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Cross-Modal Mechanisms: Perceptual Multistability in Audition and Vision

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Cross-Modal Mechanisms: Perceptual Multistability in Audition and Vision Cross-modal mechanisms: perceptual multistability in audition and vision von der Fakultät für Naturwissenschaften der Technischen Universität Chemnitz genehmigte Dissertation zur Erlangung des akademischen Grades doctor rerum naturalium (Dr. rer. nat.) vorgelegt von M.Sc. Jan Grenzebach geboren am 06.07.1988 in Rotenburg an der Fulda eingereicht am 13.01.2021 Gutachter: Prof. Dr. rer. nat. habil. Alexandra Bendixen Prof. Dr. sc. nat. Wolfgang Einhäuser-Treyer Tag der Verteidigung: 04.05.2021 Veröffentlicht unter: https://nbn-resolving.org/urn:nbn:de:bsz:ch1-qucosa2-749013 ! 1! BIBLIOGRAPHISCHE BESCHREIBUNG Autor: Grenzebach, Jan Titel: “Cross-modal mechanisms: perceptual multistability in audition and vision” Dissertation an der Fakultät für Naturwissenschaften der Technischen Universität Chemnitz, Institut für Physik, Dissertation, 2021 189 Seiten; 46 Abbildungen; 5 Tabellen; 297 Literaturzitate; 13 digitale Medien (online) Referat: Perceptual multistability is a phenomenon that is mostly studied in all modalities separately. The phenomenon reveals fundamental principles of the perceptual system in the formation of an emerging cognitive representation in the consciousness. The momentary perceptual organizations evoked during the stimulation with ambiguous stimuli switches between several perceptual organizations or percepts: The auditory streaming stimulus in audition and the moving plaids stimulus in vision, elicit different at least two percepts that dominate awareness exclusively for a random phase or dominance duration before an inevitable switch to another percept occurs. The similarity in the perceptual experience has led to propose a global mechanism contributing to the perceptual multistability phenomena crossmodally. Contrary, the difference in the perceptual experience has led to propose a distributed mechanism that is modality-specific. The development of a hybrid model has synergized both approaches. We accumulate empirical evidence for the contribution of a global mechanism, albeit distributed mechanisms play an indispensable role in this cross-modal interplay. The overt report of the perceptual experience in our experiments is accompanied by the recording of objective, cognitive markers of the consciousness: Reflexive movements of the eyes, namely the dilation of the pupil and the optokinetic nystagmus, correlate with the unobservable perceptual switches and perceptual states respectively and have their neuronal rooting in the brainstem. We complement earlier findings on the sensitivity of the pupil to visual multistability: It was shown in two independent experiments that the pupil dilates at the time of reported perceptual switches in auditory multistability. A control condition on confounding effects from the reporting process confines the results. Endogenous, evoked internally by the unchanged stimulus ambiguity, and exogenous, evoked externally by the changes in the physical properties of the stimulus, perceptual switches could be discriminated based on the maximal amplitude of the dilation. The effect of exogenous perceptual has on the pupil were captured in a report and no-report task to detect confounding perceptual effects. In two additional studies, the moment-by-moment coupling and coupling properties of percepts between concurrent multistable processes in audition, evoked by auditory streaming, and in vision, evoked by moving plaids, were found crossmodally. In the last study, the externally induced percept in the visual multistable process was not relayed to the simultaneous auditory multistable process: Still, the observed general coupling is fragile but existent. The requirement for the investigation of a moment-by-moment coupling of the multistable perceptual processes was the application of a no-report paradigm in vision: The visual stimulus evokes an optokinetic nystagmus that has machine learnable different properties when following either of the two percepts. In combination with the manually reported auditory percept, attentional bottlenecks due to a parallel report were circumvented. The two main findings, the dilation of the pupil along reported auditory perceptual switches and the crossmodal coupling of percepts in bimodal audiovisual multistability, speak in favor of a partly global mechanism being involved in control of perceptual multistability; the global mechanism is incarcerated by the, partly independent, distributed competition of percepts on modality level. Potentially, supramodal attention-related modulations consolidate the outcome of locally distributed perceptual competition in all modalities. Schlagworte: perception, multistability, multimodal, pupil dilation, optokinetic nystagmus, hybrid mechanism, auditory streaming, moving plaids, no-report, support vector machine 2! ! ACKNOWLEDGEMENTS Foremost, I want to thank both my supervisors Prof. Dr. Alexandra Bendixen and Prof. Dr. Wolfgang Einhäuser-Treyer equally, for their enduring trust and guidance, granting me in their feedback the opportunity to develop new and improve skills and gain insights into their research methodology. In the modern scientific environment, they allowed me access to, I was able to unfold, learn, and contribute to open research questions in human perception. Thank you for providing me with what I could not ask for. For his knowledgeable technical support and tireless effort keeping the excellent state of the laboratory, Thomas Baumann has my gratitude. My project partner and co-author in several studies, Thomas Wegner, must be thanked for his well-commented code. The gathered empirical data depended on the thoughtful instruction of participants in our Experiments, which was executed superbly by Christiane Breitkreutz, Susann Lerche, and Fabian Parth. I appreciate our generous project funding from the German Research Foundation (DFG). For comments and discussion on earlier drafts of this Thesis I thank Dr. Bahareh Kiani and Caroline Hübner. Lastly, Dr. Karl Kopiske and Dr. Jan Drewes have been a rich source of know-how in practical statistical inference and the implementation of machine-learning algorithms, respectively. The last three years would have been a less enjoyable journey without family, friends, and colleagues near-and-far: my sisters Lisa and Hannah, Alexander Jarka, Marius Bartholmai, Felix Beyer, Sophie Metz, Monique Michl, Daniel Walper, Daniel Backhaus, Max Theisen, Christiane Neubert and Tobias Müller… Thank you! Chemnitz, 13th January, 2021, Jan Grenzebach ! 3! CONTENTS COVER . 1 BIBLIOGRAPHISCHE BESCHREIBUNG . 2 ACKNOWLEDGEMENTS . 3 CONTENTS . 4 CHAPTER 1: Introduction . 6 C1.1: Stability and uncertainty in perception . 6 C1.2: Auditory, visual and audio-visual multistability . 14 C1.3: Capturing the subjective perceptual experience . 25 C1.4: Limitations of preceding studies, objectives, and outline of the Thesis. 33 CHAPTER 2: Study 1 “Pupillometry in auditory multistability” . 36 C2.1.1 Experiment 1: Introduction . 36 C2.1.2 Experiment 1: Material and Methods . 38 C2.1.3 Experiment 1: Data analysis . 44 C2.1.4 Experiment 1: Results . 48 C2.1.5 Experiment 1: Discussion . 52 C2.2.1 Experiment 2: Introduction . 54 C2.2.2 Experiment 2: Material and Methods . 54 C2.2.3 Experiment 2: Data analysis . 56 C2.2.4 Experiment 2: Results . 57 C2.3 Experiment 1 & 2: Discussion . 61 C2.4 Supplement Study 1 . 65 CHAPTER 3: Study 2 “Multimodal moment-by-moment coupling in perceptual bistability” . 71 C3.1.1 Experiment 1: Introduction . 71 C3.1.2 Experiment 1: Results . 74 C3.1.3 Experiment 1: Discussion . 80 C3.1.4 Experiment 1: Material and Methods . 84 C3.1.5 Experiment 1: Data analysis . 87 C3.2 Supplement Study 2 . 92 4! ! CHAPTER 4: Study 3 “Boundaries of bimodal coupling in perceptual bistability” . 93 C4.1.1 Experiment 1: Introduction . 93 C4.1.2 Experiment 1: Material and Methods . 98 C4.1.3 Experiment 1: Data analysis . 102 C4.1.4 Experiment 1: Results . 108 C4.1.5 Experiment 1: Discussion . 114 C4.2.1 Experiment 2: Introduction . 116 C4.2.2 Experiment 2: Material and Methods . 119 C4.2.3 Experiment 2: Data analysis . 125 C4.2.4 Experiment 2: Results . 133 C4.3 Experiment 1 & 2: Discussion . 144 C4.4 Supplement Study 3 . 151 CHAPTER 5: General Discussion. 154 C5.1 Significance for models of multistability and implications for the perceptual architecture 162 C5.2 Recommendations for future research . 166 C5.3 Conclusion . 168 REFERENCES . 170 APPENDIX . 186 A1: List of Figures . 186 A2: List of Tables . 188 A3: List of Abbreviations and Symbols . 189 ! ! ! ! ! ! ! 5! C1.1 Stability and uncertainty in perception CHAPTER 1 Introduction C1.1 Stability and uncertainty in perception Interactions with the surrounding scenery rely on a coherent and meaningful cognitive representation of physical attributes emitted or reflected by the distal surfaces such as light or sound waves. Sensory cells in the ear and the eye organs capture the radiated waves (electromagnetic or compressed air, respectively), transduce them into neuronal signals, and forward the sensations to higher-level processing (Yoshioka & Sakakibara, 2013). The raw neuronal signals need to be organized to form a consciously available perceptual representation (Y. C. Chen & Spence, 2017). The formative process begins on low sensory levels and is carried upwards to larger populations of neurons that interact with each other. Instead of passive transmission of the information stimulating the sensors by the neuronal structure, it was discovered that perception is an active process. In the course of the process, the neuronal signals
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