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T H E F R O N T I E R S C O L L E C T I O N Sonya Bahar T H E E S SE N T I A L T E N S ION Competition, Cooperation and Multilevel Selection in Evolution 123 THE FRONTIERS COLLECTION Series editors Avshalom C. Elitzur, Physics, Iyar Institute, Haifa, Israel Zeeya Merali, Foundational Questions Institute, Decatur, Georgia, USA T. Padmanabhan, for Astronomy and Astrophysics (IUC), Inter University Centre, Pune, India Maximilian Schlosshauer, Information Austrian Ac. of Science, Institute for Quantum Optics and Quantum Information, Portland, Oregon, USA Mark P. Silverman, Dept. Physics, Trinity College, Hartford, Connecticut, USA Jack A. Tuszynski, Department of Physics, CCIS 4-181, University of Alberta, Edmonton, Alberta, Canada Rüdiger Vaas, Redaktion Astronomie, Physik, bild der wissenschaft, Leinfelden-Echterdingen, Baden-Württemberg, Germany THE FRONTIERS COLLECTION Series Editors A.C. Elitzur Z. Merali T. Padmanabhan M. Schlosshauer M.P. Silverman J.A. Tuszynski R. Vaas The books in this collection are devoted to challenging and open problems at the forefront of modern science, including related philosophical debates. In contrast to typical research monographs, however, they strive to present their topics in a manner accessible also to scientifically literate non-specialists wishing to gain insight into the deeper implications and fascinating questions involved. Taken as a whole, the series reflects the need for a fundamental and interdisciplinary approach to modern science. Furthermore, it is intended to encourage active scientists in all areas to ponder over important and perhaps controversial issues beyond their own speciality. Extending from quantum physics and relativity to entropy, consciousness and complex systems—the Frontiers Collection will inspire readers to push back the frontiers of their own knowledge. More information about this series at http://www.springer.com/series/5342 For a full list of published titles, please see springer.com/series/5342 Sonya Bahar The Essential Tension Competition, Cooperation and Multilevel Selection in Evolution Sonya Bahar Department of Physics & Astronomy and Center for Neurodynamics University of Missouri at St Louis St Louis, MO, USA ISSN 1612-3018 ISSN 2197-6619 (electronic) The Frontiers Collection ISBN 978-94-024-1052-5 ISBN 978-94-024-1054-9 (eBook) DOI 10.1007/978-94-024-1054-9 Library of Congress Control Number: 2017949274 © Springer Science+Business Media B.V. 2018 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Printed on acid-free paper This Springer imprint is published by Springer Nature The registered company is Springer Science+Business Media B.V. The registered company address is: Van Godewijckstraat 30, 3311 GX Dordrecht, The Netherlands for Frank Moss 1934–2011 & for C & D Acknowledgements There are many people I could acknowledge, but I don’t want this to sound like the Oscars, so I will limit myself to the essential thanks that are due to the wonderful Maria Bellantone, for urging me to write this book and for her infinite patience in waiting for the manuscript to finally be completed. vii Contents Introduction ..................................................................................................... xi Part I The History of an Idea 1 Crowds ..................................................................................................... 3 References ................................................................................................. 12 2 Classification............................................................................................ 13 References ................................................................................................. 26 3 Time, Just Time: Integrating Up the Great Chain of Being ............... 29 References ................................................................................................. 50 4 The Battle of the Parts ............................................................................ 51 References ................................................................................................. 63 5 Synthesis?................................................................................................. 65 References ................................................................................................. 83 6 Selfish Creatures, Huddled Together for Warmth ............................... 85 References ................................................................................................. 97 7 The Vanishing Point Appears ................................................................. 99 References ................................................................................................. 122 Part II At the Frontier 8 Flocking, Swarming, and Communicating ........................................... 127 References ................................................................................................. 150 9 Biofilms .................................................................................................... 153 References ................................................................................................. 170 10 Multicellularity: Dictyostelium ............................................................... 175 References ................................................................................................. 202 ix x Contents 11 Multicellularity: Volvox .......................................................................... 207 References ................................................................................................. 231 12 Experimental Evolution .......................................................................... 235 References ................................................................................................. 266 Part III Beyond the Barricade 13 Cooperation and Competition: One Level Sitting on Another ........... 271 References ................................................................................................. 303 14 Evol = f(Evol) ........................................................................................... 309 References ................................................................................................. 330 15 Spandrels, Exaptations, and Raw Material .......................................... 333 References ................................................................................................. 358 16 The Essential Tension ............................................................................. 361 References ................................................................................................. 370 Index ................................................................................................................. 373 Introduction I…suggest that something like ‘convergent thinking’ is just as essential to scientific advance as is divergent. Since these two modes of thought are inevitably in conflict, it will follow that the ability to support a tension that can occasionally become unbearable is one of the prime requirements for the very best sort of scientific research. Thomas S. Kuhn Je juge cette longue querelle de la tradition et de l’invention, de l’Ordre et de l’Aventure. Guillaume Apollinaire I MUST begin with an apology to Thomas Kuhn. The title of this book has been borrowed – lifted, liberated, stolen, as you prefer – though my editor assures me that titles cannot be copyrighted and that I commit no actionable offense – from the title essay of a volume of Thomas Kuhn’s writings subtitled Selected Studies in Scientific Tradition and Change. The essay was initially delivered as a lecture at a 1959 University of Utah conference on, of all things, the “identification of scientific tal- ent.” Kuhn was already well known as a self-described “ex-physicist now working in the history of science,” though the publication of his masterpiece, The Structure of Scientific Revolutions, still lay 3 years in the future. Invited to speak at a confer- ence – run by educational psychologists – on the key characteristics of the creative personality and how to identify such a personality early in the educational process, Kuhn addressed a core issue in scientific education, in terms that clearly show his evolving thought on the themes of creativity, “normal science,” and scientific revo- lutions. Is iconoclasm, he asked, the only characteristic needed for a successful and creative scientist? Kuhn argued that a thorough grounding in tradition is as impor- tant to scientific advance as the ability to strike out into the unknown. He said: Normal research, even the best of it, is a highly convergent activity based firmly upon a settled consensus acquired from scientific education and reinforced by subsequent life in the profession. Typically, to be sure, this convergent or consensus-bound research ulti- xi

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