific findings to examine such a prospect. First and foremost, it will be demonstrated that considerable evidence of a common role of empathy and predictive processes now exists for the two domains. However, it will also be noted that an important open question remains: namely, whether the mechanisms underlying the subjective experience of uncertainty differ between the two domains with respect to recruitment of phylogenetically ancient emotion areas. It will be concluded that a comparative approach may not only help to reveal general mechanisms underlying our responses to music and literature, but may also help us better understand any idiosyncrasies in their capacity for affective impact.

Exploring the musical taste of expert listeners: musicology students reveal tendency toward omnivorous taste.

Musicology students are engaged with music on an academic level and usually have an extensive musical background. They have a considerable knowledge of music history and theory and listening to music may be regarded as one of their primary occupations. Taken together, these factors qualify them as ≫expert listeners≪, who may be expected to exhibit a specific profile of musical taste: interest in a broad range of musical styles combined with a greater appreciation of ≫sophisticated≪ styles. The current study examined the musical taste of musicology students as compared to a control student group. Participants (n = 1003) completed an online survey regarding the frequency with which they listened to 22 musical styles. A factor analysis revealed six underlying dimensions of musical taste. A hierarchical cluster analysis then grouped all participants, regardless of their status, according to their similarity on these dimensions. The employed exploratory approach was expected to reveal potential differences between musicology students and controls. A three-cluster solution was obtained. Comparisons of the clusters in terms of musical taste revealed differences in the listening frequency and variety of appreciated music styles: the first cluster (51% musicology students/27% controls) showed the greatest musical engagement across all dimensions although with a tendency toward ≫sophisticated≪ musical styles. The second cluster (36% musicology students/46% controls) exhibited an interest in ≫conventional≪ music, while the third cluster (13% musicology students/27% controls) showed a strong liking of rock music. The results provide some support for the notion of specific tendencies in the musical taste of musicology students and the contribution of familiarity and knowledge toward musical omnivorousness. Further differences between the clusters in terms of social, personality, and sociodemographic factors are discussed.

Dopamine and epistemic curiosity in music listening.

Elucidating the cognitive, affective, and reward processes that take place during music listening is the aim of a growing number of researchers. Several authors have used the Bayesian brain framework and existing models of reward to interpret neural activity observed during musical listening. The claims from Friston and colleagues regarding the role of dopamine, as well as the demonstration that salience-seeking behavior naturally emerges from minimizing free energy, will be of potential interest to those seeking to understand the general principles underlying our motivation to hear music.

Destination memory and familiarity: better memory for conversations with Elvis Presley than with unknown people.

BACKGROUND AND AIMS: Familiarity is assumed to exert a beneficial effect on memory in older adults. Our paper investigated this issue specifically for destination memory, that is, memory of the destination of previously relayed information. METHODS: Young and older adults were told familiar (Experiment 1) and unfamiliar (Experiment 2) proverbs associated with pictures depicting faces of celebrities (e.g., Elvis Presley) or unknown people, with a specific proverb assigned to each face. In a later recognition task, participants were presented with the previously exposed proverb-face pairs and for each pair had to decide whether they had previously relayed the given proverb to the given face. RESULTS: In general, destination performance was found to be higher for familiar than for unfamiliar faces. However while there was no difference between the two groups when the proverbs being relayed were unfamiliar, the advantage of face familiarity on destination memory was present only for older adults when the proverbs being relayed were familiar. DISCUSSION AND CONCLUSIONS: Our results show that destination memory in older adults is sensitive to familiarity of both destination and output information.

An Intracranial EEG Study of the Neural Dynamics of Musical Valence Processing.

The processing of valence is known to recruit the amygdala, orbitofrontal cortex, and relevant sensory areas. However, how these regions interact remains unclear. We recorded cortical electrical activity from 7 epileptic patients implanted with depth electrodes for presurgical evaluation while they listened to positively and negatively valenced musical chords. Time-frequency analysis suggested a specific role of the orbitofrontal cortex in the processing of positively valenced stimuli while, most importantly, Granger causality analysis revealed that the amygdala tends to drive both the orbitofrontal cortex and the auditory cortex in theta and alpha frequency bands, during the processing of valenced stimuli. Results from the current study show the amygdala to be a critical hub in the emotion processing network: specifically one that influences not only the higher order areas involved in the evaluation of a stimulus's emotional value but also the sensory cortical areas involved in the processing of its low-level acoustic features.