Bioluminescence L I V I N G L I G H TS , L I G H TS F O R L I V I N G Thérèse Wilson J. Woodland Hastings B I O L U M I N E S C E N C E B I O L U M I N E S C E N C E Living Lights, Lights for Living Thérèse Wilson J. Woodland Hastings Harvard University Press Cambridge, Massachusetts London, En gland 2013 Copyright © 2013 by the President and Fellows of Harvard College All rights reserved Printed in Canada Library of Congress Cataloging-i n-P ublication Data Wilson, Therese, 1925– Bioluminescence : living lights, lights for living / Therese Wilson, J. Woodland Hastings. p. cm. Includes index. ISBN 978- 0- 674- 06716- 5 (alk. paper) 1. Bioluminescence. I. Hastings, J. Woodland ( John Woodland), 1927–. II. Title. QH641.W55 2013 572'.4358—dc23 2012022227 CONTENTS Ac know ledgm ents vii Introduction 1 PART I : FIVE DIFFERENT BIOLUMINESCENCE SYSTEMS 7 1. A Marine Crustacean 9 Bioluminescent f shes as plagiarists and thieves 2. J ellyf sh and Green Fluorescent Protein 2 1 A soft coral, a calcium-s ensitive protein, and f sh with related b ioluminescence systems 3. F iref ies and Other Beetles 3 1 Luciferase- dependent bioluminescence color and rhythmic displays 4. Dinof agellates and Krill 4 5 The sparkling clocks of the oceans and bioluminescent shrimp 5. B acteria 6 1 Bacterial “communication,” symbioses, and milky seas PART II : D IVERSITY, FUNCTIONS, AND EVOLUTIONARY ORIGINS OF BIOLUMINESCENCE 77 6. S hort Accounts of Other Luminous Organisms 7 9 Having different and not well- characterized biochemistries 7. B ioluminescence in the Oceans 1 05 Anglerf sh, dragonf sh, and a Lake Baikal parenthesis 8. T he Many Functions of Bioluminescence 1 19 Defense, offense, communication, and propagation 9. T he Origins and Evolution of Bioluminescence 1 25 How did luciferases originate? PART III : BOOKENDS 133 10. A pplications 1 35 Tools for biology, medicine, and public health 11. H ow Does Life Make Light? 1 47 “Excited molecules” and bioluminescence vi C O N T E N T S Glossary 1 63 Further Reading 1 69 Illustration Credits 1 77 Index 181 ACK NOWL EDGM ENTS It is our pleas ure to acknowledge our debt to the many who have been critical to the birth of this book. Foremost among those are the students, postdoctoral fellows, and visitors to the Hastings and Wilson lab, who, over the years, created a unique and unforgettable atmosphere of free exchange of ideas, and joy in exploring all sorts of topics related to the emission of light by living (and non- living) things. Our debt to all of them cannot be expressed adequately. We are especially grateful to Jim Morin, who read the entire manu- script and pointed out its weak points, and to Anne Goldizen for her editorial suggestions throughout the manuscript. Many friends did us the favor of reading chapters or parts of chapters. Their comments and suggestions w ere universally appreciated. Among those to whom we are especially grateful are Etelvino Bechara, Arthur Halpern, Andy Knoll, Neil Krieger, Ken Nealson, John Paxton, Deb Robert- son, Vadim Viviani, and Edith Widder. Thérèse Wilson is espe- cially grateful to Meredithe Applebury for her advice and encour- agement at the very early stages of the work. Without illustrations, a book like this would not see the light of day. We thank all those who have given us permission to use photos and illustrations, especially Milton Cormier, Larry Fritz, Helen Ghir- adella, Karsten Hartel, Peter Herring, and Christopher Kenaley. Thanks also to Marvin Morales for the chemical formulae. For all other f gures, we were exceedingly lucky to be able to rely on Renate Hellmiss, whose artistry, competence, and patience made working with her a pleas ure at all stages of the work. Finally, we are grateful for f nancial help from the Faculty of Arts and Sciences and the Department of Molecular and Cellular Biology of Harvard University.