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)

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)

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)

The philosophical theory of scientific explanation proposed here involves a radically new treatment of causality that accords with the pervasively statistical character of contemporary science. Wesley C. Salmon describes three fundamental conceptions of scientific explanation--the epistemic, modal, and ontic. He argues that the prevailing view is untenable and that the modal conception is scientifically out-dated. Significantly revising aspects of his earlier work, he defends a causal/mechanical theory that is a version of the ontic conception. Professor Salmon's theory furnishes a robust (...) argument for scientific realism akin to the argument that convinced twentieth-century physical scientists of the existence of atoms and molecules. To do justice to such notions as irreducibly statistical laws and statistical explanation, he offers a novel account of physical randomness. The transition from the "reviewed view" of scientific explanation to the causal/mechanical model requires fundamental rethinking of basic explanatory concepts. (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)

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)