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Perceptual Functions of Auditory Neural Oscillation Entrainment Perceptual Functions of Auditory Neural Oscillation Entrainment Perceptual functions of auditory neural oscillation entrainment Perceptual functions of auditory neural oscillation entrainment By Andrew Chang, Bachelor of Science A Thesis Submitted to the School of Graduate Studies in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy McMaster University © Copyright by Andrew Chang July 29, 2019 McMaster University Doctor of Philosophy (2019) Hamilton, Ontario (Psychology, Neuroscience & Behaviour) TITLE: Perceptual functions of auditory neural oscillation entrainment AUTHOR: Andrew Chang (McMaster University) SUPERVISOR: Dr. Laurel J. Trainor NUMBER OF PAGES: xviii, 134 ii Lay Abstract Perceiving speech and musical sounds in real time is challenging, because they occur in rapid succession and each sound masks the previous one. Rhythmic timing regularities (e.g., musical beats, speech syllable onsets) may greatly aid in overcoming this challenge, because timing regularity enables the brain to make temporal predictions and, thereby, anticipatorily prepare for perceiv- ing upcoming sounds. This thesis investigated the perceptual and neural mechanisms for tracking auditory rhythm and enhancing perception. Per- ceptually, rhythmic regularity in streams of tones facilitates pitch percep- tion. Neurally, multiple neural oscillatory activities (high-frequency power, low-frequency phase, and their coupling) track auditory inputs, and they are associated with distinct perceptual mechanisms (enhancing sensitivity or de- creasing reaction time), and these mechanisms are coordinated to proactively track rhythmic regularity and enhance audition. The findings start the dis- cussion of answering how the human brain is able to process and understand the information in rapid speech and musical streams. iii Abstract Humans must process fleeting auditory information in real time, such as speech and music. The amplitude modulation of the acoustic waveforms of speech and music is rhythmically organized in time, following, for example, the beats of music or the syllables of speech, and this property enables temporal pre- diction and proactive perceptual optimization. At the neural level, external rhythmic sensory input entrains internal neural oscillatory activities, includ- ing low-frequency (e.g., delta, 1-4 Hz) phase, high-frequency (e.g., beta, 15-25 Hz) power, and their phase-amplitude coupling. These neural entrainment activities represent internal temporal prediction and proactive perceptual op- timization. The present thesis investigated two critical but previously unsolved questions. First, do these multiple entrainment mechanisms for tracking au- ditory rhythm have distinct but coordinated perceptual functions? Second, does regularity in the temporal (when) domain associate with prediction and perception in the orthogonal spectral (what) domain of audition? This the- sis addressed these topics by combining electroencephalography (EEG), psy- chophysics, and statistical modeling approaches. Chapter II shows that beta power entrainment reflects both rhythmic temporal prediction (when events are expected) and violation of spectral information prediction (what events are expected). Chapter III further demonstrates that degree of beta power en- trainment prior to a pitch change reflects how well an upcoming pitch change will be predicted. Chapter IV reveals that rhythmic organization of sensory input proactively facilitates pitch perception. Trial-by-trial behavioural-neural associations suggested that delta phase entrainment reflects temporal expec- tation, beta power entrainment reflects temporal attention, and their phase- amplitude coupling reflects the alignment of these two perceptual mechanisms and is associated with auditory-motor communication. Together, this thesis advanced our understanding of how neural entrainment mechanisms relate to perceptual functions for tracking auditory events in time, which are essential for perceiving speech and music. iv Acknowledgements I am grateful to complete my Ph.D. in a very supportive and kind labora- tory filled with joy. There are many people whom I would like to thank. First of all, thank you very much to my supervisor Laurel J. Trainor from the very bottom of my heart. Thanks to her great support on my research, professional development, and networking (I must have spent over millions of dollars at- tending conferences). I was very lucky to freely explore the research topics I am interested in while being able to have her guidance on how to implement them. She built a perfect lab (actually, “labs”) composed with skilled and nice colleagues that I could never dream of. Beyond professional aspects, I feel that Laurel is my friend. She never questioned me for attending rehearsals for Symphony on the Bay instead of working late in the lab. Doing my Ph.D. with Laurel was one of the best decisions I have ever made, even thought I was merely tempted by the Niagara Falls and winery tour she took me on when she tried to recruit me. Thanks to my progress committee members, Dan Bosnyak and Ian Bruce, my comprehensive exam committee members, Bruce Milliken and Larry Roberts, and my external examiner, Sidney Segalowitz. Their opinions and input have always made my research better. They were always there when I needed help, and have always guided me through technical and scientific challenges. Thanks to my lab-mates, Haley Kragness, Laura Cirelli, Chris Slugocki, Kate Einarson, Rayna Friendly, Sima Hoseingholizade, Sarah Lade, Hector Orozco Perez, David Prete, Chantal Carrillo, Dobri Dotov, Dan Cameron, Erica Flaten, and Debanjan Borthakur. They made doing science much more fun. Thanks to the lab staff members, Dave Thompson, Dan Bosnyak, Elaine Whiskin, Susan Marsh-Rollo, Carl Karichian, and Steven Living- stone. Research would have been much more challenging without their assis- tance. Thank you very much to my undergraduate assistants, Alexandra Rice, Michael Wan, Jessica Empringham, Michael Ku, Keeyeon Mark Hwang, Jen- nifer Chan, Katie Clayworth, Emily Kaunismaa, Brittany Ung, Elger Baraku, Jasmine Zhang, Jessica Otoo-Appiah, Taylor Barton, and Tessa Dickison. The data collection process would have been much much much more painful without them. Their hard work and their consent of slavery are deeply appreciated. v Specifically, I am truly indebted to Dave Thompson for making all my ex- periments possible, no matter how technically complicated they were. Thanks to Elaine and Susan for always being so enthusiastic. Having lunch with them was my daily retreat. Also, I enjoyed playing squash with Chris, even though failing to defeat him before completing my Ph.D. might be my biggest regret forever. Last but not least, I appreciate Haley and Laura for dragging me out of my cave to hang out with people and explore the city. They have helped me appreciate the North American bad jokes. Thanks to the music cognition gang of my cohort, Blair Ellis, Lorraine Chuen, and Haley (again). It was truly fun to hang out with these guys, no matter if we were discussing science, music, or any crazy topic, although I still cannot appreciate the weird indie movie we have watched together (picked by Blair, btw). Thanks to all my friends in the program, Brandon Paul, Fiona Manning, Blake Anderson, Jessica Cali, Mike Galang (GO Raptors!), Mike Slugocki (don’t support the Warriors), Ali Hashemi, Kiret Dhindsa, Lux Li, Ye Yuan, Anna Siminoski, Aimee Battcock, Sharmila Sreetharan, and many others. They all have made my graduate life special. Thanks to all the Taiwanese friends I have had in Hamilton and Toronto, Fred Liao, N曜h, 賴£安, Leo Hsu, Allison Yeh, sUh, Darren Wang, I ww, Johnson Chen, Mei-Ju Shih, Mei-Cheng Shih, 5è勛, Lucia Huang, Yu-Sian Li, 唐U, m斌, Mandy Chu, Ming-Feng Chiang, Amy Liao, and many others. Thanks for their hospitality when I first arrived Hamilton. They all made me feel at home, except once we got lost in the woods in Dundas Peak. Thanks to my parents, I am extremely lucky to be in a family that fully supports my career path and has a background of working in academia. They supported me to do my Ph.D. 12,104 km away from them. They have offered me great and honest guidance of academic survival skills, not to mention all the free statistical consulting sessions offered by my dad. I cannot enjoy doing research without their unconditional support and love, especially that they fed me quite a lot every time I went back home. Thanks to my parents-in-law, again, I am extremely lucky to have their unconditional trust and support, as they never questioned why the husband of their lovely daughter is still a poor graduate student at the age of 29. Most importantly, I cannot express how grateful I am for the love of my wife Wei Vivian Tsou. She supported my goal to pursue my Ph.D. abroad, vi even when we just met in Taiwan and I planned to leave in less than a year. She defied the challenges of our 40-month long-distance relationship between Hamilton and Taipei as well as Hamilton and New York. She relocated to Hamilton to start our family together after we were married. She was happy and sad for all the accomplishments and obstacles I have had, and she cheered me up whenever I was too anxious to make any moves. Her unconditional love and endless support made me a better person and that I can better enjoy this career path, which should also be granted with a Ph.D. (Push husband to Doctorate). This thesis could not be completed without her, not only because she is my proofreader, but she is also my perfect companion on this journey. vii Table of Contents Lay Abstract iii Abstract iv Acknowledgements v Table of Contents ix List of Figures xiii List of Tables xv Declaration of Academic Achievement xvii I General introduction 1 I.1 Perceptual mechanisms of auditory rhythm tracking . 3 I.2 Dynamic neural mechanisms of auditory rhythm tracking . 4 I.2.1 Definition of neural entrainment . 5 I.2.2 Neural entrainment for tracking auditory rhythm . 5 I.3 Aims of this thesis . 8 II Unpredicted pitch modulates beta oscillatory power during rhythmic entrainment to a tone sequence 11 II.1 Preface .
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