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The Effects of Ovarian Enzyme Modulators on Folliculogenesis and Cyst Development in the Porcine Ovary

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The Effects of Ovarian Enzyme Modulators on Folliculogenesis and Cyst Development in the Porcine Ovary The effects of ovarian enzyme modulators on folliculogenesis and cyst development in the porcine ovary A thesis submitted by Neera Sunak for the degree of Doctor of Philosophy University of London University College London 2 0 0 7 Department of Biochemistry and Molecular Biology Gower Street London WC1E6BT United Kingdom 1 UMI Number: U592428 All rights reserved INFORMATION TO ALL USERS The quality of this reproduction is dependent upon the quality of the copy submitted. In the unlikely event that the author did not send a complete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. Dissertation Publishing UMI U592428 Published by ProQuest LLC 2013. Copyright in the Dissertation held by the Author. Microform Edition © ProQuest LLC. All rights reserved. This work is protected against unauthorized copying under Title 17, United States Code. ProQuest LLC 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106-1346 To my family: Dad, Mum and Renu 2 Abstract The first aim of the research conducted as part of this thesis was to investigate the expression and activities of 11PHSD enzymes in porcine mural granulosa cells and COCs from small, medium and large antral follicles, as well as from ovarian cysts. Both cloned lipHSD enzymes (11 PHSD 1 and 11PHSD2) were expressed in porcine granulosa cells however the results of the expression studies in COCs were inconclusive. In granulosa cells from antral follicles, the 11 p-dehydrogenase (11P-DH) activities of the lipHSD enzymes increased by approximately 3-fold with antral follicle growth (P<0.01). Similarly, COCs isolated from large follicles had approximately 10-fold higher lip-D H activities than COCs from small follicles (PO.OOl). These results suggested that antral follicle growth occurred alongside increasing levels of intracellular cortisol metabolism in both granulosa cells and COCs. In granulosa cells from ovarian cysts, there were significantly decreased net lip-DH activities compared to cells from large antral follicles (P<0.01), suggesting that decreased cortisol metabolism occurred in the cells of porcine ovarian cysts. The next aim of this thesis was to investigate the levels of ovarian enzyme modulators in FF from small, medium and large antral follicles and in the fluid from ovarian cysts. The levels of inhibitors appeared to decrease in FF with follicle growth but were significantly increased in ovarian cysts (PO.Ol). These intrafollicular lipH SD l inhibitors in FF and cyst fluid were also able to modulate lipHSD activities in the granulosa cells and COCs. Furthermore, cyst fluid, and the hydrophobic components thereof (which were likely to have included the ovarian enzyme modulators), were shown to significantly increase the rates of porcine oocyte maturation (PO.OOl). In summary, the findings of the studies conducted in this thesis suggest that the intrafollicular modulators of lipHSDl could influence local cortisol-cortisone inter-conversion in granulosa cells and oocytes during antral follicle growth. In addition the ovarian enzyme inhibitors could possibly influence oocyte maturation with follicle growth. In ovarian cysts however, the high levels of intra-follicular lipH SD l inhibitors in the cyst fluid, and the decreased levels of intracellular cortisol metabolism observed in the granulosa cells, could be factors contributing to cyst development. 3 Acknowledgements Firstly, I would like to acknowledge the financial support I received while doing this PhD, both from the BBSRC and from my CASE sponsor Genus Pic. To my supervisors, Dr. Tony Michael and Dr. Lisa Thurston, you have both given me amazing support and encouragement throughout my years of study with you, and for that I am very grateful. Many thanks to you both for the time you have spent with me teaching me all you can and for the belief you had in me. I am also indebted to Dr. Rachel Webb; chapter 6 would not have been possible without your help and input. To each of my friends, I am so thankful to have all of you, particularly during the last few months of my study which have undoubtedly been the most difficult. Having said that, I am so glad to have been writing up alongside my fellow final year PhD students, Nana and Abdul. You have provided me so much entertainment and help, and you have really spurred me on to finish my writing. I am glad that your thesis writing has gone really well, you both deserve to go on and do great things after getting your PhDs! To Kim and Vicky, my former lab partners, I am so grateful for your advice and words of encouragements. You have always been there when I needed someone to talk to. I hope that we will always share our love for music festivals and that we keep up the traditions of attending one every year! I would also like to thank my other friends, in and around London, for their patience when I have been at my most busy, I now have time to spend with each and every one of you and that is exactly what I intend to do! Finally, to my family: I owe you so much for the past few years, you have truly provided me with amazing amounts of encouragement and belief (and financial backing!!). Mum and dad, our chats have really helped me to keep a clear head and ease my stress and Renu, I now have so much more time to spend in the flat with you! During my study time you have always been there with your love and support and I have done this for you as much as for myself. I dedicate this to you. 4 Table of Contents Abstract ................................................................................................................................3 Acknowledgements .............................................................................................................4 Table of Contents................................................................................................................ 5 List of Figures..................................................................................................................... 9 List of Tables.....................................................................................................................12 Publications........................................................................................................................13 Abbreviations .....................................................................................................................15 Chapter 1 General Introduction........................................................................................20 1.1 The Ovary ................................................................................................................21 1.1.1 Overview of oogenesis.................................................................................. 21 1.1.2 The porcine oestrous cycle............................................................................22 1.2 The Endocrine Regulation of Folliculogenesis................................................... 27 1.2.1 The “two-cell-two-gonadotrophin” model.................................................. 28 1.2.2 Oocyte maturation and the role of LH.........................................................30 1.2.3 Follicular atresia.............................................................................................33 1.2.4 Abnormalities of folliculogenesis - ovarian cyst development.................34 1.3 The Adrenal Glands and HPA Axis ..................................................................... 36 1.3.1 Morphology of the adrenal glands and steroidogenesis............................. 36 1.3.2 Production of cortisol by the zona fasciculata............................................ 36 1.3.3 Effects of cortisol...........................................................................................38 1.3.4 Metabolism of cortisol..................................................................................42 1.4 11 p-Hydroxysteroid Dehydrogenase (1 lpHSD) ................................................. 42 1.4.1 Genes encoding the 11 pHSD enzymes ....................................................... 42 1.4.2 Biochemistry and molecular biology of 1 lpHSD actions.........................43 1.4.3 Sites of expression for the 1 lpHSD isoforms.............................................47 1.4.4 Physiological functions of the 11 PHSD enzymes.......................................49 1.4.5 Regulation of 1 lpHSD activity by enzyme inhibitors ................................52 1.4.6 The importance of 11 pHSD in the ovary.....................................................55 1.4.7 Regulation of 11 PHSD activity in the ovary............................................... 59 Chapter 2 General Materials and Methods .................................................................... 63 2.1 Chemicals and reagents ........................................................................................64 5 2.2 Collection of porcine ovarian samples................................................................. 65 2.3 Aspiration and storage of porcine ovarian fluids................................................ 65 2.4 Reverse phase Cl 8 column chromatography of porcine ovarian fluids.............66 2.5 Isolation of porcine granulosa cells.......................................................................67 2.5.1 RT-PCR in porcine granulosa cells.............................................................. 67 2.5.2 11 PHSD dehydrogenase and reductase activities in porcine granulosa
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