This paper investigates the conception of causation required in order to make sense of natural selection as a causal explanation of changes in traits or allele frequencies. It claims that under a counterfactual account of causation, natural selection is constituted by the causal relevance of traits and alleles to the variation in traits and alleles frequencies. The “statisticalist” view of selection (Walsh, Matthen, Ariew, Lewens) has shown that natural selection is not a cause superadded to the causal interactions between individual (...) organisms. It also claimed that the only causation at work is those aggregated individual interactions, natural selection being only predictive and explanatory, but it is implicitly committed to a process-view of causation. I formulate a counterfactual construal of the causal statements underlying selectionist explanations, and show that they hold because of the reference they make to ecological reliable factors. Considering case studies, I argue that this counterfactual view of causal relevance proper to natural selection captures more salient features of evolutionary explanations than the statisticalist view, and especially makes sense of the difference between selection and drift. I eventually establish equivalence between causal relevance of traits and natural selection itself as a cause. (shrink)

This paper argues that besides mechanistic explanations, there is a kind of explanation that relies upon “topological” properties of systems in order to derive the explanandum as a consequence, and which does not consider mechanisms or causal processes. I first investigate topological explanations in the case of ecological research on the stability of ecosystems. Then I contrast them with mechanistic explanations, thereby distinguishing the kind of realization they involve from the realization relations entailed by mechanistic explanations, and explain how both (...) kinds of explanations may be articulated in practice. The second section, expanding on the case of ecological stability, considers the phenomenon of robustness at all levels of the biological hierarchy in order to show that topological explanations are indeed pervasive there. Reasons are suggested for this, in which “neutral network” explanations are singled out as a form of topological explanation that spans across many levels. Finally, I appeal to the distinction of explanatory regimes to cast light on a controversy in philosophy of biology, the issue of contingence in evolution, which is shown to essentially involve issues about realization. (shrink)

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.

Recent papers by a number of philosophers have been concerned with the question of whether natural selection is a causal process, and if it is, whether the causes of selection are properties of individuals or properties of populations. I shall argue that much confusion in this debate arises because of a failure to distinguish between causal productivity and causal relevance. Causal productivity is a relation that holds between events connected via continuous causal processes, while causal relevance is a relationship that (...) can hold between a variety of different kinds of facts and the events that counterfactually depend upon them. I shall argue that the productive character of natural selection derives from the aggregation of individual processes in which organisms live, reproduce and die. At the same time, a causal explanation of the distribution of traits will necessarily appeal both to causally relevant properties of individuals and to causally relevant properties that exist only at the level of the population. (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)

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)

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)

According to a prominent view of evolutionary theory, natural selection and the processes of development compete for explanatory relevance. Natural selection theory explains the evolution of biological form insofar as it is adaptive. Development is relevant to the explanation of form only insofar as it constrains the adaptation-promoting effects of selection. I argue that this view of evolutionary theory is erroneous. I outline an alternative, according to which natural selection explains adaptive evolution by appeal to the statistical structure of populations, (...) and development explains the causes of adaptive evolution at the level of individuals. Only together can a statistical theory of selection and a mechanical theory of development explain why populations of organisms comprise individuals that are adapted to their conditions of existence. (shrink)

Philosophers of biology have been absorbed by the problem of defining evolutionary fitness since Darwin made it central to biological explanation. The apparent problem is obvious. Define fitness as some biologists implicitly do, in terms of actual survival and reproduction, and the principle of natural selection turns into an empty tautology: those organisms which survive and reproduce in larger numbers, survive and reproduce in larger numbers. Accordingly, many writers have sought to provide a definition for ‘fitness’ which avoid this outcome. (...) In particular the definition of fitness as a probabilistic propensity has been widely favored.1 Others, recognizing that no definition both correct and complete can actually be provided, have accepted the consequence that the leading principle of the theory is a definitional truth and attempted to mitigate the impact of this outcome for the empirical character of the theory.2 Still others have argued that ‘fitness’ is properly viewed as a term undefined in the theory of natural selection (on the model of mass—a term undefined in Newtonian mechanics).3 But few have contemplated the solution to this problem proposed by Mohan Matthen and André Ariew (hereafter, MA), in.. (shrink)

argue that natural selection does not explain the genotypic arid phenotypic properties of individuals. On this view, natural selection explains the adaptedness of individuals, not by explaining why the individuals that exist have the adaptations they do, but rather by explaining why the individuals that exist are the ones with those adaptations. This paper argues that this ‘Negative’ view of natural selection ignores the fact that natural selection is a cumulative selection process. So understood, it explains how the genetic sequences (...) that individuals inherit and that are responsible for their complex (and co-adapted) adaptations first arose in the gene-pool. (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 “Two Ways of Thinking About Fitness and Natural Selection” (Matthen and Ariew [2002]; henceforth “Two Ways”), we asked how one should think of the relationship between the various factors invoked to explain evolutionary change – selection, drift, genetic constraints, and so on. We suggested that these factors are not related to one another as “forces” are in classical mechanics. We think it incoherent, for instance, to think of natural selection and drift as separate and opposed “forces” in evolutionary change (...) – that it makes sense to say, for instance, that selection contributed 80% to the actual evolutionary history of the human eye, and drift only 20%. We proposed instead a statistical view of the Theory of Evolution, a view in which fitness is not a cause of evolution, but rather a measure of growth. We also argued for a “hierarchical realization model” for thinking about the relationship between evolutionary factors such as those mentioned above, and suggested that in a “fully specified model”, as we call it below, there is no distinction between natural selection and evolution. (shrink)

In a recent paper (1994) Wesley Salmon has replied to criticisms (e.g., Dowe 1992c, Kitcher 1989) of his (1984) theory of causality, and has offered a revised theory which, he argues, is not open to those criticisms. The key change concerns the characterization of causal processes, where Salmon has traded "the capacity for mark transmission" for "the transmission of an invariant quantity." Salmon argues against the view presented in Dowe (1992c), namely that the concept of "possession of a conserved quantity" (...) is sufficient to account for the difference between causal and pseudo processes. Here that view is defended, and important questions are raised about the notion of transmission and about gerrymandered aggregates. (shrink)

This paper examines contemporary attempts to explicate the explanatory role of mathematics in the physical sciences. Most such approaches involve developing so-called mapping accounts of the relationships between the physical world and mathematical structures. The paper argues that the use of idealizations in physical theorizing poses serious difficulties for such mapping accounts. A new approach to the applicability of mathematics is proposed.

Among the liveliest disputes in evolutionary biology today are disputes concerning the role of chance in evolution--more specifically, disputes concerning the relative evolutionary importance of natural selection vs. so-called "random drift". The following discussion is an attempt to sort out some of the broad issues involved in those disputes. In the first half of this paper, I try to explain the differences between evolution by natural selection and evolution by random drift. On some common construals of "natural selection", those two (...) modes of evolution are completely indistinguishable. Even on a proper construal of "natural selection", it is difficult to distinguish between the "improbable results of natural selection" and evolution by random drift. In the second half of this paper, I discuss the variety of positions taken by evolutionists with respect to the evolutionary importance of random drift vs. natural selection. I will then consider the variety of issues in question in terms of a conceptual distinction often used to describe the rise of probabilistic thinking in the sciences. I will argue, in particular, that what is going on here is not, as might appear at first sight, just another dispute about the desirability of "stochastic" vs. "deterministic" theories. Modern evolutionists do not argue so much about whether evolution is stochastic, but about how stochastic it is. (shrink)

The so-called "adaptationism" of mainstream evolutionary biology has been criticized from a variety of sources. One, which has received relatively little philosophical attention, is developmental biology. Developmental constraints are said to be neglected by adaptationists. This paper explores the divergent methodological and explanatory interests that separate mainstream evolutionary biology from its embryological and developmental critics. It will focus on the concept of constraint itself; even this central concept is understood differently by the two sides of the dispute.

Some properties are causally relevant for a certain effect, others are not. In this paper we describe a problem for our understanding of this notion and then offer a solution in terms of the notion of a program explanation.

When philosophers defend epiphenomenalist doctrines, they often do so by way of a priori arguments. Here we suggest an empirical approach that is modeled on August Weismann’s experimental arguments against the inheritance of acquired characters. This conception of how epiphenomenalism ought to be developed helps clarify some mistakes in two recent epiphenomenalist positions – Jaegwon Kim’s (1993) arguments against mental causation, and the arguments developed by Walsh (2000), Walsh, Lewens, and Ariew (2002), and Matthen and Ariew (2002) that natural selection (...) and drift are not causes of evolution. A manipulationist account of causation (Woodward 2003) leads naturally to an account of how macro- and micro-causation are related and to an understanding of how epiphenomenalism at different levels of organization should be understood. (shrink)

Biologists studying complex causal systems typically identify some factors as causes and treat other factors as background conditions. For example, when geneticists explain biological phenomena, they often foreground genes and relegate the cellular milieu to the background. But factors in the milieu are as causally necessary as genes for the production of phenotypic traits, even traits at the molecular level such as amino acid sequences. Gene-centered biology has been criticized on the grounds that because there is parity among causes, the (...) “privileging” of genes reflects a reductionist bias, not an ontological difference. The idea that there is an ontological parity among causes is related to a philosophical puzzle identified by John Stuart Mill: what, other than our interests or biases, could possibly justify identifying some causes as the actual or operative ones, and other causes as mere background? The aim of this paper is to solve this conceptual puzzle and to explain why there is not an ontological parity among genes and the other factors. It turns out that solving this puzzle helps answer a seemingly unrelated philosophical question: what kind of causal generality matters in biology? (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)