This dissertation concerns the active role of the organism in evolutionary theory. In particular, it concerns how our conception of the relationship between organism and environment, and the nature of natural selection, influences the causal and explanatory role of organismic activity and behavior in evolutionary explanations. The overarching aim is to argue that the behaviors and activities of organisms can serve both as the explananda (that which is explained) and the explanantia (that which explains) in evolutionary explanations. I attempt to (...) achieve this aim by offering three central arguments. First, that the organism-environment relation is the ontologically basic unit of biology. A common way of conceiving the relationship between organism and environment is as a duality—as two causally independent systems that overlap through interaction. I think this is a mistake. There cannot be organisms without environments and vice versa. They are codependent and codetermined. Second, that natural selection is an ecological process. By this I mean that natural selection acts primarily on organism-environment interactions. It is only in virtue of organisms interacting with their environments that there can be fitness differences amongst individual organisms in a population. The ecological approach to natural selection also entails that the nature of the system(s) of inheritance involved in the reoccurrence of favorable organism-environment interactions is, in principle, immaterial to the process of selection. It only matters for selection that organism-environment interactions actually do reoccur in subsequent generations. Third, that niche construction theory—the most well-developed theoretical and conceptual framework for studying how the activities of organisms influences evolutionary dynamics—can be seen as fully a compatible and integrated part of evolutionary theory. Together, these three arguments constitute an overarching argument. Namely that we both can, and should, take organismic activity and behavior to be crucial explanatory elements in evolutionary theory. Throughout the dissertation I also show how these three arguments have consequences for other debates in the philosophy of evolutionary theory, such as the nature of evolutionary processes, the structure of selection-based explanations, the validity and utility of the proximate-ultimate distinction, and the nature of teleology in evolutionary systems. (shrink)

Traditionally, Thomas S. Kuhn’s The Structure of Scientific Revolutions (1962) is largely identified with his analysis of the structure of scientific revolutions. Here, we contribute to a minority tradition in the Kuhn literature by interpreting the history of evolutionary biology through the prism of the entire historical developmental model of sciences that he elaborates in The Structure. This research not only reveals a certain match between this model and the history of evolutionary biology but, more importantly, also sheds new light (...) on several episodes in that history, and particularly on the publication of Charles Darwin’s On the Origin of Species (1859), the construction of the modern evolutionary synthesis, the chronic discontent with it, and the latest expression of that discontent, called the extended evolutionary synthesis. Lastly, we also explain why this kind of analysis hasn’t been done before. (shrink)

Contemporary evolutionary biology comprises a plural landscape of multiple co-existent conceptual frameworks and strenuous voices that disagree on the nature and scope of evolutionary theory. Since the mid-eighties, some of these conceptual frameworks have denounced the ontologies of the Modern Synthesis and of the updated Standard Theory of Evolution as unfinished or even flawed. In this paper, we analyze and compare two of those conceptual frameworks, namely Niles Eldredge’s Hierarchy Theory of Evolution (with its extended ontology of evolutionary entities) and (...) the Extended Evolutionary Synthesis (with its proposal of an extended ontology of evolutionary processes), in an attempt to map some epistemic bridges (e.g. compatible views of causation; niche construction) and some conceptual rifts (e.g. extra-genetic inheritance; different perspectives on macroevolution; contrasting standpoints held in the “externalism–internalism” debate) that exist between them. This paper seeks to encourage theoretical, philosophical and historiographical discussions about pluralism or the possible unification of contemporary evolutionary biology. (shrink)

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. (shrink)

In this paper, I will conduct three interrelated analyses. First, I will develop an analysis of various concepts in the history of biology that used to refer to individual-level phenomena but were then reinterpreted by the Modern Synthesis in terms of populations. Second, I argue that a similar situation can be found in contemporary biological theory. While different approaches reflect on the causal role of developing organisms in evolution, proponents of the Modern Synthesis avoid any substantial change by reinterpreting and (...) explaining individual-level phenomena from a population perspective. Finally, I will approach this debate by advocating the statistical reading of natural selection, which holds that explanations by natural selection are statistical. I will argue that the above historical conceptual reinterpretations belong to a new explanatory strategy developed by the Modern Synthesis based on population thinking. The reinterpretation of concepts at the individual level is part of the explanatory framework of the Modern Synthesis and the empty role of development within this framework. Moreover, the statistical perspective adopted here allows for the integration of two explanatory models: population-statistical and individual-causal. Finally, I will argue that this pluralistic framework can help to define the explanatory scope of the different biological approaches in order to achieve a coherent integration of development into evolution without rejecting population thinking. (shrink)

In this paper, we analyze the debate between the Modern Synthesis and the Extended Evolutionary Synthesis in light of the concept of incommensurability developed by Thomas Kuhn. In order to do so, first we briefly present both the Modern Synthesis and the Extended Evolutionary Synthesis. Then, we clarify the meaning and interpretations of incommensurability throughout Kuhn’s works, concluding that the version of this concept deployed in The Structure of Scientific Revolutions is the best suited to the analysis of scientific disputes. (...) After discussing incommensurability in Kuhn’s works, we address the question of whether the Modern Synthesis and the Extended Evolutionary Synthesis can be considered semantically, methodologically, and ontologically incommensurable, concluding that they can. Finally, we discuss three problems that arise from such a conclusion: firstly, what are the consequences of incommensurability; secondly, which mode of scientific change better explains this current dispute in evolutionary biology; and thirdly, whether rational theory comparison is possible given incommensurability. We suggest that the main consequence of incommensurability is profound disagreement, that the kind of scientific change that better explains the current dispute between the MS and the EES may be scientific specialization, and that incommensurability does not preclude rational theory comparison. (shrink)

Synthesising arguments motivate changes to the conceptual tools, theoretical structure, and evaluatory framework employed in a given scientific domain. Recently, a broad coalition of researchers has put forward a synthesising argument in favour of an Extended Evolutionary Synthesis (‘EES’). Often this synthesising argument is evaluated using a virtue-based approach, which construes the EES as a wholesale alternative to prevailing practice. Here I argue this virtue-based approach is not fit for purpose. Taking the central concept of niche construction as a case (...) study, I show that an agenda-based approach better captures the pragmatic and epistemological goals of the EES synthesising argument and diagnoses areas of empirical disagreement with prevailing practice. (shrink)

In this article I argue that we should pay extra attention to the ecological dimension of natural selection. By this I mean that we should view natural selection primarily as acting on the outcomes of the interactions organisms have with their environment, which influences their relative reproductive output. A consequence of this view is that natural selection is not sensitive to what system of inheritance ensures reoccurrences of organism-environment interactions over generations. I end by showing the consequences of this view (...) when looking at how processes like niche construction and the Baldwin effect relate to natural selection. (shrink)

Biologists and philosophers of science have recently called for an extension of evolutionary theory. This so-called ‘extended evolutionary synthesis’ seeks to integrate developmental processes, extra-genetic forms of inheritance, and niche construction into evolutionary theory in a central way. While there is often agreement in evolutionary biology over the existence of these phenomena, their explanatory relevance is questioned. Advocates of EES posit that their perspective offers better explanations than those provided by ‘standard evolutionary theory’. Still, why this would be the case (...) is unclear. Usually, such claims assume that EES’s superior explanatory status arises from the pluralist structure of EES, its different problem agenda, and a growing body of evidence for the evolutionary relevance of developmental phenomena. However, what is usually neglected in this debate is a discussion of what the explanatory standards of EES actually are, and how they differ from prevailing standards in SET. In other words, what is considered to be a good explanation in EES versus SET? To answer this question, we present a theoretical framework that evaluates the explanatory power of different evolutionary explanations of the same phenomena. This account is able to identify criteria for why and when evolutionary explanations of EES are better than those of SET. Such evaluations will enable evolutionary biology to find potential grounds for theoretical integration. (shrink)

Kevin Laland and colleagues have put forward a number of arguments motivating an extended evolutionary synthesis. Here I examine Laland et al.'s central concept of reciprocal causation. Reciprocal causation features in many arguments supporting an expanded evolutionary framework, yet few of these arguments are clearly delineated. Here I clarify the concept and make explicit three arguments in which it features. I identify where skeptics can—and are—pushing back against these arguments, and highlight what I see as the empirical, explanatory, and methodological (...) issues at stake. (shrink)

The traditional practice of establishing morphological types and investigating morphological organization has found new support from evolutionary developmental biology (evo-devo), especially with respect to the notion of body plans. Despite recurring claims that typology is at odds with evolutionary thinking, evo-devo offers mechanistic explanations of the evolutionary origin, transformation, and evolvability of morphological organization. In parallel, philosophers have developed non-essentialist conceptions of natural kinds that permit kinds to exhibit variation and undergo change. This not only facilitates a construal of species (...) and higher taxa as natural kinds, but also broadens our perspective on the diversity of kinds found in biology. There are many different natural kinds relevant to the investigative and explanatory aims of evo-devo, including homologues and developmental modules. (shrink)

Traditionally, Thomas S. Kuhn’s The Structure of Scientific Revolutions (1962) is largely identified with his analysis of the structure of scientific revolutions. Here, we contribute to a minority tradition in the Kuhn literature by interpreting the history of evolutionary biology through the prism of the entire historical developmental model of sciences that he elaborates in The Structure. This research not only reveals a certain match between this model and the history of evolutionary biology but, more importantly, also sheds new light (...) on several episodes in that history, and particularly on the publication of Charles Darwin’s On the Origin of Species (1859), the construction of the modern evolutionary synthesis, the chronic discontent with it, and the latest expression of that discontent, called the extended evolutionary synthesis. Lastly, we also explain why this kind of analysis hasn’t been done before. (shrink)

Niche construction theory concerns how organisms can change selection pressures by altering the feature–factor relationship between themselves and their environment. These alterations are standardly understood to be brought about through two kinds of organism–environment interaction: perturbative and relocational niche construction. We argue that a reconceptualization is needed on the grounds that if a niche is understood as the feature–factor relationship, then there are three fundamental ways in which organisms can engage in niche construction: constitutive, relational, and external niche construction. We (...) further motivate our reconceptualization by showing some examples of organismic activities that fall outside of the current categorization of niche construction, but nonetheless should be included. We end by discussing two objections to niche construction and show how our reconceptualization helps to undercut these objections. 1Introduction2Niche Construction Theory 2.1The standard account of niche construction2.2Two problems with the standard account of niche construction3Three Kinds of Niche Construction 3.1Constitutive niche construction3.2Relational niche construction3.3External niche construction4Conceptual Improvement on Niche Construction Theory 4.1Constitutive niche construction4.2Relational niche construction4.3External niche construction5The Status of Niche Construction Theory within Evolutionary Theory 5.1Is niche construction a helpful addition to evolutionary theory?5.2Is niche construction an evolutionary process?6Conclusion. (shrink)

Scientific conflicts often stem from differences in the conceptual framework through which scientists view and understand their own field. In this chapter, I analyze the ontological and methodological assumptions of three traditions in evolutionary biology, namely, Ernst Mayr’s population thinking, the gene-centered view of the Modern Syn thesis, and the Extended Evolutionary Synthesis. Each of these frameworks presupposes a different account of "evolutionary causes," and this discrepancy prevents mutual understanding and objective evaluation in the recent contention surrounding the EES. From (...) this perspective, the chapter characterizes the EES research program as an attempt to introduce causal structures beyond genes as additional units of evolution, and compares its research methodology and objectives with those of the traditional MS framework. (shrink)

Welch :263–279, 2017) has recently proposed two possible explanations for why the field of evolutionary biology is plagued by a steady stream of claims that it needs urgent reform. It is either seriously deficient and incapable of incorporating ideas that are new, relevant and plausible or it is not seriously deficient at all but is prone to attracting discontent and to the championing of ideas that are not very relevant, plausible and/or not really new. He argues for the second explanation. (...) This paper presents a twofold critique of his analysis: firstly, the main calls for reform do not concern the field of evolutionary biology in general but rather, or more specifically, the modern evolutionary synthesis. Secondly, and most importantly, these calls are not only inspired by the factors, enumerated by Welch, but are also, and even primarily, motivated by four problematic characteristics of the modern synthesis. This point is illustrated through a short analysis of the latest reform challenge to the modern synthesis, the so-called extended evolutionary synthesis. We conclude with the suggestion that the modern synthesis should be amended, rather than replaced. (shrink)

A notion of delegated causality is introduced here. This subtle kind of causality is dual to interventional causality. Delegated causality elucidates the causal role of dynamical systems at the “edge of chaos”, explicates evident cases of downward causation, and relates emergent phenomena to Gödel’s incompleteness theorem. Apparently rich implications are noticed in biology and Chinese philosophy. The perspective of delegated causality supports cognitive interpretations of self-organization and evolution.

One of the major benefits of interdisciplinary research is the chance to swap tools between fields, to save having to reinvent the wheel. The fields of language evolution and evolutionary biology have been swapping tools for centuries to the enrichment of both. Here I will discuss three categories of tool swapping: conceptual tools, where analogies are drawn between hypotheses, patterns or processes, so that one field can take advantage of the path cut through the intellectual jungle by the other; theoretical (...) tools, where the machinery developed to process the data in one field is adapted to be applied to the data of the other; and analytical tools, where common problems encountered in both fields can be solved using useful tricks developed by one or the other. I will argue that conceptual tools borrowed from linguistics contributed to the Darwinian revolution in biology; that theoretical tools of evolutionary change can in some cases be applied to both genetic and linguistic data without having to assume the underlying evolutionary processes are exactly the same; and that there are practical problems that have long been recognised in historical linguistics that may be solved by borrowing some useful analytical tools from evolutionary biology. (shrink)