Neurocognitive signatures of phonemic sequencing in expert backward speakers

Despite its prolific growth, neurolinguistic research on phonemic sequencing has largely neglected the study of individuals with highly developed skills in this domain. To bridge this gap, we report multidimensional signatures of two experts in backward speech, that is, the capacity to produce utterances by reversing the order of phonemes while retaining their identity. Our approach included behavioral assessments of backward and forward speech alongside neuroimaging measures of voxel-based morphometry, diffusion tensor imaging, and resting-state functional connectivity. Relative to controls, both backward speakers exhibited behavioral advantages for reversing words and sentences of varying complexity, irrespective of working memory skills. These patterns were accompanied by increased grey matter volume, higher mean diffusivity, and enhanced functional connectivity along dorsal and ventral stream regions mediating phonological and other linguistic operations, with complementary support of areas subserving associative-visual and domain-general processes. Still, the specific loci of these neural patterns differed between both subjects, suggesting individual variability in the correlates of expert backward speech. Taken together, our results offer new vistas on the domain of phonemic sequencing, while illuminating neuroplastic patterns underlying extraordinary language abilities

Functional MRI investigations of cortical mechanisms of auditory spatial attention

In everyday settings, spatial attention helps listeners isolate and understand individual sound sources. However, the neural mechanisms of auditory spatial attention (ASpA) are only partially understood. This thesis uses within-subject analysis of functional magnetic resonance imaging (fMRI) data to address fundamental questions regarding cortical mechanisms supporting ASpA by applying novel multi-voxel pattern analysis (MVPA) and resting-state functional connectivity (rsFC) approaches. A series of fMRI studies of ASpA were conducted in which subjects performed a one-back task in which they attended to one of two spatially separated streams. Attention modulated blood oxygenation level-dependent (BOLD) activity in multiple areas in the prefrontal, temporal, and parietal cortex, including non-visuotopic intraparietal sulcus (IPS), but not the visuotopic maps in IPS. No spatial bias was detected in any cortical area using standard univariate analysis; however, MVPA revealed that activation patterns in a number of areas, including the auditory cortex, predicted the attended direction. Furthermore, we explored how cognitive task demands and the sensory modality of the inputs influenced activity with a visual one-back task and a visual multiple object tracking (MOT) task. Activity from the visual and auditory one-back tasks overlapped along the fundus of IPS and lateral prefrontal cortex (lPFC). However, there was minimal overlap of activity in the lPFC between the visual MOT task and the two one-back tasks. Finally, we endeavored to identify visual and auditory networks using rsFC. We identified a dorsal visual attention network reliably within individual subjects using visuotopic seeds. Using auditory seeds, we found a prefrontal area nested between segments of the dorsal visual attention network. These findings mark fundamental progress towards elucidating the cortical network controlling ASpA. Our results suggest that similar lPFC structures support both ASpA and its visual counterpart during a spatial one-back task, but that ASpA does not drive visuotopic IPS in the parietal cortex. Furthermore, rsFC reveals that visual and auditory seed regions are functionally connected with non-overlapping lPFC regions, possibly reflecting spatial and temporal cognitive processing biases, respectively. While we find no evidence for a spatiotopic map, the auditory cortex is sensitive to direction of attention in its patterns of activation

FMRI and ERP investigations of body representations in the human lateral and ventral occipitotemporal cortex

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From dysfunctional to extraordinary verbal repetition abilities: clinical implications and neural features

Three cases are presented to index the hypothesis that mitigated echolalia emerges from overreliance on the dorsal language stream, through the arcuate fasciculus, when the ventral stream is damaged; whereas conduite d’approche ensues when the ventral stream attempts to compensate a dorsal damage. The role of the right hemisphere and other alternative pathways in both cerebral hemispheres in the successful compensation of brain injury is also discussed. Further, Study 2 reconceptualizes different types of echolalia within a continuous of severity and communication capacity. To accomplish this new instantiation, it is proposed that different types of echolalia may be associated to failure in distinctive linguistic and non-linguistic cognitive functions. Recommendations for its evaluation and treatment are provided, suggesting that echolalia interfering with functional communication should be treated. Further, complementing the previous one, Study 3 reports a comprehensive single case study exploring response to treatment, and behavioral and neuroimaging features of a person with mitigated echolalia associated to a chronic fluent aphasia. Findings from such case include a reduction of mitigated echolalia after two weeks of intensive aphasia therapy as well as the maintenance of these gains with memantine alone for at least 6 months. Importantly, reduction of mitigated echolalia instances in response to treatment speeded up the time needed to complete comprehension tasks. Neuroimaging results, although indirectly, suggested that mitigated echolalia may be supported by the activity of the remaining components of the left dorsal stream and compensatory right hemisphere recruitment. Additionally, to further explore the neural and cognitive mechanisms involved in verbal repetition in a model of language expertise, Study 4 tackles cognitive features and neural correlates of verbal expertise in two healthy adult subjects displaying an extraordinary ability to orally reverse language, a condition referred to as backward speech. Results suggest that phonological expertise, as shown in backward speech, involves reshaping (or pre-existent differences) of cortical areas and tracts relevant for auditory-motor integration and semantic processing. Greater functional coupling between critical language areas and domain-general and high-order visual areas may further support reversing processes. Lastly, Study 5 presents a systematic review of the literature aimed to examine sex differences in the prevalence of repetition deficits in persons with post-stroke aphasia. Results show that the proportion of females in the group of aphasia characterized by repetition deficits (i.e., conduction aphasia) is lower than the expected by the prevalence of stroke among them. It is suggested that sex-related differences in the volume of areas of the right hemisphere homologues to the ones subserving repetition in the left hemisphere may be at the base of this difference. This finding poses sex as a relevant variable to account for variance in repetition abilities, and as a relevant factor to consider in future studies of language acquisition, maturation, and relearning promoted by aphasia therapy. Fecha de lectura de Tesis Doctoral: 16 de diciembre 2019Verbal repetition and audio-visual imitation stand as crucial functions for the acquisition and maturation of language in childhood, language learning in adulthood, and a major resource for language recovery after brain damage. Although modern neuroimaging techniques have allowed the identification of the brain areas involved in repetition tasks in healthy subjects, many clinical and neural aspects of this linguistic function are still overlooked in persons with aphasia and in emerging models of language expertise. Therefore, the present dissertation aims to explore cognitive correlates and neural features of verbal repetition from different perspectives including models of dysfunctional repetition (i.e., people with aphasia) and language expertise (i.e., healthy backward speakers). Generally, this thesis explores the potential of the dorsal and ventral components of the neural network supporting verbal repetition to assume, under certain circumstances (e.g., brain damage or extraordinary abilities), non-canonical functions. Further, this dissertation addresses clinical issues of some aphasic symptoms characterized by uncontrolled repetition (i.e., echolalia), as well as reviews sex as a source of variability in verbal repetition outcomes after brain damage. This dissertation includes five studies that are part of this dissertation. First, it reviews the mechanisms involved in dysfunctional repetition, especially in two repetitive verbal behaviors named conduite d’approche and mitigated echolalia (Study 1) and addresses clinical issues of the last one (Study 2 and 3). In this regard, Study 1 proposes that in the context of aphasia these symptoms (i.e., conduite d’approche and mitigated echolalia) may represent active attempts of verbal communication, rather than inconsequential repetitive verbal behaviors resulting from maladaptive neural changes

The functional anatomy of white matter pathways for visual configuration learning

The role of the medial temporal lobes (MTL) in visuo-spatial learning has been extensively studied and documented in the neuroscientific literature. Numerous animal and human studies have demonstrated that the parahippocampal place area (PPA), which sits at the confluence of the parahippocampal and lingual gyri, is particularly important for learning the spatial configuration of objects in visually presented scenes. In current visuo-spatial processing models, the PPA sits downstream from the parietal lobes which are involved in multiple facets of spatial processing. Yet, direct input to the PPA from early visual cortex (EVC) is rarely discussed and poorly understood. This thesis adopted a multimodal neuroimaging analysis approach to study the functional anatomy of these connections. First, the pattern of structural connectivity between EVC and the MTL was explored by means of surface-based ‘connectomes’ constructed from diffusion MRI tractography in a cohort of 200 healthy young adults from the Human Connectome Project. Through this analysis, the PPA emerged as a primary recipient of EVC connections within the MTL. Second, a data-driven clustering analysis of the PPA’s connectivity to an extended cortical region (including EVC, retrosplenial cortex, and other areas) revealed multiple clusters with different connectivity profiles within the PPA. The two main clusters were located in the posterior and anterior portions of the PPA, with the posterior cluster preferentially connected to EVC. Motivated by this result, virtual tractography dissections were used to delineate the medial occipital longitudinal tract (MOLT), the white matter bundle connecting the PPA with EVC. The properties of this bundle and its relation to visual configuration learning were verified in a different, cross-sectional adult cohort of 90 subjects. Finally, the role of the MOLT in the visuo-spatial learning domain was further confirmed in the case of a stroke patient who, after bilateral occipital injury, exhibited deficits confined to this domain. The results presented in this work suggest that the MOLT should be included in current visuo-spatial processing models as it offers additional insight into how the MTL acquires and processes information for spatial learning

Word contexts enhance the neural representation of individual letters in early visual cortex

Visual context facilitates perception, but how this is neurally implemented remains unclear. One example of contextual facilitation is found in reading, where letters are more easily identified when embedded in a word. Bottom-up models explain this word advantage as a post-perceptual decision bias, while top-down models propose that word contexts enhance perception itself. Here, we arbitrate between these accounts by presenting words and nonwords and probing the representational fidelity of individual letters using functional magnetic resonance imaging. In line with top-down models, we find that word contexts enhance letter representations in early visual cortex. Moreover, we observe increased coupling between letter information in visual cortex and brain activity in key areas of the reading network, suggesting these areas may be the source of the enhancement. Our results provide evidence for top-down representational enhancement in word recognition, demonstrating that word contexts can modulate perceptual processing already at the earliest visual regions

Multivariate pattern analysis of input and output representations of speech

Repeating a word or nonword requires a speaker to map auditory representations of incoming sounds onto learned speech items, maintain those items in short-term memory, interface that representation with the motor output system, and articulate the target sounds. This dissertation seeks to clarify the nature and neuroanatomical localization of speech sound representations in perception and production through multivariate analysis of neuroimaging data. The major portion of this dissertation describes two experiments using functional magnetic resonance imaging (fMRI) to measure responses to the perception and overt production of syllables and multivariate pattern analysis to localize brain areas containing associated phonological/phonetic information. The first experiment used a delayed repetition task to permit response estimation for auditory syllable presentation (input) and overt production (output) in individual trials. In input responses, clusters sensitive to vowel identity were found in left inferior frontal sulcus (IFs), while clusters responsive to syllable identity were found in left ventral premotor cortex and left mid superior temporal sulcus (STs). Output-linked responses revealed clusters of vowel information bilaterally in mid/posterior STs. The second experiment was designed to dissociate the phonological content of the auditory stimulus and vocal target. Subjects were visually presented with two (non)word syllables simultaneously, then aurally presented with one of the syllables. A visual cue informed subjects either to repeat the heard syllable (repeat trials) or produce the unheard, visually presented syllable (change trials). Results suggest both IFs and STs represent heard syllables; on change trials, representations in frontal areas, but not STs, are updated to reflect the vocal target. Vowel identity covaries with formant frequencies, inviting the question of whether lower-level, auditory representations can support vowel classification in fMRI. The final portion of this work describes a simulation study, in which artificial fMRI datasets were constructed to mimic the overall design of Experiment 1 with voxels assumed to contain either discrete (categorical) or analog (frequency-based) vowel representations. The accuracy of classification models was characterized by type of representation and the density and strength of responsive voxels. It was shown that classification is more sensitive to sparse, discrete representations than dense analog representations

Neural Correlates of Repetition Priming: Changes in fMRI Activation and Synchrony

The neural mechanisms of behavioural priming remain unclear. Recent studies have suggested that category-preferential regions in ventral occipitotemporal cortex (VOTC) play an important role; some have reported increased neural synchrony between prefrontal cortex and temporal cortex associated with stimulus repetition. Based on these results, I hypothesized that increased neural synchrony, as measured by functional connectivity analysis using functional MRI, between category-preferential regions in VOTC and broader category-related networks would underlie behavioural priming. To test this hypothesis, I localized several category-preferential regions in VOTC using an independent functional localizer. Then, Seed Partial Least Squares was used to assess task-related functional connectivity of these regions across repetition of stimuli from multiple categories during an independent semantic classification task. While the results did not show the hypothesized differences in functional connectivity across stimulus repetition, evidence of category-specificity of neural priming and novel insights about the nature of category-related organization of VOTC were revealed

Effects of Temporal and Spatial Context Within the Macaque Face-Processing System

Temporal and spatial context play a key role in vision as a whole, and in face perception specifically. However, little is known about the neurophysiological mechanisms by which contextual cues exert their effects. Anatomically distinct face patches in the macaque brain analyze facial form, and studies of the activity within these patches have begun to clarify the neural machinery that underlies facial perception. This system provides a uniquely valuable opportunity to study how context affects the perception of form. We used functional magnetic resonance imaging (fMRI) to investigate the brain activity of macaque monkeys while they viewed faces placed in either temporal or spatial context. Facial motion transmits rich and ethologically vital information, but the way that the brain interprets such natural temporal context is poorly understood. Facial motion activates the face patches and surrounding areas, yet it is not known whether this motion is processed by its own specialized neural machinery, and if so, what that machinery’s organization might be. To address these questions, we monitored the brain activity of macaque monkeys while they viewed low- and high-level motion and form stimuli. We found that, beyond classical motion areas and the known face patch system, moving faces recruited a heretofore-unrecognized face patch. Although all face patches displayed distinctive selectivity for face motion over object motion, only two face patches preferred naturally moving faces, while three others preferred randomized, rapidly varying sequences of facial form. This functional divide was anatomically specific, segregating dorsal from ventral face patches, thereby revealing a new organizational principle of the macaque face-processing system. Like facial motion, bodies can provide valuable social context, revealing emotion and identity. Little is known about the joint processing of faces and bodies, even though there is reason to believe that their neural representations are intertwined. To identify interaction between the neural representations of face and body, we monitored the brain activity of the same monkeys while they viewed pictures of whole monkeys, isolated monkey heads, and isolated monkey bodies. We found that certain areas, including anterior face patches, responded more to whole monkeys than would be predicted by summing the separate responses to isolated heads and isolated bodies. The supralinear response was specific to viewing the conjunction of head and body; heads placed atop nonbody objects did not evoke this activity signature. However, a supralinear context response was elicited by pixelated, ambiguous faces presented on bodies. The size of this response suggests that the supralinear signal in this case did not result from the disambiguation of the ambiguous faces. These studies of contextually evoked activity within the macaque face processing system deepen our understanding of the cortical organization of both visual context and face processing, and identify promising sites for future research into the mechanisms underlying these critical aspects of perception

Multimodal MRI characterization of visual word recognition: an integrative view

228 p.The ventral occipito-temporal (vOT) association cortex contributes significantly to recognize different types of visual patterns. It is widely accepted that a subset of this circuitry, including the visual word form area (VWFA), becomes trained to perform the task of rapidly identifying word forms. An important open question is the computational role of this circuitry: To what extent is part of a bottom-up hierarchical processing of information on visual word recognition and/or is involved in processing top-down signals from higher-level language regions. This doctoral dissertation thesis proposal is aimed at characterizing the vOT reading circuitry using behavioral, functional, structural and quantitative MRI indexes, and linking its computations to the other two important regions within the language network: the posterior parietal cortex (pPC) and the inferior frontal gyrus (IFG). Results revealed that two distinct word-responsive areas can be segregated in the vOT: one responsible for visual feature extraction that is connected to the intraparietal sulcus via the vertical occipital fasciculus and a second one responsible for semantic processing that is connected to the angular gyrus via the posterior arcuate fasciculus and to the IFG via the anterior arcuate fasciculus. Importantly, reading behavior was predicted by functional activation in regions identified along the vOT, pPC and IFG, as well as by structural properties of the white matter fiber tracts linking them. The present work constitutes a critical step in the creation of a highly detailed characterization of the early stages of reading at the individual-subject level and to establish a baseline model and parameter range that might serve to clarify functional and structural differences between typical, poor and atypical readers.BCBL: basque center on cognition, brain and languag