Developmental refinement of cortical systems for speech and voice processing

Development typically leads to optimized and adaptive neural mechanisms for the processing of voice and speech. In this fMRI study we investigated how this adaptive processing reaches its mature efficiency by examining the effects of task, age and phonological skills on cortical responses to voice and speech in children (8-9years), adolescents (14-15years) and adults. Participants listened to vowels (/a/, /i/, /u/) spoken by different speakers (boy, girl, man) and performed delayed-match-to-sample tasks on vowel and speaker identity. Across age groups, similar behavioral accuracy and comparable sound evoked auditory cortical fMRI responses were observed. Analysis of task-related modulations indicated a developmental enhancement of responses in the (right) superior temporal cortex during the processing of speaker information. This effect was most evident through an analysis based on individually determined voice sensitive regions. Analysis of age effects indicated that the recruitment of regions in the temporal-parietal cortex and posterior cingulate/cingulate gyrus decreased with development. Beyond age-related changes, the strength of speech-evoked activity in left posterior and right middle superior temporal regions significantly scaled with individual differences in phonological skills. Together, these findings suggest a prolonged development of the cortical functional network for speech and voice processing. This development includes a progressive refinement of the neural mechanisms for the selection and analysis of auditory information relevant to the ongoing behavioral task

Examining phonological processing in the healthy and damaged brain

In this thesis I explore the neural signature of different types of speech sound processing, in the healthy brain and after damage through stroke. The first two Experiments applied a newly developed fMRI language paradigm in healthy controls to study phonological retrieval from speech, orthography and semantics. This showed that there are at least two types of phonological processing that can be dissociated on a neuronal level. Bilateral superior temporal gyri were associated with processing auditory (phonological) representations of speech, consistent with the notion of input to phonology. In contrast, left putamen and precentral cortex/pars opercularis were associated with pre-articulatory activity, and thus with outputs from phonology. The validation of the results in a separate, larger sample increased confidence that these findings are robust rather than false positives. Experiment 3 was concerned with examining the role of a “key player” in phonological processing, which revealed that different parts of the supramarginal gyrus differ in their response profile during a set of language tasks. This is in accordance with cytoarchitectural and connectivity studies demonstrating the structural variability of the region, and has implications for prior imaging studies considering the supramarginal gyrus as a uniform entity in the phonological network. The final experiment revealed that the loss of supramarginal gyrus through stroke has inconsistent effects on language abilities, possibly due to other brain regions or white matter tracts that were damaged in some patients but not in others. It also showed that additional brain regions were recruited in patients compared to controls, which might reflect compensatory brain activation that supports recovery. Taken together, this work proposes a new way of interpreting phonological effects, in particular within the supramarginal gyrus, and new insights into how the brain supports phonological processing after stroke-induced damage

Neural representations used by brain regions underlying speech production

Thesis (Ph.D.)--Boston UniversitySpeech utterances are phoneme sequences but may not always be represented as such in the brain. For instance, electropalatography evidence indicates that as speaking rate increases, gestures within syllables are manipulated separately but those within consonant clusters act as one motor unit. Moreover, speech error data suggest that a syllable's phonological content is, at some stage, represented separately from its syllabic frame structure. These observations indicate that speech is neurally represented in multiple forms. This dissertation describes three studies exploring representations of speech used in different brain regions to produce speech. The first study investigated the motor units used to learn novel speech sequences. Subjects learned to produce a set of sequences with illegal consonant clusters (e.g. GVAZF) faster and more accurately than a similar novel set. Subjects then produced novel sequences that retained varying phonemic subsequences of previously learned sequences. Novel sequences were performed as quickly and accurately as learned sequences if they contained no novel consonant clusters, regardless of other phonemic content, implicating consonant clusters as important speech motor representations. The second study investigated the neural correlates of speech motor sequence learning. Functional magnetic resonance imaging (fMRI) revealed increased activity during novel sequence productions in brain regions traditionally associated with non-speech motor sequence learning - including the basal ganglia and premotor cortex - as well as regions associated with learning and updating speech motor representations based on sensory input - including the bilateral frontal operculum and left posterior superior temporal sulcus (pSTs). Behavioral learning measures correlated with increased response for novel sequences in the frontal operculum and with white matter integrity under the pSTs, implicating functional and structural connectivity of these regions in learning success

The structural correlates of statistical information processing during speech perception

The processing of continuous and complex auditory signals such as speech relies on the ability to use statistical cues (e.g. transitional probabilities). In this study, participants heard short auditory sequences composed either of Italian syllables or bird songs and completed a regularity-rating task. Behaviorally, participants were better at differentiating between levels of regularity in the syllable sequences than in the bird song sequences. Inter-individual differences in sensitivity to regularity for speech stimuli were correlated with variations in surface-based cortical thickness (CT). These correlations were found in several cortical areas including regions previously associated with statistical structure processing (e.g. bilateral superior temporal sulcus, left precentral sulcus and inferior frontal gyrus), as well other regions (e.g. left insula, bilateral superior frontal gyrus/sulcus and supramarginal gyrus). In all regions, this correlation was positive suggesting that thicker cortex is related to higher sensitivity to variations in the statistical structure of auditory sequences. Overall, these results suggest that inter-individual differences in CT within a distributed network of cortical regions involved in statistical structure processing, attention and memory is predictive of the ability to detect structural structure in auditory speech sequences

Repetition enhancement to voice identities in the dog brain

In the human speech signal, cues of speech sounds and voice identities are conflated, but they are processed separately in the human brain. The processing of speech sounds and voice identities is typically performed by non-primary auditory regions in humans and non-human primates. Additionally, these processes exhibit functional asymmetry in humans, indicating the involvement of distinct mechanisms. Behavioural studies indicate analogue side biases in dogs, but neural evidence for this functional dissociation is missing. In two experiments, using an fMRI adaptation paradigm, we presented awake dogs with natural human speech that either varied in segmental (change in speech sound) or suprasegmental (change in voice identity) content. In auditory regions, we found a repetition enhancement effect for voice identity processing in a secondary auditory region – the caudal ectosylvian gyrus. The same region did not show repetition effects for speech sounds, nor did the primary auditory cortex exhibit sensitivity to changes either in the segmental or in the suprasegmental content. Furthermore, we did not find evidence for functional asymmetry neither in the processing of speech sounds or voice identities. Our results in dogs corroborate former human and non-human primate evidence on the role of secondary auditory regions in the processing of suprasegmental cues, suggesting similar neural sensitivity to the identity of the vocalizer across the mammalian order