Juan /. Tarin • Antonio Cano (Eds.) Fertilization in Protozoa and Metazoan Animals

Springer-Verlag Berlin Heidelberg GmbH

Juan J. Tarin • Antonio Cano (Eds.)

Fertilization in Protozoa and Metazoan Animals Cellular and Molecular Aspects

With 43 Figures

IIP Springer

J. J. T A R I N , PhD Department of Animal Biology Faculty of Biological Sciences University of Valencia, Burjassot 46100 Valencia Spain

A. C A N O , M D Department of Paediatrics Obstetrics and Gynaecology Faculty of Medicine University of Valencia 46010 Valencia Spain

ISBN 978-3-642-63530-4

Library of Congress Cataloging-in-Publication Data Fertilization in protozoa and metazoan animals : cellular and molecular aspects / Juan J. Tarfn, Antonio Cano (eds.).

p. cm. Includes bibliographical references (p.). ISBN 978-3-642-63530-4 ISBN 978-3-642-58301-8 (eBook) DOI 10.1007/978-3-642-58301-8

1. Fertilization (Biology) 2. Physiology, Comparative. I. Tarin, Juan J., 1959- II. Cano, Antonio, 1954-QH485 .F476 2000 571.8/64-dc21

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Preface

Reproduction is the origination of new organisms from pre-existing ones. Among more than 35 separated forms of reproduction including several types of gamogony, parthenogenesis, agamogenesis, fission and division, and plas-motomy, the bisexual mode of reproduction via fertilization provides genetic variability that allows species to adapt quickly to competitive and constantly changing environments. Several excellent reviews and books have been written in the past to analyse the mechanisms of fertilization in different eukaryotic species. During the last few years, however, renewed attention has been paid to examining the process of oocyte fertilization at the cellular/molecular level not only within a single species/group but also through different phylogenetic lineages. As a result of this effort, knowledge of the molecular pathways used by oocytes and spermatozoa at fertilization has increased, but still many ques-tions remain to be answered. Being aware of the necessity of providing an inte-grated view of the process of fertilization, this book has been entirely devoted to reviewing the process of oocyte fertilization at the cellular/molecular level in two different and separated groups of eukaryotic organisms: protozoa and metazoan animals.

The book is organized into six sections dealing with oocyte fertilization in protozoa, invertebrates, teleost fishes, amphibians, birds and mammals. These sections are followed by a summary/concluding chapter that provides a com-parative overview of the process of fertilization in these groups of eukaryotes. As research on fertilization at the cellular/molecular level is limited and/or fragmentary when compared to the huge diversity of protozoan and animal species, we chose a single or a reduced number of species (those from whom more data were available) as the representative(s) of each group. Although the strategy of extrapolating the mechanism(s) used at fertilization by a single or a reduced number of species to the rest of the species of the same phylogenetic group must be taken with caution because it overlooks the existent interspecies variability, it can be used for integrative purposes. Each of the six major sec-tions begins with a general description of the sexual and asexual reproduction strategies of the species belonging to a particular group. Then, a description of the reproductive cycle as well as the design of the body and reproductive system (including gametogenesis) of the representative species is given. These form the introductory steps before focusing on the cellular/molecular aspects of fertilization. Special care was taken to analyse all the fertilization steps from sperm capacitation to karyogamy. However, in many cases, a scarce or total

VI Preface

absence of information precluded us from following this stratagem and some groups were partially treated or even not included in this monograph.

A major inspiration for this book was the work of Prof. Juan Brines, Uni-versity of Valencia, on some paediatric topics from an evolutionary perspec-tive. His insight on infant and child feeding from a biological point of view impacted so deeply on our understanding as undergraduate students that it spurred us on to produce this book. We thank Prof. Brines, not only for his inspiring work, but also for his constant support and helpful discussions on evolutionary issues. We hope this book will be the first step in a series of addi-tional thoughts and analyses of oocyte fertilization from a comparative and evolutionary viewpoint.

University of Valencia, Spain JUAN J. TARIN and ANTONIO CANO

Contents

1 Fertilization in Protozoa KAZUYUKI MIKAMI

1 1.1 1.2 1.3

2 2.1 2.2 2.3 2.3.1 2.3.2 2.3.3 2.3.4 2.3.5 2.3.6

2.3.7 2.4

3 3.1 3.2 3.3

3.4

4 4.1 4.2 4.3

5 5.1 5.2 5.3

Reproduction Strategies in Protozoa ..................... . Gametogamy ......................................... . Autogamy ........ . .................................. . Gamontogamy ........... , ........................... .

Fertilization in a Flagellate ............................. . Reproduction in Chlamydomonas reinhardtii .............. . Induction of Gametogenesis ............................ . Mating and Cell Fusion ................................ . Flagellar Agglutination ................................ . Losing Cell Wall and Protoplast Formation ................ . Activation of Mating Structures ......................... . Formation of a Cytoplasmic Bridge ...................... . Protoplasmic Fusion .................................. . Nuclear Fusion, Flagellar Retraction and Zygotic Wall Formation ........................................... . Chloroplast .......................................... . Meiosis and Germination .............................. .

Sexual Phase and Fertilization in Plasmodium ............. . Reproduction in Plasmodium falciparum ................. . Gametogenesis and Fertilization in the Mosquito .......... . Asexual Development in the Mosquito and in the Human Body ................................ . The Sexual Phase ..................................... .

Gamogony and Agamogony in Foraminiferans ............ . Life Cycle of Foraminiferans ............................ . Heterokaryotic Species Rotaliella heterocaryotica .......... . Homokaryotic Species Patellina corrugata ................ .

Conjugation in Ciliates ................................ . Life Cycle of Paramecium caudatum ..................... . Sexual Interaction at the Beginning of Conjugation ........ . Nuclear Events During Conjugation ..................... .

1 1 2 3

4 4 4 6 6 6 6 7 7

7 7 8

8 8 8

10 11

11 11 13 13

15 15 16 17

VIII Contents

5.3.1 Nuclear Behavior at Early Stages of Conjugation and Meiosis .......................................... 17

5.3.2 Random Selection of Meiotic Products .................... 18 5.3.3 Gametogenesis and Fertilization ......................... 18 5.3.4 Macro- and Micronuclear Differentiation .................. 20 5.3.5 Development of a New Macronucleus. . . . . . . . . . . . . . . . . . . ... 20 5.3.6 Determination of the Germinal Micronucleus .............. 21

References ............................................ 22

2 Fertilization in Invertebrates

1

2 2.1 2.1.1 2.1.2 2.2 2.2.1 2.2.2 2.2.3

3 3.1 3.1.1 3.1.2 3.2

4 4.1 4.1.1 4.1.2 4.2

5 5.1 5.1.1 5.1.2 5.2

6 6.1 6.1.1 6.1.2 6.2

7 7.1

HEIDE SCHATTEN and AMITABHA CHAKRABARTI

Introduction ......................................... .

Porifera ............................................. . Reproductive Strategies and Reproductive Cycle ............ . Design of Reproductive System .......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............ . Sperm ............................................... . Oocytes ............................................. . Fertilization ......................................... .

Cnidaria (Coelenterata) ................................ . Reproductive Strategies and Reproductive Cycle ........... . Design of Reproductive System ......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............. .

Ctenophora .......................................... . Reproductive Strategies and Reproductive Cycle ........... . Design of Reproductive System .......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............ .

Platyhelminthes ...................................... . Reproductive Strategies and Reproductive Cycle ........... . Design of Reproductive System ......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............ .

Rhynchocoela (Nemer tea, Nemertinea) ................... . Reproductive Strategies and Reproductive Cycle ........... . Design of Reproductive System ......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............ .

Nematoda ........................................... . Reproductive Strategies and Reproductive Cycle ........... .

27

28 28 29 29 30 30 30 30

31 32 32 32 33

34 34 34 34 34

36 36 36 37 37

38 38 39 39 39

40 40

7.1.1 7.1.2 7.2 7.2.1 7.2.2 7.2.3

8 8.1 8.1.1 8.1.2 8.2 8.2.1 8.2.2 8.2.3

9 9.1 9.1.1 9.1.2 9.2

10 10.1 10.1.1 10.1.2 10.2 10.2.1 10.2.2 10.2.3

11 11.1 11.1.1 11.1.2 11.2 11.2.1 11.2.2 11.2.3 11.2.4 11.3

12

Contents IX

Design of Reproductive System ......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............ . Sperm .............................................. . Oocytes ............................................. . Fertilization ......................................... .

Mollusca ............................................ . Reproductive Strategies and Reproductive Cycle ........... . Design of Reproductive System ......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............ . Sperm .............................................. . Oocytes ............................................. . Fertilization ......................................... .

Annelida ............................................ . Reproductive Strategies and Reproductive Cycle ........... . Design of Reproductive System ......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............ .

Arthropoda ......................................... . Reproductive Strategies and Reproductive Cycle ........... . Design of Reproductive System ......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............ . Sperm .............................................. . Oocytes ............................................. . Fertilization ......................................... .

Echinodermata ....................................... . Reproductive Strategies and Reproductive Cycle ........... . Design of Reproductive System ......................... . Gametogenesis and Fertilization ........................ . Cellular and Molecular Aspects of Fertilization ............ . The Unfertilized Egg .................................. . Sperm .............................................. . Sperm Activation ..................................... . Fertilization ......................................... . MPF and MAP Kinases During Meiotic Maturation and Fertilization ...................................... .

Proto chordata

41 41 42 42 43 43

44 44 44 45 45 45 45 46

47 47 48 48 49

51 51 52 52 53 53 54 54

56 56 57 57 57 58 58 60 61

67

67 12.1 Reproductive Strategies and Reproductive Cycle ............ 67 12.1.1 Design of Reproductive System .......................... 68 12.1.2 Gametogenesis and Fertilization ......................... 68 12.2 Cellular and Molecular Aspects of Fertilization ............. 68

X Contents

12.2.1 12.2.2 12.2.3 12.2.4 12.2.5 12.3

Sperm .............................................. . Oocytes ............................................. . Spawning of Gametes and Fertilization .................. . Calcium Oscillations .................................. . Block to Polyspermy .................................. . MPF and MAP Kinases During Meiotic Maturation and Fertilization ...................................... .

69 69 70 71 72

73

References ............................................ 74

3 Fertilization in Fishes T AKASHI IWAMATSU

1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

2 Structural Preparation of Oocyte and Spermatozoon for Fertilization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90

2.1 Egg. . . . . . . . . .. . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 2.1.1 Chorion.............................................. 91 2.1.2 Micropyle ............................................ 96 2.1.3 Cortical Layer ......................................... 97 2.1.4 Sperm Entry Site ...................................... 99 2.1.5 Nucleus .............................................. 100 2.2 Spermatozoon 101

3 Oocyte Maturation for Acquisition of Fertilizability ......... 103

4 Interaction and Membrane Fusion Between Gametes ........ 104 4.1 Behavior and Fertilizability of Spermatozoa ............... 104 4.2 Sperm-Egg Interaction ................................. 106 4.3 Sperm-Egg Membrane Fusion ........................... 106

5 Responses of Egg to Membrane Fusion with Spermatozoon ... 108 5.1 Change in Membrane Potential. . . . . . . . . . . . . . . . . . . . . . . . . . . 108 5.2 Fertilization Wave and Wave of Increase

in Cytoplasmic Ca2+ • • • • • . • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 109 5.3 Exocytosis of Cortical Alveoli ........................... 111 5.4 Mechanism of Propagative Increase in Cytoplasmic Ca2+

and pH . ............................................. 114 5.5 Relationship Between Wave of Increase in Cytoplasmic Ca2+

and Exocytosis Wave . . ................................. 115 5.6 Perivitelline Space Formation and Change in Chorion. . . . . . . . 115 5.7 Fertilization Cone and Sperm Incorporation ............... 117

6

7 7.1

Polyspermy-Preventing Mechanism

Formation of Male and Female Pronuclei ................. . Progression of Second Meiotic Metaphase to Interphase ......................................... .

119

122

122

7.2

7.3 7.4

8

9

Contents XI

Breakdown of Nuclear Envelope and Chromatin Decondensation ....................................... . Pronucleus Formation and MPF Activity ................. . DNA Synthesis and Pronucleus Formation ................ .

Migration and Karyogamy of Female and Male Pronuclei

Fertilization Without an Increase in Cytoplasmic Ca2+ and Exocytosis of Cortical Alveoli ....................... .

124 125 126

127

127

References ............................................ 129

4 Fertilization in Amphibians YASUHIRO IWAO

Introduction .......................................... 147

2 Interactions of Sperm with Oocyte Investments and Oocyte Membrane. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150

2.1 Interactions of Sperm with Jelly Layers .................... 150 2.2 Sperm-Binding to the Vitelline Envelope ................... 153 2.3 Acrosome Reaction .................................... 155 2.4 Interactions Between Sperm-and Oocyte-Membranes ........ 156

3 Mechanism of Oocyte Activation ......................... 157 3.1 [Ca2+L Increase ........................................ 157 3.2 Fertilization Potentials ................................. 160 3.3 Cortical Reactions and Fertilization Envelope Formation ..... 162 3.4 Resumption of Meiosis and Sperm Nuclear

Decondensation ....................................... 164

4 Mechanisms for Polyspermy Block ....................... 165 4.1 Role of Polyspermy Block ............................... 165 4.2 Reduction in Number of Sperm in Oocyte Investments ...... 166 4.3 Fast, Electrical Block to Polyspermy ...................... 167 4.4 Slow Polyspermy Block by the Fertilization Envelope ........ 169 4.5 Acquisition of Polyspermy Block During Oocyte Maturation

in Anurans ........................................... 169 4.6 Cytoplasmic Block to Polyspermy in Physiologically

Polyspermic Oocytes ................................... 170

5 Variations in Amphibian Fertilization ..................... 176 5.1 Polyspermy in Anurans ................................. 176 5.2 Monospermy in Primitive Urodeles ....................... 177 5.3 Sperm Nuclear Behavior in Gynogenetic Urodele Oocytes .... 178

References ............................ . ............... 179

XII Contents

5 Fertilization in Birds

2 2.1 2.2 2.2.1 2.2.2 2.3 2.3.1 2.3.2 2.4 2.4.1 2.4.2 2.4.3 2.5 2.5.1 2.5.2

3

4 4.1 4.1.1 4.1.2 4.2 4.2.1 4.2.2 4.2.3

4.2.4

4.2.5

4.3 4.3.1 4.3.2 4.3.3

5

GRAHAM J. WISHART and A. JANET HORROCKS

Introduction to Avian Breeding Systems

Gametes, Gameteogenesis and Reproductive Systems ....... . Primordial Germ Cells and Establishment of Gonads ....... . Male Reproduction System and Sperm Production ......... . Male Tract Anatomy ................................... . Maturation of Sperm Function .......................... . Female Reproductive System ........................... . The Ovary and Ovulation .............................. . The Oviduct and Egg Formation ........................ . Spermatozoa ......................................... . Sperm Morphology .................................... . Sperm Motility ....................................... . The Acrosome and Acrosomal Enzymes .................. . The Ovum .......................... . ................ . Morphology of the Ovum at Ovulation ................... . The Inner Perivitelline Layer ........................... .

Oviducal Sperm Transport and Storage ................... .

Sperm:Egg Interaction ................................. . Fertilization in Vivo and in Vitro ........................ . Site of Fertilization in Vivo ............................. . Capacitation ......................................... . Sperm: Perivitelline Layer Interaction .................... . Sperm: Inner Perivitelline Layer Interaction ............... . The Avian Acrosome Reaction .......................... . Inner Perivitelline Layer Molecules Mediating Interaction with Spermatozoa ..................................... . Preferential Attraction of Spermatozoa to the Germinal Disc ................................... . The Outer Perivitelline Layer and Cessation of Sperm: Inner Perivitelline Layer Interaction ...................... . Sperm: Ovum Interaction and Syngamy ................... . Sperm Engulfment by Oolemma ........................ . Pronuclear Formation and Syngamy ..................... . Supernumerary Spermatozoa and Embryonic Development ......................................... .

Quantitative Aspects of Sperm: Egg Interaction, Fertility and the Fertile Period ............ .

193

195 195 196 196 197 198 198 198 200 200 202 202 203 203 204

205

206 206 206 207 207 207 209

209

211

212 213 213 214

215

216

References ............................................ 217

Contents XIII

6 Fertilization in Monotreme, Marsupial and Eutherian Mammals

1 1.1 1.2 1.3 1.3.1 1.4 1.5 1.6

2 2.1 2.2

3 3.1 3.1.1 3.1.2 3.1.3 3.2 3.2.1 3.2.2 3.3 3.4

4

4.1 4.2

5 5.1 5.1.1 5.2 5.2.1 5.2.2 5.3 5.4

6 6.1 6.2 6.3

7 7.1

K.E. MATE, M.S. HARRIS, and J.e. RODGER

Introduction ......................................... . The Three Groups of Mammals ......................... . Monotremes ......................................... . Marsupials .......................................... . Marsupial Diversity .................................... . Eutherians ........................................... . Mammalian Relationships and Evolution ................. . Fertilization in Mammals .............................. .

Sperm Production .................................... . Marsupials and Eutherians ............................. . Monotremes ......................................... .

Epididymal Maturation of Spermatozoa .................. . Eutherians .......................................... . Morphological Maturation ............................. . Development of Progressive Motility ..................... . Maturation of Sperm Surface ........................... . Marsupials .......................................... . Morphological Changes ............................... . Molecular Changes .................................... . Monotremes ......................................... . Common Themes and General Overview ................. .

Sperm Deposition, Storage and Transport in the Female Tract .................................... . Marsupials and Eutherians ............................. . Monotremes ......................................... .

Capacitation ......................................... . Eutherians ........................................... . Molecular Basis ...................................... . Marsupials ........................................... . Morphological Correlates .............................. , Differences Between Marsupials ......................... . Monotremes ......................................... . Common Themes and General Overview .................. .

Hyperactivation of Spermatozoa ........................ . Eutherians ........................................... . Marsupials and Monotremes ............................ . Common Themes and General Overview .................. .

The Acrosome ....................................... . Mammalian Acrosome Morphology and Enzymes .......... .

223 223 223 224 226 227 227 227

228 228 228

230 230 230 230 232 233 233 234 235 235

236 236 237

237 237 238 238 239 239 241 241

242 242 243 243

244 244

XIV Contents

7.2 7.2.1 7.2.2 7.2.3 7.3 7.4

8 8.1 8.Ll 8.1.2 8.1.3 8.2 8.3 8.3.1 8.3.2 8.4

9 9.1 9.2 9.3 9.4

10 10.1 10.2 10.3 10.4

11 ILl

Acrosome Reaction in Eutherians ....................... . Morphology and Functional Significance in Vivo ........... . Mechanism of the Acrosome Reaction in Vivo ............. . Mechanism of the Acrosome Reaction in Vitro Acrosome Reaction in Marsupials and Monotremes ........ . Common Themes and General Overview ................. .

Sperm Interaction with the Zona Pellucida ................ . Eutherians .......................................... . Structure and Function of the ZP ........................ . Primary Sperm-ZP Binding ............................ . Penetration of the ZP and Secondary Sperm-ZP Binding .... . Monotremes ......................................... . Marsupials .......................................... . Structure and Function of the ZP ........................ . Sperm Interaction with the ZP .......................... . Common Themes and General Overview ................. .

Sperm-Oocyte Fusion ................................. . Eutherians ........................................... . Monotremes ......................................... . Marsupials ........................................... . Common Themes and General Overview ................. .

Oocyte Activation .................................... . Eutherians ........................................... . Monotremes ......................................... . Marsupials ........................................... . Common Themes and General Overview ................. .

Cortical Reaction and Block to Polyspermy ............... . Eutherians ........................................... .

l1.Ll Cortical Granules

244 244 246 246 247 247

248 248 248 251 251 252 252 252 253 254

254 254 255 255 255

256 256 257 258 258

258 258 259

11.1.2 Cortical Reaction ...................................... 259 11.2 Monotremes .......................................... 259 11.3 Marsupials ........................................... 260 11.3.1 Cortical Granules ......... . ............................ 260 11.3.2 Cortical Reaction ...................................... 261 11.4 Common Themes and General Overview .................. 261

12 Decondensation of the Sperm Nucleus .................... 262 12.1 Eutherians. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262 12.2 Monotremes.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262 12.3 Marsupials ........................................... 262 12.4 Common Themes and Overview ......................... 263

13 Completion of Meiosis, Pronuclei Development and Syngamy .... . .................................... 264

13.1 13.1.1 13.1.2 13.2 13.3 13.4

14

Contents XV

Eutherians ........................................... . Completion of Oocyte Meiosis .......................... . Pronuclei Development and Syngamy .................... . Completion of Fertilization in Monotremes ............... . Completion of Fertilization in Marsupials ................ . Common Themes and Overview ........................ .

Concluding Remarks .................................. .

264 264 264 265 265 266

266

References ............................................ 267

7 Fertilization in Protozoa and Metazoan Animals: A Comparative Overview JUAN J. TARiN

1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277

2 Definition of Fertilization ............................... 277

3 Somatic and Germ Cell Lines in Protozoa and Metazoa ...... 278

4 Comparative Analysis of Fertilization in Metazoa ........... 280 4.1 Sperm Capacitation and the Environment of Fertilization . . . . . 280 4.2 Composition of Oocyte Envelope(s) ...................... 280 4.3 Sperm Binding to Oocyte Envelope(s) .................... 292 4.4 Acrosome Reaction .................................... 294 4.5 Binding and Fusion of Spermatozoon to Oocyte Plasma

Membrane ........................................... 294 4.6 Oocyte Activation ..................................... 295 4.6.1 Changes in Plasma Membrane Potential ................... 297 4.6.2 Increase in [Ca2+L ..................................... 299 4.6.3 Cortical Alveolus/Granule Exocytosis ..................... 303 4.6.4 Ooplasmic Segregation ................................. 304 4.6.5 Increase in pHi ........................................ 307 4.6.6 Resumption of the Cell Cycle ............................ 307 4.6.7 Development of Male Pronucleus ......................... 309 4.6.8 Pronuclear Apposition and Karyogamy .................... 310

References ............................................ 311

Subject Index ............................................... 315

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