Predictions and the brain: how musical sounds become rewarding.

Music has always played a central role in human culture. The question of how musical sounds can have such profound emotional and rewarding effects has been a topic of interest throughout generations. At a fundamental level, listening to music involves tracking a series of sound events over time. Because humans are experts in pattern recognition, temporal predictions are constantly generated, creating a sense of anticipation. We summarize how complex cognitive abilities and cortical processes integrate with fundamental subcortical reward and motivation systems in the brain to give rise to musical pleasure. This work builds on previous theoretical models that emphasize the role of prediction in music appreciation by integrating these ideas with recent neuroscientific evidence.

Familiarity mediates the relationship between emotional arousal and pleasure during music listening.

Emotional arousal appears to be a major contributing factor to the pleasure that listeners experience in response to music. Accordingly, a strong positive correlation between self-reported pleasure and electrodermal activity (EDA), an objective indicator of emotional arousal, has been demonstrated when individuals listen to familiar music. However, it is not yet known to what extent familiarity contributes to this relationship. In particular, as listening to familiar music involves expectations and predictions over time based on veridical knowledge of the piece, it could be that such memory factors plays a major role. Here, we tested such a contribution by using musical stimuli entirely unfamiliar to listeners. In a second experiment we repeated the novel music to experimentally establish a sense of familiarity. We aimed to determine whether (1) pleasure and emotional arousal would continue to correlate when listeners have no explicit knowledge of how the tones will unfold, and (2) whether this could be enhanced by experimentally-induced familiarity. In the first experiment, we presented 33 listeners with 70 unfamiliar musical excerpts in two sessions. There was no relationship between the degree of experienced pleasure and emotional arousal as measured by EDA. In the second experiment, 7 participants listened to 35 unfamiliar excerpts over two sessions separated by 30 min. Repeated exposure significantly increased EDA, even though individuals did not explicitly recall having heard all the pieces before. Furthermore, increases in self-reported familiarity significantly enhanced experienced pleasure and there was a general, though not significant, increase in EDA. These results suggest that some level of expectation and predictability mediated by prior exposure to a given piece of music play an important role in the experience of emotional arousal in response to music.

From perception to pleasure: music and its neural substrates.

Music has existed in human societies since prehistory, perhaps because it allows expression and regulation of emotion and evokes pleasure. In this review, we present findings from cognitive neuroscience that bear on the question of how we get from perception of sound patterns to pleasurable responses. First, we identify some of the auditory cortical circuits that are responsible for encoding and storing tonal patterns and discuss evidence that cortical loops between auditory and frontal cortices are important for maintaining musical information in working memory and for the recognition of structural regularities in musical patterns, which then lead to expectancies. Second, we review evidence concerning the mesolimbic striatal system and its involvement in reward, motivation, and pleasure in other domains. Recent data indicate that this dopaminergic system mediates pleasure associated with music; specifically, reward value for music can be coded by activity levels in the nucleus accumbens, whose functional connectivity with auditory and frontal areas increases as a function of increasing musical reward. We propose that pleasure in music arises from interactions between cortical loops that enable predictions and expectancies to emerge from sound patterns and subcortical systems responsible for reward and valuation.

Interactions between the nucleus accumbens and auditory cortices predict music reward value.

We used functional magnetic resonance imaging to investigate neural processes when music gains reward value the first time it is heard. The degree of activity in the mesolimbic striatal regions, especially the nucleus accumbens, during music listening was the best predictor of the amount listeners were willing to spend on previously unheard music in an auction paradigm. Importantly, the auditory cortices, amygdala, and ventromedial prefrontal regions showed increased activity during listening conditions requiring valuation, but did not predict reward value, which was instead predicted by increasing functional connectivity of these regions with the nucleus accumbens as the reward value increased. Thus, aesthetic rewards arise from the interaction between mesolimbic reward circuitry and cortical networks involved in perceptual analysis and valuation.

Anatomically distinct dopamine release during anticipation and experience of peak emotion to music.

Music, an abstract stimulus, can arouse feelings of euphoria and craving, similar to tangible rewards that involve the striatal dopaminergic system. Using the neurochemical specificity of [(11)C]raclopride positron emission tomography scanning, combined with psychophysiological measures of autonomic nervous system activity, we found endogenous dopamine release in the striatum at peak emotional arousal during music listening. To examine the time course of dopamine release, we used functional magnetic resonance imaging with the same stimuli and listeners, and found a functional dissociation: the caudate was more involved during the anticipation and the nucleus accumbens was more involved during the experience of peak emotional responses to music. These results indicate that intense pleasure in response to music can lead to dopamine release in the striatal system. Notably, the anticipation of an abstract reward can result in dopamine release in an anatomical pathway distinct from that associated with the peak pleasure itself. Our results help to explain why music is of such high value across all human societies.

Neural basis of repetition priming during mathematical cognition: repetition suppression or repetition enhancement?

We investigated the neural basis of repetition priming (RP) during mathematical cognition. Previous studies of RP have focused on repetition suppression as the basis of behavioral facilitation, primarily using word and object identification and classification tasks. More recently, researchers have suggested associative stimulus-response learning as an alternate model for behavioral facilitation. We examined the neural basis of RP during mathematical problem solving in the context of these two models of learning. Brain imaging and behavioral data were acquired from 39 adults during novel and repeated presentation of three-operand mathematical equations. Despite wide-spread decreases in activation during repeat, compared with novel trials, there was no direct relation between behavioral facilitation and the degree of repetition suppression in any brain region. Rather, RT improvements were directly correlated with repetition enhancement in the hippocampus and the posteromedial cortex [posterior cingulate cortex, precuneus, and retrosplenial cortex; Brodmann's areas (BAs) 23, 7, and 30, respectively], regions known to support memory formation and retrieval, and in the SMA (BA 6) and the dorsal midcingulate ("motor cingulate") cortex (BA 24d), regions known to be important for motor learning. Furthermore, improvements in RT were also correlated with increased functional connectivity of the hippocampus with both the SMA and the dorsal midcingulate cortex. Our findings provide novel support for the hypothesis that repetition enhancement and associated stimulus-response learning may facilitate behavioral performance during problem solving.

The rewarding aspects of music listening are related to degree of emotional arousal.

BACKGROUND: Listening to music is amongst the most rewarding experiences for humans. Music has no functional resemblance to other rewarding stimuli, and has no demonstrated biological value, yet individuals continue listening to music for pleasure. It has been suggested that the pleasurable aspects of music listening are related to a change in emotional arousal, although this link has not been directly investigated. In this study, using methods of high temporal sensitivity we investigated whether there is a systematic relationship between dynamic increases in pleasure states and physiological indicators of emotional arousal, including changes in heart rate, respiration, electrodermal activity, body temperature, and blood volume pulse. METHODOLOGY: Twenty-six participants listened to self-selected intensely pleasurable music and "neutral" music that was individually selected for them based on low pleasure ratings they provided on other participants' music. The "chills" phenomenon was used to index intensely pleasurable responses to music. During music listening, continuous real-time recordings of subjective pleasure states and simultaneous recordings of sympathetic nervous system activity, an objective measure of emotional arousal, were obtained. PRINCIPAL FINDINGS: Results revealed a strong positive correlation between ratings of pleasure and emotional arousal. Importantly, a dissociation was revealed as individuals who did not experience pleasure also showed no significant increases in emotional arousal. CONCLUSIONS/SIGNIFICANCE: These results have broader implications by demonstrating that strongly felt emotions could be rewarding in themselves in the absence of a physically tangible reward or a specific functional goal.