Live music stimulates the affective brain and emotionally entrains listeners in real time.

Music is powerful in conveying emotions and triggering affective brain mechanisms. Affective brain responses in previous studies were however rather inconsistent, potentially because of the non-adaptive nature of recorded music used so far. Live music instead can be dynamic and adaptive and is often modulated in response to audience feedback to maximize emotional responses in listeners. Here, we introduce a setup for studying emotional responses to live music in a closed-loop neurofeedback setup. This setup linked live performances by musicians to neural processing in listeners, with listeners' amygdala activity was displayed to musicians in real time. Brain activity was measured using functional MRI, and especially amygdala activity was quantified in real time for the neurofeedback signal. Live pleasant and unpleasant piano music performed in response to amygdala neurofeedback from listeners was acoustically very different from comparable recorded music and elicited significantly higher and more consistent amygdala activity. Higher activity was also found in a broader neural network for emotion processing during live compared to recorded music. This finding included observations of the predominance for aversive coding in the ventral striatum while listening to unpleasant music, and involvement of the thalamic pulvinar nucleus, presumably for regulating attentional and cortical flow mechanisms. Live music also stimulated a dense functional neural network with the amygdala as a central node influencing other brain systems. Finally, only live music showed a strong and positive coupling between features of the musical performance and brain activity in listeners pointing to real-time and dynamic entrainment processes.

Neurocognitive processing efficiency for discriminating human non-alarm rather than alarm scream calls.

Across many species, scream calls signal the affective significance of events to other agents. Scream calls were often thought to be of generic alarming and fearful nature, to signal potential threats, with instantaneous, involuntary, and accurate recognition by perceivers. However, scream calls are more diverse in their affective signaling nature than being limited to fearfully alarming a threat, and thus the broader sociobiological relevance of various scream types is unclear. Here we used 4 different psychoacoustic, perceptual decision-making, and neuroimaging experiments in humans to demonstrate the existence of at least 6 psychoacoustically distinctive types of scream calls of both alarming and non-alarming nature, rather than there being only screams caused by fear or aggression. Second, based on perceptual and processing sensitivity measures for decision-making during scream recognition, we found that alarm screams (with some exceptions) were overall discriminated the worst, were responded to the slowest, and were associated with a lower perceptual sensitivity for their recognition compared with non-alarm screams. Third, the neural processing of alarm compared with non-alarm screams during an implicit processing task elicited only minimal neural signal and connectivity in perceivers, contrary to the frequent assumption of a threat processing bias of the primate neural system. These findings show that scream calls are more diverse in their signaling and communicative nature in humans than previously assumed, and, in contrast to a commonly observed threat processing bias in perceptual discriminations and neural processes, we found that especially non-alarm screams, and positive screams in particular, seem to have higher efficiency in speeded discriminations and the implicit neural processing of various scream types in humans.

Functional brain changes in sarcopenia: evidence for differential central neural mechanisms in dynapenic older women.

BACKGROUND: The European Working Group on Sarcopenia in Older People (EWGSOP2) recently revised its definition and diagnostic criteria for sarcopenia, placing muscle strength at the forefront. The pathogenesis of dynapenia (or low muscle strength) is still not fully understood, but there is emerging evidence that central neural factors constitute critical determinants. METHODS: Our cross-sectional study included 59 community-dwelling older women (mean age 73.1 ± 4.9 years). Participants underwent detailed skeletal muscle assessments for muscle strength defined by handgrip strength and chair rise time measurements using the recently published EWGSOP2 cut-off points. Functional magnetic resonance imaging (fMRI) was assessed during the performance of a cognitive dual-task paradigm, consisting of a baseline, two single-tasks (motor and arithmetic) and one dual-task (motor and arithmetic combined). RESULTS: Forty-seven percent (28/59) of participants were classified as dynapenic. fMRI results revealed a differential recruitment of motor circuits in the brain during the dual-task condition in dynapenic as compared with non-dynapenic participants. In particular, while the brain activity during the single-tasks did not differ between the two groups, only during the dual-task non-dynapenic participants showed significant increased activation in dorsolateral prefrontal and premotor cortex, and in supplementary motor area as compared to dynapenic participants. CONCLUSION: Our results point to a dysfunctional involvement of brain networks associated with motor control in dynapenia in a multi-tasking paradigm. A better knowledge of the link between dynapenia and brain functions could provide new impulses in the diagnosis and interventions for sarcopenia.

Neural oscillations in human auditory cortex revealed by fast fMRI during auditory perception.

Previous work pointed to the neural and functional significance of infraslow neural oscillations below 1 â€‹Hz that can be detected and precisely located with fast functional magnetic resonance imaging (fMRI). While previous work demonstrated this significance for brain dynamics during very low-level sensory stimulation, we here provide the first evidence for the detectability and functional significance of infraslow oscillatory blood oxygenation level-dependent (BOLD) responses to auditory stimulation by the sociobiological relevant and more complex category of voices. Previous work pointed to a specific area of the mammalian auditory cortex (AC) that is sensitive to vocal signals as quantified by activation levels. Here we show, by using fast fMRI, that the human voice-sensitive AC prioritizes vocal signals not only in terms of activity level but also in terms of specific infraslow BOLD oscillations. We found unique sustained and transient oscillatory BOLD patterns in the AC for vocal signals. For transient oscillatory patterns, vocal signals showed faster peak oscillatory responses across all AC regions. Furthermore, we identified an exclusive sustained oscillatory component for vocal signals in the primary AC. Fast fMRI thus demonstrates the significance and richness of infraslow BOLD oscillations for neurocognitive mechanisms in social cognition as demonstrated here for the sociobiological relevance of voice processing.

Whispering - The hidden side of auditory communication.

Whispering is a unique expression mode that is specific to auditory communication. Individuals switch their vocalization mode to whispering especially when affected by inner emotions in certain social contexts, such as in intimate relationships or intimidating social interactions. Although this context-dependent whispering is adaptive, whispered voices are acoustically far less rich than phonated voices and thus impose higher hearing and neural auditory decoding demands for recognizing their socio-affective value by listeners. The neural dynamics underlying this recognition especially from whispered voices are largely unknown. Here we show that whispered voices in humans are considerably impoverished as quantified by an entropy measure of spectral acoustic information, and this missing information needs large-scale neural compensation in terms of auditory and cognitive processing. Notably, recognizing the socio-affective information from voices was slightly more difficult from whispered voices, probably based on missing tonal information. While phonated voices elicited extended activity in auditory regions for decoding of relevant tonal and time information and the valence of voices, whispered voices elicited activity in a complex auditory-frontal brain network. Our data suggest that a large-scale multidirectional brain network compensates for the impoverished sound quality of socially meaningful environmental signals to support their accurate recognition and valence attribution.

Getting the beat: entrainment of brain activity by musical rhythm and pleasantness.

Rhythmic entrainment is an important component of emotion induction by music, but brain circuits recruited during spontaneous entrainment of attention by music and the influence of the subjective emotional feelings evoked by music remain still largely unresolved. In this study we used fMRI to test whether the metric structure of music entrains brain activity and how music pleasantness influences such entrainment. Participants listened to piano music while performing a speeded visuomotor detection task in which targets appeared time-locked to either strong or weak beats. Each musical piece was presented in both a consonant/pleasant and dissonant/unpleasant version. Consonant music facilitated target detection and targets presented synchronously with strong beats were detected faster. FMRI showed increased activation of bilateral caudate nucleus when responding on strong beats, whereas consonance enhanced activity in attentional networks. Meter and consonance selectively interacted in the caudate nucleus, with greater meter effects during dissonant than consonant music. These results reveal that the basal ganglia, involved both in emotion and rhythm processing, critically contribute to rhythmic entrainment of subcortical brain circuits by music.

Mapping aesthetic musical emotions in the brain.

Music evokes complex emotions beyond pleasant/unpleasant or happy/sad dichotomies usually investigated in neuroscience. Here, we used functional neuroimaging with parametric analyses based on the intensity of felt emotions to explore a wider spectrum of affective responses reported during music listening. Positive emotions correlated with activation of left striatum and insula when high-arousing (Wonder, Joy) but right striatum and orbitofrontal cortex when low-arousing (Nostalgia, Tenderness). Irrespective of their positive/negative valence, high-arousal emotions (Tension, Power, and Joy) also correlated with activations in sensory and motor areas, whereas low-arousal categories (Peacefulness, Nostalgia, and Sadness) selectively engaged ventromedial prefrontal cortex and hippocampus. The right parahippocampal cortex activated in all but positive high-arousal conditions. Results also suggested some blends between activation patterns associated with different classes of emotions, particularly for feelings of Wonder or Transcendence. These data reveal a differentiated recruitment across emotions of networks involved in reward, memory, self-reflective, and sensorimotor processes, which may account for the unique richness of musical emotions.

Nonverbal memory tests revisited: Neuroanatomical correlates and differential influence of biasing cognitive functions.

The detection of right temporal lobe dysfunction with nonverbal memory tests has remained difficult in the past. Reasons for this might be the potential influence of other biasing cognitive functions such as executive functions or the verbalisability of nonverbal material. The aim of this study was to investigate three classic nonverbal memory tests by identifying their neuroanatomical correlates with lesion-symptom mapping (LSM) and by probing their independence from verbal encoding abilities and executive functions. In a cohort of 119 patients with first-time cerebrovascular accident, memory performance was assessed in the Nonverbal Learning and Memory Test for Routes (NLMTR), the Rey Complex Figure Test (RCFT), and the Visual Design Learning Test (VDLT). Calculating multivariate LSM, we identified crucial brain structures for these three nonverbal memory tests. Behavioural analyses were performed to assess the impact of executive functions and verbal encoding abilities with regression analyses and likelihood-ratio tests. LSM revealed for the RCFT mainly right-hemispheric frontal, insular, subcortical, and white matter structures and for the NLMTR right-hemispheric temporal (hippocampus), insular, subcortical, and white matter structures. The VDLT did not reach significance in LSM analyses. Behavioural results showed that amongst the three nonverbal memory tests the impact of executive functions was most pronounced for RCFT, and the impact of verbal encoding abilities was most important in VDLT. Likelihood-ratio tests confirmed that only for NLMTR did the goodness of fit not significantly improve by adding executive functions or verbal encoding abilities. These results suggest that amongst the three nonverbal memory tests the NLMTR, as a spatial navigation test, could serve as the most suitable marker of right-hemispheric temporal lobe functioning, with the right hippocampus being involved only in this test. In addition, the behavioural results propose that only NLMTR seems mostly unaffected by executive functions and verbal encoding abilities.

The German Revised version of the Niigata PPPD Questionnaire (NPQ-R): Development with patient interviews and an expert Delphi consensus.

BACKGROUND: Persistent postural-perceptual dizziness (PPPD) is a functional disorder of the nervous system and currently one of the most common types of chronic dizziness. Currently existing questionnaires do not fully assess patients' specific symptoms of PPPD. The Japanese Niigata PPPD Questionnaire (NPQ) was recently developed following consensus-based diagnosis criteria. The aim of this study was to translate it into German, evaluate its content with the help of experts and patients and, if necessary, revise the original version to allow for a comprehensive assessment of patients' PPPD-related symptoms. METHODS: A 3-round expert Delphi survey and semi-structured patient interviews were conducted. 28 experts from Switzerland, Germany and Austria working in hospitals or outpatient centres were asked to complete a first questionnaire on various aspects of PPPD, on the translated, original NPQ and their own related experiences (Round one), a second questionnaire with statements regarding PPPD they could agree or disagree with using a 6-point Likert-scale (Round two), and a third survey to finally reach a consensus on statements to be integrated into the NPQ. In addition, eleven patients (mean age of 64.6±12.6 years; 6 females) were selected according to the criteria for the diagnosis of PPPD proposed by the Bárány Society and participated in a semi-structured interview asking for their opinion on the content of the original NPQ. All collected data were analysed using a descriptive evaluation and a qualitative content analysis based on verbatim transcripts. RESULTS: Seven new items were added to the NPQ based on expert and patient comments and ratings. Its revised version (NPQ-R) comprises 19 items divided into five subscales using a 7-point Likert-scale with two additional subscales relating to associated symptoms and symptom behaviour in PPPD. The new maximal score is 114 points compared to 72 for the NPQ. CONCLUSION: The NPQ-R is the first patient-reported outcome measurement for patients with PPPD in German. It should help to provide a comprehensive assessment of the intensity of PPPD in affected patients.

Lesion-symptom mapping corroborates lateralization of verbal and nonverbal memory processes and identifies distributed brain networks responsible for memory dysfunction.

Memory disorders are a common consequence of cerebrovascular accident (CVA). However, uncertainties remain about the exact anatomical correlates of memory impairment and the material-specific lateralization of memory function in the brain. We used lesion-symptom mapping (LSM) in patients with first-time CVA to identify which brain structures are pivotal for verbal and nonverbal memory and to re-examine whether verbal and nonverbal memory functions are lateralized processes in the brain. The cognitive performance of a relatively large cohort of 114 patients in five classic episodic memory tests was analysed with factor analysis. Two factors were extracted that distinguished the verbal and nonverbal components of these memory tests, and their scores were subsequently tested for anatomical correlates by combining univariate and multivariate LSM. LSM analysis revealed for the verbal factor exclusively left-hemispheric insular, subcortical and adjacent white matter regions and for the nonverbal factor exclusively right-hemispheric temporal, occipital, insular, subcortical and adjacent white matter structures. These results corroborate the long-standing hypothesis of a material-specific lateralization of memory function in the brain and confirm a robust association between right temporal lobe lesions and nonverbal memory dysfunction. The right-hemispheric correlates for the nonverbal aspects of episodic memory include not only classic memory structures in the medial temporal lobe but also a more distributed network that includes cortical and subcortical structures also known for implicit memory processes.

Neural Dynamics of Karaoke-Like Voice Imitation in Singing Performance.

Beyond normal and non-imitative singing, the imitation of the timbre of another singer's voice, such as in Karaoke singing, involves the demanding reproduction of voice quality features and strongly depends on singing experience and practice. We show that precise voice imitation in a highly proficient and experienced vocal imitator, even in the absence of external auditory voice feedback, largely drew on internal cortico-subcortical auditory resources to control voicing errors based on imagined voice performance. Compared to the experienced vocal imitator, singers of a control group without experience in voice imitation used only sensorimotor feedback and demanding monitoring resources for imitation in the absence of voice feedback, a neural strategy that led, however, to a significantly poorer vocal performance. Thus, only long-term vocal imitation experience allows for the additional use of internal auditory brain resources, which result from training-induced brain plasticity, and which enable accurate vocal performance even under difficult performance conditions.

Brain networks mediating the influence of background music on selective attention.

Prevalent across societies and times, music has the ability to enhance attention, a property relevant to clinical applications, but the underlying brain mechanisms remain unknown. It is also unclear whether music produces similar or differential effects with advancing age. Here, we used event-related functional magnetic resonance imaging to investigate the influence of music exposure evoking four types of emotions on distinct attentional components measured with a modified attention network test, across 19 young (21 ± 2.6) and 33 old participants (72 ± 5.4). We then determined whether music-related effects differed across age groups and whether they were associated with particular acoustic features. Background music during selective attention requiring distractor conflict resolution was associated with faster response times and greater activations of fronto-parietal areas during happy and high-arousing music, whereas sad and low-valence music was associated with slower responses and greater occipital recruitment. Shifting and altering components of attention were unaffected. The influence of music on performance and brain networks was similar between age groups. These behavioral and neuroimaging results demonstrate the importance of affective music dimensions, particularly arousal, in enhancing selective attention processes. This study adds novel support to the benefits of music in the rehabilitation of attention functions.

The sound of emotions-Towards a unifying neural network perspective of affective sound processing.

Affective sounds are an integral part of the natural and social environment that shape and influence behavior across a multitude of species. In human primates, these affective sounds span a repertoire of environmental and human sounds when we vocalize or produce music. In terms of neural processing, cortical and subcortical brain areas constitute a distributed network that supports our listening experience to these affective sounds. Taking an exhaustive cross-domain view, we accordingly suggest a common neural network that facilitates the decoding of the emotional meaning from a wide source of sounds rather than a traditional view that postulates distinct neural systems for specific affective sound types. This new integrative neural network view unifies the decoding of affective valence in sounds, and ascribes differential as well as complementary functional roles to specific nodes within a common neural network. It also highlights the importance of an extended brain network beyond the central limbic and auditory brain systems engaged in the processing of affective sounds.

Alteration of complex negative emotions induced by music in euthymic patients with bipolar disorder.

BACKGROUND: Research has shown bipolar disorder to be characterized by dysregulation of emotion processing, including biases in facial expression recognition that is most prevalent during depressive and manic states. Very few studies have examined induced emotions when patients are in a euthymic phase, and there has been no research on complex emotions. We therefore set out to test emotional hyperreactivity in response to musical excerpts inducing complex emotions in bipolar disorder during euthymia. METHODS: We recruited 21 patients with bipolar disorder (BD) in a euthymic phase and 21 matched healthy controls. Participants first rated their emotional reactivity on two validated self-report scales (ERS and MAThyS). They then rated their music-induced emotions on nine continuous scales. The targeted emotions were wonder, power, melancholy and tension. We used a specific generalized linear mixed model to analyze the behavioral data. RESULTS: We found that participants in the euthymic bipolar group experienced more intense complex negative emotions than controls when the musical excerpts induced wonder. Moreover, patients exhibited greater emotional reactivity in daily life (ERS). Finally, a greater experience of tension while listening to positive music seemed to be mediated by greater emotional reactivity and a deficit in executive functions. LIMITATIONS: The heterogeneity of the BD group in terms of clinical characteristics may have influenced the results. CONCLUSIONS: Euthymic patients with bipolar disorder exhibit more complex negative emotions than controls in response to positive music.

Music and emotions: from enchantment to entrainment.

Producing and perceiving music engage a wide range of sensorimotor, cognitive, and emotional processes. Emotions are a central feature of the enjoyment of music, with a large variety of affective states consistently reported by people while listening to music. However, besides joy or sadness, music often elicits feelings of wonder, nostalgia, or tenderness, which do not correspond to emotion categories typically studied in neuroscience and whose neural substrates remain largely unknown. Here we review the similarities and differences in the neural substrates underlying these "complex" music-evoked emotions relative to other more "basic" emotional experiences. We suggest that these emotions emerge through a combination of activation in emotional and motivational brain systems (e.g., including reward pathways) that confer its valence to music, with activation in several other areas outside emotional systems, including motor, attention, or memory-related regions. We then discuss the neural substrates underlying the entrainment of cognitive and motor processes by music and their relation to affective experience. These effects have important implications for the potential therapeutic use of music in neurological or psychiatric diseases, particularly those associated with motor, attention, or affective disturbances.

Temporal dynamics of musical emotions examined through intersubject synchrony of brain activity.

To study emotional reactions to music, it is important to consider the temporal dynamics of both affective responses and underlying brain activity. Here, we investigated emotions induced by music using functional magnetic resonance imaging (fMRI) with a data-driven approach based on intersubject correlations (ISC). This method allowed us to identify moments in the music that produced similar brain activity (i.e. synchrony) among listeners under relatively natural listening conditions. Continuous ratings of subjective pleasantness and arousal elicited by the music were also obtained for the music outside of the scanner. Our results reveal synchronous activations in left amygdala, left insula and right caudate nucleus that were associated with higher arousal, whereas positive valence ratings correlated with decreases in amygdala and caudate activity. Additional analyses showed that synchronous amygdala responses were driven by energy-related features in the music such as root mean square and dissonance, while synchrony in insula was additionally sensitive to acoustic event density. Intersubject synchrony also occurred in the left nucleus accumbens, a region critically implicated in reward processing. Our study demonstrates the feasibility and usefulness of an approach based on ISC to explore the temporal dynamics of music perception and emotion in naturalistic conditions.

The role of the medial temporal limbic system in processing emotions in voice and music.

Subcortical brain structures of the limbic system, such as the amygdala, are thought to decode the emotional value of sensory information. Recent neuroimaging studies, as well as lesion studies in patients, have shown that the amygdala is sensitive to emotions in voice and music. Similarly, the hippocampus, another part of the temporal limbic system (TLS), is responsive to vocal and musical emotions, but its specific roles in emotional processing from music and especially from voices have been largely neglected. Here we review recent research on vocal and musical emotions, and outline commonalities and differences in the neural processing of emotions in the TLS in terms of emotional valence, emotional intensity and arousal, as well as in terms of acoustic and structural features of voices and music. We summarize the findings in a neural framework including several subcortical and cortical functional pathways between the auditory system and the TLS. This framework proposes that some vocal expressions might already receive a fast emotional evaluation via a subcortical pathway to the amygdala, whereas cortical pathways to the TLS are thought to be equally used for vocal and musical emotions. While the amygdala might be specifically involved in a coarse decoding of the emotional value of voices and music, the hippocampus might process more complex vocal and musical emotions, and might have an important role especially for the decoding of musical emotions by providing memory-based and contextual associations.

Rhythm implicitly affects temporal orienting of attention across modalities.

Here we present two experiments investigating the implicit orienting of attention over time by entrainment to an auditory rhythmic stimulus. In the first experiment, participants carried out a detection and discrimination tasks with auditory and visual targets while listening to an isochronous, auditory sequence, which acted as the entraining stimulus. For the second experiment, we used musical extracts as entraining stimulus, and tested the resulting strength of entrainment with a visual discrimination task. Both experiments used reaction times as a dependent variable. By manipulating the appearance of targets across four selected metrical positions of the auditory entraining stimulus we were able to observe how entraining to a rhythm modulates behavioural responses. That our results were independent of modality gives a new insight into cross-modal interactions between auditory and visual modalities in the context of dynamic attending to auditory temporal structure.

How musical training affects cognitive development: rhythm, reward and other modulating variables.

Musical training has recently gained additional interest in education as increasing neuroscientific research demonstrates its positive effects on brain development. Neuroimaging revealed plastic changes in the brains of adult musicians but it is still unclear to what extent they are the product of intensive music training rather than of other factors, such as preexisting biological markers of musicality. In this review, we synthesize a large body of studies demonstrating that benefits of musical training extend beyond the skills it directly aims to train and last well into adulthood. For example, children who undergo musical training have better verbal memory, second language pronunciation accuracy, reading ability and executive functions. Learning to play an instrument as a child may even predict academic performance and IQ in young adulthood. The degree of observed structural and functional adaptation in the brain correlates with intensity and duration of practice. Importantly, the effects on cognitive development depend on the timing of musical initiation due to sensitive periods during development, as well as on several other modulating variables. Notably, we point to motivation, reward and social context of musical education, which are important yet neglected factors affecting the long-term benefits of musical training. Further, we introduce the notion of rhythmic entrainment and suggest that it may represent a mechanism supporting learning and development of executive functions. It also hones temporal processing and orienting of attention in time that may underlie enhancements observed in reading and verbal memory. We conclude that musical training uniquely engenders near and far transfer effects, preparing a foundation for a range of skills, and thus fostering cognitive development.