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An Investigation of Tactile Localization and Skin-Based Maps

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An Investigation of Tactile Localization and Skin-Based Maps Where was I touched? – An investigation of tactile localization and skin-based maps Jack Brooks Doctor of Philosophy Neuroscience Research Australia School of Medical Sciences, Faculty of Medicine University of New South Wales November 2017 ii Preface The coding of the position of touch on the skin and of the size and shape of the body are both fundamental for interacting with our surrounds. The aim of this thesis was to learn more about the mechanisms of tactile localization and to characterize the principles by which skin-based representations of the body update. It is commonly accepted that skin-based representations of the body are generated from the statistics of touch and other inputs. My studies required skin stimulation customised to account for inter-individual differences in touch sensitivity and forearm shape. Within the constraints of these methodological challenges, the central questions of this thesis were addressed by performing multiple behavioural experiments. In my first study, I tested how touch intensity and history influence touch localization. The study showed that reducing touch intensity increases the variability of pointing responses to touch and results in spatial biases to the middle of the recent history of touch. Thus, I showed that when uncertain about perceived touch location, a strategy is used that minimises localization errors over time. This error minimisation mechanism stabilises our perception of events on the skin and their sensory features. Next, I investigated uncertainty in a motion stimulus by fragmenting it. Studies in vision suggest that missing sensory inputs are filled-in from the surrounds, while previous tactile studies suggest fragmented motion could influence skin-based representations. In my second study, I used a motion stimulus which instantly skipped a spatial gap on the forearm, as if the skin surface had been rearranged. As expected I observed mislocalization toward the spatial gap, consistent with changes to receptive fields of neurons corresponding to the skipped patch. The results of my third study were discordant with expectations about the perceived extent of fragmented motion. However, I identified a novel judgment-dependent perceived shortening of a tactile motion stimulus. My final study found that this judgment contingent effect was accompanied by a perceived shortening of the forearm. Future studies might further explore this linkage between skin-based and higher-order representations of the limbs. The findings from this thesis may be clinically relevant for correcting disordered representations. iii Contents Commitment to open science .......................................................................................... ix Originality statement ........................................................................................................ x Copyright statement ........................................................................................................ xi Authenticity statement .................................................................................................... xi Acknowledgements ........................................................................................................ xii Publications from this thesis ......................................................................................... xiii Selected presentations from this thesis ......................................................................... xiv Selected other publications during this candidature ...................................................... xv Abbreviations ............................................................................................................... xvii Chapter 1: General Introduction ............................................... 1 1.1 Overview .................................................................................................................... 1 1.2 Touch localization ...................................................................................................... 2 1.2.1 Body representations .................................................................................. 4 1.2.2 Touch detection and localization ............................................................... 6 1.2.3 Touch localization: Acuity is better than predicted ................................... 8 1.2.4 Uncertainty about touch location ............................................................... 9 1.2.5 Localization tasks .................................................................................... 12 1.2.6 Localization in external space .................................................................. 12 1.2.7 Tactile extent ............................................................................................ 14 1.2.8 Extent reporting methods ......................................................................... 17 iv 1.3 Contextual influences on touch localization ............................................................ 19 1.3.1 Filling-in: Vision ...................................................................................... 19 1.3.2 Filling-in: Skin-based maps ..................................................................... 22 1.3.3 Filling-in: On the body............................................................................. 24 1.3.4 Spatiotemporal context modulates localization ....................................... 25 1.3.5 The spatiotemporal conditions of the CRE .............................................. 26 1.3.6 Saltation: Models ..................................................................................... 28 1.3.7 Saltation: Neural mechanisms .................................................................. 29 1.4 Tactile perception of objects and the body .............................................................. 31 1.4.1 Perceptual binding ................................................................................... 31 1.4.2 Binding: Occluded objects ....................................................................... 32 1.4.3 Space gives to time: The abridging effect ................................................ 34 1.4.4 Cortical reorganization............................................................................. 35 1.4.5 Organizing somatosensation: Analysing the statistics of touch to the skin .......................................................................................................................... 36 1.4.6 Summary .................................................................................................. 37 1.5 Aims of the present studies ...................................................................................... 38 Chapter 2: Perceived Position of Touch on the Skin ............. 40 2.1 Summary .................................................................................................................. 40 2.2 Introduction .............................................................................................................. 41 2.3 Materials and methods ............................................................................................. 44 2.4 Results ...................................................................................................................... 50 2.4.1 Experiment 1: Stimulus distribution centred on forearm ......................... 50 v 2.4.2 Experiment 2: Localization of Strong and Super strong stimuli .............. 51 2.4.3 Experiment 3: Stimulus distribution offset from middle of forearm ....... 52 2.5 Discussion ................................................................................................................ 55 Chapter 3: Artificial Tactile Scotoma ...................................... 59 3.1 Summary .................................................................................................................. 59 3.2 Introduction .............................................................................................................. 60 3.3 Materials and methods ............................................................................................. 64 3.4 Results ...................................................................................................................... 71 3.4.1 Preliminary Experiment 1: Effect of nearby pressure on localization ..... 71 3.4.2 Preliminary Experiment 2: Intensity adaptation ...................................... 72 3.4.3 Experiment 1: Artificial tactile scotoma .................................................. 73 3.5 Discussion ................................................................................................................ 78 Chapter 4: Tactile Motion Extent ............................................ 85 4.1 Summary .................................................................................................................. 85 4.2 Introduction .............................................................................................................. 86 4.3 Materials and general methods ................................................................................ 89 4.4 Experimental studies ................................................................................................ 92 4.4.1 Experiment 1: Perceived extent of fragmented tactile motion ................. 92 4.5 Control experiments ................................................................................................. 97 4.5.1 Control experiment 1a: Extent
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