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The Regulation of Tissue N-Acylethanolamine and Arachidonoylglycerol Concentrations By The regulation of tissue n-acylethanolamine and arachidonoylglycerol concentrations by diet, ischemia, n-acyl phosphatidylethanolamine-specific phospholipase D and fatty acid amide hydrolase By Lin Lin A thesis submitted in conformity with the requirements for the degree of Doctorate of Philosophy Department of Nutritional Sciences University of Toronto © Copyright by Lin Lin, 2018 The regulation of tissue n-acylethanolamine and arachidonoylglycerol levels by diet, ischemia, n-acyl phosphatidylethanolamine-specific phospholipase D and fatty acid amide hydrolase Lin Lin Doctor of Philosophy Department of Nutritional Sciences University of Toronto 2018 Abstract Dietary fatty acids (DFAs) can modulate levels of n-acylethanolamines (NAEs) and arachidonoylglycerols (AGs) via the synthetic enzyme, n-acylphosphatidylethanolamine- specific phospholipase D (NAPE-PLD) and degradation enzyme, fatty acid amide hydrolase (FAAH). However, other parallel pathways maybe also involved. This research hypothesizes that the absence of NAPE-PLD will lower tissue NAEs; while the absence of FAAH will increase NAEs. Also, basal NAE and AG levels are lower than ischemia-induced levels, while DFA can modulate basal NAE levels. In study 1, wild-type (C57BL/6), NAPE-PLD (-/+) and NAPE-PLD (-/-) mice were fed AIN-93G diets enriched with beef tallow, canola oil, corn oil or fish oil for nine weeks then killed by microwave fixation. Results showed that NAPE-PLD (-/-) had lower levels of plasma and jejunum oleoylethanolamide (OEA), lower food intake, body weight, and body fat composition than wild-type. Also, corn oil diet elevated arachidonoylethanolamide (AEA) and AGs; while fish oil elevated docosahexaenoylethanolamide (DHEA) in all genotypes. Therefore, NAPE-PLD is specific to OEA, but not other NAE, which may play a role in regulating body composition and weight management. Also, DFA can still elevate NAE levels without NAPE-PLD suggesting it is not necessary for dietary increased NAEs. In study 2, wild-type and FAAH-KO mice were fed a standard chow diet for 9 weeks; then killed by control (microwave fixation) or partial CO2- induced ischemia or CO2-induced ischemia. Brain NAE levels were higher in the FAAH-KO than the wild-type. Interestingly, CO2-induced ischemia increased all NAEs and AGs in the wild-type mice, but only DHEA was increased in the FAAH-KO mice. Thus, FAAH may play II an essential role in regulating lipid metabolism in response to ischemia. In conclusion, this work provides evidence that diet, NAPE-PLD, FAAH and ischemia are independently involved in maintaining tissue NAE levels. III Acknowledgment Beyond the persistent, cogent and passionate scientific research, this thesis is also fulfilled with my inner growth and the spirit of precious intellectual, psychological and emotional support provided by many people. I would like to share many thanks and many moments, which built the backbone of my strength to accomplish my Ph.D. degree. Dr. Richard Bazinet: I feel honored to share my sincere gratitude to my supervisor, Dr. Richard Bazinet, for the opportunity you provided me to learn and grow under your guidance, advice, and support. I have been fortunate to have you as my supervisor who genuinely cared about my research and who taught me how to think critically and write effectively to become a better scientist. I also inspired by your research ethics and philosophy: “If you followed every step carefully to test your hypothesis, then don’t worry about what the results might say. A result is a result; this is science.” Dr. Peter Jones: I sincerely thank my co-supervisor, Dr. Peter Jones, for your continuous support and the guidance you gave to me during my Ph.D. Your countless encouragement and mentorship since my Master degree at the University of Manitoba. Your support and belief in my ability on researching the field of nutrition and endocannabinoids for the past ten years have been invaluable. I become a better researcher since your first advice: “Be on time; use more English, and get your work published.” Dr. Harvey Anderson: I sincerely thank my advisory committee member, Dr. Harvey Anderson for the knowledge and experience you guided to me and the tough questions you challenged me to understand my project and the general concept of science in an exceptional training. I thank Dr. Elena Comelli, my advisory committee member, for generously given your time and insights to guide me on my graduate studies and thesis writing during the past five years. I thank Dr. Deborah O’Connor and Dr. Mary R. L’Abbé for acting as my Departmental Appraiser. Dr. Thomas Wolever for serving as my Departmental Chair; and Dr. Harold Aukema for acting as my External Appraiser from the University of Manitoba. I feel honored to have been among such a distinguished group of scientists. IV “Few are those who see with their own eyes and feel with their own hearts” – Albert Einstein My lab family: Everyone in Dr. Bazinet lab lent their helping hands to my research. I especially would like to thank Shoug Alashmali, who has been a fantastic supportive lab mate and a wonderful friend during my research journey. I would like to thank Vanessa Guiliano, Scott Lacombe, Adam Metherel, Raphael Chounard-Watkins and Maha Irfan Sarah Orr, Chuck Chen, Marco Trepanier, Kathyrn Hopperton Alex Kitson, Lauren Lin, Kayla Hildebrand, for the fun time, supportive research environment and advice to help me become a confident speaker and researcher. I also gave my special thanks to Louisa Kung and Emelia D’Souza, who has given the best administrative support in such to ensure my research journey on time and well-performed. I want to thank my teaching mentor Debbie Gurfinkel, for guiding me to be a better teaching assistant and an instructor to deliver research in the past four years. I want to thank Fiona Wallace, who is a definite hard-working role model for me as my mentor. I would like to thank Kate Banks, Tracy McCook, Nancy Tomas, AJ Wang Warren Foltz, Michael Leadley and Ashley St.Pierre for ensuring my experiments went well. Also, I want to thank many collaborators, including Sophie Laye, Cobol Su, Laura Best, Mathieu Di-Miceli, Mandy H, Ruslan Kubant, Ryan Bradley, Cigdem Sahin, Ivana Prce who shared their knowledge and expertise in the different area of research, which opened my view on conducting research containing diverse programs. In addition, I want to thank Shirley Vien, Diana Sanchez Hernandez, Amel Taibi, Paraskevi Masssara, Kit-Yi Yam and many others who are my colleagues and friends going through this fantastic post-graduate journey together. I want to thank Daniela D’Aniello and my friends, Aleese Smith, Melanie A, Marianthe Maroulis, Tad Ferreira, Gursimran Sethi, Anamika Ray Elie Lee Livia Li, Dobrachina Zubek and others who lived in Knox College and New College. We shared so many wonderful conversations, meals, events, and parties which made our experience in the University of Toronto enjoyable. I want to thank my piano teacher Arsha Nersessian and Ed Jesus and my piano students for sharing our joyful weekly music sections in the past five years V “When I walk along with two others, from at least one I will be able to learn”-Confucius I want to thank my dear friends Alex (Chen) Wang, Yanlin Zhao, Xin Yi, Ray Zhang, Dom Kowk, Linda (Dan) Li, Dongfang Chao, and many others for sharing your wise words, life and work experience with me through our numerous conversations. Life is a continuous line filled with different colors of flowers that we observe together. To my amazing parents, thank you so much for providing mental, emotional and material support to cherish my life. You taught me to how to work hard while balanced, how to respect people and keep smiling even in the face of adversity. Mother, you have a kind heart and a strong spirit to lift me up in any circumstance. Whenever I had doubt, I remember you said: “Don’t be afraid.” Father, you taught me how to be a hard worker with gratitude. You are a great listener and a patient achiever. I learn from you and make you proud. Through my fascinating Ph.D. journey, I inspired by some famous scholars and many ordinary but fantastic people by their perception of life through their experience. I also came up with my own understanding of science and life. These invaluable inspiration quotations are shared in the footnotes of the acknowledgment and reference sections, that opened my mind and heart. VI “There is always a deeper emotion which triggers all the motives seeing in people's ostensible needs.” - Agatha Christie, 《Five Little Pigs》 Financial Disclosure This research was funded by the Natural Sciences and Engineering Research Council of Canada (NSERC). Lin Lin also received scholarships from Ontario Graduate Scholarship, Ontario Student Opportunity Trust Fund and Doctoral Completion Award. VII “Pleasure in the job puts perfection in the work”-Aristotle Contributions Lin Lin contributed to and/or performed all experimental designs, method development, procedures, data collection, analyses, and wrote the first draft, with the exception of the following contributions: Co-authors for submitted / published manuscripts Dr. Adam Metherel - Contributed to statistical analysis and provided feedback on manuscript editions (Chapter 4 & 5). Dr. Alex Kitson - Contributed to database extraction and confirmation (Chapter 2); Data validation (Chapter 4) Dr. Kathryn E Hopperton and Dr. Marc-Olivier Trépanier - Contributed to animal tissue collections and provided feedback on manuscript editions (Chapter 4). Dr. Peter J
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