We offer a systematic examination of propensity interpretations of fitness, which emphasizes the role that fitness plays in evolutionary theory and takes seriously the probabilistic character of evolutionary change. We distinguish questions of the probabilistic character of fitness from the particular interpretations of probability which could be incorporated. The roles of selection and drift in evolutionary models support the view that fitness must be understood within a probabilistic framework, and the specific character of organism/environment interactions supports the conclusion that fitness (...) must be understood as a propensity rather than as a limiting frequency. (shrink)

Do the sciences aim to uncover the structure of nature, or are they ultimately a practical means of controlling our environment? In Instrumental Biology, or the Disunity of Science, Alexander Rosenberg argues that while physics and chemistry can develop laws that reveal the structure of natural phenomena, biology is fated to be a practical, instrumental discipline. Because of the complexity produced by natural selection, and because of the limits on human cognition, scientists are prevented from uncovering the basic structure of (...) biological phenomena. Consequently, biology and all of the disciplines that rest upon it--psychology and the other human sciences--must aim at most to provide practical tools for coping with the natural world rather than a complete theoretical understanding of it. (shrink)

The Nature of Selection is a straightforward, self-contained introduction to philosophical and biological problems in evolutionary theory. It presents a powerful analysis of the evolutionary concepts of natural selection, fitness, and adaptation and clarifies controversial issues concerning altruism, group selection, and the idea that organisms are survival machines built for the good of the genes that inhabit them. "Sober's is the answering philosophical voice, the voice of a first-rate philosopher and a knowledgeable student of contemporary evolutionary theory. His book merits (...) broad attention among both communities. It should also inspire others to continue the conversation."-Philip Kitcher, Nature "Elliott Sober has made extraordinarily important contributions to our understanding of biological problems in evolutionary biology and causality. The Nature of Selection is a major contribution to understanding epistemological problems in evolutionary theory. I predict that it will have a long lasting place in the literature."-Richard C. Lewontin. (shrink)

Recent work on self organization promises an explanation of complex order which is independent of adaptation. Self-organizing systems are complex systems of simple units, projecting order as a consequence of localized and generally nonlinear interactions between these units. Stuart Kauffman offers one variation on the theme of self-organization, offering what he calls a ``statistical mechanics'' for complex systems. This paper explores the explanatory strategies deployed in this ``statistical mechanics,'' initially focusing on the autonomy of statistical explanation as it applies in (...) evolutionary settings and then turning to Kauffman's analysis. Two primary morals emerge as a consequence of this examination: first, the view that adaptation and self-organization should be seen as competing theories or models is misleading and simplistic; and second, while we need a synthesis treating self-organization and adaptation as geared toward different problems, at different levels of organization, and deploying different methods, we do not yet have such a synthesis. (shrink)

The latter half of the twentieth century has been marked by debates in evolutionary biology over the relative significance of natural selection and random drift: the so-called “neutralist/selectionist” debates. Yet John Beatty has argued that it is difficult, if not impossible, to distinguish the concept of random drift from the concept of natural selection, a claim that has been accepted by many philosophers of biology. If this claim is correct, then the neutralist/selectionist debates seem at best futile, and at worst, (...) meaningless. I reexamine the issues that Beatty raises, and argue that random drift and natural selection, conceived as processes, can be distinguished from one another. (shrink)

This paper takes a critical look at the idea that evolutionary theory is a statistical theory. It argues that despite the strong instrumental motivation for statistical theories, they are not necessary to explain deterministic systems. Biological evolution is fundamentally a result of deterministic processes. Hence, a statistical theory is not necessary for describing the evolutionary forces of genetic drift and natural selection, nor is it needed for describing the fitness of organisms. There is a computational advantage to the statistical theory (...) of population genetics, but population genetics succeeds only by eliminating causes from its account of evolutionary change. (shrink)

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)

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 a small handful of papers in theoretical population genetics, John Gillespie (2000a, 2000b, 2001) argues that a new stochastic process he calls "genetic draft" is evolutionarily more significant than genetic drift. This case study of chance in evolution explores Gillespie's proposed stochastic evolutionary force and sketches the implications of Gillespie's argument for philosophers' explorations of genetic drift.

I argue that results from foraging theory give us good reason to think some evolutionary phenomena are indeterministic and hence that evolutionary theory must be probabilistic. Foraging theory implies that random search is sometimes selectively advantageous, and experimental work suggests that it is employed by a variety of organisms. There are reasons to think such search will sometimes be genuinely indeterministic. If it is, then individual reproductive success will also be indeterministic, and so too will frequency change in populations of (...) organisms employing such search. (shrink)

In this paper we first briefly review Bell's (1964, 1966) Theorem to see how it invalidates any deterministic "hidden variable" account of the apparent indeterminacy of quantum mechanics (QM). Then we show that quantum uncertainty, at the level of DNA mutations, can "percolate" up to have major populational effects. Interesting as this point may be it does not show any autonomous indeterminism of the evolutionary process. In the next two sections we investigate drift and natural selection as the locus of (...) autonomous biological indeterminacy. Here we conclude that the population-level indeterminacy of natural selection and drift are ultimately based on the assumption of a fundamental indeterminacy at the level of the lives and deaths of individual organisms. The following section examines this assumption and defends it from the determinists' attack. Then we show that, even if one rejects the assumption, there is still an important reason why one might think evolutionary theory (ET) is autonomously indeterministic. In the concluding section we contrast the arguments we have mounted against a deterministic hidden variable account of ET with the proof of the impossibility of such an account of QM. (shrink)

How, according to Darwin, does chance variation affect evolutionary outcomes? In his 1866 book, On the Various Contrivances by which British and Foreign Orchids are Fertilised by Insects, Darwin developed an argument that played an important role in his overall case for evolution by natural selection, as articulated in later editions of the Origin. This argument also figured significantly in Darwin's reflections on the theological dimensions of evolution by natural selection.

We argue that Brandon and Carson's (1996) "The Indeterministic Character of Evolutionary Theory" fails to identify any indeterminism that would require evolutionary theory to be a statistical or probabilistic theory. Specifically, we argue that (1) their demonstration of a mechanism by which quantum indeterminism might "percolate up" to the biological level is irrelevant; (2) their argument that natural selection is indeterministic because it is inextricably connected with drift fails to join the issue with determinism; and (3) their view that experimental (...) methodology in botany assumes indeterminism is both false and incompatible with the commitment to discoverable causal mechanisms underlying biological processes. We remain convinced that the probabilism of the theory of evolution is epistemically, not ontologically, motivated. (shrink)

This article offers three contrasting cases of the use of neutrality and drift in molecular evolution. In the first, neutrality is assumed as a simplest case for modeling. In the second and third, concepts of drift and neutrality are developed within the context of population genetics testing and the development and application of the molecular clock.