Ecdysone: Structures and Functions Guy Smagghe Editor
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Ecdysone: Structures and Functions Guy Smagghe Editor Ecdysone: Structures and Functions Editor Guy Smagghe Laboratory of Agrozoology Faculty of Bioscience Engineering Ghent University Belgium ISBN 978-1-4020-9111-7 e-ISBN 978-1-4020-9112-4 Library of Congress Control Number: 2008938015 © 2009 Springer Science + Business Media B.V. No part of this work may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording or otherwise, without written permission from the Publisher, with the exception of any material supplied specifically for the purpose of being entered and executed on a computer system, for exclusive use by the purchaser of the work. Printed on acid-free paper springer.com Preface The 16th International Ecdysone Workshop took place at Ghent University in Belgium, July 10–14, 2006 and drew some 150 attendees, many of these young students and postdoctoral associates. These young scientists had the opportunity to dis- cuss their work with many senior scientists at meals, breaks and during the several social events, and were encouraged to do so. This book resulting from the meeting is more up-to-date than might be expected since manuscripts were not delivered to the editor until 2007. The workshop itself had 54 oral presentations as well as many posters. This book, and the meeting itself, is comprised of 23 contributed chapters falling into five general categories: Fundamental Aspects of Ecdysteroid Research: The Distribution and Diversity of Ecdysteroids in Animals and Plants; Ecdysteroid Genetic Hierarchies in Insect Growth and Reproduction; Role of Cross Talk and Growth Factors in Ecdysteroid Titers and Signaling; Ecdysteroid Function Through Nuclear and Membrane Receptors; Ecdysteroids in Modern Agriculture, Medicine, Doping and Ecotoxicology. Clearly, all that was presented at the meeting and in this volume cannot be sum- marized here in a single page, but the reader is cordially invited to explore this rela- tively large volume that attempts to synthesize the most current knowledge base for this important category of steroid hormones. I use the plural because the nomencla- ture has indeed undergone a metamorphosis akin to what our experimental animals undergo. In the 1950s when the Butenandt and Karlson laboratory first crystallized and characterized ecdysone, it was thought to be the insect molting hormone since when injected into experimental insects the result was molting. Later, it was found that ecdysone was converted to 20-hydroxyecdysone in tissues peripheral to the prothoracic glands through the mediation of an ecdysone monooxygenase and we then believed that ecdysone was only the precursor for 20-hydroxyecdysone. However, a reading of the older literature as well as new work has demonstrated convincingly that ecdysone does have regulatory roles of its own, and perhaps joins with 20-hydroxyecdysone to initiate the molting process. Further, it must be remembered that the research of Ulrich Clever and Peter Karlson on Chironomus was the very first to demonstrate that a steroid hormone acts at the nuclear level and v vi Preface indeed Karlson wrote several theoretical papers subsequently urging endocrinolo- gists working on mammals to accept that view. In time they did! Despite the molecular focus of this meeting there remains much research to do at the physio- logical and biochemical levels, let alone in the areas of chemistry and crystallogra- phy, to further define the roles of these hormones that serve more organisms on this planet than any other steroid hormone. For at least current usage, I suggest that we use the term principal molting hormone for 20-hydroxyecdysone and the terms substrate for the molting hormone and/or hormone for ecdysone (although many Drosophila geneticists still use the term ecdysone for 20-hydroxecdysone; but that is another problem). The Ghent meeting was beautifully organized by Professor Smagghe who seemed to be around 24 h a day insuring that everything proceeded correctly and on schedule. He deserves our thanks for that, and also for this volume that he lab- ored over for many months. I personally believe that this was among the very best of the many ecdysone workshops I have attended and the book is certainly among the most comprehensive outcomes of any of the workshops. I was most honored to have been chosen to give the Karlson lecture on our work on the Halloween genes. I realize that this is not the usual preface in that I haven’t discussed any of the con- tributions, but they are so good that I would do them a disservice by summarizing each in a sentence – they deserve sustained reading and I believe that they will be cited for many years to come. As I approach my 80th birthday after more than half century in this field, I am very pleased to have my new work published here, that I had the chance to attend this very successful meeting in such a very beautiful city, and the opportunity to enjoy the amazing varieties of such fine beers offered to the ‘ecdysonists’ in Ghent. Chapel Hill, North Carolina, USA Lawrence I. Gilbert September 2008 Contents Preface ............................................................................................................. v Color Plates .................................................................................................... xi Part I Fundamental Aspects of Ecdysteroid Research: The Distribution and Diversity of Ecdysteroids in Animals and Plants 1 Phytoecdysteroids: Diversity, Biosynthesis and Distribution ....................................................................................... 3 Laurence Dinan, Juraj Harmatha, Vladimir Volodin, and René Lafont 2 Diversity of Ecdysteroids in Animal Species ......................................... 47 René Lafont and Jan Koolman 3 Crustacean Ecdysteroids and Their Receptors ..................................... 73 Penny M. Hopkins 4 Synthetic Ecdysteroidal Compounds ..................................................... 99 Josep Coll Toledano Part II In the Post-Genomic Era, Ecdysteroid Genetic Hierarchies in Insect Growth and Reproduction 5 Ecdysteroids and Ecdysteroid Signaling Pathways During Insect Oogenesis ........................................................ 127 Luc Swevers and Kostas Iatrou 6 Regulation of Sciarid DNA Puffs by Ecdysone: Mechanisms and Perspectives ................................................................. 165 Nadia Monesi, Juliana Aparecida Candido-Silva, Maria Luísa Paçó-Larson, and Jorge Cury de Almeida vii viii Contents 7 The Ecdysteroids’ Effects in the Control of Cell Proliferation and Differentiation ............................................. 185 David Siaussat, Patrick Porcheron, and Stépahne Debernard 8 Applications of RNA Interference in Ecdysone Research ................. 205 Garry N. Hannan, Ronald J. Hill, Skarlatos G. Dedos, Luc Swevers, Kostas Iatrou, Anjiang Tan, R. Parthasarathy, Hua Bai, Zhaolin Zhang, and Subba R. Palli Part III Role of Cross Talk of Genes and Growth Factors in Ecdysteroid Titers and Signalling 9 The Function and Evolution of the Halloween Genes: The Pathway to the Arthropod Molting Hormone ............................. 231 Lawrence I. Gilbert and Kim F. Rewitz 10 Recent Studies on Prothoracic Gland Cell Growth and Ecdysteroidogenesis in the Silkworm, Bombyx mori .................. 271 Shi-Hong Gu and Ju-Ling Lin 11 Diversity in Factors Regulating Ecdysteroidogenesis in Insects ....... 283 Sandrien Van de Velde, Liesbeth Badisco, Elisabeth Marchal, Jozef Vanden Broeck, and Guy Smagghe 12 20-Hydroxyecdysone, Juvenile Hormone and Biogenic Amines: Mechanisms of Interaction in Control of Drosophila Reproduction Under Normal and Stressful Conditions ..................... 317 Nataly Gruntenko and Inga Rauschenbach Part IV Ecdysteroids Function Through Nuclear and Membrane Receptors 13 The Structure and Function of Ecdysone Receptors ......................... 335 Isabelle M.L. Billas, Christopher Browning, Michael C. Lawrence, Lloyd D. Graham, Dino Moras, and Ronald J. Hill 14 The Multidimensional Partnership of EcR and USP ......................... 361 Vincent C. Henrich, Josh Beatty, Heike Ruff, Jenna Callender, Marco Grebe, and Margarethe Spindler-Barth 15 Functional Analysis of Ecdysteroid Receptor from Drosophila melanogaster “In Vitro” ............................................ 377 Anca Azoitei, Heike Ruff, Christian Tremmel, Sabine Braun, and Margarethe Spindler-Barth Contents ix 16 Intracellular Localization of the Ecdysteroid Receptor ..................... 389 Klaus-Dieter Spindler, Katarzyna Betan´ska, Claudia Nieva, Tomasz Gwóz´dz´, Joanna Dutko-Gwóz´dz´, Andrzej Oz˙yhar, and Margarethe Spindler-Barth 17 Genomic and Nongenomic Actions of 20-Hydroxyecdysone in Programmed Cell Death ................................................................... 411 Masatoshi Iga and Sho Sakurai 18 Rapid, Non-Genomic Responses to Ecdysteroids and Catecholamines Mediated by a Novel Drosophila G-Protein-Coupled Receptor............................................. 425 Peter D. Evans, D.P. Srivastava, and V. Reale Part V Ecdysteroids in Modern Agriculture, Medicine, Doping and Ecotoxicology 19 Ecdysone Receptors of Pest Insects – Molecular Cloning, Characterisation, and a Ligand Binding Domain-Based Fluorescence Polarization Screen ......................................................... 447 Lloyd D. Graham, Wynona M. Johnson, Donya Tohidi-Esfahani, Anna Pawlak-Skrzecz, Marianne Bliese, George O. Lovrecz, Louis Lu,