Prelims-P088465 2/28/06 3:58 PM Page i Human Reproductive Biology THIRD EDITION Prelims-P088465 2/28/06 3:58 PM Page ii Prelims-P088465 2/28/06 3:58 PM Page iii Human Reproductive Biology THIRD EDITION Richard E. Jones Professor of Biology Emeritus University of Colorado Boulder, Colorado Kristin H. Lopez Department of Integrative Physiology University of Colorado Boulder, Colorado AMSTERDAM • BOSTON • HEIDELBERG • LONDON NEW YORK • OXFORD • PARIS • SAN DIEGO SAN FRANCISCO • SINGAPORE • SYDNEY • TOKYO Academic Press is an imprint of Elsevier Prelims-P088465 2/28/06 3:58 PM Page iv Senior Acquisitions Editor: Nancy Maragioglio Project Manager: Philip Korn Associate Editor: Kelly Sonnack Marketing Manager: Trevor Daul Cover Design: Cate Barr Composition: Integra Software Pvt. Ltd. Cover Printer: Transcontinental Interglobe Interior Printer: Transcontinental Interglobe Academic Press is an imprint of Elsevier 30 Corporate Drive, Suite 400, Burlington, MA 01803, USA 525 B Street, Suite 1900, San Diego, California 92101-4495, USA 84 Theobald’s Road, London WC1X 8RR, UK This book is printed on acid-free paper. Copyright © 2006, Elsevier Inc. All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone: (ϩ44) 1865 843830, fax: (ϩ44) 1865 853333, E-mail: [email protected]. You may also complete your request on-line via the Elsevier homepage (http://elsevier.com), by selecting “Customer Support” and then “Obtaining Permissions.” Library of Congress Cataloging-in-Publication Data Jones, Richard E. (Richard Evan), 1940- Human reproductive biology / by Richard E. Jones. — 3rd ed. p. ; cm. Includes bibliographical references and index. ISBN-13: 978-0-12-088465-0 (hardcover : alk. paper) ISBN-10: 0-12-088465-8 (hardcover : alk. paper) 1. Human reproduction. 2. Sex. I. Title. [DNLM: 1. Reproduction—physiology. 2. Sexual Behavior. WQ 205 J78ha 2006] QP251.J636 2006 612.6—dc22 2005024432 British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library ISBN 13: 978-0-12-088465-0 ISBN 10: 0-12-088465-8 For all information on all Elsevier Academic Press publications visit our Web site at www.books.elsevier.com Printed in the United States of America 0607080910987654321 Working together to grow libraries in developing countries www.elsevier.com | www.bookaid.org | www.sabre.org Prelims-P088465 2/28/06 3:58 PM Page v To my wife, Betty, and my four sons (Evan, Ryan, Peter, and Christopher) R.E.J. To Tom and Jessica K.H.L. Prelims-P088465 2/28/06 3:58 PM Page vi Prelims-P088465 2/28/06 3:58 PM Page vii Contents Preface xvii Box 2: Kallmann’s Syndrome and the Embryological Origin and Migration of GnRH Cells 22 PART ONE Pineal Gland 23 Adult Female and Male Feedback Control of Gonadotropin Secretion 24 Reproductive Systems 1 Feedback Systems 24 Regulation of Gonadotropin Secretion by Negative Feedback 25 CHAPTER ONE Positive Feedback 26 Endocrinology, Brain Control of Prolactin Secretion 27 and Pituitary Gland 3 Chapter Summary 28 Further Reading 29 Introduction 3 Advanced Reading 29 Endocrine System 3 Science of Endocrinology 4 CHAPTER TWO Hormones 6 The Female Reproductive Receptors 6 System 31 Synthetic Hormones 9 The Pituitary Gland 10 Introduction 31 Hypothalamo–Neurohypophysial Ovaries 31 Connection 10 Ovarian Gross Anatomy 31 Adenohypophysis 14 Ovarian Microanatomy 31 Hypothalamo–Adenohypophysial Connection 15 Stages of Follicular Growth 32 Releasing and Release-Inhibiting Follicular Atresia 36 Hormones 18 Ovarian Steroid Hormone Synthesis 36 Gonadotropin-Releasing Hormone 19 Hormonal Control of Follicular Growth Box 1: GnRH Analogs 20 and Steroidogenesis 39 The GnRH Pulse Generator and Oocyte Maturation and Ovulation 40 Surge Center 21 Corpus Luteum 43 vii Prelims-P088465 2/28/06 3:58 PM Page viii viii Contents Ovarian Disorders 44 CHAPTER THREE Ovarian Cysts 44 The Menstrual Cycle 73 Ovarian Cancer 45 Box 1: The Estrogen Epidemic 46 Introduction 73 Oviducts 47 Reproductive Cycles in Mammals 73 Uterus 48 Major Events in the Menstrual Cycle 75 Uterine Functional Anatomy 48 The Menstrual Cycle in Detail 76 Uterine Disorders 49 Menstrual Phase 76 Pelvic Infection 49 Follicular Phase 77 Cervical Cancer 49 Box 1: Why Women Menstruate 79 Cervical Cysts and Polyps 50 Luteal Phase 83 Endometrial Cancer 50 Variations in Length of Menstrual Endometriosis 51 Cycle Phases 85 Endometrial Polyps, Hyperplasia, Methods for Detecting Ovulation 85 and Fibroids 51 Home Methods 86 “Tipped” Uterus 52 Laboratory Methods 86 Prolapsed Uterus 52 Premenstrual Syndrome 87 Vagina 53 Symptoms 87 Structure 53 Possible Causes and Treatments 87 The Vaginal Environment 53 Cyclic Changes in Mood and Skill Level 88 Female External Genitalia 54 Menstrual Difficulties 88 Mons Pubis 54 Dysmenorrhea 88 Labia Majora 54 Absence of Menstruation 89 Labia Minora 55 Box 2: Body Fat and Secondary Vestibule 55 Amenorrhea 89 Urethral Orifice 56 Menstrual Taboos 91 Clitoris 56 Chapter Summary 91 Mammary Glands 56 Further Reading 92 Mammary Gland Functional Anatomy 56 Advanced Reading 93 Hormonal Control of Mammary Gland Function 58 CHAPTER FOUR Suckling 58 The Male Reproductive System 97 Noncancerous Breast Disorders 59 Breast Cancer 59 Introduction 97 Risk 59 Testes 97 Inherited Breast Cancer 60 Seminiferous Tubules 99 Noninherited Breast Cancer 61 Spermatogenesis and Spermiogenesis 99 Box 2: Xenoestrogens and Breast Cancer 62 Sertoli Cells 101 Detection 65 Testicular Interstitial Tissue 102 Treatment of Breast Cancer 66 Box 1: Estrogens Are Male Sex Hormones 104 Chapter Summary 67 Hormonal Control of Testicular Function 105 Further Reading 68 Control of Gonadotropin Secretion Advanced Reading 70 in the Male 107 Prelims-P088465 2/28/06 3:58 PM Page ix Contents ix Other Factors That Influence Testicular Box 1: Why the Y Chromosome? 130 Function 108 Sex-Linked Inheritance 132 Other Hormones 108 Development of the Reproductive System 133 Nutrition and Exercise 108 The Sexually Indifferent Stage 133 Chemicals, Radiation, and Infections 109 Gonadal Sex Differentiation 133 Anabolic Steroids 109 Ovarian Development 134 Testicular Cancer 109 Testicular Development 135 Testosterone and Behavior 110 Testis-Determining Factor 137 Libido 110 Differentiation of Sex Accessory Ducts Aggression 110 and Glands 137 Sperm Count and Endocrine Disruptors 111 Differentiation of External Genitalia 138 Male Sex Accessory Ducts and Glands 112 Summary of Sexual Determination and Sex Accessory Ducts 112 Development 138 Epididymis 112 Disorders of Sexual Determination and Development 140 Vas Deferens 113 True Hermaphroditism 140 Urethra 113 Pseudohermaphroditism 140 Sex Accessory Glands 113 Box 2: Guevedoces 141 Seminal Vesicles 113 Chromosomal Errors and Sex Prostate Gland 114 Determination 144 Bulbourethral Glands 114 Other Problems in Sex Development 145 Prostate Disorders 114 Reproductive System in the Newborn 145 Hormonal Control of Sex Accessory Chapter Summary 146 Structures 117 Further Reading 147 Penis 117 Advanced Reading 147 Scrotum 118 Box 2: Why a Scrotum? 119 CHAPTER SIX Chapter Summary 121 Puberty 149 Further Reading 121 Advanced Reading 122 Introduction 149 Puberty and Its Timing 149 The Pubertal Process 149 PART TWO Pubertal Changes in Females 149 Sexual Differentiation Pubertal Changes in Males 151 and Development 125 Classification of Pubertal Changes 153 Gonadal Changes from Birth to Puberty 156 Ovarian Changes 156 CHAPTER FIVE Testicular Changes 157 Sexual Differentiation 127 Cryptorchid Testes 157 Hormone Levels from Birth to Puberty 158 Introduction 127 Hormones in Females 160 Chromosomal Sex 127 Menarche and First Ovulation 160 X Chromosome 127 Estrogens 160 Y Chromosome 129 Androgens 160 Prelims-P088465 2/28/06 3:58 PM Page x x Contents Hormone Levels in Males 160 PART THREE Gonadotropins and Androgens 160 Procreation 191 Estrogens 161 Androgens and Acne 161 What Mechanisms Cause Puberty? 161 CHAPTER EIGHT Environmental Factors and Puberty 163 The Human Sexual Response 193 Nutrition 164 Day Length and Season 165 Introduction 193 Stressors 165 Sex Roles 193 Box 1: Early Childhood Stress and Sexual Maturation 166 Factors Influencing Sex Roles 193 Environmental Pollutants 167 Gender Identity 193 Climate and Altitude 168 Nature or Nurture 194 Inheritance and Age of Puberty 168 Sexual Arousal 194 Box 2: Puberty Genes 168 Cultural Influence and Individual Variation 194 Puberty and Psychosocial Adjustment 170 Erotic Stimuli 195 Chapter Summary 171 Erogenous Zones 195 Further Reading 171 Proceptive Behavior 195 Advanced Reading 172 The Sexual Response Cycle 196 196 CHAPTER SEVEN The Female Sexual Response Cycle Excitement Phase 197 Reproductive Aging 175 Plateau Phase 198 Orgasmic Phase 198 Introduction 175 Resolution Phase 200 Menopause 175 Individual Variation 201 Timing of Menopause 176 The Male Sexual Response Cycle 201 Female Reproductive Age 176 Excitement Phase 201 Box 1: Egg Aging 177 Plateau Phase 203 Perimenopause 179 Orgasmic Phase 204 Premature Menopause 179 Resolution Phase 205 Symptoms of Menopause 179 Why Did Orgasm Evolve? 205 Endocrine Changes during Menopause 180 Coitus (Sexual Intercourse) 205 Chance of Pregnancy 182 Hormones and Sexual Behavior 207 Box 2: The Evolution of Menopause in Humans 183 Hormones