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The Effect of Steroids on Sexual Differentiation Of THE EFFECT OF STEROIDS ON SEXUAL DIFFERENTIATION OF THE RAT CENTRAL NERVOUS SYSTEM Thesis submitted for the degree of Master of Science in the Faculty of Medicine M.A. TAHER, M.B.B.S. Institute of Physiology University of Glasgow November 1992 C M.A. TAHER, 1992. ProQuest Number: 13815482 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 com plete manuscript and there are missing pages, these will be noted. Also, if material had to be removed, a note will indicate the deletion. uest ProQuest 13815482 Published by ProQuest LLC(2018). Copyright of the Dissertation is held by the Author. All rights reserved. This work is protected against unauthorized copying under Title 17, United States C ode Microform Edition © ProQuest LLC. ProQuest LLC. 789 East Eisenhower Parkway P.O. Box 1346 Ann Arbor, Ml 48106- 1346 GLASGOW \ (UNIVERSITY 1 l ib r a r y 1 IN THE NAME OF ALLAH THE MOST BENEFICENT AND MERCIFUL ACKNOWLEDGEMENTS I have much pleasure in expressing my gratitude to Professor S. Jennett and Professor J.C. McGrath of the Institute of Physiology and to Dr. A.P. Payne, Acting Head of the Anatomy Department, for allowing me to carry out this research work in their Departments. I am most grateful to my supervisors, Dr. D.P. Gilmore and Dr. A.P. Payne for their guidance, encouragement, support, constructive criticism and patience throughout the period of my research and careful reading of this manuscript. I thank all the technical staff for their help during my time in these two Departments. In particular, I am especially indebted to Mr. N. Bennett and Mr. D. Gormal for their assistance with animals, and Mr. J. McGadey and Mr. D. Russell for their excellent technical help. My sincere thanks also to Dr. F. Burton and Dr. R. Orchardson for their helpful advice and comments. Special thanks go to Dr. Heather M. Johnston and Mr. C.H. Brown for their guidance and practical help during the work with the HPLC, and thanks to Mrs. S. Byars for all her helpful assistance. I am grateful to Mrs. Caroline A. Morris and Ms. Margaret Hughes for their expert assistance in photography and art work. I warmly thank Mrs. L. Peedle and Mr. G.A. Gillespie of the Anatomy Department and Mr. R. Campbell of the Institute of Physiology for extending their helping hands during my course. I especially thank Miss Brenda Robinson for her help in typing the manuscript with great patience. Thanks are also due to Dr. N. Islam and Dr. N.A. Latif for their good companionship during the period of my research work. I am especially grateful to my wife Syeda Nasima Khanam and both of my sons Iqbal and Rizwan for their patience and peaceful companionship during my work. With sincere gratitude I wish to acknowledge my parents, parents-in-law, brothers and sisters for their continued encouragement, affection and moral support throughout this period. I would also like to thank W.H.O. and the Ministry of Health and Family Welfare, Bangladesh, for granting me a Fellowship to complete this research. Finally, any one else that I may have missed - thank you. M.A. TAHER ABBREVIATIONS ADR - Adrenaline AFP - Alpha fetoprotein CNS - Central nervous system DA - Dopamine DHBA - Dihydroxy benzylamine DHTB -50C Dihydroxy testosterone benzoate DLN - Dorsolateral nucleus HC1 - Hydrochloric acid 5HIAA - 5 Hydroxy indoleacetic acid HPLC-ECD - High performance liquid chromatography with Electrochemical detection. HRP — Horse radish peroxidase 5HT - 5 Hydroxytryptamine mCPP - m-Chlorophenylpiperaz ine MHPG - 3 methyl 4 hydroxy phenyl glycol NA - Noradrenaline p-CPA - P-chlorophenyl alanine RDLN - Retrodorsolateral nucleus SNB - Spinal nucleus of bulbocavernosus TP - Testosterone propionate VM _ Ventromedial nucleus (iii) CONTENTS Page No. ACKNOWLEDGEMENTS .................................... (i) ABBREVIATIONS ...................................... (iii) SUMMARY ............................................ (xiii) GENERAL INTRODUCTION ................................ 1 1. Introduction ................................. 2 2. Sexual differentiation of central nervous system (CNS) ................................. 5 3. Possible sites of sexual differentiation in the CNS ...................................... 7 4. Sexual dimorphism in central nervous system .. 9 5. Development of spinal cord .................. 11 6. Structure and sex differences in the rat spinal cord .................................. 12 7. Sexually dimorphic nuclei in the spinal cord .......................................... 12 8. Steroidogenesis .............................. 16 9. Mechanism of action of steroid hormone ...... 17 10. Mode of action of steroid hormone in CNS .... 17 11. Brain steroid receptors and their role ...... 19 12. Role of aromatization in the sexual differentiation of the brain ................. 19 13. Oestradiol binding protein (DC- fetoprotein) in the brain and its role .................... 21 14. Serotonin pathway in the brain .............. 21 15. Serotoninergic influence on sexual differentiation of rat brain and sexual behaviour .................................... 23 (iv) Page No. 16. Sex differences in neurotransmitters in the central nervous system ....................... 27 17. Aims of the study ............................ 29 MATERIALS AND METHODS ............................. 30 1. Animals ....................................... 31 2. Experiment 1: Sex differences in motor neuron numbers in the lumbosacral spinal cord..................... 31 3. Experiment 2: Perinatal androgen treatment and its effect on sexually dimorphic spinal motor neurons in the lumbosacral spinal cord ........................... 32 4. Experiment 3: Effects of a non-steroidal anti-androgen (flutamide) on sexually dimorphic spinal motor neurons in male rats ........... 32 5. Experiment 4: Postnatal effects of TP and p-CPA on sexually dimorphic spinal motor neurons ............ 33 6. Experiment 5: The effects of 5HT agonists and antagonists administered postnatally on the sexually dimorphic spinal motor neurons . 33 7. Experiment 6: Sex differences in the concentration of serotonin and 5HIAA in the lumbosacral and thoracic region of the spinal cord in rats ................... 34 8. Removal of the spinal cord .................... 34 9. Sample preparation for motorneuron count ...... 35 10. Sample preparation for measurement of biogenic amines using high performance liquid chromatography with electrochemical detection (HPLC-ECD) ......... 35 11. High performance liquid chromatography with electrochemical detection ..................... 36 12. Statistical Analyses ......................... 45 (v) Page No. RESULTS ............................................ 46 1. Experiment 1: Sex differences in motor neuron numbers in the lumbosacral spinal cord of control male and female rats ................ 47 2. Experiment 2: The effects of perinatal androgen treatment on sexually dimorphic spinal motor neuron groups in male rats ........... 55 3. Experiment 3: Effects of a non-steroidal anti-androgen (flutamide) on sexually dimorphic spinal motor neurons in male rats ..... 55 4. Experiment 4: Postnatal effects of TP and p-CPA on sexually dimorphic spinal motor neuron numbers .... 58 5. Experiment 5: The effects of serotonin agonists and antagonists administered postnatally on the sexually dimorphic spinal motor neurons .................. 62 6. Experiment 6: a) Sex differences in the concentrations of serotonin and 5HIAA in the lumbosacral region of the spinal cord in rats ....... 63 b) Sex differences in the concentrations of indoleamines in the thoracic region of the spinal cord of rats .................. 63 DISCUSSION......................................... 67 1. Sex specific neuronal numbers in the lumbosacral spinal cord sections of the male and female Albino-Swiss rat ................... 67 2. Perinatal androgen treatment and its effects on the sexually dimorphic motor neuron s y s t e m ......................................... 68 3. Postnatal DHTB and its role on sexually dimorphic motor neurons in the SNB and DLN .... 73 (vi) Page No. 4. Postnatal effects of androgens and of the serotonin (5HT) synthesis inhibitor p-CPA on the sexually dimorphic motor neurons of the spinal cord of rats .................... 74 5. Postnatal effects of serotonin agonists and antagonists on sexually dimorphic motor neurons in the male lumbosacral spinal cord ........... 80 6. Biogenic amine levels in the lumcosacral and thoracic regions of the spinal cord in Albino-Swiss rats ............................. 84 7. Mechanisms of control of CNS development by humoral factors such as androgens and amines ......................................... 86 CONCLUSIONS ........................................ 92 BIBLIOGRAPHY ...................................... 93 APPENDIX 1. Statistical analyses of Experiment 4 ........... 124 (vii) LIST OF FIGURES Page No. FIGURE 1 Mammalian sexual differentiation ...... 3 FIGURE 2 Schematic diagram of HPLC apparatus 38 FIGURE 3 Trace produced when 20 jil of a mixture of standards is injected into the HPLC system ................... 42 FIGURE 4 Photograph of a section of spinal cord from an intact male Albino-Swiss rat at level L5 showing the spinal nucleus of bulbocavernosus, dorsolateral nucleus, retrodorsolateral nucleus and ventromedial nucleus ............. 48 FIGURE 5 Photograph of a section of spinal cord from an intact female Albino-Swiss rat at level L5 showing the spinal nucleus of
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