Inclusive Fitness Teleology and Darwinian Explanatory Pluralism: A Theoretical Sketch and Application to Current Controversies

Abstract

Formal Darwinism (FD) [Grafen (2002) J Theor Biol 217:75–91; (2007) J Evol Biol 20:1243–1254.] is a theoretical framework for articulating optimization models in behavioral ecology and allele dynamics modeling in population genetics. It yields a teleology centered on inclusive fitness maximization (“IF teleology”), which captures the many aspects of teleology in Darwinian thinking [Huneman (2019b) Stud Hist Philos Sci Part C 76:101188. 10.1016/j.shpsc.2019.101188] and supports an explanatory pluralism in evolutionary biology. Based on this framework, the present chapter intends to show how the major distinctions regarding kinds of explanation identified in evolutionary biology can be connected and systematized through such explanatory pluralism. Then I will show that it can be redescribed in terms of Aristotle’s four causes, and finally, it makes sense of the use of two distinct notions of causation. The rest of the paper analyses two examples where this FD-based pluralism and the correlated use of IF teleology allow one to cast a light on current controversies regarding evolutionary theory: the disputed need to overcome the Modern Synthesis of evolution because of non-genetic inheritance, biased variation, or niche construction; and the opposition of kin selection and multilevel selection regarding the evolution of altruism.

Editorial Note: The Evolution of Meaning—From Neo-Darwinism to Biosemiotics

Richard Theisen Simanke

Huneman’s chapter presents an original, thoughtful, and compelling argument for explanatory pluralism in the biological sciences. Assuming Formal Darwinism as a theoretical framework and focusing on the crucial concept of inclusive fitness maximization, the author argues that this perspective provides a consistent strategy for articulating the different causal pluralisms proposed within the field of evolutionary theory. Aristotle’s theory of causation—arguably the first systematic model of explanatory pluralism in the history of philosophy of science—is called upon as a historical point of comparison, and the author presents his argument as reconstruction and updating of the Aristotelian views, especially on the complementary role played by teleology and mechanism in causation. Underlying the author’s views is the conviction that explanatory pluralism is not an epistemic virtue by itself and that it is not philosophically or scientifically productive unless one provides a formal and conceptual articulation for the different modalities of causation. The chapter still contains a consistent proposal not only to articulate the different kinds of causes within each pluralistic model but also to relate these models to one another.

Since this collection’s general subject matter addresses both evolutionary biology and biosemiotics, it seemed adequate to add here some remarks concerning how an argument for explanatory pluralism and the integration of different theoretical models relates to the biosemiotic view of living beings as meaning-producing entities and natural interpretive systems.

First of all, one must note that Neo-Darwinian evolutionism, however prevalent it may be in the context of contemporary life sciences, is but one of the great paradigms or theoretical models that one can distinguish in this field. Biosemiotics is another model of comparable scope and complexity, with all the doctrinal and methodological implications entailed by such condition. These models foster the emergence of research programs on specific questions in the field of biological knowledge and the formation of research communities endorsing these views of life and the corresponding conceptions of science. Other such models can be mentioned, like those defined by the core concepts of autopoiesis or artificial life.

All these theories organize biological research, with repercussions extending from the more concrete and applied issues to the more general and abstract ones. These more abstract questions touch the borders between biological sciences and the philosophy of biology, including the problem of defining life as such. Many authors (El-Hani 2008; Emmeche 1997) have argued that these paradigms provide the possibility for a situated and circumscribed definition of the very concept of life, relatively to the theoretical model informing the investigation of its phenomena. This strategy could overcome the supposed intractability of defining life, often rejected as a metaphysical and unscientific problem. Thus, from the viewpoint of artificial life, being alive is defined as a property of systems capable of reacting automatically and adaptively, in an open-ended way, to unpredictable changes in their environments (Bedau 1996). For autopoiesis, living beings are organizationally closed but structurally open (both materially and energetically) networks, whose components produce the network itself and its boundaries and are, in turn, recursively produced by them (Maturana and Varela 1980). From a Neo-Darwinian perspective, if one wishes to define life in accordance to the major theoretical principles (which is not something mandatory to do in this framework), life would consist in the property of being self-replicating entities likely to evolve through random variation of inheritable traits and a posteriori natural selection of those traits favoring survival and differential reproduction (Dawkins 1976, 1983). Finally, biosemiotics sees life as meaning-production through interpretation of natural sign-systems or, in Claus Emmeche’s (1998, p. 11) synthetic formulation, as the “functional interpretation of signs in self-organised material code-systems making their own Umwelts” (author’s emphases).

One of these models’ common denominators is that they all recognize the evolution of species, even if departing from the orthodox Neo-Darwinian view of evolution, at least in some of its aspects. Biosemiotics also acknowledges the species transformation over time but changes the focus from the evolution of structures, functions, and behavior to the evolution of systems of signs and the organisms’ interpretation capacities. At least since Terrence Deacon’s works, the integration of evolutionary and semiotic perspectives has come to the foreground of the debate concerning human evolution and its distinctive features (Deacon 1997; Schilhab et al. 2012). In this context, Huneman’s equally integrative approach, even if formulated from within the Darwinian evolutionary framework, can contribute to bringing closer two of the main theoretical models in contemporary biological sciences with their respective conceptions about the fundamental nature of living beings, as seen above.

Particularly significant is Huneman’s recovery and updating of Aristotelian biological philosophy. The rediscovery of Aristotle’s philosophy of nature is a striking feature in contemporary science, especially biology, although not exclusively (Feser 2019; Simpson et al. 2018). Besides providing a representative historical illustration for the kind of explanatory pluralism claimed by Huneman, Aristotelian naturalism makes it possible to consider the teleological dimension of biological explanation in terms compatible with a scientific attitude and thus reconcile two types of explanatory strategies—mechanism and finalism—often regarded as incompatible.

As intentional acts, the symbolic communication and sign interpretation privileged by biosemiotics contain a dimension of intentionality in the phenomenological sense of the term, even if considered natural phenomena (Hoffmeyer 2012). Thus, they require some teleology modality, even if it is not the transcendent finality presupposed by vitalist and metaphysical views of life, evolutionary or otherwise. Again, reference to classical conceptions of life and nature can provide a model to conceive of the immanence of meaning in the lifeworld without simply anthropomorphising it. As Merleau-Ponty remarks in the opening of his courses on nature at the Collège de France, referring to classical Greek thought:

There is nature wherever there is a life that has a meaning but where, however, there is not thought (…). Nature is what has a meaning, without this meaning being posited by thought; it is the autoproduction of a meaning. Nature is thus different from a simple thing. It has an interior, is determined from within (…). Yet nature is different from man; it is not instituted by him (…). (Merleau-Ponty 2003, p. 3)

There are elements in this passage that allow for adding a biosemiotic and autopoietic perspective to the scientific knowledge of nature—and of life, in particular. The evolutionary element—generally absent from classical thought—complements and makes this multidimensional theoretical model more comprehensive. However, if Neo-Darwinian evolutionism must participate in this process, one must trim some of its edges and challenge the necessity of some of its doctrinal commitments. It is to this task that Huneman’s work presented here represents an invaluable contribution.