HANDBOOK OF ANDROLOGY

HANDBOOK OF
ANDROLOGY
Second Edition
The American Society of Andrology
Copyright © Marc
Printed by Allen Press, In

F
ogy
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
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
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 Institutions
Baltimore, MA 21287
USA
Mary S. Croughan, Ph.D
Ob/Gyn Women’s Hlth C
University of California
at San Francisco
Box 1793, 1635 Divisade
San Francisco, CA 9414
USA
Janice P. Evans, Ph.D.
Division of Reproductive
Department of Biochemis
Molecular Biology
Bloomberg School of Pub
Baltimore, MD 21205
USA
Marc Goldstein, M.D.
Cornell Institute for Repr
Medicine
Weill-Cornell Medical Co
Box 580, 525 E 68th Stre
New York, NY 10065
Mary Ann Handel, Ph.D.
The Jackson Laboratory
600 Main Street
Bar Harbor, Maine 0460
USA
Barry T. Hinton, Ph.D.*
Department of Cell Biolo
University of Virginia Hea
Sciences Center
PO Box 800732
School Of Medicine #439
Charlottesville, VA 2290
USA

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ery
tal
est, S6.95
1A1
in
ontraception
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gton
6429
M.D., Ph.D.*
gton School
et
6420
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itute
cal Institutions
7
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.
Therapeutics
McGill University
3655 Prom. Sir-William-Osler
Canada
Janice P. Evans, Ph.D.
Molecular Biology
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.
Therapeutics
McGill University
3655 Promenade Sir-William-Osler
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 – L
!"
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.
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.
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,
Departments of Medicine
Pharmacology and The
The Research Institute of
McGill University Healt
McGill University
1650 Cedar Avenue
Montreal, QC H3G 1A4
Canada
Sally D. Perreault, Ph.D.*
US Environmental Protec
Agency
Mail Drop E205-09
Research Triangle Park N
USA
Tony M. Plant, Ph.D.
University of Pittsburgh S
of Medicine
Dept. of Cell Biology and
Magee-Womens Researc
204 Craft Avenue, Room
Pittsburgh, PA 15213
USA
Mathew C. Raynor, M.D.
Tulane Health Sciences C
1430 Tulane Avenue
USA
Susan A. Rothmann, Ph.
Fertility Solutions Inc.
11811 Shaker Blvd., Suit
Cleveland, OH 44120
USA

Handbook of Andrology – List of Contributors
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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.
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 27711
USA
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 Physiology
Magee-Womens Research Institute
204 Craft Avenue, Room B311
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.
Tulane Health Sciences Center
1430 Tulane Avenue
USA
Kenneth Roberts, Ph.D.
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.
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 – Li
VI
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
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.
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-CHUL
Quebec, QC G1V 4G2
Canada
Thomas J. Walsh, M.D.
of Medicine
1959 NE Pacific Street, # Bb667
USA
Christina Wang, M.D.*
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 Pitt
USA
Barry R. Zirkin, Ph.D.*
Division of Reproductive
Department of Biochemis
Molecular Biology
Bloomberg School of Pub
Baltimore, MD 21205
USA
*Past President, America

Handbook of Andrology – List of Contributors
VII
des, Ph.D.
terinary
treal
J2S 7C6
off, M.D.*
rinology
edicine
dical Center
es Biomedical
er
Street
509
M.D.
ology
Texas Health
San Antonio
Drive
78229
Ph.D.
evard, T1-49
che CHUQ-CHUL
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M.D.*
rinology
edicine
dical Center and
omedical
er
Street
509
Selma F. Witchel, M.D.
Children’s Hospital of Pittsburgh
USA
William W. Wright, Ph.D.
Molecular Biology
Baltimore, MD 21205
USA
Barry R. Zirkin, Ph.D.*
Molecular Biology
Baltimore, MD 21205
USA
*Past President, American Society of Andrology

VIII
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
areas have been made
and scientific discoverie
biochemistry, neuroscien
each new stage of under
the stimulus that I and my
of the multidisciplinary
themselves and the oppo
expand. As Alexander A
with the reproductive s
opportunity. Andrology c
aging. As you peruse this
of the field and share our

IX
ndrology
316
ublished to bring
handbooks are
subject for those
d to codify the
each of these
e is to present to
mportance, and
drology is to the
eals with matters
se of the term
en in 1891 in the
the formation of
heard from that
ntury that there
ogy organization,
er century since,
ions, of societies
ed to andrology.
knowledge and
logy is strikingly
book. Written by
en to indicate the
ds for graduate
s, and medical
irst of all, tries to
y to succeed and
r 40 years, I can
has held for me.
owth so that our
nges in the male
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
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 product
Hormonal regulati
and FSH
R.I. McLachlan
9. Are there epigeneti
formation? What is
cells?
Imprinting, DNA m
J.M. Trasler
10. What does the epid
B.T. Hinton
11. What is the prostat
G.S. Prins
12. How does semen a
status of the male?
C. Wang
13. How is sperm chro
doing such assess
Range of methods
B. Robaire
14. What is sperm ban
in humans? Anima
Sperm banking, c
practice
S.A. Rothmann
15. How does the sper
fertilization take pl
Capacitation, acro
egg activation
J.P. Evans and J.
16. What determines m
Sex determination
phenotype
D.W. Silversides a

XI
system?
esicles, scrotum,
ne components
ps
productive
e testis? Where
trol their
ation of a
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 – Co
XII
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
disruptors on sperm
Chemical exposur
spermatozoa
S.D. Perreault
29. Can spermatozoa b
consequences of s
conception counse
Drugs that affect s
developmental tox
spermatozoa
B.F. Hales and B.
30. Can we protect spe
Male reproductive
M.L. Meistrich
31. What male contrace
for new male contra
J.K. Amory and W
32. What are the conse
male reproduction?
F. Ochsendorf
33. What types of testic
prognosis? How ef
N.E. Skakkebaek
34. How do erections o
what is its etiology,
problem?
P.M. Pierorazio an
35. How to treat erectil
M.C. Raynor and W
36. Is there an andropa
Syndrome, and if so
Fertility, androgen
function in aging m
A. Morales

Handbook of Andrology – Contents
XIII
reproductive
ms?
RH/LH/FSH,
ly or delayed
uation
male
hniques
ility?
ceived
on and food-
of animals?
e?
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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HANDBOOK OF ANDROLOGY

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