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INVESTIGATING THE RELATIONS BETWEEN OBJECT AFFORDANCE AND PERCEPTION USING BEHAVIOURAL AND BRAIN IMAGING EXPERIMENTS by MELANIE WULFF A thesis submitted to the University of Birmingham for the degree of DOCTOR OF PHILOSOPHY School of Psychology College of Life and Environmental Sciences University of Birmingham September 2015 University of Birmingham Research Archive e-theses repository This unpublished thesis/dissertation is copyright of the author and/or third parties. The intellectual property rights of the author or third parties in respect of this work are as defined by The Copyright Designs and Patents Act 1988 or as modified by any successor legislation. Any use made of information contained in this thesis/dissertation must be in accordance with that legislation and must be properly acknowledged. Further distribution or reproduction in any format is prohibited without the permission of the copyright holder. ABSTRACT Previous research has shown facilitatory effects on attention and perception when object pairs are positioned for action compared to when they are not positioned for action. The present thesis aimed to better understand the mechanisms underlying this paired-object affordance effect. Chapters 2 and 3 showed that different but interacting parieto-frontal networks contribute to the effects of paired-object affordance in healthy participants. Chapters 4 and 5 explored the effects of paired-object affordance on visual extinction; the data showed that recovery from extinction was sensitive to the familiarity of the object pair and the completeness of the active object rather than the passive object within an interacting pair. Finally, the role of contextual information and task demands on the automatic perception of paired-object affordance effects was directly explored. The results indicate that only explicit but not implicit task demands (searching for an action vs. a colour pair, respectively) had facilitatory effects on performance and that semantic information in a scene also modulates the automatic perception of paired-object affordance. The findings provide novel behavioural and neuroimaging evidence that paired-object affordance is influenced by contextual information and task demands, with the active object (the tool) within a pair modulating the allocation of attention. To my parents, Norbert and Marianne, Without whom none of my journey would be possible ACKNOWLEDGEMENTS I would like to thank my wonderful supervisors, Professor Glyn Humphreys and Dr Pia Rotshtein for their guidance, patience, encouragement, inspiration, and unwavering support beyond academia – I cannot thank you enough for that. I am grateful to my closest friends, in particular Maliheh Taheri, Robin Green, Rainer Bögle, Yuanyuan Zhao, and Nabeela Akhtar for your friendship, love and support. A big thank you also goes to my friends and colleagues at the University of Birmingham to make my stay in Birmingham an once-in-a-lifetime experience. I would also like to thank my previous housemates Daisy Jing Lyu and Ed Corless for their “distraction” and the big hugs when needed. I would also like to thank Ed for proofreading the final version. I would also like to thank all the study participants for their time. Most importantly, I would like to say Danke schön to my parents for their love, unconditional support, and encouragement throughout my life for which I am overwhelmingly grateful. I would also like to give special thanks to Helen Grant-Taylor, my Australian mum, for being there for me during the most difficult time in my life. I would also like to thank my best friend, Angela Ritter, for always being there for me. Finally, special thanks to Jamie Barrett, for your patience, hugs, and so many laughs throughout the last couple of months. TABLE OF CONTENT CHAPTER 1: GENERAL INTRODUCTION ...................................................................... 1 1.1 Introduction ........................................................................................................................ 1 1.2 Affordance ........................................................................................................................... 1 1.3 Paired-object affordance .................................................................................................... 7 1.3.1 Mechanisms underlying paired-object affordance .......................................................... 10 1.4 Dual-route model from vision to action .......................................................................... 12 1.5 Neurocognitive models related to affordance ................................................................ 14 1.5.1 Tool-selective network .................................................................................................... 14 1.5.2 Action observation network ............................................................................................ 16 1.5.3 Mirror neuron system ...................................................................................................... 17 1.6 Overview of the thesis ...................................................................................................... 20 CHAPTER 2: DISTINCT NEURAL EFFECTS OF PERSPECTIVE AND HAND ALIGNMENT ON PAIRED-OBJECT AFFORDANCE: AN FMRI STUDY ................. 22 2.1 Abstract ............................................................................................................................. 22 2.2 Introduction ...................................................................................................................... 23 2.3 Methods ............................................................................................................................. 29 2.3.1 Participants ...................................................................................................................... 29 2.3.2 Stimuli ............................................................................................................................. 29 2.3.3 Design and procedure ...................................................................................................... 32 2.3.4 fMRI data acquisition ...................................................................................................... 33 2.3.5 fMRI data analysis ........................................................................................................... 33 2.4 Results ................................................................................................................................ 35 2.4.1 Behavioural results .......................................................................................................... 35 2.4.2 Imaging data .................................................................................................................... 36 2.4.2.1 Visual-based affordance cues ....................................................................................... 37 2.4.2.2 Motor-based affordance cues........................................................................................ 39 2.4.2.3 Combined visual- and motor-based affordance cues .................................................... 40 2.5 Discussion .......................................................................................................................... 44 2.5.1 Neural correlates for visual-based affordance cues ......................................................... 45 2.5.2 Neural correlates for motor-based affordance cues ......................................................... 47 2.5.3 Effects of visual perspective – combined visual- and motor-based affordance cues ...... 49 2.6 Study limitations ............................................................................................................... 51 2.7 Conclusion ......................................................................................................................... 53 CHAPTER 3: PAIRED-OBJECT AFFORDANCE AFFECTS MOTOR CORTEX EXCITABILITY ..................................................................................................................... 54 3.1 Abstract ............................................................................................................................. 54 3.2 Introduction ...................................................................................................................... 55 3.3 Method ............................................................................................................................... 59 3.3.1 Participants ...................................................................................................................... 59 3.3.2 TMS and MEP recording ................................................................................................. 59 3.3.3 Apparatus and stimuli ...................................................................................................... 61 3.3.4 Design and procedure ...................................................................................................... 64 3.3.5 Data analysis .................................................................................................................... 65 3.4 Results ................................................................................................................................ 66 3.4.1 Effects of hand presence .................................................................................................. 66 3.4.2 Effects of action relation, hand posture and hand alignment .......................................... 67 3.5 Discussion .......................................................................................................................... 70 3.5.1 Effects of action relation, hand posture, and hand alignment on MEP response ............ 70 3.5.2 Differences between left and right M1 excitability ......................................................... 74 3.5.3 Effects of hand presence on MEP response .................................................................... 76 3.6 Conclusion ......................................................................................................................... 77 CHAPTER 4: VISUAL RESPONSES TO ACTION BETWEEN UNFAMILIAR OBJECT PAIRS MODULATE EXTINCTION .................................................................. 78 4.1 Abstract ............................................................................................................................. 78 4.2 Introduction ...................................................................................................................... 79 4.3 Methods ............................................................................................................................. 83 4.3.1 Patients ............................................................................................................................ 83 4.3.2 Apparatus and stimuli ...................................................................................................... 87 4.3.3 Design and procedure ...................................................................................................... 90 4.4 Results ................................................................................................................................ 92 4.4.1 The presence of extinction ............................................................................................... 92 4.4.2 Effects of action relation on two-item report................................................................... 95 4.4.3 Effects of action relation on identification errors on two-item trials .............................. 97 4.5 Discussion ........................................................................................................................ 102 4.6 Conclusion ....................................................................................................................... 105 CHAPTER 5: EFFECTS OF BROKEN AFFORDANCE ON VISUAL EXTINCTION ................................................................................................................................................ 107 5.1 Abstract ........................................................................................................................... 107 5.2 Introduction .................................................................................................................... 108 5.3 Methods ........................................................................................................................... 112 5.3.1 Patients .......................................................................................................................... 112 5.3.2 Apparatus and stimuli .................................................................................................... 114 5.3.3 Design and procedure .................................................................................................... 117 5.4 Results .............................................................................................................................. 119 5.4.1 The presence of extinction ............................................................................................. 121 5.4.2 Effects of object pair type on two-item report (intact handles) ..................................... 124 5.4.3 Role of broken handles on two-item trial performance ................................................. 126 5.4.4 Effect of stimulus type on one-item report .................................................................... 132 5.5 Discussion ........................................................................................................................ 133 5.5.1 Effects independent of the broken handle ..................................................................... 134 5.5.2 Effects when a handle was broken ................................................................................ 135 5.6 Study limitations ............................................................................................................. 139 5.7 Conclusion ....................................................................................................................... 140 CHAPTER 6: EFFECTS OF PAIRED-OBJECT AFFORDANCE IN SEARCH TASKS ACROSS THE ADULT LIFESPAN ................................................................................... 141 6.1 Abstract ........................................................................................................................... 141 6.2 Introduction .................................................................................................................... 142 6.3 Method ............................................................................................................................. 153 6.3.1 Participants .................................................................................................................... 153 6.3.2 Stimuli and design ......................................................................................................... 154 6.3.3 Procedure ....................................................................................................................... 158 6.3.4 Data analysis .................................................................................................................. 159 6.4 Results .............................................................................................................................. 159 6.5 Discussion ........................................................................................................................ 169 6.5.1 Effects of procedural knowledge – selection priority for the active object – direct route for action retrieval .............................................................................................................. 170 6.5.2 Effects of semantic distracters – indirect semantic route for action retrieval ............... 174 6.5.3 Retrieval of action knowledge across the lifespan ........................................................ 176 6.6 Study limitations ............................................................................................................. 178 6.7 Conclusion ....................................................................................................................... 179 CHAPTER 7: GENERAL DISCUSSION AND CONCLUSIONS .................................. 180 7.1 Summary of thesis .......................................................................................................... 181 7.2 Paired-object affordance ................................................................................................ 185 7.2.1 Automaticity and the role of context and task demands ................................................ 185 7.2.2 The role of attention in paired-object affordance .......................................................... 187 7.2.3 Mechanisms underlying paired-object affordance ........................................................ 188 7.2.4 Affordance in relation to other perception-action theories ............................................ 191 7.3 Future studies .................................................................................................................. 194 7.4 Conclusion ....................................................................................................................... 196 APPENDICES ....................................................................................................................... 197 REFERENCES ..................................................................................................................... 201 * Chapters 4 and 5 are published journal articles. LIST OF FIGURES Figure 2.1. Examples of two-item stimuli presented from a first-person perspective (a), and from a third-person perspective (b). The objects were positioned as interacting pairs (left panels) or as non-interacting pairs (right panels). The active object (the bottle opener) was grasped either by the right hand (right-handed action; upper panels) or the left hand (left-handed action; lower panels). ..................................................... 31 Figure 2.2. Activation foci for main effect of action relation (AR > NAR) were overlaid on a single-participant structural MNI-template (FWE-corrected p < .05; cluster defining threshold p < .01, uncorrected). The bar graph shows the averaged effect size extracted from a 6-mm sphere centred on the peak coordinate of inferior parietal lobule (IPL; circled in red), precuneus and superior temporal gyrus (STG)/middle temporal gyrus (MTG). Error bars show SEMs. Abbreviations: AR = action relation; NAR = no action relation; 1PP = first-person perspective; 3PP = third- person perspective; RH = right-handed action; LH = left-handed action. ............... 38 Figure 2.3. Activation foci for the main effect of hand alignment (LH > RH) were overlaid on a single-participant structural MNI-template (FWE-corrected p < .05; cluster defining threshold p < .01, uncorrected). The bar graph shows the averaged effect size extracted from a 6-mm sphere centred on the peak coordinate of superior medial frontal gyrus (SmFG; circled in red). Error bars show SEMs. Abbreviations: AR = action relation; NAR = no action relation; 1PP = first-person perspective; 3PP = third-person perspective; RH = right-handed action; LH = left-handed action. .................................................................................................................................. 39

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