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How conceptual knowledge is represented in the human brain remains to be determined. To address the differential role of low-level sensory-based and high-level abstract features in semantic processing, we combined behavioral studies of linguistic production and brain activity measures by functional magnetic resonance imaging in sighted and congenitally blind individuals while they performed a property-generation task with concrete nouns from eight categories, presented through visual and/or auditory modalities. Patterns of neural activity within a large semantic cortical network that comprised parahippocampal, lateral occipital, temporo-parieto-occipital and inferior parietal cortices correlated with linguistic production and were independent both from the modality of stimulus presentation (either visual or auditory) and the (lack of) visual experience. In contrast, selected modality-dependent differences were observed only when the analysis was limited to the individual regions within the semantic cortical network. We conclude that conceptual knowledge in the human brain relies on a distributed, modality-independent cortical representation that integrates the partial category and modality specific information retained at a regional level., (Copyright © 2016 The Authors. Published by Elsevier Inc. All rights reserved.)

While there is ample evidence that the structure and function of visual cortical areas are affected by early visual deprivation, little is known of how early blindness modifies subcortical relay and association thalamic nuclei, as well as mesencephalic structures. Therefore, in the present multicenter study, we used MRI to measure volume of the superior and inferior colliculi, as well as of the thalamic nuclei relaying sensory and motor information to the neocortex, parcellated according to atlas-based thalamo-cortical connections, in 29 individuals with congenital blindness of peripheral origin (17 M, age 35.7 ± 14.3 years) and 29 sighted subjects (17 M, age 31.9 ± 9.0). Blind participants showed an overall volume reduction in the left (p = 0.008) and right (p = 0.007) thalami, as compared to the sighted individuals. Specifically, the lateral geniculate (i.e., primary visual thalamic relay nucleus) was 40% reduced (left: p = 4 × 10(-6), right: p < 1 × 10(-6)), consistent with findings from animal studies. In addition, associated thalamic nuclei that project to temporal (left: p = 0.005, right: p = 0.005), prefrontal (left: p = 0.010, right: p = 0.014), occipital (left: p = 0.005, right: p = 0.023), and right premotor (p = 0.024) cortical regions were also significantly reduced in the congenitally blind group. Conversely, volumes of the relay nuclei directly involved in auditory, motor, and somatosensory processing were not affected by visual deprivation. In contrast, no difference in volume was observed in either the superior or the inferior colliculus between the two groups. Our findings indicate that visual loss since birth leads to selective volumetric changes within diencephalic, but not mesencephalic, structures. Both changes in reciprocal cortico-thalamic connections or modifications in the intrinsic connectivity between relay and association nuclei of the thalamus may contribute to explain these alterations in thalamic volumes. Sparing of the superior colliculi is in line with their composite, multisensory projections, and with their not exclusive visual nature.

Although vision offers distinctive information to space representation, individuals who lack vision since birth often show perceptual and representational skills comparable to those found in sighted individuals. However, congenitally blind individuals may result in impaired spatial analysis, when engaging in 'visual' spatial features (e.g., perspective or angle representation) or complex spatial mental abilities. In the present study, we measured behavioral and brain responses using functional magnetic resonance imaging in sighted and congenitally blind individuals during spatial imagery based on a modified version of the mental clock task (e.g., angle discrimination) and a simple recognition control condition, as conveyed across distinct sensory modalities: visual (sighted individuals only), tactile and auditory. Blind individuals were significantly less accurate during the auditory task, but comparable-to-sighted during the tactile task. As expected, both groups showed common neural activations in intraparietal and superior parietal regions across visual and non-visual spatial perception and imagery conditions, indicating the more abstract, sensory independent functional organization of these cortical areas, a property that we named supramodality. At the same time, however, comparisons in brain responses and functional connectivity patterns across experimental conditions demonstrated also a functional lateralization, in a way that correlated with the distinct behavioral performance in blind and sighted individuals. Specifically, blind individuals relied more on right parietal regions, mainly in the tactile and less in the auditory spatial processing. In sighted, spatial representation across modalities relied more on left parietal regions. In conclusions, intraparietal and superior parietal regions subserve supramodal spatial representations in sighted and congenitally blind individuals. Differences in their recruitment across non-visual spatial processing in sighted and blind individuals may be related to distinctive behavioral performance and/or mental strategies adopted when they deal with the same spatial representation as conveyed through different sensory modalities., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

We examined the effects of visual deprivation at birth on the development of the corpus callosum in a large group of congenitally blind individuals. We acquired high-resolution T1-weighted MRI scans in 28 congenitally blind and 28 normal sighted subjects matched for age and gender. There was no overall group effect of visual deprivation on the total surface area of the corpus callosum. However, subdividing the corpus callosum into five subdivisions revealed significant regional changes in its three most posterior parts. Compared to the sighted controls, congenitally blind individuals showed a 12% reduction in the splenium, and a 20% increase in the isthmus and the posterior part of the body. A shape analysis further revealed that the bending angle of the corpus callosum was more convex in congenitally blind compared to the sighted control subjects. The observed morphometric changes in the corpus callosum are in line with the well-described cross-modal functional and structural neuroplastic changes in congenital blindness.

While most of the research in blind individuals classically has focused on the compensatory plastic rearrangements that follow loss of sight, novel behavioral, anatomical and functional brain studies in individuals born deprived of sight represent a powerful tool to understand to what extent the brain functional architecture is programmed to develop independently from any visual experience. Here we review work from our lab and others, conducted in sighted and congenitally blind individuals, whose results indicate that vision is not a mandatory prerequisite for the brain cortical organization to develop and function. Similar cortical networks subtend visual and/or non-visual perception of form, space and movement, as well as action recognition, both in sighted and in congenitally blind individuals. These findings support the hypothesis of a modality independent, supramodal cortical organization. Visual experience, however, does play a role in shaping specific cortical sub-regions, as loss of sight is accompanied also by cross-modal plastic phenomena. Altogether, studying the blind brain is opening our eyes on how the brain develops and works., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Cross-modal responses in occipital areas appear to be essential for sensory processing in visually deprived subjects. However, it is yet unclear whether this functional recruitment might be dependent on the sensory channel conveying the information. In order to characterize brain areas showing task-independent, but sensory specific, cross-modal responses in blind individuals, we pooled together distinct brain functional studies in a single based meta-analysis according only to the modality conveying experimental stimuli (auditory or tactile). Our approach revealed a specific functional cortical segregation according to the sensory modality conveying the non-visual information, irrespectively from the cognitive features of the tasks. In particular, dorsal and posterior subregions of occipital and superior parietal cortex showed a higher cross-modal recruitment across tactile tasks in blind as compared to sighted individuals. On the other hand, auditory stimuli activated more medial and ventral clusters within early visual areas, the lingual and inferior temporal cortex. These findings suggest a modality-specific functional modification of cross-modal responses within different portions of the occipital cortex of blind individuals. Cross-modal recruitment can thus be specifically influenced by the intrinsic features of sensory information.

Vision is important for avoiding encounters with objects in the environment that may imperil physical integrity. We tested whether, in the absence of vision, a lower pain threshold would arise from an adaptive shift to other sensory channels. We therefore measured heat and cold pain thresholds and responses to suprathreshold heat stimuli in 2 groups of congenitally blind and matched normal-sighted participants. We also assessed detection thresholds for innocuous warmth and cold, and participants' attitude toward painful encounters in daily life. Our results show that, compared to sighted subjects, congenitally blind subjects have lower heat pain thresholds, rate suprathreshold heat pain stimuli as more painful, and have increased sensitivity for cold pain stimuli. Thresholds for nonpainful thermal stimulation did not differ between groups. The results of the pain questionnaires further indicated that blind subjects are more attentive to signals of external threats. These findings indicate that the absence of vision from birth induces a hypersensitivity to painful stimuli, lending new support to a model of sensory integration of vision and pain processing., (Copyright © 2013 International Association for the Study of Pain. Published by Elsevier B.V. All rights reserved.)

The representation of actions within the action-observation network is thought to rely on a distributed functional organization. Furthermore, recent findings indicate that the action-observation network encodes not merely the observed motor act, but rather a representation that is independent from a specific sensory modality or sensory experience. In the present study, we wished to determine to what extent this distributed and 'more abstract' representation of action is truly supramodal, i.e. shares a common coding across sensory modalities. To this aim, a pattern recognition approach was employed to analyze neural responses in sighted and congenitally blind subjects during visual and/or auditory presentation of hand-made actions. Multivoxel pattern analyses-based classifiers discriminated action from non-action stimuli across sensory conditions (visual and auditory) and experimental groups (blind and sighted). Moreover, these classifiers labeled as 'action' the pattern of neural responses evoked during actual motor execution. Interestingly, discriminative information for the action/non action classification was located in a bilateral, but left-prevalent, network that strongly overlaps with brain regions known to form the action-observation network and the human mirror system. The ability to identify action features with a multivoxel pattern analyses-based classifier in both sighted and blind individuals and independently from the sensory modality conveying the stimuli clearly supports the hypothesis of a supramodal, distributed functional representation of actions, mainly within the action-observation network.

Previous studies in early blind individuals posited a possible role of parieto-occipital connections in conveying nonvisual information to the visual occipital cortex. As a consequence of blindness, parietal areas would thus become able to integrate a greater amount of multimodal information than in sighted individuals. To verify this hypothesis, we compared fMRI-measured BOLD signal temporal variability, an index of efficiency in functional information integration, in congenitally blind and sighted individuals during tactile spatial discrimination and motion perception tasks. In both tasks, the BOLD variability analysis revealed many cortical regions with a significantly greater variability in the blind as compared to sighted individuals, with an overlapping cluster located in the left inferior parietal/anterior intraparietal cortex. A functional connectivity analysis using this region as seed showed stronger correlations in both tasks with occipital areas in the blind as compared to sighted individuals. As BOLD variability reflects neural integration and processing efficiency, these cross-modal plastic changes in the parietal cortex, even if described in a limited sample, reinforce the hypothesis that this region may play an important role in processing nonvisual information in blind subjects and act as a hub in the cortico-cortical pathway from somatosensory cortex to the reorganized occipital areas.

Purpose of Review: In this review, we discuss findings from some recent brain imaging studies that shed new light on our understanding of the role of visual experience on the development of the brain morphological and functional architecture in humans. To what extent is vision truly necessary to 'see' the world around us?, Recent Findings: Congenitally blind and sighted individuals present analogous cognitive and social performances. Findings from structural and functional brain studies in both sighted and congenitally blind individuals have shown the existence of supramodal brain regions able to process external information regardless of the sensory modality through which such an information has been acquired. This more abstract nature of functional cortical organization may enable congenitally blind individuals to acquire knowledge, form mental representations of and interact effectively with an external world that they have never seen., Summary: Altogether, findings from both behavioural and imaging studies indicate that the brain functional organization is to a large extent independent from visual experience and able to process information in a supramodal fashion. The study of the blind brain is a very powerful approach to understanding not only the cross-modal plastic, adaptative modifications that occur in the 'visual' regions but primarily the functional architecture of the human brain itself.

The motion-sensitive middle temporal cortex (hMT+ complex) responds also to non-visual motion stimulation conveyed through the tactile and auditory modalities, both in sighted and in congenitally blind individuals. This indicates that hMT+ is truly responsive to motion-related information regardless of visual experience and the sensory modality through which such information is carried to the brain. Here we determined whether the hMT+ complex responds to motion perception per se, that is, motion not perceived through the visual, haptic or aural modalities. Using functional magnetic resonance imaging (fMRI), we investigated brain responses in eight congenitally blind and nine sighted volunteers who had been trained to use the tongue display unit (TDU), a sensory substitution device which converts visual information into electrotactile pulses delivered to the tongue, to resolve a tactile motion discrimination task. Stimuli consisted of either static dots, dots moving coherently or dots moving in random directions. Both groups learned the task at the same rate and activated the hMT+ complex during tactile motion discrimination, although at different anatomical locations. Furthermore, the congenitally blind subjects showed additional activations within the dorsal extrastriate cortical pathway. These results extend previous data in support of the supramodal functional organization of hMT+ complex by showing that this cortical area processes motion-related information per se, that is, motion stimuli that are not visual in nature and that are administered to body structures that, in humans, are not primarily devoted to movement perception or spatial location, such as the tongue. In line with previous studies, the differential activations between sighted and congenitally blind individuals indicate that lack of vision leads to functional rearrangements of these supramodal cortical areas., (Copyright 2010 Elsevier Inc. All rights reserved.)

Observing and learning actions and behaviors from others, a mechanism crucial for survival and social interaction, engages the mirror neuron system. To determine whether vision is a necessary prerequisite for the human mirror system to develop and function, we used functional magnetic resonance imaging to compare brain activity in congenitally blind individuals during the auditory presentation of hand-executed actions or environmental sounds, and the motor pantomime of manipulation tasks, with that in sighted volunteers, who additionally performed a visual action recognition task. Congenitally blind individuals activated a premotor-temporoparietal cortical network in response to aurally presented actions that overlapped both with mirror system areas found in sighted subjects in response to visually and aurally presented stimuli, and with the brain response elicited by motor pantomime of the same actions. Furthermore, the mirror system cortex showed a significantly greater response to motor familiar than to unfamiliar action sounds in both sighted and blind individuals. Thus, the mirror system in humans can develop in the absence of sight. The results in blind individuals demonstrate that the sound of an action engages the mirror system for action schemas that have not been learned through the visual modality and that this activity is not mediated by visual imagery. These findings indicate that the mirror system is based on supramodal sensory representations of actions and, furthermore, that these abstract representations allow individuals with no visual experience to interact effectively with others.

The objective of this review is to examine and evaluate recent findings on cognitive functioning (in particular imagery processes) in individuals with congenital visual impairments, including total blindness, low-vision and monocular vision. As one might expect, the performance of blind individuals in many behaviours and tasks requiring imagery can be inferior to that of sighted subjects; however, surprisingly often this is not the case. Interestingly, there is evidence that the blind often employ different cognitive mechanisms than sighted subjects, suggesting that compensatory mechanisms can overcome the limitations of sight loss. Taken together, these studies suggest that the nature of perceptual input on which we commonly rely strongly affects the organization of our mental processes. We also review recent neuroimaging studies on the neural correlates of sensory perception and mental imagery in visually impaired individuals that have cast light on the plastic functional reorganization mechanisms associated with visual deprivation.

We investigated whether the visual hMT+ cortex plays a role in supramodal representation of sensory flow, not mediated by visual mental imagery. We used functional magnetic resonance imaging to measure neural activity in sighted and congenitally blind individuals during passive perception of optic and tactile flows. Visual motion-responsive cortex, including hMT+, was identified in the lateral occipital and inferior temporal cortices of the sighted subjects by response to optic flow. Tactile flow perception in sighted subjects activated the more anterior part of these cortical regions but deactivated the more posterior part. By contrast, perception of tactile flow in blind subjects activated the full extent, including the more posterior part. These results demonstrate that activation of hMT+ and surrounding cortex by tactile flow is not mediated by visual mental imagery and that the functional organization of hMT+ can develop to subserve tactile flow perception in the absence of any visual experience. Moreover, visual experience leads to a segregation of the motion-responsive occipitotemporal cortex into an anterior subregion involved in the representation of both optic and tactile flows and a posterior subregion that processes optic flow only.

Research in blind individuals has primarily focused for a long time on the brain plastic reorganization that occurs in early visual areas. Only more recently, scientists have developed innovative strategies to understand to what extent vision is truly a mandatory prerequisite for the brain's fine morphological architecture to develop and function. As a whole, the studies conducted to date in sighted and congenitally blind individuals have provided ample evidence that several "visual" cortical areas develop independently from visual experience and do process information content regardless of the sensory modality through which a particular stimulus is conveyed: a property named supramodality. At the same time, lack of vision leads to a structural and functional reorganization within "visual" brain areas, a phenomenon known as cross-modal plasticity. Cross-modal recruitment of the occipital cortex in visually deprived individuals represents an adaptative compensatory mechanism that mediates processing of non-visual inputs. Supramodality and cross-modal plasticity appears to be the "yin and yang" of brain development: supramodal is what takes place despite the lack of vision, whereas cross-modal is what happens because of lack of vision. Here we provide a critical overview of the research in this field and discuss the implications that these novel findings have for the development of educative/rehabilitation approaches and sensory substitution devices (SSDs) in sensory-impaired individuals. [ABSTRACT FROM AUTHOR]

The organization of semantic information in the brain has been mainly explored through category-based models, on the assumption that categories broadly reflect the organization of conceptual knowledge. However, the analysis of concepts as individual entities, rather than as items belonging to distinct superordinate categories, may represent a significant advancement in the comprehension of how conceptual knowledge is encoded in the human brain. Here, we studied the individual representation of thirty concrete nouns from six different categories, across different sensory modalities (i.e., auditory and visual) and groups (i.e., sighted and congenitally blind individuals) in a core hub of the semantic network, the left angular gyrus, and in its neighboring regions within the lateral parietal cortex. Four models based on either perceptual or semantic features at different levels of complexity (i.e., low- or high-level) were used to predict fMRI brain activity using representational similarity encoding analysis. When controlling for the superordinate component, high-level models based on semantic and shape information led to significant encoding accuracies in the intraparietal sulcus only. This region is involved in feature binding and combination of concepts across multiple sensory modalities, suggesting its role in high-level representation of conceptual knowledge. Moreover, when the information regarding superordinate categories is retained, a large extent of parietal cortex is engaged. This result indicates the need to control for the coarse-level categorial organization when performing studies on higher-level processes related to the retrieval of semantic information. [ABSTRACT FROM AUTHOR]

• The large-scale brain architecture develops even in the absence of visual experience. • The sensory-deprived brain maintains a lifelong sensitivity to environmental inputs. • The functional 'fate' of V1 in congenitally blind people is still disputed. • V1 may retain most of its intrinsic functioning also in congenitally blind individuals. [ABSTRACT FROM AUTHOR]