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Behavioural and Neural Correlates of Individual Differences in Episodic Behavioural and Neural Correlates of Individual Differences in Episodic Memory by Daniela Jesseca Palombo A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Department of Psychology University of Toronto © Copyright by Daniela Jesseca Palombo, 2013 Behavioural and Neural Correlates of Individual Differences in Episodic Memory Daniela Jesseca Palombo Doctor of Philosophy Department of Psychology University of Toronto 2013 Abstract Episodic autobiographical memory (AM) refers to the real-life recollection of personal events that are contextually-bound to a particular time and place. Anecdotally, individuals differ widely in their ability to remember these types of experiences, yet little cognitive neuroscience research exists to support this idea. By contrast, there is a growing body of literature demonstrating that individual differences in episodic memory for laboratory experiences, intended to serve as a proxy for real life, are associated with brain-biomarkers. The present studies provide a starting point for exploring individual differences in the real-life expression of memory; which is more complex, multifaceted and has longer retention intervals, than laboratory memory (LM), thus allowing for the assessment of remote memory. While there are many factors that contribute to individual differences in episodic AM, the focus of this dissertation is on genetic influences. In particular, the KIBRA gene has been associated with episodic LM in a replicated genome- wide association study; T-carriers showing enhanced performance relative to individuals who lack this allele. The present series of studies explored the association of KIBRA with ii episodic AM. An ancillary goal was to clarify the association between KIBRA and episodic LM. Accordingly, in Chapter 2, episodic LM and AM were probed in a large sample of healthy adults in relation to KIBRA genotype. Next, in a subset of participants, functional and structural magnetic resonance imaging (MRI) were used to explore KIBRA-dependent differences in hippocampal and medial temporal lobe (MTL) blood- oxygen-level-dependent (BOLD) response (Chapter 3) and structure (Chapter 4). Akin to prior research, Chapter 2 showed that KIBRA is associated with episodic LM, albeit not on all measures. Moreover, a KIBRA association was observed for remote aspects of episodic AM on some measures. Chapter 3 showed mainly greater hippocampal and MTL cortex BOLD response in T-carriers relative to non-carriers during the recollection of both LM and AM. Moreover, during episodic AM, T-carriers showed a distinct pattern of hippocampal-neocortical connectivity. Finally, Chapter 4 showed that T-carriers have a larger cornu ammonis and dentate gyrus relative to non-carriers, suggesting a potential neural locus for the effects of KIBRA on episodic memory. iii Acknowledgments Writing a thesis is an extremely challenging process. When I doubted myself, I found strength in all the people who supported me throughout this journey. At the very top of this list was my supervisor, Dr. Brian Levine. Before I began graduate school, I knew that my future supervisor was a brilliant and accomplished scientist. But it was only during graduate school that I discovered his warmth, patience and depth as a mentor. Brian has been fundamental in shaping my academic development and instilling confidence in me. I am extremely proud of the relationship I have maintained with Brian over the last few years and I am grateful to have had an advisor who cared so much about my success (and cautioned me repeatedly to get more sleep and eat less junk food). I would next like to thank my thesis committee members, Drs. Tomáš Paus and Morris Moscovitch, for not only inspiring me with their passion for science, but for their constant guidance, encouragement, and theoretical contributions to the work presented here. I am also grateful to my external committee members, Drs. Mary Pat McAndrews and Morgan Barense for their constructive feedback on my thesis as well as my Appraiser, Dr. Donna Rose Addis for her useful and informative comments. Next, I would like to thank my DNA Affect and Memory Project (DAMP) collaborators, Drs. Rebecca Todd, Adam Anderson, Daniel Müller, as well as Natalie Freeman, who were instrumental in the inception of these projects. Rebecca has not only been a mentor to me but also a friend who always puts a smile on my face. Robert Amaral and Rosanna Olsen (fondly remembered as “the work triangle”) also made a significant contribution to the work presented here. I extend special thanks to Dr. Malcolm Binns (“the stats whisperer”) for statistical consulting. I will never forget the bonds I formed in the magical land of “Levinia.” I am extremely grateful to its members, both current and alumni, and affiliated Rotman colleagues, who have assisted me in many capacities along this journey: Anjali Beharelle, Namita Kumar, Wayne Khuu, Aggie Bacopulos, Priya Kumar, Amanda Robertson, Louis Renoult, Mike Armson, Marjorie Green (“the lab mom”), Carrie Esopenko, Signy Sheldon, Leann Lapp, iv Patty Hsu, Alice Kim, and Ryan Gosling (our fictional lab member). In particular, I am especially grateful to one of my first lab friends, and now one of my dearest friends, Charlene O’Connor, who has always been there for me for every academic milestone I faced, and who represents the better half of “superdesk,” an enchanted place where we spent countless hours working, snacking, and laughing. Before I began graduate school, I had the fortunate opportunity to work in the developmental psychobiology laboratory of Drs. Alison Fleming and Gary Kraemer. Under their mentorship, I successfully complete an undergraduate thesis, where I studied maternal behaviour in rats. My time in their laboratory was short, but the lessons I learned there stood the test of time. I thank both of them for preparing me well for the (long) road ahead and, in particular, for providing me with an appreciation and understanding of the fundamentals of research in non-human species. Next, I would like to thank my family, my parents, Paola and Tony Palombo for love, support, encouragement, and my amazing sisters, Jolene Conti and Sandra Palombo, for always being there for me and helping me get through graduate school-related stress (and for reminding me about the importance of fashion even on a graduate student budget). I thank my life-long friend, Cristina Bianco, for laminating all of my special Ph.D-related achievements on colourful construction paper (she is a grade school teacher); my dear friend, Marlene Schmidt, for teaching me to take life “one cookie at a time”-advice I took very seriously; and her husband, my dear friend, Alvaro De La Torre, for teaching me drive and determination. Finally, I would like to thank my husband (who I married less than three weeks before defending my thesis), and the love of my life, Juan Pablo de la Torre, for patience, support, laughter, and inspiration - and for teaching me computer programming and helping me format my thesis. J.P. has always encouraged me to follow my dreams and has shown me that wherever I go, he will be there by my side. I look forward to our next chapter in life - the Boston Chapter! v Table of Contents Acknowledgments iv Table of Contents vi List of Tables viii List of Figures x List of Acronyms xiv Chapter 1 General Introduction 1 1.1 What is Episodic Memory? 3 1.2 Methods of Investigation 6 1.3 The Extreme Case Approach 8 1.4 Neural Correlates of Individual Differences in Episodic LM: Neuroimaging and Genetics 11 1.4.1 Structural Imaging 11 1.4.2 Functional Imaging 13 1.4.3 Genetics 14 1.5 Individual Differences in AM 16 1.5.1 Personality and AM 17 1.5.2 Summary and Future Directions 19 Chapter 2 The effects of KIBRA Polymorphism on Episodic AM 21 2 Introduction 21 2.1 Method 25 2.1.1 Participants 25 2.1.2 Genotyping 26 2.1.3 Data Analyses 31 2.2 Results 31 2.2.2 Study 1: Episodic LM 32 2.2.3 Study 2: Episodic AM 33 2.3 Discussion 35 Chapter 3 The effects of KIBRA Polymorphism on BOLD Response during Episodic AM 40 vi 3 Introduction 40 3.1 Method 43 3.1.1 Participants 43 3.1.2 Genotyping 44 3.1.3 Image Acquisition 44 3.1.4 Scanner Task 1: Recognition LM for Faces 45 3.1.5 Scanner Task 2: Episodic AM 46 3.1.6 Data Analysis 48 3.2 Results 52 3.2.1 In Scanner Behavioral Responses 52 3.2.2 Scanner Task 1: Recognition LM for Faces 52 3.2.3 Scanner Task 2: Episodic AM 53 3.2.4 Univariate Imaging Analysis 53 3.2.5 Multivariate Imaging Analysis 55 3.3 Discussion 56 Chapter 4 The effects of KIBRA Polymorphism on Hippocampal Sub-regions using High-resolution MRI 62 4 Introduction 62 4.1 Method 63 4.1.1 Participants 63 4.1.2 Genotyping 64 4.1.3 MRI Acquisition 64 4.1.4 Segmentation 64 4.2 Results 65 4.3 Discussion 66 Chapter 5 General Discussion 70 References 78 Tables and Figures 100 vii List of Tables Table 1. Number of participants is shown for each KIBRA genotype group (CC and TT/TC) for each of the tasks used in Chapter 2 (IAPS; International Affective Picture System). Table 2. Means and standard deviations are shown for each KIBRA genotype group (CC and TT/TC) for the objects task (FA; false alarms) used in Chapter 2. Table 3. Means and standard deviations are shown for each KIBRA genotype group (CC and TT/TC) for the International Affective Picture System (IAPS) task used in Chapter 2 (FA; false alarms). Table 4. Means and standard deviations are shown for each KIBRA genotype group (CC and TT/TC) for each of the scanner tasks used in Chapter 3 (FA; false alarms).
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