We distinguish dynamical and statistical interpretations of evolutionary theory. We argue that only the statistical interpretation preserves the presumed relation between natural selection and drift. On these grounds we claim that the dynamical conception of evolutionary theory as a theory of forces is mistaken. Selection and drift are not forces. Nor do selection and drift explanations appeal to the (sub-population-level) causes of population level change. Instead they explain by appeal to the statistical structure of populations. We briefly discuss the implications (...) of the statistical interpretation of selection for various debates within the philosophy of biologythe `explananda of selection' debate and the `units of selection' debate. (shrink)

One of the most influential arguments against the claim that computers can think is that while our intentionality is intrinsic, that of computers is derived: it is parasitic on the intentionality of the programmer who designed the computer-program. Daniel Dennett chose a surprising strategy for arguing against this asymmetry: instead of denying that the intentionality of computers is derived, he endeavours to argue that human intentionality is derived too. I intend to examine that biological plausibility of Dennett’s suggestion and show (...) that Dennett’s argument for the claim that human intentionality is derived because it was designed by natural selection is based on the misunderstanding of how natural selection works. (shrink)

In ‘Are biological traits explained by their “selected effect” functions?’, Christie, Brusse, Bourrat, Takacs, and Griffiths argue that selected effect functions only explain the presence of a trait (or the frequency of a trait in a population) in cases in which the selective environment has been uniform, illustrating their point with cases of coevolution, frequency-dependent selection, and bet-hedging. This commentary suggests that selected effect functions are explanatory even in those cases, and that Christie, Brusse, et al. are mistaken about the (...) explanatory aims of the selected effects theory, criticizing it for failing to do something it never set out to do. (shrink)

Explanatory Externalism states that the only adaptive force in evolution is natural selection. Explanatory Externalism is a central thesis of the Modern Evolutionary Synthesis. The etiological theory of natural selected-effect functions also advocates Explanatory Externalism. According to this theory, natural selection is the process responsible for determining the proper natural functions of traits. However, I will point out several challenges to Explanatory Externalism that are proposed primarily by developmental biology and its various subfields. Based on these challenges, this paper will (...) argue why biological functions cannot be fully explained by the selected-effect theory, i.e., that a theory of biological functions must adopt some kind of Explanatory Internalism. At the end of this paper, I will discuss whether or not Explanatory Internalism entails a pluralistic view of biological functions, and how this discussion is related to the different explanations of fit and diversity found in evolutionary biology. (shrink)

This MPhil dissertation presents a novel account of teleological explanations in biology. I outline the “shorthand approach” to such explanations, on which they are taken to convey implicit evolutionary explanations. “Selected effects” accounts of teleological explanation dominate recent literature, but they struggle to accommodate teleological explanations of complex traits built through cumulative selection. I articulate the general notion of a landscape explanation, which, applied to biology, explains the evolution of complex features in a population by citing salient features of the (...) population’s fitness landscape. I show that such explanations lend themselves to a teleological shorthand. I close with remarks concerning when a teleological explanation of a trait is legitimate, and why teleological language strikes us as appropriate when we give evolutionary explanations. (shrink)

This paper analyzes recent attempts to reject reproduction with lineage formation as a necessary condition for evolution by means of natural selection :560–570, 2008; Stud Hist Philos Sci Part C Stud Hist Philos Biol Biomed Sci 42:106–114, 2011; Bourrat in Biol Philos 29:517–538, 2014; Br J Philos Sci 66:883–903, 2015; Charbonneau in Philos Sci 81:727–740, 2014; Doolittle and Inkpen in Proc Natl Acad Sci 115:4006–4014, 2018). Building on the strengths of these attempts and avoiding their pitfalls, it is argued that (...) a robust formulation of evolution by natural selection without reproduction can be established. The main contribution of this paper is a reformulation of Lewontin’s three principles stating that minimal evolution by natural selection occurs when two conditions are met in a population: fitness-related variation and memory. Paradigmatic evolution by natural selection, which can generate adaptations, takes place when an additional condition is present, namely regeneration. (shrink)

I distinguish two roles for a fitness concept in the context of explaining cumulative adaptive evolution: fitness as a predictor of gene frequency change, and fitness as a criterion for phenotypic improvement. Critics of inclusive fitness argue, correctly, that it is not an ideal fitness concept for the purpose of predicting gene-frequency change, since it relies on assumptions about the causal structure of social interaction that are unlikely to be exactly true in real populations, and that hold as approximations only (...) given a specific type of weak selection. However, Hamilton took this type of weak selection, on independent grounds, to be responsible for cumulative assembly of complex adaptations. In this special context, I argue that inclusive fitness is distinctively valuable as a criterion for improvement and a standard for optimality. Yet to call inclusive fitness a criterion for improvement and a standard for optimality is not to make any claim about the frequency with which inclusive fitness optimization actually occurs in nature. This is an empirical question that cannot be settled by theory alone. I close with some reflections on the place of inclusive fitness in the long running clash between ‘causalist’ and ‘statisticalist’ conceptions of fitness. (shrink)

This PhD dissertation examines the conceptual and theoretical foundations of the most general and most widely used framework for understanding social evolution, W. D. Hamilton's theory of kin selection. While the core idea is intuitive enough (when organisms share genes, they sometimes have an evolutionary incentive to help one another), its apparent simplicity masks a host of conceptual subtleties, and the theory has proved a perennial source of controversy in evolutionary biology. To move towards a resolution of these controversies, we (...) need a careful and rigorous analysis of the philosophical foundations of the theory. My aim in this work is to provide such an analysis. I begin with an examination of the concepts behavioural ecologists employ to describe and classify types of social behaviour. I stress the need to distinguish concepts that are often conflated: for example, we need to distinguish simple cooperation from collaboration in collective tasks, behaviours from strategies, and control from manipulation and coercion. I proceed from here to the formal representation of kin selection via George R. Price’s covariance selection mathematics. I address a number of interpretative issues the Price formalism raises, including the vexed question of whether kin selection theory is ‘formally equivalent’ to multi-level selection theory. In the second half of the dissertation, I assess the uses and limits of Hamilton’s rule for the evolution of social behaviour; I provide a precise statement of the conditions under which the rival neighbour-modulated fitness and inclusive fitness approaches in contemporary kin selection theory are equivalent (and describe cases in which they are not); and I criticize recent formal attempts to establish the controversial claim that kin selection leads to organisms behaving as if maximizing their inclusive fitness. (shrink)

It is commonly thought that natural selection explains the rise of adaptive complexity. Razeto-Barry and Frick have recently argued in favour of this view, dubbing it the Creative View. I argue that the Creative View is mistaken if it claims that natural selection serves to answer Paley’s question. This is shown by a case that brings out the contrastive structure inherent in this demand for explanation. There is, however, a rather trivial sense in which specific environmental conditions are crucial for (...) the rise of specific adaptations, but this is hardly what opponents of the Creative View are denying. (shrink)

The canonical version of the history of twentieth century philosophy of science tells us that Lakatos was Popper’s disciple, but it is rarely mentioned that Popper would have learned anything from Lakatos. The aim of this paper is to examine Lakatos’ influence on Popper’s philosophical system and to argue that Lakatos did have an important, yet somewhat unexpected, impact on Popper’s thinking: he influenced Popper’s evolutionary model for ‘progress’ in science. And Lakatos’ influence sheds new light on why and how (...) Popper continually revised one of the central claims of his philosophy of science: the evolutionary account of scientific theory change. (shrink)

The explanatory role of natural selection is one of the long-term debates in evolutionary biology. Nevertheless, the consensus has been slippery because conceptual confusions and the absence of a unified, formal causal model that integrates different explanatory scopes of natural selection. In this study we attempt to examine two questions: (i) What can the theory of natural selection explain? and (ii) Is there a causal or explanatory model that integrates all natural selection explananda? For the first question, we argue that (...) five explananda have been assigned to the theory of natural selection and that four of them may be actually considered explananda of natural selection. For the second question, we claim that a probabilistic conception of causality and the statistical relevance concept of explanation are both good models for understanding the explanatory role of natural selection. We review the biological and philosophical disputes about the explanatory role of natural selection and formalize some explananda in probabilistic terms using classical results from population genetics. Most of these explananda have been discussed in philosophical terms but some of them have been mixed up and confused. We analyze and set the limits of these problems. (shrink)

Many moral philosophers accept the Debunking Thesis, according to which facts about natural selection provide debunking explanations for certain of our moral beliefs. I argue that philosophers who accept the Debunking Thesis beg important questions in the philosophy of biology. They assume that past selection can explain why you or I hold certain of the moral beliefs we do. A position advanced by many prominent philosophers of biology implies that this assumption is false. According to the Negative View, natural selection (...) cannot explain the traits of individuals. Hence, facts about past selection cannot provide debunking explanations for any of our moral beliefs. The aim of this paper is to explore the conflict between the Debunking Thesis and the Negative View. (shrink)

A long-running dispute concerns which adaptation-related explananda natural selection can be said to explain. At issue are explananda of the form: why a given individual organism has a given adaptation rather than that same individual having another trait. It is broadly agreed that one must be ready to back up a “no” answer with an appropriate theory of trans-world identity for individuals. I argue, against the conventional wisdom, that the same is true for a “yes” answer. My conclusion recasts the (...) landscape and opens the door to a potential resolution. (shrink)

An influential argument due to Elliott Sober, subsequently strengthened by Denis Walsh and Joel Pust, moves from plausible premises to the bold conclusion that natural selection cannot explain the traits of individual organisms. If the argument were sound, the explanatory scope of selection would depend, surprisingly, on metaphysical considerations concerning origin essentialism. I show that the Sober-Walsh-Pust argument rests on a flawed counterfactual criterion for explanatory relevance. I further show that a more defensible criterion for explanatory relevance recently proposed by (...) Michael Strevens lends support to the view that natural selection can be relevant to the explanation of individual traits. (shrink)

The concept of population thinking was introduced by Ernst Mayr as the right way of thinking about the biological domain, but it is difficult to find an interpretation of this notion that is both unproblematic and does the theoretical work it was intended to do. I argue that, properly conceived, Mayr’s population thinking is a version of trope nominalism: the view that biological property-types do not exist or at least they play no explanatory role. Further, although population thinking has been (...) traditionally used to argue against essentialism about biological kinds, recently it has been suggested that it may be consistent with at least some forms of essentialism—ones that construe essential properties as relational. I argue that if population thinking is a version of trope nominalism, then, as Mayr originally claimed, it rules out any version of essentialism about biological kinds. (shrink)

I have recently argued that origin essentialism regarding individual organisms entails that natural selection does not explain why individual organisms have the traits that they do. This paper defends this and related theses against Mohan Matthen's recent objections.

Spencer’s heritage, while almost a forgotten chapter in the history of biology, lives on in psychology and the philosophy of mind. I particularly discuss externalist views of meaning, on which meaning crucially depends on a notion of reference, and ask whether reference should be thought of as cause or effect. Is the meaning of a word explained by what it refers to, or should we say that what we use a word to refer to is explained by what concept it (...) expresses? I argue for the latter view, which I call ‘Darwinian’, and against the former, ‘Spencerian’ one, assuming conceptual structures in humans to be an instance of adaptive structures, and adaptive relations to an environment to be the effect rather than the cause of evolutionary novelties. I conclude with the deficiency – both empirically and methodologically – of a functionalist study of human concepts and the languages they are embedded in, as it would be undertaken in a paradigm that identifies meaning with reference or that gives reference an explanatory role to play for what concepts we have. (shrink)

The extensive philosophical discussions and analyses in recent decades of function-talk in biology have done much to clarify what biologists mean when they ascribe functions to traits, but the basic metaphysical question—is there genuine teleology and design in the natural world, or only the appearance of this?—has persisted, as recent work both defending, and attacking, teleology from a Darwinian perspective, attest. I argue that in the context of standard contemporary evolutionary theory, this is for the most part a verbal, rather (...) than a substantive dispute: the disputants are talking past one another. To justify this claim I develop a general framework within which reductionist views, such as the standard ‘etiological’ account of biofunctions, occupy an intermediate position between what I call full-blooded realism and full-blooded anti-realism, and suggest that whether such views count as ‘realist’ views has no objective, theory-neutral answer. (shrink)

This article defends the Negative View of natural selection explanation, according to which natural selection cannot explain of any given individual why it has the traits it does. Over the years, this view has been criticized on empirical, metaphysical, and explanatory grounds. I review the debate and offer additional reasons for rejecting the empirical and metaphysical objections. The explanatory objection, which holds that the Negative View is rooted in a flawed account of contrastive explanation, initially seems plausible. However, I argue (...) that a closer consideration of the desiderata of contrastive explanation shows that this objection fails as well. (shrink)

Can selection explain why individuals have the traits they do? This question has generated significant controversy. I will argue that the debate encompasses two separable aspects, to detrimental effect: (1) the role of selection in explaining the origin and evolution of biological traits and (2) the implications this may have for explaining why individuals have the traits they do. (1) can be settled on the basis of evolutionary theory while (2) requires additional, extra-scientific assumptions. By making a distinction between traits (...) affected by a single factor and traits affected by multiple factors I show that selection can, under certain conditions, help explain the origin of traits. Resolving the first aspect enables us to critically assess the various incompatible and independent philosophical commitments made within the second aspect of the debate. (shrink)

There are ongoing debates in philosophy of biology about what falls within natural selection's explanatory scope. These include debates about whether selection can explain individual-level traits, the extent to which selection can explain distributions of trait frequencies, and whether selection can explain the origin of novel traits. Here I'll survey these debates, suggest which views seem most plausible, and describe some useful conceptual frameworks for thinking about the issues involved.

The notion of biological function is fraught with difficulties—intrinsically and irremediably so, we argue. The physiological practice of functional ascription originates from a time when organisms were thought to be designed and remained largely unchanged since. In a secularized worldview, this creates a paradox which accounts of functions as selected effect attempt to resolve. This attempt, we argue, misses its target in physiology and it brings problems of its own. Instead, we propose that a better solution to the conundrum of (...) biological functions is to abandon the notion altogether, a prospect not only less daunting than it appears, but arguably the natural continuation of the naturalisation of biology. (shrink)

There is a substantial literature on evolutionary debunking arguments (EDAs) in metaethics. According to these arguments, evolutionary explanations of our moral beliefs pose a significant problem for moral realism, specifically by committing the realist to an unattractive pessimism about the prospects of our having moral knowledge. In this paper, I argue that EDAs exploit an equivocation between two distinct readings of their central claim. One is plausibly true but has no epistemic relevance, and the other would have epistemic consequences for (...) realism, but is false. If I'm right, this undermines attempts to use evolutionary explanations to debunk belief in other domains too. (shrink)

‘Natural selection’ is, it seems, an ambiguous term. It is sometimes held to denote a consequence of variation, heredity, and environment, while at other times as denoting a force that creates adaptations. I argue that the latter, the force interpretation, is a redundant notion of natural selection. I will point to difficulties in making sense of this linguistic practise, and argue that it is frequently at odds with standard interpretations of evolutionary theory. I provide examples to show this; one example (...) involving the relation between adaptations and other traits, and a second involving the relation between selection and drift. (shrink)

Ruth Millikan and others advocate theories which attempt to naturalize wide mental content (e.g. beliefs.

There is still a great deal of debate over what counts as a constraint and about how to assess experimentally the relative importance of constraints and selection in evolutionary history. I will argue that the notion of a constraint on variation, and thus the selection-constraint distinction, depends on two specifications: (1) what counts as a variant -- constraints limit or bias the production of what? and (2) range of assessment -- over what range of times or conditions is the variation (...) assessed? Specifications 1 and 2 help us to understand empirical work on the relative importance of constraint and selection in evolution. (shrink)

It is commonly thought that natural selection explains the rise of adaptive complexity. Razeto-Barry and Frick have recently argued in favour of this view, dubbing it the Creative View. I argue that the Creative View is mistaken if it claims that natural selection serves to answer Paley’s question. This is shown by a case that brings out the contrastive structure inherent in this demand for explanation. There is, however, a rather trivial sense in which specific environmental conditions are crucial for (...) the rise of specific adaptations, but this is hardly what opponents of the Creative View are denying. (shrink)

Darwin's theory of evolution by natural selection provided the first, and only, causal-mechanistic account of the existence of adaptations in nature. As such, it provided the first, and only, scientific alternative to the “argument from design”. That alone would account for its philosophical significance. But the theory also raises other philosophical questions not encountered in the study of the theories of physics. Unfortunately the concept of natural selection is intimately intertwined with the other basic concepts of evolutionary theory—such as the (...) concepts of fitness and adaptation —that are themselves philosophically controversial. Fortunately we can make considerable headway in getting clear on natural selection without solving all of those outstanding problems. (shrink)

In this paper, using a multilevel approach, we defend the positive role of natural selection in the generation of organismal form. Despite the currently widespread opinion that natural selection only plays a negative role in the evolution of form, we argue, in contrast, that the Darwinian factor is a crucial (but not exclusive) factor in morphological organization. Analyzing some classic arguments, we propose incorporating the notion of ‘downward causation’ into the concept of ‘natural selection.’ In our opinion, this kind of (...) causation is fundamental to the operation of selection as a creative evolutionary process. (shrink)

This is the second of a two-part essay on the history of debates concerning the creativity of natural selection, from Darwin through the evolutionary synthesis and up to the present. In the first part, I focussed on the mid-late nineteenth century to the early twentieth, with special emphasis on early Darwinism and its critics, the self-styled “mutationists.” The second part focuses on the evolutionary synthesis and some of its critics, especially the “neutralists” and “neo-mutationists.” Like Stephen Gould, I consider the (...) creativity of natural selection to be a key component of what has traditionally counted as “Darwinism.” I argue that the creativity of natural selection is best understood in terms of selection initiating evolutionary change, and selection directing evolutionary change, for example by creating the variation that it subsequently acts upon. I consider the respects in which both of these claims sound non-Darwinian, even though they have long been understood by supporters and critics alike to be virtually constitutive of Darwinism. (shrink)

The aim of this paper is to outline a typologyof selection processes, and show that differentsub-categories have different explanatorypower. The basis of this typology of selectionprocesses is argued to be the difference ofreplication processes involved in them. Inorder to show this, I argue that: 1.Replication is necessary for selection and 2.Different types of replication lead todifferent types of selection. Finally, it isargued that this typology is philosophicallysignificant, since it contrasts cases ofselection (on the basis of the replicationprocesses involved in them) (...) whereby selectioncauses adaptation – and, therefore, can beused in explanations of the (real or apparent)teleology of Nature – and cases in whichselection lacks such explanatory power. (shrink)