Denis Walsh has written a striking new defense in this journal of the statisticalist (i.e., noncausalist) position regarding the forces of evolution. I defend the causalist view against his new objections. I argue that the heart of the issue lies in the nature of nonadditive causation. Detailed consideration of that turns out to defuse Walsh’s ‘description‐dependence’ critique of causalism. Nevertheless, the critique does suggest a basis for reconciliation between the two competing views. *Received December 2009; revised December 2009. †To contact (...) the author, please write to: Department of Philosophy, 599 Lucas Hall, One University Boulevard, University of Missouri, St. Louis, MO 63121; e‐mail: [email protected]. (shrink)

In evolutionary biology changes in population structure are explained by citing trait fitness distribution. I distinguish three interpretations of fitness explanations—the Two‐Factor Model, the Single‐Factor Model, and the Statistical Interpretation—and argue for the last of these. These interpretations differ in their degrees of causal commitment. The first two hold that trait fitness distribution causes population change. Trait fitness explanations, according to these interpretations, are causal explanations. The last maintains that trait fitness distribution correlates with population change but does not cause (...) it. My defense of the Statistical Interpretation relies on a distinctive feature of causation. Causes conform to the Sure Thing Principle. Trait fitness distributions, I argue, do not. *Received July 2009; revised October 2009. †To contact the author, please write to: Department of Philosophy/Institute for the History, Philosophy of Science and Technology, University of Toronto, Victoria College, 91 Charles Street West, Toronto, ON M5S 1K7, Canada; e‐mail: [email protected]. (shrink)

A hitherto neglected form of explanation is explored, especially its role in population genetics. “Statistically abstractive explanation” (SA explanation) mandates the suppression of factors probabilistically relevant to an explanandum when these factors are extraneous to the theoretical project being pursued. When these factors are suppressed, the explanandum is rendered uncertain. But this uncertainty traces to the theoretically constrained character of SA explanation, not to any real indeterminacy. Random genetic drift is an artifact of such uncertainty, and it is therefore wrong (...) to reify it as a cause of evolution or as a process in its own right. *Received July 2009. †To contact the author, please write to: Department of Philosophy, University of Toronto, 170 St. George St., Toronto, ON M5R 2M8, Canada; e‐mail: [email protected]. (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.

The central point of this essay is to demonstrate the incommensurability of ‘Darwinian fitness’ with the numeric values associated with reproductive rates used in population genetics. While sometimes both are called ‘fitness’, they are distinct concepts coming from distinct explanatory schemes. Further, we try to outline a possible answer to the following question: from the natural properties of organisms and a knowledge of their environment, can we construct an algorithm for a particular kind of organismic life-history pattern that itself will (...) allow us to predict whether a type in the population will increase or decrease relative to other types? Introduction Darwinian fitness Reproductive fitness and genetical models of evolution The models of reproductive fitness 4.1 The Standard Viability Model 4.2 Frequency-dependent selection 4.3 Fertility models 4.4 Overlapping generations Fitness as outcome 5.1 Fitness as actual increase in type 5.2 Fitness as expected increase in type 5.2.1 Expected increase within a generation 5.2.2 Expected increase between generations 5.2.3 Postponed reproductive fitness effects The book-keeping problem Conclusion. (shrink)

Issues concerning scientific explanation have been a focus of philosophical attention from Pre- Socratic times through the modern period. However, recent discussion really begins with the development of the Deductive-Nomological (DN) model. This model has had many advocates (including Popper 1935, 1959, Braithwaite 1953, Gardiner, 1959, Nagel 1961) but unquestionably the most detailed and influential statement is due to Carl Hempel (Hempel 1942, 1965, and Hempel & Oppenheim 1948). These papers and the reaction to them have structured subsequent discussion concerning (...) scientific explanation to an extraordinary degree. After some general remarks by way of background and orientation (Section 1), this entry describes the DN model and its extensions, and then turns to some well-known objections (Section 2). It next describes a variety of subsequent attempts to develop alternative models of explanation, including Wesley Salmon's Statistical Relevance (Section 3) and Causal Mechanical (Section 4) models and the Unificationist models due to Michael Friedman and Philip Kitcher (Section 5). Section 6 provides a summary and discusses directions for future work. (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)

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)

Ernst Mayr has argued that Darwinian theory discredited essentialist modes of thought and replaced them with what he has called "population thinking". In this paper, I characterize essentialism as embodying a certain conception of how variation in nature is to be explained, and show how this conception was undermined by evolutionary theory. The Darwinian doctrine of evolutionary gradualism makes it impossible to say exactly where one species ends and another begins; such line-drawing problems are often taken to be the decisive (...) reason for thinking that essentialism is untenable. However, according to the view of essentialism I suggest, this familiar objection is not fatal to essentialism. It is rather the essentialist's use of what I call the natural state model for explaining variation which clashes with evolutionary theory. This model implemented the essentialist's requirement that properties of populations be defined in terms of properties of member organisms. Requiring such constituent definitions is reductionistic in spirit; additionally, evolutionary theory shows that such definitions are not available, and, moreover, that they are not needed to legitimize population-level concepts. Population thinking involves the thesis that population concepts may be legitimized by showing their connections with each other, even when they are not reducible to concepts applying at lower levels of organization. In the paper, I develop these points by describing Aristotle's ideas on the origins of biological variation; they are a classic formulation of the natural state model. I also describe how the development of statistical ideas in the 19th century involved an abandoning of the natural state model. (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)

In the epigraph, Fisher is blaming two generations of theoretical biologists, from Darwin on, for ignoring Quetelet's statistical techniques and hence harboring confusions about evolution and natural selection. He is right to imply that Darwin and his contemporaries were aware of the core of Quetelet's work. Quetelet's seminal monograph, Sur L'homme, was widely discussed in Darwin's academic circles. We know that Darwin owned a copy (Schweber 1977). More importantly, we have in Darwin's notebooks two entries referring to Quetelet's work on (...) the cause of a large-scale global phenomenon where each year more boys were born than girls. The first entry is written sometime between April and July 1838. Darwin writes "Find out from the Statistical Society—where M. Quetelet has published his laws about sexes relative to age of Marriages" (C 268, Barrett et al., 1987, p. 324). The second is written sometime after October 16, 1838: "In the Atheneum Numbers 406, 407, 409, Quetelet papers are given, & I think facts there mentioned about proportion of sexes, at birth & causes" (Ibid, p. 379). So, even if Darwin did not read Sur L'homme directly it is likely (though not certain) that he read its review in the Atheneum. There is no doubt that Darwin eventually became familiar with Quetelet's work in statistics. The smoking gun is an essay that Darwin writes in 1874, entitled, "On the Males and Complemental Males of Certain Cirripedes, and on Rudimentary Structures" where he discusses Quetelet's laws of variation. (shrink)

Several philosophers of science have advanced an instrumentalist thesis about the use of probabilities in evolutionary biology. I investigate the consequences of instrumentalism on evolutionary explanations. I take issue with Barbara Horan's (1994) argument that probabilities are unnecessary to explain evolutionary change given the underlying deterministic character of evolutionary processes. First, I question Horan's deterministic assumption. Then, I attempt to undermine her Laplacian argument by demonstrating that whether probabilities are necessary depends upon the sort of questions one is asking.

Evolutionary processes such as natural selection and random drift are commonly regarded as causes of population-level change. We respond to a recent challenge that drift and selection are best understood as statistical trends, not causes. Our reply appeals to manipulation as a strategy for uncovering causal relationships: if you can systematically manipulate variable A to bring about a change in variable B, then A is a cause of B. We argue that selection and drift can be systematically manipulated to produce (...) different kinds of population-level change. They should therefore be regarded as causes. (shrink)

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 (...) of explanations from portfolio theory and population genetics that meet this characterisation. In each case the invariance relation holds between a statistical property of an ensemble and a change in structure of the ensemble. In neither case, however, does the statistical property cause the outcome it explains. There are genuine statistical, non-causal scientific explanations. (shrink)

Carl Hempel1 set the tone for subsequent philosophical work on scientific explanation by resolutely locating the problem he wanted to address outside of epistemology. “Hempel’s problem,” as I will call it, was not to say what counts as evidence that X is the explanation of Y. Rather, the question was what it means for X to explain Y. Hempel’s theory of explanation and its successors don’t tell you what to believe; instead, they tell you which of your beliefs (if any) (...) can be said to explain a given target proposition. (shrink)

The debate between the Mendelians and the (largely Darwinian) biometricians has been referred to by R. A. Fisher as ‘one of the most needless controversies in the history of science’ and by David Hull as ‘an explicable embarrassment’. The literature on this topic consists mainly of explaining why the controversy occurred and what factors prevented it from being resolved. Regrettably, little or no mention is made of the issues that figured in its resolution. This paper deals with the latter topic (...) and in doing so reorients the focus of the debate as one between Karl Pearson and R. A. Fisher rather than between the biometricians and the Mendelians. One reason for this reorientation is that Pearson's own work in 1904 and 1909 suggested that Mendelism and biometry could, to some extent, be made compatible, yet he remained steadfast in his rejection of Mendelism. The interesting question then is why Fisher, who was also a proponent of biometric methods, was able to synthesise the two traditions in a way that Pearson either could not or would not. My answer to this question involves an analysis of the ways in which different kinds of assumptions were used in modelling Mendelian populations. I argue that it is these assumptions, which lay behind the statistical techniques of Pearson and Fisher, that can be isolated as the source of Pearson's rejection of Mendelism and Fisher's success in the synthesis. (shrink)