More than skin deep: body representation beyond primary somatosensory cortex

The neural circuits underlying initial sensory processing of somatic information are relatively well understood. In contrast, the processes that go beyond primary somatosensation to create more abstract representations related to the body are less clear. In this review, we focus on two classes of higher-order processing beyond somatosensation. Somatoperception refers to the process of perceiving the body itself, and particularly of ensuring somatic perceptual constancy. We review three key elements of somatoperception: (a) remapping information from the body surface into an egocentric reference frame (b) exteroceptive perception of objects in the external world through their contact with the body and (c) interoceptive percepts about the nature and state of the body itself. Somatorepresentation, in contrast, refers to the essentially cognitive process of constructing semantic knowledge and attitudes about the body, including: (d) lexical-semantic knowledge about bodies generally and one’s own body specifically, (e) configural knowledge about the structure of bodies, (f) emotions and attitudes directed towards one’s own body, and (g) the link between physical body and psychological self. We review a wide range of neuropsychological, neuroimaging and neurophysiological data to explore the dissociation between these different aspects of higher somatosensory function

Sensory augmentation and the tactile sublime

This paper responds to recent developments in the field of sensory augmentation by analysing several technological devices that augment the sensory apparatus using the tactile sense. First, I will define the term sensory augmentation, as the use of technological modification to enhance the sensory apparatus, and elaborate on the preconditions for successful tactile sensory augmentation. These are the adaptability of the brain to unfamiliar sensory input and the specific qualities of the skin lending themselves to be used for the perception of additional sensory information. Two devices, Moon Ribas’ Seismic Sense and David Eagleman’s vest, will then be discussed as potential facilitators of aesthetic experiences in virtue of the tactile sensory augmentation that these devices allow. I will connect the experiences afforded by these devices to the Kantian categories of the mathematical and the dynamical sublime, and to existing accounts of tactile sublimity. Essentially, the objects these devices make sensible, earthquakes for the Seismic Sense and digital information for the vest, produce pleasurable feelings of potential danger, awe, and respect. The subsequent acclimation to this new way of sensing and the aim to comprehend its sensed object are then discussed as possible objections to the interpretation of these experiences as sublime, and as aesthetic in general. To exemplify these issues and concretise my thesis of tactile sensory augmentation as a trigger of the sublime, I will outline an experiment to use the vest as an aid for faster decision making on the stock market

The Speed, Precision and Accuracy of Human Multisensory Perception following Changes to the Visual Sense

Human adults can combine information from multiple senses to improve their perceptual judgments. Visual and multisensory experience plays an important role in the development of multisensory integration, however it is unclear to what extent changes in vision impact multisensory processing later in life. In particular, it is not known whether adults account for changes to the relative reliability of their senses, following sensory loss, treatment or training. Using psychophysical methods, this thesis studied the multisensory processing of individuals experiencing changes to the visual sense. Chapters 2 and 3 assessed whether patients implanted with a retinal prosthesis (having been blinded by a retinal degenerative disease) could use this new visual signal with non-visual information to improve their speed or precision on multisensory tasks. Due to large differences between the reliabilities of the visual and non-visual cues, patients were not always able to benefit from the new visual signal. Chapter 4 assessed whether patients with degenerative visual loss adjust the weight given to visual and non-visual cues during audio-visual localization as their relative reliabilities change. Although some patients adjusted their reliance on vision across the visual field in line with predictions based on cue relative reliability, others - patients with visual loss limited to their central visual field only - did not. Chapter 5 assessed whether training with either more reliable or less reliable visual feedback could enable normally sighted adults to overcome an auditory localization bias. Findings suggest that visual information, irrespective of reliability, can be used to overcome at least some non-visual biases. In summary, this thesis documents multisensory changes following changes to the visual sense. The results improve our understanding of adult multisensory plasticity and have implications for successful treatments and rehabilitation following sensory loss

Embodied Precision : Intranasal Oxytocin Modulates Multisensory Integration

© 2018 Massachusetts Institute of Technology.Multisensory integration processes are fundamental to our sense of self as embodied beings. Bodily illusions, such as the rubber hand illusion (RHI) and the size-weight illusion (SWI), allow us to investigate how the brain resolves conflicting multisensory evidence during perceptual inference in relation to different facets of body representation. In the RHI, synchronous tactile stimulation of a participant's hidden hand and a visible rubber hand creates illusory body ownership; in the SWI, the perceived size of the body can modulate the estimated weight of external objects. According to Bayesian models, such illusions arise as an attempt to explain the causes of multisensory perception and may reflect the attenuation of somatosensory precision, which is required to resolve perceptual hypotheses about conflicting multisensory input. Recent hypotheses propose that the precision of sensorimotor representations is determined by modulators of synaptic gain, like dopamine, acetylcholine, and oxytocin. However, these neuromodulatory hypotheses have not been tested in the context of embodied multisensory integration. The present, double-blind, placebo-controlled, crossover study ( N = 41 healthy volunteers) aimed to investigate the effect of intranasal oxytocin (IN-OT) on multisensory integration processes, tested by means of the RHI and the SWI. Results showed that IN-OT enhanced the subjective feeling of ownership in the RHI, only when synchronous tactile stimulation was involved. Furthermore, IN-OT increased an embodied version of the SWI (quantified as estimation error during a weight estimation task). These findings suggest that oxytocin might modulate processes of visuotactile multisensory integration by increasing the precision of top-down signals against bottom-up sensory input.Peer reviewedFinal Accepted Versio

Implicit body representations and tactile spatial remapping

To perceive the location of a tactile stimulus in external space (external tactile localisation), information about the location of the stimulus on the skin surface (tactile localisation on the skin) must be combined with proprioceptive information about the spatial location of body parts (position sense) - a process often referred to as ‘tactile spatial remapping’. Recent research has revealed that both of these component processes rely on highly distorted implicit body representations. For example, on the dorsal hand surface position sense relies on a squat, wide hand representation. In contrast, tactile localisation on the same skin surface shows large biases towards the knuckles. These distortions can be seen as behavioural ‘signatures’ of these respective perceptual processes. Here, we investigated the role of implicit body representation in tactile spatial remapping by investigating whether the distortions of each of the two component processes (tactile localisation and position sense) also appear when participants localise the external spatial location of touch. Our study reveals strong distortions characteristic of position sense (i.e., overestimation of distances across vs along the hand) in tactile spatial remapping. In contrast, distortions characteristic of tactile localisation on the skin (i.e., biases towards the knuckles) were not apparent in tactile spatial remapping. These results demonstrate that a common implicit hand representation underlies position sense and external tactile localisation. Furthermore, the present findings imply that tactile spatial remapping does not require mapping the same signals in a frame of reference centered on a specific body part

Sensory and cognitive factors in multi-digit touch, and its integration with vision

Every tactile sensation – an itch, a kiss, a hug, a pen gripped between fingers, a soft fabric brushing against the skin – is experienced in relation to the body. Normally, they occur somewhere on the body’s surface – they have spatiality. This sense of spatiality is what allows us to perceive a partner’s caress in terms of its changing location on the skin, its movement direction, speed, and extent. How this spatiality arises and how it is experienced is a thriving research topic, compelled by growing interest in the nature of tactile experiences from product design to brain-machine interfaces. The present thesis adds to this flourishing area of research by examining the unified spatial quality of touch. How does distinct spatial information converge from separate areas of the body surface to give rise to our normal unified experience of touch? After explaining the importance of this question in Chapter 1, a novel paradigm to tackle this problem will be presented, whereby participants are asked to estimate the average direction of two stimuli that are simultaneously moved across two different fingerpads. This paradigm is a laboratory analogue of the more ecological task of representing the overall movement of an object held between multiple fingers. An EEG study in Chapter 2 will reveal a brain mechanism that could facilitate such aggregated perception. Next, by characterising participants’ performance not just in terms of error rates, but by considering perceptual sensitivity, bias, precision, and signal weighting, a series of psychophysical experiments will show that this aggregation ability differs for within- and between-hand perception (Chapter 3), is independent from somatotopically-defined circuitry (Chapter 4) and arises after proprioceptive input about hand posture is accounted for (Chapter 5). Finally, inspired by the demand for integrated tactile and visual experience in virtual reality and the potential of tactile interface to aid navigation, Chapter 6 will examine the contribution of tactile spatiality on visual spatial experience. Ultimately, the present thesis will reveal sensory factors that limit precise representation of concurrently occurring dynamic tactile events. It will point to cognitive strategies the brain may employ to overcome those limitations to tactually perceive coherent objects. As such, this thesis advances somatosensory research beyond merely examining the selectivity to and discrimination between experienced tactile inputs, to considering the unified experience of touch despite distinct stimulus elements. The findings also have practical implications for the design of functional tactile interfaces

A Soft Robotic Cover with Dual Thermal Display and Sensing Capabilities

We propose a new robotic cover prototype that achieves thermal display while also being soft. We focus on the thermal cue because previous human studies have identified it as part of the touch pleasantness. The robotic cover surface can be regulated to the desired temperature by circulating water through a thermally conductive pipe embedded in the cover, of which temperature is controlled. Besides, an observer for estimating heat from human contact is implemented; it can detect human interaction while displaying the desired temperature without temperature sensing on the surface directly. We assessed the validity of the prototype in experiments of temperature control and contact detection by human hand

Enkinaesthetic polyphony: the underpinning for first-order languaging

We contest two claims: (1) that language, understood as the processing of abstract symbolic forms, is an instrument of cognition and rational thought, and (2) that conventional notions of turn-taking, exchange structure, and move analysis, are satisfactory as a basis for theorizing communication between living, feeling agents. We offer an enkinaesthetic theory describing the reciprocal affective neuro-muscular dynamical flows and tensions of co- agential dialogical sense-making relations. This “enkinaesthetic dialogue” is characterised by a preconceptual experientially recursive temporal dynamics forming the deep extended melodies of relationships in time. An understanding of how those relationships work, when we understand and are ourselves understood, when communication falters and conflict arises, will depend on a grasp of our enkinaesthetic intersubjectivity

Two-photon all-optical interrogation of mouse barrel cortex during sensory discrimination

The neocortex supports a rich repertoire of cognitive and behavioural functions, yet the rules, or neural ‘codes’, that determine how patterns of cortical activity drive perceptual processes remain enigmatic. Experimental neuroscientists study these codes through measuring and manipulating neuronal activity in awake behaving subjects, which allows links to be identified between patterns of neural activity and ongoing behaviour functions. In this thesis, I detail the application of novel optical techniques for simultaneously recording and manipulating neurons with cellular resolution to examine how tactile signals are processed in sparse neuronal ensembles in mouse somatosensory ‘barrel’ cortex. To do this, I designed a whisker-based perceptual decision-making task for head-fixed mice, that allows precise control over sensory input and interpretable readout of perceptual choice. Through several complementary experimental approaches, I show that task performance is exquisitely coupled to barrel cortical activity. Using two- photon calcium imaging to simultaneously record from populations of barrel cortex neurons, I demonstrate that different subpopulations of neurons in layer 2/3 (L2/3) show selectivity for contralateral and ipsilateral whisker input during behaviour. To directly test whether these stimulus-tuned groups of neurons differentially impact perceptual decision-making I performed patterned photostimulation experiments to selectively activate these functionally defined sets of neurons and assessed the resulting impact on behaviour and the local cortical network in layer 2/3. In contrast with the expected results, stimulation of sensory-coding neurons appeared to have little perceptual impact on task performance. However, activation of non- stimulus coding neurons did drive decision biases. These results challenge the conventional view that strongly sensory responsive neurons carry more perceptual weight than non-responsive sensory neurons during perceptual decision-making. Furthermore, patterned photostimulation revealed and imposed potent surround suppression in L2/3, which points to strong lateral inhibition playing a dominant role in shaping spatiotemporally sparse activity patterns. These results showcase the utility of combined patterned photostimulation methods and population calcium imaging for revealing and testing neural circuit function during sensorimotor behaviour and provide new perspectives on sensory coding in barrel cortex