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Biological Role of Conceptus Derived Factors During Early Pregnancy In Biological Role of Conceptus Derived Factors During Early Pregnancy in Ruminants A dissertation submitted in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY IN ANIMAL SCIENCES UNIVERSITY OF MISSOURI- COLUMBIA Division of Animal Science By KELSEY BROOKS Dr. Thomas Spencer, Dissertation Supervisor August 2016 The undersigned have examined the dissertation entitled, BIOLOGICAL ROLE OF CONCEPTUS DERIVED FACTORS DURING EARLY PREGNANCY IN RUMINANTS presented by Kelsey Brooks, a candidate for the degree of doctor of philosophy, and hereby certify that, in their opinion, it is worthy of acceptance. __________________________________ Chair, Dr. Thomas Spencer ___________________________________ Dr. Rodney Geisert ___________________________________ Dr. Randall Prather ___________________________________ Dr. Laura Schulz ACKNOWLEDGMENTS I would like to acknowledge all the students, faculty and staff at Washington State University and the University of Missouri for their help and support throughout my doctoral program. I am grateful for the opportunity to work with Dr. Thomas Spencer, and thank him for his input and guidance not only in planning experiments and completing projects but for helping me turn my love of science into a career in research. I would also like to acknowledge the members of my graduate committee at Washington State University for their help and input during the first 3 years of my studies. A special thanks to Dr. Jim Pru and Cindy Pru for providing unlimited entertainment, and the occasional missing reagent. Thank you to my committee members at the University of Missouri for adopting me late in my program and helping shape my future as an independent scientist. Thanks are also extended to members of the Prather lab and Wells lab for letting me in on the secrets of success using the CRISPR/Cas9 system. None of this work would have been possible without support from my fellow graduate students in the Spencer lab. In particular, I thank Greg Burns for his unwavering support throughout our time together as Team Uterus. A million thank yous to the current Spencer lab; Andrew Kelleher, Joao Moraes, Wang Peng, and Eleanore O’Neil. As well as past members; Carolyn Allen, Piotr Dorniak and Justyna Filant. Finally, special thanks to my family and friends for understanding that in the fall, the sheep always come first. ii TABLE OF CONTENTS ACKNOWLEDGMENTS ...................................................................................... ii LIST OF FIGURES AND TABLES ...................................................................... vi ABSTRACT ......................................................................................................... xi SECTION I: .......................................................................................................... 1 LITERATURE REVIEW ........................................................................................ 1 Early Pregnancy ........................................................................................................ 1 Uterine Histoarchitecture .......................................................................................... 1 Conceptus Development .......................................................................................... 4 Phases of implantation in sheep ............................................................................... 8 Progesterone ............................................................................................................13 Actions of Progesterone on the Endometrium ........................................................ 13 Interferon Tau ...........................................................................................................16 Actions of IFNT on the Endometrium ...................................................................... 16 Interferon Tau (IFNT) .............................................................................................. 18 Pregnancy Recognition in Ruminants ..................................................................... 18 IFNT stimulated genes in the endometrium ............................................................ 19 Effects of IFNT on the LE/sGE ............................................................................... 20 Prostaglandins .........................................................................................................22 Prostaglandin synthesis .......................................................................................... 22 Prostaglandin Receptors ........................................................................................ 25 PPAR mechanisms of action .................................................................................. 25 Peroxisome Proliferator Activator Gamma .............................................................. 29 Peroxisome Proliferator Activator Delta .................................................................. 29 Glucocorticoids........................................................................................................33 Glucocorticoid Synthesis ........................................................................................ 33 GC interconversion by hydroxysteroid (11-beta) dehydrogenases .......................... 34 Structure and function of GR .................................................................................. 37 GCs during pregnancy ............................................................................................ 40 Gene Knockdown Approaches ...............................................................................41 Morpholino antisense oligonucleotides (MAO) ........................................................ 41 iii Short interfering RNA (siRNA) ................................................................................ 45 Clustered Regularly Interspaced Short Palindromic Repeats (CRISPRs) ............... 50 SECTION II: ....................................................................................................... 57 Biological roles of interferon tau (IFNT) and Type I IFN receptors in elongation of the ovine conceptus ................................................................. 57 ABSTRACT ............................................................................................................ 58 INTRODUCTION .................................................................................................... 59 MATERIALS AND METHODS ................................................................................ 61 RESULTS ............................................................................................................... 72 DISCUSSION ......................................................................................................... 81 ACKNOWLEDGMENTS ......................................................................................... 89 SECTION III: ...................................................................................................... 90 Peroxisome proliferator activator receptor gamma (PPARG) regulates conceptus elongation in sheep ....................................................................... 90 ABSTRACT ............................................................................................................ 91 INTRODUCTION .................................................................................................... 92 MATERIALS AND METHODS ................................................................................ 95 RESULTS ............................................................................................................. 106 DISCUSSION ....................................................................................................... 123 ACKNOWLEDGMENTS ....................................................................................... 128 SECTION IV: .................................................................................................... 130 Biological Roles of Hydroxysteroid (11-Beta) Dehydrogenase 1 (HSD11B1), HSD11B2, and Glucocorticoid Receptor (NR3C1) in Sheep Conceptus Elongation ....................................................................................................... 130 ABSTRACT .......................................................................................................... 131 INTRODUCTION .................................................................................................. 132 MATERIALS AND METHODS .............................................................................. 134 RESULTS ............................................................................................................. 146 DISCUSSION ....................................................................................................... 161 ACKNOWLEDGMENTS ....................................................................................... 166 SECTION V: ..................................................................................................... 167 iv Lentiviral targeting of PTGS2 in the ovine conceptus trophectoderm ...... 167 ABSTRACT .......................................................................................................... 168 INTRODUCTION .................................................................................................. 169 MATERIALS AND METHODS .............................................................................. 171 RESULTS ............................................................................................................
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