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The emerging structure of the Extended Evolutionary Synthesis: where does Evo-Devo fit in?

Abstract

The Extended Evolutionary Synthesis (EES) debate is gaining ground in contemporary evolutionary biology. In parallel, a number of philosophical standpoints have emerged in an attempt to clarify what exactly is represented by the EES. For Massimo Pigliucci, we are in the wake of the newest instantiation of a persisting Kuhnian paradigm; in contrast, Telmo Pievani has contended that the transition to an EES could be best represented as a progressive reformation of a prior Lakatosian scientific research program, with the extension of its Neo-Darwinian core and the addition of a brand-new protective belt of assumptions and auxiliary hypotheses. Here, we argue that those philosophical vantage points are not the only ways to interpret what current proposals to ‘extend’ the Modern Synthesis-derived ‘standard evolutionary theory’ (SET) entail in terms of theoretical change in evolutionary biology. We specifically propose the image of the emergent EES as a vast network of models and interweaved representations that, instantiated in diverse practices, are connected and related in multiple ways. Under that assumption, the EES could be articulated around a paraconsistent network of evolutionary theories (including some elements of the SET), as well as models, practices and representation systems of contemporary evolutionary biology, with edges and nodes that change their position and centrality as a consequence of the co-construction and stabilization of facts and historical discussions revolving around the epistemic goals of this area of the life sciences. We then critically examine the purported structure of the EES—published by Laland and collaborators in 2015—in light of our own network-based proposal. Finally, we consider which epistemic units of Evo-Devo are present or still missing from the EES, in preparation for further analyses of the topic of explanatory integration in this conceptual framework.

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Fig. 1

Image conceived by Fátima Sofía Ávila-Cascajares and Alejandro Fábregas-Tejeda; image design by Casandra Lizbeth Méndez-Martínez, Diana Martínez Almaguer and Julio César Montero Rojas (color figure online)

Fig. 2
Fig. 3

Notes

  1. 1.

    Before moving forward, we point out that our discussion will be centered mainly around the ideas and concepts espoused in the book edited by Pigliucci and Müller (2010a, b) and in Laland et al. (2015), which constitute the mainstream usage of the framework dubbed ‘Extended Evolutionary Synthesis,’ but we recognize that many other authors in recent years have proposed diverse views as to how to ‘extend’ (or ‘expand’) the current evolutionary synthesis (e.g. Gould 2002; Kutschera and Niklas 2004; Brooks 2011 and citations therein; Tëmkin and Eldredge 2015 for an updated version of the ‘Hierarchy Theory of Evolution’ (HTE) that incorporates network analysis; see also Pievani 2016a; Eldredge et al. 2016). In the literature focused on the integration of non-genetic heritable variation into wider evolutionary frameworks, somewhat interchangeable terms to refer to some of the themes and motifs of the EES are being used, e.g. ‘Inclusive Evolutionary Synthesis’ (Danchin 2013) or ‘Pluralistic Model of Inheritance’ (Bonduriansky 2012). Differences are more explicit in Bonduriansky and Day (2018).

  2. 2.

    The visual tradition for depicting the EES as a nested set of ellipses (a Venn diagram), initiated by Pigliucci (2009), has been influential for many evolutionary biologists and philosophers. For example, taking Pigliucci´s diagram as a starting point, the British physiologist Denis Noble recently published his take on the contours of the EES (Noble 2015), for him dubbed ‘Integrative Synthesis’ as it would be based on the integration of a variety of interacting mechanisms of evolutionary change; he sees it as “a nuanced multi mechanism theory of evolution” (Noble 2015; p. 7). The differences between the two diagrams are conspicuous: at variance with Pigliucci, Noble does not portray the ‘new synthesis’ as a perfectly continuous extension of the MS: several elements of its gene-centric stance will be left out, being replaced by wider views of inheritance, multilevel selection and a different view of genomic evolution. Instead of using a nested set of enlarging ellipses, Noble depicts the EES as a set of partially overlapping ellipses (with elements of the MS not in contact with the elements of the purported EES).

  3. 3.

    It should be mentioned that Pigliucci’s stance on the current juncture and the future of evolutionary thought has become more nuanced in recent years (see Pigliucci and Finkelman 2014; Pigliucci 2018).

  4. 4.

    In the traditional narratives, it was not until a few decades later that what we properly call ‘philosophy of biology’ was born (Palma 2015). Philosophy of biology integrated aspects, reformulations and critiques to the blind spots and flaws of twentieth-century philosophy of science—e.g. the rejection to strict determinism, ‘universal laws’ and physicalist epistemologies, accepting, instead, probabilistic predictions and historical explanations (see Mayr 2004)—as well as the unique theoretical and empirical developments of a mature biology (Palma 2015). For a different and robust historiography on the origins of philosophy of biology, see Nicholson and Gawne (2015).

  5. 5.

    Other possibilities would include conceptualizing the EES, just as the MS was conceived in past decades, as a multi-field theory (sensu Darden 1986) or, following Wassermann (1981), as a hypertheory that subsumes subordinate theories that deal with different but complementary evolutionary mechanisms; for additional discussions, see Love (2013, 2017).

  6. 6.

    This idea emerged in the context of an academic collaboration with Mexican philosopher of science Mario Casanueva López (UAM-Cuajimalpa, Mexico).

  7. 7.

    A possibility that may be worth exploring further is that, for a given moment in time, there could be more than one topological configuration of the EES existing simultaneously. For Love (2013; p. 332), in evolutionary biology we can see a “(…) multiplicity of theory structures constructed from the heterogeneity of methodological and epistemological goals” (italics added); in the future, if the EES reaches far-ranging acceptance, we could have many EES structures in the cases where distinct situated communities of evolutionary biologists follow different epistemic goals.

  8. 8.

    Theories can also be epistemic units for wider conceptual frameworks (e.g. hypertheories, see Wassermann 1981) that embrace several complementary theories or models.

  9. 9.

    A quote from Love (2013; p. 328) is insightful in that regard: “(…) new findings and conceptual developments may involve the reorganization or rearrangement of theory components, even if they do not ‘overthrow’ existing foundations. Part of the call for an EES could be a complaint about how evolutionary theory organizes biological knowledge and guides inquiry, not simply about whether it contains all of the relevant parts”.

  10. 10.

    We think the notion of ‘evolutionary theories’ in that context is adequate. Niche construction theory is the prominent example of one (see Odling-Smee et al. 2003; Laland et al. 2016). Eva Jablonka and Marion Lamb have also described the general picture of their book Evolution in Four Dimensions as a “sort of Darwinian theory (…) that sees DNA as a crucial heritable developmental resource, but recognizes that DNA is not the only resource that contributes to heredity” (Jablonka and Lamb 2007; p. 356; see Jablonka and Lamb 2014 for a revised edition of the book).

  11. 11.

    Although one could interpret Fig. 2 as something akin to a construction chain.

  12. 12.

    For three different accounts of the tensions that exist between Evo-Devo and the Modern Synthesis, consult Minelli (2010), Laubichler (2010), Craig (2015). They address, respectively, the viable possibility of integration, the insurmountable obstacles for doing so, and the possible pluralist co-existence of both as individual conceptual frameworks in evolutionary biology. Amundson (2005) is a good historical complement to these debates.

  13. 13.

    Fusco (2015) places the blame on the flawed and insufficient theoretical work being conducted; other scientific or sociohistorical reasons will not be addressed here.

  14. 14.

    We have selected, for the consideration of the reader, three additional fragments of Laland et al. (2015) where the prominence of developmental processes and developmental frameworks is patent: “Parent–offspring similarity occurs not only because of transmission of DNA, but because parents transfer a variety of developmental resources that enable reconstruction of developmental niches”(p. 4); “Niche construction also influences ontogeny and constitutes an important way in which environmental factors are incorporated into normal development, sometimes to become as dependable as genomic factors” (p. 4); “Too much causal significance is afforded to genes and selection, and not enough to the developmental processes that create novel variants, contribute to heredity, generate adaptive fit, and thereby direct the course of evolution” (p. 6).

  15. 15.

    Laland and collaborators consider that Evo-Devo is in no position to subsume or engulf the study of developmental plasticity (see Laland et al. 2015).

  16. 16.

    Eva Jablonka and Marion Lamb (pers. comm, July 2017) raised the concern that, by focusing on “just one paper”, we may be acting uncharitable or unfairly to gauge the contributions of Evo-Devo (or of any other research field) to the EES; that to settle the matter of the relative contributions of different fields of research to the structure of the EES, we should wait for a full-length exposition, similar to Huxley’s 1942 book, to appear. Although we agree with them (and encourage them to take up to task to write such book) in the sense that our philosophical analysis about the structure of the EES is preliminary and somewhat sketchy, we need to stress that it is just a response to one of the objectives laid out explicitly in Laland et al. (2015: p. 3): “Our objective here is to add substance to these debates by providing a clear statement of the structure, assumptions and predictions of the EES that is useful to both enthusiasts and skeptics, allowing its status as an alternative conceptual framework to be evaluated”. We shall add that theory presentations (see Griesemer 1984) are not philosophically innocuous as they reflect how scientists use their frameworks in practice and further demonstrate “how different structures are adopted to achieve specific methodological and epistemological goals in evolutionary theory” (Love 2017; p. 161) by means of idealizing some features while ignoring or concealing to the background other properties or variables of the systems under scrutiny (see also Love 2013).

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Acknowledgements

The authors acknowledge the advice and assistance of Fátima Sofía Ávila-Cascajares during the conception of Fig. 1, and thank Casandra Lizbeth Méndez-Martínez for her aid in the design of Fig. 1. Diana Martínez Almaguer and Julio César Montero Rojas (Graphic Design Unit, Instituto de Biología, UNAM) assisted in the design of all figures of the article. We also thank Mario Casanueva for discussions and his critical remarks about the structure of the EES. The comments and feedback received from Alan Love and Kevin Laland during the presentation of this work at the 2017 Meeting of the International Society for History, Philosophy and Social Studies of Biology (ISHPSSB) in São Paulo, Brazil, were valuable for the preparation of revised versions of this work; at the same meeting, we also benefited from conversations with Eva Jablonka, Marion Lamb and Jan Baedke about the sociological, epistemological and political dimensions involved in the EES debate. AFT is indebted to participants in the poster session of the Sixth Meeting of the European Society for Evolutionary Developmental Biology (Uppsala, July 2016), especially to Mark Jonas. AFT also thanks Francesco Suman for comments on an earlier version of this manuscript and discussions on the structure of the EES and the pluralist landscape of contemporary evolutionism (Washington D.C., September 2016). FVS acknowledges facilities provided by libraries at the Instituto de Biología, UNAM and other academic institutions in Mexico City, Uppsala and London during his long-term research projects on the historiography, epistemology and sociology of biology. Finally, we thank an anonymous reviewer for critical comments which substantially improved the original manuscript. The authors declare that they have no conflict of interest, and that they did not receive any specific funding during the research and writing of this work.

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Correspondence to Alejandro Fábregas-Tejeda or Francisco Vergara-Silva.

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Fábregas-Tejeda, A., Vergara-Silva, F. The emerging structure of the Extended Evolutionary Synthesis: where does Evo-Devo fit in?. Theory Biosci. 137, 169–184 (2018). https://doi.org/10.1007/s12064-018-0269-2

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Keywords

  • Extended Evolutionary Synthesis
  • Evolutionary biology
  • Paradigm
  • Scientific research program
  • Epistemic units
  • Evo-Devo