Dissociating the functions of superior and inferior parts of the left ventral occipito-temporal cortex during visual word and object processing

During word and object recognition, extensive activation has consistently been observed in the left ventral occipito-temporal cortex (vOT), focused around the occipito-temporal sulcus (OTs). Previous studies have shown that there is a hierarchy of responses from posterior to anterior vOT regions (along the y-axis) that corresponds with increasing levels of recognition - from perceptual to semantic processing, respectively. In contrast, the functional differences between superior and inferior vOT responses (i.e. along the z-axis) have not yet been elucidated. To investigate, we conducted an extensive review of the literature and found that peak activation for reading varies by more than 1 cm in the z-axis. In addition, we investigated functional differences between superior and inferior parts of left vOT by analysing functional MRI data from 58 neurologically normal skilled readers performing 8 different visual processing tasks. We found that group activation in superior vOT was significantly more sensitive than inferior vOT to the type of task, with more superior vOT activation when participants were matching visual stimuli for their semantic or perceptual content than producing speech to the same stimuli. This functional difference along the z-axis was compared to existing boundaries between cytoarchitectonic areas around the OTs. In addition, using dynamic causal modelling, we show that connectivity from superior vOT to anterior vOT increased with semantic content during matching tasks but not during speaking tasks whereas connectivity from inferior vOT to anterior vOT was sensitive to semantic content for matching and speaking tasks. The finding of a functional dissociation between superior and inferior parts of vOT has implications for predicting deficits and response to rehabilitation for patients with partial damage to vOT following stroke or neurosurgery

Improving our understanding of speech and language outcome in neurosurgery patients

Malignant gliomas remain incurable and result in more years of life lost than any other tumours. Surgical resection is strongly recommended but carries a risk of causing functional impairment. This thesis aims to demonstrate how state-of-the-art functional magnetic resonance imaging (fMRI) language paradigms can contribute to neurosurgical planning. The first three experiments use a multitask fMRI language paradigm to functionally segregate left posterior temporal and left posterior frontal regions involved in the perception and production of speech. Experiment 1 demonstrated three functionally distinct responses in the left posterior superior temporal sulcus (STS), left temporo-parietal junction and anterior ascending terminal branch of the left STS. Experiment 2 validates these findings in an independent group of participants, increasing confidence that they are robust. Experiment 3 dissociates the response of three different parts of the left premotor cortex during speech production. Experiment 4 shows that left posterior temporal regions are more consistently activated, in neurotypical controls, when a picture naming task presents pairs of objects rather than single objects. Further work could therefore test whether paired object naming is a more sensitive task for pre- and intra-operative language mapping. Finally, Experiment 5 found that successful reading before and after surgery, in two patients with gliomas affecting the left temporo-parietal junction, enhanced activation in bilateral perirhinal regions that were associated with semantic identification of visually presented objects in neurotypical controls. Future studies can now test whether patients who undergo resection of the left temporo-parietal junction have better reading, post-surgery, when bilateral perirhinal activation is enhanced prior to surgery. Taken together, this work expands our knowledge of the functional anatomy of language, proposes a new way of utilising fMRI data from neurotypical controls to tailor pre- and intra-operative language mapping strategies and provides an insight into how the reading system reorganises itself after brain damage

Dissociating frontal regions that co-lateralize with different ventral occipitotemporal regions during word processing

The ventral occipitotemporal sulcus (vOT) sustains strong interactions with the inferior frontal cortex during word processing. Consequently, activation in both regions co-lateralize towards the same hemisphere in healthy subjects. Because the determinants of lateralisation differ across posterior, middle and anterior vOT subregions, we investigated whether lateralisation in different inferior frontal regions would co-vary with lateralisation in the three different vOT subregions. A whole brain analysis found that, during semantic decisions on written words, laterality covaried in (1) posterior vOT and the precentral gyrus; (2) middle vOT and the pars opercularis, pars triangularis, and supramarginal gyrus; and (3) anterior vOT and the pars orbitalis, middle frontal gyrus and thalamus. These findings increase the spatial resolution of our understanding of how vOT interacts with other brain areas during semantic categorisation on words

Dissociating object familiarity from linguistic properties in mirror word reading

<p>Abstract</p> <p>Background</p> <p>It is known that the orthographic properties of linguistic stimuli are processed within the left occipitotemporal cortex at about 150–200 ms. We recorded event-related potentials (ERPs) to words in standard or mirror orientation to investigate the role of visual word form in reading. Word inversion was performed to determine whether rotated words lose their linguistic properties.</p> <p>Methods</p> <p>About 1300 Italian words and legal pseudo-words were presented to 18 right-handed Italian students engaged in a letter detection task. EEG was recorded from 128 scalp sites.</p> <p>Results</p> <p>ERPs showed an early effect of word orientation at ~150 ms, with larger N1 amplitudes to rotated than to standard words. Low-resolution brain electromagnetic tomography (LORETA) revealed an increase in N1 to rotated words primarily in the right occipital lobe (BA 18), which may indicate an effect of stimulus familiarity. N1 was greater to target than to non-target letters at left lateral occipital sites, thus reflecting the first stage of orthographic processing. LORETA revealed a strong focus of activation for this effect in the left fusiform gyrus (BA 37), which is consistent with the so-called visual word form area (VWFA). Standard words (compared to pseudowords) elicited an enhancement of left occipito/temporal negativity at about 250–350 ms, followed by a larger anterior P3, a reduced frontal N400 and a huge late positivity. Lexical effects for rotated strings were delayed by about 100 ms at occipito/temporal sites, and were totally absent at later processing stages. This suggests the presence of implicit reading processes, which were pre-attentive and of perceptual nature for mirror strings.</p> <p>Conclusion</p> <p>The contrast between inverted and standard words did not lead to the identification of a purely linguistic brain region. This finding suggests some caveats in the interpretation of the inversion effect in subtractive paradigms.</p

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

Activity in Face-Responsive Brain Regions is Modulated by Invisible, Attended Faces: Evidence from Masked Priming

It is often assumed that neural activity in face-responsive regions of primate cortex correlates with conscious perception of faces. However, whether such activity occurs without awareness is still debated. Using functional magnetic resonance imaging (fMRI) in conjunction with a novel masked face priming paradigm, we observed neural modulations that could not be attributed to perceptual awareness. More specifically, we found reduced activity in several classic face-processing regions, including the “fusiform face area,” “occipital face area,” and superior temporal sulcus, when a face was preceded by a briefly flashed image of the same face, relative to a different face, even when 2 images of the same face differed. Importantly, unlike most previous studies, which have minimized awareness by using conditions of inattention, the present results occurred when the stimuli (the primes) were attended. By contrast, when primes were perceived consciously, in a long-lag priming paradigm, we found repetition-related activity increases in additional frontal and parietal regions. These data not only demonstrate that fMRI activity in face-responsive regions can be modulated independently of perceptual awareness, but also document where such subliminal face-processing occurs (i.e., restricted to face-responsive regions of occipital and temporal cortex) and to what extent (i.e., independent of the specific image)

The neural basis of object perception: dissociating action and semantic processing

This thesis has evaluated the roles of dorsal and ventral processing streams in recognition and use of objects. Four main empirical studies are presented. First, to investigate how the cortical brain processes semantic and action knowledge in different object-related tasks, I examined structural data from stroke patients (Chapter 2) and functional data from healthy individuals (Chapter 3) using a voxel-wise statistical analysis method. Using data of different modalities (structural CT, fMRI) from different sources (patients’ lesions; healthy subjects’ functional activity) handled with a systematic analysis approach, I attempted to find convergent evidence to support the dissociation of semantic and action processing. Second, I also looked into the potential differentiation within the mechanisms underlying object-related action (Chapter 4) and object naming (Chapter 5) separately. Overall, comparable findings were provided from the voxel-based morphometric analysis of patients’ lesion data and the fMRI study with healthy participants: an association was observed between ventral brain structures and the retrieval of semantic knowledge/object recognition while a dorsal fronto-parietal-occipital network was found to support the processing of action knowledge/object-oriented action. Specific dissociations were also observed within the representations for object-oriented actions as well as the mechanisms underlying naming of objects

Ventral occipito-temporal cortex function and anatomical connectivity in reading

Previous functional neuroimaging studies of reading in skilled readers, acquired dyslexia and developmental dyslexia have all shown that the left ventral occipito-temporal cortex (vOT) is involved in visual word recognition. Specifically, a region in the left posterior occipito-temporal sulcus lateral to fusiform gyrus and medial to inferior temporal gyrus has been reported to play an important role. However, the precise functional contribution of this area in reading is yet to be fully explored. In this thesis, I empirically evaluated a claim that vOT responds not only to bottom-up processing demands of the visual stimuli but is also influenced by automatic, top-down non-visual processing demands, as proposed by the Interactive Account of vOT functioning. The first part of this thesis investigated the functional properties of vOT during reading, using functional magnetic resonance imaging. In the first project, the top-down influences on vOT were investigated, teasing apart visual and non-visual properties of written stimuli. In the second project, using the Japanese orthography I disentangled a word’s lexical frequency from the frequency of its visual form – an important distinction for understanding the neural information processing in regions engaged by reading and further explored the interactive nature of the vOT responses. The second part then investigated the anatomical basis of these functional interactions between vOT and other cortical regions. I used diffusion-weighted magnetic resonance imaging and tractography, the only method currently available to identify and measure white matter fibre pathways non-invasively and in vivo. My research has demonstrated that vOT integrates bottom-up visual information and top-down predictions from regions encoding non-visual attributes of the stimulus in an interactive fashion. It also illustrated the putative anatomical basis for functional connectivity during reading, which is consistent with the parallel cortical visual pathways seen in other primates. Altogether, the results provide strong support for the Interactive Account

The role of the left inferior parietal lobule in reading.

One of the regions that have consistently been included in the neurological models of reading is the left inferior parietal lobule (IPL), however, the precise functional and temporal contributions of this region to reading have not yet been fully established. There are three hypotheses concerning IPL contributions to visual word recognition. The first one claims that the IPL is the site of stored visual word forms although it remains unclear whether these are stored in supramarginal (SMG) or angular (ANG) fields of the IPL. The second hypothesis argues that the procedures for converting spelling-to-sound are a function of the IPL, but it is unclear whether these are specifically located in SMG or ANG, or both. Finally, a third hypothesis suggests that SMG and ANG preferentially contribute to phonological and semantic processing of written words, respectively. In this thesis, I empirically evaluated these hypotheses using repetitive transcranial magnetic stimulation (rTMS) to temporarily and selectively disrupt processing in left SMG and ANG during visual word recognition and measure the effect on reading behaviour. I also investigated the time course of SMG and ANG involvement to visual word recognition using double-pulse TMS. My research demonstrates that SMG contributes preferentially to phonological aspects of word processing and the processing begins early and over a sustained period of time (between 80 to 200 msec post-stimulus onset). ANG contributes preferentially to semantic aspects of word processing but the temporal dynamics of this contribution were not successfully revealed in this thesis and require further investigation. In addition, I empirically evaluated the efficiency of using functional magnetic resonance (fMRI) and TMS to functionally localize a target site for TMS experiments. I demonstrated that both methods are similarly accurate in identifying stimulation site but neither of them is 100% accurate