The Nature of Consciousness in the Visually Deprived Brain

Vision plays a central role in how we represent and interact with the world around us. The primacy of vision is structurally imbedded in cortical organization as about one-third of the cortical surface in primates is involved in visual processes. Consequently, the loss of vision, either at birth or later in life, affects brain organization and the way the world is perceived and acted upon. In this paper, we address a number of issues on the nature of consciousness in people deprived of vision. Do brains from sighted and blind individuals differ, and how? How does the brain of someone who has never had any visual perception form an image of the external world? What is the subjective correlate of activity in the visual cortex of a subject who has never seen in life? More in general, what can we learn about the functional development of the human brain in physiological conditions by studying blindness? We discuss findings from animal research as well from recent psychophysical and functional brain imaging studies in sighted and blind individuals that shed some new light on the answers to these questions

Neuroplasticity, neural reuse, and the language module

What conception of mental architecture can survive the evidence of neuroplasticity and neural reuse in the human brain? In particular, what sorts of modules are compatible with this evidence? I aim to show how developmental and adult neuroplasticity, as well as evidence of pervasive neural reuse, forces us to revise the standard conception of modularity and spells the end of a hardwired and dedicated language module. I argue from principles of both neural reuse and neural redundancy that language is facilitated by a composite of modules (or module-like entities), few if any of which are likely to be linguistically special, and that neuroplasticity provides evidence that (in key respects and to an appreciable extent) few if any of them ought to be considered developmentally robust, though their development does seem to be constrained by features intrinsic to particular regions of cortex (manifesting as domain-specific predispositions or acquisition biases). In the course of doing so I articulate a schematically and neurobiologically precise framework for understanding modules and their supramodular interactions

How input modality and visual experience affect the representation of categories in the brain

The general aim of the present dissertation was to participate in the progress of our understanding of how sensory input and sensory experience impact on how the human brain implements categorical knowledge. The goal was twofold: (1) understand whether there are brain regions that encode information about different categories regardless of input modality and sensory experience (study 1); (2) deepen the investigation of the mechanisms that drive cross-modal and intra-modal plasticity following early blindness and the way they express during the processing of different categories presented as real-world sounds (study 2). To address these fundamental questions, we used fMRI to characterize the brain responses to different conceptual categories presented acoustically in sighted and early blind individuals, and visually in a separate sighted group. In study 1, we observed that the right posterior middle temporal gyrus (rpMTG) is the region that most reliably decoded categories and selectively correlated with conceptual models of our stimuli space independently of input modality and visual experience. However, this region maintains separate the representational format from the different modalities, revealing a multimodal rather than an amodal nature. In addition, we observed that VOTC showed distinct functional profiles according to the hemispheric side. The left VOTC showed an involvement in the acoustical categorization processing at the same degree in sighted and in blind individuals. We propose that this involvement might reflect an engagement of the left VOTC in more semantic/linguistic processing of the stimuli potentially supported by its enhanced connection with the language system. However, paralleling our observation in rpMTG, the representations from different modalities are maintained segregated in VOTC, showing little evidence for sensory-abstraction. On the other side, the right VOTC emerged as a sensory-related visual region in sighted with the ability to rewires itself toward acoustical stimulation in case of early visual deprivation. In study 2, we observed opposite effects of early visual deprivation on auditory decoding in occipital and temporal regions. While occipital regions contained more information about sound categories in the blind, the temporal cortex showed higher decoding in the sighted. This unbalance effect was stronger in the right hemisphere where we, also, observed a negative correlation between occipital and temporal decoding of sound categories in EB. These last results suggest that the intramodal and crossmodal reorganizations might be inter-connected. We therefore propose that the extension of non-visual functions in the occipital cortex of EB may trigger a network-level reorganization that reduce the computational load of the regions typically coding for the remaining senses due to the extension of such computation in occipital regions

Neural and socio-cognitive sequelae of congenital visual impairment during midchildhood

Previous studies identified cognitive difficulties in individuals with congenital visual impairment that significantly impacted on wellbeing and educational attainment. However, factors leading to adverse outcome remained unclear. The current study aimed to identify associations and mechanisms of specific cognitive deficits associated with visual impairment from a neurodevelopmental perspective. Based on recent theoretical advances (Johnson, 2011), it was assumed that visual impairment leads to differences in cognition by influencing experience-driven brain maturational processes, which support the integration between cortical areas to support cognitive processes. In order to explore this hypothesis, children with visual impairment due to disorders that were thought to only affect peripheral sensory parts of the visual system were assessed on neuropsychological instruments covering a range of functional domains. Further, structural and functional neurophysiological methods were employed to establish the impact of visual impairment on brain organisation. The results of the present work confirm earlier reports of specific deficits in spatial memory, social understanding, and aspects of executive function, despite typical performance in other domains. In addition, the current study is the first study to indicate dosage-dependence with some aspects of social communication being even impaired in children with only mild to moderate visual impairment, while aspects of executive function and spatial memory were only found to be deficient in children with more severe forms of visual impairment. Further, neurophysiological investigations indicated differences in structural and functional brain organisation in children with VI that related to differences in behavioural performance. In general, the results of the present study suggest that visual sensory impairment impacts on brain and cognitive development with important implications for education and clinical treatment of children with visual impairment

Mechanisms of top-down visual spatial attention: computational and behavioral investigations

This thesis examines the mechanisms underlying visual spatial attention. In particular I focused on top-­‐down or voluntary attention, namely the ability to select relevant information and discard the irrelevant according to our goals. Given the limited processing resources of the human brain, which does not allow to process all the available information to the same degree, the ability to correctly allocate processing resources is fundamental for the accomplishment of most everyday tasks. The cost of misoriented attention is that we could miss some relevant information, with potentially serious consequences. In the first study (chapter 2) I will address the issue of the neural substrates of visual spatial attention: what are the neural mechanisms that allow the deployment of visual spatial attention? According to the premotor theory orienting attention to a location in space is equivalent to planning an eye movement to the same location, an idea strongly supported by neuroimaging and neurophysiological evidence. Accordingly, in this study I will present a model that can account for several attentional effects without requiring additional mechanisms separate from the circuits that perform sensorimotor transformations for eye movements. Moreover, it includes a mechanism that allows, within the framework of the premotor theory, to explain dissociations between attention and eye movements that may be invoked to disprove it. In the second model presented (chapter 3) I will further investigate the computational mechanisms underlying sensorimotor transformations. Specifically I will show that a representation in which the amplitude of visual responses is modulated by postural signal is both efficient and plausible, emerging also in a neural network model trained through unsupervised learning (i.e., using only signals locally available at the neuron level). Ultimately this result gives additional support to the approach adopted in the first model. Next, I will present a series of behavioral studies: in the first (chapter 4) I will show that spatial constancy of attention (i.e., the ability to sustain attention at a spatial location across eye movements) is dependent on some properties of the image, namely the presence of continuous visual landmarks at the attended locations. Importantly, this finding helps resolve contrasts between several recent results. In the second behavioral study (chapter 5), I will investigate an often neglected aspect of spatial cueing paradigms, probably the most widely used technique in studies of covert attention: the role of cue predictivity (i.e. the extent to which the spatial cue correctly indicates the location where the target stimulus will appear). Results show that, independently of participant’s awareness, changes  in predictivity result in changes in spatial validity effects, and that reliable shifts of attention can take place also in the absence of a predictive cue. In sum the results question the appropriateness of using predictive cues for delineating pure voluntary shifts of spatial attention. Finally, in the last study I will use a psychophysiological measure, the diameter of the eye’s pupil, to investigate intensive aspects of attention. Event-­‐related pupil dilations accurately mirrored changes in visuospatial awareness induced by a dual-­‐task manipulation that consumed attentional resources. Moreover, results of the primary spatial monitoring task revealed a significant rightward bias, indicated by a greater proportion of missed targets in the left hemifield. Interestingly this result mimics the extinction to double simultaneous stimulation (i.e., the failure to respond to a stimulus when it is presented simultaneously with another stimulus) which is often found in patients with unilateral brain damage. Overall, these studies present an emerging picture of attention as a complex mechanism that even in its volitional aspects is modulated by other non-­‐volitional factors, both external and internal to the individua

Complexity, the auditory system, and perceptual learning in naïve users of a visual-to-auditory sensory substitution device.

PhDSensory substitution devices are a non-invasive visual prostheses that use sound or touch to aid functioning in the blind. Algorithms informed by natural crossmodal correspondences convert and transmit sensory information attributed to an impaired modality back to the user via an unimpaired modality and utilise multisensory networks to activate visual areas of cortex. While behavioural success has been demonstrated in non-visual tasks suing SSDs how they utilise a metamodal brain, organised for function is still a question in research. While imaging studies have shown activation of visual cortex in trained users it is likely that naïve users rely on auditory characteristics of the output signal for functionality and that it is perceptual learning that facilitates crossmodal plasticity. In this thesis I investigated visual-to-auditory sensory substitution in naïve sighted users to assess whether signal complexity and processing in the auditory system facilitates and limits simple recognition tasks. In four experiments evaluating; signal complexity, object resolution, harmonic interference and information load I demonstrate above chance performance in naïve users in all tasks, an increase in generalized learning, limitations in recognition due to principles of auditory scene analysis and capacity limits that hinder performance. Results are looked at from both theoretical and applied perspectives with solutions designed to further inform theory on a multisensory perceptual brain and provide effective training to aid visual rehabilitation.Queen Mary University of Londo

The influence of attentional focus on neuroplasticity following a seven-day balance training intervention

It is well established that focusing on the external effect of one’s movement (an external focus of attention) results in enhanced motor learning and produces superior motor performance compared to focusing inward on the body’s own physical execution of the motor movement (an internal focus). While the benefits of an external focus in motor learning, and the detriments of an inward or ‘internal’ focus have been highly replicated, there is still little mechanistic understanding pertaining to the brain-related changes that may result from these two different foci of attention during motor training. Since the brain is highly malleable and has been shown to adapt in response to motor training (i.e., neuroplasticity), it is postulated that attentional focus may change the brain’s structure and function. However, no direct examination exploring the influence of attentional focus on neuroplasticity (structural or functional) exists. The primary objective of this study was to determine the effects of balance training with different attentional foci on brain-related neuroplasticity in a young healthy population. Participants (n = 33) were randomly assigned to a control, internal focus, or external focus condition. Functional and structural brain connectivity analyses was conducted using neuroimaging data collected through functional magnetic resonance imaging (fMRI) and diffusion tensor imaging (DTI) prior to (baseline) and following a seven-day balance training intervention (retention). Between baseline and retention data collection, participants in the internal and external focus training groups practiced a dynamic balance task for one hour per day, each day for seven consecutive days (acquisition). For the internal focus trials, participants were asked to, ‘focus on keeping their feet level;’ whereas, for the external focus trials participants were asked to, ‘focus on keeping the board level.’ The control group did not complete any balance training, but completed baseline and retention balance measurements. An inertial measurement unit was attached to the center of the balance board to assess the performance and learning of the balance task. Resting-state brain connectivity analyses were performed on the fMRI data to contrast connectivity differences for each group at retention relative to baseline, and, for the diffusion data (DTI), fractional anisotropy analyses (a metric to quantify water diffusion within a voxel of white-matter) was performed to quantify the relationship between changes in balance and water diffusivity within white-matter tracts. Classical attentional focus effects were observed for acquisition, with those in the external focus condition producing significantly less mean and standard deviation velocity compared to the internal focus group (both p .05). These results suggest that a seven-day balance training program with attentional focus in a young healthy population influences brain function (specifically correlated activity at rest), but longer training programs or more rest may be needed to influence brain structure (as measured by fractional anisotropy). These findings have important implications for a variety of clinical populations who show altered resting-sate connectivity and deteriorations in balance control (e.g., Alzheimer’s disease, stroke survivors). Seven days of balance training with an external focus may be useful in improving balance control and may influence correlated brain activity at rest, but longer training programs or more rest may be needed to influence brain structure. We discuss these findings in the context of the constrained-action hypothesis and OPTIMAL theory

Diagnostic Palpation in Osteopathic Medicine: A Putative Neurocognitive Model of Expertise

This thesis examines the extent to which the development of expertise in diagnostic palpation in osteopathic medicine is associated with changes in cognitive processing. Chapter 2 and Chapter 3 review, respectively, the literature on the role of analytical and non-analytical processing in osteopathic and medical clinical decision making; and the relevant research on the use of vision and haptics and the development of expertise within the context of an osteopathic clinical examination. The two studies reported in Chapter 4 examined the mental representation of knowledge and the role of analogical reasoning in osteopathic clinical decision making. The results reported there demonstrate that the development of expertise in osteopathic medicine is associated with the processes of knowledge encapsulation and script formation. The four studies reported in Chapters 5 and 6 investigate the way in which expert osteopaths use their visual and haptic systems in the diagnosis of somatic dysfunction. The results suggest that ongoing clinical practice enables osteopaths to combine visual and haptic sensory signals in a more efficient manner. Such visuo-haptic sensory integration is likely to be facilitated by top-down processing associated with visual, tactile, and kinaesthetic mental imagery. Taken together, the results of the six studies reported in this thesis indicate that the development of expertise in diagnostic palpation in osteopathic medicine is associated with changes in cognitive processing. Whereas the experts’ diagnostic judgments are heavily influenced by top-down, non-analytical processing; students rely, primarily, on bottom-up sensory processing from vision and haptics. Ongoing training and clinical practice are likely to lead to changes in the clinician’s neurocognitive architecture. This thesis proposes an original model of expertise in diagnostic palpation which has implications for osteopathic education. Students and clinicians should be encouraged to appraise the reliability of different sensory cues in the context of clinical examination, combine sensory data from different channels, and consider using both analytical and nonanalytical reasoning in their decision making. Importantly, they should develop their skills of criticality and their ability to reflect on, and analyse their practice experiences in and on action

Investigating structural plasticity in musicians’ brains using structural magnetic resonance and diffusion tensor imaging techniques

Neuroplasticity is the ability of the brain to change its structure and/or function in response to environmental stimuli. It is implicated in many processes, such as learning, maturation, skill acquisition, and rehabilitation following brain injury. With the advent of neuroimaging techniques, the study of neuroplasticity and its mechanisms have fascinated researchers given the wide scope with which this process is involved. Musicians have long been considered an ideal model to study neuroplasticity in humans. It has been shown that musicians with their early, intensive, and multimodal skilful practice have structural plasticity in different brain regions. The objective of this work was to extend these structural studies through examining different cohorts of musicians, using a multitude of imaging and morphometric techniques, and performing novel examinations of brain regions essential for enabling high level musical performance such as Broca’s area, corpus callosum (CC), and cerebellum. Three age-, gender- and handedness-matched cohorts were examined. The first cohort included 26 orchestral musicians and 26 non-musicians. High resolution T1-weighted structural MR images were acquired to measure gray and white matter volumes and cortical surface area of Broca’s area subparts: pars opercularis/BA44 and pars triangularis/BA45. The second cohort included 12/12/12 professional musicians/amateur musicians/non-musicians. High resolution T1-weighted MR images were acquired to measure cross-sectional areas of four regions of the midsagittal CC: CC1 (rostrum/ genu/anterior body), CC2 (anterior midbody), CC3 (posterior midbody), and CC4 (isthmus and splenium). In the third cohort, 12/12 musicians and non-musicians were examined. High resolution T1-weighted structural MR images were acquired to measure cross-sectional areas of CC1-CC4 regions; and diffusion tensor imaging-based tractography was used to measure average fractional anisotropy (FA), mean diffusivity (MD), tract volume, and number of streamlines of the same regions. In a subset (10/10) of this cohort, high resolution structural scans were used to measure gray and white matter volumes of cerebellar hemispheres; and diffusion tensor imaging-based tractography was used to measure average FA, tract volume, and number of streamlines of superior (SCP) and middle (MCP) cerebellar peduncles. Outcome measures were compared between groups. Compared to controls, musicians possessed greater gray matter volume and cortical surface area of left pars opercularis/BA44 in the first cohort. The volume of left pars opercularis was positively correlated with years of musical performance. Professional musicians possessed greater cross-sectional area of CC1 and CC4 regions compared to amateurs and non-musicians in the second cohort. In the third cohort, musicians possessed greater cross-sectional area, average FA/tract volume/number of streamlines, and lower MD in CC4 region. There was a negative correlation between cross-sectional area of CC4 region and age of starting musical training. There was a positive correlation between average FA values and cross-sectional area of CC4 region in all subjects. In addition, musicians had increased white matter volume of the right cerebellar hemisphere, increased tract volume and number of streamlines of right SCP, and tract volume of right MCP. I hypothesize that these findings represent use-dependent structural plasticity imposed by musical performance. At the microscopic level, these macroanatomical changes may reflect increased synaptogenesis and dendritic growth, generation of new axon collaterals, and formation of new neurons, which would support enhanced functional demands on musicians’ brains