Correction: AVPR1a and SLC6A4 Gene Polymorphisms Are Associated with Creative Dance Performance.

[This corrects the article DOI: 10.1371/journal.pgen.0010042.].

Robust inter-subject audiovisual decoding in functional magnetic resonance imaging using high-dimensional regression.

Major methodological advancements have been recently made in the field of neural decoding, which is concerned with the reconstruction of mental content from neuroimaging measures. However, in the absence of a large-scale examination of the validity of the decoding models across subjects and content, the extent to which these models can be generalized is not clear. This study addresses the challenge of producing generalizable decoding models, which allow the reconstruction of perceived audiovisual features from human magnetic resonance imaging (fMRI) data without prior training of the algorithm on the decoded content. We applied an adapted version of kernel ridge regression combined with temporal optimization on data acquired during film viewing (234 runs) to generate standardized brain models for sound loudness, speech presence, perceived motion, face-to-frame ratio, lightness, and color brightness. The prediction accuracies were tested on data collected from different subjects watching other movies mainly in another scanner. Substantial and significant (QFDR<0.05) correlations between the reconstructed and the original descriptors were found for the first three features (loudness, speech, and motion) in all of the 9 test movies (R¯=0.62, R¯ = 0.60, R¯ = 0.60, respectively) with high reproducibility of the predictors across subjects. The face ratio model produced significant correlations in 7 out of 8 movies (R¯=0.56). The lightness and brightness models did not show robustness (R¯=0.23, R¯ = 0). Further analysis of additional data (95 runs) indicated that loudness reconstruction veridicality can consistently reveal relevant group differences in musical experience. The findings point to the validity and generalizability of our loudness, speech, motion, and face ratio models for complex cinematic stimuli (as well as for music in the case of loudness). While future research should further validate these models using controlled stimuli and explore the feasibility of extracting more complex models via this method, the reliability of our results indicates the potential usefulness of the approach and the resulting models in basic scientific and diagnostic contexts.

Common modulation of limbic network activation underlies musical emotions as they unfold.

Music is a powerful means for communicating emotions among individuals. Here we reveal that this continuous stream of affective information is commonly represented in the brains of different listeners and that particular musical attributes mediate this link. We examined participants' brain responses to two naturalistic musical pieces using functional Magnetic Resonance imaging (fMRI). Following scanning, as participants listened to the musical pieces for a second time, they continuously indicated their emotional experience on scales of valence and arousal. These continuous reports were used along with a detailed annotation of the musical features, to predict a novel index of Dynamic Common Activation (DCA) derived from ten large-scale data-driven functional networks. We found an association between the unfolding music-induced emotionality and the DCA modulation within a vast network of limbic regions. The limbic-DCA modulation further corresponded with continuous changes in two temporal musical features: beat-strength and tempo. Remarkably, this "collective limbic sensitivity" to temporal features was found to mediate the link between limbic-DCA and the reported emotionality. An additional association with the emotional experience was found in a left fronto-parietal network, but only among a sub-group of participants with a high level of musical experience (>5years). These findings may indicate two processing-levels underlying the unfolding of common music emotionality; (1) a widely shared core-affective process that is confined to a limbic network and mediated by temporal regularities in music and (2) an experience based process that is rooted in a left fronto-parietal network that may involve functioning of the 'mirror-neuron system'.

Spatial vision is superior in musicians when memory plays a role.

Musicians' perceptual advantage in the acoustic domain is well established. Recent studies show that musicians' verbal working memory is also superior. Additionally, some studies report that musicians' visuospatial skills are enhanced although others failed to find this enhancement. We now examined whether musicians' spatial vision is superior, and if so, whether this superiority reflects refined visual skills or a general superiority of working memory. We examined spatial frequency discrimination among musicians and nonmusician university students using two presentation conditions: simultaneous (spatial forced choice) and sequential (temporal forced choice). Musicians' performance was similar to that of nonmusicians in the simultaneous condition. However, their performance in the sequential condition was superior, suggesting an advantage only when stimuli need to be retained, i.e., working memory. Moreover, the two groups showed a different pattern of correlations: Musicians' visual thresholds were correlated, and neither was correlated with their verbal memory. By contrast, among nonmusicians, the visual thresholds were not correlated, but sequential thresholds were correlated with verbal memory scores, suggesting that a general working memory component limits their performance in this condition. We propose that musicians' superiority in spatial frequency discrimination reflects an advantage in a domain-general aspect of working memory rather than a general enhancement in spatial-visual skills.

Feeling the Beat: Temporal Predictability is Associated with Ongoing Changes in Music-Induced Pleasantness.

Music is a complex phenomenon that elicits a range of emotional responses, influenced by numerous variables, such as rhythm, melody and harmony. One interesting aspect of music is listeners' ability to predict its continuation as it unfolds - an inherent attribute hypothesized to contribute to our emotional response to music. In this study, we investigated this link by examining the relationship between temporal predictability - the ability to predict the timing of the next event - and the ongoing changes in music-induced pleasantness. Temporal predictability was operationalized as the degree to which taps of 20 musically trained participants, who tapped to the beat along three naturalistic and highly contrastive musical pieces, were aligned. We then examined the degree to which this measure could explain the ongoing emotional experience, as reflected in continuous measures of arousal and valence, in a separate group of 40 participants that listened to these pieces. Our findings reveal a positive correlation between fluctuations in reported valence and temporal predictability, even when controlling for a set of other musical features, in four out of five musical sections. The only exception being a lyrical slow section. These findings were further supported by a large online database of annotated musical emotions (n = 1780 songs), where a consistent and robust correlation between valence ratings and an automatically extracted feature of pulse clarity was demonstrated. Overall, our findings shed light on the significance of temporal predictability as a contributing factor to the hedonic experience of music, especially within the tempo range of salient beat perception.

The rediscovered motor-related area 55b emerges as a core hub of music perception.

Passive listening to music, without sound production or evident movement, is long known to activate motor control regions. Nevertheless, the exact neuroanatomical correlates of the auditory-motor association and its underlying neural mechanisms have not been fully determined. Here, based on a NeuroSynth meta-analysis and three original fMRI paradigms of music perception, we show that the long-ignored pre-motor region, area 55b, an anatomically unique and functionally intriguing region, is a core hub of music perception. Moreover, results of a brain-behavior correlation analysis implicate neural entrainment as the underlying mechanism of area 55b's contribution to music perception. In view of the current results and prior literature, area 55b is proposed as a keystone of sensorimotor integration, a fundamental brain machinery underlying simple to hierarchically complex behaviors. Refining the neuroanatomical and physiological understanding of sensorimotor integration is expected to have a major impact on various fields, from brain disorders to artificial general intelligence.

The Efficacy of Music for Emotional Wellbeing During the COVID-19 Lockdown in Spain: An Analysis of Personal and Context-Related Variables.

The strict lockdown experienced in Spain during March-June 2020 as a consequence of the COVID-19 crisis has led to strong negative emotions. Music can contribute to enhancing wellbeing, but the extent of this effect may be modulated by both personal and context-related variables. This study aimed to analyze the impact of the two types of variables on the perceived efficacy of musical behaviors to fulfill adults' emotional wellbeing-related goals during the lockdown established in Spain. Personal variables included age, gender, musical training, personality, resilience, and perception of music's importance. Contextual variables referred to living in a region with a high COVID-19 impact, perception of belonging to a risk group, being alone, having caring responsibilities during confinement, and amount of time of music listening as compared to prior to the crisis. The study was conducted retrospectively during August-December 2020, when the strict lockdown was over in Spain. An online survey was disseminated among the general population and groups of musicians, and the answers of 507 adults (from 18 years on, 73.9% females, 51.3% musically trained adults) were analyzed. Only personal, but not COVID-19 context-related variables, showed an impact on music's efficacy. The youngest age group of adults and those with musical training reported the highest efficacy of music for wellbeing enhancement, and music's importance was found to be the main significant predictor of music's perceived efficacy. Our findings suggest that the people who have been reported to be emotionally more vulnerable during the lockdown, due to either a strong impact on their daily lives or their lower resilience, perceive a higher benefit from musical behaviors. Being musically trained, even for a small number of years, also leads to a perception of higher efficacy of music for the achievement of emotional wellbeing goals. However, this effect is explained by the musically trained individuals' higher perception of music's importance. Although musical behaviors can be generally considered as important for wellbeing enhancement, our study highlights who are the potential individuals who could benefit the most from music-related activities for obtaining better levels of wellbeing, at least within the current context of the COVID-19 crisis.

"Help! I Need Somebody": Music as a Global Resource for Obtaining Wellbeing Goals in Times of Crisis.

Music can reduce stress and anxiety, enhance positive mood, and facilitate social bonding. However, little is known about the role of music and related personal or cultural (individualistic vs. collectivistic) variables in maintaining wellbeing during times of stress and social isolation as imposed by the COVID-19 crisis. In an online questionnaire, administered in 11 countries (Argentina, Brazil, China, Colombia, Italy, Mexico, the Netherlands, Norway, Spain, the UK, and USA, N = 5,619), participants rated the relevance of wellbeing goals during the pandemic, and the effectiveness of different activities in obtaining these goals. Music was found to be the most effective activity for three out of five wellbeing goals: enjoyment, venting negative emotions, and self-connection. For diversion, music was equally good as entertainment, while it was second best to create a sense of togetherness, after socialization. This result was evident across different countries and gender, with minor effects of age on specific goals, and a clear effect of the importance of music in people's lives. Cultural effects were generally small and surfaced mainly in the use of music to obtain a sense of togetherness. Interestingly, culture moderated the use of negatively valenced and nostalgic music for those higher in distress.

Surprise-related activation in the nucleus accumbens interacts with music-induced pleasantness.

How can music-merely a stream of sounds-be enjoyable for so many people? Recent accounts of this phenomenon are inspired by predictive coding models, hypothesizing that both confirmation and violations of musical expectations associate with the hedonic response to music via recruitment of the mesolimbic system and its connections with the auditory cortex. Here we provide support for this model, by revealing associations of music-induced pleasantness with musical surprises in the activity and connectivity patterns of the nucleus accumbens (NAcc)-a central component of the mesolimbic system. We examined neurobehavioral responses to surprises in three naturalistic musical pieces using fMRI and subjective ratings of valence and arousal. Surprises were associated with changes in reported valence and arousal, as well as with enhanced activations in the auditory cortex, insula and ventral striatum, relative to unsurprising events. Importantly, we found that surprise-related activation in the NAcc was more pronounced among individuals who experienced greater music-induced pleasantness. These participants also exhibited stronger surprise-related NAcc-auditory cortex connectivity during the most pleasant piece, relative to participants who found the music less pleasant. These findings provide a novel demonstration of a direct link between musical surprises, NAcc activation and music-induced pleasantness.

Is there a prediction network? Meta-analytic evidence for a cortical-subcortical network likely subserving prediction.

Predictive coding is an increasingly influential and ambitious concept in neuroscience viewing the brain as a 'hypothesis testing machine' that constantly strives to minimize prediction error, the gap between its predictions and the actual sensory input. Despite the invaluable contribution of this framework to the formulation of brain function, its neuroanatomical foundations have not been fully defined. To address this gap, we conducted activation likelihood estimation (ALE) meta-analysis of 39 neuroimaging studies of three functional domains (action perception, language and music) inherently involving prediction. The ALE analysis revealed a widely distributed brain network encompassing regions within the inferior and middle frontal gyri, anterior insula, premotor cortex, pre-supplementary motor area, temporoparietal junction, striatum, thalamus/subthalamus and the cerebellum. This network is proposed to subserve domain-general prediction and its relevance to motor control, attention, implicit learning and social cognition is discussed in light of the predictive coding scheme. Better understanding of the presented network may help advance treatments of neuropsychiatric conditions related to aberrant prediction processing and promote cognitive enhancement in healthy individuals.

AVPR1a and SLC6A4 gene polymorphisms are associated with creative dance performance.

Dancing, which is integrally related to music, likely has its origins close to the birth of Homo sapiens, and throughout our history, dancing has been universally practiced in all societies. We hypothesized that there are differences among individuals in aptitude, propensity, and need for dancing that may partially be based on differences in common genetic polymorphisms. Identifying such differences may lead to an understanding of the neurobiological basis of one of mankind's most universal and appealing behavioral traits--dancing. In the current study, 85 current performing dancers and their parents were genotyped for the serotonin transporter (SLC6A4: promoter region HTTLPR and intron 2 VNTR) and the arginine vasopressin receptor 1a (AVPR1a: promoter microsatellites RS1 and RS3). We also genotyped 91 competitive athletes and a group of nondancers/nonathletes (n = 872 subjects from 414 families). Dancers scored higher on the Tellegen Absorption Scale, a questionnaire that correlates positively with spirituality and altered states of consciousness, as well as the Reward Dependence factor in Cloninger's Tridimensional Personality Questionnaire, a measure of need for social contact and openness to communication. Highly significant differences in AVPR1a haplotype frequencies (RS1 and RS3), especially when conditional on both SLC6A4 polymorphisms (HTTLPR and VNTR), were observed between dancers and athletes using the UNPHASED program package (Cocaphase: likelihood ratio test [LRS] = 89.23, p = 0.000044). Similar results were obtained when dancers were compared to nondancers/nonathletes (Cocaphase: LRS = 92.76, p = 0.000024). These results were confirmed using a robust family-based test (Tdtphase: LRS = 46.64, p = 0.010). Association was also observed between Tellegen Absorption Scale scores and AVPR1a (Qtdtphase: global chi-square = 26.53, p = 0.047), SLC6A4 haplotypes (Qtdtphase: chi-square = 2.363, p = 0.018), and AVPR1a conditional on SCL6A4 (Tdtphase: LRS = 250.44, p = 0.011). Similarly, significant association was observed between Tridimensional Personality Questionnaire Reward Dependence scores and AVPR1a RS1 (chi-square = 20.16, p = 0.01). Two-locus analysis (RS1 and RS3 conditional on HTTLPR and VNTR) was highly significant (LRS = 162.95, p = 0.001). Promoter repeat regions in the AVPR1a gene have been robustly demonstrated to play a role in molding a range of social behaviors in many vertebrates and, more recently, in humans. Additionally, serotonergic neurotransmission in some human studies appears to mediate human religious and spiritual experiences. We therefore hypothesize that the association between AVPR1a and SLC6A4 reflects the social communication, courtship, and spiritual facets of the dancing phenotype rather than other aspects of this complex phenotype, such as sensorimotor integration.

Brain responses to regular and octave-scrambled melodies: A case of predictive-coding?

Melody recognition is an online process of evaluating incoming information and comparing this information to an existing internal corpus, thereby reducing prediction error. The predictive-coding model postulates top-down control on sensory processing accompanying reduction in prediction error. To investigate the relevancy of this model to melody processing, the current study examined early magnetoencephalogram (MEG) auditory responses to familiar and unfamiliar melodies in 25 participants. The familiar melodies followed and primed an octave-scrambled version of the same melody. The retrograde version of theses melodies served as the unfamiliar control condition. Octave-transposed melodies were included to examine the influence of pitch representation (pitch-height/pitch-chroma representation) on brain responses to melody recognition. Results demonstrate a reduction of the M100 auditory response to familiar, as compared with unfamiliar, melodies regardless of their form of presentation (condensed vs. octave-scrambled). This trend appeared to begin after the third tone of the melody. An additional behavioral study with the same melody corpus showed a similar trend-namely, a significant difference between familiarity rating for familiar and unfamiliar melodies, beginning with the third tone of the melody. These results may indicate a top-down inhibition of early auditory responses to melodies that is influenced by pitch representation. (PsycINFO Database Record

The enigma of dyslexic musicians.

Musicians are known to have exceptional sensitivity to sounds, whereas poor phonological representations (or access to these representations) are considered a main characteristic of dyslexic individuals. Though these two characteristics refer to different abilities that are related to non-verbal and verbal skills respectively, the recent literature suggests that they are tightly related. However, there are informal reports of dyslexic musicians. To better understand this enigma, two groups of musicians were recruited, with and without a history of reading difficulties. The pattern of reading difficulties found among musicians was similar to that reported for non-musician dyslexics, though its magnitude was less severe. In contrast to non-musician dyslexics, their performance in pitch and interval discrimination, synchronous tapping and speech perception tasks, did not differ from the performance of their musician peers, and was superior to that of the general population. However, the auditory working memory scores of dyslexic musicians were consistently poor, including memory for rhythm, melody and speech sounds. Moreover, these abilities were inter-correlated, and highly correlated with their reading accuracy. These results point to a discrepancy between their perceptual and working memory skills rather than between sensitivity to speech and non-speech sounds. The results further suggest that in spite of intensive musical training, auditory working memory remains a bottleneck to the reading accuracy of dyslexic musicians.

Effects of arginine vasopressin on musical working memory.

Previous genetic studies showed an association between variations in the gene coding for the 1a receptor of the neuro-hormone arginine vasopressin (AVP) and musical working memory (WM). The current study set out to test the influence of intranasal administration (INA) of AVP on musical as compared to verbal WM using a double blind crossover (AVP-placebo) design. Two groups of 25 males were exposed to 20 IU of AVP in one session, and 20 IU of saline water (placebo) in a second session, 1 week apart. In each session subjects completed the tonal subtest from Gordon's "Musical Aptitude Profile," the interval subtest from the "Montreal Battery for Evaluation of Amusias (MBEA)," and the forward and backward digit span tests. Scores in the digit span tests were not influenced by AVP. In contrast, in the music tests there was an AVP effect. In the MBEA test, scores for the group receiving placebo in the first session (PV) were higher than for the group receiving vasopressin in the first session (VP) (p < 0.05) with no main Session effect nor Group × Session interaction. In the Gordon test there was a main Session effect (p < 0.05) with scores higher in the second as compared to the first session, a marginal main Group effect (p = 0.093) and a marginal Group × Session interaction (p = 0.88). In addition we found that the group that received AVP in the first session scored higher on scales indicative of happiness, and alertness on the positive and negative affect scale, (PANAS). Only in this group and only in the music test these scores were significantly correlated with memory scores. Together the results reflect a complex interaction between AVP, musical memory, arousal, and contextual effects such as session, and base levels of memory. The results are interpreted in light of music's universal use as a means to modulate arousal on the one hand, and AVP's influence on mood, arousal, and social interactions on the other.

Accuracy of pitch matching significantly improved by live voice model.

Singing is, undoubtedly, the most fundamental expression of our musical capacity, yet an estimated 10-15% of Western population sings "out-of-tune (OOT)." Previous research in children and adults suggests, albeit inconsistently, that imitating a human voice can improve pitch matching. In the present study, we focus on the potentially beneficial effects of the human voice and especially the live human voice. Eighteen participants varying in their singing abilities were required to imitate in singing a set of nine ascending and descending intervals presented to them in five different randomized blocked conditions: live piano, recorded piano, live voice using optimal voice production, recorded voice using optimal voice production, and recorded voice using artificial forced voice production. Pitch and interval matching in singing were much more accurate when participants repeated sung intervals as compared with intervals played to them on the piano. The advantage of the vocal over the piano stimuli was robust and emerged clearly regardless of whether piano tones were played live and in full view or were presented via recording. Live vocal stimuli elicited higher accuracy than recorded vocal stimuli, especially when the recorded vocal stimuli were produced in a forced vocal production. Remarkably, even those who would be considered OOT singers on the basis of their performance when repeating piano tones were able to pitch match live vocal sounds, with deviations well within the range of what is considered accurate singing (M=46.0, standard deviation=39.2 cents). In fact, those participants who were most OOT gained the most from the live voice model. Results are discussed in light of the dual auditory-motor encoding of pitch analogous to that found in speech.

Electrophysiological evidence for a two-stage process underlying single chord priming.

In this study, we examine through electrophysiological measures three alternative mechanisms underlying musical chord priming: psychoacoustic distance, common parent-key, and distance along the circle of fifths. In contrast with previous behavioral studies, we present complex tones which do not blur the melodic component, we present various chord arrangements, and we focus on nonmusicians. Target major chords, in three different harmonic conditions (1, 2, and 4 steps along the circle of fifths between prime and target chords), elicited two centro-anterior negativities labeled N5E (early) and N5L (late) suggesting a dissociation between an earlier psychoacoustic process based on pitch commonality and proximity and a later cognitive process based on a common parent-key.

Memory for tonal pitches: a music-length effect hypothesis.

One of the most studied effects of verbal working memory (WM) is the influence of the length of the words that compose the list to be remembered. This work aims to investigate the nature of musical WM by replicating the word length effect in the musical domain. Length and rate of presentation were manipulated in a recognition task of tone sequences. Results showed significant effects for both factors (length and presentation rate) as well as their interaction, suggesting the existence of different strategies (e.g., chunking and rehearsal) for the immediate memory of musical information, depending upon the length of the sequences.

Molecular genetic studies of the arginine vasopressin 1a receptor (AVPR1a) and the oxytocin receptor (OXTR) in human behaviour: from autism to altruism with some notes in between.

Converging evidence from both human and animal studies has highlighted the pervasive role of two neuropeptides, oxytocin (OXT) and arginine vasopressin (AVP), in mammalian social behaviours. Recent molecular genetic studies of the human arginine vasopressin 1a (AVPR1a) and oxytocin (OXTR) receptors have strengthened the evidence regarding the role of these two neuropeptides in a range of normal and pathological behaviours. Significant association between both AVPR1a repeat regions and OXTR single nucleotide polymorphisms (SNPs) with risk for autism has been provisionally shown which was mediated by socialization skills in our study. AVPR1a has also been linked to eating behaviour in both clinical and non-clinical groups, perhaps reflecting the social and ritualistic side of eating behaviour. Evidence also suggests that repeat variations in AVPR1a are associated with two other social domains in Homo sapiens: music and altruism. AVPR1a was associated with dance and musical cognition which we theorize as reflecting the ancient role of this hormone in social interactions executed by vocalization, ritual movement and dyadic (mother-offspring) and group communication. Finally, we have shown that individual differences in allocation of funds in the dictator game, a laboratory game of pure altruism, is predicted by length of the AVPR1a RS3 promoter-region repeat echoing the mechanism of this hormone's action in the vole model of affiliative behaviours and facilitation of positive group interactions. While still in its infancy, the current outlook for molecular genetic investigations of AVP-OXT continues to be fascinating. Future studies should profitably focus on pharmacogenomic and genomic imaging strategies facilitated by the ease and efficacy of manipulating AVP-OXT neurotransmission by intranasal administration. Importantly, physiological measures, behavioural paradigms and brain activation can be informed by considering between-group and also within-group individual differences defined by common polymorphisms. Ultimately, investigators should strive to develop a cohesive model explaining how genomic variations are translated into individual and group differences in higher-order social behaviours.

Neural sensitivity to human voices: ERP evidence of task and attentional influences.

In an earlier study, we found that human voices evoked a positive event-related potential (ERP) peaking at approximately 320 ms after stimulus onset, distinctive from those elicited by instrumental tones. Here we show that though similar in latency to the Novelty P3, this Voice-Sensitive Response (VSR) differs in antecedent conditions and scalp distribution. Furthermore, when participants were not attending to stimuli, the response to voices was undistinguished from other harmonic stimuli (strings, winds, and brass). During a task requiring attending to a feature other than timbre, voices were not distinguished from voicelike stimuli (strings), but were distinguished from other harmonic stimuli. We suggest that the component elicited by voices and similar sounds reflects the allocation of attention on the basis of stimulus significance (as opposed to novelty), and propose an explanation of the task and attentional factors that contribute to the effect.