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The Role of Bdnf in the Survival and Morphological THE ROLE OF BDNF IN THE SURVIVAL AND MORPHOLOGICAL DEVELOPMENT OF ADULT-BORN OLFACTORY NEURONS by Brittnee McDole A Dissertation Submitted to the Faculty of The Charles E. Schmidt College of Science In Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy Florida Atlantic University Boca Raton, FL December 2018 Copyright 2018 by Brittnee McDole ii ACKNOWLEDGEMENTS I would like to thank my committee members, Dr. Ken Dawson-Scully, Dr. Ceylan Isgor, and Dr. Robert W. Stackman for their guidance and invaluable input in the preparation of this dissertation. I would also like to acknowledge my adviser, Dr. Kathleen Guthrie for her mentorship and support through graduate school. Thank you! I would like to thank all my past and current colleagues in our lab for always being willing to help, and especially Crystal Gilkes for her countless attendance at 6am surgeries and her selflessness in providing me with any help I needed that allowed me to complete my project. I am forever in your debt! I would like to thank my family and friends for their overwhelming support and understanding. I would especially like to acknowledge my mother, thank you for believing in me and providing me with the strength and encouragement I needed to succeed. I would like to thank the National Institutes of Health for the support provided by grant NIH: DC012425 iv ABSTRACT Author: Brittnee McDole Title: The role of BDNF in the survival and morphological development of adult-born olfactory neurons Institution: Florida Atlantic University Dissertation Advisor: Dr. Kathleen Guthrie Degree: Doctor of Philosophy Year: 2018 Olfactory Granule cells (GCs) are a population of inhibitory interneurons responsible for maintaining normal olfactory bulb (OB) function and circuitry. Through dendrodendritic synapses with the OBs projection neurons, the GCs regulate information sent to the olfactory cortices. Throughout adulthood, GCs continue to integrate into the OB and contribute to olfactory circuitry. However, only ~50% will integrate and survive long- term. Factors aiding in the survival and morphological development of these neurons are still being explored. The neurotrophin brain-derived neurotrophic factor (BDNF) aids in the survival and dendritic spine maturation/maintenance in several populations of CNS neurons. Investigators show that increasing BDNF in the adult-rodent SVZ stimulates proliferation and increases numbers of new OB GCs. However, attempts to replicate these experiments failed to find that BDNF affects proliferation or survival of adult-born granule cells (abGCs). BDNFs regulation of dendritic spines in the CNS is well characterized. In the OB, absence of BDNF’s receptor on abGCs hinders normal spine development and v demonstrates a role for BDNF /TrkB signaling in abGCs development. In this study, we use transgenic mice over-expressing endogenous BDNF in the OB (TgBDNF) to determine how sustained increased in BDNF affect the morphology of olfactory GCs and the survival and development of abGCs. Using protein assays, we discovered that TgBDNF mice have higher BDNF protein levels in their OB. We employed a Golgi-cox staining technique to show that increased BDNF expression leads to an increase in dendritic spines, mainly the mature, headed-type spine on OB GCs. With cell birth-dating using 5-bromo-2’- deoxyuridine (BrdU), immunofluorescent cell markers, TUNEL staining and confocal microscopy, we demonstrate that over-expression of BDNF in the OB does not increase survival of abGCs or reduce cell death in the GC population. Using virally labeled abGCs, we concluded that abGCs in TgBDNF mice had similar integration patterns compared to wild-type (WT) mice, but maintained increases in apical headed-type spine density from 12 to 60 days PI. The evidence combined demonstrates that although increased BDNF does not promote cell survival, BDNF modifies GC morphology and abGC development through its regulation of dendritic spine development, maturation and maintenance in vivo. vi DEDICATION This manuscript is dedicated to my best-friend and mother, Pamela McDole. THE ROLE OF BDNF IN THE SURVIVAL AND MORPHOLOGICAL DEVELOPMENT OF ADULT-BORN OLFACTORY NEURONS FIGURES …………………………………………………………………………..xiii ABBREVIATIONS .......................................................................................................... xv PART I: INTRODUCTION ................................................................................................ 1 1.1 Olfactory Bulb Anatomy and Circuitry .................................................................. 1 1.2 Adult Neurogenesis ................................................................................................. 4 1.2.1 Subventricular zone ....................................................................................... 5 1.3 Integration of adult born olfactory granule cells..................................................... 6 1.4 Neurotrophic Factors .......................................................................................... 10 1.4.1 Brain-Derived Neurotrophic Factor in the CNS .......................................... 11 1.5 BDNF’s role in adult neurogenesis ....................................................................... 14 1.7 BDNF’s role in dendrite and spine morphology ................................................... 16 1.9 Mouse background ................................................................................................ 19 1.8 Dissertation aims and objectives ........................................................................... 19 1.9.1 Chapter 2 ...................................................................................................... 20 1.9.2 Chapter 3 ...................................................................................................... 21 1.9.3 Chapter 4 ...................................................................................................... 21 viii PART II: BDNF OVER-EXPRESSION INCREASES OLFACTORY BULB GRANULE CELL DENDRITIC SPINE IN VIVO .......................................................... 23 2.1 Abstract ................................................................................................................. 23 2.2 Introduction ........................................................................................................... 24 2.3 Materials Methods ................................................................................................ 28 2.3.1 Animals ........................................................................................................ 28 2.3.2 BDNF Enzyme-Linked Immunosorbent Assay (ELISA) ............................ 29 2.3.3 Western blotting ........................................................................................... 30 2.3.4 Golgi–Cox staining ...................................................................................... 31 2.3.5 Analyses of dendritic morphology ............................................................... 32 2.3.6 Analyses of dendritic spines ........................................................................ 34 2.4 Results ................................................................................................................... 35 2.4.1 BDNF protein levels are increased in the transgenic olfactory bulb ........... 35 2.4.2 GC dendritic lengths and branch complexity ............................................... 36 2.4.3 Over-expression of BDNF by GCs increases their dendritic spine number and density ................................................................................................... 37 2.4.4 BDNF-mediated changes in spine density occur in proximal and distal dendritic domains ......................................................................................... 38 2.5 Discussion ............................................................................................................. 39 PART III: OVER-EXPRESSION OF BDNF IN THE OLFACTORY BULB DOES NOT INCREASE ADULT-BORN GRANULE CELL SURVIVAL .............................. 46 ix 3.1 Abstract ................................................................................................................. 46 3.2 Introduction ........................................................................................................... 47 3.3 Materials and methods .......................................................................................... 50 3.3.1 Animals ........................................................................................................ 50 3.3.2 BrdU injections ............................................................................................ 51 3.3.3 Immunofluorescence .................................................................................... 52 3.3.4 Imaging ........................................................................................................ 52 3.3.5 Terminal deoxynucelotidyl transferase dUTP nick end labeling (TUNEL) ...................................................................................................... 53 3.4 Results ................................................................................................................... 54 3.5 Discussion ............................................................................................................. 55 PART IV- VIRAL-MEDIATED LABELING AND MORPHOMETRIC ANALYSIS OF ADULT-BORN GRANULE CELLS IN TGBDNF MICE .......................................
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