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1 Developmental Biology

2 Springer New York Berlin Heidelberg Barcelona Budapest Hong Kong London Milan Paris Santa Clara Singapore Tokyo

3 Werner A. Miiller Developmental Biology With 123 Figure Springer

4 Werner A. Muller Zoologisches Institut-Physiologie University of Heidelberg D Heidelberg Germany Cover: Foreground: a 13.5-day-old mouse embryo showing the expression of ptc mrna by in situ hybridization. Courtesy of Ljiljana Milenkovic and Matthew P. Scott. Background: double immunofluorescence of wild type Drosophila eye imaginal disk cells stained for dtfiia-s (red) and Elav (green). Image provided by Martin P. Zeidler and Mark Mlodzik. This edition is published by arrangement with Gustav Fischer Verlag, Stuttgart. Original copyright Gustav Fischer Verlag. Title of German original: Werner A Miiller, EntwickIungsbiologie Einfuhrung in die kiassische und molekulare EntwickIungsbiologie von Mensch und Tier Library of Congress Cataloging-in-Publication Data Miiller, Werner A., Developmental biology / Werner A. MUller. p. cm. Includes bibliographical references and index. ISBN-13: DOI: / e-isbn-13: Developmental biology. I. Title. QH491.M dc Printed on acid-free paper. English Translation 1997 Springer-Verlag New York, Inc. Softcover reprint of the hardcover 15t edition 1997 All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag New York, Inc., 175 Fifth Avenue, New York, NY 10010, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use of general descriptive names, trade names, trademarks, etc., in this publication, even if the former are not especially identified, is not to be taken as a sign that such names, as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Production supervised by Natalie Johnson and managed by Impressions Book and Journal Services, Inc. Manufacturing supervised by Jeffrey Taub. Typeset by Impressions Book and Journal Services, Inc., Madison, WI

5 Preface T his book is addressed to students of biology and medicine whose interests encompass human development as well as the most important model organisms currently being investigated with methods from genetics, biochemistry, and molecular biology. Developmental Biology has been written for advanced undergraduates as well as graduate students and instructors who seek a succinct but thorough introduction to contemporary developmental biology. A particular aim of this book is to bring together the rich heritage and pivotal ideas of the first "golden age" of developmental biology ( ) with discoveries and hypotheses of recent research. Modern research in developmental biology is based mainly on biochemical and molecular methods, but increasingly incorporates computer modeling to cope with such intriguing and complex phenomena as biological pattern formation. The awarding of the 1995 Nobel prize in physiology or medicine for research on embryo development in Drosophila highlights a new "golden era" for developmental biology. This book takes full advantage of the molecular revolution that has swept through the biological sciences, but it also presents, in addition to data and observations, basic principles of development and points out the historical roots of our interpretations. v

6 vi PREFACE Many years of teaching experience have prompted me to follow an approach different from that used in other textbooks. After an introduction to terminology used in descriptive developmental biology, I introduce several "model" organisms of importance in research and teaching. I then introduce selected developmental processes and present them from a comparative perspective. Using this approach, students do not have to piece together the development of, for example, the frog (Xenopus) or the fruit fly (Drosophila), from fragmentary information scattered throughout many chapters (such as "Cleavage," "Gastrulation," and so forth). Nevertheless, the student will also find overviews of major themes, such as fertilization (Chapter 6), develop-' mental genetics (Chapter 10), or sexual development (Chapter 20). It is my experience that any redundancy arising from this approach will serve to reinforce the concepts. An additional feature of the book is the presentation of seven boxed essays, each providing a brief overview of a single topic, such as the history of developmental biology (Box 1), signal transduction (Box 3), or current methods of research in developmental biology (Box 7). Developmental biology is a discipline undergoing extremely rapid development: an explosion of papers appears daily in an ever-growing number of journals and monographs devoted to this field. It is my hope that reading this book will prepare a student to understand and appreciate information presented in more extensive (but perhaps less clearly structured) textbooks, and to follow the present state of knowledge by reading original research articles and current reviews. This book is a revision and translation of a volume published in Germany in To illustrate the book, I drew most of the figures with my own hands; a minority were prepared on a computer using a drawing program. Approximately one-third of the figures were redrawn by Peter Adam, reproducing and modifying my drafts or the figures cited in the legends. Following recent rules, the name of a gene is written in italics. To designate proteins derived from defined genes, this book uses capital letters. This enables the reader to easily distinguish, for example, the WINGLESS and EYE LESS proteins from wingless and eyeless flies. The following people have assisted in preparing this book for publication: the staff of Springer-Verlag, particularly the Editor, Robert C. Garber; Senior Production Editor, Natalie Johnson; and, in addition, the staff at Impressions Book and Journal Services, Inc. I also express my gratitude to all those who read (parts of) the original German edition and gave hints for improving the book. In particular, I would like to name Kurt Baumann, Giinter F achbach, Horst Grunz, Klaus Sander, and Einhard Schierenberg. Heidelberg, June 1996

7 Contents ze. _ Preface v 1. Development: Organisms Construct and Organize Themselves on the Basis of Inherited Information 1 2. Basic Stages, Principles, and Terms of Developmental Biology 9 3. Model Organisms in Developmental Biology Comparative Review: The Phylotypic Stage of Vertebrates, Common versus Distinct Features, and Aspects of Evolution The Egg Cell and the Sperm Get a Dowry The Start: Fertilization and Activation of the Egg 146 vii

8 viii CONTENTS 7. Precisely Patterned Cleavage Divisions Are Driven by an Oscillator Detennination: Cells Are Programmed and Committed to Their Fates Epigenetic Pattern Fonnation: New Patterns Are Created During Development Differentiation Is Based upon Differential Gene Expression that Is Programmed during Detennination Cell Differentiation Frequently Is Irreversible and Causes Cell Death; Early Cell Death Can Be Programmed Animal Morphogenesis Is Shaped Actively by Adhesion and Cell Migration Cell Journeys: Even Genn Cells and Cells of the Peripheral Nervous System Originate from Emigrant Precursors Development of the Nervous System: Cell Migration, Pathfinding, and Self-Organization Heart and Blood Vessels: Divergent Developmental Roads but One System in the End Stem Cells Enable Continuous Growth and Renewal Signal Molecules Control Development and Growth Cancer Comes from Disturbed Growth and Differentiation Control Metamorphosis: A Second Embryogenesis Creates a Second Phenotype Sex and the Single Gene 301

9 Contents IX 21. Regeneration and Renewal versus Loss and Death Life and Death: What Is the Major Mystery? 323 Bibliography 335 Index 373 Boxes Box 1 History: From the Soul to Information 3 Box 2 Famous Experiments with Eggs and Embryos: Cloning, Chimeras, Teratomas, and Transgenic Mice 106 Box 3 The PI Signal Transduction System 153 Box 4 Models of Biological Pattern Formation 184 Box 5 Signal Molecules Acting through Nuclear Receptors 200 Box 6 How Cells Communicate and Interact 271 Box 7 Contemporary Techniques in Developmental Biology 330