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Wolpert text

Lewis Wolpert, Rosa Beddington, Thomas Jessell, Peter Lawrence, Elliot Meyerowitz, Jim Smith
Principles of Development, Second Edition
Oxford University Press
ISBN 0-19-924939-3


Developmental biology is at the core of all biology. It deals with the process by which the genes in the fertilized egg control cell behavior in the embryo and so determine its pattern, its form, and much of its behavior. The progress in developmental biology in recent years with the application of advances in cell and molecular biology has been remarkable, and an enormous amount of information is now available.

In this second edition, we have included many recent advances, for example in the understanding of somite formation, and this new material is complemented by over 30 additional illustrations. Sections on the development of the heart, the vascular system, and teeth have been added, and we have given more attention, for example, to stem cells, signal transduction, and evolution.

Principles of Development is designed for undergraduates and the emphasis is on principles and key concepts. Central to our approach is that development can be best understood by understanding how genes control cell behavior. We have assumed that the students have some very basic familiarity with cell biology and genetics, but all key concepts, such as the control of gene activity, are explained in the text.

Conscious of the pressures on students, we have tried to make the principles as clear as possible and to provide numerous summaries, in both words and pictures. The illustrations in this book are a special feature and have been carefully designed and chosen to illuminate both experiments and mechanisms.

We have resisted the temptation to cover every aspect of development and have, instead, focused on those systems that best illuminate common principles. Indeed, a theme that runs throughout the book is that universal principles govern the process of development. At all stages, what we included has been guided by what we believe undergraduates should know about development.

We have thus concentrated our attention on vertebrates and Drosophila, but not to the exclusion of other systems, such as the nematode and the sea urchin, where they best illustrate a concept. An important feature of our book is the inclusion of the development of plants, which is usually neglected in textbooks. There have been striking advances in plant developmental biology in recent times, and some unique and important features have emerged. As knowing the basic features of the embryology of the main organisms used to study development is essential for an understanding of molecular mechanisms, we have introduced embryology at an early stage.

Whereas our emphasis has been on the laying down of the body plans and organ systems, such as limbs and the nervous system, we have also included later aspects of development, including growth and regeneration. The book concludes with a consideration of evolution and development.

In providing references, our prime concern has been to guide the students to helpful papers rather than to give credit to all the scientists who have made major contributions: to those whom we have neglected, we apologize.

For this new edition we welcome Jim Smith as a co-author and say goodbye to Jeremy Brockes, with thanks for all his help. Each chapter has also been reviewed by a number of experts (see page xix), to whom we give thanks. As with the first edition, I did all the writing-and I mean writingin consultation with my co-authors; it was typed by Maureen Maloney and my revisions were deciphered, edited, and incorporated by our editor Eleanor Lawrence, whose expertise and influence pervades the book. Most of the new illustrations were brilliantly created or adapted by Matthew McClements, who created the illustrations for the first edition.

We are indebted to Jonathan Crowe, John Grandidge, and the illustrators at Oxford University Press, our new publishers, for their help and patience throughout the preparation of this new edition. My thanks to Vitek Tracz and Peter Newmark of the Current Science Group, without whom the first edition of this book would never have been started, let alone completed.

Finally, and sadly, we are acutely conscious of the loss of Rosa Beddington, who died last year, after a long illness. She was helpful to the end and we miss her terribly.

L. W.
September 2001


List of headings
Text acknowledgments
Figure acknowledgments
Chapter 1: History and basic concepts
The origins of developmental biology
A conceptual tool kit
Chapter 2: Model systems
Model organisms: vertebrates
Model organisms: invertebrates
Model systems: plants
Identifying developmental genes
Chapter 3: Patterning the vertebrate body plan I: axes and germ layers
Setting up the body axes
The origin and specification of the germ layers
Chapter 4: Patterning the vertebrate body plan II: the mesoderm and early nervous system
Somite formation and patterning
Neural induction and the role of the organizer
Chapter 5: Development of the Drosophila body plan
Maternal genes set up the body axes
Polarization of the body axes during oogenesis
Zygotic genes pattern the early embryo
Segmentation: activation of the pair-rule genes
Segment polarity genes and compartments
Segmentation: selector and homeotic genes
Chapter 6: Development of nematodes, sea urchins, ascidians, and slime molds
Cellular slime molds
Chapter 7: Plant development
Embryonic development
Flower development
Chapter 8: Morphogenesis: change in form in the early embryo
Cell adhesion
Cleavage and formation of the blastula
Neural tube formation
Cell migration
Directed dilation
Chapter 9: Cell differentiation
The control of gene expression
Models of cell differentiation
The reversibility of patterns of gene activity
Chapter 10: Organogenesis
The chick limb
Insect wings and legs
The nematode vulva
Internal organs: heart, blood vessels, lungs, kidneys, and teeth
Chapter 11: Development of the nervous system
Specification of cell identity in the nervous system
The insect compound eye
Axonal guidance
Neuronal survival, synapse formation, and refinement
Chapter 12: Germ cells and sex
Determination of the sexual phenotype
The development of germ cells
Chapter 13: Regeneration
Limb regeneration
Regeneration in Hydra
Chapter 14: Growth and post-embryonic development
Molting and metamorphosis
Aging and senescence
Chapter 15: Evolution and development
Modification of development in evolution
Changes in the timing of developmental processes during evolution
Evolution off development


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Developmental Biology
Published by Elsevier Science under Auspices of Society for Developmental Biology
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