Isabelle Deschamps

On the role of the supramarginal gyrus in phonological processing and verbal working memory: Evidence from rTMS studies

The supramarginal gyrus (SMG) is activated for phonological processing during both language and verbal working memory tasks. Using rTMS, we investigated whether the contribution of the SMG to phonological processing is domain specific (specific to phonology) or more domain general (specific to verbal working memory). A measure of phonological complexity was developed based on sonority differences and subjects were tested after low frequency rTMS on a same/different judgment task and an n-back verbal memory task. It was reasoned that if the phonological processing in the SMG is more domain general, i.e., related to verbal working memory demands, performance would be more affected by the rTMS during the n-back task than during the same/different judgment task. Two auditory experiments were conducted. The first experiment demonstrated that under conditions where working memory demands are minimized (i.e. same/different judgment), repetitive stimulation had no effect on performance although performance varied as a function of phonological complexity. The second experiment demonstrated that during a verbal working memory task (n-back task), where phonological complexity was also manipulated, subjects were less accurate and slower at performing the task after stimulation but the effect of phonology was not affected. The results confirm that the SMG is involved in verbal working memory but not in the encoding of sonority differences.

Regional heterogeneity in the processing and the production of speech in the human planum temporale

INTRODUCTION The role of the left planum temporale (PT) in auditory language processing has been a central theme in cognitive neuroscience since the first descriptions of its leftward neuroanatomical asymmetry. While it is clear that PT contributes to auditory language processing there is still some uncertainty about its role in spoken language production. METHODS Here we examine activation patterns of the PT for speech production, speech perception and single word reading to address potential hemispheric and regional functional specialization in the human PT. To this aim, we manually segmented the left and right PT in three non-overlapping regions (medial, lateral and caudal PT) and examined, in two complementary experiments, the contribution of exogenous and endogenous auditory input on PT activation under different speech processing and production conditions. RESULTS Our results demonstrate that different speech tasks are associated with different regional functional activation patterns of the medial, lateral and caudal PT. These patterns are similar across hemispheres, suggesting bilateral processing of the auditory signal for speech at the level of PT. CONCLUSIONS Results of the present studies stress the importance of considering the anatomical complexity of the PT in interpreting fMRI data.

Sequencing at the syllabic and supra-syllabic levels during speech perception: an fMRI study

The processing of fluent speech involves complex computational steps that begin with the segmentation of the continuous flow of speech sounds into syllables and words. One question that naturally arises pertains to the type of syllabic information that speech processes act upon. Here, we used functional magnetic resonance imaging to profile regions, using a combination of whole-brain and exploratory anatomical region-of-interest (ROI) approaches, that were sensitive to syllabic information during speech perception by parametrically manipulating syllabic complexity along two dimensions: (1) individual syllable complexity, and (2) sequence complexity (supra-syllabic). We manipulated the complexity of the syllable by using the simplest syllable template—a consonant and vowel (CV)-and inserting an additional consonant to create a complex onset (CCV). The supra-syllabic complexity was manipulated by creating sequences composed of the same syllable repeated six times (e.g., /pa-pa-pa-pa-pa-pa/) and sequences of three different syllables each repeated twice (e.g., /pa-ta-ka-pa-ta-ka/). This parametrical design allowed us to identify brain regions sensitive to (1) syllabic complexity independent of supra-syllabic complexity, (2) supra-syllabic complexity independent of syllabic complexity and, (3) both syllabic and supra-syllabic complexity. High-resolution scans were acquired for 15 healthy adults. An exploratory anatomical ROI analysis of the supratemporal plane (STP) identified bilateral regions within the anterior two-third of the planum temporale, the primary auditory cortices as well as the anterior two-third of the superior temporal gyrus that showed different patterns of sensitivity to syllabic and supra-syllabic information. These findings demonstrate that during passive listening of syllable sequences, sublexical information is processed automatically, and sensitivity to syllabic and supra-syllabic information is localized almost exclusively within the STP.

The processing of polar quantifiers, and numerosity perception

We investigated the course of language processing in the context of a verification task that required numerical estimation and comparison. Participants listened to sentences with complex quantifiers that contrasted in Polarity, a logical property (e.g., more-than-half, less-than-half), and then performed speeded verification on visual scenarios that displayed a proportion between 2 discrete quantities. We varied systematically not only the sentences, but also the visual materials, in order to study their effect on the verification process. Next, we used the same visual scenarios with analogous non-verbal probes that featured arithmetical inequality symbols (<, >). This manipulation enabled us to measure not only Polarity effects, but also, to compare the effect of different probe types (linguistic, non-linguistic) on processing. Like many previous studies, our results demonstrate that perceptual difficulty affects error rate and reaction time in keeping with Webers Law. Interestingly, these performance parameters are also affected by the Polarity of the quantifiers used, despite the fact that sentences had the exact same meaning, sentence structure, number of words, syllables, and temporal structure. Moreover, an analogous contrast between the non-linguistic probes (<, >) had no effect on performance. Finally, we observed no interaction between performance parameters governed by Webers Law and those affected by Polarity. We consider 4 possible accounts of the results (syntactic, semantic, pragmatic, frequency-based), and discuss their relative merit.

Structural brain aging and speech production: a surface-based brain morphometry study

While there has been a growing number of studies examining the neurofunctional correlates of speech production over the past decade, the neurostructural correlates of this immensely important human behaviour remain less well understood, despite the fact that previous studies have established links between brain structure and behaviour, including speech and language. In the present study, we thus examined, for the first time, the relationship between surface-based cortical thickness (CT) and three different behavioural indexes of sublexical speech production: response duration, reaction times and articulatory accuracy, in healthy young and older adults during the production of simple and complex meaningless sequences of syllables (e.g., /pa–pa-pa/ vs. /pa-ta-ka/). The results show that each behavioural speech measure was sensitive to the complexity of the sequences, as indicated by slower reaction times, longer response durations and decreased articulatory accuracy in both groups for the complex sequences. Older adults produced longer speech responses, particularly during the production of complex sequence. Unique age-independent and age-dependent relationships between brain structure and each of these behavioural measures were found in several cortical and subcortical regions known for their involvement in speech production, including the bilateral anterior insula, the left primary motor area, the rostral supramarginal gyrus, the right inferior frontal sulcus, the bilateral putamen and caudate, and in some region less typically associated with speech production, such as the posterior cingulate cortex.

Neural sensitivity to syllable frequency and mutual information in speech perception and production

Many factors affect our ability to decode the speech signal, including its quality, the complexity of the elements that compose it, as well as their frequency of occurrence and co-occurrence in a language. Syllable frequency effects have been described in the behavioral literature, including facilitatory effects during speech production and inhibitory effects during word recognition, but the neural mechanisms underlying these effects remain largely unknown. The objective of this study was to examine, using functional neuroimaging, the neurobiological correlates of three different distributional statistics in simple 2-syllable nonwords: the frequency of the first and second syllables, and the mutual information between the syllables. We examined these statistics during nonword perception and production using a powerful single-trial analytical approach. We found that repetition accuracy was higher for nonwords in which the frequency of the first syllable was high. In addition, brain responses to distributional statistics were widespread and almost exclusively cortical. Importantly, brain activity was modulated in a distinct manner for each statistic, with the strongest facilitatory effects associated with the frequency of the first syllable and mutual information. These findings show that distributional statistics modulate nonword perception and production. We discuss the common and unique impact of each distributional statistic on brain activity, as well as task differences.

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.

Phonological processing in speech perception: What do sonority differences tell us?

Previous research has associated the inferior frontal and posterior temporal brain regions with a number of phonological processes. In order to identify how these specific brain regions contribute to phonological processing, we manipulated subsyllabic phonological complexity and stimulus modality during speech perception using fMRI. Subjects passively attended to visual or auditory pseudowords. Similar to previous studies, a bilateral network of cortical regions was recruited during the presentation of visual and auditory stimuli. Moreover, pseudowords recruited a similar network of regions as words and letters. Few regions in the whole-brain results revealed neural processing differences associated with phonological complexity independent of modality of presentation. In an ROI analysis, the only region sensitive to phonological complexity was the posterior part of the inferior frontal gyrus (IFGpo), with the complexity effect only present for print. In sum, the sensitivity of phonological brain areas depends on the modality of stimulus presentation and task demands.

Age differences in the motor control of speech: An fMRI study of healthy aging: Age Differences in Speech Production

Healthy aging is associated with a decline in cognitive, executive, and motor processes that are concomitant with changes in brain activation patterns, particularly at high complexity levels. While speech production relies on all these processes, and is known to decline with age, the mechanisms that underlie these changes remain poorly understood, despite the importance of communication on everyday life. In this cross‐sectional group study, we investigated age differences in the neuromotor control of speech production by combining behavioral and functional magnetic resonance imaging (fMRI) data. Twenty‐seven healthy adults underwent fMRI while performing a speech production task consisting in the articulation of nonwords of different sequential and motor complexity. Results demonstrate strong age differences in movement time (MT), with longer and more variable MT in older adults. The fMRI results revealed extensive age differences in the relationship between BOLD signal and MT, within and outside the sensorimotor system. Moreover, age differences were also found in relation to sequential complexity within the motor and attentional systems, reflecting both compensatory and de‐differentiation mechanisms. At very high complexity level (high motor complexity and high sequence complexity), age differences were found in both MT data and BOLD response, which increased in several sensorimotor and executive control areas. Together, these results suggest that aging of motor and executive control mechanisms may contribute to age differences in speech production. These findings highlight the importance of studying functionally relevant behavior such as speech to understand the mechanisms of human brain aging. Hum Brain Mapp 38:2751–2771, 2017.

Neurobiology of Speech Production: A Motor Control Perspective

Speaking is a dynamic process in which neural representations associated with language and the constantly changing vocal tract configurations used to produce speech are seamlessly integrated to communicate. This integrative process requires a stable repertoire of motor routines and associated flexible sensorimotor and cognitive processes for speech preparation, execution, and monitoring. The development of neurobiologically plausible models of speech production requires an understanding of the role and contribution of both central and peripheral factors in the acquisition, maintenance, and reorganization of the brain-behavior patterns underlying speech production. Recent advances in brain imaging and neuromodulation methods, signal processing, and data analysis techniques have provided new insights into the organization of the brain networks and brain-behavior relations, leading to more comprehensive and more realistic models of speech production. In this chapter we identify the set of processes that are involved in producing speech and their neural substrate.