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HANDBOOK OF ANDROLOGY Second Edition The American Society of Andrology Copyright © March 2010, by The American Society of Andrology Printed by Allen Press, Inc.; 1041 New Hampshire Street, Lawrence, KS 66044 HANDBOOK OF ANDROLOGY Second Edition The American Society of Andrology Copyright © March 2010, by The American Society of Andrology Printed by Allen Press, Inc.; 1041 New Hampshire Street, Lawrence, KS 66044 Preface Since the first edition of the Handbook of Andrology, published by the American Society of Andrology 15 years ago, over 20,000 printed copies, as well as uncounted numbers of electronic copies, have been distributed to colleagues and trainees around the world. The first edition was translated into Spanish, Italian and Chinese. Most of the fundamental information provided in the first edition is still valid; however, the volume of scientific literature on the various facets of the subject of andrology has expanded tremendously. This has resulted in a constantly growing body of knowledge, not only in basic science but also in the clinical management of men’s health issues. In this edition, we have increased the number of chapters from 24 to 41, but have made every effort to retain a style that will allow trainees to be introduced to the field of Andrology and become as excited about working in this field as are the contributing authors. Our chapters encompass the wide range of topics that characterizes the field of Andrology, from molecular biology to veterinary and human medicine, from applied research to ethics. We once again have been extremely fortunate to have as authors world- renowned Andrologists who are experts in the various subjects included in the Handbook; we wish to thank each of them for their valuable contributions. With the advances in information technology, we feel that it is time to drop the traditional format of a paper bound book and move to an electronic only version for this edition of the Handbook. With the support of the American Society of Andrology, this edition of the Handbook will be freely available to all members of the Society. The electronic version is available as a single PDF or as PDFs of individual chapters. We believe that this approach will allow for a wider scope of circulation of the Handbook, permit more frequent updates of the content, and help save a forest of trees. It is our hope that this second edition of the Handbook of Andrology lays the foundation for basic scientists, clinician scientists, healthcare professionals, trainees, policy makers and anyone who has an interest in the discipline to acquire the relevant knowledge that they seek. Finally, we would like to acknowledge the dedicated secretarial support of Ms. Elise Boivin-Ford and to thank all the contributors and various members of the American Society of Andrology for their assistance and support in making this Handbook possible. Bernard Robaire and Peter Chan Co-Editors February 2010 I List of Contributors Mary S. Croughan, Ph.D. Géraldine Delbès, Ph.D. Ob/Gyn Women’s Hlth Clinical Res Department of Pharmacology and University of California Therapeutics Co-editors at San Francisco McGill University Box 1793, 1635 Divisadero St. 601 3655 Prom. Sir-William-Osler Bernard Robaire, Ph.D.* Peter Chan, M.D. San Francisco, CA 94143-1793 Montreal, QC H3G 1Y6 Department of Pharmacology and McGill University USA Canada Therapeutics Department of Surgery McGill University Royal Victoria Hospital 3655 Promenade Sir-William-Osler 687 Pine Avenue West, S6.95 Janice P. Evans, Ph.D. Duane L. Garner, Ph.D. Montreal, QC H3G 1Y6 Montreal, QC H3A 1A1 Division of Reproductive Biology Department of Animal Canada Canada Department of Biochemistry and Biotechnology Molecular Biology University of Nevada, Reno Bloomberg School of Public Health Reno, NV 89557 Authors Johns Hopkins University USA Baltimore, MD 21205 Rupert P. Amann, Ph.D.* John K. Amory, M.D. USA Animal Reproduction and Center for Research in Biotechnology Laboratory Reproduction & Contraception Colorado State University Department of Medicine Marc Goldstein, M.D. Barbara F. Hales, Ph.D. Fort Collins, CO 80523-1683 University of Washington Cornell Institute for Reproductive Department of Pharmacology and USA Box 356429 Medicine Therapeutics Seattle, WA 98195-6429 Weill-Cornell Medical College McGill University USA Box 580, 525 E 68th Street 3655 Promenade Sir-William-Osler New York, NY 10065 Montreal, QC H3G 1Y6 Janice L. Bailey, Ph.D. William J. Bremner M.D., Ph.D.* Canada Centre de recherche en biologie University of Washington School de la reproduction of Medicine Mary Ann Handel, Ph.D. Wayne J.G. Hellstrom. M.D.* Faculté des sciences de 1959 NE Pacific Street The Jackson Laboratory Tulane University School l'agriculture et de l'alimentation UW Mailbox 356420 600 Main Street of Medicine Pavillon Paul-Comtois, local 4147 Seattle, WA 98195-6420 Bar Harbor, Maine 04609 Department of Urology Université Laval USA USA New Orleans, LA 70112 Québec, QC G1K 7P4 USA Canada Terry R. Brown, Ph.D.* Arthur L. Burnett, M.D., Ph.D. Barry T. Hinton, Ph.D.* Ilpo Huhtaniemi, M.D., Ph.D. Division of Reproductive Biology Brady Urological Institute Department of Cell Biology Department of Reproductive Department of Biochemistry and Johns Hopkins Medical Institutions University of Virginia Health Biology Molecular Biology Baltimore, MA 21287 Sciences Center Imperial College London Bloomberg School of Public Health USA PO Box 800732 London W120NN Johns Hopkins University School Of Medicine #439 UK Baltimore, MD 21205 Charlottesville, VA 22908 USA USA (cid:3) II III List of Contributors Mary S. Croughan, Ph.D. Géraldine Delbès, Ph.D. Ob/Gyn Women’s Hlth Clinical Res Department of Pharmacology and University of California Therapeutics Co-editors at San Francisco McGill University Box 1793, 1635 Divisadero St. 601 3655 Prom. Sir-William-Osler Bernard Robaire, Ph.D.* Peter Chan, M.D. San Francisco, CA 94143-1793 Montreal, QC H3G 1Y6 Department of Pharmacology and McGill University USA Canada Therapeutics Department of Surgery McGill University Royal Victoria Hospital 3655 Promenade Sir-William-Osler 687 Pine Avenue West, S6.95 Janice P. Evans, Ph.D. Duane L. Garner, Ph.D. Montreal, QC H3G 1Y6 Montreal, QC H3A 1A1 Division of Reproductive Biology Department of Animal Canada Canada Department of Biochemistry and Biotechnology Molecular Biology University of Nevada, Reno Bloomberg School of Public Health Reno, NV 89557 Authors Johns Hopkins University USA Baltimore, MD 21205 Rupert P. Amann, Ph.D.* John K. Amory, M.D. USA Animal Reproduction and Center for Research in Biotechnology Laboratory Reproduction & Contraception Colorado State University Department of Medicine Marc Goldstein, M.D. Barbara F. Hales, Ph.D. Fort Collins, CO 80523-1683 University of Washington Cornell Institute for Reproductive Department of Pharmacology and USA Box 356429 Medicine Therapeutics Seattle, WA 98195-6429 Weill-Cornell Medical College McGill University USA Box 580, 525 E 68th Street 3655 Promenade Sir-William-Osler New York, NY 10065 Montreal, QC H3G 1Y6 Janice L. Bailey, Ph.D. William J. Bremner M.D., Ph.D.* Canada Centre de recherche en biologie University of Washington School de la reproduction of Medicine Mary Ann Handel, Ph.D. Wayne J.G. Hellstrom. M.D.* Faculté des sciences de 1959 NE Pacific Street The Jackson Laboratory Tulane University School l'agriculture et de l'alimentation UW Mailbox 356420 600 Main Street of Medicine Pavillon Paul-Comtois, local 4147 Seattle, WA 98195-6420 Bar Harbor, Maine 04609 Department of Urology Université Laval USA USA New Orleans, LA 70112 Québec, QC G1K 7P4 USA Canada Terry R. Brown, Ph.D.* Arthur L. Burnett, M.D., Ph.D. Barry T. Hinton, Ph.D.* Ilpo Huhtaniemi, M.D., Ph.D. Division of Reproductive Biology Brady Urological Institute Department of Cell Biology Department of Reproductive Department of Biochemistry and Johns Hopkins Medical Institutions University of Virginia Health Biology Molecular Biology Baltimore, MA 21287 Sciences Center Imperial College London Bloomberg School of Public Health USA PO Box 800732 London W120NN Johns Hopkins University School Of Medicine #439 UK Baltimore, MD 21205 Charlottesville, VA 22908 USA USA (cid:3) II III Handbook of Andrology – List of Contributors Handbook of Andrology – List of Contributors Nigel Hunt, Ph.D. Dolores Lamb, Ph.D.* Vassilios Papadopoulos, Ph.D. D.J. Parekh, M.D. Institute of Work, Health and Scott Department of Urology Departments of Medicine and of Department of Urology Organisations Baylor College of Medicine Pharmacology and Therapeutics The University of Texas Health University of Nottingham Houston, Texas 77030 The Research Institute of the Science Center at San Antonio Floor B, International House USA McGill University Health Centre San Antonio TX 78229 Jubilee Campus, Wollaton Road McGill University USA Nottingham NG8 1BB 1650 Cedar Avenue UK Montreal, QC H3G 1A4 Canada Sue McHale, Ph.D. Robert I. McLachlan, Ph.D. Faculty of Development Prince Henry’s Inst of Med Res Sally D. Perreault, Ph.D.* Phillip M. Pierorazio, M.D. and Society Level 4, Block E US Environmental Protection James Buchanan Brady Sheffield Hallam University Monash Medical Centre Agency Urological Institute Collegiate Crescent Campus 24 Clayton Rd Mail Drop E205-09 Johns Hopkins Medical Inst. Sheffield S10 2BP Clayton, Victoria 3168 Research Triangle Park NC 27711 Baltimore, MD 21287 UK Australia USA USA Kevin T. McVary, M.D. Marvin L. Meistrich, Ph.D. Tony M. Plant, Ph.D. Gail S. Prins, Ph.D.* Department of Urology Department of Experimental University of Pittsburgh School Department of Urology, MC 955 Feinberg School of Medicine Radiation Oncology of Medicine University of Illinois at Chicago Northwestern University School University of Texas M. D. Dept. of Cell Biology and Physiology 820 South Wood Street of Medicine Anderson Cancer Center Magee-Womens Research Institute Chicago, IL 60612 Unit 066, 1515 Holcombe Blvd. Chicago, IL 60611 204 Craft Avenue, Room B311 USA Houston, TX 77030-4009 USA Pittsburgh, PA 15213 USA USA Alvaro Morales, M.D. Abraham Morgentaler, M.D. Mathew C. Raynor, M.D. Kenneth Roberts, Ph.D. Department of Urology Men’s Health Boston Department of Urology University of Washington School Centre for Applied Urological Res Harvard Medical School Tulane Health Sciences Center of Medicine Queen's University One Brookline Place, Suite #624 1430 Tulane Avenue 320N Health Sciences Building Kingston, ON K7L 2V7 Brookline, MA 02445-7224 New Orleans, LA 70112 PO Box 1495 Canada USA USA Spokane, WA 9 9210 USA Makoto Nagano, Ph.D. Falk Ochsendorf, M.D., Ph.D. Department of Obstetrics and Zentrum Dermatologie und Gynecology Venerologie Susan A. Rothmann, Ph.D. Peter N. Schlegel, M.D. The Research Institute of the McGill Klinikum der J.W.Goethe- Fertility Solutions Inc. Department of Urology University Health Centre Universität Frankfurt 11811 Shaker Blvd., Suite 330 New York Presbyterian Hospital/ McGill University Theodor-Stern-Kai 7 Cleveland, OH 44120 Weill Cornell Medical Center 687 Pine Avenue West 60590 Frankfurt USA 525 East 68th Street Montreal QC H3A 1A1 Germany New York, NY 10021 Canada USA (cid:3) !"(cid:3) "(cid:3) Handbook of Andrology – List of Contributors Handbook of Andrology – List of Contributors Nigel Hunt, Ph.D. Dolores Lamb, Ph.D.* Vassilios Papadopoulos, Ph.D. D.J. Parekh, M.D. Institute of Work, Health and Scott Department of Urology Departments of Medicine and of Department of Urology Organisations Baylor College of Medicine Pharmacology and Therapeutics The University of Texas Health University of Nottingham Houston, Texas 77030 The Research Institute of the Science Center at San Antonio Floor B, International House USA McGill University Health Centre San Antonio TX 78229 Jubilee Campus, Wollaton Road McGill University USA Nottingham NG8 1BB 1650 Cedar Avenue UK Montreal, QC H3G 1A4 Canada Sue McHale, Ph.D. Robert I. McLachlan, Ph.D. Faculty of Development Prince Henry’s Inst of Med Res Sally D. Perreault, Ph.D.* Phillip M. Pierorazio, M.D. and Society Level 4, Block E US Environmental Protection James Buchanan Brady Sheffield Hallam University Monash Medical Centre Agency Urological Institute Collegiate Crescent Campus 24 Clayton Rd Mail Drop E205-09 Johns Hopkins Medical Inst. Sheffield S10 2BP Clayton, Victoria 3168 Research Triangle Park NC 27711 Baltimore, MD 21287 UK Australia USA USA Kevin T. McVary, M.D. Marvin L. Meistrich, Ph.D. Tony M. Plant, Ph.D. Gail S. Prins, Ph.D.* Department of Urology Department of Experimental University of Pittsburgh School Department of Urology, MC 955 Feinberg School of Medicine Radiation Oncology of Medicine University of Illinois at Chicago Northwestern University School University of Texas M. D. Dept. of Cell Biology and Physiology 820 South Wood Street of Medicine Anderson Cancer Center Magee-Womens Research Institute Chicago, IL 60612 Unit 066, 1515 Holcombe Blvd. Chicago, IL 60611 204 Craft Avenue, Room B311 USA Houston, TX 77030-4009 USA Pittsburgh, PA 15213 USA USA Alvaro Morales, M.D. Abraham Morgentaler, M.D. Mathew C. Raynor, M.D. Kenneth Roberts, Ph.D. Department of Urology Men’s Health Boston Department of Urology University of Washington School Centre for Applied Urological Res Harvard Medical School Tulane Health Sciences Center of Medicine Queen's University One Brookline Place, Suite #624 1430 Tulane Avenue 320N Health Sciences Building Kingston, ON K7L 2V7 Brookline, MA 02445-7224 New Orleans, LA 70112 PO Box 1495 Canada USA USA Spokane, WA 9 9210 USA Makoto Nagano, Ph.D. Falk Ochsendorf, M.D., Ph.D. Department of Obstetrics and Zentrum Dermatologie und Gynecology Venerologie Susan A. Rothmann, Ph.D. Peter N. Schlegel, M.D. The Research Institute of the McGill Klinikum der J.W.Goethe- Fertility Solutions Inc. Department of Urology University Health Centre Universität Frankfurt 11811 Shaker Blvd., Suite 330 New York Presbyterian Hospital/ McGill University Theodor-Stern-Kai 7 Cleveland, OH 44120 Weill Cornell Medical Center 687 Pine Avenue West 60590 Frankfurt USA 525 East 68th Street Montreal QC H3A 1A1 Germany New York, NY 10021 Canada USA (cid:3) !"(cid:3) "(cid:3) Handbook of Andrology – List of Contributors Handbook of Andrology – List of Contributors Richard J. Sherins, M.D.* David W. Silversides, Ph.D. Selma F. Witchel, M.D. William W. Wright, Ph.D. Columbia Fertility Associates Department of Veterinary Children’s Hospital of Pittsburgh Division of Reproductive Biology 2440 M Street North West, # 401 Biomedicine Division of Endocrinology Department of Biochemistry and Washington, DC 20037 University of Montreal Pittsburgh, PA 15213 Molecular Biology USA St-Hyacinthe, QC J2S 7C6 USA Bloomberg School of Public Health Canada Johns Hopkins University Baltimore, MD 21205 Niels E. Skakkebæk, M.D. Ronald S. Swerdloff, M.D.* USA Rigshospitalet, Department of Division of Endocrinology Growth & Reproduction Department of Medicine Barry R. Zirkin, Ph.D.* GR 5064, Blegdamsvej 9 Harbor-UCLA Medical Center Division of Reproductive Biology DK-2100 Copenhagen and Los Angeles Biomedical Department of Biochemistry and Denmark Research Center Molecular Biology 1000 W. Carson Street Bloomberg School of Public Health Torrance, CA 90509 Johns Hopkins University USA Baltimore, MD 21205 USA J. Lisa Tenover, M.D., Ph.D.* I.M. Thompson, M.D. Division of Geriatric Medicine Department of Urology Wesley Woods Health Center The University of Texas Health *Past President, American Society of Andrology 1841 Clifton Road NE Science Ctr at San Antonio Atlanta, GA 30329-5102 7703 Floyd Curl Drive USA San Antonio, TX 78229 USA Jacquetta M. Trasler, M.D., Ph.D. Robert S. Viger, Ph.D. Department of Pediatrics Laval University McGill University, MCH MUHC-RI Dept. of Ob/Gyn 2300 rue Tupper 2705 Laurier boulevard, T1-49 Montréal, QC H3H 1P3 Centre de recherche CHUQ-CHUL Canada Quebec, QC G1V 4G2 Canada Thomas J. Walsh, M.D. Christina Wang, M.D.* Department of Urology Division of Endocrinology University of Washington School Department of Medicine of Medicine Harbor-UCLA Medical Center and 1959 NE Pacific Street, # Bb667 Los Angeles Biomedical Seattle, WA 98195-0001 Research Center USA 1000 W. Carson Street Torrance, CA 90509 USA VI VII Handbook of Andrology – List of Contributors Handbook of Andrology – List of Contributors Richard J. Sherins, M.D.* David W. Silversides, Ph.D. Selma F. Witchel, M.D. William W. Wright, Ph.D. Columbia Fertility Associates Department of Veterinary Children’s Hospital of Pittsburgh Division of Reproductive Biology 2440 M Street North West, # 401 Biomedicine Division of Endocrinology Department of Biochemistry and Washington, DC 20037 University of Montreal Pittsburgh, PA 15213 Molecular Biology USA St-Hyacinthe, QC J2S 7C6 USA Bloomberg School of Public Health Canada Johns Hopkins University Baltimore, MD 21205 Niels E. Skakkebæk, M.D. Ronald S. Swerdloff, M.D.* USA Rigshospitalet, Department of Division of Endocrinology Growth & Reproduction Department of Medicine Barry R. Zirkin, Ph.D.* GR 5064, Blegdamsvej 9 Harbor-UCLA Medical Center Division of Reproductive Biology DK-2100 Copenhagen and Los Angeles Biomedical Department of Biochemistry and Denmark Research Center Molecular Biology 1000 W. Carson Street Bloomberg School of Public Health Torrance, CA 90509 Johns Hopkins University USA Baltimore, MD 21205 USA J. Lisa Tenover, M.D., Ph.D.* I.M. Thompson, M.D. Division of Geriatric Medicine Department of Urology Wesley Woods Health Center The University of Texas Health *Past President, American Society of Andrology 1841 Clifton Road NE Science Ctr at San Antonio Atlanta, GA 30329-5102 7703 Floyd Curl Drive USA San Antonio, TX 78229 USA Jacquetta M. Trasler, M.D., Ph.D. Robert S. Viger, Ph.D. Department of Pediatrics Laval University McGill University, MCH MUHC-RI Dept. of Ob/Gyn 2300 rue Tupper 2705 Laurier boulevard, T1-49 Montréal, QC H3H 1P3 Centre de recherche CHUQ-CHUL Canada Quebec, QC G1V 4G2 Canada Thomas J. Walsh, M.D. Christina Wang, M.D.* Department of Urology Division of Endocrinology University of Washington School Department of Medicine of Medicine Harbor-UCLA Medical Center and 1959 NE Pacific Street, # Bb667 Los Angeles Biomedical Seattle, WA 98195-0001 Research Center USA 1000 W. Carson Street Torrance, CA 90509 USA VI VII Forward by Philip Troen prostate, sexual function and contraception. Advances in these and other Reprinted from the first edition of the Handbook of Andrology areas have been made possible by a remarkable series of clinical studies and scientific discoveries using the classical disciplines of physiology, biochemistry, neuroscience, and molecular biology. As we have entered “What a piece of work is a man” each new stage of understanding and science, there has been no waning of the stimulus that I and my colleagues experience. At the same time, because Shakespeare, Hamlet, Act ii, scene 2, line 316 of the multidisciplinary nature of andrology, unsolved problems present themselves and the opportunities for advancement and success continue to expand. As Alexander Albert has noted, "Nature has experimented lavishly Why a handbook of andrology? Some handbooks are published to bring with the reproductive system." This fact provides both challenge and together multiple aspects of a diversified subject. Some handbooks are opportunity. Andrology covers a wide spectrum from before conception to designed to present an overview of a rapidly expanding subject for those aging. As you peruse this handbook, we hope you will appreciate the scope working in the field while other handbooks are intended to codify the of the field and share our excitement in the study of andrology.(cid:3) progress already made. Although there are elements of each of these approaches in this handbook of andrology, our main purpose is to present to scientists/clinicians early in their careers the scope, importance, and excitement of our discipline. What is andrology? Simplistically, one might say that andrology is to the male what gynecology is to the female. That is, andrology deals with matters affecting the male reproductive system. The earliest use of the term andrology, as reported by Mikko Niemi, appears to have been in 1891 in the Journal of the American Medical Association, reporting on the formation of the American Andrological Association. Little more was heard from that association and it was not until the latter half of this century that there emerged an andrology journal in 1969 and an active andrology organization, Comite Internacional de Andrologia, in 1970. In the quarter century since, there has been a veritable explosion of journals and publications, of societies and congresses, and of workshops and symposia devoted to andrology. Thus, we appear to be on a rapidly rising growth curve of knowledge and application in andrology. The scope of modern day andrology is strikingly indicated by the range of topics discussed in this handbook. Written by distinguished leaders in their fields, these topics were chosen to indicate the exciting breadth of andrology and the opportunity it holds for graduate students in the biological sciences, veterinary students, and medical students. Why become an andrologist? In choosing a career one, first of all, tries to identify an area of interest. Then one looks at the opportunity to succeed and the likelihood of making a contribution. As an andrologist for 40 years, I can testify to the continued excitement and interest the discipline has held for me. As noted above, there has been a rapid escalation of growth so that our discipline now ranges from genetic studies to pubertal changes in the male and from infertility and assisted reproduction techniques to disorders of the VIII IX

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