The notion that natural selection is a process of fitness maximization gets a bad press in population genetics, yet in other areas of biology the view that organisms behave as if attempting to maximize their fitness remains widespread. Here I critically appraise the prospects for reconciliation. I first distinguish four varieties of fitness maximization. I then examine two recent developments that may appear to vindicate at least one of these varieties. The first is the ‘new’ interpretation of Fisher's fundamental theorem (...) of natural selection, on which the theorem is exactly true for any evolving population that satisfies some minimal assumptions. The second is the Formal Darwinism project, which forges links between gene frequency change and optimal strategy choice. In both cases, I argue that the results fail to establish a biologically significant maximization principle. I conclude that it may be a mistake to look for universal maximization principles justified by theory alone. A more promising approach may be to find maximization principles that apply conditionally and to show that the conditions were satisfied in the evolution of particular traits. (shrink)

The propensity interpretation of fitness draws on the propensity interpretation of probability, but advocates of the former have not attended sufficiently to problems with the latter. The causal power of C to bring about E is not well-represented by the conditional probability Pr. Since the viability fitness of trait T is the conditional probability Pr, the viability fitness of the trait does not represent the degree to which having the trait causally promotes surviving. The same point holds for fertility fitness. (...) This failure of trait fitness to capture causal role can also be seen in the fact that coextensive traits must have the same fitness values even if one of them promotes survival and the other is neutral or deleterious. Although the fitness of a trait does not represent the trait’s causal power to promote survival and reproduction, variation in fitness in a population causally promotes change in trait frequencies; in this sense, fitness variation is a population-level propensity. (shrink)

Abstract Gene-selectionists define fundamental terms in non-standard ways. Genes are determinants of difference. Phenotypes are defined as a gene’s effects relative to some alternative whereas the environment is defined as all parts of the world that are shared by the alternatives being compared. Environments choose among phenotypes and thereby choose among genes. By this process, successful gene sequences become stores of information about what works in the environment. The strategic gene is defined as a set of gene tokens that combines (...) ‘actor’ tokens responsible for an effect with ‘recipient’ tokens whose replication is thereby enhanced. This set of tokens can extend across the boundaries of individual organisms, or other levels of selection, as these are traditionally defined. Content Type Journal Article Pages 1-19 DOI 10.1007/s10539-012-9315-5 Authors David Haig, Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA Journal Biology and Philosophy Online ISSN 1572-8404 Print ISSN 0169-3867. (shrink)

This essay examines the claim “path dependence entails irreversibility” from the point of view of evolutionary biology. I argue that evolutionary irreversibility possesses many faces, sometimes conflicting with path dependence. I propose an account of path dependence that does not rely on irreversibility and explains why it more naturally coexists with the notion of (contingent) irreversibility developed by the Belgian paleontologist Louis Dollo. However, I argue that we should not conceive of this relationship as necessary.

Wright’s “adaptive landscape” has been influential in evolutionary thinking but controversial, especially because the landscape that organisms encounter is altered by the evolutionary process itsel...

The aim of this paper is to offer a new biological interpretation of Fisher’s ‘Fundamental Theorem of Natural Selection’ and from this to consider optimality properties of gene frequency changes. These matters are of continuing interest to biologists and philosophers alike. In particular, the extent to which biological evolution can be calculated from the ‘gene’s-eye’ point of view is also discussed. In this sense, the paper bears indirectly on the concepts of the unit of selection and of the ‘selfish gene’. (...) A new biological significance for the Fundamental Theorem, not previously found in the literature, is offered, together with an optimality principle connected with this theorem. (shrink)

In this paper I critically evaluate Reisman and Forber’s :1113–1123, 2005) arguments that drift and natural selection are population-level causes of evolution based on what they call the manipulation condition. Although I agree that this condition is an important step for identifying causes for evolutionary change, it is insufficient. Following Woodward, I argue that the invariance of a relationship is another crucial parameter to take into consideration for causal explanations. Starting from Reisman and Forber’s example on drift and after having (...) briefly presented the criterion of invariance, I show that once both the manipulation condition and the criterion of invariance are taken into account, drift, in this example, should better be understood as an individual-level rather than a population-level cause. Later, I concede that it is legitimate to interpret natural selection and drift as population-level causes when they rely on genuinely indeterministic events and some cases of frequency-dependent selection. (shrink)

This article is a systematic critical survey of work done in the philosophy of biology within the logical empiricist tradition, beginning in the 1930s and until the end of the 1950s. It challenges a popular view that the logical empiricists either ignored biology altogether or produced analyses of little value. The earliest work on the philosophy of biology within the logical empiricist corpus was that of Philipp Frank, Ludwig von Bertalanffy, and Felix Mainx. Mainx, in particular, provided a detailed analysis (...) of biology in the 1930s and 1940s in his contribution to the logical empiricists’ Encyclopedia of Unified Science. However, the most important contributions to the philosophy of biology were those of Joseph Henry Woodger and Ernest Nagel. Woodger is primarily remembered for deploying the axiomatic method in biology but he also used semiformal methods for the analysis of many biological problems. While Woodger’s axiomatic work was often derided by some later philosophers of biology (e.g., David Hull and Michael Ruse), this article defends both the biological and the philosophical significance of some of that work, for instance, those aspects that led to the recognition of the conceptual complexity of mereology and temporal identity in biological systems. Woodger’s semiformal analyses were even more important, for instance, his explication of the concepts of the Bauplan and of innateness. Nagel’s importance lies in his analyses of reduction and emergence in the context of all empirical sciences and his use of these analyses in a careful exploration of biological problems. While Nagel’s model of reduction was generally rejected by philosophers of science in the 1970s and 1980s, particularly for biological contexts, it has recently been sympathetically reconstructed by many commentators; this article defends its continued relevance for the philosophy of biology. (shrink)

Is there any way to reconcile the adaptationist’s image of natural selection as an engine of optimality with the more complex image of its dynamics we get from population genetics? This has long been an important strand in the controversy surrounding adaptationism, yet debate has been hampered by a tendency to conflate various different ways of thinking about maximization. Here I distinguish four varieties of maximization principle. I then discuss the logical relations between these varieties, arguing that, although they may (...) seem similar at face value, none entails any of the others. I then turn briefly to the status of each variety, arguing that, while each type of maximization principle faces serious problems, the problems are subtly different for each type. In the last section, I reflect on what is at stake in the debate. (shrink)

The semantic view of theories is normally considered to be an ac-count of theories congenial to Scientific Realism. Recently, it has been argued that Ontic Structural Realism could be fruitfully applied, in combination with the semantic view, to some of the philosophical issues peculiarly related to bi-ology. Given the central role that models have in the semantic view, and the relevance that mathematics has in the definition of the concept of model, the fo-cus will be on population genetics, which is (...) one of the most mathematized areas in biology. We will analyse some of the difficulties which arise when trying to use Ontic Structural Realism to account for evolutionary biology. (shrink)

Selectionist evolutionary theory has often been faulted for not making novel predictions that are surprising, risky, and correct. I argue that it in fact exhibits the theoretical virtue of predictive capacity in addition to two other virtues: explanatory unification and model fitting. Two case studies show the predictive capacity of selectionist evolutionary theory: parallel evolutionary change in E. coli, and the origin of eukaryotic cells through endosymbiosis.

This article proposes two conditions to assess whether an entity at a level of description is a unit of selection qua interactor. These two conditions make it possible to (1) distinguish biologically relevant entities from arbitrary ones and (2) distinguish units that can _potentially_ enter a selection process from those that have already done so. I show that the classical approaches used in the literature on units and levels of selection do not fare well with respect to either or both (...) of these desiderata. (shrink)

In this paper I propose a new interpretation of the British evolutionary synthesis. The synthetic work of J. B. S. Haldane, R. A. Fisher and J. S. Huxley was characterized by both an integration of Mendelism and Darwinism and the unification of different biological subdisciplines within a coherent framework. But it must also be seen as a bold and synthetic Darwinian program in which the biosciences served as a utopian blueprint for the progress of civilization. Describing the futuristic visions of (...) these three scientists in their synthetic heydays, I show that, despite a number of important divergences, their biopolitical ideals could be biased toward a controlled and regimented utopian society. Their common ideals entailed a social order where liberal and democratic principles were partially or totally suspended in favor of bioscientific control and planning for the future. Finally, I will argue that the original redefinition of Darwinism that modern synthesizers proposed is a significant historical example of how Darwinism has been used and adapted in different contexts. The lesson I draw from this account is a venerable one: that, whenever we wish to define Darwinism, we need to recognize not only its scientific content and achievements but expose the other traditions and ideologies it may have supported. (shrink)

Although classical evolutionary theory, i.e., population genetics and the Modern Synthesis, was already implicitly ‘gene-centred’, the organism was, in practice, still generally regarded as the individual unit of which a population is composed. The gene-centred approach to evolution only reached a logical conclusion with the advent of the gene-selectionist or gene’s eye view in the 1960s and 1970s. Whereas classical evolutionary theory can only work with (genotypically represented) fitness differences between individual organisms, gene-selectionism is capable of working with fitness differences (...) among genes within the same organism and genome. Here, we explore the explanatory potential of ‘intra-organismic’ and ‘intra-genomic’ gene-selectionism, i.e., of a behavioural-ecological ‘gene’s eye view’ on genetic, genomic and organismal evolution. First, we give a general outline of the framework and how it complements the—to some extent—still ‘organism-centred’ approach of classical evolutionary theory. Secondly, we give a more in-depth assessment of its explanatory potential for biological evolution, i.e., for Darwin’s ‘common descent with modification’ or, more specifically, for ‘historical continuity or homology with modular evolutionary change’ as it has been studied by evolutionary developmental biology (evo-devo) during the last few decades. In contrast with classical evolutionary theory, evo-devo focuses on ‘within-organism’ developmental processes. Given the capacity of gene-selectionism to adopt an intra-organismal gene’s eye view, we outline the relevance of the latter model for evo-devo. Overall, we aim for the conceptual integration between the gene’s eye view on the one hand, and more organism-centred evolutionary models (both classical evolutionary theory and evo-devo) on the other. (shrink)

Frank has recently argued for an information-theoretic interpretation of natural selection. This interpretation is based on the identification of a measure related to the Malthusian parameter (for population change) with the Jeffreys divergence between the present allelic distribution of the population and that distribution in the next generation. It is pointed out in this analysis that this identification only holds if the mean fitness of the population is a constant, that is, there is no selection. This problem is used to (...) argue for the superiority of the standard dynamical interpretation of natural selection over its information-theoretic counterpart. (shrink)

This paper contains replies to the reviews of my book by Steven Downes, Massimo Pigliucci and Deborah Shelton & Rick Michod.

Although classical evolutionary theory, i.e., population genetics and the Modern Synthesis, was already implicitly ‘gene-centred’, the organism was, in practice, still generally regarded as the individual unit of which a population is composed. The gene-centred approach to evolution only reached a logical conclusion with the advent of the gene-selectionist or gene’s eye view in the 1960s and 1970s. Whereas classical evolutionary theory can only work with fitness differences between individual organisms, gene-selectionism is capable of working with fitness differences among genes (...) within the same organism and genome. Here, we explore the explanatory potential of ‘intra-organismic’ and ‘intra-genomic’ gene-selectionism, i.e., of a behavioural-ecological ‘gene’s eye view’ on genetic, genomic and organismal evolution. First, we give a general outline of the framework and how it complements the—to some extent—still ‘organism-centred’ approach of classical evolutionary theory. Secondly, we give a more in-depth assessment of its explanatory potential for biological evolution, i.e., for Darwin’s ‘common descent with modification’ or, more specifically, for ‘historical continuity or homology with modular evolutionary change’ as it has been studied by evolutionary developmental biology during the last few decades. In contrast with classical evolutionary theory, evo-devo focuses on ‘within-organism’ developmental processes. Given the capacity of gene-selectionism to adopt an intra-organismal gene’s eye view, we outline the relevance of the latter model for evo-devo. Overall, we aim for the conceptual integration between the gene’s eye view on the one hand, and more organism-centred evolutionary models on the other. (shrink)

The idea that natural selection can operate on cultural as well as genetic variation is central to recent theories of cultural evolution. This raises an overarching question: how much of traditional evolutionary theory, which was formulated in population-genetic terms, can survive intact once the possibility of cultural inheritance is taken into account? This question is addressed in relation to R. A. Fisher’s “fundamental theorem” of natural selection. Though Fisher’s theorem may appear to be an essentially genetic result, a version of (...) the theorem can be derived that applies to any evolving population, with an arbitrary inheritance mechanism. However, I argue that the biological significance of this “generalized” version of the fundamental theorem is less than that of the Fisherian original. (shrink)

I examine the consistency between contemporary environmentalist ideals and Buddhist philosophy, focusing, first, on the problem of value in nature. I argue that the teachings found in the Pāli canon cannot easily be reconciled with a belief in the intrinsic value of life, whether human or otherwise. This is because all existence is regarded as inherently unsatisfactory, and all beings are seen as impermanent and insubstantial, while the ultimate spiritual goal is often viewed, in early Buddhism, as involving a deep (...) renunciation of the world. Therefore, the discussion focuses mostly on the Mahāyāna vehicle, which, I suggest has better resources for environmentalism because enlightenment and the ordinary world are not conceived as antithetical. Still, many contemporary green ideas do not sit well with classical Mahāyāna doctrines. Mahāyāna philosophers coincide in equating ultimate reality with ‘emptiness,’ and propose knowledge of this reality as a final soteriological purpose. Emptiness is generally said to be ineffable, and to involve the negation of all views. An important question is how to reconcile environmentalism with the relinquishing of views. I consider several prevalent themes in environmentalism, including the philosophy of ‘Oneness,’ and other systems that are often compared with Buddhism, like process thought. Many of these turn out to have more in common with an extreme view that Buddhism seeks to avoid, namely, eternalism. I attempt to outline an environmental position that, like the doctrine of emptiness, traverses a Middle Path between eternalism and nihilism. I conclude by proposing that emptiness could be regarded as the source of value in nature, if it is seen in its more positive aspect, as ‘pliancy.’ This would imply that what Buddhist environmentalists should seek to protect is not any being in its current form, nor any static natural system, but the possibility of adaptation and further evolution. (shrink)

The scientific status of evolutionary theory seems to be more or less perennially under question. I am not referring here (just) to the silliness of young Earth creation- ism (Pigliucci 2002; Boudry and Braeckman 2010), or even of the barely more intel- lectually sophisticated so-called Intelligent Design theory (Recker 2010; Brigandt this volume), but rather to discussions among scientists and philosophers of science concerning the epistemic status of evolutionary theory (Sober 2010). As we shall see in what follows, this debate (...) has a long history, dating all the way back to Darwin, and it is in great part rooted in the fundamental dichotomy between what French biologist and Nobel laureate Jacques Monod (1971) called chance and necessity—i.e., the inevitable and inextricable interplay of deterministic and stochastic mechanisms operating during the course of evolution. (shrink)

The relationship between statistical mechanics and population genetics has a long history. Both take advantage of statistics to address the behavior of large groups of entities. The main objective of this article is to assess the obstacles population genetics is meeting in its claim to explain biological phenomena from the conceptual apparatus of statistical mechanics according to two recent articles. Several tools available to the latter are missing in the former. Thus, in the absence of an adequate justification of the (...) relevance of the analogy and without the satisfaction of certain basic assumptions, I argue that the explanatory strategy of the analogously based model is seriously compromised. La relation entre mécanique statistique et génétique des populations a une longue histoire. À l’instar de la première, la seconde tire profit des statistiques afin de traiter le comportement des grands ensembles d’entités. L’objectif principal du présent article est d’identifier les obstacles que rencontre la génétique des populations dans sa prétention d’explication des phénomènes biologiques à partir de l’appareillage conceptuel de la mécanique statistique selon deux articles récents. Or, plusieurs outils dont dispose cette dernière manque à la première. Ainsi, en l’absence d’une justification adéquate de la pertinence de l’analogie et sans la satisfaction de certaines hypothèses fondamentales, je soutiens que la stratégie explicative du modèle analogue est sérieusement compromise. (shrink)

A number of areas of biology raise questions about what is of value in the natural environment and how we ought to behave towards it: conservation biology, environmental science, and ecology, to name a few. Based on my experience teaching students from these and similar majors, I argue that the field of environmental ethics has much to teach these students. They come to me with pent-up questions and a feeling that more is needed to fully engage in their subjects, and (...) I believe some exposure to environmental ethics can help focus their interests and goals. I identify three primary areas in which environmental ethics can con- tribute to their education. The first is an examination of who (or what) should be considered to be part of our moral community (i.e., the community to whom we owe direct duties). Is it humans only? Or does it include all sentient life? Or all life? Or ecosystems considered holistically? Often, readings implicitly assume one or more of these answers; the goal is to make the student more sensitive to these implicit claims and to get them to think about the different reasons that support them. The second area, related to the first, is the application of the different answers concerning the extent of the ethical community to real environmental issues and problems. Students need to be aware of how the different answers concerning the moral community can imply conflicting answers for how we should act in certain cases and to think about ways to move toward conflict resolution. The third area in which environmental ethics can contribute is a more conceptual one, focusing on central concepts such as biodiversity, sustainability, species, and ecosystems. Exploring and evaluating various meanings of these terms will make students more reflective and thoughtful citizens and biologists, sensitive to the implications that different conceptual choices make. (shrink)