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Switch to: References
Citations of:
The Confusions of Fitness
British Journal for the Philosophy of Science 55 (2):347-363 (2004)
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I argue that the propensity interpretation of fitness, properly understood, not only solves the explanatory circularity problem and the mismatch problem, but can also withstand the Pandora’s box full of problems that have been thrown at it. Fitness is the propensity (i.e., probabilistic ability, based on heritable physical traits) for organisms or types of organisms to survive and reproduce in particular environments and in particular populations for a specified number of generations; if greater than one generation, “reproduction” includes descendants of (...) |
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Since the introduction of mathematical population genetics, its machinery has shaped our fundamental understanding of natural selection. Selection is taken to occur when differential fitnesses produce differential rates of reproductive success, where fitnesses are understood as parameters in a population genetics model. To understand selection is to understand what these parameter values measure and how differences in them lead to frequency changes. I argue that this traditional view is mistaken. The descriptions of natural selection rendered by population genetics models are (...) |
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The most compelling extant accounts of explanation casts all explanations as causal. Yet there are sciences, theoretical population biology in particular, that explain their phenomena by appeal to statistical, non-causal properties of ensembles. I develop a generalised account of explanation. An explanation serves two functions: metaphysical and cognitive. The metaphysical function is discharged by identifying a counterfactually robust invariance relation between explanans event and explanandum. The cognitive function is discharged by providing an appropriate description of this relation. I offer examples (...) |
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Over the past fifteen years there has been a considerable amount of debate concerning what theoretical population dynamic models tell us about the nature of natural selection and drift. On the causal interpretation, these models describe the causes of population change. On the statistical interpretation, the models of population dynamics models specify statistical parameters that explain, predict, and quantify changes in population structure, without identifying the causes of those changes. Selection and drift are part of a statistical description of population (...) |
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The niche is allegedly the conceptual bedrock underpinning the most prominent, and some would say most important, theorizing in ecology. We argue this point of view is more aspirational than veridical. Rather than critically dissect existing definitions of the concept, the supposedly significant work it is thought to have done in ecology is our evaluative target. There is no denying the impressive mathematical sophistication and theoretical ingenuity of the ecological modeling that invokes ‘niche’ terminology. But despite the pervasive labeling, we (...) |
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The most consistent definition of fitness makes it a static property of organisms. However, this is not how fitness is used in many evolutionary models. In those models, fitness is permitted to vary with an organism’s circumstances. According to this second conception, fitness is dynamic. There is consequently tension between these two conceptions of fitness. One recently proposed solution suggests resorting to conditional properties. We argue, however, that this solution is unsatisfactory. Using a very simple model, we show that it (...) |
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Bourrat and Griffiths :33, 2018) have recently argued that most of the evidence presented by holobiont defenders to support the thesis that holobionts are evolutionary individuals is not to the point and is not even adequate to discriminate multispecies evolutionary individuals from other multispecies assemblages that would not be considered evolutionary individuals by most holobiont defenders. They further argue that an adequate criterion to distinguish the two categories is fitness alignment, presenting the notion of fitness boundedness as a criterion that (...) |
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The propensity nature of evolutionary fitness has long been appreciated and is nowadays amply discussed. The discussion has, however, on occasion followed long standing conflations in the philosophy of probability literature between propensities, probabilities, and frequencies. In this paper, I apply a more recent conception of propensities in modelling practice to some of the key issues, regarding the mathematical representation of fitness and how it may be regarded as explanatory. The ensuing complex nexus of fitness emphasises the distinction between biological (...) |
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We look to mitonuclear ecology and the phenomenon of Mother’s Curse to argue that the sex of parents and offspring among populations of eukaryotic organisms, as well as the mitochondrial genome, ought to be taken into account in the conceptualization of evolutionary fitness. Subsequently, we show how characterizations of fitness considered by philosophers that do not take sex and the mitochondrial genome into account may suffer. Last, we reflect on the debate regarding the fundamentality of trait versus organism fitness and (...) |
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Much has been made of how Darwinian thinking destroyed proofs for the existence of God from ‘design’ in the universe. I challenge that prevailing view by looking closely at classical ‘teleological’ arguments for the existence of God. One version championed by Aristotle and Thomas Aquinas stems from how chance is not a sufficient kind of ultimate explanation of the universe. In the course of constructing this argument, I argue that the classical understanding of teleology is no less necessary in modern (...) |
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This paper explores how work in the philosophy of science can be used when teaching scientific content to science students and when training future science teachers. I examine the debate on the concept of fitness in biology and in the philosophy of biology to show how conceptual pluralism constitutes a problem for the conceptual change model, and how philosophical work on conceptual clarification can be used to address that problem. The case of fitness exemplifies how the philosophy of science offers (...) |
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Fitness is a central concept in evolutionary theory. Just as it is central to biological evolution, so, it seems, it should be central to cultural evolutionary theory. But importing the biological fitness concept to CET is no straightforward task—there are many features unique to cultural evolution that make this difficult. This has led some theorists to argue that there are fundamental problems with cultural fitness that render it hopelessly confused. In this essay, we defend the coherency of cultural fitness against (...) |
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In this paper, I argue (contra some recent philosophical work) that an objective distinction between natural selection and drift can be drawn. I draw this distinction by conceiving of drift, in the most fundamental sense, as an individual-level phenomenon. This goes against some other attempts to distinguish selection from drift, which have argued either that drift is a population-level process or that it is a population-level product. Instead of identifying drift with population-level features, the account introduced here can explain these (...) |
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Work throughout the history and philosophy of biology frequently employs ‘chance’, ‘unpredictability’, ‘probability’, and many similar terms. One common way of understanding how these concepts were introduced in evolution focuses on two central issues: the first use of statistical methods in evolution (Galton), and the first use of the concept of “objective chance” in evolution (Wright). I argue that while this approach has merit, it fails to fully capture interesting philosophical reflections on the role of chance expounded by two of (...) |
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Philosophers of biology have worked extensively on how we ought best to interpret the probabilities which arise throughout evolutionary theory. In spite of this substantial work, however, much of the debate has remained persistently intractable. I offer the example of Bayesian models of divergence time estimation as a case study in how we might bring further resources from the biological literature to bear on these debates. These models offer us an example in which a number of different sources of uncertainty (...) |
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The propensity interpretation of fitness (PIF) is commonly taken to be subject to a set of simple counterexamples. We argue that three of the most important of these are not counterexamples to the PIF itself, but only to the traditional mathematical model of this propensity: fitness as expected number of offspring. They fail to demonstrate that a new mathematical model of the PIF could not succeed where this older model fails. We then propose a new formalization of the PIF that (...) |
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Over the past decade philosophers of biology have discussed whether evolutionary theory is a causal theory or a phenomenological study of evolution based solely on the statistical features of a population. This article reviews this controversy from three aspects, respectively concerning the assumptions, applications, and explanations of evolutionary theory, with a view to arriving at a definite conclusion in each contention. In so doing I also argue that an implicit methodological assumption shared by both sides of the debate, namely the (...) |
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According to a once influential view of selection, it consists of repeated cycles of replication and interaction. It has been argued that this view is wrong: replication is not necessary for evolution by natural selection. I analyze the nine most influential arguments for this claim and defend the replication–interaction conception of selection against these objections. In order to do so, however, the replication–interaction conception of selection needs to be modified significantly. My proposal is that replication is not the copying of (...) |
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Some mental states have valence—they are pleasant or unpleasant. According to imperativism, valence depends on imperative content, while evaluativism tells us that it depends on evaluative content. We argue that if one considers valence’s informational profile, it becomes evident that imperativism is superior to evaluativism. More precisely, we show that if one applies the best available metasemantics to the role played by (un)pleasant mental states in our cognitive economy, then these states turn out to have imperative rather than evaluative content, (...) |
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There is a trend within philosophy of biology to concentrate on questions that are strongly related to particular biological research programs rather than on the general scope of the field and its relation to other sciences. Projects of the latter kind, of course, are followed as well but will not be the topic of this review. Shifting the focus to particular research programs reflects philosophers’ increased interest in knowledge of, and contribution to, actual biological research, which is organized in such (...) |
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Darwin’s claim about natural selection is reconstructed as an empirical claim about a causal connection leading from the match of the physiology of an individual and its environment to leaving surviving progeny. Variations in this match, Darwin claims, cause differences in the survival of the progeny. Modern concepts of fitness focus the survival side of this chain. Therefore, the assumption that evolutionary theory wants to explain reproductive success in terms of a modern concept of fitness has given rise to the (...) |
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Adaptation is one of the central concepts in evolutionary theory, which nonetheless has been given different definitions. Some scholars support a historical definition of adaptation, considering it as a trait that is the outcome of natural selection, whereas others support an ahistorical definition, considering it as a trait that contributes to the survival and reproduction of its possessors. Finally, adaptation has been defined as a process, as well. Consequently, two questions arise: the first is a philosophical one and focuses on (...) |
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In this article, I consider the term “environment” in various claims and models by evolutionists and ecologists. I ask whether “environment” is amenable to a philosophical explication, in the same way some key terms of evolutionary theorizing such as “fitness,” “species,” or more recently “population” have been. I will claim that it cannot. In the first section, I propose a typology of theoretical terms, according to whether they are univocal or equivocal, and whether they have been the object of formal (...) |
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‘Statisticalists’ argue that the individual interactions of organisms taken together constitute natural selection. On this view, natural selection is an aggregated effect of interactions rather than some added cause acting on populations. The statisticalists’ view entails that natural selection and drift are indistinguishable aggregated effects of interactions, so that it becomes impossible to make a difference between them. The present paper attempts to make sense of the difference between selection and drift, given the main insights of statisticalism; basically, it will (...) |
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Both biologists and philosophers often make use of simple verbal formulations of necessary and sufficient conditions for evolution by natural selection (ENS). Such summaries go back to Darwin's Origin of Species (especially the "Recapitulation"), but recent ones are more compact.1 Perhaps the most commonly cited formulation is due to Lewontin.2 These summaries tend to have three or four conditions, where the core requirement is a combination of variation, heredity, and fitness differences. The summaries are employed in several ways. First, they (...) |
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Problems raised by complex life cycles for standard summaries of evolutionary processes, and for concepts of individuality in biology, are described. I then outline a framework that can be used to compare life cycles. This framework treats reproduction as a combination of production and recurrence and organizes life cycles according to the distribution of steps in which multiplication, bottlenecks, and sex occur. I also discuss fitness and its measurement in complex life cycles and consider some phenomena that raise complications and (...) |
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The concept of fitness has generated a lot of discussion in philosophy of biology. There is, however, relative agreement about the need to distinguish at least two uses of the term: ecological fitness on the one hand, and population genetics fitness on the other. The goal of this paper is to give an explication of the concept of ecological fitness by providing a reconstruction of the theory of natural selection in which this concept was framed, that is, based on the (...) |
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Questioning the nature of individuality has a long and a rich history, both in philosophy and in biology. Because they differ in several features from the pervasive vertebrate-human model, plants have been considered as complicating the question. Here, the various plant species on which authors—whether biologists or philosophers—rely to build the picture of plant individuality are examined and tracked for their peculiarities, thus constituting an “herbiary” of plant individuality. The herbiary of plant individuality has as its members species exhibiting a (...) |
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Recent studies in the evolution of cooperation have shifted focus from altruistic to mutualistic cooperation. This change in focus is purported to reveal new explanations for the evolution of prosocial behavior. We argue that the common classification scheme for social behavior used to distinguish between altruistic and mutualistic cooperation is flawed because it fails to take into account dynamically relevant game-theoretic features. This leads some arguments about the evolution of cooperation to conflate dynamical scenarios that differ regarding the basic conditions (...) |
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We argue that ecology in general and biodiversity and ecosystem function research in particular need an understanding of functions which is both ahistorical and evolutionarily grounded. A natural candidate in this context is Bigelow and Pargetter’s evolutionary forward-looking account which, like the causal role account, assigns functions to parts of integrated systems regardless of their past history, but supplements this with an evolutionary dimension that relates functions to their bearers’ ability to thrive and perpetuate themselves. While Bigelow and Pargetter’s account (...) |
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Fitness is a central but notoriously vexing concept in evolutionary biology. The propensity interpretation of fitness is often regarded as the least problematic account for fitness. It ties an individual's fitness to a probabilistic capacity to produce offspring. Fitness has a clear causal role in evolutionary dynamics under this account. Nevertheless, the propensity interpretation faces its share of problems. We discuss three of these. We first show that a single scalar value is an incomplete summary of a propensity. Second, we (...) |
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The purpose of this paper is to defend, contra Fodor and Piattelli-Palmarini (F&PP), that the theory of natural selection (NS) is a perfectly bona fide empirical unified explanatory theory. F&PP claim there is nothing non-truistic, counterfactual-supporting, of an “adaptive” character and common to different explanations of trait evolution. In his debate with Fodor, and in other works, Sober defends NS but claims that, compared with classical mechanics (CM) and other standard theories, NS is peculiar in that its explanatory models are (...) |
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The problem of plant individuality is something which has vexed botanists throughout the ages, with fashion swinging back and forth from treating plants as communities of individuals (Darwin 1800 ; Braun and Stone 1853 ; Münch 1938 ) to treating them as organisms in their own right, and although the latter view has dominated mainstream thought most recently (Harper 1977 ; Cook 1985 ; Ariew and Lewontin 2004 ), a lively debate conducted mostly in Scandinavian journals proves that the issues (...) |
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Evolutionary transitions in individuality (ETIs) are events during which individuals at a given level of organization (particles) interact to form higher-level entities (collectives) which are then recognized as new individuals at that level. ETIs are intimately related to levels of selection, which, following Okasha, can be approached from two different perspectives. One, referred to as ‘synchronic’, asks whether selection occurs at the collective level while the partitioning of particles into collectives is taken for granted. The other, referred to as ‘diachronic’, (...) No categories |
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Yes, fitness is the central concept of evolutionary biology, but it is an elusive concept. Almost everyone who looks at it seriously comes out in a different place. |
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We argue that a fashionable interpretation of the theory of natural selection as a claim exclusively about populations is mistaken. The interpretation rests on adopting an analysis of fitness as a probabilistic propensity which cannot be substantiated, draws parallels with thermodynamics which are without foundations, and fails to do justice to the fundamental distinction between drift and selection. This distinction requires a notion of fitness as a pairwise comparison between individuals taken two at a time, and so vitiates the interpretation (...) |
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Following Wallace’s suggestion, Darwin framed his theory using Spencer’s expression “survival of the fittest”. Since then, fitness occupies a significant place in the conventional understanding of Darwinism, even though the explicit meaning of the term ‘fitness’ is rarely stated. In this paper I examine some of the different roles that fitness has played in the development of the theory. Whereas the meaning of fitness was originally understood in ecological terms, it took a statistical turn in terms of reproductive success throughout (...) |
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A basic challenge to naturalistic moral realism is that, even if moral properties existed, there would be no way to naturalistically represent or track them. Here, the basic structure for a tracking account of moral epistemology is given in empirically respectable terms, based on a eudaimonist conception of morality. The goal is to show how this form of moral realism can be seen as consistent with the details of evolutionary biology as well as being amenable to the most current understanding (...) |
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This volume focuses on various questions concerning the interpretation of probability and probabilistic reasoning in biology and physics. It is inspired by the idea that philosophers of biology and philosophers of physics who work on the foundations of their disciplines encounter similar questions and problems concerning the role and application of probability, and that interaction between the two communities will be both interesting and fruitful. In this introduction we present the background to the main questions that the volume focuses on (...) No categories |
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It has been argued that biological fitness cannot be defined as expected number of offspring in all contexts. Some authors argue that fitness therefore merely satisfies a common schema or that no unified mathematical characterization of fitness is possible. I argue that comparative fitness must be relativized to an evolutionary effect; thus relativized, fitness can be given a unitary mathematical characterization in terms of probabilities of producing offspring and other effects. Such fitnesses will sometimes be defined in terms of probabilities (...) |
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and was correct to conclude that the way a biological population is described should affect conclusions about whether natural selection occurs, but wrong to conclude that natural selection is therefore not a cause. After providing a new argument that ignored crucial biological details, I give a biological illustration that motivates a fairly extreme dependence on description. I argue that contrary to an implication of , biologists allow much flexibility in describing populations, as contemporary research on recent human evolution shows. Properly (...) |
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One controversy about the existence of so called evolutionary forces such as natural selection and random genetic drift concerns the sense in which such “forces” can be said to interact. In this paper I explain how natural selection and random drift can interact. In particular, I show how population-level probabilities can be derived from individual-level probabilities, and explain the sense in which natural selection and drift are embodied in these population-level probabilities. I argue that whatever causal character the individual-level probabilities (...) |
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It’s recently been argued that biological fitness can’t change over the course of an organism’s life as a result of organisms’ behaviors. However, some characterizations of biological function and biological altruism tacitly or explicitly assume that an effect of a trait can change an organism’s fitness. In the first part of the paper, I explain that the core idea of changing fitness can be understood in terms of conditional probabilities defined over sequences of events in an organism’s life. The result (...) |
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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 (...) |
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Fitness plays many roles throughout evolutionary theory, from a measure of populations in the wild to a central element in abstract theoretical presentations of natural selection. It has thus been the subject of an extensive philosophical literature, which has primarily centered on the way to understand the relationship between fitness values and reproductive outcomes. If fitness is a probabilistic or statistical quantity, how is it to be defined in general theoretical contexts? How can it be measured? Can a single conceptual (...) |
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