MARSHALL'S PHYSIOLOGY OF REPRODUCTION

MARSHALL'S PHYSIOLOGY OF REPRODUCTION Fourth edition

Volume 3 Pregnancy and Lactation Part One Ovulation and early pregnancy Edited by

G.E. Lamming Emeritus Professor of Animal Physiology University of Nottingham

DI]I

I

SPRINGER-SCIENCE+BUSINESS MEDIA. B.Y.

First edition 1910 Second edition 1922 Third edition 1952 Reprinted 1958, 1961, 1964, 1967 Fourth edition 1985, 1990 and 1994

© 1910,1922,1952 Longman, Green & © 1985, 1990 and 1994 G. E. Lamming

Co.

Originally published by Chapman & Hall in 1994 Softcover reprint of the hardcover 4th edition 1994 Typeset in 10112 Palatino by Florencetype, Kewstoke, Avon ISBN 978-94-010-4561-2

ISBN 978-94-011-1286-4 (eBook)

DOI 10.1007/978-94-011-1286-4

Apart from any fair dealing for the purposes of research or private study, or criticism or review, as permitted under the UK Copyright Designs and Patents Act, 1988, this publication may not be reproduced, stored, or transmitted, in any form or by any means, without the prior permission in writing of the publishers, or in the case of reproduction only in accordance with the terms of the licences issued by the Copyright Licensing Agency in the UK, or in accordance with the terms of licences issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries concerning reproduction outside the terms stated here should be sent to the publishers at the London address printed on this page. The publisher, editor and contributors make no representation, express or implied, with regard to the accuracy of the information contained in this book and cannot accept any legal responsibility or liability or omissions that may be made. A catalogue record for this book is available from the British Library Library of Congress Catalog Card Number available 00

Printed on acid-free text paper, manufactured in accordance with ANSIINISO Z39.48-1992 (Permanence of Paper). [Paper = Fineblade, 100 gsm]

CONTENTS

Consultant and advisors List of contributors Preface F.H.A. Marshall: a biographical note Sir Alan Parkes: a biographical note Acknowledgements

ix

x

xiii xiv

xvii xix

1 The ovarian egg and ovulation A. Tsafriri, R. Reich and A. Abisogun 1.1 Introduction 1.2 The primordial cells 1.3 Oogonia 1.4 The oocyte 1.5 Ovarian differentiation and folliculogenesis 1.6 Follicular growth 1. 7 Ovulatory changes 1.8 Concluding remarks Acknowledgements References

3 4 6 8 12 13 28 56 58 58

2 Pre-implantation development

93

c.R. Austin

2.1 Introduction 2.2 The cleaving eutherian egg and the blastocyst 2.3 Development in monotremes and marsupials 2.4 Experimental manipulation Acknowledgement References 3 Implantation

c.R. Finn

3.1 Introduction 3.2 Species variation 3.3 Preparation of the uterus for implantation 3.4 Implantation of the blastocyst

1

94 96 111 114

136 136

157 159 159 162 176

vi Contents

3.5 The decidual cell reaction 3.6 Evolution of the endometrial response to implantation Acknowledgement References 4

Placentation F.B.P. Wooding and A.P.F. Flint 4.1 Introduction 4.2 General characteristics and definitions 4.3 Development of embryonic membranes in the various vertebrate taxa 4.4 Placental endocrine function 4.5 Interspecific embryo transfer and hybrid pregnancies: role of the placenta 4.6 Viviparity Acknowledgements References

5 Hormones of the placenta: hCG and hPL W.R. Butt, T. Chard and R. Iles

5.1 5.2 5.3 5.4

Introduction Human chorionic gonadotrophin Chorionic gonadotrophin and the glycoprotein hormones Human placental lactogen References

6

Normal and abnormal embryo-fetal development in mammals J.P. Hearn, A.G. Hendrickx, G.E. Webley and P.E. Peterson

6.1 6.2 6.3 6.4 6.5 6.6 6.7 6.8

Introduction Embryo-maternal interactions Regulation of the corpus luteum Rates of development Prenatal mortality Development of the reproductive tract Abnormal development Concluding comments References

7

Endocrinology of pregnancy, parturition and lactation in marsupials Marilyn B. Renfree

7.1 7.2 7.3 7.4 7.5 7.6 7.7 7.8

Introduction Patterns of reproduction Early pregnancy: ovulation, cleavage and the early blastocyst Embryonic diapause Initiation and maintenance of diapause Reactivation after diapause Maintenance of pregnancy Acquisition of endocrine organs during development

186 203 205 205 233

235 236 293 407 423 424 429 429 461

463 463 464 490 505 535

537 538 553 562 573 586 615 636 637 677

678 679 683 688 691 693 704 718

Contents

7.9 Control of parturition 7.10 Lactation: lactogenesis and galactopoiesis 7.11 Control of milk secretion and concurrent, asynchronous lactation 7.12 Adaptive significance of the marsupial mode of pregnancy and lactation Acknowledgements References 8 Fetal physiology F. Broughton-Pipkin, D. Hull and T. Stephenson

vii

729 737 743 749 751 751 767

8.1 Introduction 8.2 The placenta 8.3 Fetal physiology 8.4 Response to birth 8.5 Conclusion References

769 776 786 829 842 842

9 Initiation of parturition G.c. Liggins and G.D. Thorburn

863

9.1 Introduction 9.2 Man 9.3 Monkey 9.4 Pig 9.5 Sheep 9.6 Goat 9.7 Cattle 9.8 Horse 9.9 Rat 9.10 Guinea pig 9.11 Dog 9.12 Rabbit 9.13 Conclusions Acknowledgements References 10 Role of the fetal pituitary-adrenal axis and placenta in the initiation of parturition G.D. Thorburn and G.c. Liggins

10.1 10.2 10.3 10.4 10.5 10.6 10.7 10.8 10.9

Historical aspects Plasma cortisol concentrations in the fetal lamb Adrenocorticotrophic hormone (ACTH) Development of fetal adrenal function Fetal maturation - in vitro studies Biochemical changes in the fetal adrenal associated with maturation ACTH receptors in the fetal adrenal Inhibition of fetal adrenal function Steroidogenic pathways

866 867 884 891 906 930 938 946 951 958 960 962 966 967 967 1003

1004 1005 1006 1010 1010 1012 1012 1014 1015

viii

Contents

10.10 Prostaglandin E2 10.11 The development of glucocorticoid feedback on the fetal HPA axis 10.12 Possible heterogeneity of fetal pituitary corticotrophin 10.13 What controls the release of ACTH l - 39 ? 10.14 Fetal hypothalamus: ultrastructural development 10.15 The 'trigger' for parturition 10.16 Summary Acknowledgements References

11 Regulation of post-partum fertility in lactating mammals A.S. McNeilly, I.A. Forsyth and J.R. McNeilly 11.1 Introduction 11.2 Endocrine changes associated with milk production 11.3 Oxytocin and milk ejection reflex 11.4 Ovarian activity during lactation 11.5 Gonadotrophins 11.6 Induction of ovarian activity during lactation 11.7 Mechanism of suckling/milking-induced suppression of GnRH 11.8 Conclusions References 12

Physiology and biochemistry of lactation T.B. Mepham and N.J. Kuhn 12.1 Introduction 12.2 The evolution of lactation 12.3 Morphology of the mammary gland 12.4 Milk composition 12.5 Provisioning the mammary glands 12.6 Metabolic activity of mammary cells 12.7 Secretory processes 12.8 Control of milk secretion 12.9 Trends and significance of research on lactation References Subject index Author index

1016 1019 1022 1023 1025 1026 1026 1028 1028

1037 1038 1038 1048 1051 1057 1065 1066 1070 1071

1103 1105 1105 1111 1118 1128 1134 1157 1162 1166 1168 1187 1209

CONSULTANT TO THE FOURTH EDITION

Professor Sir Alan Parkes CBE, MA, PhD, DSe, SeD, FRS (deceased) 1 The Bramleys, Shepreth, Royston, Hertfordshire, SG8 6PY, UK ADVISORS TO THE FOURTH EDITION

Professor D.T. Baird MBChB, DSe, FRCP(Ed), FRCOG

Department of Obstetrics and Gynaecology, Centre for Reproductive Biology, University of Edinburgh, Edinburgh, EH39EW, UK Professor W.R. Butt PhD, DSe, FRIC 36 Avonside, Mill Lane, Stratford upon Avon CV37 6BJ, UK Professor 1.0. Cooke MB, BS, FRCOG Jessop Hospital for Women, Sheffield, S3 7RE, UK Professor R.O. Greep BS, MS, PhD Harvard Medical School, Boston, Massachusetts, USA

Professor H.D. Hafs BS, MS, PhD Merck, Sharpe & Dohme, Rahway, New Jersey 07065/0912, USA Professor H. Karg Dr met vet Technische Universitat Munich, Munich, Germany Professor C.G. Liggins MB, ChB, PhD, FRCS(Ed), FRACS, FRCOG, FRSNZ

Postgraduate School of Obstetrics and Gynaecology, University of Auckland, Auckland 3, New Zealand Professor R. V. Short BVSe, MSe, PhD, SeD, FRCVS, FRS, FRSE

Monash University, Victoria 3168, Australia Professor G.M.H. Waites

BSe, MA, PhD, SeD,

FIBiol

World Health Organisation, 211 Geneva 27, Switzerland Dr Barbara J. Weir MA, PhD 56 Church Road, Hauxton, Cambridge, CB25HS, UK

CONTRIBUTORS

A.O. Abisogun MD, PhD The Bernhard Zondek Laboratory of Hormone Research Department of Hormone Research The Weizmann Institute of Science 76 100 Rehovot Israel

T. Chard MD, FRCOG Professor of Obstetrics, Gynaecology and Reproductive Physiology Department of Reproductive Physiology St Bartholomew's Hospital Medical College West Smithfield London EClA 7BE

C.R. Austin BVSe, DSe, MA, FAIBioJ, FAA Emeritus Professor 47 Dixon Road Buderim Queensland 4556 Australia

C.A. Finn BSe, PhD, MRCVS Professor of Veterinary Physiology Department of Veterinary Pre-Clinical Sciences University of Liverpool Leahurst Chester High Road Neston, South Wirral L64 7fE

UK

UK

F. Broughton-Pipkin MA DPhil Professor of Perinatal Physiology Department of Obstetrics & Gynaecology University of Nottingham Queen's Medical Centre Nottingham NG7 2UH

UK

A.P.F. Flint BSe, PhD, FIBiol Professor of Animal Physiology Department of Physiology and Environmental Science University of Nottingham Sutton Bonington Campus Loughborough Leicestershire LEl2 5RD

UK

W.R. Butt PhD, DSe, CChem, FRlC, FRSC Emeritus Professor 36 Avonside Mill Lane Stratford upon Avon CV37 6BJ

UK

I.A. Forsyth BA, DPhil, FIBiol Research Scientist AFRC Babraham Institute Babraham Cambridge CB2 4AT

UK

Contributors xi J.P. Hearn BSc, MSc, PhD Professor in Physiology, School of Medicine Director of Wisconsin Regional Primate Research Center University of Wisconsin Madison, Wisconsin 53715-1299 USA A. G. Hendrickx PhD Director of California Regional Primate Research Center University of California Davis, California 95616--8542 USA Sir D. Hull BSc, MB, ChB, FRCP, DCH, DO bst RCOG Professor and Head of Child Health Department of Child Health University of Nottingham Queen's Medical Centre Nottingham NG7 2UH

UK

R. lIes BSc, MSc, PhD, CBio!, MIBioI Honorary Lecturer and Senior Scientist Williamson Laboratory for Molecular Oncology Joint Academic Unit of Obstetrics, Gynaecology and Reproductive Physiology St Bartholomew's Hospital Medical College West Smithfield London EC1A 7BE

UK

N.J. Kuhn BA, DPhii Reader in Biochemistry School of Biochemistry University of Birmingham Birmingham B15 2TT

UK

Sir G.C. Liggins MB, ChB, PhD, FRCS (ED), FRACS, FRCOG, FRSNZ, CBE

Professor, Department of Obstetrics & Gynaecology University of Auckland National Women's Hospital Claude Road Auckland 3 New Zealand

A.S. McNeilly BSc, PhD, DSc Deputy Director and Honorary Professor MRC Reproductive Biology Unit University of Edinburgh Centre for Reproductive Biology 37 Chalmers Street Edinburgh EH3 9EW

UK

J.R. McNeilly BSc, PhD Research Scientist Centre for Genome Research University of Edinburgh Kings Buildings Edinburgh

UK

T.B. Mepham BSc, PhD Reader in Lactational Physiology Department of Physiology & Environmental Science University of Nottingham Sutton Bonington Campus Loughborough Leicestershire LE12 5RD

UK

P.E. Peterson Research Scientist California Regional Primate Research Center University of California Davis, California 95616--8542 USA R. Reich BSc, MSc, PhD Department of Pharmacology School of Pharmacy Faculty of Medicine The Hebrew University of Jerusalem Israel M.B. Renfree BSc (Hons), PhD, DSc Professor and Head of Department Department of Zoology University of Melbourne Parkville Victoria 3052 Australia

xii

Contributors

T. Stephenson BSc, BM, BCh, MRCP, DM Senior Lecturer in Child Health Department of Child Health University of Nottingham Queen's Medical Centre Nottingham NG7 2UH

UK

G.D. Thorburn BSc (Med), MB, BS, MD, FRACP, FAA

Professor and Head of Physiology Department of Physiology Monash University Clayton, Melbourne Victoria, 3168 Australia A. Tsafriri BSc, MSc, PhD Professor, The Bernhard Zondek Laboratory of Hormone Research The Weizmann Institute of Science 76100 Rehovot Israel

G. Webley BSc, PhD Research Fellow Institute of Zoology Regent's Park London NW1 4RY

UK

F.B.P. Wooding BA, MA, PhD Research Scientist AFRC Babraham Institute Babraham Hall Cambridge CB2 4AT

UK

PREFACE

In 1910 F.H.A. Marshall established a landmark in the history of science when he published the first edition of The Physiology of Reproduction. A revised second edition was published by Marshall in 1927 and in this he was assisted by a young graduate, A.S. Parkes (later Sir Alan), who accepted an invitation to organize and edit the third edition, the preparation of which was planned in 1939 and then delayed by the advent of the Second World War. It was subsequently completed in three volumes, the first of which appeared in 1956 and the final volume in 1966. For nearly a century succeeding editions of Marshall's Physiology of Reproduction have provided an advanced reference text for reproductive biologists. We hope the current volume will maintain this role. Organizing and editing the work of the Fourth Edition has involved an extension of the scope of each succeeding text while retaining a comprehensive philosophical approach to the review of literature. The coverage demanded for such an advanced text has expanded exponentially during the last decade. For the Fourth Edition, Volume 1 The Reproductive Cycles of Vertebrates published in 1984, provided a synopsis of the various reproductive patterns. It contains major contributions concerning fishes, reptiles, amphibia, birds, marsupials and eutherian mammals. The volume included special chapters on Non-human Primates and Humans. Volume 2 on Reproduction in the Male, published in 1990, provided a detailed analysis of research on male reproductive physiology,

male life cycles and seasonality, the physiology of the testis and accessory sexual organs, artificial insemination and the preservation of semen. The current volume aims to maintain Marshall's traditional comparative approach with particular emphasis on reviewing research data derived from studies of farm and laboratory animal species, non-human primates and humans. A major effort has been made by contributors in the final revision of their texts to present the results of the most recent research. However, given the time required to receive and edit the manuscripts, check the proofs and index the volume, it is as fully up-to-date as possible. The increasing impact of new information on the reproductive physiology of humans is well reviewed and this aspect amplifies the valuable interrelationship which exists between those working experimentally with farm and laboratory species and those engaged in the control of reproduction and assisted conception in humans. The comparative aspects of placentation, embryo and fetal development, the initiation of parturition and lactation are extensively reviewed. It is our established editorial policy to seek contributions from the most eminent international authorities. This I trust we have achieved in full measure. I acknowledge and welcome the contributions made by the authors from Australia, New Zealand, Israel, the United States of America, as well as those from the UK. G.E. Lamming University of Nottingham

A BIOGRAPHICAL NOTE PREPARED BY SIR ALAN PARKES (for the Third Edition)

F. H. A. MARSHALL

CBE, MA, PhD, DSe, SeD, FRS. 11th July 1878-Sth February 1949

Marshall's name is associated, mainly or even exclusively, with the study of the physiology of reproduction in its many branches. He was preceded in this field by another pioneer, Walter Heape, who was completing his classic studies of the primate uterus and collecting material for his analysis of the oestrous cycle at the time when Marshall was an undergraduate in Cambridge. The two men did not meet until later, but they corresponded freely, and Heape's morphological studies and extensive knowledge of living animals were of great value to Marshall in his early days. Heape, however, was not a physiologist - he seems, for instance, to have regarded the ovary merely as a producer of eggs - and it is to Marshall that we owe the foundations of our present knowledge of the

role of internal secretions in the reproductive processes. Marshall's debut was characteristic of an age in which research was a calling rather than a profession; a precarious calling to which young men were drawn by interest, not by ambition. In 1900, the late Lord Carmichael gave £200 to Prof. Cossar Ewart, Professor of Natural History in the University of Edinburgh, to enable a Research Student to assist with his long-term experiments on telegony, and to undertake work on reproduction in farm animals. Cossar Ewart's inquiries resulted in Marshall accepting an invitation to go to Edinburgh. The work on telegony, which involved a detailed study of the hair of the Equidae, was uninteresting and unproductive, and

A biographical note xv Marshall turned his attention to research on the sexual cycle in the sheep, for which he was to make use of Ewart's farm at Penicuik. This work was the subject of his first important scientific publication, which appeared in the Transactions of the Royal Society in 1903. Similar studies followed with Schafer on the oestrous cycle in the ferret (1904), and with Jolly on the oestrous cycle in the dog (1905). A communication on the ovary as an organ of internal secretion, written in collaboration with Jolly and published in the Transactions of the Royal Society in 1905, constituted the first serious attempt to correlate the changes in the uterus during the reproductive cycle with the cyclic production of different internal secretions by the ovary. Marshall returned to Cambridge in 1908, and continued his work on the physiology of reproduction. During the war, 1914-18, he studied, on behalf of the Ministries of Food and Agriculture, such problems as the most economic age and condition at which to slaughter cattle. Subsequently, Marshall's researches have dealt mainly with the influence of exteroceptive factors - light, climate, etc. - on the reproductive processes, and in 1936 he made this the subject of his Croonian Lecture. He was also intensely interested in the courtship and breeding behaviour of birds, on which he was a considerable authority. Marshall spent a great deal of time preparing the first edition of The Physiology of Reproduction in which he gathered together all relevant knowledge concerning the breeding season, cyclic changes in the reproductive organs, fertilization, fertility, lactation and so on. The book was published by Longmans, Green & Co. in 1910, and immediately achieved success. A second edition was called for after the 1914-18 war. This second edition, published in 1922, was necessarily much larger than the first, for since 1910 knowledge of the reproductive processes in mammals and lower vertebrates had increased considerably. A third edition, largely

rewritten by Marshall's pupils and colleagues, and again much enlarged, is now in the course of publication. Marshall's main contribution to biology has been twofold. In the first place, the stimulating synthesis of existing knowledge embodied in the first edition of The Physiology of Reproduction gave impetus and balance to a branch of biology sadly lacking in both. The fact that between the wars the physiology of reproduction became a subject of intensive study by scientists in all countries was due in no small measure to the stimulus given by Marshall's writings. In the second place, Marshall's remarkable prescience in deducing correctly the existence and respective roles of follicular and luteal hormones long before either was isolated, or even obtained in active extracts, had a profound influence on the development of the subject. During the early years of the century Ludwig Fraenkel in Breslau and P. Bouin in Strasbourg, working on the rabbit, concentrated attention on the corpus luteum; in the 1920s intensive study of the follicular hormone by Edgar Allen and his colleagues in the USA swung attention to the follicular apparatus. It fell to the English school of workers, under Marshall's influence, to hold the balance until the final demonstration and isolation of the corpus luteum hormone in C.W. Corner's Rochester laboratory completed the working out of his early predictions. Scientists are of many kinds, but inspiration flows most fruitfully from those who are able, by some gift withheld from lesser men, to divine the richness of uncharted country and sense the vital landmarks. Thus do they avoid the barren places and the morasses of unimportant detail which engulf so many. To these, discovery is an art rather than a science, a matter of instinct rather than of intellectual machinery. Such was Marshall. 20 October, 1948 Revised 10 February, 1949.

A.S.P.

xvi A biographical note

F.H.A. MARSHALL CURRICULUM VITAE

Francis Hugh Adam Marshall was the younger son of the late Thomas Marshall, J.P., and was born at High Wycombe on 11 July, 1878. He was educated at St. Mark's School, Windsor, and privately. After a short time at University College, London, he went to Christ's College, Cambridge, in 1896, and took the Natural Science Tripos in 1899. During the latter part of the time in Edinburgh Marshall became Carnegie Fellow and Lecturer on the physiology of reproduction. After his return to Cambridge he was Lecturer and then Reader in Physiology, in the School of Agriculture, for more than 35 years, until 1943. For a short period, Marshall followed the late Prof. T.B. Wood in directing the Institute of Animal Nutrition. He became in succession, Fellow, Tutor, Dean, and finally, in 1940, Vice-Master of Christ's

College. He was a Proctor in 1911-12 and afterwards served on the Council of the Senate. Marshall was an original member of the Council of Management of the Journal of Endocrinology and was one of the four honorary members of the Society for Endocrinology. For many years he was an editor of the Journal of Agricultural Science, and finally became principal editor. Marshall's scientific distinctions included D.Sc. (Edin.), Sc.D. (Cantab.), D.5c. (Hon. Manch.), LL.D. (Hon. Edin.), F.R.S. (1920), C.B.E. (1933), Council of the Royal Society (1933-5), Baly Medal of the Royal College of Physicians (1935), Croonian Lecturer (Royal Society) 1936, and Royal Medal (Royal Society) 1940. Reprinted from THE ENDOCRINOLOGY

JOURNAL

OF

A BIOGRAPHICAL NOTE PREPARED BY G. E. LAMMING (for the Fourth Edition)

PROFESSOR SIR ALAN PARKES CBE, FRS.

Sir Alan Parkes' association with F.H.A. Marshall began when he enrolled as an undergraduate at Christ's College, Cambridge and Marshall became his tutor. Parkes describes this first association with Marshall as no more than formal since he preferred to avoid undue contact with those in authority, no doubt prompted by previous experience at school and in the army. Parkes read agriculture, obtaining a pass degree in 1921 and an MA in 1924. After graduating in 1921 he applied for a position at the Shirley Cotton Research Institute in Manchester which (fortunately for reproductive physiology) he failed to obtain. However, fortified by his father's encouragement and support he entered Manchester University as a PhD student in the Zoology Department to

research on the mammalian sex ratio. During this period Parkes accepted a request from Marshall to help him with indexing the second edition of MPR which was published in 1922. No doubt Marshall had been impressed, not so much by Parkes' enthusiasm as an undergraduate, but by the initiative he had shown in carrying out a literature search for his first published paper on 'Sex heredity', which appeared in Science Progress in 1921. At Manchester, Parkes completed his PhD in two years having A.V. Hill as an internal examiner and Marshall as the external. The inclusion of A.V. Hill was fortunate for it resulted in Parkes moving with Hill, now a Professor, to University College, London, where Parkes remained until 1931, holding a

xviii

A biographical note

series of three research fellowships. Here he co-operated in researches with a number of famous contemporaries including Jack (later Sir Jack) Drummond, Professors F. W.R. Brambell, Guy Marrion and Solly (later Lord) Zuckerman. From 1932 to 1961 Parkes was a member of the MRC's National Institute of Medical Research, first at Hampstead and later at Mill Hill. He was responsible, with others, for organizing a number of research programmes in reproductive endocrinology, many of which had major influences on developments in reproductive physiology as applied to agriculture. Detailed research on important topics included the induction and control of ovulation and lactation, and the cryo-preservation of animal tissues. In 1961 Parkes returned to Cambridge, fittingly as the first holder of the newly created Mary Marshall Chair in the Physiology of Reproduction, established under the terms of Marshall's will. Here the nature of his work changed, for the Marshall position was vastly different from his previous MRC post; it lacked the fabric for Parkes to continue his detailed researches. Nevertheless, the change produced benefits which history may well judge of equal merit, for Parkes continued to influence the course of reproductive physiology by virtue of major roles in organizing societies and scientific journals including the Society for the Study of Fertility, its associated Journal of Reproduction and Fertility, and also the Journal of Biosocial Sciences, acting for a number of years in addition as executive editor for both journals. The following is abstracted (by kind permission of the publishers, A & C Black (Publishers) Limited) from his entry in the 1990 edition of Who's Who.

PARKES, Sir Alan (Sterling), Kt 1968; CBE 1956; FRS 1933; MA, PhD, DSc, ScD; Fellow of Christ's College, Cambridge, 1961--69, Hon. Fellow 1970; Fellow of University College, London; Chairman, Galton Foundation, since 1969; b 1900; Y s of E. T. Parkes, Purley; m 1933, Ruth d of Edward Deanesly, FRCS, Cheltenham; one s two d. Educ: Willaston School; Christ's College, Cambridge; BA Cantab, 1921, ScD 1931; PhD Manchester, 1923; Sharpey Scholar, University College, London, 1923--24; Beit Memorial Research Fellow, 192430; MA Cantab 1925; Schafer Prize in Physiology, 1926; DSc London 1927; Julius Mickle Fellowship, University of London, 1929; Hon. Lecturer, University College, London, 1929-31; Member of the Staff of the National Institute for Medical Research, London, 1932--61; Mary Marshall Prof. of the Physiology of Reproduction, Univ. of Cambridge, 1961--67, Professor Emeritus 1968. Consultant, Cayman Turtle Farm Ltd, Grand Cayman, BW1, 1973--. Foulerton Student of the Royal Society, 1930--34. Mem., BioI. and Med. Cttee, Royal Commn on Population, 1944-46. President: Section of Endocrinology, Roy. Soc. Med., 1949-50, Section of Comparative Medicine, 1962--63; Section 0 Brit. Assoc. for the Advancement of Science, 1958; Eugenics Soc., 1968-70; Inst. of Biology, 1959--61; Assoc. of Scientific Workers, 1960--62. Chairman: Soc. for Endocrinology, 1944-51; Soc. for Study of Fertility, 1950--52, 1963--66; Nuffield Unit of Tropical Animal Ecology, 1966--69; Breeding Policy Cttee, Zool Soc. of London, 1960--67; Scientific Adv. Cttee, Brit. Egg Mkting Bd, 1961-70. Mem. Adv. Cttee on Med. Research of the WHO, 1968-71. Executive Editor, JI Biosocial Science, 1969-; Sec., Jls of Reproduction & Fertility Ltd, 1970--76. Consultant, IPPF, 1969-79. Cameron Prize, 1962; Sir Henry Dale Medal, Soc. for Endocrinology, 1965; John Scott Award (jtly with Dr A. U. Smith and Dr C. Polge), City of Philadelphia, 1969; Marshall Medal, Soc. Stud. Fert., 1970; Oliver Bird medal, FPA, 1970. Publications: The Internal Secretions of the Ovary, 1929; Sex, Science and Society, 1966; Patterns of Sexuality and Reproduction, 1976; Off-beat BiolOgist, 1985; Biologist at Large, 1988; papers on the Physiology of Reproduction, on Endocrinology and on the behaviour of living cells at low temperatures in JI of Physiology, Proc. Royal Society and other scientific jls. Ed. Marshall's Physiology of Reproduction, 3rd edn, 1952, consultant 4th edn, 1994. Address: 1 The Bramleys, Shepreth, Royston, Hertfordshire SG8 6PY.

Professor Parkes died on July 17th 1990.

ACKNOWLEDGEMENTS

Fortunately, few will have experienced the volume of work involved in commissioning, editing and preparing a text of this size. It can only be completed by competent teamwork. My main task has been to persuade the team members to participate and then organize their work. My special thanks goes initially to the contributors but also to their secretaries. It is well known that the best authors are busy individuals heavily committed both to their research and to allied commitments, (including travel abroad). Thus, many of my queries concerning the texts are handled initially (and, in some cases, entirely) by secretaries. I applaud their tenacity and tact, and also their skill, especially in checking reference citations. I owe them a special vote of thanks. Mrs Pamela Blythe completed the initial secretarial work prior to her move to a senior managerial post elsewhere. The main bulk of the editorial work was completed by Mrs Linda Faillace during her husband's (Dr L. Faillace) postdoctoral research work in this department. I acknowledge her skill in the preparation of texts for printing and thank her husband and family for their forebearance in tolerating the many hours of work she completed, both in the office and at home. I also thank Mrs Monica Bagshaw for providing additional secretarial help, and Dr Karen Shelton for her contribution to chapter 2. I again thank collectively the advisers to the Fourth Edition. Professor Parkes assisted,

prior to his death, in the initial plans. Several of the advisers were helpful not only in planning the volume, but also in the detailed discussions involving our move to publication by Chapman & Hall. I acknowledge the commitment, skill and speed of the Chapman & Hall staff (Miss Rachel Young, Mr Martin Tribe and Mr Jeremy Macdonald) in publishing this volume. Finally, I reserve special praise for those who have completed the indexing of volume 3 with a total of about 10 000 reference citations. What a pleasure it is to have the continued help of Dr Barbara Weir as an adviser, and to take advantage of her skill in the preparation of the subject index. Her knowledge of world literature in reproductive physiology, gained from years of experience as Editor of the Journal of Reproduction and Fertility, is invaluable. I am sure many who use these texts, and who know Barbara personally, will support my expression of gratitude. The author index was ably and generously completed by my young staff colleagues. (They may comment that they had little choice since I organize their work schedule). I am most fortunate to have young staff colleagues who cheerfully tackled these time-consuming chores, and I record my sincere appreciation to Misses Morag Batten, Sarah Machin, Jane Plowright and Alison Clarkson. G.E. Lamming University of Nottingham