Ovarian Follicles in Reptiles and Birds

296 Pages·1989·9.401 MB·English

by  Professor Dr. Sardul S. Guraya (auth.)

Zoophysiology Volume 24
Coordinating Editor: D. S. Farner
Editors:
w.
Burggren S. Ishii H. Langer
O. Neuweiler D. J. Randall
Zoophysiology
Volumes already published in the series:
Volume 1: P.J. Bentley Volume 13: S. Nilsson
Endocrines and Osmoregulation (1971) Autonomic Nerve Function in the
Vertebrates (1983)
Volume 2: L. Irving
Volume 14: A.D. Hasler
Arctic Life of Birds and Mammals In
Olfactory Imprinting and Homing
cluding Man (1972)
in Salmon (1983)
Volume 3: A. E. Needham
Volume 15: T. Mann
The Significance of Zoochromes (1974)
Spermatophores (1984)
Volume 4/5: A. C. Neville Volume 16: P. Bouverot
Biology of the Arthropod Cuticle Adaptation to Altitude-Hypoxia
(1975) in Vertebrates (1985)
Volume 6: K. Schmidt-Koenig Volume 17: R.J.F. Smith
The Control of Fish Migration (1985)
Migration and Homing in Animals
(1975) Volume 18: E. Gwinner
Circannual Rhythms (1986)
Volume 7: E. Curio
The Ethology of Predation (1976) Volume 19: J. C. RUegg
Calcium in Muscle Activation (1986)
Volume 8: W. Leuthold
Volume 20: J.-R. 1I-uchot
African Ungulates (1977)
Comparative Aspects of Extracellular
Volume 9: E. B. Edney Acid-Base Balance (1987)
Water Balance in Land Arthropods Volume 21: A. Epple and J.E. Brinn
(1977) The Comparative Physiology of the
Pancreatic Islets (1987)
Volume 10: H.-u. Thiele
Volume 22: W. H. Dantzler
Carabid Beetles in Their Environments
Comparative Physiology of the
(1977)
Vertebrate Kidney (1988)
Volume 11: M.H.A. Keenleyside
Volume 23: G.L. Kooyman
Diversity and Adaptation in
Diverse Divers (1989)
Fish Behaviour (1979)
Volume 24: S. S. Guraya
Volume 12: E. Skadhauge Ovarian Follicles in Reptiles and Birds
Osmoregulation in Birds (1981) (1989)
Sardul S. Guraya
Ovarian Follicles
in Reptiles and Birds
With 72 Figures
Springer-Verlag
Berlin Heidelberg New York
London Paris Tokyo
Professor Dr. SARDUL S. GURAYA
ICMR Regional Advance Research Centre
in Reproductive Biology
Department of Zoology
College of Basic Sciences and Humanities
Punjab Agd€ultural University
Ludhiana, India
ISBN- 13: 978-3-642-83630-5 e-ISBN- 13: 978-3-642-83628-2
DOl: 10.1007/978-3-642-83628-2
Library of Congress Cataloging-in-Publication Data. Guraya, Sardul S., 1930- Ovarian
follicles in reptiles and birds/Sardul S. Guraya. p. em. - (Zoophysiology; v. 24)
Bibliography: p. Includes index. 1. Ovaries. 2. GraafIan follicle. 3. Birds - Physiology.
4. Reptiles - Physiology. I. Title. II. Series. QP261.G87 1989596'.03'2 - dc 19 88-36896
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© Springer-Verlag BerOO Heidelberg 1989
Softcover reprint of the hardcover 1st edition 1989
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Dedicated to Dr. M. S. Randhawa
Preface
In 1954, when I obtained my B. Sc. (Honours School) degree in
Zoology, I was asked by my advisor, the late Professor Vishwa
Nath, to make comparative cytological studies of oogenesis in
birds for my M.Sc. (Honours School) thesis submitted to the
Panjab University in 1956. Using the classical techniques of
cytology, I studied the various aspects of follicle growth and
atresia in five species of birds. Professor Vishwa Nath, who
himself devoted most of his life to the study of the comparative
cytology of gametogenesis in animals, was so much impressed
by my findings on avian oogenesis that he asked me to continue
my research work for the Ph.D. degree under his supervision.
During my research work on avian oogenesis, I became very
much interested in further comparative study of ovarian
follicles in vertebrates and thus decided to make comparative,
morphological and histochemical studies of ovarian follicles in
reptiles, birds and mammals. Actually, very little information
was available in this regard in 1957 when I started working for
the Ph.D. degree. In subsequent years, I and my numerous
dedicated and highly motivated M.Sc. and Ph.D. students con
tinued to make comparative, morphological, histochemical,
biochemical and physiological studies of ovarian follicles in all
groups of vertebrates. In connection with the subject of this
book, I would like to make special mention of my two brilliant
students, namely Dr. S. K. Varma and Dr. R. K. Chalana (now
Parshad), who carried out excellent studies on the ovarian
follicles of reptiles and birds, respectively. Many other workers
from allover the world, using techniques of transmission and
scanning electron microscopy, histochemistry, autoradiography,
biochemistry and physiology, have also studied the ovarian
follicles of different species of vertebrates. The major objective
of all of these studies has been to reveal differences at the
cellular, subcellular and molecular levels in ovarian follicles of
vertebrates for the better understanding of their structure,
chemistry, function and evolution. In other words, how the rep
tilian and avian follicles producing large cleidoic eggs differ
from those of other vertebrates, such as the fish and amphi
bians that produce numerous small yolky eggs, and eutherian
mammals that produce a small number of alecithal eggs. Ac
tually, a thorough understanding of the comparative biology of
VII
ovarian follicles in vertebrates is of great current interest to a
wide variety of academic and scientific disciplines, as the in
vestigation of their structure, function and control involves
morphology (including ultrastructure), cell, molecular and
developmental biology, physiology, endocrinology, biochemis
try, immunology, neurobiology and pharmacology. Zoologists
study comparative and evolutionary aspects of biology of
ovarian follicles in different classes of Vertebrata. Agricultural
scientists, wildlife biologists, physiologists and toxicologists
study the biology of ovarian follicles for the better understan
ding and management of fertility, sterility and pathological
problems in domestic and endangered species of vertebrates.
Keeping in view the importance of investigating various
aspects of biology of ovarian follicles in vertebrates, numerous
papers and review chapters on their different aspects continue
to be published annually in different journals and books. It is
not possible for a scientist, teacher and student to study even
a small fraction of these publications. Very little effort was
made previously to review and integrate the vast amount of
literature available and then to point out the gaps in our
knowledge about the physiological, endocrinological, biochem
ical, histochemical and morphological aspects of ovarian follic
les in vertebrates. Although much needed, a multidisciplinary
approach in the discussion of biology of ovarian follicles in
vertebrates is very difficult, and thus was lacking in previous
publications. Therefore, by critically reviewing and integrating
the results from a wide range of morphological, biochemical
and physiological techniques, as well as by comparison of
species, I have already published two books on ovarian follicles.
The first book Biology of Ovarian Follicles in Mammals was
published by Springer-Verlag in 1985. The second The Cell and
Molecular Biology of Fish Oogenesis was published as Volume
18 of the series Monographs in Developmental Biology by S.
Karger in 1986. The purpose of the present book is also to pre
sent a timely, thorough review on various aspects of com
parative biology of ovarian follicles in reptiles and birds, which
produce large cleidoic eggs, so that the students, zoologists,
biologists, and scientists from other disciplines can review the
current state of knowledge in the biology of ovarian follicles in
reptiles and birds. These three books have grown out of our
numerous comparative studies on the biology of ovarian
follicles in vertebrates. The present book is organized into nine
chapters dealing with comparative, cellular, molecular, physio
logical (including endocrinological) and evolutionary aspects
of ovarian normal, atretic and postovulatory follicles in reptiles
and birds and also presenting thorough, up-to-date reviews and
bibliography on various aspects of the biology of the ovarian
VIn
follicles; future research needs related to each chapter are also
clearly pointed out. The various chapters are contemporary
reviews which will serve as an important source for investigators
of the comparative biology of ovarian follicles in reptiles and
birds for years to come.
These three books clearly show that there still exist great
gaps in our knowledge of the comparative biology of ovarian
follicles in vertebrates. It is hoped that this book and the previ
ous two will serve as a stimulus for zoologists, reproductive
biologists, wildlife biologists, animal breeders, physiologists,
endocrinologists, and cellular, molecular and developmental
biologists to fill these voids, so that we come in a better position
to solve fertility and sterility problems in wild and domesticated
vertebrates, including reptiles and birds, as well as in man.
For this book, I am greatly indebted to editors of the series
Zoophysiology for making useful comments on its proposed
contents and Introduction submitted for their approval. My
special thanks are due to the late Professor Donald S. Farner
for taking special interest and acting as the responsible editor
for this book and to my students for their helpful cooperation.
I must thank authors and copyright holders for permission to
republish some of their illustrations. Thanks are due to Sardar
Inerjit Singh for typing the manuscript and to Sardar Chain
Singh for helping me in the mounting and labelling of figures.
Thanks are particularly due to the staff, especially Dr. Dieter
Czeschlik, of the Biology Editorial Department, Springer
Verlag, Heidelberg, for their helpful cooperation during the
completion and publication ofthis book. I would like to extend
appreciation and thanks to my wife Surinder and my children,
Gurmeet, Harmeet and Rupa for providing encouragement, in
spiration and patience during the time I was writing this book.
Ludhiana, Winter 1988/89 SARDUL S. GURAYA
IX
Contents
Introduction .................................... .
Chapter 1. Germinal Beds, Oogenesis and
Folliculogenesis .................................. 6
1.1 Reptiles ..................................... 6
1.2 Birds ........................................ 12
1.3 Concluding Remarks .......................... 19
Chapter 2. Nucleus or Germinal Vesicle 22
2.1 Reptiles ..................................... 22
2.2 Birds ........................................ 26
2.3 Concluding Remarks .......................... 33
Chapter 3. Ooplasmic Components .. . . . . . . . . . . . . . . . 35
3.1 Reptiles ..................................... 35
3.1.1 Morphology, Chemistry and Function ...... 35
3.1.2 Seasonal Changes in Ooplasmic Components 42
3.1.3 Cortical Granules ........................ 46
3.2 Birds ........................................ 46
3.2.1 Morphology, Chemistry and Function ...... 46
3.2.2 Cortical Granules ........................ 60
3.3 Concluding Remarks .......................... 60
Chapter 4. Vitellogenesis .......................... 63
4.1 Reptiles ..................................... 63
4.1.1 Fatty Yolk .............................. 65
4.1.2 Proteid Yolk ............................ 68
4.2 Birds ........................................ 77
4.2.1 Fatty Yolk .............................. 78
4.2.2 Proteid Yolk ............................ 79
4.2.3 Chemistry .............................. 86
XI
4.2.4 Hormonal Regulation of Yolk Synthesis in
the Liver and its 'Itansport ............... 92
4.3 Concluding Remarks .......................... 94
Chapter 5;·Follicle Wall... . .... .. . .... . ........ . .. 96
5.1 Reptiles ..................................... 96
5.1.1 Follicular Epithelium. . . . . . . . . . . . . . . . . . . . . 97
5.1.2 Zona Pellucida .......................... 109
5.1.3 Basement or Basal Lamina ............... 111
5.1.4 Theca .................................. 111
5.1.5 Epithelium ............................. 114
5.2 Birds. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
5.2.1 Follicular Epithelium. . . . . . . . . . . . . . . . . . . . . 116
5.2.2 Zona Pellucida .......................... 134
5.2.3 Basal Lamina ........................... 137
5.2.4 Theca .................................. 138
5.2.5 Epithelium ............................. 145
5.3 Concluding Remarks .......................... 146
Chapter 6. Maturation oj Ovum ................... 148
Chapter 7. Ovulation ............................. 153
7.1 Reptiles ..................................... 154
7.1.1 Hormonal Regulation .................... 154
7.1.2 Mechanisms ............................ 156
7.2 Birds ........................................ 159
7.2.1 Hormonal Regulation .................... 160
7.2.2 Mechanism ............................. 167
7.3 Concluding Remarks.......................... 171
Chapter 8. Postovulatory Follicles .................. 174
8.1 Reptiles ..................................... 174
8.1.1 Morphology and Histochemistry .......... 174
8.1.2 Biochemistry and Physiology.. . . .. ... . . . . . 181
8.2 Birds ........................................ 187
8.2.1 Morphology and Histochemistry .......... 187
8.2.2 Biochemistry and Physiology .............. 194
8.3 Comparisons of Postovulatory Follicles in Reptiles
and Birds and Their Evolutionary Significance 197
XII