RECENT PROGRESS IN HORMONE RESEARCH Edited by ANTHONY R. MEANS VOLUME 57 Reproductive Hormones & Human Health “Recent Progress in Hormone Research” is an annual publication of The Endocrine Society that is published under the editorial auspices of Endocrine Reviews. The Endocrine Society 4350 East West Highway, Suite 500 Bethesda, Maryland 20814 Copyright 2002 by The Endocrine Society All Rights Reserved The reproduction or utilization of this work in any form or in any electronic, mechanical, or other means, now known or hereafter invented, including photocopying or recording and in any information storage or retrieval system, is forbidden, except as may be expressly permitted by the 1976 Copyright Act or by permission of the publisher. ISBN 0–879225-45-X CONTENTS Senior Author Correspondence Information v 1. Cellular and Molecular Pathways Regulating Mammalian Sex Determination 1 Christopher Tilmann and Blanche Capel 2. Steroidogenic Factor 1: An Essential Mediator of Endocrine Development 19 Keith L. Parker, Douglas A. Rice, Deepak S. Lala, Yayoi Ikeda, Xunrong Luo, Margaret Wong, Marit Bakke, Liping Zhao, Claudia Frigeri, Neil A. Hanley, Nancy Stallings, and Bernard P. Schimmer 3. Molecular Basis of Mammalian Gamete Binding 37 David J. Miller, Xudong Shi, and Heather Burkin 4. Regulation of the Mitotic and Meiotic Cell Cycles in the Male Germ Line 75 Debra J. Wolgemuth, Erika Laurion, and Karen M. Lele 5. Male Germ Cell Gene Expression 103 Edward M. Eddy 6. The Expression of the Follicle-stimulating Hormone Receptor in Spermatogenesis 129 Leslie L. Heckert and Michael D. Griswold 7. Identification of Specific Sites of Hormonal Regulation in Spermatogenesis in Rats, Monkeys, and Man 149 R.I. McLachlan, L. O’Donnell, S.J. Meachem, P.G. Stanton, D.M. de Kretser, K. Pratis, and D.M. Robertson 8. Molecular Defects of the Androgen Receptor 181 Michael J. McPhaul 9. Novel Signaling Pathways That Control Ovarian Follicular Development, Ovulation, and Luteinization 195 JoAnne S. Richards, Darryl L. Russell, Scott Ochsner, Minnie Hsieh, Kari H. Doyle, Allison E. Falender, Yuet K. Lo, and S. Chidananda Sharma 10. Transcription Factors Underlying the Development and Endocrine Functions of the Placenta 221 James C. Cross, Lynn Anson-Cartwright, and Ian C. Scott 11. Neuroendocrine Modulation and Repercussions of Female Reproductive Aging 235 Phyllis M. Wise, Matthew J. Smith, Dena B. Dubal, Melinda E. Wilson, Shane W. Rau, Adrienne B. Cashion, Martina Bo¨ttner, and Katherine L. Rosewell iii iv CONTENTS 12. Hormonal Changes in the Menopause Transition 257 Henry G. Burger, Emma C. Dudley, David M. Robertson, and Lorraine Dennerstein 13. Role of Hormonal and Reproductive Factors in the Etiology and Treatment of Uterine Leiomyoma 277 Cheryl L. Walker 14. Definition of the Molecular and Cellular Mechanisms Underlying the Tissue-selective Agonist/Antagonist Activities of Selective Estrogen Receptor Modulators 295 Donald P. McDonnell, Caroline E. Connor, Ashini Wijayaratne, Ching-yi Chang, and John D. Norris 15. Aromatase and Its Inhibitors: Significance for Breast Cancer Therapy 317 Evan R. Simpson and Mitch Dowsett 16. Reproductive Functions of Progesterone Receptors 339 Orla M. Conneely, Biserka Mulac-Jericevic, Francesco DeMayo, John P. Lydon, and Bert W. O’Malley 17. Estrogen Actions Throughout the Brain 357 Bruce McEwen 18. Sex Steroids and Bone 385 S.C. Manolagas, S. Kousteni, and R.L. Jilka 19. Anabolic Steroids 411 Cynthia M. Kuhn 20. Prolactin Modulation of Immune and Inflammatory Responses 435 Li-yuan Yu-Lee Author Index 457 Subject Index 459 Senior Author Correspondence Information Henry G. Burger James Cross Prince Henry’s Institute of Medical Research Department of Biochemistry & Molecular Level 4, Block E Biology Monash Medical Centre University of Calgary Faculty of Medicine 246 Clayton Road HSC Room 2279 Clayton, Victoria 3168 Australia 3330 Hospital Drive, NW Phone: 011-61-3-9594-3553 Calgary, Alberta T2N 4N1 Canada Fax: 011-61-3-9594-3558 Phone: 403-220-6876 E-mail: [email protected] Fax: 403-270-0737 E-mail: [email protected] Blanche Capel Department of Cell Biology Edward M. Eddy Duke University Medical Center Laboratory of Reproductive and 340 Nanaline Duke Developmental Toxicology Box 3709 National Institute of Environmental Health Durham, NC 27710 Sciences Phone: 919-684-6390 National Institutes of Health Fax: 919-681-7978 111 T.W. Alexander Drive, Building 101, E-mail: [email protected] Room C422a Research Triangle Park, NC 27709-2233 Phone: 919-541-3015 Fax: 919-541-3800 Orla M. Conneely E-mail: [email protected] Department of Molecular and Cellular Biology M-511 Debakey Building Baylor College of Medicine Michael D. Griswold Houston, TX 77030 School of Molecular Biosciences Phone: 713-798-6233 Center for Reproductive Biology E-mail: [email protected] Washington State University 630 Fulmer Hall Pullman, WA 99164-4660 Phone: 509-335-6281 Fax: 509-335-9688 E-mail: [email protected] v vi CORRESPONDENCE INFORMATION Cynthia M. Kuhn Robert I. McLachlan Department of Pharmacology and Cancer Male Reproductive Endocrinology Group Biology Prince Henry’s Institute of Medical Research Duke University Medical Center Level 4, Block E 4011 Bryan Research Building Monash Medical Centre Durham, NC 27710 246 Clayton Road Phone: 919-684-8828 Clayton, Victoria, 3168, Australia Fax: 919-681-8609 Phone: 011-61-3-9594-4372 E-mail: [email protected] Fax: 011-61-3-9594-6125 E-mail: [email protected] S.C. Manolagas Center for Osteoporosis and Metabolic Bone Michael J McPhaul Diseases Department of Internal Medicine Division of Endocrinology and Metabolism Division of Endocrinology and Metabolism University of Arkansas Medical School University of Texas Southwestern Medical 4301 W. Markham Street, Slot 587 Center Little Rock, AR 72205 5323 Harry Hines Boulevard Phone: 501-686-5130 Dallas, Texas 75390-8857 Fax: 501-296-1002 Phone: 214-648-3494 E-mail: [email protected] Fax: 214-648-8917 E-mail: [email protected] Donald P. McDonnell Department of Pharmacology and Cancer Biology David J Miller Duke University Medical Center Department of Animal Sciences Box 3813 University of Illinois at Urbana-Champaign Levine Science Research Center, C259 328 Animal Sciences Laboratory Research Drive 1207 West Gregory Drive Durham, NC 27710 Urbana, Illinois 61801 Phone: 919-684-6035 Phone: 217-333-3408 Fax: 919-681-7139 Fax: 217-333-8286 E-mail: [email protected] E-mail: [email protected] Bruce S. McEwen Keith L Parker Harold and Margaret Milliken Hatch Division of Endocrinology Laboratory of Neuroendocrinology Room J6.106 The Rockefeller University University of Texas Southwestern Medical 1230 York Avenue Center New York, NY 10021 5323 Harry Hines Boulevard Phone: 212-327-8624 Dallas, TX 75390-8857 Fax: 212-327-8634 Phone: 214-648-5027 E-mail: [email protected] Fax: 214-648-5044 E-mail: [email protected] CORRESPONDENCE INFORMATION vii JoAnne S Richards Phyllis M Wise Department of Molecular and Cellular Division of Biological Sciences Biology University of California-Davis Baylor College of Medicine One Shields Avenue One Baylor Plaza Davis, CA 95616-8536 Houston, TX 77030 Phone: 530-752-4460 Phone: 713-798-6238 Fax: 530-752-2604 Fax: 713-790-1275 E-mail: [email protected] E-mail: [email protected] Debra J Wolgemuth Evan R. Simpson Department of Genetics and Development Prince Henry’s Institute of Medical Research Columbia University College of Physicians Monash Medical Centre & Surgeons Level 4, Block E 630 West 168th Street 246 Clayton Road Black Building 1613 Clayton, Victoria 3168 Australia New York, NY 10032 Phone: 011-61-3-9594-4397 Phone: 212-305-7900 Fax: 011-61-3-9594-6376 Fax: 212-305-6084 E-mail: [email protected] E-mail: [email protected] Cheryl L Walker Li-yuan Yu-Lee Department of Carcinogenesis Department of Medicine University of Texas MD Anderson Cancer Section on Immunology, Allergy and Center Rheumatology Science Park-Research Division Baylor College of Medicine Park Road 1C 1 Baylor Plaza Smithville, TX 78957 Houston, TX 77030 Phone: 512-237-9550 Phone: 713-798-4770 Fax: 512-237-2475 Fax: 713-798-5780 E-mail: [email protected] E-mail: [email protected] Cellular and Molecular Pathways Regulating Mammalian Sex Determination CHRISTOPHER TILMANN AND BLANCHE CAPEL Department of Cell Biology, Duke University Medical Center, Durham, North Carolina 27710 ABSTRACT In mammals, sex is determined by the presence or absence of a single gene on the Y chromosome, Sry. Sry, a member of the high mobility group family of transcription factors, is required to initiate male-specific pathways and repress female-specific pathways. Expression of Sry in the gonad, beginning at 10.5 days postcoitum, leads to the differentiation of the somatic supporting cell precursors as Sertoli cells. These cells direct the other cells of the gonad into their respective lineages. Currently, no direct targets of Sry are known. A number of cellular pathways initiated by Sry are required for testis development. These include the proliferation of pre-Sertoli cells and commitment to the Sertoli lineage, migration of cells from the adjacent mesonephros, and formation of a male-specific vasculature. Work is underway to identify genes controlling these processes. These genes will then be linked to Sry. I. Introduction Organisms that display sexually dimorphic phenotypes have evolved a variety of mechanisms for determining sex. In worms and flies, sex is determined by the ratio of X chromosomes to autosomes (Parkhurst and Meneely, 1994; Cline and Meyer, 1996). Reptiles and fish use hormones, temperature, or environmental cues to determine sex (Shapiro, 1990; Pieau et al., 2001). In mammals, sex is determined genetically by the presence or absence of a Y chromosome (Swain and Lovell-Badge, 1999; Capel, 2000). In 1947, Alfred Jost showed that removal of the gonads from fetal rabbits resulted in the development of female secondary sex characteristics, whether the animal was XX or XY.