Overlapping but separate number representations in the intraparietal sulcus-Probing format- and modality-independence in sighted Braille readers.

The Triple-Code Model stipulates that numerical information from different formats and modalities converges on a common magnitude representation in the Intraparietal Sulcus (IPS). To what extent the representations of all numerosity forms overlap remains unsolved. It has been postulated that the representation of symbolic numerosities (for example, Arabic digits) is sparser and grounded in an existing representation that codes for non-symbolic numerosity information (i.e., sets of objects). Other theories argue that numerical symbols represent a separate number category that emerges only during education. Here, we tested a unique group of sighted tactile Braille readers with numerosities 2, 4, 6 and 8 in three number notations: Arabic digits, sets of dots, tactile Braille numbers. Using univariate methods, we showed a consistent overlap in activations evoked by these three number notations. This result shows that all three used notations are represented in the IPS, which may suggest at least a partial overlap between the representations of the three notations used in this experiment. Using MVPA, we found that only non-automatized number information (Braille and sets of dots) allowed successful number classification. However, the numerosity of one notation could not be predicted above chance from the brain activation patterns evoked by another notation (no cross-classification). These results show that the IPS may host independent number codes in overlapping cortical circuits. In addition, they suggest that the level of training in encoding a given type of number information is an important factor that determines the amount of exploitable information and needs to be controlled for in order to identify the neural code underlying numerical information per se.

Temporal Dynamics of Brain White Matter Plasticity in Sighted Subjects during Tactile Braille Learning: A Longitudinal Diffusion Tensor Imaging Study.

The white matter (WM) architecture of the human brain changes in response to training, though fine-grained temporal characteristics of training-induced white matter plasticity remain unexplored. We investigated white matter microstructural changes using diffusion tensor imaging at five different time points in 26 sighted female adults during 8 months of training on tactile braille reading. Our results show that training-induced white matter plasticity occurs both within and beyond the trained sensory modality, as reflected by fractional anisotropy (FA) increases in somatosensory and visual cortex, respectively. The observed changes followed distinct time courses, with gradual linear FA increase along the training in the somatosensory cortex and sudden visual cortex cross-modal plasticity occurring after braille input became linguistically meaningful. WM changes observed in these areas returned to baseline after the cessation of learning in line with the supply-demand model of plasticity. These results also indicate that the temporal dynamics of microstructural plasticity in different cortical regions might be modulated by the nature of computational demands. We provide additional evidence that observed FA training-induced changes are behaviorally relevant to tactile reading. Together, these results demonstrate that WM plasticity is a highly dynamic process modulated by the introduction of novel experiences.SIGNIFICANCE STATEMENT Throughout the lifetime the human brain is shaped by various experiences. Training-induced reorganization in white matter (WM) microstructure has been reported, but we know little about its temporal dynamics. To fill this gap, we scanned sighted subjects five times during tactile braille reading training. We observed different dynamics of WM plasticity in the somatosensory and visual cortices implicated in braille reading. The former showed a continuous increase in WM tissue anisotropy along with tactile training, while microstructural changes in the latter were observed only after the participants learned to read braille words. Our results confirm the supply-demand model of brain plasticity and provide evidence that WM reorganization depends on distinct computational demands and functional roles of regions involved in the trained skill.

Brain plasticity dynamics during tactile Braille learning in sighted subjects: Multi-contrast MRI approach.

A growing body of empirical evidence supports the notion of diverse neurobiological processes underlying learning-induced plasticity changes in the human brain. There are still open questions about how brain plasticity depends on cognitive task complexity, how it supports interactions between brain systems and with what temporal and spatial trajectory. We investigated brain and behavioural changes in sighted adults during 8-months training of tactile Braille reading whilst monitoring brain structure and function at 5 different time points. We adopted a novel multivariate approach that includes behavioural data and specific MRI protocols sensitive to tissue properties to assess local functional and structural and myelin changes over time. Our results show that while the reading network, located in the ventral occipitotemporal cortex, rapidly adapts to tactile input, sensory areas show changes in grey matter volume and intra-cortical myelin at different times. This approach has allowed us to examine and describe neuroplastic mechanisms underlying complex cognitive systems and their (sensory) inputs and (motor) outputs differentially, at a mesoscopic level.

Tactile to visual number priming in the left intraparietal cortex of sighted Braille readers.

Numbers can be presented in different notations and sensory modalities. It is currently debated to what extent these formats overlap onto a single representation. We asked whether such an overlap exists between symbolic numbers represented in two sensory modalities: Arabic digits and Braille numbers. A unique group of sighted Braille readers underwent extensive Braille reading training and was tested in an fMRI repetition-suppression paradigm with tactile Braille digit primes and visual Arabic digit targets. Our results reveal cross-modal priming: compared to repetition of two different quantities (e.g., Braille "5" and Arabic "2"), repetition of the same quantity presented in two modalities (e.g., Braille "5" and Arabic "5") led to a reduction of activation in several sub-regions of the Intraparietal Sulcus (IPS), a key cortical region for magnitude processing. Thus, in sighted Braille readers, the representations of numbers read by sight and by touch overlap to a degree sufficient to cause repetition suppression. This effect was modulated by the numerical prime-probe distance. Altogether this indicates that the left parietal cortex hosts neural assemblies that are sensitive to numerical information from different notations (number words or Arabic digits) and modalities (tactile and visual).

A shared code for Braille and Arabic digits revealed by cross-modal priming in sighted Braille readers.

Quantities can be represented by different formats (e.g. symbolic or non-symbolic) and conveyed via different modalities (e.g. tactile or visual). Despite different priming curves: V-shape and step-shape for place and summation coded representation, respectively, the occurrence of priming effect supports the notion of different format overlap on the same mental number line. However, little is known about tactile-visual overlap of symbolic numerosities i.e. Braille numbers to Arabic digits on the magnitude number representation. Here, in a priming experiment, we tested a unique group of sighted Braille readers to investigate whether tactile Braille digits would activate a place-coding type of mental number representation (V-shape), analogous to other symbolic formats. The primes were either tactile Braille digits presented on a Braille display or number words presented on a computer screen. The targets were visually presented Arabic digits, and subjects performed a naming task. Our results reveal a V-shape priming function for both prime formats: tactile Braille and written words representing numbers, with strongest priming for primes of identical value (e.g. "four" and "4"), and a symmetrical decrease of priming strength for neighboring numbers, which indicates that the observed priming is due to identity priming. We thus argue that the magnitude information is processed according to a shared phonological code, independent of the input modality.

Functional hierarchy for tactile processing in the visual cortex of sighted adults.

Perception via different sensory modalities was traditionally believed to be supported by largely separate brain systems. However, a growing number of studies demonstrate that the visual cortices of typical, sighted adults are involved in tactile and auditory perceptual processing. Here, we investigated the spatiotemporal dynamics of the visual cortex's involvement in a complex tactile task: Braille letter recognition. Sighted subjects underwent Braille training and then participated in a transcranial magnetic stimulation (TMS) study in which they tactually identified single Braille letters. During this task, TMS was applied to their left early visual cortex, visual word form area (VWFA), and left early somatosensory cortex at five time windows from 20 to 520 ms following the Braille letter presentation's onset. The subjects' response accuracy decreased when TMS was applied to the early visual cortex at the 120-220 ms time window and when TMS was applied to the VWFA at the 320-420 ms time window. Stimulation of the early somatosensory cortex did not have a time-specific effect on the accuracy of the subjects' Braille letter recognition, but rather caused a general slowdown during this task. Our results indicate that the involvement of sighted people's visual cortices in tactile perception respects the canonical visual hierarchy-the early tactile processing stages involve the early visual cortex, whereas more advanced tactile computations involve high-level visual areas. Our findings are compatible with the metamodal account of brain organization and suggest that the whole visual cortex may potentially support spatial perception in a task-specific, sensory-independent manner.

Structural reorganization of the early visual cortex following Braille training in sighted adults.

Training can induce cross-modal plasticity in the human cortex. A well-known example of this phenomenon is the recruitment of visual areas for tactile and auditory processing. It remains unclear to what extent such plasticity is associated with changes in anatomy. Here we enrolled 29 sighted adults into a nine-month tactile Braille-reading training, and used voxel-based morphometry and diffusion tensor imaging to describe the resulting anatomical changes. In addition, we collected resting-state fMRI data to relate these changes to functional connectivity between visual and somatosensory-motor cortices. Following Braille-training, we observed substantial grey and white matter reorganization in the anterior part of early visual cortex (peripheral visual field). Moreover, relative to its posterior, foveal part, the peripheral representation of early visual cortex had stronger functional connections to somatosensory and motor cortices even before the onset of training. Previous studies show that the early visual cortex can be functionally recruited for tactile discrimination, including recognition of Braille characters. Our results demonstrate that reorganization in this region induced by tactile training can also be anatomical. This change most likely reflects a strengthening of existing connectivity between the peripheral visual cortex and somatosensory cortices, which suggests a putative mechanism for cross-modal recruitment of visual areas.

Universal Visual Features Might Be Necessary for Fluent Reading. A Longitudinal Study of Visual Reading in Braille and Cyrillic Alphabets.

It has been hypothesized that efficient reading is possible because all reading scripts have been matched, through cultural evolution, to the natural capabilities of the visual cortex. This matching has resulted in all scripts being made of line-junctions, such as T, X, or L. Our aim was to test a critical prediction of this hypothesis: visual reading in an atypical script that is devoid of line-junctions (such as the Braille alphabet read visually) should be much less efficient than reading in a "normal" script (e.g., Cyrillic). Using a lexical decision task, we examined Visual Braille reading speed and efficiency in sighted Braille teachers. As a control, we tested learners of a natural visual script, Cyrillic. Both groups participated in a two semester course of either visual Braille or Russian while their reading speed and accuracy was tested at regular intervals. The results show that visual Braille reading is slow, prone to errors and highly serial, even in Braille readers with years of prior reading experience. Although subjects showed some improvements in their visual Braille reading accuracy and speed following the course, the effect of word length on reading speed (typically observed in beginning readers) was remained very sizeable through all testing sessions. These results are in stark contrast to Cyrillic, a natural script, where only 3 months of learning were sufficient to achieve relative proficiency. Taken together, these results suggest that visual features such as line junctions and their combinations might be necessary for efficient reading.

Braille in the Sighted: Teaching Tactile Reading to Sighted Adults.

Blind people are known to have superior perceptual abilities in their remaining senses. Several studies suggest that these enhancements are dependent on the specific experience of blind individuals, who use those remaining senses more than sighted subjects. In line with this view, sighted subjects, when trained, are able to significantly progress in relatively simple tactile tasks. However, the case of complex tactile tasks is less obvious, as some studies suggest that visual deprivation itself could confer large advantages in learning them. It remains unclear to what extent those complex skills, such as braille reading, can be learnt by sighted subjects. Here we enrolled twenty-nine sighted adults, mostly braille teachers and educators, in a 9-month braille reading course. At the beginning of the course, all subjects were naive in tactile braille reading. After the course, almost all were able to read whole braille words at a mean speed of 6 words-per-minute. Subjects with low tactile acuity did not differ significantly in braille reading speed from the rest of the group, indicating that low tactile acuity is not a limiting factor for learning braille, at least at this early stage of learning. Our study shows that most sighted adults can learn whole-word braille reading, given the right method and a considerable amount of motivation. The adult sensorimotor system can thus adapt, to some level, to very complex tactile tasks without visual deprivation. The pace of learning in our group was comparable to congenitally and early blind children learning braille in primary school, which suggests that the blind's mastery of complex tactile tasks can, to a large extent, be explained by experience-dependent mechanisms.

Massive cortical reorganization in sighted Braille readers.

The brain is capable of large-scale reorganization in blindness or after massive injury. Such reorganization crosses the division into separate sensory cortices (visual, somatosensory...). As its result, the visual cortex of the blind becomes active during tactile Braille reading. Although the possibility of such reorganization in the normal, adult brain has been raised, definitive evidence has been lacking. Here, we demonstrate such extensive reorganization in normal, sighted adults who learned Braille while their brain activity was investigated with fMRI and transcranial magnetic stimulation (TMS). Subjects showed enhanced activity for tactile reading in the visual cortex, including the visual word form area (VWFA) that was modulated by their Braille reading speed and strengthened resting-state connectivity between visual and somatosensory cortices. Moreover, TMS disruption of VWFA activity decreased their tactile reading accuracy. Our results indicate that large-scale reorganization is a viable mechanism recruited when learning complex skills.