Computing geodesic paths encoding a curvature prior for curvilinear structure tracking.

In this paper, we introduce an efficient method for computing curves minimizing a variant of the Euler-Mumford elastica energy, with fixed endpoints and tangents at these endpoints, where the bending energy is enhanced with a user-defined and data-driven scalar-valued term referred to as the curvature prior. In order to guarantee that the globally optimal curve is extracted, the proposed method involves the numerical computation of the viscosity solution to a specific static Hamilton-Jacobi-Bellman (HJB) partial differential equation (PDE). For that purpose, we derive the explicit Hamiltonian associated with this variant model equipped with a curvature prior, discretize the resulting HJB PDE using an adaptive finite difference scheme, and solve it in a single pass using a generalized fast-marching method. In addition, we also present a practical method for estimating the curvature prior values from image data, designed for the task of accurately tracking curvilinear structure centerlines. Numerical experiments on synthetic and real-image data illustrate the advantages of the considered variant of the elastica model with a prior curvature enhancement in complex scenarios where challenging geometric structures appear.

Subtitled speech: the neural mechanisms of ticker-tape synaesthesia.

The acquisition of reading modifies areas of the brain associated with vision and with language, in addition to their connections. These changes enable reciprocal translation between orthography and the sounds and meaning of words. Individual variability in the pre-existing cerebral substrate contributes to the range of eventual reading abilities, extending to atypical developmental patterns, including dyslexia and reading-related synaesthesias. The present study is devoted to the little-studied but highly informative ticker-tape synaesthesia, in which speech perception triggers the vivid and irrepressible perception of words in their written form in the mind's eye. We scanned a group of 17 synaesthetes and 17 matched controls with functional MRI, while they listened to spoken sentences, words, numbers or pseudowords (Experiment 1), viewed images and written words (Experiment 2) or were at rest (Experiment 3). First, we found direct correlates of the ticker-tape synaesthesia phenomenon: during speech perception, as ticker-tape synaesthesia was active, synaesthetes showed over-activation of left perisylvian regions supporting phonology and of the occipitotemporal visual word form area, where orthography is represented. Second, we provided support to the hypothesis that ticker-tape synaesthesia results from atypical relationships between spoken and written language processing: the ticker-tape synaesthesia-related regions overlap closely with cortices activated during reading, and the overlap of speech-related and reading-related areas is larger in synaesthetes than in controls. Furthermore, the regions over-activated in ticker-tape synaesthesia overlap with regions under-activated in dyslexia. Third, during the resting state (i.e. in the absence of current ticker-tape synaesthesia), synaesthetes showed increased functional connectivity between left prefrontal and bilateral occipital regions. This pattern might reflect a lowered threshold for conscious access to visual mental contents and might imply a non-specific predisposition to all synaesthesias with a visual content. These data provide a rich and coherent account of ticker-tape synaesthesia as a non-detrimental developmental condition created by the interaction of reading acquisition with an atypical cerebral substrate.

Color Naming and Categorization Depend on Distinct Functional Brain Networks.

Naming a color can be understood as an act of categorization, that is, identifying it as a member of a category of colors that are referred to by the same name. But are naming and categorization equivalent cognitive processes and consequently rely on same neural substrates? Here, we used task and resting-state functional magnetic resonance imaging as well as behavioral measures to identify functional brain networks that modulated naming and categorization of colors. We first identified three bilateral color-sensitive regions in the ventro-occipital cortex. We then showed that, across participants, color naming and categorization response times (RTs) were correlated with different resting state connectivity networks seeded from the color-sensitive regions. Color naming RTs correlated with the connectivity between the left posterior color region, the left middle temporal gyrus, and the left angular gyrus. In contrast, color categorization RTs correlated with the connectivity between the bilateral posterior color regions, and left frontal, right temporal and bilateral parietal areas. The networks supporting naming and categorization had a minimal overlap, indicating that the 2 processes rely on different neural mechanisms.

A mesial-to-lateral dissociation for orthographic processing in the visual cortex.

Efficient reading requires a fast conversion of the written word to both phonological and semantic codes. We tested the hypothesis that, within the left occipitotemporal cortical regions involved in visual word recognition, distinct subregions harbor slightly different orthographic codes adapted to those 2 functions. While the lexico-semantic pathway may operate on letter or open-bigram information, the phonological pathway requires the identification of multiletter graphemes such as "ch" or "ou" in order to map them onto phonemes. To evaluate the existence of a specific stage of graphemic encoding, 20 adults performed lexical decision and naming tasks on words and pseudowords during functional MRI. Graphemic encoding was facilitated or disrupted by coloring and spacing the letters either congruently with multiletter graphemes (ch-ai-r) or incongruently with them (c-ha-ir). This manipulation affected behavior, primarily during the naming of pseudowords, and modulated brain activity in the left midfusiform sulcus, at a site medial to the classical visual word form area (VWFA). This putative grapheme-related area (GRA) differed from the VWFA in being preferentially connected functionally to dorsal parietal areas involved in letter-by-letter reading, while the VWFA showed effects of lexicality and spelling-to-sound regularity. Our results suggest a partial dissociation within left occipitotemporal cortex: the midfusiform GRA would encode orthographic information at a sublexical graphemic level, while the lateral occipitotemporal VWFA would contribute primarily to direct lexico-semantic access.

Reading music and words: The anatomical connectivity of musicians' visual cortex.

Musical score reading and word reading have much in common, from their historical origins to their cognitive foundations and neural correlates. In the ventral occipitotemporal cortex (VOT), the specialization of the so-called Visual Word Form Area for word reading has been linked to its privileged structural connectivity to distant language regions. Here we investigated how anatomical connectivity relates to the segregation of regions specialized for musical notation or words in the VOT. In a cohort of professional musicians and non-musicians, we used probabilistic tractography combined with task-related functional MRI to identify the connections of individually defined word- and music-selective left VOT regions. Despite their close proximity, these regions differed significantly in their structural connectivity, irrespective of musical expertise. The music-selective region was significantly more connected to posterior lateral temporal regions than the word-selective region, which, conversely, was significantly more connected to anterior ventral temporal cortex. Furthermore, musical expertise had a double impact on the connectivity of the music region. First, music tracts were significantly larger in musicians than in non-musicians, associated with marginally higher connectivity to perisylvian music-related areas. Second, the spatial similarity between music and word tracts was significantly increased in musicians, consistently with the increased overlap of language and music functional activations in musicians, as compared to non-musicians. These results support the view that, for music as for words, very specific anatomical connections influence the specialization of distinct VOT areas, and that reciprocally those connections are selectively enhanced by the expertise for word or music reading.

Impact of literacy on the functional connectivity of vision and language related networks.

Learning to read leads to functional and structural changes in cortical brain areas related to vision and language. Previous evidence suggests that the Visual Word Form Area (VWFA), a region devoted to the recognition of letter strings in literate persons, acts as an interface between both systems. While different studies have performed univariate analyses to study the effects of literacy on brain function, little is known about its impact on whole functional networks, especially when literacy is acquired during adulthood. We investigated functional connectivity in three groups of adults with different literacy status: illiterates, ex-illiterates (i.e., who learned to read during adulthood), and literates (i.e., who learned to read in childhood). We used a data-driven, multivariate whole brain approach (Independent Component Analysis [ICA]) combined with a region of interest (ROI) analysis in order to explore the functional connectivity of the VWFA with four ICA networks related to vision and language functions. ICA allowed for the identification of four networks of interest: left fronto-parietal, auditory, medial visual and lateral visual functional networks, plus a control right fronto-parietal network. We explored the effects literacy on the connectivity between the VWFA and these networks, trying furthermore to disentangle the roles of reading proficiency and age of acquisition (i.e., literacy status) in these changes. Results showed that functional connectivity between the VWFA and the left fronto-parietal and lateral visual networks increased and decreased, respectively, with literacy. Moreover, the functional coupling of the VWFA and the auditory network decreased with literacy. This study provides novel insights in the mechanisms of reading acquisition and brain plasticity, putting to light the emergence of the VWFA as a bridge between language and vision. Further studies are required to characterize the interplay of proficiency and age of reading acquisition, and its relevance to models of brain plasticity across lifespan.

Illiterate to literate: behavioural and cerebral changes induced by reading acquisition.

The acquisition of literacy transforms the human brain. By reviewing studies of illiterate subjects, we propose specific hypotheses on how the functions of core brain systems are partially reoriented or 'recycled' when learning to read. Literacy acquisition improves early visual processing and reorganizes the ventral occipito-temporal pathway: responses to written characters are increased in the left occipito-temporal sulcus, whereas responses to faces shift towards the right hemisphere. Literacy also modifies phonological coding and strengthens the functional and anatomical link between phonemic and graphemic representations. Literacy acquisition therefore provides a remarkable example of how the brain reorganizes to accommodate a novel cultural skill.

When colours split from objects: The disconnection of colour perception from colour language and colour knowledge.

We investigated object-colour knowledge in RDS, a patient with impaired colour naming after a left occipito-temporal stroke. RDS's colour perception, object naming and verbal colour-knowledge (the ability to verbally say the typical colour of an object) were relatively spared. RDS was also able to state if an object was appropriately coloured or not. However, he could neither match colour names to coloured objects, nor match colour patches to grey-scale objects. Thus, RDS's colour-naming deficit was associated with an impaired ability to conceptually relate visually presented object shapes and colours. These results suggest that objects in their typical colour are processed holistically in the visual modality, and that abilities important for colour naming may also be involved in abstracting colours from visual objects. We discuss these findings in the context of developmental psychology and linguistic anthropology, and propose a model of neuro-functional organization of object-colour knowledge.

Distinctive Interaction Between Cognitive Networks and the Visual Cortex in Early Blind Individuals.

In early blind individuals, brain activation by a variety of nonperceptual cognitive tasks extends to the visual cortex, while in the sighted it is restricted to supramodal association areas. We hypothesized that such activation results from the integration of different sectors of the visual cortex into typical task-dependent networks. We tested this hypothesis with fMRI in blind and sighted subjects using tasks assessing speech comprehension, incidental long-term memory and both verbal and nonverbal executive control, in addition to collecting resting-state data. All tasks activated the visual cortex in blind relative to sighted subjects, which enabled its segmentation according to task sensitivity. We then assessed the unique brain-scale functional connectivity of the segmented areas during resting state. Language-related seeds were preferentially connected to frontal and temporal language areas; the seed derived from the executive task was connected to the right dorsal frontoparietal executive network; and the memory-related seed was uniquely connected to mesial frontoparietal areas involved in episodic memory retrieval. Thus, using a broad set of language, executive, and memory tasks in the same subjects, combined with resting state connectivity, we demonstrate the selective integration of different patches of the visual cortex into brain-scale networks with distinct localization, lateralization, and functional roles.

Neurophysiological dynamics of phrase-structure building during sentence processing.

Although sentences unfold sequentially, one word at a time, most linguistic theories propose that their underlying syntactic structure involves a tree of nested phrases rather than a linear sequence of words. Whether and how the brain builds such structures, however, remains largely unknown. Here, we used human intracranial recordings and visual word-by-word presentation of sentences and word lists to investigate how left-hemispheric brain activity varies during the formation of phrase structures. In a broad set of language-related areas, comprising multiple superior temporal and inferior frontal sites, high-gamma power increased with each successive word in a sentence but decreased suddenly whenever words could be merged into a phrase. Regression analyses showed that each additional word or multiword phrase contributed a similar amount of additional brain activity, providing evidence for a merge operation that applies equally to linguistic objects of arbitrary complexity. More superficial models of language, based solely on sequential transition probability over lexical and syntactic categories, only captured activity in the posterior middle temporal gyrus. Formal model comparison indicated that the model of multiword phrase construction provided a better fit than probability-based models at most sites in superior temporal and inferior frontal cortices. Activity in those regions was consistent with a neural implementation of a bottom-up or left-corner parser of the incoming language stream. Our results provide initial intracranial evidence for the neurophysiological reality of the merge operation postulated by linguists and suggest that the brain compresses syntactically well-formed sequences of words into a hierarchy of nested phrases.

The cerebral bases of the bouba-kiki effect.

The crossmodal correspondence between some speech sounds and some geometrical shapes, known as the bouba-kiki (BK) effect, constitutes a remarkable exception to the general arbitrariness of the links between word meaning and word sounds. We have analyzed the association of shapes and sounds in order to determine whether it occurs at a perceptual or at a decisional level, and whether it takes place in sensory cortices or in supramodal regions. First, using an Implicit Association Test (IAT), we have shown that the BK effect may occur without participants making any explicit decision relative to sound-shape associations. Second, looking for the brain correlates of implicit BK matching, we have found that intermodal matching influences activations in both auditory and visual sensory cortices. Moreover, we found stronger prefrontal activation to mismatching than to matching stimuli, presumably reflecting a modulation of executive processes by crossmodal correspondence. Thus, through its roots in the physiology of object categorization and crossmodal matching, the BK effect provides a unique insight into some non-linguistic components of word formation.

Musical literacy shifts asymmetries in the ventral visual cortex.

The acquisition of literacy has a profound impact on the functional specialization and lateralization of the visual cortex. Due to the overall lateralization of the language network, specialization for printed words develops in the left occipitotemporal cortex, allegedly inducing a secondary shift of visual face processing to the right, in literate as compared to illiterate subjects. Applying the same logic to the acquisition of high-level musical literacy, we predicted that, in musicians as compared to non-musicians, occipitotemporal activations should show a leftward shift for music reading, and an additional rightward push for face perception. To test these predictions, professional musicians and non-musicians viewed pictures of musical notation, faces, words, tools and houses in the MRI, and laterality was assessed in the ventral stream combining ROI and voxel-based approaches. The results supported both predictions, and allowed to locate the leftward shift to the inferior temporal gyrus and the rightward shift to the fusiform cortex. Moreover, these laterality shifts generalized to categories other than music and faces. Finally, correlation measures across subjects did not support a causal link between the leftward and rightward shifts. Thus the acquisition of an additional perceptual expertise extensively modifies the laterality pattern in the visual system.

Timing the impact of literacy on visual processing.

Learning to read requires the acquisition of an efficient visual procedure for quickly recognizing fine print. Thus, reading practice could induce a perceptual learning effect in early vision. Using functional magnetic resonance imaging (fMRI) in literate and illiterate adults, we previously demonstrated an impact of reading acquisition on both high- and low-level occipitotemporal visual areas, but could not resolve the time course of these effects. To clarify whether literacy affects early vs. late stages of visual processing, we measured event-related potentials to various categories of visual stimuli in healthy adults with variable levels of literacy, including completely illiterate subjects, early-schooled literate subjects, and subjects who learned to read in adulthood (ex-illiterates). The stimuli included written letter strings forming pseudowords, on which literacy is expected to have a major impact, as well as faces, houses, tools, checkerboards, and false fonts. To evaluate the precision with which these stimuli were encoded, we studied repetition effects by presenting the stimuli in pairs composed of repeated, mirrored, or unrelated pictures from the same category. The results indicate that reading ability is correlated with a broad enhancement of early visual processing, including increased repetition suppression, suggesting better exemplar discrimination, and increased mirror discrimination, as early as ∼ 100-150 ms in the left occipitotemporal region. These effects were found with letter strings and false fonts, but also were partially generalized to other visual categories. Thus, learning to read affects the magnitude, precision, and invariance of early visual processing.

Exploring the anatomical encoding of voice with a mathematical model of the vocal system.

The faculty of language depends on the interplay between the production and perception of speech sounds. A relevant open question is whether the dimensions that organize voice perception in the brain are acoustical or depend on properties of the vocal system that produced it. One of the main empirical difficulties in answering this question is to generate sounds that vary along a continuum according to the anatomical properties the vocal apparatus that produced them. Here we use a mathematical model that offers the unique possibility of synthesizing vocal sounds by controlling a small set of anatomically based parameters. In a first stage the quality of the synthetic voice was evaluated. Using specific time traces for sub-glottal pressure and tension of the vocal folds, the synthetic voices generated perceptual responses, which are indistinguishable from those of real speech. The synthesizer was then used to investigate how the auditory cortex responds to the perception of voice depending on the anatomy of the vocal apparatus. Our fMRI results show that sounds are perceived as human vocalizations when produced by a vocal system that follows a simple relationship between the size of the vocal folds and the vocal tract. We found that these anatomical parameters encode the perceptual vocal identity (male, female, child) and show that the brain areas that respond to human speech also encode vocal identity. On the basis of these results, we propose that this low-dimensional model of the vocal system is capable of generating realistic voices and represents a novel tool to explore the voice perception with a precise control of the anatomical variables that generate speech. Furthermore, the model provides an explanation of how auditory cortices encode voices in terms of the anatomical parameters of the vocal system.

Event-Related Potential, Time-frequency, and Functional Connectivity Facets of Local and Global Auditory Novelty Processing: An Intracranial Study in Humans.

Auditory novelty detection has been associated with different cognitive processes. Bekinschtein et al. (2009) developed an experimental paradigm to dissociate these processes, using local and global novelty, which were associated, respectively, with automatic versus strategic perceptual processing. They have mostly been studied using event-related potentials (ERPs), but local spiking activity as indexed by gamma (60-120 Hz) power and interactions between brain regions as indexed by modulations in beta-band (13-25 Hz) power and functional connectivity have not been explored. We thus recorded 9 epileptic patients with intracranial electrodes to compare the precise dynamics of the responses to local and global novelty. Local novelty triggered an early response observed as an intracranial mismatch negativity (MMN) contemporary with a strong power increase in the gamma band and an increase in connectivity in the beta band. Importantly, all these responses were strictly confined to the temporal auditory cortex. In contrast, global novelty gave rise to a late ERP response distributed across brain areas, contemporary with a sustained power decrease in the beta band (13-25 Hz) and an increase in connectivity in the alpha band (8-13 Hz) within the frontal lobe. We discuss these multi-facet signatures in terms of conscious access to perceptual information.

Anatomical connections of the visual word form area.

The visual word form area (VWFA), a region systematically involved in the identification of written words, occupies a reproducible location in the left occipitotemporal sulcus in expert readers of all cultures. Such a reproducible localization is paradoxical, given that reading is a recent invention that could not have influenced the genetic evolution of the cortex. Here, we test the hypothesis that the VWFA recycles a region of the ventral visual cortex that shows a high degree of anatomical connectivity to perisylvian language areas, thus providing an efficient circuit for both grapheme-phoneme conversion and lexical access. In two distinct experiments, using high-resolution diffusion-weighted data from 75 human subjects, we show that (1) the VWFA, compared with the fusiform face area, shows higher connectivity to left-hemispheric perisylvian superior temporal, anterior temporal and inferior frontal areas; (2) on a posterior-to-anterior axis, its localization within the left occipitotemporal sulcus maps onto a peak of connectivity with language areas, with slightly distinct subregions showing preferential projections to areas respectively involved in grapheme-phoneme conversion and lexical access. In agreement with functional data on the VWFA in blind subjects, the results suggest that connectivity to language areas, over and above visual factors, may be the primary determinant of VWFA localization.

Large scale screening of neural signatures of consciousness in patients in a vegetative or minimally conscious state.

In recent years, numerous electrophysiological signatures of consciousness have been proposed. Here, we perform a systematic analysis of these electroencephalography markers by quantifying their efficiency in differentiating patients in a vegetative state from those in a minimally conscious or conscious state. Capitalizing on a review of previous experiments and current theories, we identify a series of measures that can be organized into four dimensions: (i) event-related potentials versus ongoing electroencephalography activity; (ii) local dynamics versus inter-electrode information exchange; (iii) spectral patterns versus information complexity; and (iv) average versus fluctuations over the recording session. We analysed a large set of 181 high-density electroencephalography recordings acquired in a 30 minutes protocol. We show that low-frequency power, electroencephalography complexity, and information exchange constitute the most reliable signatures of the conscious state. When combined, these measures synergize to allow an automatic classification of patients' state of consciousness.

Learning to read improves the structure of the arcuate fasciculus.

The acquisition of literacy results from an effortful learning process that leads to functional changes in several cortical regions. We explored whether learning to read also leads to anatomical changes within the left intrahemispheric white matter pathways that interconnect these regions. Using diffusion tensor imaging tractography, we compared illiterates with ex-illiterates who learned to read during adulthood and literates who learned to read during their childhood. Literacy related to an increase in fractional anisotropy and a decrease in perpendicular diffusivity in the temporo-parietal portion of the left arcuate fasciculus. The microstructure within this pathway correlated with the reading performance and the degree of functional activation within 2 dominant brain regions involved in reading: The Visual Word Form Area in response to letter strings, and the posterior superior temporal cortex in response to spoken language. Thus, the acquisition of literacy is associated with a reinforcement of left temporo-parietal connections whose microstructure predicts overall reading performance and the functional specialization of the Visual Word Form Area. This anatomical magnetic resonance imaging marker may be useful to predict developmental reading disorders.

Universal brain systems for recognizing word shapes and handwriting gestures during reading.

Do the neural circuits for reading vary across culture? Reading of visually complex writing systems such as Chinese has been proposed to rely on areas outside the classical left-hemisphere network for alphabetic reading. Here, however, we show that, once potential confounds in cross-cultural comparisons are controlled for by presenting handwritten stimuli to both Chinese and French readers, the underlying network for visual word recognition may be more universal than previously suspected. Using functional magnetic resonance imaging in a semantic task with words written in cursive font, we demonstrate that two universal circuits, a shape recognition system (reading by eye) and a gesture recognition system (reading by hand), are similarly activated and show identical patterns of activation and repetition priming in the two language groups. These activations cover most of the brain regions previously associated with culture-specific tuning. Our results point to an extended reading network that invariably comprises the occipitotemporal visual word-form system, which is sensitive to well-formed static letter strings, and a distinct left premotor region, Exner's area, which is sensitive to the forward or backward direction with which cursive letters are dynamically presented. These findings suggest that cultural effects in reading merely modulate a fixed set of invariant macroscopic brain circuits, depending on surface features of orthographies.