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Male Reproductive
Dysfunction
Pathophysiology and
Treatment

Edited by
FOUAD R. KANDEEL
City of Hope National Medical Center, Duarte, California, USA
David Geffen School of Medicine at the University of California, Los Angeles, USA

Associate Editors
RONALD S. SWERDLOFF
Harbor-UCLA Medical Center,Torrance, California, USA
David Geffen School of Medicine at the University of California, Los Angeles, USA
JON L. PRYOR
University of Minnesota Medical School, Minneapolis, USA
Informa Healthcare USA, Inc.
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© 2007 by Informa Healthcare USA, Inc.

Informa Healthcare is an Informa business

No claim to original U.S. Government works

Printed in the United States of America on acid-free paper
10 9 8 7 6 5 4 3 2 1

International Standard Book Number-10: 1-57444-848-X (Hardcover)

International Standard Book Number-13: 978-1-57444-848-1 (Hardcover)

This book contains information obtained from authentic and highly regarded sources. Reprinted
material is quoted with permission, and sources are indicated. A wide variety of references are listed.
Reasonable efforts have been made to publish reliable data and information, but the author and
the publisher cannot assume responsibility for the validity of all materials or for the consequence of
their use.

No part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any
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Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are
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Library of Congress Cataloging-in-Publication Data

Male reproductive dysfunction : pathophysiology and treatment / [edited by]

Fouad R. Kandeel.
p. ; cm.
Includes bibliographical references.
ISBN-13: 978-1-57444-848-1 (hardcover : alk. paper)
ISBN-10: 1-57444-848-X (hardcover : alk. paper)
1. Infertility, Male. 2. Impotence. I. Kandeel, Fouad R.
[DNM: 1. Infertility, Male. 2. Genital Diseases, Male--Complications.
3. Infertility, Male--therapy. WJ 709 M24555 2007]

RC889.M3558 2007
616.6'92--dc22 2007005802

Visit the Informa Web site at

www.informa.com

and the Informa Healthcare Web site at

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 This volume is dedicated to those who have helped shape the
path of my academic and professional career in the hope that the
information contained herein will enrich the knowledge of others.
Preface

Given the finite nature of the human lifespan, it is possible to argue that an
individual man’s most precious and lasting gift to the world is the legacy created
by his offspring. It is no wonder, then, that the purely reproductive function of
fertility has become inextricably tied to the human male’s impression of his own
masculinity. The fact of being childless is often an extremely emotional issue for
patients and their families, and ameliorating such a cause is certainly worthy of
our efforts. Each day, great strides are made in the ever-evolving body of knowl-
edge that comprises the whole science and practice of medicine, allowing men to
live longer, healthier lives; thus, retaining sexual and reproductive health even
well into their later years has taken on increased importance for many men, and,
consequently, for the physicians who treat them.
Yet, with the benefit of increased longevity, there must simultaneously exist
an increased likelihood that the male reproductive tract will be subjected to
greater levels of stress resulting from longer exposure to harmful environmental
agents, poor lifestyle choices, and new diseases and infections. Additionally, the
therapeutic drugs and regimens prescribed for these conditions frequently dimin-
ish sexual and reproductive capacities to an even greater degree. For example, a
male patient who survives cancer and its treatment often falls victim to sexual,
reproductive, and interpersonal difficulties that may further compromise his
quality of life.
Due to the wide range of impairments that may impact a man’s reproductive
health throughout his lifespan, by necessity, the field of reproductive medicine is
a multidisciplinary specialty comprised of andrology, endocrinology, urology,
gynecology, radiology, psychology, and sociology. The vast array of data and the
complexity of new technologies constantly being produced in so many different
areas are often bewildering. Therefore, the chief goal in creating this first edition
of Male Reproductive Dysfunction: Pathophysiology and Treatment has been to assimi-
late the most pertinent and important scientific knowledge gained from the
complementary expertise of more than 70 individual leaders in all related disci-
plines into a single, comprehensive source, and to present this information from a
clinical viewpoint, not only for the benefit of those actively involved in managing
male patients with reproductive disorders, but also for the considerable numbers
of physicians and healthcare providers who are likely to encounter and care for
such men, regardless of etiology or precipitating factors. We hope to educate
our readers with both the science behind these complex disorders and to equip
them with practical tools for the effective and rational management of these
conditions.
Based on the most recent data available, Male Reproductive Dysfunction:
Pathophysiology and Treatment presents historical, psychosocial, behavioral, medical,
and surgical perspectives in an unparalleled 43-chapter guide to the assessment
and treatment of the male patient with reproductive dysfunction. Logically divided
into five parts, the book begins with an extensive review of the normal anatomy
and physiology of the male reproductive system, including a chapter on the effect
of aging on testosterone production. Building on this foundation, the second part
details the many etiologies, mechanisms, and clinical manifestations of male
reproductive disorders. Both common and rare conditions of male reproductive
dysfunction are covered, from genetic defects and childhood developmental abnor-
malities to other causes of hypogonadism, including the adverse effects of
substance abuse, malnutrition, environmental toxins, and the iatrogenic effects
of cancer treatments and other pharmacotherapies on the hypothalamic-pitui-
tary-testicular axis. Also highlighted are problems that occur later in life, such as
vi Preface

the complications of systemic diseases, obstructive and mass lesions of the

reproductive tract, and sperm allergy.
With this information well in hand, the third part invaluably outlines
strategies for the complete assessment of the hypogonadal and/or infertile male
patient, from obtaining the initial history and physical exam, to ordering blood-
work and semen analyses, to interpreting advanced imaging studies and per-
forming testicular biopsies. The multitude of illustrations, images, algorithms, and
tables elucidate these complex topics even more clearly. Of special note are the
chapters dedicated to the evaluation of immunological factors that may be contrib-
uting to infertility and the effective use of imaging technology in the diagnosis of
male reproductive tract pathology.
The fourth part then provides extensive guidelines for the treatment of male
reproductive dysfunction depending on the individual pathology, including spe-
cialized situations such as patients who are cancer survivors or have spinal cord or
other neurological injury. The details of various surgical sperm retrieval methods
as well as assisted reproduction techniques (ART) are described step-by-step.
Several chapters feature additional information that is unique to this book, such as
the latest progress in male contraception, micromanipulation procedures for the
treatment of the subfertile male, the psychological effects of infertility and its
treatment, and the management of immunological factors in male infertility.
The final, fifth part is devoted to a full discussion of the mass lesions of the
male reproductive tract, including testicular, penile, prostate, and male breast
cancers. The problems of prostate hyperplasia, prostatitis, and gynecomastia are
also addressed.
Most importantly, each chapter in this volume considers the future aims of its
particular topic, opening the doorway for continued thought and discussion as
new research is added to the current armamentarium. It is hoped that this book
will become an essential, self-contained desk reference for both new and seasoned
clinicians involved in the care of male patients with all forms of reproductive
dysfunction, and that by enriching the current body of knowledge, our work will
provide the starting point for future generations of medical professionals to learn,
study, develop, and discover.

Fouad R. Kandeel
Acknowledgments

The editors are very grateful to the following individuals and would like to acknowl-
edge their assistance in the preparation of this volume: Chakriya Anunta, Jeannette
Hacker, Bernard de la Cruz, Karen Ramos, and Angela Hacker.
Contents

Preface . . . . v
Acknowledgments . . . . vii
Contributors . . . . xiii

Part I. Physiology of Male Reproductive Function

Section I: History
1. Male Reproduction: Evolving Concepts of Procreation and
Infertility Through the Ages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Jeannette Hacker and Fouad R. Kandeel

Section II: Anatomy

2. Anatomy of the Male Reproductive Tract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Fouad R. Kandeel, Vivien Koussa, and Ewa Kuligowska

Section III: Physiology

3. Neuroendocrine Control of Testicular Function . . . . . . . . . . . . . . . . . . . . . . . 21
Stephen J. Winters
4. Analysis of the Neuroendocrine Control of the Human
Hypothalamo–Pituitary–Testicular Axis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Johannes D. Veldhuis and Daniel M. Keenan
5. Normal Spermatogenesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Jeffrey B. Kerr
6. Physiology of the Epididymis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Russell C. Jones and Jean-Louis Dacheux
7. Physiology of the Prostate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
Helen D. Nicholson and Stephen J. Assinder

Section IV: Aging

8. Testosterone and Aging . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Horace M. Perry III, John E. Morley, and Gary A. Wittert
9. Effects of Aging on Spermatogenesis and Sperm Function . . . . . . . . . . . . 101
Bernard Robaire, Christopher C. Oakes, and Ekaterina V. Zubkova

Section V: Psychological and Social Factors

10. Androgen Effects on Cognitive Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Monique M. Cherrier
11. Social and Psychological Influences on Male Reproductive Function . . . 129
William Marsiglio and Ramon Hinojosa

Part II. Pathophysiology of Male Reproductive Dysfunction

Section I: General Aspects
12. Male Hypogonadism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Ronald S. Swerdloff and Christina Wang

Section II: Developmental Disorders

13. Cryptorchidism, Testicular Torsion, and Torsion of the
Testicular Appendages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
Bartley G. Cilento, Jr. and Anthony Atala
14. Disorders of Male Puberty . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Erick J. Richmond and Alan D. Rogol
15. Male Varicoceles. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Benjamin K. Canales, Nissrine Nakib, Ann Lavers, and Jon L. Pryor
x Contents

Section III: Infections

16. Male Genital Tract Infections and Infertility . . . . . . . . . . . . . . . . . . . . . . . . . 197
Durwood E. Neal, Jr. and Stephen H. Weinstein
17. HIV and Male Fertility . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Timothy B. Hargreave and Chhanda Ghosh

Section IV: Ejaculatory Duct Disorder

18. Ejaculatory Duct Obstruction in the Infertile Male . . . . . . . . . . . . . . . . . . . 213
Erik T. Goluboff and Harry Fisch

Section V: Impact of Other Diseases

19. Male Hormones and Systemic Disease . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221
John E. Morley and Gary A. Wittert
20. The Effects of Chemotherapy and Radiotherapy on Testicular
Function . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235
Simon J. Howell and Steve M. Shalet

Section VI: Environmental Factors

21. Environmental Toxins and Male Fertility . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245
Rebecca Z. Sokol
22. Micronutrients and Male Fertility . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
Nan B. Oldereid

Part III. Investigation of Male Reproductive Dysfunction

Section I: General Assessment

23. Clinical Assessment of the Infertile Male . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
Chakriya D. Anunta and Fouad R. Kandeel
24. Semen Analysis and Sperm Function Testing . . . . . . . . . . . . . . . . . . . . . . . . 271
H. W. Gordon Baker, Gary N. Clarke, De Yi Liu,
Tanya M. Stewart, and Claire Garrett

Section II: Specialized Investigations

25. Testicular Biopsy of the Infertile Male . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 285
Brett C. Mellinger
26. Immunological Evaluation of Male Infertility . . . . . . . . . . . . . . . . . . . . . . . . 293
Gary N. Clarke and H. W. Gordon Baker
27. Genetic Evaluation of the Infertile Male . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
Edward D. Kim
28. Imaging of the Reproductive Tract in Male Infertility . . . . . . . . . . . . . . . . . 313
Ewa Kuligowska, Erik N. Nelson, and Helen M. Fenlon

Part IV. Treatment of Male Reproductive Dysfunction

Section I: Pharmacological Interventions

29. Androgen Replacement Therapy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 333
Ronald S. Swerdloff
30. Hormonal Induction of Spermatogenesis in Males with
Hypogonadotropic Hypogonadism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
Peter Y. Liu
31. Progress in Male Contraception . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 349
Christina Wang and Ronald S. Swerdloff

Section II: Treatment of Seminal Transport Disorders

32. Retrograde Ejaculation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359
Avner Hershlag and Sarah K. Girardi
33. Surgical Treatment of Male Infertility. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 365
Peter N. Schlegel and Jeremy Kaufman
34. Vasectomy and Vasectomy Reversal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Peter T. K. Chan and Marc Goldstein
 Contents xi

Section III: Psychological Issues in Male Infertility

35. Psychological Effects of Infertility and Its Treatment in Males . . . . . . . . . 407
Kenneth Gannon, Lesly Glover, and Paul David Abel

Section IV: Treatment of Special Problems in Male Infertility

36. Management of Immunological Factors in Male Infertility . . . . . . . . . . . . 419
H. W. Gordon Baker and Gary N. Clarke
37. Micromanipulation of Male Gametes in the Treatment of the
Subfertile Male . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 431
Soon-Chye Ng and Swee-Lian Liow
38. Fertility Preservation in Males with Cancer . . . . . . . . . . . . . . . . . . . . . . . . . . 453
Paul J. Turek
39. Treatment of Reproductive Dysfunction in Males with Spinal
Cord Injury and Other Neurologically Disabling Diseases . . . . . . . . . . . . 465
Khashayar Hematpour and Carol Bennett

Part V. Mass Lesions of the Male Reproductive System

Section I: Mass Lesions of the Penis and Testes

40. Testicular and Penile Cancer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481
John W. Davis, Donald F. Lynch, and Paul F. Schellhammer
41. Prostate Disease: Prostate Hyperplasia, Prostate Cancer, and Prostatitis . . . . 499
Stephen J. Assinder and Helen D. Nicholson

Section II: Mass Lesions of the Male Breast

42. Gynecomastia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 519
Ronald S. Swerdloff, Jason Ng, and Fouad R. Kandeel
43. Male Breast Cancer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 529
Sandeep K. Reddy and Lucille Leong

Index . . . . 537
Contributors

Paul David Abel Division of Surgery, Oncology, Reproduction and Anesthesia,

Imperial College School of Medicine, London, U.K.
Chakriya D. Anunta Department of Diabetes, Endocrinology and Metabolism,
City of Hope National Medical Center, Duarte, California, U.S.A.
Stephen J. Assinder Discipline of Physiology, School of Medical Sciences,
University of Sydney, New South Wales, Australia
Anthony Atala Department of Urology, Wake Forest University School of
Medicine, Wake Forest Institute for Regenerative Medicine, Winston-Salem,
North Carolina, U.S.A.
H. W. Gordon Baker Department of Obstetrics and Gynaecology, University of
Melbourne and The Royal Women’s Hospital, Carlton, Victoria, Australia
Carol Bennett Department of Urology, Greater Los Angeles VA Medical System
and David Geffen School of Medicine, University of California, Los Angeles,
California, U.S.A.
Benjamin K. Canales Department of Urologic Surgery, University of Minnesota
Medical School, Minneapolis, Minnesota, U.S.A.
Peter T. K. Chan Department of Urology, McGill University Health Centre,
Montreal, Quebec, Canada
Monique M. Cherrier Department of Psychiatry and Behavioral Sciences,
University of Washington School of Medicine and Veterans Administration Puget
Sound Health Care System, Seattle, Washington, U.S.A.
Bartley G. Cilento, Jr. Department of Urology, Children’s Hospital and
Department of Surgery, Harvard Medical School, Boston, Massachusetts, U.S.A.
Gary N. Clarke Andrology Laboratory Division of Laboratory Services,
The Royal Women’s Hospital, Carlton, Victoria, Australia
Jean-Louis Dacheux Equipe “Spermatozoides,” UMR INRA-CNRS 6073,
Institut National de la Recherche Agronomique, Nouzilly, France
John W. Davis Department of Urology, The University of Texas M.D. Anderson
Cancer Center, Houston, Texas, U.S.A.
Helen M. Fenlon Department of Radiology, Mater Misericordiae Hospital,
Dublin, Ireland
Harry Fisch College of Physicians and Surgeons, Columbia University and
Male Reproductive Center, Columbia-Presbyterian Medical Center, New York,
New York, U.S.A.
Kenneth Gannon School of Psychology, University of East London, Stratford,
London, U.K.
Claire Garrett Department of Obstetrics and Gynaecology, University of
Melbourne, and The Royal Women’s Hospital, Carlton, Victoria, Australia
Chhanda Ghosh Department of Oncology, University of Edinburgh,
Edinburgh, Scotland, U.K.
xiv Contributors

Sarah K. Girardi Department of Urology, Weill Medical College of Cornell

University, North Shore University Hospital, Manhasset, New York, U.S.A.
Lesly Glover Department of Clinical Psychology, The University of Hull,
Hull, U.K.
Marc Goldstein Cornell Institute for Reproductive Medicine, Department of
Urology, New York-Presbyterian Hospital and Weill Medical College of Cornell
University, New York, New York, U.S.A.
Erik T. Goluboff College of Physicians and Surgeons, Columbia University and
New York-Presbyterian Hospital, New York, New York, U.S.A.
Jeannette Hacker Department of Diabetes, Endocrinology and Metabolism,
City of Hope National Medical Center, Duarte, California, U.S.A.
Timothy B. Hargreave Department of Oncology, University of Edinburgh,
Western General Hospital, Fertility Problems Clinic, Edinburgh, Scotland, U.K.
Khashayar Hematpour St. Luke’s-Roosevelt Hospital Center, Columbia
University College of Physicians and Surgeons, New York, New York, U.S.A.
Avner Hershlag Department of Obstetrics and Gynecology, North Shore
University Hospital, New York University School of Medicine, Manhasset,
New York, U.S.A.

Ramon Hinojosa Department of Sociology, University of Florida, Gainesville,

Florida, U.S.A.
Simon J. Howell Department of Endocrinology, Christie Hospital NHS Trust,
Withington, Manchester, U.K.
Russell C. Jones Discipline of Biological Sciences, The University of Newcastle,
New South Wales, Australia
Fouad R. Kandeel Department of Diabetes, Endocrinology and
Metabolism, City of Hope National Medical Center, Duarte,
California and David Geffen School of Medicine, University of California,
Los Angeles, California, U.S.A.
Jeremy Kaufman Department of Urology, The New York-Presbyterian
Hospital, New York Center for Biomedical Research and The Rockefeller
University Hospital, New York, New York, U.S.A.
Daniel M. Keenan Department of Statistics, University of Virginia,
Charlottesville, Virginia, U.S.A.
Jeffrey B. Kerr Department of Anatomy and Cell Biology, Monash University,
Victoria, Australia
Edward D. Kim Department of Surgery, Division of Urology, University of
Tennessee Medical Center, Knoxville, Tennessee, U.S.A.
Vivien Koussa H.C. Healthcare Management and Development, Inc.,
Long Beach, California, U.S.A.
Ewa Kuligowska Department of Radiology, Boston Medical Center,
Boston, Massachusetts, U.S.A.
Ann Lavers Department of Urologic Surgery, University of Minnesota Medical
School, Minneapolis, Minnesota, U.S.A.
Lucille Leong Division of Medical Oncology, City of Hope National Medical
Center, Duarte, California, U.S.A.
Swee-Lian Liow Embryonics International, Gleneagles Hospital, Singapore
 Contributors xv

De Yi Liu Department of Obstetrics and Gynaecology, University of

Melbourne, and The Royal Women’s Hospital, Carlton, Victoria, Australia
Peter Y. Liu Division of Endocrinology, Harbor-UCLA Medical Center,
Torrance, California, U.S.A. and Department of Andrology, ANZAC Research
Institute, University of Sydney, New South Wales, Australia
Donald F. Lynch Department of Urology, East Virginia Medical School,
Norfolk, Virginia, U.S.A.
William Marsiglio Department of Sociology, University of Florida, Gainesville,
Florida, U.S.A.
Brett C. Mellinger Department of Clinical Urology, SUNY Stony Brook,
Stony Brook, New York, U.S.A.
John E. Morley Department of Medicine, Adelaide University, Australia and
Division of Geriatric Medicine, St. Louis University School of Medicine and
GRECC, St. Louis VA Medical Center, St. Louis, Missouri, U.S.A.
Nissrine Nakib Department of Urologic Surgery, University of Minnesota
Medical School, Minneapolis, Minnesota, U.S.A.
Durwood E. Neal, Jr. Department of Urology, University of Missouri
Medical School, Columbia, Missouri, U.S.A.
Erik N. Nelson Department of Radiology, Massachusetts General Hospital,
Boston, Massachusetts, U.S.A.
Jason Ng Division of Endocrinology, Department of Medicine, Queen Elizabeth
Hospital, Hong Kong, Special Administration Region, P.R. China
Soon-Chye Ng O & G Partners Fertility, Gleneagles Hospital, Singapore
Helen D. Nicholson Department of Anatomy and Structural Biology, Otago
School of Medical Sciences, University of Otago, Dunedin, New Zealand
Christopher C. Oakes Departments of Pharmacology and Therapeutics, and
Obstetrics and Gynecology, McGill University, Montreal, Quebec, Canada
Nan B. Oldereid Department of Obstetrics and Gynecology,
Rikshospitalet–Radiumhospitalet Medical Center, Oslo, Norway
Horace M. Perry III Department of Internal Medicine–Geriatrics, St. Louis
University School of Medicine, St. Louis, Missouri, U.S.A.
Jon L. Pryor Department of Urologic Surgery, University of Minnesota Medical
School, Minneapolis, Minnesota, U.S.A.
Sandeep K. Reddy Los Alamitos Hematology/Oncology Medical Group, Inc.,
Los Alamitos, California, U.S.A.
Erick J. Richmond Pediatric Oncology, National Children’s Hospital, San Jose,
Costa Rica
Bernard Robaire Departments of Pharmacology and Therapeutics, and
Obstetrics and Gynecology, McGill University, Montreal, Quebec, Canada
Alan D. Rogol Clinical Pediatrics, University of Virginia, Charlottesville,
Virginia, U.S.A.
Paul F. Schellhammer Department of Urology, Eastern Virginia Medical School,
Norfolk, Virginia and Prostate Cancer Center, Sentara Cancer Institute, Hampton,
Virginia, U.S.A.
Peter N. Schlegel Departments of Urology and Reproductive Medicine,
Weill Medical College of Cornell University, New York, New York, U.S.A.
xvi Contributors

Steve M. Shalet Department of Endocrinology, Christie Hospital NHS Trust,

Withington, Manchester, U.K.
Rebecca Z. Sokol Keck School of Medicine, University of Southern California,
Los Angeles, California, U.S.A.
Tanya M. Stewart Department of Obstetrics and Gynaecology, University of
Melbourne, and The Royal Women’s Hospital, Carlton, Victoria, Australia
Ronald S. Swerdloff Division of Endocrinology and Metabolism,
Harbor-UCLA Medical Center, Torrance, California and Department of
Medicine, David Geffen School of Medicine, University of California,
Los Angeles, California, U.S.A.
Paul J. Turek Departments of Urology, Obstetrics, Gynecologic and
Reproductive Sciences, Male Reproductive Laboratory, University of
California, San Francisco, California, U.S.A.
Johannes D. Veldhuis Endocrine Research Unit, Mayo Clinic College of
Medicine, Rochester, Minnesota, U.S.A.
Christina Wang General Clinical Research Center, Harbor-UCLA Medical
Center, Torrance, California and Department of Medicine, David Geffen School
of Medicine, University of California, Los Angeles, California, U.S.A.

Stephen H. Weinstein Director of Clinical Skills, University of Missouri

Medical School, Columbia, Missouri, U.S.A.

Stephen J. Winters Division of Endocrinology and Metabolism, University of

Louisville, Louisville, Kentucky, U.S.A.

Gary A. Wittert School of Medicine, Discipline of Medicine, University of

Adelaide; Hanson Research Institute; and Discipline of Medicine,
Royal Adelaide Hospital, Adelaide, Australia

Ekaterina V. Zubkova Departments of Pharmacology and Therapeutics, and

Obstetrics and Gynecology, McGill University, Montreal, Quebec, Canada
Part I
Physiology of Male Reproductive Function

R eproduction is perhaps the most evolutionarily

significant activity a man can perform in his
lifetime. It is no wonder, then, that the successful
has been made in the understanding of the physiology
of male reproductive function, resulting in a greater
appreciation of the intricate neurohormonal mechanisms
achievement of new life from the combined events of that govern sexual maturation and spermatogenesis.
spermatogenesis, sexual arousal, sexual intercourse, Similar innovations in surgical and microsurgical tech-
and the penetration and fertilization of the female niques have enabled previously infertile men to have
ovum is so tightly regulated at every step in the children, via, e.g., the operative relief of ejaculatory
process—a feat nothing short of biologically extraordi- duct obstruction (including vasectomy reversal), and
nary. Yet, because intact reproductive function requires the ability to harvest healthy sperm for in vitro fertil-
the synchronous coordination of myriad anatomical, ization or intracytoplasmic sperm injection. Therefore,
vascular, neurologic, psychologic, endocrine, immu- it is imperative that the reproductive health profes-
nological, systemic, and even environmental and sional remains abreast of the ever-evolving data in this
nutritional factors, defects may easily occur in any one exciting field. The chapters in Part I of this volume
or more of these individual components, resulting in carefully describe the most up-to-date knowledge
some type of solely male-factor reproductive dys- regarding the physiology of male reproductive health
function, as detailed in Part II of this volume. Due to at all stages of the life cycle in the hopes of providing
breakthroughs in bench research studies and the a thorough scientific and psychosocial understanding
development of state-of-the-art anatomical and func- for any clinician involved in the care of the infertile
tional imaging techniques, however, much progress male patient.
 Section I: History
1
Male Reproduction: Evolving Concepts of Procreation
and Infertility Through the Ages
Jeannette Hacker
Department of Diabetes, Endocrinology and Metabolism, City of Hope National Medical Center, Duarte, California, U.S.A.

Fouad R. Kandeel
Department of Diabetes, Endocrinology and Metabolism, City of Hope National Medical Center, Duarte, California and
David Geffen School of Medicine, University of California, Los Angeles, California, U.S.A.

† INTRODUCTION to serve only as a nesting ground within which this

new life could be nurtured; this may be due to the fact
Human perspectives on reproductive function seem to that the man’s contribution to the reproductive process
have evolved along with an increased capacity for (the emission of semen) could be tangibly perceived,
learning and the ability to reason. As brain function whereas the woman’s contribution of the egg could
and thought processes began to advance, so too did not. The only visible physical sign of the woman’s con-
our ancestors’ understanding of their roles in repro- tribution was the monthly flow of menstrual blood,
duction, albeit with some interesting misnomers along which came to be interpreted as another indicator of
the way. A detailed review of evolving concepts of her purely nutritive function in pregnancy. Metaphors
human reproduction through the ages would require such as the “seed” and “soil” were symbols commonly
the dedication of an entire text and is beyond the scope used to represent male semen and the female womb.
of the current volume. Therefore, this chapter seeks to Intercourse was sometimes poetically described as
provide a synopsis of milestones that have shaped “cultivating the soil.”
mankind’s understanding of procreation and inferti-
lity and inevitably influenced the practice of modern Greco-Roman Philosophy
reproductive medicine (Table 1). Western concepts of human reproduction were largely
shaped by theories presented by the prominent Greek
philosophers/physicians Aristotle, Hippocrates, and
† EVOLVING CONCEPTS OF HUMAN Galen from the fifth century b.c. through the first cen-
REPRODUCTION tury a.d. In addition to their many other contributions
to philosophy and medicine, these men individually
Early humans likely perceived childbirth as a sponta- strove to answer more complex questions about how
neous process. From the beginning, the female role in human life began.
reproductive events was highly regarded, probably The first of the three philosophers, Aristotle (384–
due to the obvious physical growth witnessed during 322 b.c.), promoted what has been referred to as the
gestation and the nature of the birth process itself. “one-seed theory” of conception. As in previous male-
Therefore, the worship of “Mother Earth” and other dominated preconceptions of procreation, Aristotle
personified female objects was predominant in ancient thought male seed (semen) was responsible for the
cultures. Originally, many cultures believed that the generation of new life, while women provided merely
earth was born from the female sea. Some cultures the matter (menstrual blood) that nourished the seed
thought that the sun itself fathered human children, once it was implanted in the womb through sexual
because it was observed that the warm rays of the sun intercourse. Because women only donate nutrient
could make things grow. Others surmised that the matter, Aristotle held that women were able to con-
moon, or a male personification of it, impregnated a ceive in the absence of orgasm, and thus concluded
woman (1). that their role was less significant. He likened the
Sexual interaction with a man was eventually reproductive process to that of curdling cheese, com-
recognized as being essential for the “magical” feat of paring the menstrual blood to inert milk and active
procreation, and the male role in reproduction was semen to the rennin that activates it (2).
given precedence over the female one. For more than The Greek physician Hippocrates (460–377 b.c.),
1000 years, common belief held that the male seed car- the “Father of Medicine,” was one of the first Western
ried the power of new life, and women were thought philosophers to recognize the contributions of both
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