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University of Bradford Ethesis University of Bradford eThesis This thesis is hosted in Bradford Scholars – The University of Bradford Open Access repository. Visit the repository for full metadata or to contact the repository team © University of Bradford. This work is licenced for reuse under a Creative Commons Licence. The role of sensory history and stimulus context in human time perception Adaptive and integrative distortions of perceived duration Corinne FULCHER Submitted for the Degree of Doctor of Philosophy Faculty of Life Sciences University of Bradford 2017 Abstract Corinne Fulcher The Role of Sensory History and Stimulus Context in Human Time Perception Adaptive and Integrative Distortions of Perceived Duration Keywords: Time perception, Duration, Human, Adaptation, Integration, Subsecond, Vision, Audition, Psychophysics, Modelling This thesis documents a series of experiments designed to investigate the mechanisms subserving sub-second duration processing in humans. Firstly, duration aftereffects were generated by adapting to consistent duration information. If duration aftereffects represent encoding by neurons selective for both stimulus duration and non-temporal stimulus features, adapt-test changes in these features should prevent duration aftereffect generation. Stimulus characteristics were chosen which selectively target differing stages of the visual processing hierarchy. The duration aftereffect showed robust interocular transfer and could be generated using a stimulus whose duration was defined by stimuli invisible to monocular mechanisms, ruling out a pre- cortical locus. The aftereffects transferred across luminance-defined visual orientation and facial identity. Conversely, the duration encoding mechanism was selective for changes in the contrast-defined envelope size of a Gabor and showed broad spatial selectivity which scaled proportionally with adapting stimulus size. These findings are consistent with a second stage visual spatial mechanism that pools input across proportionally smaller, i spatially abutting filters. A final series of experiments investigated the pattern of interaction between concurrently presented cross-modal durations. When duration discrepancies were small, multisensory judgements were biased towards the modality with higher precision. However, when duration discrepancies were large, perceived duration was compressed by both longer and shorter durations from the opposite modality, irrespective of unimodal temporal reliability. Taken together, these experiments provide support for a duration encoding mechanism that is tied to mid-level visual spatial processing. Following this localised encoding, supramodal mechanisms then dictate the combination of duration information across the senses. ii Acknowledgements I wish to firstly thank Dr James Heron, for guiding me through the intricacies of human psychophysics and for his wealth of advice and support throughout my postgraduate studies. In addition, I am indebted to my collaborators and colleagues Professor David Whitaker, Dr Neil Roach, Dr Andrew Logan and Professor Paul McGraw for their professional expertise and help, particularly in regards to mathematics and computational modelling. I am incredibly grateful to my fellow postgraduate colleagues in G39, for your continued friendship and support, for steering me through the twists and turns of vision research, but also for giving up your time to participate in experiments. Thank you in particular to Dr Samantha Strong, who not only helped me to negotiate Matlab, SPSS and a host of neuroimaging literature, but who also became a fantastic friend over the last four years. Lastly I must thank my wonderful husband Jacko, and my parents Jen and Richard, for giving me unconditional love and support and the courage to undertake a PhD in the first place. You never stopped believing in me, and for that I will be eternally grateful. iii Table of Contents Abstract.................................................................................................. i Acknowledgements............................................................................... iii Table of Contents.................................................................................. iv Table of Figures..................................................................................... x Chapter 1 – Introduction....................................................................... 1 Chapter 2 – Literature Review.............................................................. 8 2.1 – Introduction.......................................................................... 8 2.2 – Classical models of temporal processing............................ 8 2.2.1 – Scalar timing.......................................................... 8 2.2.2 – Internal clock models............................................. 12 2.3 – Alternative models of temporal processing.......................... 18 2.3.1 – The striatal beat frequency model.......................... 18 2.3.2 – The state dependent network model...................... 23 2.3.3 – The channel-based processing model................... 29 2.3.4 – Summary................................................................ 35 2.4 – Psychophysical time............................................................ 36 2.4.1 – Conformity to the scalar property........................... 36 2.4.2 – Non-temporal magnitude....................................... 40 2.4.3 – Sensory modality................................................... 47 2.4.4 – Temporal adaptation.............................................. 53 2.4.5 – Attention and time.................................................. 59 iv 2.4.6 – Timing multiple independent intervals.................... 66 2.4.7 – Integrating multisensory durations......................... 70 2.4.8 – Summary................................................................ 85 2.5 – The neurophysiological basis of temporal processing......... 86 2.5.1 – Electrophysiological studies................................... 87 2.5.2 – Neuroimaging......................................................... 98 2.5.3 – Lesion studies........................................................ 102 2.5.3.1 – Transcranial magnetic stimulation............ 103 2.5.3.2 – Clinical populations.................................. 107 2.5.4 – Summary................................................................ 110 Chapter 3 – General methods............................................................... 112 3.1 – Introduction.......................................................................... 112 3.2 – Signal detection theory and decision types.......................... 112 3.3 – Psychophysical measures................................................... 117 3.3.1 – Sensory threshold.................................................. 117 3.3.2 – Perceptual bias...................................................... 118 3.4 – Methods used for the study of duration................................ 118 3.4.1 – Method of limits...................................................... 119 3.4.2 – Staircase method................................................... 122 3.4.3 – Magnitude estimation methods.............................. 124 3.4.4 – The method of single stimulus............................... 126 3.4.4.1 – Temporal bisection................................... 126 v 3.4.4.2 – Temporal generalisation........................... 128 3.4.5 – Method of constant stimuli...................................... 131 3.5 – The psychometric function................................................... 132 3.6 – Curve fitting.......................................................................... 136 3.7 – Bootstrapping....................................................................... 138 3.8 – Apparatus............................................................................ 141 3.9 – Experimental calibration...................................................... 142 3.9.1 – Gamma correction.................................................. 142 3.9.2 – Verifying stimulus timing........................................ 145 Chapter 4 – An investigation into the spatial tuning of the duration 148 aftereffect............................................................................................... Introduction................................................................................... 148 Experiment 4.1.............................................................................. 151 4.1.1 – Methods................................................................. 151 4.1.1.1 – Observers................................................. 151 4.1.1.2 – Stimuli and apparatus............................... 151 4.1.1.3 – Procedure................................................. 152 4.1.1.4 – Modelling.................................................. 154 4.1.2 – Results and discussion.......................................... 156 Experiment 4.2.............................................................................. 161 4.2.1 – Methods.................................................................. 161 4.2.1.1 – Observers................................................. 161 4.2.1.2 – Stimuli and apparatus............................... 161 vi 4.2.1.3 – Procedure................................................. 161 4.2.2 – Results and discussion........................................... 161 Experiment 4.3.............................................................................. 165 4.3.1 –
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