Bodily maps of musical sensations across cultures.

Emotions, bodily sensations and movement are integral parts of musical experiences. Yet, it remains unknown i) whether emotional connotations and structural features of music elicit discrete bodily sensations and ii) whether these sensations are culturally consistent. We addressed these questions in a cross-cultural study with Western (European and North American, n = 903) and East Asian (Chinese, n = 1035). We precented participants with silhouettes of human bodies and asked them to indicate the bodily regions whose activity they felt changing while listening to Western and Asian musical pieces with varying emotional and acoustic qualities. The resulting bodily sensation maps (BSMs) varied as a function of the emotional qualities of the songs, particularly in the limb, chest, and head regions. Music-induced emotions and corresponding BSMs were replicable across Western and East Asian subjects. The BSMs clustered similarly across cultures, and cluster structures were similar for BSMs and self-reports of emotional experience. The acoustic and structural features of music were consistently associated with the emotion ratings and music-induced bodily sensations across cultures. These results highlight the importance of subjective bodily experience in music-induced emotions and demonstrate consistent associations between musical features, music-induced emotions, and bodily sensations across distant cultures.

Endogenous Opioid Release After Orgasm in Man: A Combined PET/Functional MRI Study.

The endogenous µ-opioid receptor (MOR) system plays a key role in the mammalian reward circuit. Human and animal experiments suggest the involvement of MORs in human sexual pleasure, yet this hypothesis currently lacks in vivo support. Methods: We used PET with the radioligand [11C]carfentanil, which has high affinity for MORs, to quantify endogenous opioid release after orgasm in man. Participants were scanned once immediately after orgasm and once in a baseline state. Hemodynamic activity was measured with functional MRI during penile stimulation. Results: The PET data revealed significant opioid release in the hippocampus. Hemodynamic activity in the somatosensory and motor cortices and in the hippocampus and thalamus increased during penile stimulation, and thalamic activation was linearly dependent on self-reported sexual arousal. Conclusion: Our data show that endogenous opioidergic activation in the medial temporal lobe is centrally involved in sexual arousal, and this circuit may be implicated in orgasmic disorders.

Decoding brain basis of laughter and crying in natural scenes.

Laughter and crying are universal signals of prosociality and distress, respectively. Here we investigated the functional brain basis of perceiving laughter and crying using naturalistic functional magnetic resonance imaging (fMRI) approach. We measured haemodynamic brain activity evoked by laughter and crying in three experiments with 100 subjects in each. The subjects i) viewed a 20-minute medley of short video clips, and ii) 30 min of a full-length feature film, and iii) listened to 13.5 min of a radio play that all contained bursts of laughter and crying. Intensity of laughing and crying in the videos and radio play was annotated by independent observes, and the resulting time series were used to predict hemodynamic activity to laughter and crying episodes. Multivariate pattern analysis (MVPA) was used to test for regional selectivity in laughter and crying evoked activations. Laughter induced widespread activity in ventral visual cortex and superior and middle temporal and motor cortices. Crying activated thalamus, cingulate cortex along the anterior-posterior axis, insula and orbitofrontal cortex. Both laughter and crying could be decoded accurately (66-77% depending on the experiment) from the BOLD signal, and the voxels contributing most significantly to classification were in superior temporal cortex. These results suggest that perceiving laughter and crying engage distinct neural networks, whose activity suppresses each other to manage appropriate behavioral responses to others' bonding and distress signals.

Functional organization of social perception networks in the human brain.

Humans rapidly extract diverse and complex information from ongoing social interactions, but the perceptual and neural organization of the different aspects of social perception remains unresolved. We showed short movie clips with rich social content to 97 healthy participants while their haemodynamic brain activity was measured with fMRI. The clips were annotated moment-to-moment for a large set of social features and 45 of the features were evaluated reliably between annotators. Cluster analysis of the social features revealed that 13 dimensions were sufficient for describing the social perceptual space. Three different analysis methods were used to map the social perceptual processes in the human brain. Regression analysis mapped regional neural response profiles for different social dimensions. Multivariate pattern analysis then established the spatial specificity of the responses and intersubject correlation analysis connected social perceptual processing with neural synchronization. The results revealed a gradient in the processing of social information in the brain. Posterior temporal and occipital regions were broadly tuned to most social dimensions and the classifier revealed that these responses showed spatial specificity for social dimensions; in contrast Heschl gyri and parietal areas were also broadly associated with different social signals, yet the spatial patterns of responses did not differentiate social dimensions. Frontal and subcortical regions responded only to a limited number of social dimensions and the spatial response patterns did not differentiate social dimension. Altogether these results highlight the distributed nature of social processing in the brain.

Pattern recognition reveals sex-dependent neural substrates of sexual perception.

Sex differences in brain activity evoked by sexual stimuli remain elusive despite robust evidence for stronger enjoyment of and interest toward sexual stimuli in men than in women. To test whether visual sexual stimuli evoke different brain activity patterns in men and women, we measured hemodynamic brain activity induced by visual sexual stimuli in two experiments with 91 subjects (46 males). In one experiment, the subjects viewed sexual and nonsexual film clips, and dynamic annotations for nudity in the clips were used to predict hemodynamic activity. In the second experiment, the subjects viewed sexual and nonsexual pictures in an event-related design. Men showed stronger activation than women in the visual and prefrontal cortices and dorsal attention network in both experiments. Furthermore, using multivariate pattern classification we could accurately predict the sex of the subject on the basis of the brain activity elicited by the sexual stimuli. The classification generalized across the experiments indicating that the sex differences were task-independent. Eye tracking data obtained from an independent sample of subjects (N = 110) showed that men looked longer than women at the chest area of the nude female actors in the film clips. These results indicate that visual sexual stimuli evoke discernible brain activity patterns in men and women which may reflect stronger attentional engagement with sexual stimuli in men.

Neural responses to biological motion distinguish autistic and schizotypal traits.

Difficulties in social interactions characterize both autism and schizophrenia and are correlated in the neurotypical population. It is unknown whether this represents a shared etiology or superficial phenotypic overlap. Both conditions exhibit atypical neural activity in response to the perception of social stimuli and decreased neural synchronization between individuals. This study investigated if neural activity and neural synchronization associated with biological motion perception are differentially associated with autistic and schizotypal traits in the neurotypical population. Participants viewed naturalistic social interactions while hemodynamic brain activity was measured with fMRI, which was modeled against a continuous measure of the extent of biological motion. General linear model analysis revealed that biological motion perception was associated with neural activity across the action observation network. However, intersubject phase synchronization analysis revealed neural activity to be synchronized between individuals in occipital and parietal areas but desynchronized in temporal and frontal regions. Autistic traits were associated with decreased neural activity (precuneus and middle cingulate gyrus), and schizotypal traits were associated with decreased neural synchronization (middle and inferior frontal gyri). Biological motion perception elicits divergent patterns of neural activity and synchronization, which dissociate autistic and schizotypal traits in the general population, suggesting that they originate from different neural mechanisms.

Chinese-English bilinguals are more sensitive to environmental sound perception than Spanish-English bilinguals through top-down cognitive mechanism.

Sensitive perception to environmental sound is important for an individual's daily life since it helps in responding to the environment quickly and avoiding potential risks. It remains unclear, however, whether and how bilingual experiences with different native language backgrounds influence brain responses reflecting environmental sound perception. The present study compared Chinese-English bilinguals, Spanish-English bilinguals, and English monolinguals on their brain response to environmental sound perception and further examined its processing mechanism. The present study did not find the differences between English monolinguals and two bilingual groups on the environmental sound perception. However, we found that compared with Spanish-English bilinguals, Chinese-English bilinguals showed significantly larger mismatch negativity (MMN), indicating that Chinese-English bilinguals were more sensitive to the environmental sounds than that of Spanish-English bilinguals. Further, compared with Spanish-English bilinguals, Chinese-English bilinguals showed significantly larger theta oscillation. In addition, the theta oscillation is significantly correlated with MMN for Chinese-English bilinguals. The results indicated that the advantage of Chinese-English bilingual experience on environmental sound perception may be due to the enhanced top-down cognitive ability. In summary, these findings suggest that compared with the Spanish-English bilinguals, Chinese-English bilinguals showed an advantage in environmental sound perception via enhanced top-down cognitive modulation.

Mu-opioid receptor system modulates responses to vocal bonding and distress signals in humans.

Laughter is a contagious prosocial signal that conveys bonding motivation; adult crying conversely communicates desire for social proximity by signalling distress. Endogenous mu-opioid receptors (MORs) modulate sociability in humans and non-human primates. In this combined PET-fMRI study (n = 17), we tested whether central MOR tone is associated with regional brain responses to social signals of laughter and crying. MOR availability was measured with positron emission tomography (PET) using the high-affinity agonist radioligand [11C]carfentanil. Haemodynamic responses to social laughter and crying vocalizations were measured using functional magnetic resonance imaging (fMRI). Social laughter evoked activation in the auditory cortex, insula, cingulate cortex, amygdala, primary and secondary somatosensory cortex, and primary and secondary motor cortex; crying sounds led to more restricted activation in the auditory cortex and nearby areas. MOR availability was negatively correlated with the haemodynamic responses to social laughter in the primary and secondary somatosensory cortex, primary and secondary motor cortex, posterior insula, posterior cingulate cortex, precuneus, cuneus, temporal gyri and lingual gyrus. For crying-evoked activations, MOR availability was negatively correlated with medial and lateral prefrontal haemodynamic responses. Altogether our findings highlight the role of the MOR system in modulating acute brain responses to both positive and negative social signals. This article is part of the theme issue 'Cracking the laugh code: laughter through the lens of biology, psychology and neuroscience'.

Brain structural alterations in autism and criminal psychopathy.

The goal of this study was to elucidate the anatomical brain basis of social cognition through two disorders with distinctively different phenotypes of social interaction. We compared structural MR images of 20 individuals with autism spectrum disorder (ASD), 19 violent offenders with high psychopathic traits, and 19 control participants using voxel-based morphometry (VBM). Our earlier study showed lower grey matter volume (GMV) values in the insula, frontal cortex, and sensorimotor cortex of the offender group compared to controls. In the present study, the images of the ASD group revealed lower GMV in the left precuneus, right cerebellum, and right precentral gyrus in comparison with controls. The comparison between the offender and ASD groups showed lower GMV values for the right temporal pole and left inferior frontal gyrus in the offender group. There was also an overlap of both disorders in the right pre-central cortex, showing lower GMV compared to controls. Our findings suggest structural differences between violent offenders with high psychopathy traits and ASD individuals in the frontotemporal social brain network areas, previously associated with empathy. We also provide evidence of similar abnormal structures in the motor cortex for both of these disorders, possibly related to uniting issues of social cognition.

Inter-brain synchronization occurs without physical co-presence during cooperative online gaming.

Inter-brain synchronization during social interaction has been linked with several positive phenomena, including closeness, cooperation, prosociality, and team performance. However, the temporal dynamics of inter-brain synchronization during collaboration are not yet fully understood. Furthermore, with collaboration increasingly happening online, the dependence of inter-brain phase synchronization of oscillatory activity on physical presence is an important but understudied question. In this study, physically isolated participants performed a collaborative coordination task in the form of a cooperative multiplayer game. We measured EEG from 42 subjects working together as pairs in the task. During the measurement, the only interaction between the participants happened through on-screen movement of a racing car, controlled by button presses of both participants working with distinct roles, either controlling the speed or the direction of the car. Pairs working together in the task were found to have elevated neural coupling in the alpha, beta, and gamma frequency bands, compared to performance matched false pairs. Higher gamma synchrony was associated with better momentary performance within dyads and higher alpha synchrony was associated with better mean performance across dyads. These results are in line with previous findings of increased inter-brain synchrony during interaction, and show that phase synchronization of oscillatory activity occurs during online real-time joint coordination without any physical co-presence or video and audio connection. Synchrony decreased during a playing session, but was found to be higher during the second session compared to the first. The novel paradigm, developed for the measurement of real-time collaborative performance, demonstrates that changes in inter-brain EEG phase synchrony can be observed continuously during interaction.

Atlas of type 2 dopamine receptors in the human brain: Age and sex dependent variability in a large PET cohort.

BACKGROUND: The dopamine system contributes to a multitude of functions ranging from reward and motivation to learning and movement control, making it a key component in goal-directed behavior. Altered dopaminergic function is observed in neurological and psychiatric conditions. Numerous factors have been proposed to influence dopamine function, but due to small sample sizes and heterogeneous data analysis methods in previous studies their specific and joint contributions remain unresolved. METHODS: In this cross-sectional register-based study we investigated how age, sex, body mass index (BMI), as well as cerebral hemisphere and regional volume influence striatal type 2 dopamine receptor (D2R) availability in the human brain. We analyzed a large historical dataset (n=156, 120 males and 36 females) of [11C]raclopride PET scans performed between 2004 and 2018. RESULTS: Striatal D2R availability decreased through age for both sexes (2-5 % in striatal ROIs per 10 years) and was higher in females versus males throughout age (7-8% in putamen). BMI and striatal D2R availability were weakly associated. There was no consistent lateralization of striatal D2R. The observed effects were independent of regional volumes. These results were validated using two different spatial normalization methods, and the age and sex effects also replicated in an independent sample (n=135). CONCLUSIONS: D2R availability is dependent on age and sex, which may contribute to the vulnerability of neurological and psychiatric conditions involving altering D2R expression.

Aberrant motor contagion of emotions in psychopathy and high-functioning autism.

Psychopathy and autism are both associated with aberrant social skills and empathy, yet only psychopaths are markedly antisocial and violent. Here, we compared the functional neural alterations underlying these two groups that both have aberrant empathetic abilities but distinct behavioral phenotypes. We studied 19 incarcerated male offenders with high psychopathic traits, 20 males with high-functioning autism, and 19 age-matched healthy controls. All groups underwent functional magnetic resonance imaging while they viewed dynamic happy, angry, and disgusted faces or listened to laughter and crying sounds. Psychopathy was associated with reduced somatomotor responses to almost all expressions, while participants with autism demonstrated less marked and emotion-specific alterations in the somatomotor area. These data suggest that psychopathy and autism involve both common and distinct functional alterations in the brain networks involved in the socioemotional processing. The alterations are more profound in psychopathy, possibly reflecting the more severely disturbed socioemotional brain networks in this population.

µ-opioid receptor availability is associated with sex drive in human males.

The endogenous mu-opioid receptor (MOR) system modulates a multitude of social and reward-related functions, and exogenous opiates also influence sex drive in humans and animals. Sex drive shows substantial variation across humans, and it is possible that individual differences in MOR availability underlie interindividual of variation in human sex drive. We measured healthy male subjects' (n = 52) brain's MOR availability with positron emission tomography (PET) using an agonist radioligand, [11C]carfentanil, that has high affinity for MORs. Sex drive was measured using self-reports of engaging in sexual behaviour (sex with partner and masturbating). Bayesian hierarchical regression analysis revealed that sex drive was positively associated with MOR availability in cortical and subcortical areas, notably in caudate nucleus, hippocampus, and cingulate cortices. These results were replicated in full-volume GLM analysis. These widespread effects are in line with high spatial autocorrelation in MOR expression in human brain. Complementary voxel-based morphometry analysis (n = 108) of anatomical MR images provided limited evidence for positive association between sex drive and cortical density in the midcingulate cortex. We conclude that endogenous MOR tone is associated with individual differences in sex drive in human males.

Improved Auditory Function Caused by Music Versus Foreign Language Training at School Age: Is There a Difference?

In adults, music and speech share many neurocognitive functions, but how do they interact in a developing brain? We compared the effects of music and foreign language training on auditory neurocognition in Chinese children aged 8-11 years. We delivered group-based training programs in music and foreign language using a randomized controlled trial. A passive control group was also included. Before and after these year-long extracurricular programs, auditory event-related potentials were recorded (n = 123 and 85 before and after the program, respectively). Through these recordings, we probed early auditory predictive brain processes. To our surprise, the language program facilitated the children's early auditory predictive brain processes significantly more than did the music program. This facilitation was most evident in pitch encoding when the experimental paradigm was musically relevant. When these processes were probed by a paradigm more focused on basic sound features, we found early predictive pitch encoding to be facilitated by music training. Thus, a foreign language program is able to foster auditory and music neurocognition, at least in tonal language speakers, in a manner comparable to that by a music program. Our results support the tight coupling of musical and linguistic brain functions also in the developing brain.

Being moved by listening to unfamiliar sad music induces reward-related hormonal changes in empathic listeners.

Many people enjoy sad music, and the appeal for tragedy is widespread among the consumers of film and literature. The underlying mechanisms of such aesthetic experiences are not well understood. We tested whether pleasure induced by sad, unfamiliar instrumental music is explained with a homeostatic or a reward theory, each of which is associated with opposite patterns of changes in the key hormones. Sixty-two women listened to sad music (or nothing) while serum was collected for subsequent measurement of prolactin (PRL) and oxytocin (OT) and stress marker (cortisol and adrenocorticotropic hormone) concentrations. Two groups of participants were recruited on the basis of low and high trait empathy. In the high empathy group, PRL and OT levels were significantly lower with music compared with no music. And compared to the low empathy group, the high empathy individuals reported an increase of positive mood and higher ratings of being moved with music. None of the stress markers showed any changes across the conditions or the groups. These hormonal changes, inconsistent with the homeostatic theory proposed by Huron, exhibit a pattern expected of general reward. Our findings illuminate how unfamiliar and low arousal music may give rise to pleasurable experiences.

Faster maturation of selective attention in musically trained children and adolescents: Converging behavioral and event-related potential evidence.

Previous work suggests that musical training in childhood is associated with enhanced executive functions. However, it is unknown whether this advantage extends to selective attention-another central aspect of executive control. We recorded a well-established event-related potential (ERP) marker of distraction, the P3a, during an audio-visual task to investigate the maturation of selective attention in musically trained children and adolescents aged 10-17 years and a control group of untrained peers. The task required categorization of visual stimuli, while a sequence of standard sounds and distracting novel sounds were presented in the background. The music group outperformed the control group in the categorization task and the younger children in the music group showed a smaller P3a to the distracting novel sounds than their peers in the control group. Also, a negative response elicited by the novel sounds in the N1/MMN time range (~150-200 ms) was smaller in the music group. These results indicate that the music group was less easily distracted by the task-irrelevant sound stimulation and gated the neural processing of the novel sounds more efficiently than the control group. Furthermore, we replicated our previous finding that, relative to the control group, the musically trained children and adolescents performed faster in standardized tests for inhibition and set shifting. These results provide novel converging behavioral and electrophysiological evidence from a cross-modal paradigm for accelerated maturation of selective attention in musically trained children and adolescents and corroborate the association between musical training and enhanced inhibition and set shifting.

Brain Basis of Psychopathy in Criminal Offenders and General Population.

Psychopathy is characterized by persistent antisocial behavior, impaired empathy, and egotistical traits. These traits vary also in normally functioning individuals. Here, we tested whether such antisocial personalities are associated with similar structural and neural alterations as those observed in criminal psychopathy. Subjects were 100 non-convicted well-functioning individuals, 19 violent male offenders, and 19 matched controls. Subjects underwent T1-weighted magnetic resonance imaging and viewed movie clips with varying violent content during functional magnetic resonance imaging. Psychopathic traits were evaluated with Levenson Self-Report Psychopathy Scale (controls) and Psychopathy Checklist-Revised (offenders). Psychopathic offenders had lower gray matter density (GMD) in orbitofrontal cortex and anterior insula. In the community sample, affective psychopathy traits were associated with lower GMD in the same areas. Viewing violence increased brain activity in periaqueductal grey matter, thalamus, somatosensory, premotor, and temporal cortices. Psychopathic offenders had increased responses to violence in thalamus and orbitofrontal, insular, and cingulate cortices. In the community sample, impulsivity-related psychopathy traits were positively associated with violence-elicited responses in similar areas. We conclude that brain characteristics underlying psychopathic spectrum in violent psychopathy are related to those observed in well-functioning individuals with asocial personality features.

Arcuate fasciculus architecture is associated with individual differences in pre-attentive detection of unpredicted music changes.

The mismatch negativity (MMN) is an event related brain potential (ERP) elicited by unpredicted sounds presented in a sequence of repeated auditory stimuli. The neural sources of the MMN have been previously attributed to a fronto-temporo-parietal network which crucially overlaps with the so-called auditory dorsal stream, involving inferior and middle frontal, inferior parietal, and superior and middle temporal regions. These cortical areas are structurally connected by the arcuate fasciculus (AF), a three-branch pathway supporting the feedback-feedforward loop involved in auditory-motor integration, auditory working memory, storage of acoustic templates, as well as comparison and update of those templates. Here, we characterized the individual differences in the white-matter macrostructural properties of the AF and explored their link to the electrophysiological marker of passive change detection gathered in a melodic multifeature MMN-EEG paradigm in 26 healthy young adults without musical training. Our results show that left fronto-temporal white-matter connectivity plays an important role in the pre-attentive detection of rhythm modulations within a melody. Previous studies have shown that this AF segment is also critical for language processing and learning. This strong coupling between structure and function in auditory change detection might be related to life-time linguistic (and possibly musical) exposure and experiences, as well as to timing processing specialization of the left auditory cortex. To the best of our knowledge, this is the first time in which the relationship between neurophysiological (EEG) and brain white-matter connectivity indexes using DTI-tractography are studied together. Thus, the present results, although still exploratory, add to the existing evidence on the importance of studying the constraints imposed on cognitive functions by the underlying structural connectivity.

Effects of Music Training on the Auditory Working Memory of Chinese-Speaking School-Aged Children: A Longitudinal Intervention Study.

Music expertise is known to be beneficial for cognitive function and development. In this study, we conducted 1-year music training for school children (n = 123; 7-11 years of age before training) in China. The children were assigned to music or second-language after-class training groups. A passive control group was included. We aimed to investigate whether music training could facilitate working memory (WM) development compared to second-language training and no training. Before and after the training, auditory WM was measured via a digit span (DS) task, together with the vocabulary and block tests of the Wechsler Intelligence Scale for Child IV (WISC-IV). The results of the DS task revealed superior development in the music group compared to the other groups. However, further analysis of DS forward and backward tasks indicated that the performance of the three training/non-training groups only differed significantly in DS backward scores, but not in the DS forward scores. We conclude that music training may benefit the central executive system of WM, as reflected by the DS backward task.

Decoding Music-Evoked Emotions in the Auditory and Motor Cortex.

Music can induce strong subjective experience of emotions, but it is debated whether these responses engage the same neural circuits as emotions elicited by biologically significant events. We examined the functional neural basis of music-induced emotions in a large sample (n = 102) of subjects who listened to emotionally engaging (happy, sad, fearful, and tender) pieces of instrumental music while their hemodynamic brain activity was measured with functional magnetic resonance imaging (fMRI). Ratings of the four categorical emotions and liking were used to predict hemodynamic responses in general linear model (GLM) analysis of the fMRI data. Multivariate pattern analysis (MVPA) was used to reveal discrete neural signatures of the four categories of music-induced emotions. To map neural circuits governing non-musical emotions, the subjects were scanned while viewing short emotionally evocative film clips. The GLM revealed that most emotions were associated with activity in the auditory, somatosensory, and motor cortices, cingulate gyrus, insula, and precuneus. Fear and liking also engaged the amygdala. In contrast, the film clips strongly activated limbic and cortical regions implicated in emotional processing. MVPA revealed that activity in the auditory cortex and primary motor cortices reliably discriminated the emotion categories. Our results indicate that different music-induced basic emotions have distinct representations in regions supporting auditory processing, motor control, and interoception but do not strongly rely on limbic and medial prefrontal regions critical for emotions with survival value.