It's not what you say but the way that you say it: an fMRI study of differential lexical and non-lexical prosodic pitch processing

<p>Abstract</p> <p>Background</p> <p>This study aims to identify the neural substrate involved in prosodic pitch processing. Functional magnetic resonance imaging was used to test the premise that prosody pitch processing is primarily subserved by the right cortical hemisphere.</p> <p>Two experimental paradigms were used, firstly pairs of spoken sentences, where the only variation was a single internal phrase pitch change, and secondly, a matched condition utilizing pitch changes within analogous tone-sequence phrases. This removed the potential confounder of lexical evaluation. fMRI images were obtained using these paradigms.</p> <p>Results</p> <p>Activation was significantly greater within the right frontal and temporal cortices during the tone-sequence stimuli relative to the sentence stimuli.</p> <p>Conclusion</p> <p>This study showed that pitch changes, stripped of lexical information, are mainly processed by the right cerebral hemisphere, whilst the processing of analogous, matched, lexical pitch change is preferentially left sided. These findings, showing hemispherical differentiation of processing based on stimulus complexity, are in accord with a 'task dependent' hypothesis of pitch processing.</p

Results of a pilot study on the involvement of bilateral inferior frontal gyri in emotional prosody perception: an rTMS study

<p>Abstract</p> <p>Background</p> <p>The right hemisphere may play an important role in paralinguistic features such as the emotional melody in speech. The extent of this involvement however is unclear. Imaging studies have shown involvement of both left and right inferior frontal gyri in emotional prosody perception. The present pilot study examined whether these brain areas are critically involved in the processing of emotional prosody and of semantics in 9 healthy subjects. Repetitive transcranial magnetic stimulation was used with a coil centred over left and right inferior frontal gyri, as localized by neuronavigation based on the subject's MRI. A sham condition was included. An online-TMS approach was applied; an emotional language task was completed during stimulation. This computerized task consisted of sentences pronounced by actors. In the semantics condition an emotion (fear, anger or neutral) was expressed in the content pronounced with a neutral intonation. In the prosody condition the emotion was expressed in the intonation, while the content was neutral.</p> <p>Results</p> <p>Reaction times on the emotional prosody task condition were significantly longer after rTMS over both the right and the left inferior frontal gyrus as compared to sham stimulation and after controlling for learning effects associated with order of condition. When taking all emotions together, there was no difference in effect on reaction times between the right and left stimulation. For the emotion Fear, reaction times were significantly longer after stimulating the left inferior frontal gyrus as compared to the right inferior frontal gyrus. Reaction times in the semantics task condition were not significantly different between the three TMS conditions.</p> <p>Conclusions</p> <p>The data indicate a critical involvement of both the right and the left inferior frontal gyrus in emotional prosody perception. The findings of this pilot study need replication. Future studies should include more subjects and examine whether the left and right inferior frontal gyrus play a differential role and complement each other, e.g. in the integrated processing of linguistic and prosodic aspects of speech, respectively.</p

The hearing hippocampus

The hippocampus has a well-established role in spatial and episodic memory but a broader function has been proposed including aspects of perception and relational processing. Neural bases of sound analysis have been described in the pathway to auditory cortex, but wider networks supporting auditory cognition are still being established. We review what is known about the role of the hippocampus in processing auditory information, and how the hippocampus itself is shaped by sound. In examining imaging, recording, and lesion studies in species from rodents to humans, we uncover a hierarchy of hippocampal responses to sound including during passive exposure, active listening, and the learning of associations between sounds and other stimuli. We describe how the hippocampus' connectivity and computational architecture allow it to track and manipulate auditory information – whether in the form of speech, music, or environmental, emotional, or phantom sounds. Functional and structural correlates of auditory experience are also identified. The extent of auditory-hippocampal interactions is consistent with the view that the hippocampus makes broad contributions to perception and cognition, beyond spatial and episodic memory. More deeply understanding these interactions may unlock applications including entraining hippocampal rhythms to support cognition, and intervening in links between hearing loss and dementia

Hidden sources of joy, fear, and sadness : Explicit versus implicit neural processing of musical emotions

Music is often used to regulate emotions and mood. Typically, music conveys and induces emotions even when one does not attend to them. Studies on the neural substrates of musical emotions have, however, only examined brain activity when subjects have focused on the emotional content of the music. Here we address with functional magnetic resonance imaging (fMRI) the neural processing of happy, sad, and fearful music with a paradigm in which 56 subjects were instructed to either classify the emotions (explicit condition) or pay attention to the number of instruments playing (implicit condition) in 4-s music clips. In the implicit vs. explicit condition, stimuli activated bilaterally the inferior parietal lobule, premotor cortex, caudate, and ventromedial frontal areas. The cortical dorsomedial prefrontal and occipital areas activated during explicit processing were those previously shown to be associated with the cognitive processing of music and emotion recognition and regulation. Moreover, happiness in music was associated with activity in the bilateral auditory cortex, left parahippocampal gyrus, and supplementary motor area, whereas the negative emotions of sadness and fear corresponded with activation of the left anterior cingulate and middle frontal gyrus and down-regulation of the orbitofrontal cortex. Our study demonstrates for the first time in healthy subjects the neural underpinnings of the implicit processing of brief musical emotions, particularly in frontoparietal, dorsolateral prefrontal, and striatal areas of the brain. (C) 2016 Elsevier Ltd. All rights reserved.Peer reviewe

Multiple prosodic meanings are conveyed through separate pitch ranges: Evidence from perception of focus and surprise in Mandarin Chinese

F0 variation is a crucial feature in speech prosody, which can convey linguistic information such as focus and paralinguistic meanings such as surprise. How can multiple layers of information be represented with F0 in speech: are they divided into discrete layers of pitch or overlapped without clear divisions? We investigated this question by assessing pitch perception of focus and surprise in Mandarin Chinese. Seventeen native Mandarin listeners rated the strength of focus and surprise conveyed by the same set of synthetically manipulated sentences. An fMRI experiment was conducted to assess neural correlates of the listeners’ perceptual response to the stimuli. The results showed that behaviourally, the perceptual threshold for focus was 3 semitones and that for surprise was 5 semitones above the baseline. Moreover, the pitch range of 5-12 semitones above the baseline signalled both focus and surprise, suggesting a considerable overlap between the two types of prosodic information within this range. The neuroimaging data positively correlated with the variations in behavioural data. Also, a ceiling effect was found as no significant behavioural differences or neural activities were shown after reaching a certain pitch level for the perception of focus and surprise respectively. Together, the results suggest that different layers of prosodic information are represented in F0 through different pitch ranges: paralinguistic information is represented at a pitch range beyond that used by linguistic information. Meanwhile, the representation of paralinguistic information is achieved without obscuring linguistic prosody, thus allowing F0 to represent the two layers of information in parallel

Music in the brain

Music is ubiquitous across human cultures — as a source of affective and pleasurable experience, moving us both physically and emotionally — and learning to play music shapes both brain structure and brain function. Music processing in the brain — namely, the perception of melody, harmony and rhythm — has traditionally been studied as an auditory phenomenon using passive listening paradigms. However, when listening to music, we actively generate predictions about what is likely to happen next. This enactive aspect has led to a more comprehensive understanding of music processing involving brain structures implicated in action, emotion and learning. Here we review the cognitive neuroscience literature of music perception. We show that music perception, action, emotion and learning all rest on the human brain’s fundamental capacity for prediction — as formulated by the predictive coding of music model. This Review elucidates how this formulation of music perception and expertise in individuals can be extended to account for the dynamics and underlying brain mechanisms of collective music making. This in turn has important implications for human creativity as evinced by music improvisation. These recent advances shed new light on what makes music meaningful from a neuroscientific perspective

Social relationship-dependent neural response to speech in dogs: Social relationship-based neural responses in dogs

In humans, social relationship with the speaker affects neural processing of speech, as exemplified by children's auditory and reward responses to their mother's utterances. Family dogs show human analogue attachment behavior towards the owner, and neuroimaging revealed auditory cortex and reward center sensitivity to verbal praises in dog brains. Combining behavioral and non-invasive fMRI data, we investigated the effect of dogs' social relationship with the speaker on speech processing. Dogs listened to praising and neutral speech from their owners and a control person. We found positive correlation between dogs' behaviorally measured attachment scores towards their owners and neural activity increase for the owner's voice in the caudate nucleus; and activity increase in the secondary auditory caudal ectosylvian gyrus and the caudate nucleus for the owner's praise. Through identifying social relationship-dependent neural reward responses, our study reveals similarities in neural mechanisms modulated by infant-mother and dog-owner attachment

Predictive cognition in dementia: the case of music

The clinical complexity and pathological diversity of neurodegenerative diseases impose immense challenges for diagnosis and the design of rational interventions. To address these challenges, there is a need to identify new paradigms and biomarkers that capture shared pathophysiological processes and can be applied across a range of diseases. One core paradigm of brain function is predictive coding: the processes by which the brain establishes predictions and uses them to minimise prediction errors represented as the difference between predictions and actual sensory inputs. The processes involved in processing unexpected events and responding appropriately are vulnerable in common dementias but difficult to characterise. In my PhD work, I have exploited key properties of music – its universality, ecological relevance and structural regularity – to model and assess predictive cognition in patients representing major syndromes of frontotemporal dementia – non-fluent variant PPA (nfvPPA), semantic-variant PPA (svPPA) and behavioural-variant FTD (bvFTD) - and Alzheimer’s disease relative to healthy older individuals. In my first experiment, I presented patients with well-known melodies containing no deviants or one of three types of deviant - acoustic (white-noise burst), syntactic (key-violating pitch change) or semantic (key-preserving pitch change). I assessed accuracy detecting melodic deviants and simultaneously-recorded pupillary responses to these deviants. I used voxel-based morphometry to define neuroanatomical substrates for the behavioural and autonomic processing of these different types of deviants, and identified a posterior temporo-parietal network for detection of basic acoustic deviants and a more anterior fronto-temporo-striatal network for detection of syntactic pitch deviants. In my second chapter, I investigated the ability of patients to track the statistical structure of the same musical stimuli, using a computational model of the information dynamics of music to calculate the information-content of deviants (unexpectedness) and entropy of melodies (uncertainty). I related these information-theoretic metrics to performance for detection of deviants and to ‘evoked’ and ‘integrative’ pupil reactivity to deviants and melodies respectively and found neuroanatomical correlates in bilateral dorsal and ventral striatum, hippocampus, superior temporal gyri, right temporal pole and left inferior frontal gyrus. Together, chapters 3 and 4 revealed new hypotheses about the way FTD and AD pathologies disrupt the integration of predictive errors with predictions: a retained ability of AD patients to detect deviants at all levels of the hierarchy with a preserved autonomic sensitivity to information-theoretic properties of musical stimuli; a generalized impairment of surprise detection and statistical tracking of musical information at both a cognitive and autonomic levels for svPPA patients underlying a diminished precision of predictions; the exact mirror profile of svPPA patients in nfvPPA patients with an abnormally high rate of false-alarms with up-regulated pupillary reactivity to deviants, interpreted as over-precise or inflexible predictions accompanied with normal cognitive and autonomic probabilistic tracking of information; an impaired behavioural and autonomic reactivity to unexpected events with a retained reactivity to environmental uncertainty in bvFTD patients. Chapters 5 and 6 assessed the status of reward prediction error processing and updating via actions in bvFTD. I created pleasant and aversive musical stimuli by manipulating chord progressions and used a classic reinforcement-learning paradigm which asked participants to choose the visual cue with the highest probability of obtaining a musical ‘reward’. bvFTD patients showed reduced sensitivity to the consequence of an action and lower learning rate in response to aversive stimuli compared to reward. These results correlated with neuroanatomical substrates in ventral and dorsal attention networks, dorsal striatum, parahippocampal gyrus and temporo-parietal junction. Deficits were governed by the level of environmental uncertainty with normal learning dynamics in a structured and binarized environment but exacerbated deficits in noisier environments. Impaired choice accuracy in noisy environments correlated with measures of ritualistic and compulsive behavioural changes and abnormally reduced learning dynamics correlated with behavioural changes related to empathy and theory-of-mind. Together, these experiments represent the most comprehensive attempt to date to define the way neurodegenerative pathologies disrupts the perceptual, behavioural and physiological encoding of unexpected events in predictive coding terms