handbook of andrology
TRANSCRIPT
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
I
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
II III
List of Contributors
Co-editors
Bernard Robaire, Ph.D.* Department of Pharmacology and Therapeutics McGill University 3655 Promenade Sir-William-Osler Montreal, QC H3G 1Y6 Canada
Peter Chan, M.D. McGill University Department of Surgery Royal Victoria Hospital 687 Pine Avenue West, S6.95 Montreal, QC H3A 1A1 Canada
Authors
Rupert P. Amann, Ph.D.* Animal Reproduction and Biotechnology Laboratory Colorado State University Fort Collins, CO 80523-1683 USA
John K. Amory, M.D. Center for Research in Reproduction & Contraception Department of Medicine University of Washington Box 356429 Seattle, WA 98195-6429 USA
Janice L. Bailey, Ph.D. Centre de recherche en biologie de la reproduction Faculté des sciences de l'agriculture et de l'alimentation Pavillon Paul-Comtois, local 4147 Université Laval Québec, QC G1K 7P4 Canada
William J. Bremner M.D., Ph.D.* University of Washington School of Medicine 1959 NE Pacific Street UW Mailbox 356420 Seattle, WA 98195-6420 USA
Terry R. Brown, Ph.D.* Division of Reproductive Biology Department of Biochemistry and Molecular Biology Bloomberg School of Public Health Johns Hopkins University Baltimore, MD 21205 USA
Arthur L. Burnett, M.D., Ph.D. Brady Urological Institute Johns Hopkins Medical InstitutionsBaltimore, MA 21287 USA
Mary S. Croughan, Ph.D. Ob/Gyn Women’s Hlth Clinical Res University of California at San Francisco Box 1793, 1635 Divisadero St. 601 San Francisco, CA 94143-1793 USA
Géraldine Delbès, Ph.D. Department of Pharmacology and Therapeutics McGill University 3655 Prom. Sir-William-Osler Montreal, QC H3G 1Y6 Canada
Janice P. Evans, Ph.D. Division of Reproductive Biology Department of Biochemistry and Molecular Biology Bloomberg School of Public Health Johns Hopkins University Baltimore, MD 21205 USA
Duane L. Garner, Ph.D. Department of Animal Biotechnology University of Nevada, Reno Reno, NV 89557 USA
Marc Goldstein, M.D. Cornell Institute for Reproductive Medicine Weill-Cornell Medical College Box 580, 525 E 68th Street New York, NY 10065
Barbara F. Hales, Ph.D. Department of Pharmacology and Therapeutics McGill University 3655 Promenade Sir-William-Osler Montreal, QC H3G 1Y6 Canada
Mary Ann Handel, Ph.D. The Jackson Laboratory 600 Main Street Bar Harbor, Maine 04609 USA
Wayne J.G. Hellstrom. M.D.* Tulane University School of Medicine Department of Urology New Orleans, LA 70112 USA
Barry T. Hinton, Ph.D.* Department of Cell Biology University of Virginia Health Sciences Center PO Box 800732 School Of Medicine #439 Charlottesville, VA 22908 USA
Ilpo Huhtaniemi, M.D., Ph.D. Department of Reproductive Biology Imperial College London London W120NN UK
II III
List of Contributors
Co-editors
Bernard Robaire, Ph.D.* Department of Pharmacology and Therapeutics McGill University 3655 Promenade Sir-William-Osler Montreal, QC H3G 1Y6 Canada
Peter Chan, M.D. McGill University Department of Surgery Royal Victoria Hospital 687 Pine Avenue West, S6.95 Montreal, QC H3A 1A1 Canada
Authors
Rupert P. Amann, Ph.D.* Animal Reproduction and Biotechnology Laboratory Colorado State University Fort Collins, CO 80523-1683 USA
John K. Amory, M.D. Center for Research in Reproduction & Contraception Department of Medicine University of Washington Box 356429 Seattle, WA 98195-6429 USA
Janice L. Bailey, Ph.D. Centre de recherche en biologie de la reproduction Faculté des sciences de l'agriculture et de l'alimentation Pavillon Paul-Comtois, local 4147 Université Laval Québec, QC G1K 7P4 Canada
William J. Bremner M.D., Ph.D.* University of Washington School of Medicine 1959 NE Pacific Street UW Mailbox 356420 Seattle, WA 98195-6420 USA
Terry R. Brown, Ph.D.* Division of Reproductive Biology Department of Biochemistry and Molecular Biology Bloomberg School of Public Health Johns Hopkins University Baltimore, MD 21205 USA
Arthur L. Burnett, M.D., Ph.D. Brady Urological Institute Johns Hopkins Medical InstitutionsBaltimore, MA 21287 USA
Mary S. Croughan, Ph.D. Ob/Gyn Women’s Hlth Clinical Res University of California at San Francisco Box 1793, 1635 Divisadero St. 601 San Francisco, CA 94143-1793 USA
Géraldine Delbès, Ph.D. Department of Pharmacology and Therapeutics McGill University 3655 Prom. Sir-William-Osler Montreal, QC H3G 1Y6 Canada
Janice P. Evans, Ph.D. Division of Reproductive Biology Department of Biochemistry and Molecular Biology Bloomberg School of Public Health Johns Hopkins University Baltimore, MD 21205 USA
Duane L. Garner, Ph.D. Department of Animal Biotechnology University of Nevada, Reno Reno, NV 89557 USA
Marc Goldstein, M.D. Cornell Institute for Reproductive Medicine Weill-Cornell Medical College Box 580, 525 E 68th Street New York, NY 10065
Barbara F. Hales, Ph.D. Department of Pharmacology and Therapeutics McGill University 3655 Promenade Sir-William-Osler Montreal, QC H3G 1Y6 Canada
Mary Ann Handel, Ph.D. The Jackson Laboratory 600 Main Street Bar Harbor, Maine 04609 USA
Wayne J.G. Hellstrom. M.D.* Tulane University School of Medicine Department of Urology New Orleans, LA 70112 USA
Barry T. Hinton, Ph.D.* Department of Cell Biology University of Virginia Health Sciences Center PO Box 800732 School Of Medicine #439 Charlottesville, VA 22908 USA
Ilpo Huhtaniemi, M.D., Ph.D. Department of Reproductive Biology Imperial College London London W120NN UK
Handbook of Andrology – List of Contributors Handbook of Andrology – List of Contributors
!" "
Nigel Hunt, Ph.D. Institute of Work, Health and Organisations University of Nottingham Floor B, International House Jubilee Campus, Wollaton Road Nottingham NG8 1BB UK
Dolores Lamb, Ph.D.* Scott Department of Urology Baylor College of Medicine Houston, Texas 77030 USA
Sue McHale, Ph.D. Faculty of Development and Society Sheffield Hallam University Collegiate Crescent Campus Sheffield S10 2BP UK
Robert I. McLachlan, Ph.D. Prince Henry’s Inst of Med Res Level 4, Block E Monash Medical Centre 24 Clayton Rd Clayton, Victoria 3168 Australia
Kevin T. McVary, M.D. Department of Urology Feinberg School of Medicine Northwestern University School of Medicine Chicago, IL 60611 USA
Marvin L. Meistrich, Ph.D. Department of Experimental Radiation Oncology University of Texas M. D. Anderson Cancer Center Unit 066, 1515 Holcombe Blvd. Houston, TX 77030-4009 USA
Alvaro Morales, M.D. Department of Urology Centre for Applied Urological Res Queen's University Kingston, ON K7L 2V7 Canada
Abraham Morgentaler, M.D. Men’s Health Boston Harvard Medical School One Brookline Place, Suite #624 Brookline, MA 02445-7224 USA
Makoto Nagano, Ph.D. Department of Obstetrics and Gynecology The Research Institute of the McGill University Health Centre McGill University 687 Pine Avenue West Montreal QC H3A 1A1 Canada
Falk Ochsendorf, M.D., Ph.D. Zentrum Dermatologie und Venerologie Klinikum der J.W.Goethe-Universität Frankfurt Theodor-Stern-Kai 7 60590 Frankfurt Germany
Vassilios Papadopoulos, Ph.D. Departments of Medicine and of Pharmacology and Therapeutics The Research Institute of the McGill University Health Centre McGill University 1650 Cedar Avenue Montreal, QC H3G 1A4 Canada
D.J. Parekh, M.D. Department of Urology The University of Texas Health Science Center at San Antonio San Antonio TX 78229 USA
Sally D. Perreault, Ph.D.* US Environmental Protection Agency Mail Drop E205-09 Research Triangle Park NC 27711USA
Phillip M. Pierorazio, M.D. James Buchanan Brady Urological Institute Johns Hopkins Medical Inst. Baltimore, MD 21287 USA
Tony M. Plant, Ph.D. University of Pittsburgh School of Medicine Dept. of Cell Biology and PhysiologyMagee-Womens Research Institute204 Craft Avenue, Room B311 Pittsburgh, PA 15213 USA
Gail S. Prins, Ph.D.* Department of Urology, MC 955 University of Illinois at Chicago 820 South Wood Street Chicago, IL 60612 USA
Mathew C. Raynor, M.D. Department of Urology Tulane Health Sciences Center 1430 Tulane Avenue New Orleans, LA 70112 USA
Kenneth Roberts, Ph.D. University of Washington School of Medicine 320N Health Sciences Building PO Box 1495 Spokane, WA 9 9210 USA
Susan A. Rothmann, Ph.D. Fertility Solutions Inc. 11811 Shaker Blvd., Suite 330 Cleveland, OH 44120 USA
Peter N. Schlegel, M.D. Department of Urology New York Presbyterian Hospital/ Weill Cornell Medical Center 525 East 68th Street New York, NY 10021 USA
Handbook of Andrology – List of Contributors Handbook of Andrology – List of Contributors
!" "
Nigel Hunt, Ph.D. Institute of Work, Health and Organisations University of Nottingham Floor B, International House Jubilee Campus, Wollaton Road Nottingham NG8 1BB UK
Dolores Lamb, Ph.D.* Scott Department of Urology Baylor College of Medicine Houston, Texas 77030 USA
Sue McHale, Ph.D. Faculty of Development and Society Sheffield Hallam University Collegiate Crescent Campus Sheffield S10 2BP UK
Robert I. McLachlan, Ph.D. Prince Henry’s Inst of Med Res Level 4, Block E Monash Medical Centre 24 Clayton Rd Clayton, Victoria 3168 Australia
Kevin T. McVary, M.D. Department of Urology Feinberg School of Medicine Northwestern University School of Medicine Chicago, IL 60611 USA
Marvin L. Meistrich, Ph.D. Department of Experimental Radiation Oncology University of Texas M. D. Anderson Cancer Center Unit 066, 1515 Holcombe Blvd. Houston, TX 77030-4009 USA
Alvaro Morales, M.D. Department of Urology Centre for Applied Urological Res Queen's University Kingston, ON K7L 2V7 Canada
Abraham Morgentaler, M.D. Men’s Health Boston Harvard Medical School One Brookline Place, Suite #624 Brookline, MA 02445-7224 USA
Makoto Nagano, Ph.D. Department of Obstetrics and Gynecology The Research Institute of the McGill University Health Centre McGill University 687 Pine Avenue West Montreal QC H3A 1A1 Canada
Falk Ochsendorf, M.D., Ph.D. Zentrum Dermatologie und Venerologie Klinikum der J.W.Goethe-Universität Frankfurt Theodor-Stern-Kai 7 60590 Frankfurt Germany
Vassilios Papadopoulos, Ph.D. Departments of Medicine and of Pharmacology and Therapeutics The Research Institute of the McGill University Health Centre McGill University 1650 Cedar Avenue Montreal, QC H3G 1A4 Canada
D.J. Parekh, M.D. Department of Urology The University of Texas Health Science Center at San Antonio San Antonio TX 78229 USA
Sally D. Perreault, Ph.D.* US Environmental Protection Agency Mail Drop E205-09 Research Triangle Park NC 27711USA
Phillip M. Pierorazio, M.D. James Buchanan Brady Urological Institute Johns Hopkins Medical Inst. Baltimore, MD 21287 USA
Tony M. Plant, Ph.D. University of Pittsburgh School of Medicine Dept. of Cell Biology and PhysiologyMagee-Womens Research Institute204 Craft Avenue, Room B311 Pittsburgh, PA 15213 USA
Gail S. Prins, Ph.D.* Department of Urology, MC 955 University of Illinois at Chicago 820 South Wood Street Chicago, IL 60612 USA
Mathew C. Raynor, M.D. Department of Urology Tulane Health Sciences Center 1430 Tulane Avenue New Orleans, LA 70112 USA
Kenneth Roberts, Ph.D. University of Washington School of Medicine 320N Health Sciences Building PO Box 1495 Spokane, WA 9 9210 USA
Susan A. Rothmann, Ph.D. Fertility Solutions Inc. 11811 Shaker Blvd., Suite 330 Cleveland, OH 44120 USA
Peter N. Schlegel, M.D. Department of Urology New York Presbyterian Hospital/ Weill Cornell Medical Center 525 East 68th Street New York, NY 10021 USA
Handbook of Andrology – List of Contributors Handbook of Andrology – List of Contributors
VI VII
Richard J. Sherins, M.D.* Columbia Fertility Associates 2440 M Street North West, # 401 Washington, DC 20037 USA
David W. Silversides, Ph.D. Department of Veterinary Biomedicine University of Montreal St-Hyacinthe, QC J2S 7C6 Canada
Niels E. Skakkebæk, M.D. Rigshospitalet, Department of Growth & Reproduction GR 5064, Blegdamsvej 9 DK-2100 Copenhagen Denmark
Ronald S. Swerdloff, M.D.* Division of Endocrinology Department of Medicine Harbor-UCLA Medical Center and Los Angeles Biomedical Research Center 1000 W. Carson Street Torrance, CA 90509 USA
J. Lisa Tenover, M.D., Ph.D.* Division of Geriatric Medicine Wesley Woods Health Center 1841 Clifton Road NE Atlanta, GA 30329-5102 USA
I.M. Thompson, M.D. Department of Urology The University of Texas Health Science Ctr at San Antonio 7703 Floyd Curl Drive San Antonio, TX 78229 USA
Jacquetta M. Trasler, M.D., Ph.D. Department of Pediatrics McGill University, MCH MUHC-RI 2300 rue Tupper Montréal, QC H3H 1P3 Canada
Robert S. Viger, Ph.D. Laval University Dept. of Ob/Gyn 2705 Laurier boulevard, T1-49 Centre de recherche CHUQ-CHULQuebec, QC G1V 4G2 Canada
Thomas J. Walsh, M.D. Department of Urology University of Washington School of Medicine 1959 NE Pacific Street, # Bb667 Seattle, WA 98195-0001 USA
Christina Wang, M.D.* Division of Endocrinology Department of Medicine Harbor-UCLA Medical Center and Los Angeles Biomedical Research Center 1000 W. Carson Street Torrance, CA 90509 USA
Selma F. Witchel, M.D. Children’s Hospital of Pittsburgh Division of Endocrinology Pittsburgh, PA 15213 USA
William W. Wright, Ph.D. Division of Reproductive Biology Department of Biochemistry and Molecular Biology Bloomberg School of Public Health Johns Hopkins University Baltimore, MD 21205 USA
Barry R. Zirkin, Ph.D.* Division of Reproductive Biology Department of Biochemistry and Molecular Biology Bloomberg School of Public Health Johns Hopkins University Baltimore, MD 21205 USA
*Past President, American Society of Andrology
Handbook of Andrology – List of Contributors Handbook of Andrology – List of Contributors
VI VII
Richard J. Sherins, M.D.* Columbia Fertility Associates 2440 M Street North West, # 401 Washington, DC 20037 USA
David W. Silversides, Ph.D. Department of Veterinary Biomedicine University of Montreal St-Hyacinthe, QC J2S 7C6 Canada
Niels E. Skakkebæk, M.D. Rigshospitalet, Department of Growth & Reproduction GR 5064, Blegdamsvej 9 DK-2100 Copenhagen Denmark
Ronald S. Swerdloff, M.D.* Division of Endocrinology Department of Medicine Harbor-UCLA Medical Center and Los Angeles Biomedical Research Center 1000 W. Carson Street Torrance, CA 90509 USA
J. Lisa Tenover, M.D., Ph.D.* Division of Geriatric Medicine Wesley Woods Health Center 1841 Clifton Road NE Atlanta, GA 30329-5102 USA
I.M. Thompson, M.D. Department of Urology The University of Texas Health Science Ctr at San Antonio 7703 Floyd Curl Drive San Antonio, TX 78229 USA
Jacquetta M. Trasler, M.D., Ph.D. Department of Pediatrics McGill University, MCH MUHC-RI 2300 rue Tupper Montréal, QC H3H 1P3 Canada
Robert S. Viger, Ph.D. Laval University Dept. of Ob/Gyn 2705 Laurier boulevard, T1-49 Centre de recherche CHUQ-CHULQuebec, QC G1V 4G2 Canada
Thomas J. Walsh, M.D. Department of Urology University of Washington School of Medicine 1959 NE Pacific Street, # Bb667 Seattle, WA 98195-0001 USA
Christina Wang, M.D.* Division of Endocrinology Department of Medicine Harbor-UCLA Medical Center and Los Angeles Biomedical Research Center 1000 W. Carson Street Torrance, CA 90509 USA
Selma F. Witchel, M.D. Children’s Hospital of Pittsburgh Division of Endocrinology Pittsburgh, PA 15213 USA
William W. Wright, Ph.D. Division of Reproductive Biology Department of Biochemistry and Molecular Biology Bloomberg School of Public Health Johns Hopkins University Baltimore, MD 21205 USA
Barry R. Zirkin, Ph.D.* Division of Reproductive Biology Department of Biochemistry and Molecular Biology Bloomberg School of Public Health Johns Hopkins University Baltimore, MD 21205 USA
*Past President, American Society of Andrology
VIII IX
Forward by Philip Troen Reprinted from the first edition of the Handbook of Andrology
“What a piece of work is a man”
Shakespeare, Hamlet, Act ii, scene 2, line 316
Why a handbook of andrology? Some handbooks are published to bring together multiple aspects of a diversified subject. Some handbooks are designed to present an overview of a rapidly expanding subject for those working in the field while other handbooks are intended to codify the 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
prostate, sexual function and contraception. Advances in these and other 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 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 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 with the reproductive system." This fact provides both challenge and opportunity. Andrology covers a wide spectrum from before conception to aging. As you peruse this handbook, we hope you will appreciate the scope of the field and share our excitement in the study of andrology.
VIII IX
Forward by Philip Troen Reprinted from the first edition of the Handbook of Andrology
“What a piece of work is a man”
Shakespeare, Hamlet, Act ii, scene 2, line 316
Why a handbook of andrology? Some handbooks are published to bring together multiple aspects of a diversified subject. Some handbooks are designed to present an overview of a rapidly expanding subject for those working in the field while other handbooks are intended to codify the 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
prostate, sexual function and contraception. Advances in these and other 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 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 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 with the reproductive system." This fact provides both challenge and opportunity. Andrology covers a wide spectrum from before conception to aging. As you peruse this handbook, we hope you will appreciate the scope of the field and share our excitement in the study of andrology.
X XI
Handbook of Andrology
Contents
PREFACE B. Robaire and P. Chan
LIST OF CONTRIBUTORS
FOREWORD P. Troen
1. What are the components of the male reproductive system? CNS, pituitary, testis, epididymis, prostate, seminal vesicles, scrotum, penis K.P. Roberts
2. What is the relationship among the various endocrine components of the male reproductive system?
Hypothalamic-Pituitary-Testicular Axis, feedback loops R. Swerdloff
3. How are communication signals read in the male reproductive system?
Receptors for gonadotropins and androgens I. Huhtaniemi
4. What compounds mediate communication within the testis? Where and how are male-associated hormones produced?
Integration of the hypothalamus, pituitary and testis B.R. Zirkin
5. How is the synthesis of testosterone regulated? V. Papadopoulos
6. How are germ cells produced and what factors control their production?
M. Nagano
7. What unique chromosomal events lead to the formation of a haploid male germ cell?
M.A. Handel
8. How is the production of spermatozoa regulated? Hormonal regulation of spermatogenesis: role of androgens and FSH R.I. McLachlan
9. Are there epigenetic events associated with male germ cell formation? What is the role of genomic imprinting in male germ cells?
Imprinting, DNA methylation J.M. Trasler
10. What does the epididymis do and how does it do it? B.T. Hinton
11. What is the prostate and what are its functions? G.S. Prins
12. How does semen analysis assist in understanding the reproductive status of the male?
C. Wang
13. How is sperm chromatin structure assessed? What is the value of doing such assessments?
Range of methods that assess sperm chromatin quality B. Robaire
14. What is sperm banking? When and how is it (or should it be) used in humans? Animals?
Sperm banking, consequences of its use in animal and clinical practice S.A. Rothmann
15. How does the sperm make its way to the egg and how does fertilization take place?
Capacitation, acrosome exocytosis, steps of sperm-egg interaction, egg activation J.P. Evans and J.L. Bailey
16. What determines maleness? Sex determination, testis formation and the development of the male phenotype D.W. Silversides and R.S. Viger
X XI
Handbook of Andrology
Contents
PREFACE B. Robaire and P. Chan
LIST OF CONTRIBUTORS
FOREWORD P. Troen
1. What are the components of the male reproductive system? CNS, pituitary, testis, epididymis, prostate, seminal vesicles, scrotum, penis K.P. Roberts
2. What is the relationship among the various endocrine components of the male reproductive system?
Hypothalamic-Pituitary-Testicular Axis, feedback loops R. Swerdloff
3. How are communication signals read in the male reproductive system?
Receptors for gonadotropins and androgens I. Huhtaniemi
4. What compounds mediate communication within the testis? Where and how are male-associated hormones produced?
Integration of the hypothalamus, pituitary and testis B.R. Zirkin
5. How is the synthesis of testosterone regulated? V. Papadopoulos
6. How are germ cells produced and what factors control their production?
M. Nagano
7. What unique chromosomal events lead to the formation of a haploid male germ cell?
M.A. Handel
8. How is the production of spermatozoa regulated? Hormonal regulation of spermatogenesis: role of androgens and FSH R.I. McLachlan
9. Are there epigenetic events associated with male germ cell formation? What is the role of genomic imprinting in male germ cells?
Imprinting, DNA methylation J.M. Trasler
10. What does the epididymis do and how does it do it? B.T. Hinton
11. What is the prostate and what are its functions? G.S. Prins
12. How does semen analysis assist in understanding the reproductive status of the male?
C. Wang
13. How is sperm chromatin structure assessed? What is the value of doing such assessments?
Range of methods that assess sperm chromatin quality B. Robaire
14. What is sperm banking? When and how is it (or should it be) used in humans? Animals?
Sperm banking, consequences of its use in animal and clinical practice S.A. Rothmann
15. How does the sperm make its way to the egg and how does fertilization take place?
Capacitation, acrosome exocytosis, steps of sperm-egg interaction, egg activation J.P. Evans and J.L. Bailey
16. What determines maleness? Sex determination, testis formation and the development of the male phenotype D.W. Silversides and R.S. Viger
Handbook of Andrology – Contents Handbook of Andrology – Contents
XII XIII
17. Are there specific genetic defects affecting the male reproductive tract? What are the underlying molecular mechanisms?
Turner’s and Klinefelter’s syndromes, genetic loci, GnRH/LH/FSH, steroid biosynthesis and androgen insensitivity T.R. Brown
18. Is there a trigger for puberty in the male? Should early or delayed puberty be treated? If so, how?
T.M. Plant and S.F. Witchel
19. Clinical male infertility: Epidemiology and basic evaluation P. Chan
20. Why evaluate the infertile male in the era of ART? Medical and surgical therapies for male infertility M. Goldstein
21. What are the assisted reproductive technologies for male infertility? Indications for IVF/ICSI/IUI, Surgical sperm retrieval techniques P.N. Schlegel
22. Are there genetic factors associated with male infertility? D. Lamb
23. How can we assess the health outcomes in ART-conceived children?
Ethical issues in andrology T.J. Walsh and M.S. Croughan
24. How is male-factor subfertility minimized in companion and food-producing animals?
R.P. Amann
25. What should I know about artificial insemination (AI) of animals? R.P. Amann and D.L. Garner
26. Is there a decline in sperm counts in men? R.J. Sherins and G. Delbès
27. How do environmental cues affect male reproductive? W.W. Wright
28. What are the effects of environmental toxicants and/or endocrine disruptors on sperm function?
Chemical exposure effects on the male, reactive oxygen species and spermatozoa S.D. Perreault
29. Can spermatozoa be targets for drugs? If so, what are the consequences of such drug exposures? Is there a need for pre-conception counselling for men?
Drugs that affect sperm structure or function, male-mediated developmental toxicity, prevention, tests to detect damage to spermatozoa B.F. Hales and B. Robaire
30. Can we protect spermatogenesis against testicular insults? Male reproductive toxicology M.L. Meistrich
31. What male contraceptives currently exist and what is the outlook for new male contraceptives?
J.K. Amory and W.J. Bremner
32. What are the consequences of sexually transmitted infections on male reproduction?
F. Ochsendorf
33. What types of testicular cancers occur in men? What is their prognosis? How effective are existing therapeutic approaches?
N.E. Skakkebaek
34. How do erections occur? How common is erectile dysfunction, what is its etiology, and how do you evaluate men with this problem?
P.M. Pierorazio and A.L. Burnett
35. How to treat erectile dysfunction? M.C. Raynor and W.J.G. Hellstrom
36. Is there an andropause, better known as Testosterone Deficiency Syndrome, and if so, what tissues are affected and how?
Fertility, androgen production and sensitivity, osteoporosis and sexual function in aging men A. Morales
Handbook of Andrology – Contents Handbook of Andrology – Contents
XII XIII
17. Are there specific genetic defects affecting the male reproductive tract? What are the underlying molecular mechanisms?
Turner’s and Klinefelter’s syndromes, genetic loci, GnRH/LH/FSH, steroid biosynthesis and androgen insensitivity T.R. Brown
18. Is there a trigger for puberty in the male? Should early or delayed puberty be treated? If so, how?
T.M. Plant and S.F. Witchel
19. Clinical male infertility: Epidemiology and basic evaluation P. Chan
20. Why evaluate the infertile male in the era of ART? Medical and surgical therapies for male infertility M. Goldstein
21. What are the assisted reproductive technologies for male infertility? Indications for IVF/ICSI/IUI, Surgical sperm retrieval techniques P.N. Schlegel
22. Are there genetic factors associated with male infertility? D. Lamb
23. How can we assess the health outcomes in ART-conceived children?
Ethical issues in andrology T.J. Walsh and M.S. Croughan
24. How is male-factor subfertility minimized in companion and food-producing animals?
R.P. Amann
25. What should I know about artificial insemination (AI) of animals? R.P. Amann and D.L. Garner
26. Is there a decline in sperm counts in men? R.J. Sherins and G. Delbès
27. How do environmental cues affect male reproductive? W.W. Wright
28. What are the effects of environmental toxicants and/or endocrine disruptors on sperm function?
Chemical exposure effects on the male, reactive oxygen species and spermatozoa S.D. Perreault
29. Can spermatozoa be targets for drugs? If so, what are the consequences of such drug exposures? Is there a need for pre-conception counselling for men?
Drugs that affect sperm structure or function, male-mediated developmental toxicity, prevention, tests to detect damage to spermatozoa B.F. Hales and B. Robaire
30. Can we protect spermatogenesis against testicular insults? Male reproductive toxicology M.L. Meistrich
31. What male contraceptives currently exist and what is the outlook for new male contraceptives?
J.K. Amory and W.J. Bremner
32. What are the consequences of sexually transmitted infections on male reproduction?
F. Ochsendorf
33. What types of testicular cancers occur in men? What is their prognosis? How effective are existing therapeutic approaches?
N.E. Skakkebaek
34. How do erections occur? How common is erectile dysfunction, what is its etiology, and how do you evaluate men with this problem?
P.M. Pierorazio and A.L. Burnett
35. How to treat erectile dysfunction? M.C. Raynor and W.J.G. Hellstrom
36. Is there an andropause, better known as Testosterone Deficiency Syndrome, and if so, what tissues are affected and how?
Fertility, androgen production and sensitivity, osteoporosis and sexual function in aging men A. Morales
Handbook of Andrology – Contents
!"#
37. Should there be hormone replacement therapy for aging men? J.L. Tenover
38. What is the importance of psychosocial issues, counseling and psychotherapy in andrology?
N. Hunt and S. McHale
39. What is BPH? K.T. McVary
40. Are some men more susceptible to prostate cancer than others and why? What are the treatments and their effectiveness? What are the possibilities for improvements in therapy?
D.J. Parekh and I.M. Thompson
41. What is the risk of testosterone therapy after prostate cancer? A. Morgentaler
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&-/01/1! /1! /+/7/*7-,! *47-9!8/707/;! =90./4-9*7/0+! 04! 1=-98*7050+/*! @>! IJ(!1>+7<-1/1!7<*7!*;;08=./1<-1!=9-;/1-!9-=./;*7/0+!04!-*;<!;<9080108-!70!4098!7:0!;<908*7/,1'!H<21! 7<-!IJ(!;0+7-+7! ALMN!6*.2-C!<*1!,[email protected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email protected]#179*+,!IJ(!@9-*E1D!8-,/*7-,!@>! 7<-!FTU$$!-+B>8-D! 7<*7! /+/7/*7-! 7<-! 80.-;2.*9! -6-+71! 04! 8-/07/;! 9-;08@/+*7/0+'! (.10!,29/+5! 7<-! .-=707-+-! =<*1-D! <080.05021! ;<9080108-1! 4/+,! -*;<! 07<-9! @>!<080.05>!1-*9;</+5!8-;<*+/181! 7<*7!*9-!+07!:-..!2+,-91700,D!@27!8*>!@-!4*;/./7*7-,! @>! 7-.08-9-! ;.217-9/+5! /+70! *! L@0232-7N! 0+! 7<-! +2;.-*9! -+6-.0=-!*+,V09! 7<-!IJ(! @9-*E1! *+,! 12@1-32-+7! 4098*7/0+! 04! 1<097! 1/+5.-#179*+,-,!-+,1'! I29/+5! 7<-! B>507-+-! 17*5-D! ;<9080108-! =*/9/+5! -K7-+,1! *+,!<080.051!@-;08-!/+7/8*7-.>!*110;/*7-,!@>!1>+*=1/1D!*!=90;-11!8-,/*7-,!@>!7<-!1>+*=70+-8*.!;08=.-K!AFMC'!
H<-!FM!/1!*!=907-/+!;08=.-K!;08=9/1-,!04!.*7-9*.!-.-8-+71!7<*7!4098!7<-!1;*440.,D!09!*K-1D!04!-*;<!<080.05D!*+,!*!;-+79*.!-.-8-+7! 7<*7! /1!1792;729*.!L5.2-N!8-,/*7/+5!;08=.-7-!1>+*=1/1'!H<-!;08=.-7/0+!04! 1>+*=1/1!8*9E1! 7<-!@-5/++/+5!04!7<-!=*;<>7-+-!17*5-D!:</;<!/1!.-+57<>!A*==90K/8*7-.>!$Q!,*>1!/+!7<-!<28*+C!*+,! ;<*9*;7-9/B-,!@>! ;0+1/,-9*@.-! 590:7<!04! 7<-! 1=-98*70;>7-!*1!:-..!*1!@>!/8=097*+7!;<9080108-!,>+*8/;1'!!J0:!7<-!;<9080108-1!*9-!6/12*./B-,! *1! <080.05021! =*/91D! ;*..-,! @/6*.-+71D! -'5'D! %W! @/6*.-+71! /+!
<28*+1'!J07*@.>D!<0:-6-9D!7<-!+0+#<080.05021!X!*+,!Y!;<9080108-1!*9-!1>+*=1-,!0+.>!/+!*!18*..!9-5/0+!04!<080.05>!A7<-!=1-2,0#*270108*.!9-5/0+C!*+,!*9-!1-32-17-9-,!/+!*!<-7-90;<908*7/;!+2;.-*9!,08*/+!E+0:+!*1!7<-!XY!@0,>! A09! 1-K! @0,>C'! I29/+5! 7<-! =*;<>7-+-! 17*5-D! 80.-;2.*9! -6-+71! 04!<080.05021! 9-;08@/+*7/0+! 2+40.,'! R+7-9-17/+5.>D! 7<-! +28@-9! 04!9-;08@/+*7/0+#/+/7/*7/+5! ,[email protected]#179*+,! @9-*E1! /1! /+! *==90K/8*7-.>! 7-+#40.,!-K;-11!70!7<-!+28@-9!04! 4/+*.!9-;/=90;*.!9-;08@/+*7/0+1!A;901106-91CD!:</;<!/1! *.:*>1! *7! .-*17! 0+-! =-9! ;<9080108-D! @27! ;*+! @-! 2=! 70! 7:0! 09! 7<9--! /+!.0+5-9!;<9080108-1'!H<-!-K;-11!IJ(!,[email protected]#179*+,!@9-*E1!*9-!7<025<7!70!@-!9-=*/9-,!@>!*!9-;08@/+*7/0+!8-;<*+/18!7<*7!/+60.6-1!+0+#;901106-9!5-+-!;0+6-91/0+D! 09! 1/7-#1=-;/4/;! -K;<*+5-! 04! /+4098*7/0+! :/7<027! -K;<*+5-! 04!129902+,/+5!;<9080108*.!9-5/0+1'!
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
!"#$%&'()*+*,-.%&/01*2"/&.)$*"3*,14%"5"'$* 60147""8*"3*,14%"5"'$*9*:(0)*;1&<;.*/(%"-"="-05*.>.1)=*5.04*)"*)(.*3"%-0)&"1*"3*0** * * (0#5"&4*-05.*'.%-*/.55?*
! "#$! "#%!
!"#$%&'()&*%+",-.-/-.#0%*1*'$/%0*#2%$-%$"*%3-,.#$(-'%-3%#%"#40-(2%.#0*%5*,.%+*006%
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
&-/01/1! /1! /+/7/*7-,! *47-9!8/707/;! =90./4-9*7/0+! 04! 1=-98*7050+/*! @>! IJ(!1>+7<-1/1!7<*7!*;;08=./1<-1!=9-;/1-!9-=./;*7/0+!04!-*;<!;<9080108-!70!4098!7:0!;<908*7/,1'!H<21! 7<-!IJ(!;0+7-+7! ALMN!6*.2-C!<*1!,[email protected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email protected]#179*+,!IJ(!@9-*E1D!8-,/*7-,!@>! 7<-!FTU$$!-+B>8-D! 7<*7! /+/7/*7-! 7<-! 80.-;2.*9! -6-+71! 04! 8-/07/;! 9-;08@/+*7/0+'! (.10!,29/+5! 7<-! .-=707-+-! =<*1-D! <080.05021! ;<9080108-1! 4/+,! -*;<! 07<-9! @>!<080.05>!1-*9;</+5!8-;<*+/181! 7<*7!*9-!+07!:-..!2+,-91700,D!@27!8*>!@-!4*;/./7*7-,! @>! 7-.08-9-! ;.217-9/+5! /+70! *! L@0232-7N! 0+! 7<-! +2;.-*9! -+6-.0=-!*+,V09! 7<-!IJ(! @9-*E1! *+,! 12@1-32-+7! 4098*7/0+! 04! 1<097! 1/+5.-#179*+,-,!-+,1'! I29/+5! 7<-! B>507-+-! 17*5-D! ;<9080108-! =*/9/+5! -K7-+,1! *+,!<080.051!@-;08-!/+7/8*7-.>!*110;/*7-,!@>!1>+*=1/1D!*!=90;-11!8-,/*7-,!@>!7<-!1>+*=70+-8*.!;08=.-K!AFMC'!
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
<28*+1'!J07*@.>D!<0:-6-9D!7<-!+0+#<080.05021!X!*+,!Y!;<9080108-1!*9-!1>+*=1-,!0+.>!/+!*!18*..!9-5/0+!04!<080.05>!A7<-!=1-2,0#*270108*.!9-5/0+C!*+,!*9-!1-32-17-9-,!/+!*!<-7-90;<908*7/;!+2;.-*9!,08*/+!E+0:+!*1!7<-!XY!@0,>! A09! 1-K! @0,>C'! I29/+5! 7<-! =*;<>7-+-! 17*5-D! 80.-;2.*9! -6-+71! 04!<080.05021! 9-;08@/+*7/0+! 2+40.,'! R+7-9-17/+5.>D! 7<-! +28@-9! 04!9-;08@/+*7/0+#/+/7/*7/+5! ,[email protected]#179*+,! @9-*E1! /1! /+! *==90K/8*7-.>! 7-+#40.,!-K;-11!70!7<-!+28@-9!04! 4/+*.!9-;/=90;*.!9-;08@/+*7/0+1!A;901106-91CD!:</;<!/1! *.:*>1! *7! .-*17! 0+-! =-9! ;<9080108-D! @27! ;*+! @-! 2=! 70! 7:0! 09! 7<9--! /+!.0+5-9!;<9080108-1'!H<-!-K;-11!IJ(!,[email protected]#179*+,!@9-*E1!*9-!7<025<7!70!@-!9-=*/9-,!@>!*!9-;08@/+*7/0+!8-;<*+/18!7<*7!/+60.6-1!+0+#;901106-9!5-+-!;0+6-91/0+D! 09! 1/7-#1=-;/4/;! -K;<*+5-! 04! /+4098*7/0+! :/7<027! -K;<*+5-! 04!129902+,/+5!;<9080108*.!9-5/0+1'!
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Copyright © Society of Andrology Handbook of Andrology – Are there epigenetic events associated with male germ cell formation?
9-1 9-2
Are there epigenetic events associated with male germ cell formation? What is the role of genomic imprinting
in male germ cells?
J.M. Trasler
Epigenetics refers to heritable mechanisms of modulating gene expression that do not involve alterations in DNA sequence. Thus epigenetics affects the phenotype of a cell without affecting its genotype and is governed by processes that permit heritability from the parental to the daughter cells but are at the same time potentially modifiable or even reversible. DNA methylation, histone modifications and small RNAs are the three main molecular mechanisms that have been associated with epigenetic regulation of genes. These three mechanisms interact and stabilize each other and disruptions of one or more can lead to inappropriate expression or silencing of genes, resulting in “epigenetic diseases” such as cancer and imprinting disorders. Epigenetic mechanisms are conserved across higher eukaryotes including humans suggesting that they act in combination to regulate higher order chromatin and ultimately genome integrity.
Epigenetic patterning begins in the germline and is essential for normal embryo and postnatal development. Epigenetic modifications occurring during germ cell development are postulated to play roles in gene expression, meiosis, genomic integrity and genomic imprinting. Male and female germ cell development is a particularly critical time for the acquisition of the differential ‘marking’ of imprinted genes to ensure parent-of-origin specific expression. Genomic imprinting involves about 100 genes to date and refers to variation in the expression of these genes according to their maternal or paternal origin; imprinted genes play key roles in growth, placental function and many aspects of development including neurobehavioral processes. Many imprinted genes have distinct sequences known as differentially methylated regions or DMRs where DNA methylation differs between the maternal and paternal alleles. Since DNA methylation is the most extensively characterized epigenetic mechanism, it will be used here to illustrate the importance of epigenetics to male germ cell biology.
DNA methylation is found at the 5-position of cytosine residues within CpG dinucleotides (where cytosine is 5’ to guanine) at 20-30 million sites throughout the mammalian genome; about 60-80% of cytosines within CpGs are methylated. Methylation of DNA is invariably associated with transcriptional silencing. Two types of DNA methylation occur, either de novo methylation or the acquisition of methylation on unmethylated cytosines, or maintenance methylation, that takes place at the time of DNA replication to
ensure the propagation of genomic methylation from parent to daughter cells. The majority of de novo methylation is targeted to transposons and their remnants and to repeats such as pericentric satellite sequences with smaller amounts at single-copy sequences and the DMRs of imprinted genes. Methylation of DNA is catalyzed by a family of DNA (cytosine-5)-methyltransferases (DNMT enzymes or DNMTs). The main DNMTs involved in the establishment and maintenance of methylation patterns are DNMT1, DNMT3A and DNMT3B. Although it lacks enzymatic activity, DNMT3L (DNMT3-like) is related to and works with DNMT3A and DNMT3B. Demethylation occurs passively when methylation is not maintained following DNA replication or actively by an as yet poorly understood process/enzyme.
Epigenetic modifications including DNA methylation are for the most part erased in primordial germ cells and then reacquired in a sex-specific manner during germ cell development (Fig. 1). The erasure is particularly important for imprinted genes as maternal ‘marks’ on imprinted genes inherited from the mother must be erased and new paternal ‘marks’ introduced. A second period of erasure occurs in the preimplantation embryo and is thought to affect most sequences with the exception of imprinted genes and some repeat sequences. Imprinted gene methylation is maintained during preimplantation development since it is only in the germline (male or female depending on the gene) that imprinted genes acquire the allele-specific methylation that will result in monoallelic expression in the embryo and postnatal individual. As most methylation will be erased in the early embryo, it is postulated that germline DNA methylation, at sequences other than imprinted genes and repeats, plays specific roles in germ cell development, gene expression or chromatin structure during gametogenesis.
Following erasure in primordial germ cells (at approximately mid-gestation in the fetus), most male germline DNA methylation is acquired premeiotically in the prenatal gonocytes or pro-spermatogonial cells in the period between mid-gestation and birth (Fig. 1). DNA methylation acquisition in male germ cells continues after birth in the mitotic and meiotic germ cells and is complete by the pachytene phase of meiosis. Mitotic spermatogonia must also maintain the DNA methylation patterns acquired in the prenatal period. The precise signals that target DNA methylation to specific sequences in male germ cells are unclear but may include RNA-directed events. Potential candidates for RNA-based targeting are the recently described germ cell-specific small RNAs, the piRNAs. Gene-targeting experiments in mice have identified DNMT3a and DNMT3L as the predominant enzymes involved in the methylation of repetitive and imprinted sequences in the male germline. Absence of these enzymes results in the failure of spermatogenesis and infertility. For example, in male mice lacking DNMT3L, male germ cells over-express retrotransposons, and there is
Copyright © Society of Andrology Handbook of Andrology – Are there epigenetic events associated with male germ cell formation?
9-1 9-2
Are there epigenetic events associated with male germ cell formation? What is the role of genomic imprinting
in male germ cells?
J.M. Trasler
Epigenetics refers to heritable mechanisms of modulating gene expression that do not involve alterations in DNA sequence. Thus epigenetics affects the phenotype of a cell without affecting its genotype and is governed by processes that permit heritability from the parental to the daughter cells but are at the same time potentially modifiable or even reversible. DNA methylation, histone modifications and small RNAs are the three main molecular mechanisms that have been associated with epigenetic regulation of genes. These three mechanisms interact and stabilize each other and disruptions of one or more can lead to inappropriate expression or silencing of genes, resulting in “epigenetic diseases” such as cancer and imprinting disorders. Epigenetic mechanisms are conserved across higher eukaryotes including humans suggesting that they act in combination to regulate higher order chromatin and ultimately genome integrity.
Epigenetic patterning begins in the germline and is essential for normal embryo and postnatal development. Epigenetic modifications occurring during germ cell development are postulated to play roles in gene expression, meiosis, genomic integrity and genomic imprinting. Male and female germ cell development is a particularly critical time for the acquisition of the differential ‘marking’ of imprinted genes to ensure parent-of-origin specific expression. Genomic imprinting involves about 100 genes to date and refers to variation in the expression of these genes according to their maternal or paternal origin; imprinted genes play key roles in growth, placental function and many aspects of development including neurobehavioral processes. Many imprinted genes have distinct sequences known as differentially methylated regions or DMRs where DNA methylation differs between the maternal and paternal alleles. Since DNA methylation is the most extensively characterized epigenetic mechanism, it will be used here to illustrate the importance of epigenetics to male germ cell biology.
DNA methylation is found at the 5-position of cytosine residues within CpG dinucleotides (where cytosine is 5’ to guanine) at 20-30 million sites throughout the mammalian genome; about 60-80% of cytosines within CpGs are methylated. Methylation of DNA is invariably associated with transcriptional silencing. Two types of DNA methylation occur, either de novo methylation or the acquisition of methylation on unmethylated cytosines, or maintenance methylation, that takes place at the time of DNA replication to
ensure the propagation of genomic methylation from parent to daughter cells. The majority of de novo methylation is targeted to transposons and their remnants and to repeats such as pericentric satellite sequences with smaller amounts at single-copy sequences and the DMRs of imprinted genes. Methylation of DNA is catalyzed by a family of DNA (cytosine-5)-methyltransferases (DNMT enzymes or DNMTs). The main DNMTs involved in the establishment and maintenance of methylation patterns are DNMT1, DNMT3A and DNMT3B. Although it lacks enzymatic activity, DNMT3L (DNMT3-like) is related to and works with DNMT3A and DNMT3B. Demethylation occurs passively when methylation is not maintained following DNA replication or actively by an as yet poorly understood process/enzyme.
Epigenetic modifications including DNA methylation are for the most part erased in primordial germ cells and then reacquired in a sex-specific manner during germ cell development (Fig. 1). The erasure is particularly important for imprinted genes as maternal ‘marks’ on imprinted genes inherited from the mother must be erased and new paternal ‘marks’ introduced. A second period of erasure occurs in the preimplantation embryo and is thought to affect most sequences with the exception of imprinted genes and some repeat sequences. Imprinted gene methylation is maintained during preimplantation development since it is only in the germline (male or female depending on the gene) that imprinted genes acquire the allele-specific methylation that will result in monoallelic expression in the embryo and postnatal individual. As most methylation will be erased in the early embryo, it is postulated that germline DNA methylation, at sequences other than imprinted genes and repeats, plays specific roles in germ cell development, gene expression or chromatin structure during gametogenesis.
Following erasure in primordial germ cells (at approximately mid-gestation in the fetus), most male germline DNA methylation is acquired premeiotically in the prenatal gonocytes or pro-spermatogonial cells in the period between mid-gestation and birth (Fig. 1). DNA methylation acquisition in male germ cells continues after birth in the mitotic and meiotic germ cells and is complete by the pachytene phase of meiosis. Mitotic spermatogonia must also maintain the DNA methylation patterns acquired in the prenatal period. The precise signals that target DNA methylation to specific sequences in male germ cells are unclear but may include RNA-directed events. Potential candidates for RNA-based targeting are the recently described germ cell-specific small RNAs, the piRNAs. Gene-targeting experiments in mice have identified DNMT3a and DNMT3L as the predominant enzymes involved in the methylation of repetitive and imprinted sequences in the male germline. Absence of these enzymes results in the failure of spermatogenesis and infertility. For example, in male mice lacking DNMT3L, male germ cells over-express retrotransposons, and there is
Handbook of Andrology – Are there epigenetic events associated with male germ cell Handbook of Andrology – Are there epigenetic events associated with male germ cell formation? formation?
! " ! #
asynapsis and non-homologous synapsis during meiotic prophase and subsequent apoptosis of germ cells prior to pachytene.
FIG. 1. Timing of the erasure (gray bar), acquisition (red bar) and maintenance (black bar) of DNA methylation patterns in the prenatal and postnatal periods of male germ cell development. PGC, primordial germ cell; G, gonocyte or prospermatogonia; Spg, spermatogonia; PL, preleptotene spermatocyte; L, leptotene spermatocyte; Z, zygotene spermatocyte; P, pachytene spermatocyte; D, diplotene spermatocyte; RS, round spermatid; ES, elongating spermatid.
Evidence is accumulating that errors in the establishment or maintenance of germ cell DNA methylation patterns can cause human diseases such as imprinting disorders and cancer. Sperm samples from oligospermic patients have been reported to contain DNA methylation defects at imprinted loci. It has been suggested that paternal age effects that are seen for schizophrenia and other human conditions may involve errors in the maintenance of genomic methylation in the spermatogonia of older men. Furthermore, there is also concern for more than one generation, since while most epigenetic errors will be erased in the germline of the progeny, there are several examples of the potential for inheritance of epigenetic defects. Ongoing studies are assessing the effects of environmental conditions, diet and drugs on DNA methylation patterns in human sperm.
!"##$%&$' ($)'*+#
Grewal SI, Moazed D. Heterochromatin and epigenetic control of gene expression. Science 2003; 17: 2162-76.
Lin H. piRNAs in the germ line. Science 2007; 316: 397. Richards EJ. Inherited epigenetic variation—revisiting soft inheritance. Nat
Rev Genet 2006; 7: 395-401. Sasaki H, Matsui Y. Epigenetic events in mammalian germ-cell development:
reprogramming and beyond. Nat Rev Genet 2008; 9: 129-40. Schaefer CB, Ooi SKT, Bestor TH, Bourc’his D. Epigenetic decisions in
mammalian germ cells. Science 2007; 316: 398-99.
Handbook of Andrology – Are there epigenetic events associated with male germ cell Handbook of Andrology – Are there epigenetic events associated with male germ cell formation? formation?
! " ! #
asynapsis and non-homologous synapsis during meiotic prophase and subsequent apoptosis of germ cells prior to pachytene.
FIG. 1. Timing of the erasure (gray bar), acquisition (red bar) and maintenance (black bar) of DNA methylation patterns in the prenatal and postnatal periods of male germ cell development. PGC, primordial germ cell; G, gonocyte or prospermatogonia; Spg, spermatogonia; PL, preleptotene spermatocyte; L, leptotene spermatocyte; Z, zygotene spermatocyte; P, pachytene spermatocyte; D, diplotene spermatocyte; RS, round spermatid; ES, elongating spermatid.
Evidence is accumulating that errors in the establishment or maintenance of germ cell DNA methylation patterns can cause human diseases such as imprinting disorders and cancer. Sperm samples from oligospermic patients have been reported to contain DNA methylation defects at imprinted loci. It has been suggested that paternal age effects that are seen for schizophrenia and other human conditions may involve errors in the maintenance of genomic methylation in the spermatogonia of older men. Furthermore, there is also concern for more than one generation, since while most epigenetic errors will be erased in the germline of the progeny, there are several examples of the potential for inheritance of epigenetic defects. Ongoing studies are assessing the effects of environmental conditions, diet and drugs on DNA methylation patterns in human sperm.
!"##$%&$' ($)'*+#
Grewal SI, Moazed D. Heterochromatin and epigenetic control of gene expression. Science 2003; 17: 2162-76.
Lin H. piRNAs in the germ line. Science 2007; 316: 397. Richards EJ. Inherited epigenetic variation—revisiting soft inheritance. Nat
Rev Genet 2006; 7: 395-401. Sasaki H, Matsui Y. Epigenetic events in mammalian germ-cell development:
reprogramming and beyond. Nat Rev Genet 2008; 9: 129-40. Schaefer CB, Ooi SKT, Bestor TH, Bourc’his D. Epigenetic decisions in
mammalian germ cells. Science 2007; 316: 398-99.
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Are there specific genetic defects affecting the male reproductive tract? What are the underlying mechanisms?
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Gonadal disorders
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Are there specific genetic defects affecting the male reproductive tract? What are the underlying mechanisms?
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Gonadal disorders
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Copyright © American Society of Andrology Handbook of Andrology – What are the assisted reproductive technologies for male infertility?
21-1 21-2
What are the assisted reproductive technologies for male infertility?
Indications for IVF/ICSI/IUI, Surgical sperm retrieval techniques
P.N. Schlegel
Male fertility problems can occur when sperm are limited in number or function. In some cases, sperm washing is used to concentrate spermatozoa and deliver them through the uterus at the time of ovulation. In order for this treatment (referred to as intrauterine insemination [IUI]) to be successful, typically 5 million motile sperm must be present in the ejaculate. A more advanced technique for enhancing the interaction of sperm and egg is in vitro fertilization (IVF). IVF is an involved process that includes treatment of the woman using a series of hormone injections to stimulate the ovaries to produce multiple eggs, egg retrieval, and fertilization of those eggs outside of the body. A limited number of the fertilized eggs (embryos) are then transferred back to the woman’s uterus after incubating in the laboratory for several days. It was initially recognized that impaired sperm would not fertilize eggs very efficiently, even when sperm are put directly next to the eggs in the laboratory. Indeed, it was recognized early on in IVF that if sperm were abnormal, then IVF was not very successful. Conditions where very impaired sperm are present include men who have very few sperm in the ejaculate, when motility of sperm is severely impaired, or the shape of the sperm was very abnormal. All of these conditions may be present for men with severely impaired sperm production. When there is severe deficiency of sperm number and/or limited ability of the sperm to fertilize during IVF, then the adjunctive treatment of intracytoplasmic sperm injection (ICSI) may be required to provide any reasonable chance of pregnancy to occur for these patients.
The solution of ICSI to enhance the ability of impaired sperm to fertilize an egg was identified in 1991 by Drs. Gianpiero Palermo and Andre Van Steirteghem in Belgium. During an attempt to place a sperm close to an egg, Dr. Palermo accidentally injected a spermatozoon directly into an egg. The injected egg fertilized, developed into an embryo, was transferred back to the woman’s uterus and resulted in a pregnancy with delivery of a normal, healthy child. With this “accident,” a revolution in fertility treatment (especially the treatment of men with severe fertility problems) was started. Subsequently, ICSI has been used to expand the spectrum of male-factor infertility cases that can be treated for infertility. Whereas sperm from the testis or epididymis were thought to be incapable of fertilization, ICSI has changed our view of the fertilizing ability of “immature” sperm retrieved from the male reproductive tract.
Currently, not only are sperm of limited number (oligozoospermia), severely impaired motility (asthenozoospermia), and sperm that are not normally shaped (teratozoospermia) possible to use for ICSI, but sperm taken from the epididymis or testis that have not passed through the male reproductive system can also be used for ICSI. The development of ICSI encouraged us to search more extensively for sperm in the testes, not only for men who have reproductive tract obstruction, but also for men with sperm production so poor that no sperm are present in the ejaculate (azoospermia).
In order to perform the single-sperm injection technique (ICSI), in vitro fertilization is necessary. In vitro fertilization involves the stimulation of a woman's egg production in a controlled manner so that multiple eggs are produced in the given month or “cycle” of treatment, rather than the usual one egg per month. Initial treatment with medications to suppress a woman’s natural control of ovulation is given, followed by a series of hormone injections over a 3-4 week period of FSH-like fertility drugs. The final ten days or so of hormonal stimulation requires close monitoring of the woman's response to these drugs with daily serial ultrasound and blood tests. When optimal development of the eggs is seen, based on the development of follicles (fluid containing structures on the ovaries) and increased female hormone levels, then a final priming injection of a natural hormone (hCG), mimicking the physiological “LH-surge” that occurs before ovulation, is given to stimulate final egg development. Eggs are retrieved using a needle, directed by ultrasound through the vagina into these follicles, under a light general ("twilight") anesthetic. Sperm may then be injected into the eggs (oocytes) outside of the body ("in vitro").
The fertilized oocytes (now embryos) are maintained in culture for 3-5 days before transfer back to the woman. The chance of an individual embryo implanting and developing into a fetus is limited to 20-50%, depending on how well the embryo developed prior to transfer. Overall, the chance of pregnancy for each IVF attempt varies based on the age of the female partner. The chance of delivering a child for an individual IVF attempt from initial stimulation of egg production averages about 30%, but the chance of pregnancy when a woman is over 40 years of age may drop to about 10%.
Since the sperm that are selected for IVF and ICSI would not have naturally fertilized eggs in the past, concern exists about the risk of potential birth defects after application of IVF/ICSI (Chapter 23). To-date, the risk of birth defects does not appear to be any higher with sperm selection during ICSI, although a small but statistically significant increase in chromosomal abnormalities occurs after ICSI. The increase of chromosomal abnormalities in children from 0.2 to 0.8% with ICSI may occur because of abnormalities in the man providing sperm or the ICSI procedure itself.
The development of the advanced reproductive techniques of IVF and ICSI have revolutionized the treatment of severe male infertility. Many men are now
Copyright © American Society of Andrology Handbook of Andrology – What are the assisted reproductive technologies for male infertility?
21-1 21-2
What are the assisted reproductive technologies for male infertility?
Indications for IVF/ICSI/IUI, Surgical sperm retrieval techniques
P.N. Schlegel
Male fertility problems can occur when sperm are limited in number or function. In some cases, sperm washing is used to concentrate spermatozoa and deliver them through the uterus at the time of ovulation. In order for this treatment (referred to as intrauterine insemination [IUI]) to be successful, typically 5 million motile sperm must be present in the ejaculate. A more advanced technique for enhancing the interaction of sperm and egg is in vitro fertilization (IVF). IVF is an involved process that includes treatment of the woman using a series of hormone injections to stimulate the ovaries to produce multiple eggs, egg retrieval, and fertilization of those eggs outside of the body. A limited number of the fertilized eggs (embryos) are then transferred back to the woman’s uterus after incubating in the laboratory for several days. It was initially recognized that impaired sperm would not fertilize eggs very efficiently, even when sperm are put directly next to the eggs in the laboratory. Indeed, it was recognized early on in IVF that if sperm were abnormal, then IVF was not very successful. Conditions where very impaired sperm are present include men who have very few sperm in the ejaculate, when motility of sperm is severely impaired, or the shape of the sperm was very abnormal. All of these conditions may be present for men with severely impaired sperm production. When there is severe deficiency of sperm number and/or limited ability of the sperm to fertilize during IVF, then the adjunctive treatment of intracytoplasmic sperm injection (ICSI) may be required to provide any reasonable chance of pregnancy to occur for these patients.
The solution of ICSI to enhance the ability of impaired sperm to fertilize an egg was identified in 1991 by Drs. Gianpiero Palermo and Andre Van Steirteghem in Belgium. During an attempt to place a sperm close to an egg, Dr. Palermo accidentally injected a spermatozoon directly into an egg. The injected egg fertilized, developed into an embryo, was transferred back to the woman’s uterus and resulted in a pregnancy with delivery of a normal, healthy child. With this “accident,” a revolution in fertility treatment (especially the treatment of men with severe fertility problems) was started. Subsequently, ICSI has been used to expand the spectrum of male-factor infertility cases that can be treated for infertility. Whereas sperm from the testis or epididymis were thought to be incapable of fertilization, ICSI has changed our view of the fertilizing ability of “immature” sperm retrieved from the male reproductive tract.
Currently, not only are sperm of limited number (oligozoospermia), severely impaired motility (asthenozoospermia), and sperm that are not normally shaped (teratozoospermia) possible to use for ICSI, but sperm taken from the epididymis or testis that have not passed through the male reproductive system can also be used for ICSI. The development of ICSI encouraged us to search more extensively for sperm in the testes, not only for men who have reproductive tract obstruction, but also for men with sperm production so poor that no sperm are present in the ejaculate (azoospermia).
In order to perform the single-sperm injection technique (ICSI), in vitro fertilization is necessary. In vitro fertilization involves the stimulation of a woman's egg production in a controlled manner so that multiple eggs are produced in the given month or “cycle” of treatment, rather than the usual one egg per month. Initial treatment with medications to suppress a woman’s natural control of ovulation is given, followed by a series of hormone injections over a 3-4 week period of FSH-like fertility drugs. The final ten days or so of hormonal stimulation requires close monitoring of the woman's response to these drugs with daily serial ultrasound and blood tests. When optimal development of the eggs is seen, based on the development of follicles (fluid containing structures on the ovaries) and increased female hormone levels, then a final priming injection of a natural hormone (hCG), mimicking the physiological “LH-surge” that occurs before ovulation, is given to stimulate final egg development. Eggs are retrieved using a needle, directed by ultrasound through the vagina into these follicles, under a light general ("twilight") anesthetic. Sperm may then be injected into the eggs (oocytes) outside of the body ("in vitro").
The fertilized oocytes (now embryos) are maintained in culture for 3-5 days before transfer back to the woman. The chance of an individual embryo implanting and developing into a fetus is limited to 20-50%, depending on how well the embryo developed prior to transfer. Overall, the chance of pregnancy for each IVF attempt varies based on the age of the female partner. The chance of delivering a child for an individual IVF attempt from initial stimulation of egg production averages about 30%, but the chance of pregnancy when a woman is over 40 years of age may drop to about 10%.
Since the sperm that are selected for IVF and ICSI would not have naturally fertilized eggs in the past, concern exists about the risk of potential birth defects after application of IVF/ICSI (Chapter 23). To-date, the risk of birth defects does not appear to be any higher with sperm selection during ICSI, although a small but statistically significant increase in chromosomal abnormalities occurs after ICSI. The increase of chromosomal abnormalities in children from 0.2 to 0.8% with ICSI may occur because of abnormalities in the man providing sperm or the ICSI procedure itself.
The development of the advanced reproductive techniques of IVF and ICSI have revolutionized the treatment of severe male infertility. Many men are now
Handbook of Andrology – What are the assisted reproductive technologies for male infertility?
21-3
routinely treated using IVF/ICSI who would not have naturally fathered children before. Many doctors believe that specific treatment of male abnormalities make these advanced reproductive treatments unnecessary, because natural pregnancy can occur if improved sperm production can be obtained. ICSI has allowed new groups of men to be treated, including men who require sperm retrieval because of a lack of sperm in the ejaculate. The outcomes of these treatments continue to be studied.
Suggested reading
Alukal JP, Lamb DJ. Intracytoplasmic sperm injection (ICSI)--what are the risks? Urol Clin North Am. 2008; 35: 277-88,
Neri QV, Takeuchi T, Palermo GD. An update of assisted reproductive technologies results in the United States. Ann N Y Acad Sci. 2008; 1127: 41-8.
Nyboe Andersen A, Carlsen E, Loft A. Trends in the use of intracytoplasmatic sperm injection marked variability between countries. Hum Reprod Update. 2008; 14: 593-604.
Sarkar NN. Intracytoplasmic sperm injection: an assisted reproductive technique and its outcome to overcome infertility. J Obstet Gynaecol. 2007; 27: 347-53.
Schlegel PN. Nonobstructive azoospermia: A revolutionary surgical approach and results. Semin Reprod Med. 2009; 27: 165-70.
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Copyright © American Society of Andrology Handbook of Andrology – What should I know about artificial insemination (AI) of animals?
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What should I know about artificial insemination (AI) of animals?
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Why does AI have great impact?
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Use of AI
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How is outcome from AI measured?
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Copyright © American Society of Andrology Handbook of Andrology – What should I know about artificial insemination (AI) of animals?
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What should I know about artificial insemination (AI) of animals?
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Why does AI have great impact?
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Use of AI
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How is outcome from AI measured?
c213:C:3B!6:*9CB!*2+,6!W52:,;!8213:C2X'
Handbook of Andrology – What should I know about artificial insemination (AI) of Handbook of Andrology – What should I know about artificial insemination (AI) of animals animals?
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Modifying sperm before AI
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Use of “sexed sperm”
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Summary
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Suggested reading
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Handbook of Andrology – What should I know about artificial insemination (AI) of Handbook of Andrology – What should I know about artificial insemination (AI) of animals animals?
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Modifying sperm before AI
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Use of “sexed sperm”
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Summary
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Suggested reading
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Copyright © American Society of Andrology Handbook of Andrology – What are the effects of environmental toxicants and/or endocrine disruptors on sperm function?
28-1 28-2
What are the effects of environmental toxicants and/or endocrine disruptors on sperm function?
Chemical exposure effects on the male, reactive oxygen species and spermatozoa
S.D. Perreault
What is the evidence that environmental contaminants may impact sperm function in humans?
Our appreciation for the potential impact of chemicals and other environmental contaminants on male reproductive function (spermatogenesis and fertility) was awakened in the 1970s when a group of women whose husbands worked in a plant manufacturing the fungicide DBCP (dibromochloropropane) discovered that they were all having trouble getting pregnant. Subsequent epidemiological investigations showed an association between the men’s exposure to DBCP and low sperm counts or even azoospermia. Furthermore, sperm production improved in most men when the exposure ceased. This rather serendipitous finding raised awareness that infertility was not necessarily a female problem and motivated regulatory agencies like the US EPA (Environmental Protection Agency), USDA (United States Department of Agriculture) and FDA (Food and Drug Administration) to improve their testing procedures and protocols in order to detect potential male reproductive toxicants. It also motivated NIOSH (National Institute of Occupational Safety and Health) and others to conduct occupational health studies for a variety of chemicals. We now know that in addition to certain pesticides, many environmental contaminants including certain industrial solvents, plasticizers and metals (notably lead and cadmium) have the potential to disrupt spermatogenesis and or specifically alter sperm function, given a sufficiently high exposure.
In general, evidence for adverse reproductive effects in humans has been found only in small cohorts of men exposed to relatively high levels of chemicals in occupational settings or from accidental chemical release. The extent to which the low levels of compounds or combinations of compounds that we encounter in ambient day-to-day environments may impact male reproduction is largely unknown and far more difficult to determine.
How do regulatory agencies test chemicals for spermatotoxicity and predict male reproductive effects?
In 1998, the US EPA updated its testing protocols required for the (http://www.epa.gov/opptsfrs/publications/OPPTS_Harmonized/870_Health Effects_Test_Guidelines/Series/870-3800.pdf) registration of pesticides. The current guidelines, that have been harmonized across federal agencies in the US and internationally, call for multigenerational testing in animal test species (primarily rats) where measures of sperm production and function (sperm motility and morphology) are obtained in parental adult animals and in their offspring. The protocol also evaluates reproductive tract development after in utero or early life exposures. Because males are dosed for the entire duration of spermatogenesis before they are evaluated and bred, adverse effects on any spermatogenic or somatic cell target should be detectable in the form of reduced testis or epididymal weights, altered testis histology (one of the most sensitive indicators of effects), reduced epididymal sperm counts, altered sperm motility or morphology and/or reduced fertility. When effects are seen, additional specific tests for sperm function such as in vitro fertilization (IVF) and evaluation of sperm chromatin structure, DNA damage, and chromosome numbers/structure can be used to further characterize the effect (Chapter 13).
Although typically not required for non-pesticidal industrial chemicals, these standard and specialized tests are being applied on a voluntary basis to evaluate chemical safety for compounds produced and used in high volume and those predicted to have reproductive effects based on their similarity to known toxicants (e.g. by structure-activity analysis) or based on in vitro screening for endocrine activity, oxidative stress, mutagenicity or cytotoxicity.
Regulatory agencies consider all available information from such tests when they perform a risk assessment, the complex process that integrates dose-response data from the test species, the chemical’s mode of action (if known), the relevance of the animal data to humans, and any available human epidemiological data with the goal of predicting risk to humans. In addition to formal risk assessments conducted by regulatory agencies, the National Toxicology Program conducts reviews of high priority reproductive toxicants through its Center for Environmental Risks to Human Reproduction (CERHR) and posts reports on the internet (http://cerhr.niehs.nih.gov/). CERHR selects chemicals based on both their occurrence (production volume and likely human exposure) and mode of action (likely reproductive
Copyright © American Society of Andrology Handbook of Andrology – What are the effects of environmental toxicants and/or endocrine disruptors on sperm function?
28-1 28-2
What are the effects of environmental toxicants and/or endocrine disruptors on sperm function?
Chemical exposure effects on the male, reactive oxygen species and spermatozoa
S.D. Perreault
What is the evidence that environmental contaminants may impact sperm function in humans?
Our appreciation for the potential impact of chemicals and other environmental contaminants on male reproductive function (spermatogenesis and fertility) was awakened in the 1970s when a group of women whose husbands worked in a plant manufacturing the fungicide DBCP (dibromochloropropane) discovered that they were all having trouble getting pregnant. Subsequent epidemiological investigations showed an association between the men’s exposure to DBCP and low sperm counts or even azoospermia. Furthermore, sperm production improved in most men when the exposure ceased. This rather serendipitous finding raised awareness that infertility was not necessarily a female problem and motivated regulatory agencies like the US EPA (Environmental Protection Agency), USDA (United States Department of Agriculture) and FDA (Food and Drug Administration) to improve their testing procedures and protocols in order to detect potential male reproductive toxicants. It also motivated NIOSH (National Institute of Occupational Safety and Health) and others to conduct occupational health studies for a variety of chemicals. We now know that in addition to certain pesticides, many environmental contaminants including certain industrial solvents, plasticizers and metals (notably lead and cadmium) have the potential to disrupt spermatogenesis and or specifically alter sperm function, given a sufficiently high exposure.
In general, evidence for adverse reproductive effects in humans has been found only in small cohorts of men exposed to relatively high levels of chemicals in occupational settings or from accidental chemical release. The extent to which the low levels of compounds or combinations of compounds that we encounter in ambient day-to-day environments may impact male reproduction is largely unknown and far more difficult to determine.
How do regulatory agencies test chemicals for spermatotoxicity and predict male reproductive effects?
In 1998, the US EPA updated its testing protocols required for the (http://www.epa.gov/opptsfrs/publications/OPPTS_Harmonized/870_Health Effects_Test_Guidelines/Series/870-3800.pdf) registration of pesticides. The current guidelines, that have been harmonized across federal agencies in the US and internationally, call for multigenerational testing in animal test species (primarily rats) where measures of sperm production and function (sperm motility and morphology) are obtained in parental adult animals and in their offspring. The protocol also evaluates reproductive tract development after in utero or early life exposures. Because males are dosed for the entire duration of spermatogenesis before they are evaluated and bred, adverse effects on any spermatogenic or somatic cell target should be detectable in the form of reduced testis or epididymal weights, altered testis histology (one of the most sensitive indicators of effects), reduced epididymal sperm counts, altered sperm motility or morphology and/or reduced fertility. When effects are seen, additional specific tests for sperm function such as in vitro fertilization (IVF) and evaluation of sperm chromatin structure, DNA damage, and chromosome numbers/structure can be used to further characterize the effect (Chapter 13).
Although typically not required for non-pesticidal industrial chemicals, these standard and specialized tests are being applied on a voluntary basis to evaluate chemical safety for compounds produced and used in high volume and those predicted to have reproductive effects based on their similarity to known toxicants (e.g. by structure-activity analysis) or based on in vitro screening for endocrine activity, oxidative stress, mutagenicity or cytotoxicity.
Regulatory agencies consider all available information from such tests when they perform a risk assessment, the complex process that integrates dose-response data from the test species, the chemical’s mode of action (if known), the relevance of the animal data to humans, and any available human epidemiological data with the goal of predicting risk to humans. In addition to formal risk assessments conducted by regulatory agencies, the National Toxicology Program conducts reviews of high priority reproductive toxicants through its Center for Environmental Risks to Human Reproduction (CERHR) and posts reports on the internet (http://cerhr.niehs.nih.gov/). CERHR selects chemicals based on both their occurrence (production volume and likely human exposure) and mode of action (likely reproductive
Handbook of Andrology – What are the effects of environmental toxicants and/or Handbook of Andrology – What are the effects of environmental toxicants and/or endocrine disruptors on sperm function? endocrine disruptors on sperm function?
28-3 28-4
toxicant), thereby evaluating compounds of high public and regulatory concern such as acrylamide, bromopropane, phthalates and bisphenol A.
Biologically plausible modes of spermatotoxicity: Oxidative stress
Sperm are unique in their metabolism and actually use oxidative stress to advantage. Reactive oxygen species (ROS) generated by metabolically active sperm play a role in the normal signaling that leads to capacitation or the capability of fertilizing eggs (Chapter 15). However, excess ROS can also cause membrane damage (lipid peroxidation) and sperm DNA damage, with undesirable consequences on fertilizing ability and genetic integrity of the ensuing embryo, respectively. Therefore, environmental contaminants that generate oxidative stress or alter REDOX status (including DBCP, acrylamide and bromopropane) are biologically plausible candidate spermatotoxicants. An understanding of oxidative stress in sperm is also directly applicable to assisted reproductive technologies. Because sperm can undergo oxidative damage during in vitro culture, media components and IVF procedures are being optimized to minimize such effects.
Few known male reproductive toxicants act only or specifically on sperm maturing in the epididymis. Alpha-chlorohydrin, a sperm metabolic poison that inhibits sperm motility, is one such chemical. However, some toxicants can have direct effects on sperm motility and/or fertilizing ability and impair earlier stages of spermatogenesis. For example, chronic exposure to acrylamide can damage spermatogonia and spermatocytes and indirectly impair fertility by arresting spermatogenesis. At levels that do not arrest spermatogenesis, the reactive metabolite of acrylamide can also alkylate sperm nuclear protamine during the final stages of spermiogenesis. As a result, the sperm may be able to fertilize eggs, but the resulting embryos die due to acrylamide-induced damage in the sperm pronucleus. The good news is that effects of chemicals that react directly with maturing sperm are as transient as the exposure. Once the exposure has ended and the affected sperm have been released from the body, the risk disappears.
Cigarette smoke and air pollution resulting from the combustion of coal and gasoline contain reactive chemicals that can cause oxidative stress and inflammatory changes in the respiratory and cardiovascular systems. Both cigarette consumption and exposure to high levels of air pollution have also been associated with altered semen quality and DNA damage in sperm. Because such sources of oxidative stress can be difficult to avoid, antioxidant dietary supplements may be of therapeutic or protective value. However, convincing benefits of this strategy have been hard to measure in human studies.
Biologically plausible modes of spermatotoxicity: Endocrine disruptors
Certain chemicals may impair sperm function by interfering with androgen action. For example, toxicology studies in rats show that the fungicide vinclozolin (and its metabolites) inhibits androgen action by binding to the androgen receptor, and some phthalates interfere with steroidogenesis resulting in reduced testosterone synthesis. Some chemicals such as bisphenol A can act as estrogens, while others are promiscuous and bind multiple receptors. Because hormones themselves are active at very low levels, these so-called “endocrine disruptors” have the potential to impact human reproductive processes, even when present at the low levels typically encountered in the environment. Furthermore, the developing male reproductive system is particularly sensitive to altered testosterone signaling. Toxicology studies in rats, where specific windows of exposure and sensitivity can be precisely defined, have shown that blocking androgen action during reproductive tract development can result in hypospadias and testicular maldescent in male offspring. Since the later stages of spermiogenesis depend on androgen, and there is increasing evidence for a role for estrogen in sperm function, endocrine disruptors may also impact sperm function in adults. At environmental levels of these contaminants, such effects are likely to be subtle and difficult to detect. However, even subtle effects may have negative impacts on subfertile men who are thought to be more susceptible to environmental stressors.
Based on growing evidence about the occurrence and potentially adverse effects of environmental endocrine disruptors, regulatory agencies have been directed by Congress to develop an effective screening and testing program for contaminants with endocrine disruption potential (http://www.epa.gov/endo/). These screening tests and other high throughput test systems currently under development will make it more feasible to test the large numbers of new chemicals introduced into our environment by commerce and thereby protect our reproductive health.
How can clinicians advise their infertility patients and couples attemp-ting pregnancy?
Clearly there is a need for patient-oriented information about the linkage between environmental contaminants and risks to male reproductive function. The Collaborative on Health and Environment makes such information readily available on the web (http://www.healthandenvironment .org/working_groups/fertility). At present, physicians can at least include a
Handbook of Andrology – What are the effects of environmental toxicants and/or Handbook of Andrology – What are the effects of environmental toxicants and/or endocrine disruptors on sperm function? endocrine disruptors on sperm function?
28-3 28-4
toxicant), thereby evaluating compounds of high public and regulatory concern such as acrylamide, bromopropane, phthalates and bisphenol A.
Biologically plausible modes of spermatotoxicity: Oxidative stress
Sperm are unique in their metabolism and actually use oxidative stress to advantage. Reactive oxygen species (ROS) generated by metabolically active sperm play a role in the normal signaling that leads to capacitation or the capability of fertilizing eggs (Chapter 15). However, excess ROS can also cause membrane damage (lipid peroxidation) and sperm DNA damage, with undesirable consequences on fertilizing ability and genetic integrity of the ensuing embryo, respectively. Therefore, environmental contaminants that generate oxidative stress or alter REDOX status (including DBCP, acrylamide and bromopropane) are biologically plausible candidate spermatotoxicants. An understanding of oxidative stress in sperm is also directly applicable to assisted reproductive technologies. Because sperm can undergo oxidative damage during in vitro culture, media components and IVF procedures are being optimized to minimize such effects.
Few known male reproductive toxicants act only or specifically on sperm maturing in the epididymis. Alpha-chlorohydrin, a sperm metabolic poison that inhibits sperm motility, is one such chemical. However, some toxicants can have direct effects on sperm motility and/or fertilizing ability and impair earlier stages of spermatogenesis. For example, chronic exposure to acrylamide can damage spermatogonia and spermatocytes and indirectly impair fertility by arresting spermatogenesis. At levels that do not arrest spermatogenesis, the reactive metabolite of acrylamide can also alkylate sperm nuclear protamine during the final stages of spermiogenesis. As a result, the sperm may be able to fertilize eggs, but the resulting embryos die due to acrylamide-induced damage in the sperm pronucleus. The good news is that effects of chemicals that react directly with maturing sperm are as transient as the exposure. Once the exposure has ended and the affected sperm have been released from the body, the risk disappears.
Cigarette smoke and air pollution resulting from the combustion of coal and gasoline contain reactive chemicals that can cause oxidative stress and inflammatory changes in the respiratory and cardiovascular systems. Both cigarette consumption and exposure to high levels of air pollution have also been associated with altered semen quality and DNA damage in sperm. Because such sources of oxidative stress can be difficult to avoid, antioxidant dietary supplements may be of therapeutic or protective value. However, convincing benefits of this strategy have been hard to measure in human studies.
Biologically plausible modes of spermatotoxicity: Endocrine disruptors
Certain chemicals may impair sperm function by interfering with androgen action. For example, toxicology studies in rats show that the fungicide vinclozolin (and its metabolites) inhibits androgen action by binding to the androgen receptor, and some phthalates interfere with steroidogenesis resulting in reduced testosterone synthesis. Some chemicals such as bisphenol A can act as estrogens, while others are promiscuous and bind multiple receptors. Because hormones themselves are active at very low levels, these so-called “endocrine disruptors” have the potential to impact human reproductive processes, even when present at the low levels typically encountered in the environment. Furthermore, the developing male reproductive system is particularly sensitive to altered testosterone signaling. Toxicology studies in rats, where specific windows of exposure and sensitivity can be precisely defined, have shown that blocking androgen action during reproductive tract development can result in hypospadias and testicular maldescent in male offspring. Since the later stages of spermiogenesis depend on androgen, and there is increasing evidence for a role for estrogen in sperm function, endocrine disruptors may also impact sperm function in adults. At environmental levels of these contaminants, such effects are likely to be subtle and difficult to detect. However, even subtle effects may have negative impacts on subfertile men who are thought to be more susceptible to environmental stressors.
Based on growing evidence about the occurrence and potentially adverse effects of environmental endocrine disruptors, regulatory agencies have been directed by Congress to develop an effective screening and testing program for contaminants with endocrine disruption potential (http://www.epa.gov/endo/). These screening tests and other high throughput test systems currently under development will make it more feasible to test the large numbers of new chemicals introduced into our environment by commerce and thereby protect our reproductive health.
How can clinicians advise their infertility patients and couples attemp-ting pregnancy?
Clearly there is a need for patient-oriented information about the linkage between environmental contaminants and risks to male reproductive function. The Collaborative on Health and Environment makes such information readily available on the web (http://www.healthandenvironment .org/working_groups/fertility). At present, physicians can at least include a
Handbook of Andrology – What are the effects of environmental toxicants and/or endocrine disruptors on sperm function?
28-5
short history on occupational and environmental exposures and advise their patients to avoid exposures to known toxicants such as cigarette smoke, solvents and pesticides.
Suggested reading
Aitken JR, Baker MA. Oxidative stress, sperm survival and fertility control. Mol Cell Endocrinol. 2006; 250: 66-9.
Hotchkiss AK, Rider CV, Blystone CR, Wilson VS, Hartig PC, Ankley GT, Foster PM, Gray CL, Gray LE. Fifteen years after "Wingspread"--environmental endocrine disrupters and human and wildlife health: where we are today and where we need to go. Toxicol Sci. 2008; 105: 235-59.
Lawson CC, Schnorr TM, Daston GP, Grajewski B, Marcus M, McDiarmid M, Murono E, Perreault SD, Schrader SM, Shelby M. An occupational reproductive research agenda for the third millennium. Environ Health Perspect. 2003; 111: 584-92.
Moline JM, Golden AL, Bar-Chama N, Smith E, Rauch ME, Chapin RE, Perreault SD, Schrader SM, Suk Wa, Landrigan PJ. Exposure to hazardous substances and male reproductive health: a research framework. Environ Health Perspect. 2000; 108: 803-13.
Perreault SD and Cancel AM. 2001. Significance of incorporating measures of sperm production and function into rat toxicology studies. Reproduction. 2001; 121: 207-16.
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Copyright © American Society of Andrology Handbook of Andrology – Can we protect spermatogenesis against testicular insults?
30-1 30-2
Stem spermatogonium
Differentiatingspermatogonium
Spermatocyte
Spermatid
Sperm
Stem spermatogonium
Differentiatingspermatogonium
Spermatocyte
Spermatid
Sperm
Serto
li ce
ll
Peritubular cell
Leydig cell
Serto
li ce
llSe
rtoli
cell
Peritubular cellPeritubular cell
Leydig cellLeydig cell
Total cell death by apoptosis
Cell killing
Anticancer Drugs(alkylating agents)
& Radiation
Anticancer Drugs(alkylating agents)
& Radiation
Thiols, antioxidants
Thiols, antioxidants
Hormone suppression
Protection or restoration of ability to support differentiation of
surviving stem spermatogoniaHormone
suppression
Protection or restoration of ability to support differentiation of
surviving stem spermatogonia
Antiapoptotic agents
Antiapoptotic agents
Can we protect spermatogenesis against testicular insults?
Male reproductive toxicology
M.L. Meistrich
Exposures to testicular insults
The germinal epithelium of the human testis is often subjected to chemical and physical insults as a result of medical, occupational, and environmental exposures. Among necessary medical treatments, anticancer agents and immunosuppressive drugs, particularly radiation and alkylating drugs, frequently cause temporary or permanent reductions in sperm count (oligospermia), and may even result in complete lack of sperm (azoospermia). Industrial solvents, such as bromopropane, and compounds released from plastics used in food containers and medical tubing, such as phthalates and bisphenol A, also damage spermatogenic function. Anabolic steroids, hormone antagonists, and environmental compounds such as bisphenol A or the fungicide vinclozolin can disrupt hormone signals essential for spermatogenesis (Chapters 28,29). Methods to prevent these damaging effects and to restore gonadal function after the toxic treatment are of great importance to men who desire to have children.
Nature of testicular insults
Some examples of the chemical, cellular, or endocrine nature of toxic insults are given below.
Radiation and alkylating agents, including those used in cancer chemotherapy, and such environmental toxicants as dibromochloropropane, induce free radicals, reactive oxygen species, and reactive alkyl groups that directly damage the DNA. These reactive intermediates can be scavenged and chemically detoxified, which prevents them from even producing the damage to DNA.
The anticancer agents preferentially kill proliferating cells, which in the testis are specific germ cells, primarily the rapidly proliferating differentiating spermatogonia, but the more slowly proliferating stem spermatogonia are somewhat susceptible (Fig. 1). As the surviving cells progress through spermatogenesis with fixed kinetics, the gap left by killing of spermatogonia will result in temporary depletion of later germ cells and reductions in sperm counts. But recovery should occur from the surviving stem cells within several months.
Killing of the stem cells reduces the potential for recovery and results in prolonged or permanent azoospermia. In addition, the somatic cells of the testis, which in adults are generally nonproliferating, survive cytotoxic exposures but suffer functional damage that may hinder the recovery of spermatogenesis from the surviving stem spermatogonia. One of these somatic cells, the Sertoli cell, is essential for the structural organization, nourishment, and regulation of the germ cells and is a direct target of some toxicants (phthalates). Damage to these somatic cells often leads to apoptotic death of the germ cells.
Endocrine disrupting chemicals can, by several different mechanisms, reduce the production or action of testosterone, which is essential in high concentrations in the testis for spermatogenesis to proceed (Chapter 8). Anabolic steroid analogs can inhibit testosterone production in the Leydig cells, 5-alpha reductase inhibitors can reduce testosterone activation, and vinclozolin and other antiandrogens can block testosterone action. These agents can affect fetal and pubertal testicular development and have irreversible effects on sperm output in later adulthood. FIG. 1. Diagrammatic representation of action of anticancer agents on specific cells of the testis and outcomes. Different targets for protection are shown: blocking the initial damage (thiols, antioxidants), restoring somatic cell function to enable surviving spermatogonia to reinitiate spermatogenesis (hormone suppression), and enabling the damaged cells to survive by blocking apoptosis (anti-apoptotic).
Various protective approaches
Methods to prevent these damaging effects and to restore gonadal function after the toxic treatment have been tested by a variety of physical, biochemical, and biological approaches in experimental animals. The only
Copyright © American Society of Andrology Handbook of Andrology – Can we protect spermatogenesis against testicular insults?
30-1 30-2
Stem spermatogonium
Differentiatingspermatogonium
Spermatocyte
Spermatid
Sperm
Stem spermatogonium
Differentiatingspermatogonium
Spermatocyte
Spermatid
Sperm
Serto
li ce
ll
Peritubular cell
Leydig cell
Serto
li ce
llSe
rtoli
cell
Peritubular cellPeritubular cell
Leydig cellLeydig cell
Total cell death by apoptosis
Cell killing
Anticancer Drugs(alkylating agents)
& Radiation
Anticancer Drugs(alkylating agents)
& Radiation
Thiols, antioxidants
Thiols, antioxidants
Hormone suppression
Protection or restoration of ability to support differentiation of
surviving stem spermatogoniaHormone
suppression
Protection or restoration of ability to support differentiation of
surviving stem spermatogonia
Antiapoptotic agents
Antiapoptotic agents
Can we protect spermatogenesis against testicular insults?
Male reproductive toxicology
M.L. Meistrich
Exposures to testicular insults
The germinal epithelium of the human testis is often subjected to chemical and physical insults as a result of medical, occupational, and environmental exposures. Among necessary medical treatments, anticancer agents and immunosuppressive drugs, particularly radiation and alkylating drugs, frequently cause temporary or permanent reductions in sperm count (oligospermia), and may even result in complete lack of sperm (azoospermia). Industrial solvents, such as bromopropane, and compounds released from plastics used in food containers and medical tubing, such as phthalates and bisphenol A, also damage spermatogenic function. Anabolic steroids, hormone antagonists, and environmental compounds such as bisphenol A or the fungicide vinclozolin can disrupt hormone signals essential for spermatogenesis (Chapters 28,29). Methods to prevent these damaging effects and to restore gonadal function after the toxic treatment are of great importance to men who desire to have children.
Nature of testicular insults
Some examples of the chemical, cellular, or endocrine nature of toxic insults are given below.
Radiation and alkylating agents, including those used in cancer chemotherapy, and such environmental toxicants as dibromochloropropane, induce free radicals, reactive oxygen species, and reactive alkyl groups that directly damage the DNA. These reactive intermediates can be scavenged and chemically detoxified, which prevents them from even producing the damage to DNA.
The anticancer agents preferentially kill proliferating cells, which in the testis are specific germ cells, primarily the rapidly proliferating differentiating spermatogonia, but the more slowly proliferating stem spermatogonia are somewhat susceptible (Fig. 1). As the surviving cells progress through spermatogenesis with fixed kinetics, the gap left by killing of spermatogonia will result in temporary depletion of later germ cells and reductions in sperm counts. But recovery should occur from the surviving stem cells within several months.
Killing of the stem cells reduces the potential for recovery and results in prolonged or permanent azoospermia. In addition, the somatic cells of the testis, which in adults are generally nonproliferating, survive cytotoxic exposures but suffer functional damage that may hinder the recovery of spermatogenesis from the surviving stem spermatogonia. One of these somatic cells, the Sertoli cell, is essential for the structural organization, nourishment, and regulation of the germ cells and is a direct target of some toxicants (phthalates). Damage to these somatic cells often leads to apoptotic death of the germ cells.
Endocrine disrupting chemicals can, by several different mechanisms, reduce the production or action of testosterone, which is essential in high concentrations in the testis for spermatogenesis to proceed (Chapter 8). Anabolic steroid analogs can inhibit testosterone production in the Leydig cells, 5-alpha reductase inhibitors can reduce testosterone activation, and vinclozolin and other antiandrogens can block testosterone action. These agents can affect fetal and pubertal testicular development and have irreversible effects on sperm output in later adulthood. FIG. 1. Diagrammatic representation of action of anticancer agents on specific cells of the testis and outcomes. Different targets for protection are shown: blocking the initial damage (thiols, antioxidants), restoring somatic cell function to enable surviving spermatogonia to reinitiate spermatogenesis (hormone suppression), and enabling the damaged cells to survive by blocking apoptosis (anti-apoptotic).
Various protective approaches
Methods to prevent these damaging effects and to restore gonadal function after the toxic treatment have been tested by a variety of physical, biochemical, and biological approaches in experimental animals. The only
Handbook of Andrology – Can we protect spermatogenesis against testicular insults? Handbook of Andrology – Can we protect spermatogenesis against testicular insults?
30-3 30-4
example of protection from testicular injury that has been reliably used in humans is shielding of the testes during radiation therapy. The testes can be well-shielded from the direct radiation beam, although scattered radiation still presents some concerns.
Among the methods that have been limited to experimental animals are free radical scavengers, antioxidants, and reducing compounds, including thiols; these can protect against the primary damage (Fig. 1). Amifostine, originally designed to selectively protect cells against free radicals produced by radiation, protects some normal tissues but not tumors against chemotherapy. Although it does partially protect mouse spermatogenesis against radiation, amifostine shows direct toxicity to stem spermatogonia, which in conventional fractionated anticancer regimens would outweigh the protective benefit.
Reducing blood flow to testes by transient ligation or cryptorchidism has been investigated to protect against damage by chemotherapy drugs with short half-lives. Indeed reversible ligation does protect animal testes against adriamycin and produced minimal direct damage, but its potential to cause ischemic damage remains a concern.
Anti-apoptotic compounds, such as sphingosine-1-phosphate, have been used against toxicants that kill germ cells by apoptosis. However, this does not offer significant protection of the testes.
Other empirical approaches, including administration of prostaglandins, vitamins, growth factors, and cytokines, have been tested in animals, and have demonstrated some protection against toxic effects. However, there is no evidence of therapeutic benefit against anticancer agents, i.e., that these treatments would provide more protection of the testis than of the tumors themselves.
Hormonal approaches to protection
The concept of hormone suppression for protecting gonadal function from cytotoxic exposure was based on the greater resistance of non-cycling cells than rapidly proliferating cells to killing by antineoplastic agents. It was erroneously thought that suppression of luteinizing hormone (LH), follicle-stimulating hormone (FSH), and testosterone levels would reduce the proliferation of the target cells and render them more resistant to the toxicant. But suppression of these hormones only blocks the completion of spermatogenesis and has no effect on the proliferation of the crucial target cells, the stem spermatogonia.
Despite this incorrect theoretical basis, hormonal suppression of rats prior to, during, and even after exposure to chemotherapy or radiation dra-
matically enhances the subsequent recovery of spermatogenesis. However, this is a result of protection or restoration of the function of the somatic cells of the testis to support the differentiation of the surviving stem spermatogonia and not protection of these spermatogonia from being killed. Hormonal sup-pression can modestly stimulate recovery of spermatogenesis from cytotoxic insults in mice but not in non-human primates.
Clinical trials of protection or restimulation of spermatogenesis by hormonal suppression have generally been unsuccessful, with only one study showing protection. We will have to know more about the mechanisms underlying the protection and recovery of spermatogenesis by hormonal suppression in rodent models, and their interspecies similarities and differences, before we can more rationally apply the results to human spermatogenesis.
Conclusions
As yet there are no proven methods for protecting the human germinal epithelium when it is directly exposed to chemical or physical insults nor proven therapeutic options to improve spermatogenesis after damage has occurred. However, protection may be achieved by removal of germ cells prior to exposure and storage for later use. Sperm banking done prior to medical exposures routinely results in successful pregnancies. In experi-mental animals, cryopreservation of spermatogonial stem cells or testicular tissue, and later reintroduction into the testicular tubules or transplantation into a subcutaneous site, respectively, result in production of viable sperm that yield live offspring. Awareness of such novel research in experimental animals designed to either protect the germinal epithelium from toxic insults or restore its function after the insult could lead to their application to humans.
Suggested reading
Jahnukainen K, Jahnukainen T, Salmi TT, Svechnikov K, Eksborg S, Soder O. Amifostine protects against early but not late toxic effects of doxorubicin in infant rats. Cancer Res. 2001; 61: 6423-7.
Meistrich ML, Shetty G. Inhibition of spermatogonial differentiation by testosterone. J Androl. 2003; 24:135-48.
Meistrich ML, Vassilopoulou-Sellin R, Lipshultz LI. Gonadal dysfunction. In: DeVita VT, Hellman S, Rosenberg SA, eds. Cancer: Principles and Practice of Oncology. Philadelphia: Lippincott Williams & Wilkins. 2005; 2560-74.
Handbook of Andrology – Can we protect spermatogenesis against testicular insults? Handbook of Andrology – Can we protect spermatogenesis against testicular insults?
30-3 30-4
example of protection from testicular injury that has been reliably used in humans is shielding of the testes during radiation therapy. The testes can be well-shielded from the direct radiation beam, although scattered radiation still presents some concerns.
Among the methods that have been limited to experimental animals are free radical scavengers, antioxidants, and reducing compounds, including thiols; these can protect against the primary damage (Fig. 1). Amifostine, originally designed to selectively protect cells against free radicals produced by radiation, protects some normal tissues but not tumors against chemotherapy. Although it does partially protect mouse spermatogenesis against radiation, amifostine shows direct toxicity to stem spermatogonia, which in conventional fractionated anticancer regimens would outweigh the protective benefit.
Reducing blood flow to testes by transient ligation or cryptorchidism has been investigated to protect against damage by chemotherapy drugs with short half-lives. Indeed reversible ligation does protect animal testes against adriamycin and produced minimal direct damage, but its potential to cause ischemic damage remains a concern.
Anti-apoptotic compounds, such as sphingosine-1-phosphate, have been used against toxicants that kill germ cells by apoptosis. However, this does not offer significant protection of the testes.
Other empirical approaches, including administration of prostaglandins, vitamins, growth factors, and cytokines, have been tested in animals, and have demonstrated some protection against toxic effects. However, there is no evidence of therapeutic benefit against anticancer agents, i.e., that these treatments would provide more protection of the testis than of the tumors themselves.
Hormonal approaches to protection
The concept of hormone suppression for protecting gonadal function from cytotoxic exposure was based on the greater resistance of non-cycling cells than rapidly proliferating cells to killing by antineoplastic agents. It was erroneously thought that suppression of luteinizing hormone (LH), follicle-stimulating hormone (FSH), and testosterone levels would reduce the proliferation of the target cells and render them more resistant to the toxicant. But suppression of these hormones only blocks the completion of spermatogenesis and has no effect on the proliferation of the crucial target cells, the stem spermatogonia.
Despite this incorrect theoretical basis, hormonal suppression of rats prior to, during, and even after exposure to chemotherapy or radiation dra-
matically enhances the subsequent recovery of spermatogenesis. However, this is a result of protection or restoration of the function of the somatic cells of the testis to support the differentiation of the surviving stem spermatogonia and not protection of these spermatogonia from being killed. Hormonal sup-pression can modestly stimulate recovery of spermatogenesis from cytotoxic insults in mice but not in non-human primates.
Clinical trials of protection or restimulation of spermatogenesis by hormonal suppression have generally been unsuccessful, with only one study showing protection. We will have to know more about the mechanisms underlying the protection and recovery of spermatogenesis by hormonal suppression in rodent models, and their interspecies similarities and differences, before we can more rationally apply the results to human spermatogenesis.
Conclusions
As yet there are no proven methods for protecting the human germinal epithelium when it is directly exposed to chemical or physical insults nor proven therapeutic options to improve spermatogenesis after damage has occurred. However, protection may be achieved by removal of germ cells prior to exposure and storage for later use. Sperm banking done prior to medical exposures routinely results in successful pregnancies. In experi-mental animals, cryopreservation of spermatogonial stem cells or testicular tissue, and later reintroduction into the testicular tubules or transplantation into a subcutaneous site, respectively, result in production of viable sperm that yield live offspring. Awareness of such novel research in experimental animals designed to either protect the germinal epithelium from toxic insults or restore its function after the insult could lead to their application to humans.
Suggested reading
Jahnukainen K, Jahnukainen T, Salmi TT, Svechnikov K, Eksborg S, Soder O. Amifostine protects against early but not late toxic effects of doxorubicin in infant rats. Cancer Res. 2001; 61: 6423-7.
Meistrich ML, Shetty G. Inhibition of spermatogonial differentiation by testosterone. J Androl. 2003; 24:135-48.
Meistrich ML, Vassilopoulou-Sellin R, Lipshultz LI. Gonadal dysfunction. In: DeVita VT, Hellman S, Rosenberg SA, eds. Cancer: Principles and Practice of Oncology. Philadelphia: Lippincott Williams & Wilkins. 2005; 2560-74.
Handbook of Andrology – Can we protect spermatogenesis against testicular insults?
30-5
Meistrich ML, Zhang Z, Porter KL, Bolden-Tiller OU, Shetty G. Prevention of adverse effects of cancer treatment on the germline. In: Anderson D, Brinkworth MH, eds. Male-Mediated Developmental Toxicity. Cambridge: Royal Society of Chemistry. 2007; 114-23.
Geens M, Goossens E, De Block G, Ning L, Van Saen D, Tournaye H. Autologous spermatogonial stem cell transplantation in man: current obstacles for a future clinical application. Hum Reprod Update. 2008; 14: 121-30.
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Copyright © Amerian Society of Andrology Handbook of Andrology – What are the consequences of sexually transmitted infections on male reproduction?
32-1 32-2
What are the consequences of sexually transmitted infections on male reproduction?
F.R. Ochsendorf
Sexually transmitted infections/diseases (STI) are caused by organisms sharing the following properties: 1) very sensitive to physical/chemical factors, hence only transmittable by direct contact, 2) only infectious for humans, 3) colonizing the genital region, 4) often causing only mild symptoms and 5) not leading to immunity. As a consequence concurrent infections with different STD-agents may occur at the same time (Table 1).
Mechanism of damage
Fertility may be impaired by organ damage due to the organism itself, cell damage via mediators of inflammation, inflammatory obstruction of the excurrent ductal system, or binding to spermatozoa. Thus, with respect to the impact on male reproductive system, chronic or inadequately treated STI’s are more relevant than the acute infections.
Relevance
The difference in the prevalence of STI's in different regions of the world as well as the access to their adequate diagnosis and treatment explain conflicting data on the relevance of these infections in male infertility. In western countries STI’s only account for a minority of inflammatory damage in the reproductive tract. However, In other parts of the world STI’s can severely impair male fertility: men with a history of penile discharge, painful micturition and genital ulcers, who did not seek adequate treatment for these symptoms, are more likely to be infertile than men without these symptoms or are adequately treated. Recently it was shown that 18.7% of semen samples (45/ 241) of asymptomatic men seeking infertility investigation contained DNA from STI-pathogens (e.g., CMV, HPV, HSV, HHV6, EBV, C. trachomatis). There was no difference in prevalence between samples with or without leukocytospermia. The presence of STI-DNA in semen samples was found to be associated with impairment in semen parameters (e.g., concentration, motile sperm concentration, total sperm count and neutral alpha-glucosidase concentration). However it has to be kept in mind that the presence of a germ does not necessarily mean “infection”, as the pathogen may just colonize the tissue without causing a clinical infection with pathological sequelae. This fact may also explain some conflicting data in the literature. Another impact of STI’s is their potential to increase the rate of
HIV-transmission. Thus, adequate diagnosis and treatment of STI’s is not only relevant for the prevention of long-term negative consequences for fertility but also for the prevention of HIV-spreading.
Relevant infections
Various common pathogens that can be isolated in the male reproductive tract are listed in Table1. Currently, literature supports the association of a significant negative impact on male fertility with infection of gonococci and HIV, while the roles of other pathogens such as chlamydia trachomatis, ureaplasma urealyticum, HSV, HPV and trichomonas vaginalis in impairing male fertility remain controversial.
Bacteria and protozoa A review of the literature could not demonstrate a general link between
urethritis and male infertility. Gonorrhoea, however, can cause urethral strictures as well as a strong impairment of testicular functions. In female, C. trachomatis infection can impair fertility by tubal obstruction. On the other hand, its impact on male fertility remains to be established although some epidemiological data points to the risk of subfertility in men (and women) with a past C. trachomatis infection, The exact role of Mycoplasmae, i.e. M. hominis, Ureaplasma urealyticum and M. genitalium has still to be elucidated. U. urealyticum may cause infertility via deleterious effects on sperm chromatin and DNA integrity, leading to impairment of embryo development. Trichomonas vaginalis was more often found in infertile men than in fertile controls; however, no effect on motility or sperm-mucus interaction has been demonstrated (Soper 2004).
Viruses At present the clinical relevance of viruses as a cause of male infertility is
largely unknown. Cytomegalovirus (CMV) and human herpes virus type 6 (HHV-6) were demonstrated in semen without an association with impaired semen parameters. The data on the relevance of herpes simplex virus (HSV) as well as human papilloma virus (HPV) and their associations with sperm parameters are conflicting (Ochsendorf 2006).
HIV. Infected leukocytes are the most relevant source of human-
immunodeficiency virus (HIV) in the male reproductive tract. In AIDS patients oligozoospermia, azoospermia, orchitis and hypogonadism were reported. Spermatozoa morphology is particularly impaired with progression of the disease. Disturbed function of the seminal vesicles and prostate gland may explain the decrease in semen volume and increase in ejaculate viscosity. With the latest advances in drug-therapies, HIV infection is considered a chronic disease. The safety of both the uninfected partner and the potential
Copyright © Amerian Society of Andrology Handbook of Andrology – What are the consequences of sexually transmitted infections on male reproduction?
32-1 32-2
What are the consequences of sexually transmitted infections on male reproduction?
F.R. Ochsendorf
Sexually transmitted infections/diseases (STI) are caused by organisms sharing the following properties: 1) very sensitive to physical/chemical factors, hence only transmittable by direct contact, 2) only infectious for humans, 3) colonizing the genital region, 4) often causing only mild symptoms and 5) not leading to immunity. As a consequence concurrent infections with different STD-agents may occur at the same time (Table 1).
Mechanism of damage
Fertility may be impaired by organ damage due to the organism itself, cell damage via mediators of inflammation, inflammatory obstruction of the excurrent ductal system, or binding to spermatozoa. Thus, with respect to the impact on male reproductive system, chronic or inadequately treated STI’s are more relevant than the acute infections.
Relevance
The difference in the prevalence of STI's in different regions of the world as well as the access to their adequate diagnosis and treatment explain conflicting data on the relevance of these infections in male infertility. In western countries STI’s only account for a minority of inflammatory damage in the reproductive tract. However, In other parts of the world STI’s can severely impair male fertility: men with a history of penile discharge, painful micturition and genital ulcers, who did not seek adequate treatment for these symptoms, are more likely to be infertile than men without these symptoms or are adequately treated. Recently it was shown that 18.7% of semen samples (45/ 241) of asymptomatic men seeking infertility investigation contained DNA from STI-pathogens (e.g., CMV, HPV, HSV, HHV6, EBV, C. trachomatis). There was no difference in prevalence between samples with or without leukocytospermia. The presence of STI-DNA in semen samples was found to be associated with impairment in semen parameters (e.g., concentration, motile sperm concentration, total sperm count and neutral alpha-glucosidase concentration). However it has to be kept in mind that the presence of a germ does not necessarily mean “infection”, as the pathogen may just colonize the tissue without causing a clinical infection with pathological sequelae. This fact may also explain some conflicting data in the literature. Another impact of STI’s is their potential to increase the rate of
HIV-transmission. Thus, adequate diagnosis and treatment of STI’s is not only relevant for the prevention of long-term negative consequences for fertility but also for the prevention of HIV-spreading.
Relevant infections
Various common pathogens that can be isolated in the male reproductive tract are listed in Table1. Currently, literature supports the association of a significant negative impact on male fertility with infection of gonococci and HIV, while the roles of other pathogens such as chlamydia trachomatis, ureaplasma urealyticum, HSV, HPV and trichomonas vaginalis in impairing male fertility remain controversial.
Bacteria and protozoa A review of the literature could not demonstrate a general link between
urethritis and male infertility. Gonorrhoea, however, can cause urethral strictures as well as a strong impairment of testicular functions. In female, C. trachomatis infection can impair fertility by tubal obstruction. On the other hand, its impact on male fertility remains to be established although some epidemiological data points to the risk of subfertility in men (and women) with a past C. trachomatis infection, The exact role of Mycoplasmae, i.e. M. hominis, Ureaplasma urealyticum and M. genitalium has still to be elucidated. U. urealyticum may cause infertility via deleterious effects on sperm chromatin and DNA integrity, leading to impairment of embryo development. Trichomonas vaginalis was more often found in infertile men than in fertile controls; however, no effect on motility or sperm-mucus interaction has been demonstrated (Soper 2004).
Viruses At present the clinical relevance of viruses as a cause of male infertility is
largely unknown. Cytomegalovirus (CMV) and human herpes virus type 6 (HHV-6) were demonstrated in semen without an association with impaired semen parameters. The data on the relevance of herpes simplex virus (HSV) as well as human papilloma virus (HPV) and their associations with sperm parameters are conflicting (Ochsendorf 2006).
HIV. Infected leukocytes are the most relevant source of human-
immunodeficiency virus (HIV) in the male reproductive tract. In AIDS patients oligozoospermia, azoospermia, orchitis and hypogonadism were reported. Spermatozoa morphology is particularly impaired with progression of the disease. Disturbed function of the seminal vesicles and prostate gland may explain the decrease in semen volume and increase in ejaculate viscosity. With the latest advances in drug-therapies, HIV infection is considered a chronic disease. The safety of both the uninfected partner and the potential
Handbook of Andrology – What are the consequences of sexually transmitted Handbook of Andrology – What are the consequences of sexually transmitted infections on male reproduction? infections on male reproduction?
32-3 32-4
offspring are of particular concern. In a sero-discordant couple the risk of acquiring HIV is dependent on the viral load in semen. Depending on the semen quality, spermatozoa can be used for intrauterine insemination, IVF or ICSI after adequate preparation of the ejaculate. The semen has to be washed free of HIV by a gradient technique and the success of this procedure has to be controlled prior to use. A thorough counselling of both partners on the therapeutic options, psychosocial and economic aspects with respect to their sexual health and reproductive health is thus necessary.
Table 1: Sexually transmitted infections: pathogens and relevance for male fertility; + = clinical relevance demonstrated;
? = relevance possible but not yet proven
Disease Pathogen Relevance for infertilityBacteria Gonorrhoea Neisseria gonorrhoea Male +
Female + Chlamydia infection Chlamydia trachomatis
(D-K) Male ?
Female + Urethritis (due to) Ureaplasma urealyticum May impair motility,
sperm chromatin ? Syphilis Treponema pallidum Co-factor for transmission of HIV
Gummatous testicular lesions: +Female +
Chancroid Haemophilus ducrey - Lymphogranuloma venereum
Chlamydia trachomatis (L1-L3)
-
Granuloma inguinale Calymmatobacterium granulomatis
-
Viruses AIDS HIV Male + Mononucleosis CMV - Asymptomatic infection HSV Impaired semen quality ? Asymptomatic infection HPV Impaired semen quality ??
Asthenozoospermia Asymptomatic infection Adenovirus Infertility ? Mumps Mumps virus Testicular atrophy + Protozoa Urethritis (Prostatis, Epididymitis) due to
Trichomonas vaginalis ? controversial
Yeasts Balanitis, Urethritis (due to) Candida albicans -
Suggested reading
Bezold G, Politch JA, Kiviat NB, Kuypers JM, Wolff H, Anderson DJ. Prevalence of sexually transmissible pathogens in semen from asymptomatic male infertility patients with and without leukocytospermia. Fertil Steril. 2007; 87: 1087-1097.
Dejucq N, Jégou B. Viruses in the mammlian male genital tract and their effects on the reproductive system. Microbiol Mol Biol Rev. 2001; 65: 208 - 231.
Karinen L, Pouta A, Hartikainen AL, et al. Association between Chlamydia trachomatis antibodies and subfertility in the Northern Finland Birth Cohort 1966 (NFBC 1966), at the age of 31 years. Epidemiol Infect. 2004; 132: 977-984.
Ness RB, Markovic N, Carlson CL, Coughlin MT. Do men become infertile after having sexually transmitted urethritis? An epidemiologic examination. Fertil Steril. 1997; 68: 205-213.
Ochsendorf FR. Reproductive tract infections/sexually transmitted diseases. In: Schill WB, Comhaire FH, Hargreave TB, eds. Andrology for the clinician Berlin - Heidelberg - New York Springer. 2006: 125 – 130.
Okonofua F, Menakaya U, Onemu SO, Omo-Aghoja LO, Bergstrom S. A case-control study of risk factors for male infertility in Nigeria. Asian J Androl. 2005; 7: 351-361.
Paavonen J, Eggert-Kruse W. Chlamydia trachomatis: impact on human reproduction. Hum Reprod Update. 1999; 5: 433-447.
Reichart M, Kahane I, Bartoov B. In vivo and in vitro impairment of human and ram sperm nuclear chromatin integrity by sexually transmitted Ureaplasma urealyticum infection. Biol Reprod. 2000; 63: 1041-1048.
Soper D. Trichomoniasis: under control or undercontrolled? Am J Obstet Gynecol. 2004; 190: 281-290.
van Leeuwen E, Prins JM, Jurriaans S, et al. Reproduction and fertility in human immunodeficiency virus type-1 infection. Hum Reprod Update. 2007; 13: 197-206.
Handbook of Andrology – What are the consequences of sexually transmitted Handbook of Andrology – What are the consequences of sexually transmitted infections on male reproduction? infections on male reproduction?
32-3 32-4
offspring are of particular concern. In a sero-discordant couple the risk of acquiring HIV is dependent on the viral load in semen. Depending on the semen quality, spermatozoa can be used for intrauterine insemination, IVF or ICSI after adequate preparation of the ejaculate. The semen has to be washed free of HIV by a gradient technique and the success of this procedure has to be controlled prior to use. A thorough counselling of both partners on the therapeutic options, psychosocial and economic aspects with respect to their sexual health and reproductive health is thus necessary.
Table 1: Sexually transmitted infections: pathogens and relevance for male fertility; + = clinical relevance demonstrated;
? = relevance possible but not yet proven
Disease Pathogen Relevance for infertilityBacteria Gonorrhoea Neisseria gonorrhoea Male +
Female + Chlamydia infection Chlamydia trachomatis
(D-K) Male ?
Female + Urethritis (due to) Ureaplasma urealyticum May impair motility,
sperm chromatin ? Syphilis Treponema pallidum Co-factor for transmission of HIV
Gummatous testicular lesions: +Female +
Chancroid Haemophilus ducrey - Lymphogranuloma venereum
Chlamydia trachomatis (L1-L3)
-
Granuloma inguinale Calymmatobacterium granulomatis
-
Viruses AIDS HIV Male + Mononucleosis CMV - Asymptomatic infection HSV Impaired semen quality ? Asymptomatic infection HPV Impaired semen quality ??
Asthenozoospermia Asymptomatic infection Adenovirus Infertility ? Mumps Mumps virus Testicular atrophy + Protozoa Urethritis (Prostatis, Epididymitis) due to
Trichomonas vaginalis ? controversial
Yeasts Balanitis, Urethritis (due to) Candida albicans -
Suggested reading
Bezold G, Politch JA, Kiviat NB, Kuypers JM, Wolff H, Anderson DJ. Prevalence of sexually transmissible pathogens in semen from asymptomatic male infertility patients with and without leukocytospermia. Fertil Steril. 2007; 87: 1087-1097.
Dejucq N, Jégou B. Viruses in the mammlian male genital tract and their effects on the reproductive system. Microbiol Mol Biol Rev. 2001; 65: 208 - 231.
Karinen L, Pouta A, Hartikainen AL, et al. Association between Chlamydia trachomatis antibodies and subfertility in the Northern Finland Birth Cohort 1966 (NFBC 1966), at the age of 31 years. Epidemiol Infect. 2004; 132: 977-984.
Ness RB, Markovic N, Carlson CL, Coughlin MT. Do men become infertile after having sexually transmitted urethritis? An epidemiologic examination. Fertil Steril. 1997; 68: 205-213.
Ochsendorf FR. Reproductive tract infections/sexually transmitted diseases. In: Schill WB, Comhaire FH, Hargreave TB, eds. Andrology for the clinician Berlin - Heidelberg - New York Springer. 2006: 125 – 130.
Okonofua F, Menakaya U, Onemu SO, Omo-Aghoja LO, Bergstrom S. A case-control study of risk factors for male infertility in Nigeria. Asian J Androl. 2005; 7: 351-361.
Paavonen J, Eggert-Kruse W. Chlamydia trachomatis: impact on human reproduction. Hum Reprod Update. 1999; 5: 433-447.
Reichart M, Kahane I, Bartoov B. In vivo and in vitro impairment of human and ram sperm nuclear chromatin integrity by sexually transmitted Ureaplasma urealyticum infection. Biol Reprod. 2000; 63: 1041-1048.
Soper D. Trichomoniasis: under control or undercontrolled? Am J Obstet Gynecol. 2004; 190: 281-290.
van Leeuwen E, Prins JM, Jurriaans S, et al. Reproduction and fertility in human immunodeficiency virus type-1 infection. Hum Reprod Update. 2007; 13: 197-206.
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