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Evolutionary Biology
New Perspectives on Its Development 6
Thomas E. Dickins
Benjamin J. A. Dickins Editors
Evolutionary
Biology:
Contemporary
and Historical
Reflections Upon
Core Theory
Evolutionary Biology – New Perspectives
on Its Development
Volume 6
Series Editor
Richard G. Delisle, Department of Philosophy and School of Liberal Education,
University of Lethbridge, Lethbridge, AB, Canada
All chapters are systematically reviewed by the series editor and respective volume
editor(s). For monographs, the editor of the book series reach out to two competent
reviewers. The editor ensures that reviews are fair and relevant. For edited volumes,
the volume’s editor selects two competent reviewers for each chapter, ensuring that
reviews are fair and relevant. The series editor oversees the whole evaluation
process.
***
Thomas E. Dickins • Benjamin J. A. Dickins
Editors
Evolutionary Biology:
Contemporary
and Historical Reflections
Upon Core Theory
Editors
Thomas E. Dickins Benjamin J. A. Dickins
Faculty of Science and Technology School of Science and Technology
Middlesex University Nottingham Trent University
London, UK Nottingham, UK
Centre for Philosophy of Natural
and Social Science
London School of Economics
London, UK
# The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland
AG 2023
Chapters 8, 20, 22 and 33 are licensed under the terms of the Creative Commons Attribution 4.0
International License (http://creativecommons.org/licenses/by/4.0/). For further details see licence infor-
mation in the chapters.
This work is subject to copyright. All rights are solely and exclusively licensed 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
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The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
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To Anne and David for their endless parental
investment.
To Nicola, Jack, and Isabella for continued
kinship.
Preface
vii
viii Preface
intellectual landscape and our hope is that this volume presents a balanced contribu-
tion that will prove useful to those interested by or engaged in current debates in
evolutionary biology.
We must first thank all the contributing authors for their dedication to the task of
writing this book. Not only did we ask them to write their own chapters, but also to
produce commentary on each other and reply to those who commented upon their
work. Richard Webb, Andy Wells, and Paul Taylor read many draft chapters and
gave useful commentary that helped to shape the book. These three, together with
Edgar Porcher and Max Steuer, are also members of the Behavior Lite discussion
group at the London School of Economics, where many matters pertaining to the
application of evolutionary theory are discussed. Those discussions helped to shape
and sharpen the thinking of both editors. Richard Delisle, the series editor for
Springer, has been tremendously supportive throughout the project, as have the
Springer team. BD thanks Axel Barlow for his input during chapter review. TD
would like to thank Oliver Curry, Daniel Nettle, Qazi Rahman, and Thom Scott
Phillips for direct as well as incidental input during various discussions. Most
importantly, TD will be forever grateful to Nicola, Jack, and Isabella for their
support and love, allowing him the time to hide away and work on this project.
Finally, both editors are grateful to Anne and David Dickins for their parental and
evolutionary inputs over the course of their life histories.
ix
Contents
1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Thomas E. Dickins and Benjamin J. A. Dickins
Part I
2 Every Evolutionist Their Own Historian: The Importance of
History, Context, and the Extended Evolutionary Synthesis . . . . . . . 25
V. Betty Smocovitis
3 Yes Indeed, Evolutionary Biologists Should Pay More
Attention to History: A Commentary on Smocovitis . . . . . . . . . . . . 55
Erik I. Svensson
4 History, Evolution, and the “Rashomon Effect”:
A Reply to Svensson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
V. Betty Smocovitis
Part II
5 The Creativity of Natural Selection and the Creativity
of Organisms: Their Roles in Traditional Evolutionary
Theory and Some Proposed Extensions . . . . . . . . . . . . . . . . . . . . . . 65
John J. Welch
6 Let there Be Light: A Commentary on Welch . . . . . . . . . . . . . . . . . 109
David Haig
7 Creative Destruction: A Reply to Haig . . . . . . . . . . . . . . . . . . . . . . 115
John J. Welch
Part III
8 The Organism in Evolutionary Explanation: From Early
Twentieth Century to the Extended Evolutionary Synthesis . . . . . . 121
Jan Baedke and Alejandro Fábregas-Tejeda
xi
xii Contents
Part IV
11 The Structure of Evolutionary Theory: Beyond Neo-Darwinism,
Neo-Lamarckism and Biased Historical Narratives About the
Modern Synthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Erik I. Svensson
12 It Is the Endless Forms, Stupid: A Commentary on Svensson . . . . . 219
David M. Shuker
13 Ecology, Agents, and the Causes of Selection:
A Reply to Shuker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
Erik I. Svensson
Part V
14 Hypertextuality of a Hyperextended Synthesis: On the
Interpretation of Theories by Means of Selective Quotation . . . . . . 231
David Haig
15 Teleology, Organisms, and Genes: A Commentary on Haig . . . . . . 249
Alejandro Fábregas-Tejeda and Jan Baedke
16 A Token Response: A Reply to Fábregas-Tejeda and Baedke . . . . . 265
David Haig
Part VI
17 The Darwinian Core of Evolutionary Theory and the Extended
Evolutionary Synthesis: Similarities and Differences . . . . . . . . . . . . 271
T. N. C. Vidya, Sutirth Dey, N. G. Prasad, and Amitabh Joshi
18 Evolution Is Bigger than All of Us: A Commentary on Vidya,
Dey, Prasad, and Joshi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 329
Vassiliki Betty Smocovitis
19 Why Evolution Is Bigger than all of Us: A Reply to Smocovitis . . . . 335
T. N. C. Vidya, Sutirth Dey, N. G. Prasad, and Amitabh Joshi
Part VII
20 Inclusive Fitness: A Scientific Revolution . . . . . . . . . . . . . . . . . . . . 343
António M. M. Rodrigues and Andy Gardner
Contents xiii
Part VIII
23 Evolution of Bacteriophage Latent Period Length . . . . . . . . . . . . . . 375
Stephen T. Abedon
24 Optimality and Idealisation in Models of Bacteriophage Evolution:
A Commentary on Abedon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427
Benjamin J. A. Dickins
25 On r-K Selection in the Evolution of Bacteriophages: A Reply to
Dickins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 433
Stephen T. Abedon
Part IX
26 Plasticity and Information . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 441
Thomas E. Dickins
27 Phenotypic Plasticity and Evolutionary Syntheses:
A Commentary on Dickins, T.E. . . . . . . . . . . . . . . . . . . . . . . . . . . . 461
Douglas J. Futuyma
28 On Rhetoric and Conceptual Frames: A Reply to Futuyma . . . . . . 467
Thomas E. Dickins
Part X
29 The Curious Incident of the Wasp in the Fig Fruit:
Sex Allocation and the Extended Evolutionary Synthesis . . . . . . . . 473
David M. Shuker
30 The Nuances of Biological Syntheses:
A Commentary on Shuker . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 505
Mitchell Ryan Distin
31 On Genetics, Ecology, and the Role of Philosophy in Evolutionary
Biology: A Reply to Distin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 511
David M. Shuker
Part XI
32 The Evolving Evolutionary Synthesis . . . . . . . . . . . . . . . . . . . . . . . 517
Douglas J. Futuyma
xiv Contents
Part XII
35 Genes and Organisms in the Legacy of the Modern Synthesis . . . . . 555
J. Arvid Ågren
36 The Parallax View: A Commentary on Ågren . . . . . . . . . . . . . . . . . 569
John J. Welch
37 Why We Disagree About Selfish Genes: A Reply to Welch . . . . . . . 581
J. Arvid Ågren
Part XIII
38 Genetic Evolvability: Using a Restricted Pluralism to
Tidy up the Evolvability Concept . . . . . . . . . . . . . . . . . . . . . . . . . . 587
Mitchell Ryan Distin
39 Pluralism and Progress in Evolutionary Biology:
A Commentary on Distin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 611
J. Arvid Ågren
40 Genetic Evolvability: A Reply to Ågren . . . . . . . . . . . . . . . . . . . . . . 617
Mitchell Ryan Distin
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 621
Contributors
xv
xvi Contributors
Abstract
In this chapter, we make some general comments about the nature of scientific
explanation and use them to contextualize recent debates within evolutionary
biology about the adequacy of what is sometimes termed standard evolutionary
theory. These comments serve to introduce the aims of the book and we then
summarize the chapters to follow, relating them to the opening themes.
Keywords
Modern synthesis · Extended evolutionary synthesis · Idealization · Abstraction
1.1 Introduction
This book is about the debate between advocates of what has been called an
Extended Evolutionary Synthesis and the defenders of the Modern Synthesis,
sometimes also referred to as the standard theory of evolution. We will come to
discuss the broad parameters of this debate shortly, but first, we wish to develop a
context within which to place this debate.
The history of evolutionary theory is one of argument. Gradualist biometricians
were locked in a dispute with saltationists about Darwinian gradualism in the late
T. E. Dickins (*)
Faculty of Science & Technology, Middlesex University, London, UK
Centre for Philosophy of Social and Natural Science, London School of Economics, London, UK
e-mail: [email protected]
B. J. A. Dickins
Department of Biosciences, Nottingham Trent University, Nottingham, UK
e-mail: [email protected]
(F)aced with the need to grapple with this complex world, scientists face cognitive, and
other, limitations. These limitations make it difficult to secure causal knowledge, to make
accurate predictions, and to pursue science’s other aims in this causally complex world of
ours. Or, perhaps better, this point can be phrased positively: simple patterns are cognitively
valuable. Simple patterns support human influence on and understanding of our world. There
is thus a basic mismatch between the cognitive value of simple patterns and the world’s
complexity... So, in the face of this mismatch, we often resort to lying a little bit: we
artificially simplify the parts of accounts that we are not interested in to improve our access
in a variety of ways to the parts we are interested in. This is one service that idealizations
provide. (Potochnik 2020: 934–935)
Causal patterns are patterns insofar as they are regularities that are limited in scope and that
may permit exceptions. The ideal gas law characterizes the approximate behavior of most
gases, although its predicted relationships break down at low temperatures and at high
pressures. It also ignores molecular size and intermolecular forces. Recall the idealization
of an ideal gas composed of noninteracting point particles; this idealization achieves that
neglect. Accordingly, even within its scope of application, the ideal gas law has
exceptions. . . (T)o represent a causal pattern is to show how changes to a system would,
over some range of circumstances, precipitate changes in other feature(s) of the system. The
ideal gas law shows, for example, how temperature increasing in a sealed container of gas
with a fixed volume increases the pressure. Mastery of causal patterns is exactly the kind of
thing that beings who prize simplicity need in order to operate in and grapple with a causally
complex world like ours. (Potochnik 2020: 935)
4 T. E. Dickins and B. J. A. Dickins
Suppose the temperature increase in a sealed container of fixed volume was in fact a can of
aerosol hair spray left in a car on a hot day. This phenomenon embodies the pattern described
in the ideal gas law. It also embodies the pattern of the greenhouse effect: the short
wavelengths of visible light can enter through the glass of the closed windows, but the
longer wavelengths of infrared light radiated by the objects in the car that absorbed the light
cannot exit through the glass as easily. These patterns relate to different aspects of the
phenomenon, and which is of interest depends on which aspects we are focused on. As these
simple examples show, different patterns embodied by some phenomenon may be closely
related to one another or wholly unrelated (or anywhere in between). (Potochnik 2020: 936)
Potochnik’s view, that there are many different projects within science each of which
may require its own idealizations, its own methods of rendering intelligibility, has
led her to develop a view on disagreements in science (Potochnik 2013). She notes
that some disagreements in biology have been or are treated as ideological debates.
Among her examples is the clash between the anti-adaptationists (Gould and
Lewontin 1979) and those defending adaptationism as an optimization approach.
She notes how Gould and Lewontin’s original paper invoked religious language to
characterize adaptationists, clearly labeling this approach as an ideology. Further-
more, Potochnik points to defenders of optimization who openly reference it as a
world view or leap of faith.
Either you subscribe to the Optimization Research Program as your worldview, or you reject
it... These positions are presented as ideological in the sense that they involve adherence to a
systematic set of ideas, a comprehensive way of looking at things. The set of ideas in
question is viewed as fundamental to the domain under investigation, and adherence to one
side or the other is taken to be a total commitment. This ideological tenor therefore suggests
that there is a rift in theory, that there is dispute regarding the basic understanding of these
types of phenomena. (Potochnik 2013: 119)
Potochnik argues that we can fruitfully move away from much ideological
grandstanding by taking what she refers to as a methodological approach. By this,
1 Introduction 5
she means that we ought to characterize science as model based, and those models as
idealizations in the manner discussed above. To this end, models adopted by groups
of scientists will contain deliberate falsehoods to facilitate intelligibility under
specific task demands. Where interests, or focal phenomena diverge, we perhaps
should expect to find different idealizations in play, and it is very easy to present
differences in idealization as distinctions in ideology. One reason for this, Potochnik
claims, is that scientists often commit to “simple causal processes with broad
domains of application” (Potochnik 2013: 121). Yet, we should really agree that
phenomena are the result of multiple, complex causal pathways and that a focus on a
particular route through such a tangle is a commitment to producing an intelligible
explanation. In this sense, a causal model is privileged above others, but that
privilege should only be seen as a pragmatic expedient, not an ontological commit-
ment. It is therefore to be seen as methodological because it is a method for
providing a workable explanation within a specific domain of enquiry.
In 2014 Nature carried a debate between two groups of scholars (Laland et al. 2014).
The question under discussion was “does evolutionary theory need a rethink?” One
group argued that yes it did, and urgently. The second that no it did not, and that all
was well.
The advocates made a core claim that mainstream evolutionary theory focused
almost entirely on gene-level explanations, a criticism aligned with the term gene-
centrism which captures the concept of privileging the gene in evolutionary
explanations. Their counter was that developmental processes should be recognized
as contributory factors in evolution. This idea is at the heart of the Extended
Evolutionary Synthesis (EES) movement and was most clearly expressed by
Pigliucci when he called for the unification of theories of genes with those of form
(Pigliucci 2007). In this way, the extended synthesis effectively promotes the
introduction of mechanistic theories of form into evolutionary theory.
While the above opening statement references a specific scientific idea, that devel-
opmental processes should be considered within evolutionary theory, the language
deployed is distinctly ideological (Potochnik 2013). Some of those opposed to the
EES are allocated emotional responses, those advocating for the EES are battling for
the fundamental essence of the discipline. By implication there can only be one
winner, there is no room for coexistence.
The advocates for a rethink establish a distinction between the Modern Synthesis,
which they see as the period in which population genetics emerged, and Standard
Evolutionary Theory (SET). It is not entirely clear what the status of SET is, but the
authors claim that it incorporates much of the Modern Synthesis. Thus, SET sees
new random variation established by genetic mutation, and natural selection as the
sole source of adaptation. In making these claims the advocates side neatly with
Gould’s view of the hardening of the Modern Synthesis into a panadaptationist,
gene-level theory (Gould 2002; Dickins 2021).
In our view, this “gene-centric” focus fails to capture the full gamut of processes that direct
evolution. Missing pieces include how physical development influences the generation of
variation (developmental bias); how the environment directly shapes organisms’ traits
(plasticity); how organisms modify environments (niche construction); and how organisms
transmit more than genes across generations (extra-genetic inheritance). For SET, these
phenomena are just outcomes of evolution. For the EES, they are also causes. (Laland et al.
2014, 162)
Here we see an explicit claim about the introduction of other causes beyond mutation
and selection. By implication, the idea is that those pursuing SET and those pursuing
the EES have the same broad target in mind—evolutionary explanation—but the
EES is seeking to proliferate causes under a development conception of evolution in
which those processes, while possibly the outcome of evolution can also affect
subsequent evolution. Put another way, there is an implicit assumption that both
groups of scholars share the same explanatory task. The EES claim is that SET is
causally inadequate to that task because they miss detail. Thus, causal explanation is
the focus in this debate.
The advocates move to an example:
(C)ichlid fishes in Lake Malawi are more closely related to other cichlids in Lake Malawi
than to those in Lake Tanganyika, but species in both lakes have strikingly similar body
shapes. In each case, some fish have large fleshy lips, others protruding foreheads, and still
others short, robust lower jaws.
SET explains such parallels as convergent evolution: similar environmental conditions
select for random genetic variation with equivalent results. This account requires extraordi-
nary coincidence to explain the multiple parallel forms that evolved independently in each
lake. A more succinct hypothesis is that developmental bias and natural selection work
together. Rather than selection being free to traverse across any physical possibility, it is
1 Introduction 7
guided along specific routes opened up by the processes of development. (Laland et al.
2014: 162)
In this quotation we see a particular causal strategy in play. The authors claim that a
SET account of morphological similarity across cichlid species challenges credulity,
as it appears to rely on multiple instances of the same mutation occurring and being
selected. Introducing developmental path dependency as a further cause is argued to
reduce search space for natural selection and render the model credible. Crucially the
EES claim is not that there is no role for selection, but rather that selection is assisted
by a narrowing of the parameters over which it must search. Theories of this kind
often rely on modular models of development (Brakefield 2006, 2011; Kirschner and
Gerhart 2010; Newman 2010) which reduce lethality effects associated with muta-
tion by reducing the number of genetic mutations required to enact a change in
morphology and enhance evolvability. Where selection does operate, in these
models, is over regulatory genes that might change where in time a developmental
module is activated, or might enhance the outcome of developmental modular
processes etc. (Dickins 2021).
The cichlid example is instructive. The first thing we might note, in keeping with
Potochnik’s clear view that our causal world is complex (Potochnik 2020), is that it
is very unlikely that either the SET or the EES approach will tell us the whole story
about the evolution of cichlid morphology and its ecological distribution. We should
accept that both accounts are idealizing. For example, a SET account might adopt the
idealization of single locus selection to mount an optimization model of cichlid
adaptations. In doing this no account of development processes would be made, but
development would be assumed. A part of that assumption would be that to all
intents and purposes the variation resulting from development was insignificant to
the adaptationist generalizations sought. Meanwhile, the EES account would high-
light developmental variation and make a case for the mechanisms of development
enabling more effective selection. But that selection may be modeled in single locus
terms again, with a focus upon regulatory genes. Moreover, the precise mechanistic
account of development is unlikely to capture all developmental causes, and further
idealizations will be introduced at some point. For example, the idea of develop-
mental modules is most likely an idealization designed to neatly capture some
dependencies in development. How encapsulated and domain specific such
dependencies are becomes a matter of empirical interest in each and every case.
The intuition we are seeking to prime is that whilst both the SET and EES
approaches, in this case, deploy idealizations it is not clear that the idealizations
are in contradiction to one another. Indeed, we might claim that the SET approach
simply assumes development, while the EES incorporates a version of it, and in that
way SET is more abstracted than EES, where abstraction is a process of reducing
detail to gain generality (Levy 2021). Moreover, the less abstracted EES account is
rendered this way simply because its project is different from that of SET. We might
say that the EES project is to reduce abstraction in evolutionary biology by
introducing proximate developmental mechanisms to show their effect upon evolu-
tionary dynamics and trajectories.
8 T. E. Dickins and B. J. A. Dickins
Table 1.1 Brown’s (2022) three types of adaptationism and structuralism summarized. In both
positions, the commitment becomes softer from empirical to methodological versions. The former
contains an ontological claim while the latter amounts to a set of guiding principles to do science
and generate findings
Adaptationism Structuralism
Empirical Selection is powerful, has causal Developmental constraint has causal
primacy, and can be used to explain primacy and can be used to explain
and predict evolutionary outcomes and predict evolutionary outcomes
Explanatory Selection has unique explanatory Developmental constraint can answer
importance as it can address the important questions of diversity,
apparent design disparity, and complexity
Methodological Adaptation is a good initial Scientists should look at disparity
hypothesis for scientists enabling and diversity and developmental
subsequent work constraint is a good initial hypothesis
A sticking point for this intuition is that the advocates have suggested that the
SET account of cichlids requires an extraordinary coincidence. In doing this they are
suggesting it is a false account. Initially, this might appear devastating to our
argument about abstractness. If the SET account is wrong, then its abstractions are
false and unrelated to any true account. However, the EES invocation of develop-
ment to reduce the search space for selection does not in fact do away with this
problem. It is still present in that account also. In the version we have presented a
mutation (presumably in a regulatory gene) is posited, and it will be selected because
it is attached to a highly conserved developmental program that is well insulated
against internal disruption by mutation. That regulatory mutation must have hap-
pened more than once. What the advocates are really arguing against is the idea that
the entire developmental suite was re-engineered by mutation and selection on two
separate occasions, which would be a non-parsimonious claim, if made. As the SET
account holds development constant this is in fact not a necessary commitment for
that position. The advocates thus fill in the detail of how that constancy might be
delivered and in so doing render a less abstract explanation, but no less idealized.
Recently, a related argument has been made. The claim is that much of the debate
between SET and EES turns on a distinction between structuralism and
adaptationism, which are seen as two separable scientific projects in their own
right (Brown 2022). Brown claims that both structuralism and adaptationism are
attempts at understanding “phenotypic diversification and the mechanisms that
generate it” (2022: 2). Structuralism is concerned with developmental constraints,
bias, and innovation as causes, while adaptationism is concerned with adaptation and
natural selection, approaching this in diverse ways from population genetics to
behavioral ecology. Brown’s innovation is to separate three different kinds of
adaptationism and structuralism: empirical, explanatory, and methodological
(Table 1.1).
Empirical commitments to adaptationism or structuralism amount to an ontologi-
cal claim for causal primacy in evolutionary explanations. This is to be contrasted
with the softer explanatory approaches which begin to make explicit claims about
1 Introduction 9
the explanatory targets and then point to specific frameworks as more useful. To this
end, adaptationism is focused upon apparent design while structuralism focuses
upon the diversity, disparity, and complexity of phenotypes. Finally, methodological
approaches are more pragmatic affairs, in which explanatory targets differ but the
framework adopted is merely seen as the best starting point for hypothesis genera-
tion and test. However, Brown notes that while methodological adaptationism seems
to be a straightforward commitment for most adaptationists, due simply to the great
success of this approach, EES advocates claim that in recent years the amount of new
developmental data calls this assumption into question. Brown suggests that the
argument is that adaptationism as a method is now eclipsed by the availability of
structuralist data. She further notes that structuralists can still see adaptation as an
explanatory target, but that their shift is away from the pure externalism of selection
toward a more interactive, or constructivist model of building those traits. In this
way, developmental causes at a minimum have explanatory parity with selection.
We think Brown has the EES claim right here, but we would note that this still
amounts to a shift of explanatory focus as described above. If the overall aim is to
explain phenotypic diversification and the mechanisms that produce it, then we can
see these two approaches as an effort toward that. However, the devil is in the detail.
For an adaptationist, the question of phenotypic diversification is one of what is
selected, what is retained in the population, and why. Selection is the mechanism,
and as inheritance is required, genetic variation is the source of novelty under
constancy assumptions about development (as above). For the structuralist, the
question of phenotypic diversity is a question of the multiple causes of variation in
the phenotype prior to any selection. This does not mean that they must deny
selection, selection is simply not a principal focus. As suggested the structuralist is
less abstracted than the adaptationist. But Brown might claim that the developmental
constancy of adaptationism is an idealization. We think this is both an idealization
and an abstraction because the variation introduced by development is in fact already
captured as an idea in SET through the concept of reaction norms. Reaction norms
are understood as the available range of phenotypic expression for a genotype, and it
is assumed this impacts upon evolutionary dynamics downstream (Stearns 1989).
Reaction norms are therefore an available explanation for SET, but they are idealized
out of the main account under constancy assumptions (in part because it is assumed
selection has operated to design conditional response in phenotypic expression
(Nettle and Bateson 2015). If EES advocates were to argue for unbounded pheno-
typic variation, with no connection to genetic variation, then they would be in want
of a new mechanism for inheritance to continue their evolutionary accounts. Of
course, some theorists have latched onto epigenetic inheritance as a possible second
mechanism but the causal dependency of epigenetic effects upon genes is strong and
this claim unsubstantiated (Dickins and Rahman 2012; Futuyma 2017).
We will stop here. Our aim is not to launch into a full analytic argument about key
claims on either side of this debate. Instead, we want to show how a range of
engagements is possible with this literature. At one pole, there are clear analytic
disagreements to be had and this tends to happen as explanatory aims overlap or are
entirely in common. At the other, there is the complete separation of explanatory
10 T. E. Dickins and B. J. A. Dickins
targets and the emergence of specific strategies to gain understanding. With this view
in mind, we turn now to discuss what is to come in this volume.
This book aims to survey various aspects of the debate between EES and SET
advocates. There is no ambition to be partisan, but rather to inspect key claims and to
place them in contemporary and historical contexts. Our authors are drawn from
science, and the history and philosophy of science, offering various perspectives on
elements of the argument as it has been played out over the last two decades. We
have also introduced a discursive element to the book, producing commentaries on
chapters, from within the author list, and allowing response. Our hope is to convey
the liveliness of debate through this method, but also to expose further lines of
thinking beyond the original chapters. In this section, we will survey the main
chapters of the book. Each main chapter is grouped with its commentary and reply
into an individual part.
In Part 1, Chap. 2, Betty Smocovitis addresses the uses of history in the debates
surrounding the Extended Evolutionary Synthesis. She opens by urging caution,
there are many pitfalls to deploying historical narrative in support of particular views
about present theories. An early example of this is her analysis of the term Modern
Synthesis. This is what historians label a trace because it indicates something about
the actor who deployed it. In this instance, that actor was Julian Huxley, and
Smocovitis discusses the mid-twentieth century mood to which he was addressed
in labeling the emergence of evolutionary biology the Modern Synthesis. What this
term is not is something derived from secondary analyses by historians. A modern
synthesis has not been unearthed as a movement. Rather Huxley’s clever coinage,
used in a book designed for a wide, if intellectually curious audience, has stuck. This
does not mean that our early twenty-first-century mood will unambiguously and
correctly interpret the term as it was intended. Smocovitis also discusses the later
adoption of evolutionary synthesis, by Mayr and others, to firmly focus on the
discipline-building activities of evolutionary biologists while controlling the histori-
cal narrative. This habit of leading scientists to write their own histories of evolu-
tionary biology was common, and Smocovitis surveys several key contributions
revealing different sub-disciplinary emphases, alternative lists of the core architects,
and even disagreement about the duration of any synthesis. These observations draw
Smocovitis to a critical analysis of how certain key terms are casually deployed in
the extended synthesis debate, without care for their origins and conceptual place in
the past, sowing confusion and much crosstalk. Her plea is for histories of science to
be produced that try to place the work of scientists in their appropriate moment, not
just through the interpretation of their scientific publications but also by understand-
ing the zeitgeist of the moment in which the work was done; partial histories are
clumsy rhetorical tools (see also Chap. 11). She ends by ably demonstrating the
inadequacy of some partisan histories deployed in the recent debates about exten-
sion, and relates them to prior arguments in the 1980s, when standard evolutionary
1 Introduction 11
theory was also declared to be in decline. A clear message from Smocovitis, and one
important for readers of this book, is that there never has been a single, monolithic
theory of evolution. But there has always been disagreement and diversity within
evolutionary biology.
Chapter 5 (Part 2), by John Welch, parses the many arguments about creativity
and natural selection that have arisen in the literature. As a result, the chapter spans a
considerable section of the history of evolutionary thought in order to position key
claims about creativity. Welch focuses the chapter on the idea that the theory of
natural selection was specifically developed to deal with apparent design in
organisms, with the concept of adaptation. This is one source of creativity in biology
and does not rule out others, but Welch makes a case for how these different forms of
creativity can be related to and separated from one another, in part dependent upon
the scientific focus brought to bear. While he does not directly discuss abstraction
and idealization, the contemporary views of modeling discussed above are evident
within his analysis. He also demonstrates how a tendency to lose ascription in the
literature has led to ambiguity and, at times, hyperbole with an inevitable loss of
explanatory traction. Ultimately Welch’s chapter provides a conceptual structure that
supports a form of pluralism, that of need. Under this account, natural selection, and
thus the standard evolutionary theory derived during the Modern Synthesis, has a
central and organizing role but is not to be considered the sole causal source of trait
variation and nor is selection isolated from the dynamics of organismic agency. The
explanatory needs of researchers should determine which aspect of the conceptual
architecture is most relevant to them, while understanding its place relative to the
whole.
In Chap. 8 (Part 3) Jan Baedke and Alejandro Fábregas-Tejeda present a histori-
cal and philosophical analysis of the role of the organism in evolutionary biology.
This chapter complements that of Welch in that it analyzes the history of the claims
about the role of organismic agency in evolution. They outline the emergence of
organicism, the non-reductionist, non-vitalist third way position that fell out of favor
in the twentieth century. Reasons for this decline are given and include a lack of
institutional support as well as specific, reductionist moves in the philosophy of
biology developed by Mayr and others during the Modern Synthesis. Baedke and
Fábregas-Tejeda then make the case that the Extended Evolutionary Synthesis is
advancing a new version of organicism, albeit without explicit reference to their
third-way forebears. The EES movement, by granting the organism a role in
evolution, looking to reciprocal interactions between organism and environment,
and by showing organisms to be agents actively shaping their environments, is
introducing the main themes of organicism as outlined by Baedke and Fábregas-
Tejeda. But Baedke and Fábregas-Tejeda conclude their chapter with a note of
philosophical caution. While the organicists and advocates for an EES have clearly
laid out what they see as missing elements in evolutionary biology, those elements
need to be drawn into a theory that displays key explanatory virtues which include
the proper deployment of abstractions and idealizations. The implication is not that
this is an impossible task, but rather that is an essential one, if any form of pluralism
12 T. E. Dickins and B. J. A. Dickins
is to be defended. For the authors this is an effort that cuts across debating sides—all
need to decide upon their explanatory standards and give reason for them.
In Chap. 11 (Part 4) Erik Svensson, in keeping with the preceding chapters, looks
to the historical antecedents of the Extended Evolutionary Synthesis and criticisms
of the ideas emerging from the Modern Synthesis. A key point from Svensson is that
these criticisms often ignore the developments in evolutionary theory since the
historical synthesis period (for example, neutral theory). Central to Svensson’s
argument is the idea that the Modern Synthesis was a period of intentional synthesis
between the subdisciplines of biology, under a shift from a natural historical,
organism-focused discipline to a process-based one. While he notes it may not
have included all elements of biology, he claims this effort toward synthesis should
not be understated. It is this synthetic ambition that continues to this day in
evolutionary biology and allows neutralism and other developments to become
incorporated. Evolutionary biology is a dynamic discipline and not one in crisis as
some critics have suggested. To this end, Svensson sees the Modern Synthesis as the
formation of a framework for doing science, not a formal theory, and he makes a case
for two broad schools of thought emerging during the synthesis to support this claim:
one in the UK and the other in the USA. This leads Svensson to make the stronger
claim that the criticisms of the Modern Synthesis are often focused upon an inaccu-
rate presentation of contemporary evolutionary theory based in partial accounts of
the past that neglect the inherent pluralism of the synthesis period. Svensson sees
evolutionary biology as encompassing several theories that are determined by the
focus of the researchers in question, and this again is in keeping with modern views
of scientific explanation outlined at the beginning of this chapter. But he also
recognizes that empirical evolutionary biology has raced ahead of theory in recent
years as many novel findings have accumulated. There is a job to be done here, but
his view is that it will not be achieved by misrepresenting the past and calling for
reform—instead, the history of evolutionary biology shows a flexible discipline that
has consistently integrated and re-engineered itself.
David Haig draws our focus to teleological and teleonomic explanations in
Chap. 14 (Part 5). He begins by discussing the odd nature of evolutionary theory
when addressing cause and effect. Where we normally hold that an effect cannot
precede its cause, evolutionary theory renders this more complex showing how the
outcomes of genetic variation (which are effects) can then cause the perpetuation of
those genetic variants and the associated traits due to selection. This is the distinction
between type and token causation. It is the treatment of cause and effect in biology
that is of concern to Haig, but before he directly addresses this issue, he argues for
differences in interpretation based on need. While his comments are founded in a
manner consistent with Derrida (an unusual move within theoretical biology) his
argument is in keeping with the idea that scientific interpretations of facts are derived
with a specific explanatory purpose in mind. Following Svensson’s comments, Haig
sees a tendency for establishing strawmen but on both sides of the debate about an
Extended Evolutionary Synthesis, in order to support specific interpretations at the
expense of others. Given this, Haig sets to analyzing, and re-interpreting the work of
Laland and colleagues on reciprocal causation and related matters, with a mind to
1 Introduction 13
modulates kin competition. For Rodrigues and Gardner, this is a textbook case of
reciprocal causation that emerged during the 1970s. They continue with such
analyses to conclude that core Extended Synthesis requirements for reciprocal
causation and a focal role for organisms were the very concepts that motivated the
development of inclusive fitness theory, while others were subsequently
incorporated as the theory developed. Given all of this, Rodrigues and Gardner
wonder why inclusive fitness theory has been ignored in the debates around the
Extended Evolutionary Synthesis. Their suggestion is that advocates of extension
have primarily focused upon increasing the realism of models, by adding parameters.
In doing this scholars are seeking to reduce the abstractions within models, while
inclusive fitness theory has been aimed at producing an abstract generalizable
explanation that cuts across many phenomena. To this end, the Extended Evolution-
ary Synthesis is less a scientific endeavor and more a set of quibbles, to borrow a
phrase from Rodrigues and Gardner.
In Chap. 23 (Part 8) Stephen Abedon provides a case study in post-Modern
Synthesis theory development in phage biology. Owing to their relatively late
discovery and starring role in the development of molecular biology, phage life
cycles have been characterized in painstaking mechanistic detail. Abedon shows
how concepts from ecology, and optimal foraging theory, can be brought to bear on
phage life cycles. This first entails re-parsing the life cycle so that variations can be
considered that modulate reproductive output, measured in whole organisms. Focus-
sing primarily on lytic cycles, Abedon shows that the intracellular development of
phages imposes an opportunity cost on phages that is traded off against the number
of offspring produced upon cell lysis (the burst size). Ecological parameters are then
considered that influence this trade-off in addition to organism-level constraints that
limit optimization. Abedon thereby reveals an array of intrinsic and extrinsic features
to be considered in formal and informal models. This exemplifies how applying
resources from evolutionary theory is both productive and attention-directing:
focussing the researcher on features of the life cycle that may, at first, appear trivial
such as the kinetics of phage adsorption by host cells. Two features that are
challenging to incorporate in formal models are reproductive variance and related-
ness. Considering the first, Abedon describes the concept of effective burst size
showing that a single parameter can be devised that incorporates multiple details
(including host cell density). Returning to the broad framework (viz., weighing time
spent within hosts in the form of a virocell against time spent extracellularly as “free
phage”) Abedon shows that this can be brought to bear on the lysis/lysogeny switch
exhibited by temperate phages. Also at the comparative level, Abedon separates lytic
and lysogenic life cycles from chronic-productive infections by mapping these to
semelparous versus iteroparous reproductive strategies, respectively. We also see
examples of reciprocal causation in this chapter with phages influencing their own
biotic environment by depleting host cells. Overall, like all good science, the
application of theory by Abedon is seen to be at least as generative of questions as
it is of answers.
In Chap. 26 (Part 9) Tom Dickins focuses upon the topic of plasticity. The chapter
begins with a discussion of Pigliucci’s argument that a mechanistic theory of form is
16 T. E. Dickins and B. J. A. Dickins
early in the history of the discipline, gaining refinement through the synthesis and
later from inclusive fitness theory—and asks what the Extended Synthesis would
change in order to explain this phenotype. Shuker begins with a comprehensive
summary of the major theoretical transitions in sex allocation from Darwin onward.
He then summarizes the core claims of the Extended Evolutionary Synthesis as a
focus upon organismic agency and an emphasis upon non-genetic inheritance. These
claims are then related to sex allocation. Unsurprisingly, Shuker sees organismic
agency as central to sex allocation, with adjustments made given sensitivities to
crucial environmental contingencies. Shuker does not make a claim about the kinds
of mechanisms one should allocate to agency but merely commits to their being
mechanisms that are facultative and thus make decisions. This is a minimal view of
agency, but one seen in the writings of the Extended Evolutionary Synthesis also.
Shuker firmly places this kind of decision architecture within the domain of plastic-
ity, another topic central to advocacy for extension, and makes a strong case for sex
allocation as a canonical form of plasticity in enabling multiple outcomes from a
genotype. Shuker continues to relate sex allocation to non-genetic transgenerational
effects, development, niche construction and many of the key topics of discussion
within extended evolutionary circles. As with Rodrigues and Gardner, Shuker asks
why the Extended Synthesis has ignored this active domain of evolutionary biology
that appears to tick all their boxes. In addressing this Shuker refers to recent
philosophical discussion about explanatory virtues and varieties of explanatory
purpose. For Shuker, this is something already enshrined within the field, thanks
to Tinbergen’s four questions. The expectation was not that each biologist addresses
all four, but rather that the collective effort of all biologists across these domains
would lead to a complete understanding. Shuker’s strong conclusion is that the
extension sought is less extension and more detachment. He argues that advocates of
the Extended Evolutionary Synthesis have a laser-like focus upon the construction of
form and in so doing are simply not addressing evolutionary questions in the first
place.
Douglas Futuyma discusses the evolving evolutionary synthesis as an interaction
between theory and empirical findings (Chap. 32, Part 11). Early in his chapter
Futuyma discusses the generality of population genetics, which was central to what
he refers to as the Evolutionary Synthesis. 1 Population genetics tells us how
evolution works, but not about specific features or taxa, it can predict short-term
outcomes such as allele fixation, but it cannot explore macroevolutionary trends,
such as diversification. Here, Futuyma shares some of the concerns of the Extended
Evolutionary Synthesis, as the biology of actual organisms is absent from population
genetics. But he continues, claiming that much of the synthetic effort was to then
accrue evidence of genetic variation from multiple biological disciplines and to look
to the evolution of morphological traits, etc. in terms compatible with population
genetics. Not all disciplines were included, and he lists physiology, development,
1
Some historians of science prefer to use the term Evolutionary Synthesis to denote the process of
theoretical synthesis and to separate it from claims to modernity (Smocovitis 1996).
18 T. E. Dickins and B. J. A. Dickins
and ecology as missing parties, noting disagreement about some aspects of exclusion
in the historical literature. Futuyma then moves to outline controversies within
evolutionary biology: neutralism, levels of selection, sympatric speciation,
punctuated equilibria, and adaptation and constraint. The lesson he draws from
analyzing these controversies is that, while some of the initial suppositions may
have proved incorrect, they each helped to improve knowledge and understanding.
Futuyma does not use this observation to defend the role of controversy but rather to
hint at the robustness of the original theory, developed during the synthesis.
Futuyma then discusses the Extended Evolutionary Synthesis, as a controversy,
and with particular focus on the 2014 exchange in Nature, that he was a part of
(Laland et al. 2014). He carefully goes through each of the major areas discussed in
that exchange—niche construction, evolutionary developmental biology, plasticity,
and inclusive inheritance—demonstrating what has been studied within these areas
prior to the advent of calls for an Extended Evolutionary Synthesis. Futuyma pulls
no punches and makes clear where claims for extension are overwrought, in particu-
lar choosing to see niche construction theory as a simple rebranding of community
ecology. But he is also charitable and argues that while controversy may not be the
best mechanism for moving science forward (he claims it is impossible to
counterfactually know whether it is), advocacy for extension may provide a useful
service in highlighting important work that requires further integration into evolu-
tionary biology. Thus, the thrust of Futuyma’s argument is that this is business as
usual, as we saw during the Evolutionary Synthesis, and not a wholesale demolition
of core theory but rather integration of empirical findings leading to improved
understanding in a manner he aligns with the Kuhnian concept of normal science.
Futuyma’s argument about the generality of the population genetic approach to
evolution amounts to a claim for high-level abstraction, and his view of normal
science to the reduction of abstraction through the addition of empirical detail. He is
quite clear that empirical details can challenge and overturn theory, but at this point
in the history of evolutionary biology he sees no evidence of this having happened.
Central to the Extended Evolutionary Synthesis claim is the criticism that Stan-
dard Evolutionary Theory, emerging from Modern Synthesis, is gene centric at the
expense of the organism (Chap. 8). In Chap. 35 (Part 12), J. Arvid Ågren discusses
the changing fortunes of both gene and organism concepts in evolutionary biology.
He surveys various views from within and without the Modern Synthesis,
demonstrating how the gene’s eye view of evolution has divided biologists and
philosophers since it first emerged in the work of Williams and then Dawkins
(Dawkins 1976; Williams 1996). Both authors made cogent arguments regarding
the problem of design as the central problem for evolutionary biology and for
shifting from the Darwinian focus upon individuals and groups to the level of the
gene to address this problem. Ågren argues that the project of understanding
adaptations requires solving a beneficiary problem—what are adaptations
good for? Williams’ and Dawkins’ answer is genes, understood as having the
properties of longevity, fecundity, and copy-fidelity. It is this understanding that
shifted the concept of the gene from a purely molecular or particulate one to that of a
replicator, in Dawkins’ terminology. In determining the replicator concept,
1 Introduction 19
organisms were assigned to the category of vehicle, lacking the essential properties
for evolutionary process, but enabling genes in their purpose through their differen-
tial survival and reproduction. The focus upon adaptation is related to Paley’s natural
theological legacy by Ågren, but he also discusses the role of Fisher’s early popula-
tion genetics. Unlike Wright, who believed he was modeling organismic evolution,
Fisher was focused upon adaptation and genes and Ågren shows how Fisher
introduced a simple segregation between genetic and environmental variation (vari-
ance) that resulted in the genome being seen as an environment, in the same way as
more standard ecological factors were. The final move that cemented the gene’s eye
view was the rejection of group selection and specifically naïve good of the species
versions of this argument. It is this decomposition of evolution to a gene-level
process, and the relegation of organisms to the vehicle category, that has led to
concerns that gene-centrism is detrimental to biology. Those advocating for an
extension to the Modern Synthesis are keen to prioritize the phenotype in evolution-
ary explanations, and therefore they seek a role for the organism. In this way, they
are reverting to a Darwinian core (Chap. 17). But as Ågren shows, this tension
existed within the Modern Synthesis also. We have already referenced Wright, but
Mayr too clashed with the gene-centric Haldane in the infamous bean-bag genetics
dispute. Moreover, inclusive fitness theory marks an organism-focused post-synthe-
sis development that is neutral about genes even though it can be cashed out in
gene’s eye terms (Chap. 20). This aspect of inclusive fitness theory has frustrated
Dawkins and Maynard Smith, as Ågren makes clear, because of the difficulties of
calculating a value at the individual level, which has led Dawkins to re-emphasize
the utility of the gene’s eye view for gaining explanatory traction. Ågren discusses
genetic conflicts and shows how this challenges the inclusive fitness whole organism
view that there is a unity of purpose in the organism. This leads to some inclusive
fitness theorists idealizing such conflicts as absent to deliver models for specific
phenotypes. It is here that Ågren concludes that inclusive fitness theory works for
specific explanatory targets and as a result this supports the ambition to keep
organisms at the center of evolutionary accounts. But this is also a clear statement
that this ambition existed before the calls for an extension of the Modern Synthesis,
as well as a pragmatic point about the differing utilities of the gene’s eye and
organism-level approaches. Under Ågren’s interpretation, inclusive fitness theory
is an idealization, claiming unity of purpose at the level of the organism. It is perhaps
a thin idealization as the major transitions in evolution are regarded as cooperative
shifts to such unity, but nonetheless it is a deliberate falsehood (Maynard-Smith and
Szathmary 1995). This does not imply, however, that the gene’s eye view is without
idealizing assumptions.
Mitchell Distin focuses upon evolvability in Chap. 38 (Part 13). He opens by
claiming that evolvability was effectively overlooked during the development of
modern evolutionary biology. This was in part due to the abstract nature of evolu-
tionary theory (Chap. 32) and a lack of connection to nature. This obscured complex
evolutionary dynamics emerging beyond the individual, and Distin claims a division
between theory and empiricism emerged that was enhanced by a commitment to the
logical positivism of the early twentieth century. The ambition was to produce
20 T. E. Dickins and B. J. A. Dickins
mathematically expressed nomic statements about biology, something that did not
suit evolvability. By implication, Distin also relates this to a neglect of ecology, and
states that the discipline of evolutionary ecology is now coming of age and enabling
a true focus upon evolvability, for example within discussions of evolutionary
rescue. He also points to other developments around genetic evolvability
mechanisms such as stress-induced mutagenesis, which he argues point to high-
level selection at the species or lineage level, something perhaps anathema to earlier
critics of naïve multi-level selection accounts (Chap. 35) and Distin aligns this move
with specific pluralist and multi-causal explanatory approaches within the philoso-
phy of science. The phenomenon of evolvability demands a different kind of causal
model than commonly deployed by Standard Evolutionary Theory. From here Distin
moves to discuss evolvability as the cornerstone of the Extended Evolutionary
Synthesis, where evolvability is conceptually embedded within evolutionary devel-
opmental biology, or evo-devo. However, Distin does not wish to support the view
that evo-devo is the focus of evolvability, instead preferring to see evo-devo as a
method of thinking about evolvability. Indeed, Distin is clear that evolvability as a
generic concept has been present in evolutionary thinking for many decades, as he
carefully documents, but as above, its traction has been impeded by the abstract
nature of the evolutionary biology emerging during the Modern Synthesis. This has
led to a high degree of conceptual fuzziness in contemporary discussions of
evolvability and Distin offers to untangle this first with an analysis of the
evo-devo view of non-genetic evolvability. He surveys the work on modular devel-
opmental programs (see Kirschner 2013 for an overview) that reduce lethality effects
associated with novel mutation by reducing the number of mutations required to
introduce novelty. While praising the innovation and progress made in this field,
Distin sees it as hindering understanding of genetic evolvability, and obscuring
distinctions between short and long-term evolvability. The kind of genetic causality
Distin favors is that which leads to differences in mutation rates between species,
something that Distin notes natural selection can operate over. For Distin, the focus
upon evo-devo makes the causal account limited and perhaps too abstract. He wants
evolvability to encompass both developmental and genetic causes and for the causal
model of evolvability to clearly explain the different roles of each component. For
Distin, then, evolvability is a unitary phenomenon with broad application thanks to a
complex, causal mechanism. Indeed, early on he labels evolvability as an emergent
dispositional property, causally relevant at higher levels of organization over long
stretches of time. Thus, evolvability is multiply caused, and in turn, causes evolu-
tionary change. Distin uses this discussion to draw out a philosophical point about
pluralism. The fuzziness of the evolvability concept in the literature ought not to
encourage an unrestricted pluralism, allowing multiple different causal idealizations
to emerge, each operating within its own explanatory bailiwick. Instead, we need a
restricted pluralism, in which we aim to minimize, but not necessarily eradicate,
plural approaches, in order to sharpen our focus upon the phenomena. The subtext of
Distin’s chapter is that the initial abstractions of the Modern Synthesis led to a formal
neglect of various phenomena, including evolvability, but also gave licence to an
anything-goes culture among those addressing evolvability. To bring evolvability
1 Introduction 21
into evolutionary biology a clear set of phenomena associated with that label need to
be decided, and a part of that decision will be made with reference to Standard
Evolutionary Theory. In this way, any extension of the Modern Synthesis is quite
simply an extension of its phenomenal reach.
Our summaries of the chapters are intended as a rough guide to what follows. They
do not do full justice to the rich, detailed work that the authors have undertaken and
presented. While we have drawn out a theme about idealization in this chapter this is
by no means the only emerging theme, nor do we expect our readers to necessarily
agree with it. Instead, our hope is that this volume will stimulate continued discus-
sion and engagement within the broader community of scholars who think about
evolution and how to account for it.
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Part I
Every Evolutionist Their Own Historian: The
Importance of History, Context, 2
and the Extended Evolutionary Synthesis
V. Betty Smocovitis
Abstract
This chapter opens with a discussion of scholarly practices within the history of
science, noting a distinct difference between professional historians and
scientists-turned-historians. History is important and how it is done has
implications not only for our understanding of the emergence of a discipline
but also for contemporary debates within it. This theme is followed with a
detailed analysis of the often-partisan uses of history to define disciplinary
boundaries, to found disciplines, and to criticize them. Parallels are drawn
between the anti-adaptationist debates of the 1980s and recent calls for an
V. B. Smocovitis (✉)
Department of Biology, University of Florida, Gainesville, FL, USA
Department of History, University of Florida, Gainesville, FL, USA
e-mail: bsmocovi@ufl.edu
Keywords
Entering the conversation about the so-called extended evolutionary synthesis, and
its use of history reminds me of a classic 1932 essay by Carl Becker, one of the most
reflective of US-based historians long associated with Cornell University in Ithaca,
New York. Titled “everyman his own historian,” Becker drew the analogy between
everyday historical reckonings that human beings perform in their daily tasks and the
work of professional historians, scholars trained in the use of historical evidence,
methods, and publications (Becker 1932). His intention was to make the case for the
importance of history in everyday life, and to diminish if not eliminate its profes-
sional pretentions with respect to scientific history, the newfangled application of
scientific methods to understanding the past in objective terms. But while he
supposedly received a standing ovation when he first delivered it as his 1931
Presidential address to the American Historical Association, the essay, and its
analogy, eventually backfired because it introduced a kind of historical relativism
that Becker himself had not intended. It also highlighted the kinds of interpretive
ambiguities few historians were comfortable embracing, and indeed, had been
working hard to eliminate from their work. People do have the tendency to recall
the past in light of the present, and to selectively ignore or forget some things, while
remembering others; the gallons of ink spilled on discussions of selective memories,
for example, speaks directly to this (Pohl 2004), so why should historians be any
different, or so Becker’s logic goes. Everyone is going to craft their own history, and
erase it when they see fit, and though there may be a price, and a big one at that, to be
paid if others do not like it, everyone is entitled, in a sense, to do this, especially
when matters of self-existence and identity come to play. This latter point needs
underscoring because boundaries and identities are determined, and bloody wars are
fought, over matters of history. It does matter, hugely.
Becker was of course addressing history, writ large, but much of what he relayed
in his essay also speaks to the history of science, at least as I see it. To be sure,
historians of science do have their own issues; we frequently refer to “scientist’s
histories,” oftentimes deriding them because they lapse into the kinds of naïve
mythologizing that celebrate great discoveries, ideas, and historically white men
(Forman 1991). Such histories, or narratives, can simultaneously marginalize or
2 Every Evolutionist Their Own Historian: The Importance of History,. . . 27
exclude alternatives, while at the same time serving to sanitize the messiness and
conceal the arduous work, or the sociopolitical dimensions, involved in the practice
of doing science, especially when something like consensus-building comes to play.
Such histories can also be blinkered, or skewed by “Whig,” presentist, or even self-
serving perspectives, and these can then shape entire disciplines once they are
embodied in textbooks as the received wisdom of the field that enroll novices to
the craft (Ashplant and Wilson 1988a, b draw these distinctions at length). Thomas
Kuhn drew attention to most of this in his celebrated The Structure of Scientific
Revolutions in 1962 (Kuhn 1962), of course, but this is why historians of science are
wary when history is evoked by scientists-participants, especially in the context of
disputes and arguments over who or what is right, and who or what is wrong, or
when deployed in an arsenal when conflicts break out over the direction of an entire
field.
Apart from pointing all this out, however, historians of science are pretty well
powerless to do much of anything about it, especially if, following Becker’s logic,
we are subject to the same kinds of interpretive ambiguities ourselves. Scientists can
and will write their own histories as they see fit especially in a science like
evolutionary biology which is a historical science that has included more than its
share of evolutionists-turned-historians like Ernst Mayr and Stephen J. Gould
(Nitecki and Nitecki 1992). As I have argued elsewhere, their use of history as
well as philosophy has accompanied their crafting of evolutionary theory, both for
scientific and wider, more popular audiences (Smocovitis 1992, 1996). And it is not
part of some devious, underhanded, or even deliberate process that serves self-
interest narrowly defined; it is part of the active, and ongoing process of constructing
disciplinary narratives that unify and lend coherence to the group and define the
identities of its members, while at the same time unwittingly serving to exclude and
marginalize others; in short, they do what sociologists of science call “boundary
work,” they determine the contours of the discipline and who or what will be inside
and who and what will be outside its boundaries (Gieryn 1983, 1999; Shapin 1992).
And when the disciplinary narrative of an area of study like evolutionary biology can
tap into, or become part of, a wider narrative of say, the Enlightenment, or of the
history of science in western cultural contexts, and when it can encompass an
originary narrative of humanity, or of life as a whole, it can become part of a
grand unifying narrative (another technical term) that undergirds understanding of
the world, structuring a worldview or Weltanschauung that may tell us something
about the “human’s place in nature” (Greene 1981; Smocovitis 1992, 1996).
I begin my chapter with these historical reflections because I think that they are at
play in many of the contemporary discussions and debates over evolutionary theory
especially pertaining to the extended synthesis. My task, as I see it, is not to weigh in
on scientific matters, but instead to draw on the historical record as revealed by the
traces left behind by the historical actors, and to engage and understand them on
their own terms, avoiding presentism to the extent that I can, and then to examine the
uses of history by contemporary evolutionary biologists, especially as understood
and deployed in some contemporary discussions and disputes. In this way, I hope to
offer some historical observations, try to reduce, if not eliminate some
28 V. B. Smocovitis
misunderstandings, and to offer some clarification in the way of moving the discus-
sion forward in a more productive direction. My firm belief is that despite the
frequent use and referencing of history, many of these disputes and discussions are
based on several historical misunderstandings, rely on a problematic, confused, and
underexamined terminology, and at times adopt a wholly ahistorical view of
evolution, especially of evolutionary theory. Indeed, they are too much from the
“every evolutionist their own historian” perspective and thus lead to unnecessary
confusion and discussions that are often at cross-purposes.
In the way of laying some groundwork, I would like to start with the word “histori-
cize”, or “historicism.” To some, it might appear like academic jargon, the kind of
thing common in the humanities, associated for example with literary theorists like
Frederic Jameson, and his well-known slogan “always historicize” (Jameson 1981).
But I would argue vigorously that there is nothing jargon-esque, or fashionable about
it to those of us studying history as well as a historical science like evolutionary
biology. It simply means to embed in history, to render a thing as an object of
history, and to give it meaning and signification within a historical web of beliefs and
practices; it means to think diachronically, processually, or indeed, in evolutionary
terms. Let me offer a concrete example: does a fossil have much meaning without a
sense of where it is found, not just in what kind of environment, but in what kind of
temporal sequence, or context, of what came before, what is found near it, and what
comes afterward? The answer is not really, because the meaning of fossils is
understood in historical sequences as well as environmental contexts. The same
holds true for ideas, beliefs, practices, and even scientific theories. But realize the
twist in accepting this; it means that ideas, beliefs, practices, and even scientific
theories are embedded, or situated in particular contexts and understood diachroni-
cally; they change, evolve, and come to be. They do not transcend history and culture
but take on a more local or historical specificity, and themselves evolve, though the
process is very different from that operating in organic evolution. The view that
theories evolve, moreover, is not much of a novel or even radical idea to reflective
evolutionary biologists; Douglas J. Futuyma has made this point more than once and
argues for it indirectly in his contribution in this collection (Chap. 32), for example
(and see CALLEBAUT 2010).
This gets me to my next word, “context” or “contextualize,” which adds the
cultural, or social, and political to the mix, especially when referring to ideas, beliefs,
practices, and even theories, which do not arise de novo, but draw on raw materials
from the past and prevailing currents of thoughts, habits, or what in the humanities
are called “circulating discourses.” To the historian, this is a kind of reframing of the
older concept of zeitgeist, though this current version may also incorporate material
practices as well as the prevailing thoughts or spirit of an age along with a closer
attention to language because thoughts—ideas—are embodied in words. Ideas
2 Every Evolutionist Their Own Historian: The Importance of History,. . . 29
themselves are not viewed as unit particles that arise from the brain of a Zeus, a
“great man of science,” or a “genius” to be transmitted and bounced off others in a
billiard-ball-like causal process, but instead exist in the forms of words comprising
languages that circulate, moving around in a cultural system not unlike materials that
cycle in ecosystem ecology, but in that process may alter in meaning. So, to
contextualize means to embed words within other words in texts that enable us to
access or understand cultures that are different from ours (Smocovitis 1996). To the
evolutionary biologist, such a view of context, also provocatively opens the door to a
plurality of legitimate theoretical perspectives, each a kind of culture, emphasizing
one or other aspect of evolution, based on particularities and specificities of methods,
training, generation, field, or organismic system. It is provocative because it
challenges the assumption that there is one singular, unifying, or grand theory, and
instead opens the door to the possibility of multiple overlapping theories emerging
from context and history.
We move then to another important word that I have already used in the
introduction and that is “presentism,” the tendency to project the present into the
past or to interpret the past in light of the present. Lapsing into presentism gives us
not just a distorted view of the past, but turns history into a kind of extractive
industry, full of moral and ethical quandaries since it often comes at the expense of
the historical actors who are rendered voiceless in the process. What is it we actually
want to know, anyways? Do we want to know something about them, or do we want
some affirmation from the past about us today? If we want to know something about
them, we have to make every effort to understand them as they were, especially if we
are trying to trace our historical roots. L. P. Hartley’s famous opening line to his
novel The Go-Between that the “past is a foreign country” was not just a pithy
statement about the past (Hartley 1953); we enter the past much as foreigners
encountering another culture, needing to learn another language. Suffice it to say
that presentism is a violation of basic historical methodology and something to be
avoided; paying careful attention to the words and phrases as we trace their occur-
rence helps us to avoid doing this. So, I end this section of my chapter underscoring
the importance of language as a way of mediating between history and context and
stating explicitly that, along with Kuhn and subsequent students of the history and
philosophy of science, science is best viewed as a historically rooted and culturally
embedded practice. Such an approach enables us not just to mediate between
historicism and context, but to also draw on a mix of approaches from history,
philosophy, sociology as well as anthropology, in a kind of multidisciplinary
approach that will help us gain a greater understanding of the past on its own
terms. That in turn, I believe, may allow us to avoid miscommunication and some
of the more contentious conversations in evolutionary biology today (Smocovitis
2021).
30 V. B. Smocovitis
Next, I would like to sort through some of the actual language of the synthesis
focused on naming and draw some useful distinctions in terminology. In the process,
I hope to also cover some ground in the history of evolutionary biology. We have for
example frequent reference to the “modern synthesis” of evolution. This term comes
straight out of the title of Julian Huxley’s well known and oft-cited comprehensive
treatment of evolution originating in the late 1930s and published in 1942 as
Evolution: The Modern Synthesis. It was a widely read book aimed at a more
general, semi-popular audience, the kind of synthetic book offering a scientific
worldview that Huxley was very good at writing. Heir to the Huxley family legacy,
Julian was a gifted writer and popularizer of science, becoming a kind of celebrity in
his own time. Putting his skills to use, Huxley wrote the 1942 book drawing on an
earlier essay of 1936 titled “Natural Selection and Evolutionary Progress” that
reflected his belief in a progressive view of evolution, that would also help ground
a secular, liberal worldview that was increasingly divided by extreme ideologies
(Smocovitis 1996, 2016). The publication of the 1942 book has traditionally
heralded that a new and modernized evolution had emerged, one that synthesized
Darwinian selection theory with Mendelian genetics (Mayr and Provine 1980). But
perhaps equally important, Huxley framed this “modern synthesis” with his intro-
duction that offered a kind of early disciplinary narrative of what became a new field
of study, namely evolutionary biology. As a disciplinary term, “evolutionary biol-
ogy,” was only just gaining currency, and Huxley was just starting to use it
interchangeably with the broadly conceived but generic “evolutionary studies”
(see Smocovitis 1996, p. 162 for more on the etymology of “evolutionary biology”
and the history of the discipline). Setting up this “modern synthesis of evolution”
Huxley stressed the preceding two decades of work that turned a mostly descriptive
natural history-oriented study of evolution into a rigorous science grounded in
observation and experiment that had drawn novel insights from population genetics
and mathematical modeling. This, according to Huxley, made possible the fusion of
genetics and Darwinian selection and served to reanimate Darwinism following a
period he named “the eclipse of Darwin” that had seen several theories that had
either built on, amended, rivaled or even challenged Darwinian selection theory,
especially after the year 1900 when Mendelian genetics began to gain currency. Like
a “mutated phoenix risen from the ashes of the pyre,” he wrote with dramatic flair,
this “reborn” Darwinism made natural selection a “fact of nature capable of verifica-
tion” and made natural selection one of the fundamental principles of biology.
Biology itself, according to Huxley, was undergoing its own “phase of synthesis”
bringing together a set of newer sub-disciplines previously isolated and often
“contradictory,” and was in the process of becoming a “more unified science,”
rivaling the “unity” of sciences like physics and chemistry (Huxley 1942 pp. 13–28).
Without getting too heavily into the details of Huxley’s intentions, and his life
and work, subjects that have been extensively explored (Keynes and Harrison 1989;
Greene 1990; Waters and Van Helden 1992; Smocovitis 1992, 1996, 2016; Bashford
2 Every Evolutionist Their Own Historian: The Importance of History,. . . 31
2022) we can think of his “modern synthesis” as an actor’s phrase or even as a kind
of actor’s category with a discrete periodization (interwar and wartime) because it is
associated with the title of Huxley’s book. The phrase “modern synthesis” thus has
direct relevance not just to understanding Huxley but also speaks to the context of
the 1930s and very early 1940s and possibly the two decades preceding that, since
Huxley was trying to trace the tortuous history, in a kind of “rise and fall narrative”
of Darwinism in setting up his argument for a modern synthesis of evolution; what I
am getting at here is that for our purposes, the term “modern synthesis” comes from a
primary source. In historical parlance it is a trace, one that directly reveals something
about the past and the historical actor who used it and gave it signification. This is in
contrast to secondary sources, works by subsequent commentators, usually
historians or people trying to reconstruct history, offering interpretations of the
past, based primarily on following traces and using primary sources; that, and the
scholarly understanding that has accumulated in the historiography, a term which
refers to an understanding gained from a given body of historical literature that has
been written or produced by scholars working in the subject area.
Consideration of secondary sources takes us to yet another term used frequently
by some evolutionary biologists, usually those who consult historical scholarship
more frequently, namely the “evolutionary synthesis.” The term is most closely
associated with Ernst Mayr and William B. Provine’s edited collection of 1980, a
now classic, foundational, and entry-level work for anyone interested in the history
of evolution. The collection grew out of two workshops held in 1974 devoted to the
subject and organized by Mayr and Provine. It included a number of original
participants, or “architects,” of the evolutionary synthesis (a self-designated or
identifying label used by Mayr and others who were still alive to reflect on their
work and its outcomes; see the index in Mayr and Provine 1980 for the list of
conference participants) but it also included a number of historians and philosophers,
many of whom, like Provine were junior scholars and keen to understand the history
and philosophy of evolutionary biology, especially at time when the physical
sciences had long held too much dominance; an examination of evolutionary biology
had the potential to yield novel insights, especially because it was a historical
science, unlike physics or chemistry.
As such, the workshop, the abundant transcripts, biographical sketches, and
correspondence (all safeguarded and deposited in archival collections) along with
the published edited collection is an important and fascinating mix of both secondary
as well as primary sources, sometimes indistinguishable, that anyone interested in
the history of evolution may consult. Indeed, one of the most interesting aspects of
the “evolutionary synthesis” is the length to which Mayr and Provine (but especially
Mayr) went to ensure that all the materials would be available for future reference. I
do not believe this was accidental but view it as part of Mayr’s strategy of drawing
attention to “the evolutionary synthesis” and to actively fashion it as a major
historical event in the history of science. His choice of terminological change from
“modern synthesis,” which had gained some currency after Huxley, was in response
to a title that appeared too generic and too vague and was getting lost to other
syntheses. Retitling it to “evolutionary synthesis” left little doubt that this was a
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his wife and not for these pages. In a month the few intimate friends
she saw had grown tired of telling her how charming she looked in
black. In the settling of the estate, despite the money owed to his
creditors, Sam had left her far from penniless. The house in Park
Avenue was sold, and all it contained—the pictures alone bringing
her a comfortable fortune that many another woman in her situation
would have been satisfied with. Rose Van Cortlandt considered it a
mere pittance. She found a bond of sympathy among other widows
who had been reduced to twenty-five thousand a year.
Lamont became a frequent visitor to her smart little studio apartment
in Washington Square—to which Sue was never invited, and where
we shall leave Rose Van Cortlandt to the care of a few so-called
Bohemians to consume her whiskey and cigarettes.