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History, Philosophy and Theory of the Life Sciences Marta Bertolaso Silvia Caianiello Emanuele Serrelli E ditors Biological Robustness Emerging Perspectives from within the Life Sciences History, Philosophy and Theory of the Life Sciences Volume 23 Editors Charles T. Wolfe, Ghent University, Belgium Philippe Huneman, IHPST (CNRS/Université Paris I Panthéon-Sorbonne), France Thomas A. C. Reydon, Leibniz Universität Hannover, Germany Editorial Board Marshall Abrams, University of Alabama at Birmingham Andre Ariew Missouri Minus van Baalen UPMC, Paris Domenico Bertoloni Meli Indiana Richard Burian Virginia Tech Pietro Corsi EHESS, Paris François Duchesneau Université de Montréal John Dupré Exeter Paul Farber Oregon State Lisa Gannett Saint Mary’s University, Halifax Andy Gardner Oxford Paul Griffiths Sydney Jean Gayon IHPST Paris Guido Giglioni Warburg Institute, London Thomas Heams INRA, AgroParisTech, Paris James Lennox Pittsburgh Annick Lesne CNRS, UPMC, Paris Tim Lewens Cambridge Edouard Machery Pittsburgh Alexandre Métraux Archives Poincaré, Nancy Hans Metz Leiden Roberta Millstein Davis Staffan Müller-Wille Exeter Dominic Murphy Sydney François Munoz Université Montpellier 2 Stuart Newman New York Medical College Frederik Nijhout Duke Samir Okasha Bristol Susan Oyama CUNY Kevin Padian Berkeley David Queller Washington University, St Louis Stéphane Schmitt SPHERE, CNRS, Paris Phillip Sloan Notre Dame Jacqueline Sullivan Western University, London, ON Giuseppe Testa IFOM-IEA, Milano J. Scott Turner Syracuse Denis Walsh Toronto Marcel Weber Geneva More information about this series at http://www.springer.com/series/8916 Marta Bertolaso • Silvia Caianiello Emanuele Serrelli Editors Biological Robustness Emerging Perspectives from within the Life Sciences Editors Marta Bertolaso Silvia Caianiello FAST Institute of Philosophy of Scientific Institute for the History of Philosophy and Practice and Faculty of Engineering Science in Modern Age (ISPF) University Campus Bio-Medico of Rome Italian National Research Council Rome, Italy Naples, Italy Emanuele Serrelli CISEPS - Center for Interdisciplinary Studies in Economics, Psychology and Social Sciences University of Milano Bicocca CISEPS Brescia, Italy ISSN 2211-1948 ISSN 2211-1956 (electronic) History, Philosophy and Theory of the Life Sciences ISBN 978-3-030-01197-0 ISBN 978-3-030-01198-7 (eBook) https://doi.org/10.1007/978-3-030-01198-7 Library of Congress Control Number: 2018962881 © Springer Nature Switzerland AG 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. This Springer imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland Foreword The papers collected in this volume are the outcome of a series of workshops orga- nized by the Bio-Techno-Practice group on the different ways in which philoso- phers, biologists, neuroscientists, and engineers employ the concept of “robustness.” The goal, which was successfully realized, was to stimulate an interactive interdis- ciplinary engagement that would highlight differences and commonalities across disciplines and perspectives. The hope was that, by this type of engagement, confu- sions would be evaporated and insights from one part of the intellectual landscape could aid those exploring robustness in another part. This volume is evidence of the success of that methodology, as the individual papers reveal the benefits of interac- tive engagement. Even more important is the reframing of future work by the par- ticipants in light of the refraction of individual disciplinary commitments in the context of cross-disciplinary connections. Robustness is a perfect subject for this type of engagement. It marks the system level property of maintaining system function in response to internal and external perturbations. We see it in evolved systems like organisms in the way body tempera- tures are maintained, or neurons are reassigned after head trauma. We see it in engi- neered systems like bridges and buildings or software algorithms whose design aims to preserve performance under a range of expected conditions. But how robust- ness is achieved varies both within and between types of systems. There is clearly intellectual traffic between disciplines studying robustness. Systems biology employs notions of control networks, feedback, and modularity and reverse engi- neering. Top down and bottom up approaches converge on how a complex system not only does what it does but continues to do it when there are changes in the external environment and loss or change in internal components. Philosophy of science can abstract away from the details of any one mechanism for achieving robustness to characterize what it is for a system to be robust. Robustness is always relative, robust with respect to this function, or that equilib- rium. Indeed, by evolving or engineering robustness for a particular function in a range of values for internal and external variables, fragility will be introduced for other functional stabilities in other conditions. Redundancy, modularity, and multi- ple pathways are features that promote robustness. v vi Foreword Biological Robustness: Emerging Perspectives from Within the Life Sciences combines the detailed explorations of robustness by engineers, biologists, neurosci- entists, and philosophers, inviting the reader into reflective engagement that the workshops promoted. I have learned much from participating in the project that gave rise to this volume. By bringing together a plurality of perspectives, this vol- ume extends the reach of interdisciplinary engagement. Distinguished Professor Sandra D. Mitchell Department of History and Philosophy of Science University of Pittsburgh Pittsburgh, PA, USA Contents 1 Introduction: Issues About Robustness in the Practice of Biological Sciences ............................................................................. 1 Marta Bertolaso, Emanuele Serrelli, and Silvia Caianiello 1.1 Biological Robustness ..................................................................... 4 1.2 The Book ......................................................................................... 7 1.3 Emerging Epistemological Perspectives from Within the Life Sciences ........................................................ 14 References ................................................................................................ 17 2 Prolegomena to a History of Robustness ............................................. 23 Silvia Caianiello 2.1 Origin of the Modern Meaning ....................................................... 25 2.2 Robustness and Control Theory ...................................................... 28 2.3 Early Inceptions of Robustness in Biology: Organizing Vs Design Principles .................................................... 37 2.4 Robustness and Complexity ............................................................ 43 References ................................................................................................ 48 3 Robustness, Mechanism, and the Counterfactual Attribution of Goals in Biology ............................................................. 55 Marco Buzzoni 3.1 Introduction ..................................................................................... 55 3.2 Robustness and Intersubjective Reproducibility ............................. 58 3.3 Robustness and the Counterfactual Attribution of Goals in Biology ......................................................................... 63 3.4 Conclusion ...................................................................................... 71 References ................................................................................................ 72 vii viii Contents 4 Multiple Realization and Robustness ................................................... 75 Worth (Trey) Boone 4.1 Introduction ..................................................................................... 75 4.2 Multiple Realization and Causal Explanation ................................. 76 4.3 Multiple Realization as Distributed Functional Robustness ........... 80 4.4 Kinds Reconsidered ........................................................................ 87 4.5 Conclusion ...................................................................................... 92 References ................................................................................................ 92 5 Robustness: The Explanatory Picture .................................................. 95 Philippe Huneman 5.1 Introduction ..................................................................................... 96 5.2 Characterizing and Situating Robustness ........................................ 97 5.3 Three Families of Explanations of Robustness ............................... 100 5.4 Robustness as Explanandum in Evolutionary Biology, and the Explanatory Reversibility Proper to Evolutionary Biology ...................................................... 107 5.5 Robustness and Other Reversible Explananda of Evolutionary Biology.................................................................. 113 5.6 Conclusion ...................................................................................... 117 References ................................................................................................ 118 6 Robustness and Autonomy in Biological Systems: How Regulatory Mechanisms Enable Functional Integration, Complexity and Minimal Cognition Through the Action of Second-Order Control Constraints .................................................. 123 Leonardo Bich 6.1 Introduction ..................................................................................... 124 6.2 Basic Concepts: Stability, Control and Signal in Autonomous Systems .................................................................. 126 6.3 Biological Regulation ..................................................................... 132 6.4 Regulation at the Crossroads Between Identity, Complexity, and Cognition ............................................................. 136 6.5 Final Remarks ................................................................................. 142 References ................................................................................................ 143 7 Robustness and Emergent Dynamics in Noisy Biological Systems .................................................................................. 149 Christian Cherubini, Simonetta Filippi, and Alessandro Loppini 7.1 Introduction: Robustness and Stability in Physics and Biology .................................................................... 149 7.2 Robustness: The Point of View of Biophysics ................................ 152 7.3 Modeling Robustness in Pancreatic β-Cells Populations ................ 152 7.4 Conclusion ...................................................................................... 159 References ................................................................................................ 161 Contents ix 8 The Robustness/Sensitivity Paradox: An Essay on the Importance of Phase Separation .............................. 163 Alessandro Giuliani 8.1 Introduction ..................................................................................... 163 8.2 Biological Networks ....................................................................... 164 8.3 Conclusion ...................................................................................... 171 References ................................................................................................ 172 9 Can Engineering Principles Help Us Understand Nervous System Robustness? ................................................................ 175 Timothy O’Leary 9.1 Feedback Control ............................................................................ 176 9.2 Feedback Control in Nervous Systems ........................................... 178 9.3 Robust Architectures: Degeneracy .................................................. 182 9.4 Conclusion ...................................................................................... 185 References ................................................................................................ 185 10 Robustness vs. Control in Distributed Systems ................................... 189 Marta Menci and Gabriele Oliva 10.1 Introduction ..................................................................................... 190 10.2 Control Theory Overview ............................................................... 191 10.3 Open-Loop vs. Closed-Loop Control ............................................. 192 10.4 Dynamical Distributed Systems ...................................................... 194 10.5 Robustness and Control .................................................................. 195 10.6 Control and Robustness in Distributed Systems ............................. 199 10.7 Conclusions ..................................................................................... 203 References ................................................................................................ 204 11 The Robustness of Musical Language: A Perspective from Complex Systems Theory .................................... 207 Flavio Keller and Nicola Di Stefano 11.1 Introduction ..................................................................................... 207 11.2 Stability and Fragility in Auditory Perception ................................ 209 11.3 Fragility and Emotional Resonance of Musical Language ............. 212 11.4 Conclusion ...................................................................................... 214 References ................................................................................................ 215 12 Dynamical Rearrangement of Symmetry and Robustness in Physics and Biology ................................................ 219 Giuseppe Vitiello 12.1 Introduction ..................................................................................... 219 12.2 A Two Level Description: Heisenberg Fields and Physical Fields ......................................................................... 220 12.3 Spontaneous Breakdown of Symmetry and Dynamical Rearrangement of Symmetry ................................. 221 12.4 Boson Condensation, Ordered Patterns and Low Energy Theorem ............................................................... 222

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