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«HIPPOCAMPAL OVERSHADOWING: EXPLORING THE UNDERLYING MECHANISMS TINE LANDEHAGEN GULBRANDSEN Bachelor of Science Kinesiology, University of Lethbridge, ...»

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HIPPOCAMPAL OVERSHADOWING: EXPLORING THE UNDERLYING MECHANISMS

TINE LANDEHAGEN GULBRANDSEN

Bachelor of Science Kinesiology, University of Lethbridge, 2009

A Thesis

Submitted to the School of Graduate Studies

Of the University of Lethbridge

in Partial Fulfilment of the

Requirements of the Degree

DOCTOR OF PHILOSOPHY

Department of Neuroscience University of Lethbridge

LETHBRIDGE, ALBERTA, CANADA

© Tine Landehagen Gulbrandsen, 2015

HIPPOCAMPAL OVERSHADOWING: EXPLORING THE UNDERLYING MECHANISMS

TINE LANDEHAGEN GULBRANDSEN

Date of Defense: Monday August 10th 2015 Dr. Robert J. Sutherland Professor Ph.D.

Supervisor Dr. Robert J. McDonald Professor Ph.D.

Thesis Examination Committee Member Dr. Igor P. Kovalchuk Professor Ph.D.

Thesis Examination Committee Member Dr. Majid Mohajerani Assistant Professor Ph.D.

Internal Examiner Dr. Jill Leutgeb Professor Ph.D.

External Examiner University of California San Diego, California Dr. Artur Luczak Associate Professor Ph.D.

Chair, Thesis Examination Committee ii ABSTRACT The aim of the current thesis was to evaluate the underlying mechanisms of hippocampal (HPC) overshadowing. One theory to explain this phenomenon predicts that perceptual representation of an event is stored in non-HPC cortical areas. Following multiple distributed exposure to the same or similar event, this representation becomes stable and the cellular activation patterns can be recalled in the absence of HPC input.

Through a series of experiments, the hypothesis that the dysgraunlar layer of the retrosplenial cortex (RSD) fulfill these requirements was tested. Using double in-situ hybridization for the mRNA of immediate-early genes Arc and Homer1a, the cellular activation patterns in HPC subregion CA1 and RSD were evaluated. In addition, temporary inactivation of HPC was used to evaluate cellular activation patterns of a HPC-independent memory in RSD. The results presented here support the idea that cellular activation patterns for may be stored – at least in part – in the RSD.

ACKNOWLEDGEMENTS

As with any major life-event, the road to getting this thesis submitted has been a long and bumpy one. I have been passionate about this project since I first suggested it to my supervisor, Dr. Robert Sutherland, almost four years ago. His guidance has been invaluable in forming a comprehensive and cohesive theory on how to best describe the phenomenon I was so fascinated about. The support of good family, friends and co-workers has made the completion of this thesis possible. I would first of all like to thank my family and my parents for all their support during the last 10 years, all on which I have spent at a university on the other side of the world from them. Secondly I would like to thank Jason Getty for all his help on conceptual thinking, keeping me motivated, and keeping me strong during the very tough last 3 months of this thesis. From the Canadian Centre for Behavioural Neuroscience I would like to thank everyone who has helped me in one way or another the last 6 years, either it be as a friend, motivator, challenger, or support. Among these I would like to especially mention Robin Keeley, Heather Bell, Erin Zelinksi, Khadaryna Hernandez, Aubrey Demchuk, Valerie Lapointe, Karen Dow-Cazal, Isabelle Gauthier, Kelsey O’Brien, Naomi Cramer, and Amanda Mauthe-Kaddoura.

Without the guidance of my supervisor, Dr. Robert J. Sutherland, and my supervisory committee, Dr. Robert J. McDonald and Dr. Igor Kovalchuk, this work would have never happened. I would like to thank all of you for serving on my committee and for guiding me through this. I especially want to thank Dr. Sutherland who let me stay in his lab after completing my M.Sc. in 2011, and to let me continue to learn from him these last four yearsit has been a pleasure! Lastly, I would like to thank my thesis examination committee, Dr.

Majid Mohajerani, who I have had the pleasure of working with the last year and from whom I have already learned a lot, and Dr. Jill Leutgeb, whose work has inspired the fundamentals of this thesis. I have admired her work throughout my graduate career, and having her travel all the way from San Diego to serve on my committee is a true honour.

There are many more who have helped me through my graduate career, and there are many that deserve far greater thanks than just a mention in the text above. However, in order to prevent writing up and index of CCBN staff – past and present – I will leave it as is.

Please know that I have not forgotten about any of you and I am truly grateful for all the people affiliated with this building and all the ways they have contributed to me getting where I am today.

Thank you.

–  –  –

ABSTRACT………………………………………………………………………………………………………. iii ACKNOWLEDGEMENTS…………………………………………………………………………………… iv LIST OF TABLES………………………………………………………………………………………………. vii LIST OF FIGURES……………………………………………………………………………………….…….. viii LIST OF ABBREVIATIONS………………………………………………………………………………… ix

CHAPTER ONE: GENERAL INTRODUCTION





General introduction……………………………………………………………………………… 1 Theories of hippocampal overshadowing………………………………………….......... 2 The hippocampus and retrosplenial cortex……………………………………………… 5 Immediate Early Genes…………………………………………………………………………… 8 The goal of the current thesis………………………………………………………………….. 10

CHAPTER TWO: EXPLORATORY BEHAVIOUR OF THE RATTUS NORVEGICUS

Introduction…………………………………………………………………………………………... 13 Materials and Methods…………………………………………………………………………… 14 Results…………………………………………………………………………………………………… 16 Discussion……………………………………………………………………………………………… 17

CHAPTER THREE: TEMPORARY INACTIVATOIN OF THE RODENT HIPPOCAMPUS USING

MUSCIMOL Introduction…………………………………………………………………………………………… 20 Materials and Methods……………………………………………………………………………. 21 Results…………………………………………………………………………………………………… 25 Discussion……………………………………………………………………………………………… 26

CHAPTER FOUR: CONTEXT DISCRIMINATION IN CA1 AS MEASURED BY IMMEDIATE

EARLY GENE ACTIVATOIN

Introduction…………………………………………………………………………………………… 27 Materials and Methods……………………………………………………………………………. 29 Results……………………………………………………………………………………………………. 31 Discussion………………………………………………………………………………………………. 36

CHAPTER FIVE: CONTEXT DISCRIMINATION IN RETROSPLENIAL CORTEX AS

INDICATED BY IMMEDIATE EARLY GENE ACTIVATION

Introduction……………………………………………………………………………………………. 41 Materials and Methods……………………………………………………………………………. 42 Results……………………………………………………………………………………………………. 42 Discussion………………………………………………………………………………………………. 45

CHAPTER SIX: CONTEXT SPECIFIC ACTIVATION IN RETROSPLENIAL CORTEX

FOLLOWING HIPPOCAMPAL INACTIVATION AS MEASURED BY IMMEDIATE EARLY

GENE ACTIVITY

Introduction………………………………………………………………………………………….. 47 Materials and Methods…………………………………………………………………………… 49 Results………………………………………………………………………………………………….. 50 Discussion……………………………………………………………………………………………… 55

CHAPTER SEVEN: GENERAL DISCUSSION

Summary of the thesis……………………………………………………………………………. 58 Further considerations…………………………………………………………………………… 60 Final conclusion……………………………………………………………………………………… 65 REFERENCES….………………………………………………………………………………………..………... 68 TABLES……………………………………………………………………………………………………………… 81 FIGURES…………………………………………………………………………………………………………….. 83

LIST OF TABLES

2.1 An outline of the training procedures for each of the behavioural groups………..…… 81

3.1 Surgical coordinates………………………………………………………………………..………………… 81

4.1 Training and testing paradigm…………………………………………………………………………... 81

6.1 Behavioural protocol for training and testing day…………………………………………….… 82

LIST OF FIGURES

2.1 Depictions of testing room 1 and 2…………………………………………………………………….. 83

2.2 The total amount of rearing on each training day, independent of group or context...……………………………………………………………………………………………………………….……… 83

2.3 Total amount of rearing on each training day in the dark vs. the light room…………. 84

2.4 Rearing behaviour by group……………………………………………………………………….……… 84

2.5 Rearing behaviour on group AA within each session across training days……..…….. 85

3.1 Arc mRNA labeling in HPC at different time-points following muscimol infusion.… 85

3.2 Pictures of dorsal HPC in control (A) and inactivated (B) hemisphere 10 min following muscimol infusion……………………………………………………………………………… 86

4.1 IEG labeling in CA1 following 3 days of habituation…………………………………………..… 86

4.2 IEG labeling in left and right CA1 following 3 days of habituation………………………... 87

4.3 IEG labeling in dorsal and ventral CA1 following 3 days of habituation………………... 88

4.4 IEG labeling in CA1 following 12 days of habituation…………………………………………... 89

4.5 Pictures of dorsal CA1 in group AA (A) and AB (B) following 12 days of habituation ………………………………………………………………………………………………………………………… 89

4.6 IEG labeling in left and right CA1 following 12 days of habituation…………………….... 90

4.7 IEG labeling in dorsal and ventral CA1 following 12 days of habituation ………….….. 90

4.8 Proportion of cells labeled with IEGs in CA1 after 3 and 12 days of habituation…… 91

4.9 Proportion of IEG labeled with cells in CA1 that are co-labeled after 3 and 12 days of habituation…………………………………………………………...………………………………………. 91

5.1 Maps of RSD as described by Wyss and van Groen (1992)………………………………...… 92

5.2 IEG labeling in RSD following 3 days of habituation……………………………………………. 92

5.3 IEG labeling in RSD after 12 days of habituation………………………………………………..... 93

5.4 Proportion of cells labeled with IEGs in RSD after 3 and 12 days of exploration…… 93

5.5 Proportion of IEG labeled cells in RSD that are co-labeled after 3 and 12 days of habituation……………………………………………………………………………………………………..… 94

5.6 Correlation between IEG labeling in CA1 and RSD after 3 days of exploration…….... 94

5.7 Correlation between IEG labeling in CA1 and RSD after 12 days of exploration……. 95

6.1 Total rearing in all groups……………………………………………………………………………….… 95

6.2 Rearing in first and second context exploration………………………………………………..… 96

6.3 HPC Arc labeling following muscimol infusion……………………………………………………. 96

6.4 IEG labeling in group AA and AxA………………………………………………………………….…… 97

6.5 Correlation between HPC Arc and cells labeling Arc alone in RSD in group AxA….... 97

6.6 IEG labeling in group AB and AxB………………………………………………………………………. 98



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