Representing melodic relationships using network science.

Music is a complex system consisting of many dimensions and hierarchically organized information-the organization of which, to date, we do not fully understand. Network science provides a powerful approach to representing such complex systems, from the social networks of people to modelling the underlying network structures of different cognitive mechanisms. In the present research, we explored whether network science methodology can be extended to model the melodic patterns underlying expert improvised music. Using a large corpus of transcribed improvisations, we constructed a network model in which 5-pitch sequences were linked depending on consecutive occurrences, constituting 116,403 nodes (sequences) and 157,429 edges connecting them. We then investigated whether mathematical graph modelling relates to musical characteristics in real-world listening situations via a behavioral experiment paralleling those used to examine language. We found that as melodic distance within the network increased, participants judged melodic sequences as less related. Moreover, the relationship between distance and reaction time (RT) judgements was quadratic: participants slowed in RT up to distance four, then accelerated; a parallel finding to research in language networks. This study offers insights into the hidden network structure of improvised tonal music and suggests that humans are sensitive to the property of melodic distance in this network. More generally, our work demonstrates the similarity between music and language as complex systems, and how network science methods can be used to quantify different aspects of its complexity.

Action-Perception Coupling and Near Transfer: Listening to Melodies after Piano Practice Triggers Sequence-Specific Representations in the Auditory-Motor Network.

Understanding how perception and action are coupled in the brain has important implications for training, rehabilitation, and brain-machine interfaces. Ideomotor theory postulates that willed actions are represented through previously experienced effects and initiated by the anticipation of those effects. Previous research has accordingly found that sensory events, if previously associated with action outcomes, can induce activity in motor regions. However, it remains unclear whether the motor-related activity induced during perception of more naturalistic sequences of actions actually represents "sequence-specific" information. In the present study, nonmusicians were firstly trained to play two melodies on the piano; secondly, they performed an fMRI experiment while listening to these melodies as well as novel, untrained melodies; thirdly, multivariate pattern analysis was used to test if voxel-wise patterns of brain activity could identify trained, but not novel melodies. The results importantly show that after associative learning, a series of sensory events can trigger sequence-specific representations in both sensory and motor networks. Interestingly, also novel melodies could be classified in multiple regions, including default mode regions. A control experiment confirmed these outcomes to be training-dependent. We discuss how action-perception coupling may enable spontaneous near transfer and action simulation during action observation.

Effects of the monoamine stabilizer (-)OSU6162 on cognitive function in alcohol dependence.

INTRODUCTION: Alcohol dependence (AD) is associated with a dysregulated mesolimbocortical dopamine system-a pathway which is also implicated in both reward and cognition. The monoamine stabilizer (-)-OSU6162 (OSU) is a novel pharmacological compound with the ability to reduce ethanol intake and ethanol seeking in long-term drinking rats as well as reducing alcohol craving in AD patients. Dopaminergic drugs can both impair and improve cognitive functions, and the aim of the current study was to investigate the effect of OSU treatment on cognitive functioning in AD patients. METHOD: In a randomized double-blind placebo-controlled study, 56 individuals with AD received 14 days of OSU or placebo treatment. Neuropsychological tasks from the Cambridge Automated Neuropsychological Test Battery (CANTAB®) and other tasks were used to evaluate treatment effect on executive function/impulsivity, working memory, attention, emotional recognition, and divergent thinking. RESULTS: Treatment with OSU did not impair neuropsychological function in any of the cognitive domains investigated (all p > 0.1). In fact, OSU treatment did, compared to placebo, improve future planning ability (F(1,46) = 6.9; p = 0.012; Cohen's d = 0.54), verbal divergent thinking (F(1,44) = 10.1; p = 0.003; d = 0.96), and response time for emotional recognition (F(1,47) = 6.7; p = 0.013; d = 0.44). CONCLUSION: OSU treatment did not cause short-term cognitive side effects, further supporting the potential of OSU as a clinically feasible pharmacological treatment in AD patients. OSU treatment might improve future planning, verbal divergent thinking, and emotional recognition latency, which in turn may have a beneficial impact on alcohol use outcomes. Future studies are needed to confirm these preliminary findings.

Same Genes, Different Brains: Neuroanatomical Differences Between Monozygotic Twins Discordant for Musical Training.

Numerous cross-sectional and observational longitudinal studies show associations between expertise and regional brain anatomy. However, since these designs confound training with genetic predisposition, the causal role of training remains unclear. Here, we use a discordant monozygotic (identical) twin design to study expertise-dependent effects on neuroanatomy using musical training as model behavior, while essentially controlling for genetic factors and shared environment of upbringing. From a larger cohort of monozygotic twins, we were able to recruit 18 individuals (9 pairs) that were highly discordant for piano practice. We used structural and diffusion magnetic resonance imaging to analyze the auditory-motor network and within-pair differences in cortical thickness, cerebellar regional volumes and white-matter microstructure/fractional anisotropy. The analyses revealed that the musically active twins had greater cortical thickness in the auditory-motor network of the left hemisphere and more developed white matter microstructure in relevant tracts in both hemispheres and the corpus callosum. Furthermore, the volume of gray matter in the left cerebellar region of interest comprising lobules I-IV + V, was greater in the playing group. These findings provide the first clear support for that a significant portion of the differences in brain anatomy between experts and nonexperts depend on causal effects of training.

On the Relationship Between Domain-Specific Creative Achievement and Sexual Orientation in Swedish Twins.

Despite the commonly held belief that homosexual males and females are more creative compared to heterosexuals, empirical studies on homosexuality and its relationship to creativity have been sparse, often with questionable methodology and very small sample sizes, reporting mixed findings. No study till date has explored the associations described above in a large population-based and genetically informative sample. Here, we examined such potential associations between sexual orientation and creative achievement in several different domains (music, writing, dance, visual arts, science, invention, and theater) using a large cohort of 4494 Swedish twins (of which 7.5 % were not exclusively heterosexual). Data were analyzed for the sexes separately as well as pooled. Results showed significant associations between sexual orientation and two of the creative domains-theater and writing-with non-heterosexuals being more creative in these domains. In all other domains, no significant differences were found between the non-heterosexual and heterosexual groups. Findings from co-twin control analyses suggested that the significant associations may not be causal in nature (i.e., homosexual orientation leads to higher creativity) but due to shared liability. However, we lacked power to differentiate between shared genetic and shared environmental influences. Results and potential implications are discussed critically.

Self-reported psychological demands, skill discretion and decision authority at work: A twin study.

OBJECTIVES: To examine the contribution of genetic factors to self-reported psychological demands (PD), skill discretion (SD) and decision authority (DA) and the possible importance of such influence on the association between these work variables and depressive symptoms. METHODS: 11,543 participants aged 27-54 in the Swedish Twin Registry participated in a web survey. First of all, in multiple regressions, phenotypic associations between each one of the three work environment variables and depressive symptoms were analysed. Secondly, by means of classical twin analysis, the genetic contribution to PD, SD and DA was assessed. After this, cross-twin cross-trait correlations were computed between PD, SD and DA, on the one hand, and depressive symptom score, on the other hand. RESULTS: The genetic contribution to self-reported PD, DS and DA ranged from 18% for decision authority to 30% for skill discretion. Cross-twin cross-trait correlations were very weak (r values < .1) and non-significant for dizygotic twins, and we lacked power to analyse the genetic architecture of the phenotypic associations using bivariate twin modelling. However, substantial genetic contribution to these associations seems unlikely. CONCLUSIONS GENETIC CONTRIBUTIONS TO THE SELF-REPORTED WORK ENVIRONMENT SCORES WERE 18-30%.

Addressing a Paradox: Dual Strategies for Creative Performance in Introspective and Extrospective Networks.

Neuroimaging studies of internally generated behaviors have shown seemingly paradoxical results regarding the dorsolateral prefrontal cortex (DLPFC), which has been found to activate, not activate or even deactivate relative to control conditions. On the one hand, the DLPFC has been argued to exert top-down control over generative thought by inhibiting habitual responses; on the other hand, a deactivation and concomitant decrease in monitoring and focused attention has been suggested to facilitate spontaneous associations and novel insights. Here, we demonstrate that prefrontal engagement in creative cognition depends dramatically on experimental conditions, that is, the goal of the task. We instructed professional pianists to perform improvisations on a piano keyboard during fMRI and play, either with a certain emotional content (happy/fearful), or using certain keys (tonal/atonal pitch-sets). We found lower activity in primarily the right DLPFC, dorsal premotor cortex and inferior parietal cortex during emotional conditions compared with pitch-set conditions. Furthermore, the DLPFC was functionally connected to the default mode network during emotional conditions and to the premotor network during pitch-set conditions. The results thus support the notion of two broad cognitive strategies for creative problem solving, relying on extrospective and introspective neural circuits, respectively.

Physiological correlates of the flow experience during computer game playing.

Flow is the subjective experience of effortless attention, reduced self-awareness, and enjoyment that typically occurs during optimal task performance. Previous studies have suggested that flow may be associated with a non-reciprocal coactivation of the sympathetic and parasympathetic systems and, on a cortical level, with a state of hypofrontality and implicit processing. Here, we test these hypotheses, using the computer game TETRIS as model task. The participants (n=77) played TETRIS under three conditions that differed in difficulty (Easy

Connecting to create: expertise in musical improvisation is associated with increased functional connectivity between premotor and prefrontal areas.

Musicians have been used extensively to study neural correlates of long-term practice, but no studies have investigated the specific effects of training musical creativity. Here, we used human functional MRI to measure brain activity during improvisation in a sample of 39 professional pianists with varying backgrounds in classical and jazz piano playing. We found total hours of improvisation experience to be negatively associated with activity in frontoparietal executive cortical areas. In contrast, improvisation training was positively associated with functional connectivity of the bilateral dorsolateral prefrontal cortices, dorsal premotor cortices, and presupplementary areas. The effects were significant when controlling for hours of classical piano practice and age. These results indicate that even neural mechanisms involved in creative behaviors, which require a flexible online generation of novel and meaningful output, can be automated by training. Second, improvisational musical training can influence functional brain properties at a network level. We show that the greater functional connectivity seen in experienced improvisers may reflect a more efficient exchange of information within associative networks of importance for musical creativity.

Individual differences in the proneness to have flow experiences are linked to dopamine D2-receptor availability in the dorsal striatum.

Flow is a subjective experience of high but effortless attention, enjoyment, and low self-awareness that can occur during the active performance of challenging tasks. The dispositional proneness to experience flow is associated with personality traits that are known to be influenced by dopaminergic neural systems. Here, for the first time, we investigated relations between flow proneness and dopaminergic function. Specifically, we tested the hypothesis that the availability of dopamine D2-receptors in the striatum is positively associated with flow proneness. Striatal D2-receptor availability was measured in a sample of 25 healthy adults using positron emission tomography and [(11)C]raclopride. Flow proneness was measured using the Swedish Flow Proneness Questionnaire. As hypothesized, there was a significant correlation (r=.41) between striatal D2-receptor availability and flow proneness. An exploratory analysis of striatal subregions showed that the relation was mainly driven by the dorsal striatum, with a significantly higher correlation in the putamen than in the ventral striatum. The findings constitute the first demonstration of an association between flow proneness and dopaminergic function. We suggest that the proneness to experience flow is related to personality dimensions that are under dopaminergic control and characterized by low impulsiveness, stable emotion, and positive affect.

Activation and connectivity patterns of the presupplementary and dorsal premotor areas during free improvisation of melodies and rhythms.

Free, i.e. non-externally cued generation of movement sequences is fundamental to human behavior. We have earlier hypothesized that the dorsal premotor cortex (PMD), which has been consistently implicated in cognitive aspects of planning and selection of spatial motor sequences may be particularly important for the free generation of spatial movement sequences, whereas the pre-supplementary motor area (pre-SMA), which shows increased activation during perception, learning and reproduction of temporal sequences, may contribute more to the generation of temporal structures. Here we test this hypothesis using fMRI and musical improvisation in professional pianists as a model behavior. We employed a 2 × 2 factorial design with the factors Melody (Specified/Improvised) and Rhythm (Specified/Improvised). The main effect analyses partly confirmed our hypothesis: there was a main effect of Melody in the PMD; the pre-SMA was present in the main effect of Rhythm, as predicted, as well as in the main effect of Melody. A psychophysiological interaction analysis of functional connectivity demonstrated that the correlation in activity between the pre-SMA and cerebellum was higher during rhythmic improvisation than during the other conditions. In summary, there were only subtle differences in activity level between the pre-SMA and PMD during improvisation, regardless of condition. Consequently, the free generation of rhythmic and melodic structures, appears to be largely integrated processes but the functional connectivity between premotor areas and other regions may change during free generation in response to sequence-specific spatiotemporal demands.

Differences in regional brain volume related to the extraversion-introversion dimension--a voxel based morphometry study.

Extraverted individuals are sociable, behaviorally active, and happy. We report data from a voxel based morphometry study investigating, for the first time, if regional volume in gray and white matter brain regions is related to extraversion. For both gray and white matter, all correlations between extraversion and regional brain volume were negative, i.e. the regions were larger in introverts. Gray matter correlations were found in regions that included the right prefrontal cortex and the cortex around the right temporo-parietal junction--regions that are known to be involved in behavioral inhibition, introspection, and social-emotional processing, e.g. evaluation of social stimuli and reasoning about the mental states of others. White matter correlations extended from the brainstem to widespread cortical regions, and were largely due to global effects, i.e. a larger total white matter volume in introverts. We speculate that these white matter findings may reflect differences in ascending modulatory projections affecting cortical regions involved in behavioral regulation.

Goal-independent mechanisms for free response generation: creative and pseudo-random performance share neural substrates.

To what extent free response generation in different tasks uses common and task-specific neurocognitive processes has remained unclear. Here, we investigated overlap and differences in neural activity during musical improvisation and pseudo-random response generation. Brain activity was measured using fMRI in a group of professional classical pianists, who performed musical improvisation of melodies, pseudo-random key-presses and a baseline condition (sight-reading), on either two, six or twelve keys on a piano keyboard. The results revealed an extensive overlap in neural activity between the two generative conditions. Active regions included the dorsolateral and dorsomedial prefrontal cortices, inferior frontal gyrus, anterior cingulate cortex and pre-SMA. No regions showed higher activity in improvisation than in pseudo-random generation. These findings suggest that the activated regions fulfill generic functions that are utilized in different types of free generation tasks, independent of overall goal. In contrast, pseudo-random generation was accompanied by higher activity than improvisation in several regions. This presumably reflects the participants' musical expertise as well as the pseudo-random generation task's high load on attention, working memory, and executive control. The results highlight the significance of using naturalistic tasks to study human behavior and cognition. No brain activity was related to the size of the response set. We discuss that this may reflect that the musicians were able to use specific strategies for improvisation, by which there was no simple relationship between response set size and neural activity.

Dissociation of brain areas associated with force production and stabilization during manipulation of unstable objects.

Multifinger dexterous manipulation of unstable or deformable objects requires control of both direction and magnitude of fingertip force vectors. Our aim was to study the neuroanatomical correlates of these two distinct control functions. Brain activity was measured using functional magnetic resonance imaging while 16 male subjects (age: 26-42, M = 32, SD ± 4 years) compressed four springs representing a 2 × 2 factorial design with two levels of force and instability requirements. Significant activations associated with higher instability were located bilaterally in the precentral gyri, the postcentral gyrus, and the cerebellum. In the main effect for high force, activity was found in areas located in the primary motor regions contralateral to the active hand and bilaterally in the cerebellum. An overlap in activation between the two main effects was found bilaterally in the cerebellum (lobule VI). This study not only confirms a recently described bilateral fronto-parieto-cerebellar network for manipulation of increasingly unstable objects, but critically extends our understanding by describing its differentiated modulation with both force magnitude and instability requirements. Our results, therefore, expose a previously unrecognized and context-sensitive system of brain regions that enable dexterous manipulation for different force magnitude and instability requirements of the task.

The psychophysiology of flow during piano playing.

Expert performance is commonly accompanied by a subjective state of optimal experience called flow. Previous research has shown positive correlations between flow and quality of performance and suggests that flow may function as a reward signal that promotes practice. Here, piano playing was used as a flow-inducing behavior in order to analyze the relationship between subjective flow reports and psychophysiological measures. Professional classical pianists were asked to play a musical piece and then rate state flow. The performance was repeated five times in order to induce a variation in flow, keeping other factors constant, while recording the arterial pulse pressure waveform, respiration, head movements, and activity from the corrugator supercilii and zygomaticus major facial muscles. A significant relation was found between flow and heart period, blood pressure, heart rate variability, activity of the zygomaticus major muscle, and respiratory depth. These findings are discussed in relation to current models of emotion, attention, and expertise, and flow is proposed to be a state of effortless attention, which arises through an interaction between positive affect and high attention.

Thinking outside a less intact box: thalamic dopamine D2 receptor densities are negatively related to psychometric creativity in healthy individuals.

Several lines of evidence support that dopaminergic neurotransmission plays a role in creative thought and behavior. Here, we investigated the relationship between creative ability and dopamine D2 receptor expression in healthy individuals, with a focus on regions where aberrations in dopaminergic function have previously been associated with psychotic symptoms and a genetic liability to schizophrenia. Scores on divergent thinking tests (Inventiveness battery, Berliner Intelligenz Struktur Test) were correlated with regional D2 receptor densities, as measured by Positron Emission Tomography, and the radioligands [(11)C]raclopride and [(11)C]FLB 457. The results show a negative correlation between divergent thinking scores and D2 density in the thalamus, also when controlling for age and general cognitive ability. Hence, the results demonstrate that the D2 receptor system, and specifically thalamic function, is important for creative performance, and may be one crucial link between creativity and psychopathology. We suggest that decreased D2 receptor densities in the thalamus lower thalamic gating thresholds, thus increasing thalamocortical information flow. In healthy individuals, who do not suffer from the detrimental effects of psychiatric disease, this may increase performance on divergent thinking tests. In combination with the cognitive functions of higher order cortical networks, this could constitute a basis for the generative and selective processes that underlie real life creativity.

Dopamine D2 receptor density in the limbic striatum is related to implicit but not explicit movement sequence learning.

A large body of literature suggests that motor sequence learning involves dopamine-modulated plastic processes in the basal ganglia. Sequence learning can occur both implicitly, without conscious awareness and intention to learn, and explicitly, i.e., under conscious control. Here, we investigated whether individual differences in implicit and explicit sequence learning of movement sequences in a group of 15 healthy participants are related to dopamine D2 receptor densities in functional subregions of the striatum. Sequence learning was assessed using the serial reaction time task, and measures of implicit and explicit knowledge were estimated using a process dissociation procedure. Correlation analyses were performed between these measures and D2 receptor densities, which had been measured previously with positron emission tomography. Striatal D2 densities were negatively related to measures of sequence learning. In the limbic subregion, D2 densities were specifically related to implicit but not explicit learning. These findings suggest that individual differences in striatal DA function underlie differences in sequence learning ability and support that implicit and explicit sequence learning depend on partly distinct neural circuitry. The findings are also in line with the general view that implicit learning systems are evolutionarily primitive and tend to rely more on phylogenetically old neural circuitry than does explicit learning and cognition.