Biologists and philosophers have been extremely pessimistic about the possibility of demonstrating random drift in nature, particularly when it comes to distinguishing random drift from natural selection. However, examination of a historical case-Maxime Lamotte's study of natural populations of the land snail, Cepaea nemoralis in the 1950s - shows that while some pessimism is warranted, it has been overstated. Indeed, by describing a unique signature for drift and showing that this signature obtained in the populations under study, Lamotte was able (...) to make a good case for a significant role for drift. It may be difficult to disentangle the causes of drift and selection acting in a population, but it is not impossible. (shrink)

Evolutionary models can explain the dynamics of populations, how genetic, genotypic, or phenotypic frequencies change with time. Models incorporating chance, or drift, predict specific patterns of change. These are illustrated using classic work on blood types by Cavalli-Sforza and his collaborators in the Parma Valley of Italy, in which the theoretically predicted patterns are exhibited in human populations. These data and the models display properties of ensembles of populations. The explanatory problem needs to be understood in terms of how likely (...) an observed change, in either a population or an ensemble, would be under drift alone; this is fundamentally a matter of chance. Understood in this way, issues of drift and chance undercut most recent philosophical, but not biological, discussions of the role of "genetic drift.". (shrink)

The four case studies on chance in evolution provide a rich source for further philosophical analysis. Among the issues raised are the following: Are there different conceptions of chance at work, or is there a common underlying conception? How can a given concept of chance be distinguished from other chance concepts and from nonchance concepts? How can the occurrence of a given chance process be distinguished empirically from nonchance processes or other chance processes? What role does chance play in evolutionary (...) theory? I argue that in order to answer these questions, a careful distinction between process and outcome must be made; however, the purpose of this essay is not to answer these questions definitively, but rather to elaborate on them and to provide a starting point for further discussion. (shrink)

Do evolutionary processes such as selection and random drift cause evolutionary change, or are they merely convenient ways of describing or summarizing it? Philosophers have lined up on both sides of this question. One recent defense (Reisman and Forber 2005) of the causal status of selection and drift appeals to a manipulability theory of causation. Yet, even if one accepts manipulability, there are still reasons to doubt that genetic drift, in particular, is genuinely causal. We will address two challenges to (...) treating drift as causal within a manipulation framework. We will argue that both challenges ultimately fail, but that they raise interesting and subtle issues about the nature of causation and the differences between selection and drift. ‡Thanks to audiences at the ‘PBDB1' Conference and the 2006 PSA Meeting for valuable feedback. †To contact the authors, please write to: Patrick Forber, Tufts University, Philosophy Department, Miner Hall, Medford, MA 02155; e-mail: [email protected]. (shrink)

The adequacy of Elliott Sober’s analogy between classical mechanics and evolutionary theory—according to which both theories explain via a zero-force law and a set of forces that alter the zero-force state—has been criticized from various points of view. I focus here on McShea and Brandon’s claim that drift shouldn’t be considered a force because it is not directional. I argue that there are a number of different theses that could be meant by this, and show that one of those theses—the (...) idea that drift cannot bias populations to be taken somewhere in the evolutionary space from one generation to the next—is actually false. Not only has this thesis been implicitly assumed in the discussion of the force analogy thus far, but it is also commonly found in a wider range of philosophical and biological texts. I argue that correcting this view, and the usual images associated with it, will thereby bring heuristic benefits that impact the force analogy discussion, but that also go beyond it. (shrink)