Table Of ContentEvolutionary Causation
Vienna Series in Theoretical Biology
Gerd B. Müller, editor- in- chief
Johannes Jäger, Thomas Pradeu, Katrin Schäfer, associate editors
The Evolution of Cognition, edited by Cecilia Heyes and Ludwig Huber, 2000
Origination of Organismal Form, edited by Gerd B. Müller and Stuart A. Newman, 2003
Environment, Development, and Evolution, edited by Brian K. Hall, Roy D. Pearson, and
Gerd B. Müller, 2004
Evolution of Communication Systems, edited by D. Kimbrough Oller and Ulrike Griebel, 2004
Modularity: Understanding the Development and Evolution of Natur al Complex Systems,
edited by Werner Callebaut and Diego Rasskin-G utman, 2005
Compositional Evolution: The Impact of Sex, Symbiosis, and Modularity on the Gradualist
Framework of Evolution, by Richard A. Watson, 2006
Biological Emergences: Evolution by Natur al Experiment, by Robert G. B. Reid, 2007
Modeling Biology: Structure, Beh av iors, Evolution, edited by Manfred D. Laubichler and
Gerd B. Müller, 2007
Evolution of Communicative Flexibility, edited by Kimbrough D. Oller and Ulrike Griebel,
2008
Functions in Biological and Artificial Worlds, edited by Ulrich Krohs and Peter Kroes, 2009
Cognitive Biology, edited by Luca Tommasi, Mary A. Peterson, and Lynn Nadel, 2009
Innovation in Cultural Systems, edited by Michael J. O’Brien and Stephen J. Shennan, 2010
The Major Transitions in Evolution Revisited, edited by Brett Calcott and Kim Sterelny, 2011
Transformations of Lamarckism, edited by Snait B. Gissis and Eva Jablonka, 2011
Convergent Evolution: Limited Forms Most Beautiful, by George McGhee, 2011
From Groups to Individuals, edited by Frédéric Bouchard and Philippe Huneman, 2013
Developing Scaffolds in Evolution, Culture, and Cognition, edited by Linnda R. Caporael,
James Griesemer, and William C. Wimsatt, 2014
Multicellularity: Origins and Evolution, edited by Karl J. Niklas and Stuart A. Newman, 2015
Vivarium: Experimental, Quantitative, and Theoretical Biology at Vienna’s Biologische
Versuchsanstalt, edited by Gerd B. Müller, 2017
Landscapes of Collectivity in the Life Sciences, edited by Snait B. Gissis, Ehud Lamm,
and Ayelet Shavit, 2017
Rethinking Human Evolution, edited by Jeffrey H. Schwartz, 2017
Evolutionary Causation
Biological and Philosophical Reflections
Edited by Tobias Uller and Kevin N. Laland
The MIT Press
Cambridge, Mas sa chu setts
London, England
© 2019 Mas sa chu setts Institute of Technology
All rights reserved. No part of this book may be reproduced in any form by any electronic or mechanical means
(including photocopying, recording, or information storage and retrieval) without permission in writing from the
publisher.
This book was set in Times Roman by Westchester Publishing Ser vices. Printed and bound in the United States
of Amer i ca.
Library of Congress Cataloging-in-Publication Data
Names: Uller, Tobias, editor. | Laland, Kevin N., editor.
Title: Evolutionary causation : biological and philosophical reflections /
edited by Tobias Uller and Kevin N. Laland.
Description: Cambridge, MA : The MIT Press, [2019] | Series: Vienna series in
theoretical biology | Includes bibliographical references and index.
Identifiers: LCCN 2018042611 | ISBN 9780262039925 (hardcover : alk. paper)
Subjects: LCSH: Evolution (Biology)—Philosophy. | Causation.
Classification: LCC QH360.5 .E9596 2019 | DDC 576.8—dc23
LC record available at https://lccn.loc.gov/2018042611
10 9 8 7 6 5 4 3 2 1
Contents
Series Foreword vii
1 Evolutionary Causation 1
Tobias Uller and Kevin N. Laland
2 Causality and the Role of Philosophy of Science 13
Massimo Pigliucci
3 Understanding Bias in the Introduction of Variation
as an Evolutionary Cause 29
Arlin Stoltzfus
4 The Shape of Things to Come: Evo Devo Perspectives on
Causes and Consequences in Evolution 63
Armin P. Moczek
5 Incorporating the Environmentally Sensitive Phenotype
into Evolutionary Thinking 81
David I. Dayan, Melissa A. Graham, John A. Baker, and Susan A. Foster
6 Genotype- Environment Interaction and the Unscripted Reaction Norm 109
Sonia E. Sultan
7 Understanding Niche Construction as an Evolutionary Pro cess 127
Kevin N. Laland, John Odling-S mee, and Marcus W. Feldman
8 Biological Dynamics and Evolutionary Causation 153
Renée A. Duckworth
9 The Causes of a Major Transition: How Social Insects Traverse
Darwinian Space 173
Heikki Helanterä and Tobias Uller
vi Contents
10 Are Developmental Plasticity, Niche Construction, and Extended
Inheritance Necessary for Evolution by Natu ral Se lection?
The Role of Active Phenotypes in the Minimal Criteria for
Darwinian Individuality 197
Richard A. Watson and Christoph Thies
11 The Paradox of Population Thinking: First Order C auses
and Higher Order Effects 227
Denis M. Walsh
12 Ontology, Causality, and Methodology of Evolutionary
Research Programs 247
Jun Otsuka
13 A Darwinian Dream: On Time, Levels, and Proc esses in Evolution 265
Arnaud Pocheville
14 Decoupling, Commingling, and the Evolutionary Significance
of Experiential Niche Construction 299
Lynn Chiu
15 Biological Information in Developmental and Evolutionary Systems 323
Karola Stotz
Contributors 345
Index 347
Series Foreword
Biology is a leading science in this c entury. As in all other sciences, prog ress in biology
depends on the interrelations between empirical research, theory building, modeling, and
societal context. But whereas molecular and experimental biology have evolved dramati-
cally in recent years, generating a flood of highly detailed data, the integration of t hese
results into useful theoretical frameworks has lagged behind. Driven largely by pragmatic
and technical considerations, research in biology continues to be less guided by theory
than it may seem.
By promoting the formulation and discussion of new theoretical concepts in the biosci-
ences, this series intends to help fill import ant gaps in our understanding of some of the
major open questions of biology, such as the origin and organi zation of organismal form,
the relationship between development and evolution, and the biological bases of cognition
and mind. Theoretical biology has import ant roots in the experimental tradition of early
twentieth-c entury Vienna. Paul Weiss and Ludwig von Bertalanffy were among the first
to use the term theoretical biology in its modern sense. In their understanding, the subject
was not limited to mathematical formalization, as is often the case today, but extended to
the conceptual foundations of biology. It is this commitment to a comprehensive and cross-
disciplinary integration of theoretical concepts that the Vienna Series intends to emphasize.
Today, theoretical biology has gen e tic, developmental, and evolutionary components, the
central connective themes in modern biology, but it also includes relevant aspects of
computational or systems biology and extends to the naturalistic philosophy of sciences.
The Vienna Series grew out of theory-o riented workshops or ga nized by the KLI, an
international institute for the advanced study of natur al complex systems. The KLI fosters
research proje cts, workshops, book proj ects, and the journal Biological Theory, all devoted
to aspects of theoretical biology, with an emphasis on—b ut not restriction to— integrating
the developmental, evolutionary, and cognitive sciences. The series editors welcome sug-
gestions for book proj ects in these domains.
Gerd B. Müller, Johannes Jäger, Thomas Pradeu, Katrin Schäfer
1
Evolutionary Causation
Tobias Uller and Kevin N. Laland
Introduction
Scientific inference typically relies on establishing causation. This is also the case in
evolutionary biology, a discipline charged with providing historical accounts of the proper-
ties of living beings, as well as an understanding of the proc esses that explain the origin
of those properties. Familiar phenomena that demand an evolutionary explanation include
the fit of form and function (adaptation) and the evolution of reproductive isolation
(speciation), but also many o thers, including the origin of novelty and the organi zation of
biological systems, including genomes and ecosystems. As expected from this diversity
of topics, the study of evolution is a quite loosely or ga nized endeavor. Nevertheless, there
is broad agreement that an explanation for why organisms have part ic u lar features com-
monly will involve a part ic u lar kind of evolutionary proc ess, namely evolution by natur al
se lection.
While the basic princip les of evolution by natur al se lection are s imple— variation among
individuals, differences in survival and reproduction, and heredity—t he biological reali ty
that instantiates these princip les is incredibly complex. Furthermore, evolutionary pro-
cesses encompass causation at dif fer ent levels of biological organi zation, from genes to
ecosystems, and at dif fer ent timescales. This complexity makes it necessary to find means
to represent biological systems in ways that leave out almost all causal detail, while ensur-
ing that models, observations, and experiments carry explanatory weight. What evolution-
ary researchers should incorporate into their explanatory accounts, and what they should
leave out, is far from a trivial issue.
While biologists are typically content to leave the nature of causation to phil os o phers,
there is scientific value in reflecting on core theoretical concepts. Dif fer ent views on
causation can lead to alternative explanatory frameworks, dictate what counts as legitimate
difference makers in evolutionary proc esses, and shape perspectives on key concepts such
as natur al sel ection and adaptation. How causation is understood thus shapes the structure
of evolutionary theory, with both historical and cont emporary debates in evolutionary
biology revolving around the nature of causation (Laland et al. 2011). Yet, t hese issues
