"Love looks not with the eyes": supranormal processing of emotional speech in individuals with late-blindness versus preserved processing in individuals with congenital-blindness.

Processing of emotional speech in the absence of visual information relies on two auditory channels: semantics and prosody. No study to date has investigated how blindness impacts this process. Two theories, Perceptual Deficit, and Sensory Compensation, yiled different expectations about the role of visual experience (or its lack thereof) in processing emotional speech. To test the effect of vision and early visual experience on processing of emotional speech, we compared individuals with congenital blindness (CB, n = 17), individuals with late blindness (LB, n = 15), and sighted controls (SC, n = 21) on identification and selective-attention of semantic and prosodic spoken-emotions. Results showed that individuals with blindness performed at least as well as SC, supporting Sensory Compensation and the role of cortical reorganisation. Individuals with LB outperformed individuals with CB, in accordance with Perceptual Deficit, supporting the role of early visual experience. The LB advantage was moderated by executive functions (working-memory). Namely, the advantage was erased for individuals with CB who showed higher levels of executive functions. Results suggest that vision is not necessary for processing of emotional speech, but early visual experience could improve it. The findings support a combination of the two aforementioned theories and reject a dichotomous view of deficiencies/enhancements of blindness.

Activation of human visual area V6 during egocentric navigation with and without visual experience.

V6 is a retinotopic area located in the dorsal visual stream that integrates eye movements with retinal and visuo-motor signals. Despite the known role of V6 in visual motion, it is unknown whether it is involved in navigation and how sensory experiences shape its functional properties. We explored the involvement of V6 in egocentric navigation in sighted and in congenitally blind (CB) participants navigating via an in-house distance-to-sound sensory substitution device (SSD), the EyeCane. We performed two fMRI experiments on two independent datasets. In the first experiment, CB and sighted participants navigated the same mazes. The sighted performed the mazes via vision, while the CB performed them via audition. The CB performed the mazes before and after a training session, using the EyeCane SSD. In the second experiment, a group of sighted participants performed a motor topography task. Our results show that right V6 (rhV6) is selectively involved in egocentric navigation independently of the sensory modality used. Indeed, after training, rhV6 of CB is selectively recruited for auditory navigation, similarly to rhV6 in the sighted. Moreover, we found activation for body movement in area V6, which can putatively contribute to its involvement in egocentric navigation. Taken together, our findings suggest that area rhV6 is a unique hub that transforms spatially relevant sensory information into an egocentric representation for navigation. While vision is clearly the dominant modality, rhV6 is in fact a supramodal area that can develop its selectivity for navigation in the absence of visual experience.

Neural substrates of spatial processing and navigation in blindness: An activation likelihood estimation meta-analysis.

Even though vision is considered the best suited sensory modality to acquire spatial information, blind individuals can form spatial representations to navigate and orient themselves efficiently in space. Consequently, many studies support the amodality hypothesis of spatial representations since sensory modalities other than vision contribute to the formation of spatial representations, independently of visual experience and imagery. However, given the high variability in abilities and deficits observed in blind populations, a clear consensus about the neural representations of space has yet to be established. To this end, we performed a meta-analysis of the literature on the neural correlates of spatial processing and navigation via sensory modalities other than vision, like touch and audition, in individuals with early and late onset blindness. An activation likelihood estimation (ALE) analysis of the neuroimaging literature revealed that early blind individuals and sighted controls activate the same neural networks in the processing of non-visual spatial information and navigation, including the posterior parietal cortex, frontal eye fields, insula, and the hippocampal complex. Furthermore, blind individuals also recruit primary and associative occipital areas involved in visuo-spatial processing via cross-modal plasticity mechanisms. The scarcity of studies involving late blind individuals did not allow us to establish a clear consensus about the neural substrates of spatial representations in this specific population. In conclusion, the results of our analysis on neuroimaging studies involving early blind individuals support the amodality hypothesis of spatial representations.

Gyrification in relation to cortical thickness in the congenitally blind.

Greater cortical gyrification (GY) is linked with enhanced cognitive abilities and is also negatively related to cortical thickness (CT). Individuals who are congenitally blind (CB) exhibits remarkable functional brain plasticity which enables them to perform certain non-visual and cognitive tasks with supranormal abilities. For instance, extensive training using touch and audition enables CB people to develop impressive skills and there is evidence linking these skills to cross-modal activations of primary visual areas. There is a cascade of anatomical, morphometric and functional-connectivity changes in non-visual structures, volumetric reductions in several components of the visual system, and CT is also increased in CB. No study to date has explored GY changes in this population, and no study has explored how variations in CT are related to GY changes in CB. T1-weighted 3D structural magnetic resonance imaging scans were acquired to examine the effects of congenital visual deprivation in cortical structures in a healthy sample of 11 CB individuals (6 male) and 16 age-matched sighted controls (SC) (10 male). In this report, we show for the first time an increase in GY in several brain areas of CB individuals compared to SC, and a negative relationship between GY and CT in the CB brain in several different cortical areas. We discuss the implications of our findings and the contributions of developmental factors and synaptogenesis to the relationship between CT and GY in CB individuals compared to SC. F.

Deductive Reasoning and Working Memory Skills in Individuals with Blindness.

Deductive reasoning and working memory are integral parts of executive functioning and are important skills for blind people in everyday life. Despite the importance of these skills, the influence of visual experience on reasoning and working memory skills, as well as on the relationship between these, is unknown. In this study, fifteen participants with congenital blindness (CB), fifteen with late blindness (LB), fifteen sighted blindfolded controls (SbfC), and fifteen sighted participants performed two tasks of deductive reasoning and two of working memory. We found that while the CB and LB participants did not differ in their deductive reasoning abilities, the CB group performed worse than the sighted controls, and the LB group performed better than the SbfC group. Those with CB outperformed all the other groups in both of the working memory tests. Working memory is associated with deductive reasoning in all three visually impaired groups, but not in the sighted group. These findings suggest that deductive reasoning is not a uniform skill, and that it is associated with visual impairment onset, the level of reasoning difficulty, and the degree of working memory load.

Brain-Machine Interfaces to Assist the Blind.

The loss or absence of vision is probably one of the most incapacitating events that can befall a human being. The importance of vision for humans is also reflected in brain anatomy as approximately one third of the human brain is devoted to vision. It is therefore unsurprising that throughout history many attempts have been undertaken to develop devices aiming at substituting for a missing visual capacity. In this review, we present two concepts that have been prevalent over the last two decades. The first concept is sensory substitution, which refers to the use of another sensory modality to perform a task that is normally primarily sub-served by the lost sense. The second concept is cross-modal plasticity, which occurs when loss of input in one sensory modality leads to reorganization in brain representation of other sensory modalities. Both phenomena are training-dependent. We also briefly describe the history of blindness from ancient times to modernity, and then proceed to address the means that have been used to help blind individuals, with an emphasis on modern technologies, invasive (various type of surgical implants) and non-invasive devices. With the advent of brain imaging, it has become possible to peer into the neural substrates of sensory substitution and highlight the magnitude of the plastic processes that lead to a rewired brain. Finally, we will address the important question of the value and practicality of the available technologies and future directions.

Spatial Competence and Brain Plasticity in Congenital Blindness via Sensory Substitution Devices.

In congenital blindness (CB), tactile, and auditory information can be reinterpreted by the brain to compensate for visual information through mechanisms of brain plasticity triggered by training. Visual deprivation does not cause a cognitive spatial deficit since blind people are able to acquire spatial knowledge about the environment. However, this spatial competence takes longer to achieve but is eventually reached through training-induced plasticity. Congenitally blind individuals can further improve their spatial skills with the extensive use of sensory substitution devices (SSDs), either visual-to-tactile or visual-to-auditory. Using a combination of functional and anatomical neuroimaging techniques, our recent work has demonstrated the impact of spatial training with both visual to tactile and visual to auditory SSDs on brain plasticity, cortical processing, and the achievement of certain forms of spatial competence. The comparison of performances between CB and sighted people using several different sensory substitution devices in perceptual and sensory-motor tasks uncovered the striking ability of the brain to rewire itself during perceptual learning and to interpret novel sensory information even during adulthood. We discuss here the implications of these findings for helping blind people in navigation tasks and to increase their accessibility to both real and virtual environments.

Neural Networks Mediating Perceptual Learning in Congenital Blindness.

Despite the fact that complete visual deprivation leads to volumetric reductions in brain structures associated with spatial learning, blind individuals are still able to navigate. The neural structures involved in this function are not fully understood. Our study aims to correlate the performance of congenitally blind individuals (CB) and blindfolded sighted controls (SC) in a life-size obstacle-course using a visual-to-tactile sensory substitution device, with the size of brain structures (voxel based morphometry-VBM-) measured through structural magnetic resonance Imaging (MRI). VBM was used to extract grey matter volumes within several a-priori defined brain regions in all participants. Principal component analysis was utilized to group brain regions in factors and orthogonalize brain volumes. Regression analyses were then performed to link learning abilities to these factors. We found that (1) both CB and SC were able to learn to detect and avoid obstacles; (2) their learning rates for obstacle detection and avoidance correlated significantly with the volume of brain structures known to be involved in spatial skills. There is a similar relation between regions of the dorsal stream network and avoidance for both SC and CB whereas for detection, SC rely more on medial temporal lobe structures and CB on sensorimotor areas.

Spatial navigation by congenitally blind individuals.

Spatial navigation in the absence of vision has been investigated from a variety of perspectives and disciplines. These different approaches have progressed our understanding of spatial knowledge acquisition by blind individuals, including their abilities, strategies, and corresponding mental representations. In this review, we propose a framework for investigating differences in spatial knowledge acquisition by blind and sighted people consisting of three longitudinal models (i.e., convergent, cumulative, and persistent). Recent advances in neuroscience and technological devices have provided novel insights into the different neural mechanisms underlying spatial navigation by blind and sighted people and the potential for functional reorganization. Despite these advances, there is still a lack of consensus regarding the extent to which locomotion and wayfinding depend on amodal spatial representations. This challenge largely stems from methodological limitations such as heterogeneity in the blind population and terminological ambiguity related to the concept of cognitive maps. Coupled with an over-reliance on potential technological solutions, the field has diffused into theoretical and applied branches that do not always communicate. Here, we review research on navigation by congenitally blind individuals with an emphasis on behavioral and neuroscientific evidence, as well as the potential of technological assistance. Throughout the article, we emphasize the need to disentangle strategy choice and performance when discussing the navigation abilities of the blind population. For further resources related to this article, please visit the WIREs website.

Navigation using sensory substitution in real and virtual mazes.

Under certain specific conditions people who are blind have a perception of space that is equivalent to that of sighted individuals. However, in most cases their spatial perception is impaired. Is this simply due to their current lack of access to visual information or does the lack of visual information throughout development prevent the proper integration of the neural systems underlying spatial cognition? Sensory Substitution devices (SSDs) can transfer visual information via other senses and provide a unique tool to examine this question. We hypothesize that the use of our SSD (The EyeCane: a device that translates distance information into sounds and vibrations) can enable blind people to attain a similar performance level as the sighted in a spatial navigation task. We gave fifty-six participants training with the EyeCane. They navigated in real life-size mazes using the EyeCane SSD and in virtual renditions of the same mazes using a virtual-EyeCane. The participants were divided into four groups according to visual experience: congenitally blind, low vision & late blind, blindfolded sighted and sighted visual controls. We found that with the EyeCane participants made fewer errors in the maze, had fewer collisions, and completed the maze in less time on the last session compared to the first. By the third session, participants improved to the point where individual trials were no longer significantly different from the initial performance of the sighted visual group in terms of errors, time and collision.

The "EyeCane", a new electronic travel aid for the blind: Technology, behavior & swift learning.

PURPOSE: Independent mobility is one of the most pressing problems facing people who are blind. We present the EyeCane, a new mobility aid aimed at increasing perception of environment beyond what is provided by the traditional White Cane for tasks such as distance estimation, navigation and obstacle detection. METHODS: The "EyeCane" enhances the traditional White Cane by using tactile and auditory output to increase detectable distance and angles. It circumvents the technical pitfalls of other devices, such as weight, short battery life, complex interface schemes, and slow learning curve. It implements multiple beams to enables detection of obstacles at different heights, and narrow beams to provide active sensing that can potentially increase the user's spatial perception of the environment. Participants were tasked with using the EyeCane for several basic tasks with minimal training. RESULTS: Blind and blindfolded-sighted participants were able to use the EyeCane successfully for distance estimation, simple navigation and simple obstacle detection after only several minutes of training. CONCLUSIONS: These results demonstrate the EyeCane's potential for mobility rehabilitation. The short training time is especially important since available mobility training resources are limited, not always available, and can be quite expensive and/or entail long waiting periods.

The effect of extended sensory range via the EyeCane sensory substitution device on the characteristics of visionless virtual navigation.

Mobility training programs for helping the blind navigate through unknown places with a White-Cane significantly improve their mobility. However, what is the effect of new assistive technologies, offering more information to the blind user, on the underlying premises of these programs such as navigation patterns? We developed the virtual-EyeCane, a minimalistic sensory substitution device translating single-point-distance into auditory cues identical to the EyeCane's in the real world. We compared performance in virtual environments when using the virtual-EyeCane, a virtual-White-Cane, no device and visual navigation. We show that the characteristics of virtual-EyeCane navigation differ from navigation with a virtual-White-Cane or no device, and that virtual-EyeCane users complete more levels successfully, taking shorter paths and with less collisions than these groups, and we demonstrate the relative similarity of virtual-EyeCane and visual navigation patterns. This suggests that additional distance information indeed changes navigation patterns from virtual-White-Cane use, and brings them closer to visual navigation.

Increasing accessibility to the blind of virtual environments, using a virtual mobility aid based on the "EyeCane": feasibility study.

Virtual worlds and environments are becoming an increasingly central part of our lives, yet they are still far from accessible to the blind. This is especially unfortunate as such environments hold great potential for them for uses such as social interaction, online education and especially for use with familiarizing the visually impaired user with a real environment virtually from the comfort and safety of his own home before visiting it in the real world. We have implemented a simple algorithm to improve this situation using single-point depth information, enabling the blind to use a virtual cane, modeled on the "EyeCane" electronic travel aid, within any virtual environment with minimal pre-processing. Use of the Virtual-EyeCane, enables this experience to potentially be later used in real world environments with identical stimuli to those from the virtual environment. We show the fast-learned practical use of this algorithm for navigation in simple environments.

Navigation with a sensory substitution device in congenitally blind individuals.

Vision allows for obstacle detection and avoidance. The compensatory mechanisms involved in maintaining these functions in blind people using their remaining intact senses are poorly understood. We investigated the ability of congenitally blind participants to detect and avoid obstacles using the tongue display unit, a sensory substitution device that uses the tongue as a portal to the brain. We found that congenitally blind were better than sighted control participants in detecting and avoiding obstacles using the tongue display unit. Obstacles size and avoidance strategy had a significant effect on performance: large obstacles were better detected than small ones and step-around obstacles were better avoided than step-over ones. These data extend our earlier findings that when using a sensory substitution device, blind participants outperform sighted controls not only in a virtual navigation task but also during effective navigation within a human-sized obstacle course.

Tactile-'visual' acuity of the tongue in early blind individuals.

This study compares the 'tactile-visual' acuity of the tongue for 15 early blind participants with that of 24 age-matched and sex-matched sighted controls. Snellen's tumbling E test was used to assess 'visual' acuity using the tongue display unit. The tongue display unit is a sensory substitution device that converts a visual stimulus grabbed by a camera into electro-tactile pulses delivered to the tongue via a grid made out of electrodes. No overall significant difference was found in thresholds between early blind (1/206) and sighted control (1/237) participants. We found, however, a larger proportion of early blind in the two highest visual acuity categories (1/150 and 1/90). These results extend earlier findings that it is possible to measure visual acuity in the blind individuals using the tongue. Moreover, our data demonstrate that a subgroup of early blind participants is more efficient than controls in conveying visual information through the tongue.

Alterations in right posterior hippocampus in early blind individuals.

This study compares hippocampal volumes of early blind and sex/age-matched sighted controls through volumetric and localization analyses. Early blind individuals showed a significantly smaller right posterior hippocampus compared with controls. No differences in total hippocampal volumes were found between groups and there were no within-group differences for left versus right hippocampus. Sex, age and total brain grey matter volume had no effect on hippocampal volumes. Although extensive navigational training results in structural enhancement of the hippocampus for the sighted, the reduction of the posterior hippocampus in early blind individuals suggests the implication of this region in visual spatial memory.

Development of the commissure of the superior colliculus in the hamster.

The development of the corpus callosum (CC) and the anterior commissure (CA) is well known in a wide variety of species. No study, however, has described the development of the commissure of the superior colliculus (CSC) from embryonic state to adulthood in mammals. In this study, by using the lipophylic tracer DiI, we investigated the ontogeny of this mesencephalic commissure in the hamster at various ages. The development of axonal terminals, growth cone morphologies, and axons branching were described for the superior colliculus (SC) contralateral to the tracer injection. The first CSC axons cross the midline at embryonic day 11 (E-11) and grow further into the intermediate layers of the contralateral SC between E-12 and E-14. There is little axon growth therein between E-14 and the day of birth (P-0). Growth cones at the tip of these axons adopt complex morphologies at E-12 and progressively simplify until P-0. Pioneer axons are clearly visible between E-14 and P-1. These are followed by other axons progressively more numerous between P-0 and P-5. Axons do not show any branching until P-2. Between P-3 and P-9, the axons progressively arborize in the intermediate layers. Some axons reach the superficial layers at P-5, and they become more numerous around P-11, and only a few axons remain therein by P-21. Myelinated axons appear at P11 and are very dense at P-21. Our results indicate that the CSC follows developmental schemes similar to those of the CC and the AC but that initial axon midline crossing occurs earlier.