Population-level theories of evolution—the stock and trade of population genetics—are statistical theories par excellence. But what accounts for the statistical character of population-level phenomena? One view is that the population-level statistics are a product of, are generated by, probabilities that attach to the individuals in the population. On this conception, population-level phenomena are explained by individual-level probabilities and their population-level combinations. Another view, which arguably goes back to Fisher but has been defended recently, is that the population-level statistics (...) are sui generis, that they somehow emerge from the underlying deterministic behavior of the individuals composing the population. Walsh et al. label this the statistical interpretation. We are not willing to give them that term, since everyone will admit that the population-level theories of evolution are statistical, so we will call this the emergentist statistical interpretation. Our goals are to show that: This interpretation is based on gross factual errors concerning the practice of evolutionary biology, concerning both what is done and what can be done; its adoption would entail giving up on most of the explanatory and predictive projects of evolutionary biology; and finally a rival interpretation, which we will label the propensity statistical interpretation succeeds exactly where the emergentist interpretation fails. (shrink)

What is natural selection? I address this question by exploring the relation between two debates: Is natural selection a mechanism? Is natural selection a causal or a statistical theory? I argue that the first can be assessed only relative to a model and that, following the second, there are two fundamentally different and independent kinds of models, Modern-Synthesis and Darwinian models. MS-models, I argue, are not mechanistic even if they are causal. D-models, in (...) contrast, are mechanistic. A causal-mechanistic interpretation of D-models is thus compatible with a statistical interpretation of MS-models. Natural selection, I conclude, lacks a single, unifying nature. (shrink)

There is a widespread philosophical interpretation of natural selection in evolutionary theory: natural selection, like mutation, migration, and drift are seen as forces that propel the evolution of populations. Natural selection is thus a population level causal process. This account has been challenged by the Statistics, claiming that natural selection is not a population level cause but rather a statistical feature of a population. This paper examines the nature of the (...) aforementioned ontological debate and the nature of statistical explanations given by population genetics. I claim that the Modern Synthesis provides good explanations of the changes in trait structure of populations without appealing to detailed causal information about the individual trajectories of the members of a population. (shrink)

How do fitness and natural selection relate to other evolutionary factors like architectural constraint, mode of reproduction, and drift? In one way of thinking, drawn from Newtonian dynamics, fitness is one force driving evolutionary change and added to other factors. In another, drawn from statistical thermodynamics, it is a statistical trend that manifests itself in natural selection histories. It is argued that the first model is incoherent, the second appropriate; a hierarchical realization model is (...) proposed as a basis for a statistical treatment. It emerges that natural selection does not cause evolution; it just is evolution. The theory incorporates relations of statistical correlation, but not the kind of causation found in fundamental physical processes. (shrink)

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)

In this paper, I answer a fundamental question facing any view according to which natural selection is a population‐level causal process—namely, how is the causal process of natural selection related to, yet not preempted by, causal processes that occur at the level of individual organisms? Without an answer to this grounding question, the population‐level causal view appears unstable—collapsing into either an individual‐level causal interpretation or the claim that selection is a purely formal, statistical (...) phenomenon. I argue that a causal account of realization provides an answer to the grounding question. By applying this account of realization to the natural selection of melanism in rock pocket mice, I show how an alternative, formal account of realization, favored by proponents of the statistical interpretation, misses biologically important features. More generally, this paper shows how metaphysical issues about realization normally discussed in the philosophy of mind apply to debates in philosophy of biology. Thus, it is a first step toward fleshing out the oft‐noted similarities between debates in these areas. (shrink)

Recently some philosophers have emphasized a potentially irreconcilable conceptual antagonism between the statistical characterization of natural selection and the standard scientific discussion of natural selection in terms of forces and causes. Other philosophers have developed an account of the causal character of selectionist statements represented in terms of counterfactuals. I examine the compatibility between such statisticalism and counterfactually based causal accounts of natural selection by distinguishing two distinct statisticalist claims: firstly the suggested impossibility (...) for natural selection to be a cause acting upon populations and secondly the conceptualization that all evolutionary causes occur at the level of interactions between individual organisms. I argue that deriving the latter from the former involves supplementary assumptions concerning precisely what causation is. I critically examine two of these assumptions purportedly preventing natural selection being regarded as a cause: the locality claim and the modularity claim. I conclude that justifying the strongest version of statisticalism—i.e. evolutionary causation only occurs at the level of individual interactions between organisms—would require further metaphysical arguments that are likely to be deemed highly problematic. Additionally, I argue that such a metaphysical position would be considered incongruous with both our scientific and ordinary use of the concepts of causality and explanation as employed within our everyday epistemological framework. (shrink)

The debate between the dynamical and the statistical interpretations of natural selection is centred on the question of whether all explanations that employ the concepts of natural selection and drift are reducible to causal explanations. The proponents of the statistical interpretation answer negatively, but insist on the fact that selection/drift arguments are explanatory. However, they remain unclear on where the explanatory power comes from. The proponents of the dynamical interpretation answer positively (...) and try to reduce selection/drift arguments to some of the most prominent accounts of causal explanation. In turn, they face the criticism raised by statisticalists that current accounts of causation have to be violated in some of their core conditions or otherwise used in a very loose manner in order to account for selection/drift explanations. We propose a reconciliation of both interpretations by conveying evolutionary explanations within the unificationist model of scientific explanation. Therefore, we argue that the explanatory power in natural selection arguments is a result of successful unification of individual- and population-level facts. A short case study based on research on sympatric speciation will be presented as an example of how population- and individual-level facts are unified to explain the morphological mosaic of bill shape in island scrub jays. (shrink)

I defend a radical interpretation of biological populations—what I call population pluralism—which holds that there are many ways that a particular grouping of individuals can be related such that the grouping satisfies the conditions necessary for those individuals to evolve together. More constraining accounts of biological populations face empirical counter-examples and conceptual difficulties. One of the most intuitive and frequently employed conditions, causal connectivity—itself beset with numerous difficulties—is best construed by considering the relevant causal relations as ‘thick’ causal concepts. (...) I argue that the fine-grained causal relations that could constitute membership in a biological population are huge in number and many are manifested by degree, and thus we can construe population membership as being defined by massively multidimensional constructs, the differences between which are largely arbitrary. I end by showing that positions in two recent debates in theoretical biology depend on a view of biological populations at odds with the pluralism defended here. 1 Introduction2 Biological Population, Broad and Narrow3 Difficulties with Narrow Biological Population Conditions3.1 Against the genealogical condition3.2 Against the conspecificity condition3.3 Against the proximity condition3.4 Against the typology condition4 Causal Connectivity5 Massively Multidimensional Population Constructs6 Population Uniqueness and Natural Selection6.1 Statisticalism and its discontents6.2 Price at what price?7 Conclusion. (shrink)

In The Structure of Biological Science I argued that the theory of natural selection is a statistical theory for reasons much like those which makes thermodynamics a statistical theory. In particular, the theory claims that fitness differences are large enough and the life span of species long enough for increases in average fitness always to appear in the long run; and this claim, I held, is of the same form as the statistical version of the (...) second law of thermodynamics.For the latter law also makes a claim about the long run, and its statistical character is due to this claim: thermodynamic systems must in the long run approach an equilibrium level of organization that maximizes entropy. Over finite times, given local boundary conditions, an isolated mechanical system, like the molecules in a container of gas, may sometimes interact so as to move the entropy of the system further from, instead of closer to the equilbrium level. But given enough interacting bodies, and enough time, the system will always eventually move in the direction prescribed by the law. Thus, we can attach much higher probabilities to the prediction that non-equilibrium systems will reflect greater entropy in future periods than we can to predictions that they will move in the opposite direction. And as we increase the amount of time and the number of bodies interacting, the strength of the probability we can attach to the prediction becomes greater and greater. (shrink)

In The Structure of Biological Science I argued that the theory of natural selection is a statistical theory for reasons much like those which makes thermodynamics a statistical theory. In particular, the theory claims that fitness differences are large enough and the life span of species long enough for increases in average fitness always to appear in the long run; and this claim, I held, is of the same form as the statistical version of the (...) second law of thermodynamics.For the latter law also makes a claim about the long run, and its statistical character is due to this claim: thermodynamic systems must in the long run approach an equilibrium level of organization that maximizes entropy. Over finite times, given local boundary conditions, an isolated mechanical system, like the molecules in a container of gas, may sometimes interact so as to move the entropy of the system further from, instead of closer to the equilbrium level. But given enough interacting bodies, and enough time, the system will always eventually move in the direction prescribed by the law. Thus, we can attach much higher probabilities to the prediction that non-equilibrium systems will reflect greater entropy in future periods than we can to predictions that they will move in the opposite direction. And as we increase the amount of time and the number of bodies interacting, the strength of the probability we can attach to the prediction becomes greater and greater. (shrink)

Shapiro and Sober claim that Walsh, Ariew, Lewens, and Matthen give a mistaken, a priori defense of natural selection and drift as epiphenomenal. Contrary to Shapiro and Sober’s claims, we first argue that WALM’s explanatory doctrine does not require a defense of epiphenomenalism. We then defend WALM’s explanatory doctrine by arguing that the explanations provided by the modern genetical theory of natural selection are ‘autonomous-statistical explanations’ analogous to Galton’s explanation of reversion to mediocrity and an (...) explanation of the diffusion ofgases. We then argue that whereas Sober’s theory of forces is an adequate description of Darwin’s theory, WALM’s explanatory doctrine is required to understand how themodern genetical theory of natural selection explains large-scale statistical regularities. 1 Introduction2 Shapiro and Sober’s ‘Epiphenomenalism Do’s and Don’ts’3 WALM’s Explanatory Doctrine4 Galton’s Autonomous-Statistical Explanation5 A Second Example: The Statistical Explanation of the Diffusion of Gases6 Distinguishing Two Theories of Evolution by Natural Selection7 A Possible Objection: Are Statistical Laws Sufficient for Explanation?8 Conclusion. (shrink)

We have argued elsewhere that: (A) Natural selection is not a cause of evolution. (B) A resolution-of-forces (or vector addition) model does not provide us with a proper understanding of how natural selection combines with other evolutionary influences. These propositions have come in for criticism recently, and here we clarify and defend them. We do so within the broad framework of our own “hierarchical realization model” of how evolutionary influences combine.

This paper gives a detailed narrative of a controversial empirical research in postwar population genetics, the analysis of the cytological polymorphisms of an Australian grasshopper, Moraba scurra. This research intertwined key technical developments in three research areas during the 1950s and 1960s: it involved Dobzhansky’s empirical research program on cytological polymorphisms, the mathematical theory of natural selection in two-locus systems, and the building of reliable estimates of natural selection in the wild. In the mid-1950s the cytologist (...) Michael White discovered an interesting case of epistasis in populations of Moraba scurra. These observations received a wide diffusion when theoretical population geneticist Richard Lewontin represented White’s data on adaptive topographies. These topographies connected the information on the genetic structure of these grasshopper populations with the formal framework of theoretical population genetics. As such, they appeared at the time as the most successful application of two-locus models of natural selection to an empirical study system. However, this connection generated paradoxical results: in the landscapes, all grasshopper populations were located on a ridge while they were expected to reach a peak. This puzzling result fueled years of research and triggered a controversy attracting contributors from Australia, the United States and the United Kingdom. While the original problem seemed, at first, purely empirical, the subsequent controversy affected the main mathematical tools used in the study of two-gene systems under natural selection. Adaptive topographies and their underlying mathematical structure, Wright’s mean fitness equations, were submitted to close scrutiny. Suspicion eventually shifted to the statistical machinery used in data analysis, reflecting the crucial role of statistical inference in applied population genetics. In the 1950s and 1960s, population geneticists were not simply in search for new generalizations about the evolutionary process and for new evidence about genetic variation: struggling against statistical artifacts, they were searching for reliable tests and descriptors to analyze their data. (shrink)

There are two competing interpretations of the modern synthesis theory of evolution: the dynamical (also know as ‘traditional’) and the statistical. The dynamical interpretation maintains that explanations offered under the auspices of the modern synthesis theory articulate the causes of evolution. It interprets selection and drift as causes of population change. The statistical interpretation holds that modern synthesis explanations merely cite the statistical structure of populations. This paper offers a defense of statisticalism. It argues (...) that a change in trait frequencies in a population can be attributed only to selection or drift against the background of a particular statistical description of the population. The traditionalist supposition that selection and drift are description‐independent causes of population change leads the dynamical interpretation into a dilemma: it must face a contradiction or accept the loss of explanatory power. (shrink)

This paper provides a philosophical analysis of the ongoing controversy surrounding R.A. Fisher's famous ‘fundamental theorem’ of natural selection. The difference between the ‘traditional’ and ‘modern’ interpretations of the theorem is explained. I argue that proponents of the modern interpretation have captured Fisher's intended meaning correctly and shown that the theorem is mathematically correct, pace the traditional consensus. However, whether the theorem has any real biological significance remains an unresolved issue. I argue that the answer depends on (...) whether we accept Fisher's non-standard notion of environmental change, on which the theorem rests; arguments for and against this notion are explored. I suggest that there is a close link between Fisher's fundamental theorem and the modern ‘gene's eye’ view of evolution. Introduction What Does the Fundamental Theorem Say? Key Concepts Explained Alleged Significance of the FTNS Traditional versus Modern Interpretations of the FTNS The Modern Interpretation Illustrated Fisher's Concept of ‘Environmental Change’ Causality and the Modern Interpretation The Significance of the FTNS Re-considered Appendix CiteULike Connotea Del.icio.us What's this? (shrink)

Recent arguments concerning the nature of causation in evolutionary theory, now often known as the debate between the 'causalist' and 'statisticalist' positions, have involved answers to a variety of independent questions – definitions of key evolutionary concepts like natural selection, fitness, and genetic drift; causation in multi-level systems; or the nature of evolutionary explanations, among others. This Element offers a way to disentangle one set of these questions surrounding the causal structure of natural selection. Doing so (...) allows us to clearly reconstruct the approach that some of these major competing interpretations of evolutionary theory have to this causal structure, highlighting particular features of philosophical interest within each. Further, those features concern problems not exclusive to the philosophy of biology. Connections between them and, in two case studies, contemporary metaphysics and philosophy of physics demonstrate the potential value of broader collaboration in the understanding of evolution. (shrink)

‘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 disentangle the various kinds of indistinguishability between selection and drift that happen within biology, by examining the epistemological and metaphysical nature of the distinction between selection and drift. It will be based on a ‘difference-making account’ of selection. The first section will explicate the inscrutability of selection and drift, its various types in the statisticalist writings, and its implications. The second section specifies concepts of natural selection and drift in the difference making account of selection I am using, and shows that one can derive from this the statistical signatures of selection and drift. On this basis I focus on one sort of indistinguishability issue about selection and drift, which I call epistemic opacity, and explain why it mostly affects small populations. The last section explains why epistemic opacity does not raise an genuine epistemic problem for evolutionary biology. (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)

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 change; they are not discrete, apportionable causes. Our objective here is to provide a definitive statement of the statistical position, so as to allay some confusions in the current literature. We outline four commitments that are central to statisticalism. They are: 1. Natural Selection is a higher order effect; 2. Trait fitness is primitive; 3. Modern Synthesis (MS)-models are substrate neutral; 4. MS-selection and drift are model-relative. (shrink)

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 of the theory as one about populations exclusively. (shrink)

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)

Recent discussion of mechanism has suggested new approaches to several issues in the philosophy of science, including theory structure, causal explanation, and reductionism. Here, I apply what I take to be the fruits of the 'new mechanical philosophy' to an analysis of a contemporary debate in evolutionary biology about the role of natural selection in speciation. Traditional accounts of that debate focus on the geographic context of genetic divergence--namely, whether divergence in the absence of geographic isolation is possible (...) (or significant). Those accounts are at best incomplete, I argue, because they ignore the mechanisms producing divergence and miss what is at stake in the biological debate. I argue that the biological debate instead concerns the scope of particular speciation mechanisms which assign different roles to natural selection at various stages of divergence. The upshot is a new interpretation of the crux of that debate--namely, whether divergence with gene flow is possible (or significant) and whether the isolating mechanisms producing it are adaptive. (shrink)

In Darwinism's Struggle for Survival Jean Gayon offers a philosophical interpretation of the history of theoretical Darwinism. He begins by examining the different forms taken by the hypothesis of natural selection in the nineteenth century and the major difficulties which it encountered, particularly with regard to its compatibility with the theory of heredity. He then shows how these difficulties were overcome during the seventy years which followed the publication of Darwin's Origin of Species, and he concludes by (...) analysing the major features of the genetic theory of natural selection, as it developed from 1920 to 1960. This rich and wide-ranging study will appeal to philosophers and historians of science and to evolutionary biologists. (shrink)

Stephen Jay Gould argued that replaying the “tape of life” would result in a radically different evolutionary outcome. Some biologists and philosophers, however, have pointed to convergent evolution as evidence for robust replicability in macroevolution. These authors interpret homoplasy, or the independent origination of similar biological forms, as evidence for the power of natural selection to guide form toward certain morphological attractors, notwithstanding the diversionary tendencies of drift and the constraints of phylogenetic inertia. In this paper, I consider (...) the implications of homoplasy for the debate over the nature of macroevolution. I argue that once the concepts of contingency and convergence are fleshed out, it becomes clear that many instances of homoplasy fail to negate Gould’s overarching thesis, and may in fact support a Gouldian view of life. My argument rests on the distinction between parallelism and convergence, which I defend against a recent challenge from developmental biology. I conclude that despite the difficulties in defining and identifying parallelism, the concept remains useful and relevant to the contingency controversy insofar as it underscores the common developmental origins of iterated evolution. (shrink)

This paper is a commentary on the focal article by Grafen and on earlier papers of his on which many of the results of this focal paper depend. Thus it is in effect a commentary on the “formal Darwinian project”, the focus of this sequence of papers. Several problems with this sequence are raised and discussed. The first of these concerns fitness maximization. It is often claimed in these papers that natural selection leads to a maximization of fitness (...) and that this view is claimed in Fisher’s “fundamental theorem of natural selection”. These claims are refuted, and various incorrect statements about the meaning and interpretation of the fundamental theorem of natural selection, in this sequence and in other papers by other authors, are discussed. Next, much of the work in this sequence rests on the first Price equation. In the deterministic (infinite population) case this equation is no more than the standard classical equation relating to changes in gene frequencies. In the stochastic case the equation gives the change in gene frequencies as the sum of two terms (the second of which vanishes in the deterministic case). These two terms are of essentially equal importance in the situation considered in the focal article, yet one of Grafen’s results ignores the second term in the stochastic analysis. This is associated with a wavering between deterministic and stochastic analyses and the use of the Price fitness concept and the classical fitness concept. These comments cast doubts on Grafen’s optimization theory. (shrink)

In this paper I propose an interpretation of classical statistical mechanics that centers on taking seriously the idea that probability measures represent complete states of statistical mechanical systems. I show how this leads naturally to the idea that the stochasticity of statistical mechanics is associated directly with the observables of the theory rather than with the microstates (as traditional accounts would have it). The usual assumption that microstates are representationally significant in the theory is therefore dispensable, (...) a consequence which suggests interesting possibilities for developing non-equilibrium statistical mechanics and investigating inter-theoretic answers to the foundational questions of statistical mechanics. (shrink)

When natural selection theory was presented, much active philosophical debate, in which Darwin himself participated, centered on its hypothetical nature, its explanatory power, and Darwin's methodology. Upon first examination, Darwin's support of his theory seems to consist of a set of claims pertaining to various aspects of explanatory success. I analyze the support of his method and theory given in the Origin of Species and private correspondence, and conclude that an interpretation focusing on the explanatory strengths of (...) natural selection theory accurately reflects neither Darwin's own self-consciously held views, nor the nature of his support. Darwin's methodological and philosophical arguments were at once consistently empiricist and more sophisticated than such interpretations credit to him. (shrink)

In Darwinism's Struggle for Survival Jean Gayon offers a philosophical interpretation of the history of theoretical Darwinism. He begins by examining the different forms taken by the hypothesis of natural selection in the nineteenth century and the major difficulties which it encountered, particularly with regard to its compatibility with the theory of heredity. He then shows how these difficulties were overcome during the seventy years which followed the publication of Darwin's Origin of Species, and he concludes by (...) analysing the major features of the genetic theory of natural selection, as it developed from 1920 to 1960. This rich and wide-ranging study will appeal to philosophers and historians of science and to evolutionary biologists. (shrink)

In this paper, I discuss the concept of complexity. I show that the principle of natural selection as acting on complexity gives a solution to the problem of reconciling the seemingly contradictory notion of generally increasing complexity and the observation that most species don’t follow such a trend. I suggest the process of evolution to be illustrated by means of a schematic diagram of complexity versus time, interpreted as a form of the Tree of Life. The suggested model (...) implies that complexity is cumulatively increasing, giving evolution a direction, an arrow of time, thus also implying that the latest emerging species will be the one with the highest level of complexity. Since the human species is the last species evolved in the evolutionary process seen at large, this means that we are the species with the highest complexity. The model implies that the human species constitutes an integral part of organic evolution, yet rendering us the exclusive status as the species of the highest complexity. (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)

The Units of Selection debate is a dispute about the causes of population change. I argue that it is generated by a particular `dynamical'' interpretation of natural selection theory, according to which natural selection causes differential survival and reproduction of individuals and natural selection explanations cite these causes. I argue that the dynamical interpretation is mistaken and offer in outline an alternative, `statistical'' interpretation, according to which natural (...) class='Hi'>selection theory is a fancy kind of `bookkeeping''. It explains by citing the statistical structure of a population and not by citing the causes of survival and reproduction. From the perspective of the statistical interpretation there is no substantive Units of Selection issue. (shrink)

Susan Mills and John Beatty proposed a propensity interpretation of fitness (1979) to show that Darwinian explanations are not circular, but they did not address the critics' chief complaint that the principle of the survival of the fittest is either tautological or untestable. I show that the propensity interpretation cannot rescue the principle from the critics' charges. The critics, however, incorrectly assume that there is nothing more to Darwin's theory than the survival of the fittest. While Darwinians all (...) scoff at this assumption, they do not agree about what role, if any, this principle plays in Darwin's theory of natural selection. I argue that the principle has no place in Darwin's theory. His theory does include the idea that some organisms are fitter than others. But greater reproductive success is simply inferred from higher fitness. There is no reason to embody this inference in the form of a special principle of the survival of the fittest. (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)

The epistemic status of Natural Selection has seemed intriguing to biologists and philosophers since the very beginning of the theory to our present times. One prominent contemporary example is Elliott Sober, who claims that NS, and some other theories in biology, and maybe in economics, are peculiar in including explanatory models/conditionals that are a priori in a sense in which explanatory models/conditionals in Classical Mechanics and most other standard theories are not. Sober’s argument focuses on some “would promote” (...) sentences that according to him, play a central role in NS explanations and are both causal and a priori. Lange and Rosenberg criticize Sober arguing that, though there may be some unspecific a priori causal claims, there are not a priori causal claims that specify particular causal factors. Although we basically agree with Lange and Rosenberg’s criticism, we think it remains silent about a second important element in Sober’s dialectics, namely his claim that, contrary to what happens in mechanics, in NS explanatory conditionals are a priori, and that this is so in quite specific explanatory models. In this paper we criticize this second element of Sober’s argument by analyzing what we take to be the four possible interpretations of Sober’s claim, and argue that, terminological preferences aside, the possible senses in which explanatory models in NS can qualify, or include elements that can qualify, as a priori, also apply to CM and other standard, highly unified theories. We conclude that this second claim is unsound, or at least that more needs to be said in order to sustain that NS explanatory models are a priori in a sense in which CM models are not. (shrink)

It is an ongoing controversy whether natural selection is a cause of population change, or a mere statistical description of how individual births and deaths accumulate. In this paper I restate the problem in terms of the reference class problem, and propose how the structure of stable equilibrium can provide a solution in continuity with biological practice. Insofar natural selection can be understood as a tendency towards equilibrium, key statisticalist criticisms are avoided. Further, in a (...) modification of the Newtonian-force analogy, it can be suggested that a better metaphor for natural selection is that of an emergent force, similar in nature to entropic forces: with magnitude and direction, but lacking a spatiotemporal origin or point of application. (shrink)

Recent discussions in the philosophy of biology have brought into question some fundamental assumptions regarding evolutionary processes, natural selection in particular. Some authors argue that natural selection is nothing but a population-level, statistical consequence of lower-level events (Matthen and Ariew [2002]; Walsh et al. [2002]). On this view, natural selection itself does not involve forces. Other authors reject this purely statistical, population-level account for an individual-level, causal account of natural selection (...) (Bouchard and Rosenberg [2004]). I argue that each of these positions is right in one way, but wrong in another; natural selection indeed takes place at the level of populations, but it is a causal process nonetheless. (shrink)

The Darwinian concept of natural selection was conceived within a set of Newtonian background assumptions about systems dynamics. Mendelian genetics at first did not sit well with the gradualist assumptions of the Darwinian theory. Eventually, however, Mendelism and Darwinism were fused by reformulating natural selection in statistical terms. This reflected a shift to a more probabilistic set of background assumptions based upon Boltzmannian systems dynamics. Recent developments in molecular genetics and paleontology have put pressure on (...) Darwinism once again. Current work on self-organizing systems may provide a stimulus not only for increased problem solving within the Darwinian tradition, especially with respect to origins of life, developmental genetics, phylogenetic pattern, and energy-flow ecology, but for deeper understanding of the very phenomenon of natural selection itself. Since self-organizational phenomena depend deeply on stochastic processes, self-organizational systems dynamics advance the probability revolution. In our view, natural selection is an emergent phenomenon of physical and chemical selection. These developments suggest that natural selection may be grounded in physical law more deeply than is allowed by advocates of the autonomy of biology, while still making it possible to deny, with autonomists, that evolutionary explanations can be modeled in terms of a deductive relationship between laws and cases. We explore the relationship between, chance, self-organization, and selection as sources of order in biological systems in order to make these points. (shrink)

In The Point of View for my Work as an Author, Kierkegaard declares that the works discussed therein move from the aesthetic to the religious, that the Postscript represents the turning point in this movement, etc. In this brief and preliminary study we use a ?change?point? version of the chi?square test on the frequencies of selected sets of ?aesthetic? and ?religious? words to determine the degree of statistical evidence for these and other related claims. Briefly, these tests show that (...) there is strong statistical evidence for all but one of these various claims and that there are therefore good statistical grounds for taking Kierkegaard's ?aesthetic/religious? interpretation seriously. (shrink)

In the recent philosophical literature, two questions have arisen concerning the status of natural selection: (1) Is it a population-level phenomenon, or is it an organism-level phenomenon? (2) Is it a causal process, or is it a purely statistical summary of lower-level processes? In an earlier work (Millstein, Br J Philos Sci, 57(4):627–653, 2006), I argue that natural selection should be understood as a population-level causal process, rather than a purely statistical population-level summation of (...) lower-level processes or as an organism-level causal process. In a 2009 essay entitled “Productivity, relevance, and natural selection,” Stuart Glennan argues in reply that natural selection is produced by causal pro- cesses operating at the level of individual organisms, but he maintains that there is no causal productivity at the population level. However, there are, he claims, many population-level properties that are causally relevant to the dynamics of evolution- ary processes. Glennan’s claims rely on a causal pluralism that holds that there are two types of causes: causal production and causal relevance. Without calling into question Glennan’s causal pluralism or his claims concerning the causal relevance of natural selection, I argue that natural selection does in fact exhibit causal production at the population level. It is true that natural selection does not fit with accounts of mechanisms that involve decomposition of wholes into parts, such as Glennan’s own. However, it does fit with causal production accounts that do not require decomposition, such as Salmon’s Mark Transmission account, given the extent to which populations act as interacting “objects” in the process of natural selection. (shrink)

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 have is then shared by the population-level probabilities, and that natural selection and random drift then have that same causal character. Moreover, natural selection and drift can then be viewed as two aspects of probability distributions over frequencies in populations of organisms. My characterization of population-level probabilities is largely neutral about what interpretation of probability is required, allowing my approach to support various positions on biological probabilities, including those which give biological probabilities one or another sort of causal character. ‡This paper has benefited from feedback on and discussions of this and earlier work. I want to thank André Ariew, Matt Barker, Lindley Darden, Patrick Forber, Nancy Hall, Mohan Matthen, Samir Okasha, Jeremy Pober, Robert Richardson, Alex Rosenberg, Eric Seidel, Denis Walsh, and Bill Wimsatt. †To contact the author, please write to: Department of Philosophy, University of Alabama at Birmingham, HB 414A, 900 13th Street South, Birmingham, AL 35294-1260; e-mail: [email protected]. (shrink)

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 way the theory was put to use in Darwin’s main texts. I will contend that this reconstruction enables us to account for the current use of the theory of natural selection. The framework presupposed in the analysis will be that of metatheoretical structuralism. This framework will provide both a better understanding of the nature of ecological fitness and a more complete reconstruction of the theory. In particular, it will provide what I think is a better way of understanding how the concept of fitness is applied through heterogeneous cases. One of the major advantages of my way of thinking about natural selection theory is that it would not have the peculiar metatheoretical status that it has in other available views. I will argue that in order to achieve these goals it is necessary to make several concepts explicit, concepts that are frequently omitted in usual reconstructions. (shrink)