Does the brain behave indeterministically? I argue that accounting for ion channels, key functional units in the brain, requires indeterministic models. These models are probabilistic, so the brain does behave indeterministically in a weak sense. I explore the implications of this point for a stronger sense of indeterminism. Ultimately I argue that it is not possible, either empirically or through philosophical argument, to show that the brain is indeterministic in that stronger sense.

There are several different ways in which chance affects evolutionary change. That all of these processes are called “random genetic drift” is in part a due to common elements across these different processes, but is also a product of historical borrowing of models and language across different levels of organization in the biological hierarchy. A history of the concept of drift will reveal the variety of contexts in which drift has played an explanatory role in biology, and will shed light (...) on some of the philosophical controversy surrounding whether drift is a cause of evolutionary change. (shrink)

I examine recent debates in the philosophy of biology over the determinism or indeterminism of the evolutionary process, focusing on two papers in particular: Glymour 2001 and Stamos 2001. I argue that neither of these papers succeeds in making the case for the indeterminism of the evolutionary process, and suggest that what is needed is a detailed analysis of the causal processes at every level from the quantum mechanical to the evolutionary.

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)

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)

Recent debate on the nature of probabilities in evolutionary biology has focused largely on the propensity interpretation of fitness (PIF), which defines fitness in terms of a conception of probability known as “propensity”. However, proponents of this conception of fitness have misconceived the role of probability in the constitution of fitness. First, discussions of probability and fitness have almost always focused on organism effect probability, the probability that an organism and its environment cause effects. I argue that much of the (...) probability relevant to fitness must be organism circumstance probability, the probability that an organism encounters particular, detailed circumstances within an environment, circumstances which are not the organism’s effects. Second, I argue in favor of the view that organism effect propensities either don’t exist or are not part of the basis of fitness, because they usually have values close to 0 or 1. More generally, I try to show that it is possible to develop a clearer conception of the role of probability in biological processes than earlier discussions have allowed. (shrink)

Naturalized metaphysics aims to establish justified metaphysical claims, where metaphysics is meant to carry its usual significance, while avoiding the traditional methods of metaphysics—a priori reasoning, conceptual analysis, intuitions, and common sense—which naturalized metaphysics argues are not epistemically probative. After offering an explication of what it means to do metaphysics, this paper argues that naturalized metaphysics, at the outset, is hospitable to doing metaphysics. The underdetermination of metaphysics by science, however, changes the picture. Naturalized metaphysics has to break this underdetermination, (...) but the criticism of the traditional methods of metaphysics leaves no resources with which to do so. Naturalized metaphysics must therefore be more restrictive than originally intended to ensure that some metaphysical features avoid underdetermination. In this restrictive naturalized metaphysics, however, metaphysicians are only left the task of surveying the opinions of scientists which, it is argued, does not qualify as doing metaphysics. Thus, to fulfill its promise to save metaphysics, naturalized metaphysics displaces the metaphysician. Furthermore, the attempt to re-employ them via the principle of naturalistic closure is argued to fail. Metaphysicians should therefore not be happier with naturalized metaphysics than they are with the more explicitly eliminative trends in contemporary metametaphysics, such as neo-Carnapian deflationism, despite the promise of naturalized metaphysics, likely to Carnap’s dismay, to deliver justified claims about ultimate reality. (shrink)

Philosophers of biology have worked extensively on how we ought best to interpret the probabilities which arise throughout evolutionary theory. In spite of this substantial work, however, much of the debate has remained persistently intractable. I offer the example of Bayesian models of divergence time estimation as a case study in how we might bring further resources from the biological literature to bear on these debates. These models offer us an example in which a number of different sources of uncertainty (...) are combined to produce an estimate for a complex, unobservable quantity. These models have been carefully analyzed in recent biological work, which has determined the relationship between these sources of uncertainty, both quantitatively and qualitatively. I suggest here that this case shows us the limitations of univocal analyses of probability in evolution, as well as the simple dichotomy between “subjective” and “objective” probabilities, and I conclude by gesturing toward ways in which we might introduce more sophisticated interpretive taxonomies of probability as a path toward advancing debates on probability in the life sciences. (shrink)

The notion of fitness is usually equated to reproductive success. However, this actualist approach presents some difficulties, mainly the explanatory circularity problem, which have lead philosophers of biology to offer alternative definitions in which fitness and reproductive success are distinguished. In this paper, we argue that none of these alternatives is satisfactory and, inspired by Mumford and Anjum’s dispositional theory of causation, we offer a definition of fitness as a causal dispositional property. We argue that, under this framework, the distinctiveness (...) that biologists usually attribute to fitness—namely, the fact that fitness is something different from both the physical traits of an organism and the number of offspring it leaves—can be explained, and the main problems associated with the concept of fitness can be solved. Firstly, we introduce Mumford and Anjum's dispositional theory of causation and present our definition of fitness as a causal disposition. We explain in detail each of the elements involved in our definition, namely: the relationship between fitness and the functional dispositions that compose it, the emergent character of fitness, and the context-sensitivity of fitness. Finally, we explain how fitness and realized fitness, as well as expected and realized fitness are distinguished in our approach to fitness as a causal disposition. (shrink)

Evolutionary theory (ET) is teeming with probabilities. Probabilities exist at all levels: the level of mutation, the level of microevolution, and the level of macroevolution. This uncontroversial claim raises a number of contentious issues. For example, is the evolutionary process (as opposed to the theory) indeterministic, or is it deterministic? Philosophers of biology have taken different sides on this issue. Millstein (1997) has argued that we are not currently able answer this question, and that even scientific realists ought to remain (...) agnostic concerning the determinism or indeterminism of evolutionary processes. If this argument is correct, it suggests that, whatever we take probabilities in ET to be, they must be consistent with either determinism or indeterminism. This raises some interesting philosophical questions: How should we understand the probabilities used in ET? In other words, what is meant by saying that a certain evolutionary change is more or less probable? Which interpretation of probability is the most appropriate for ET? I argue that the probabilities used in ET are objective in a realist sense, if not in an indeterministic sense. Furthermore, there are a number of interpretations of probability that are objective and would be consistent with ET under determinism or indeterminism. However, I argue that evolutionary probabilities are best understood as propensities of population-level kinds. (shrink)

Evolution in its contemporary meaning in biology typically refers to the changes in the proportions of biological types in a population over time (see the entry on the concept of evolution to 1872 for earlier meanings). As evolution is too large of a topic to address thoroughly in one entry, the primary goal of this entry is to serve as a broad overview of contemporary issues in evolution with links to other entries where more in-depth discussion can be found. The (...) entry begins with a brief survey of definitions of evolution, followed by a discussion of the different modes of evolution and related philosophical issues, and ends with a summary of other topics in the philosophy of evolution focusing particularly on topics covered in this encyclopedia. (shrink)

This work is a critical consideration of several arguments recently given by Elliott Sober that are aimed at undermining the Laplacean stance on probability in evolutionary theory. The Laplacean contends that the only objective probability an event has is the one assigned to it by a complete description of the relevant microparticles. Sober alleges a formal demonstration that the Laplacean stance on probability in evolutionary theory is inconsistent. But Sober’s argument contains a crucial lacuna, one that likely cannot be repaired (...) to yield the conclusion that Sober draws. He also argues that the Laplacean is committed to inferring semantic facts about probabilities from pragmatic facts about agents’ reasons for using probabilities. But Sober’s arguments against inferring semantic facts from pragmatic constraints would only undermine one basis for Laplaceanism. The Laplacean who formulates her position carefully need not leave herself vulnerable to Sober’s objection to inferring semantics from pragmatics. (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)

Classical mechanics is empirically successful because the probabilistic mean values of quantum mechanical observables follow the classical equations of motion to a good approximation (Messiah 1970, 215). We examine this claim for the one-dimensional motion of a particle in a box, and extend the idea by deriving a special case of the ideal gas law in terms of the mean value of a generalized force used to define "pressure." The examples illustrate the importance of probabilistic averaging as a method of (...) abstracting away from the messy details of microphenomena, not only in physics, but in other sciences as well. (shrink)

Discussions of “chance” and related concepts are found throughout philosophical work on evolutionary theory. By drawing attention to three very commonly-recognized distinctions, I separate four independent concepts falling under the broad heading of “chance”: randomness, epistemic unpredictability, causal indeterminism, and probabilistic causal processes. Far from a merely semantic distinction, however, it is demonstrated that conflation of these obviously distinct notions has an important bearing on debates at the core of evolutionary theory, particularly the debate over the interpretation of fitness, natural (...) selection, and genetic drift. (shrink)

Recent discussion of the statistical character of evolutionary theory has centered around two positions: (1) Determinism combined with the claim that the statistical character is eliminable, a subjective interpretation of probability, and instrumentalism; (2) Indeterminism combined with the claim that the statistical character is ineliminable, a propensity interpretation of probability, and realism. I point out some internal problems in these positions and show that the relationship between determinism, eliminability, realism, and the interpretation of probability is more complex than previously assumed (...) in this debate. Furthermore, I take some initial steps towards a more adequate account of the statistical character of evolutionary theory. (shrink)

The Darwinian theory of evolution is itself evolving and this book presents the details of the core of modern Darwinism and its latest developmental directions. The authors present current scientific work addressing theoretical problems and challenges in four sections, beginning with the concepts of evolution theory, its processes of variation, heredity, selection, adaptation and function, and its patterns of character, species, descent and life. The second part of this book scrutinizes Darwinism in the philosophy of science and its usefulness in (...) understanding ecosystems, whilst the third section deals with its application in disciplines beyond the biological sciences, including evolutionary psychology and evolutionary economics, Darwinian morality and phylolinguistics. The final section addresses anti-Darwinism, the creationist view and issues around teaching evolution in secondary schools. The reader learns how current experimental biology is opening important perspectives on the sources of variation, and thus of the very power of natural selection. This work examines numerous examples of the extension of the principle of natural selection and provides the opportunity to critically reflect on a rich theory, on the methodological rigour that presides in its extensions and exportations, and on the necessity to measure its advantages and also its limits. Scholars interested in modern Darwinism and scientific research, its concepts, research programs and controversies will find this book an excellent read, and those considering how Darwinism might evolve, how it can apply to the human sciences and other disciplines beyond its origins will find it particularly valuable. Originally produced in French (Les Mondes Darwiniens), the scope and usefulness of the book have led to the production of this English text, to reach a wider audience. This book is a milestone in the impressive penetration by Francophone scholars into the world of Darwinian science, its historiography and philosophy over the last two decades. Alex Rosenberg, R. Taylor Cole Professor of Philosophy, Duke University Until now this useful and comprehensive handbook has only been available to francophones. Thanks to this invaluable new translation, this collection of insightful and original essays can reach the global audience it deserves. Tim Lewens, University of Cambridge. (shrink)

Some have argued that chance and determinism are compatible in order to account for the objectivity of probabilities in theories that are compatible with determinism, like Classical Statistical Mechanics (CSM) and Evolutionary Theory (ET). Contrarily, some have argued that chance and determinism are incompatible, and so such probabilities are subjective. In this paper, I argue that both of these positions are unsatisfactory. I argue that the probabilities of theories like CSM and ET are not chances, but also that they are (...) not subjective probabilities either. Rather, they are a third type of probability, which I call counterfactual probability. The main distinguishing feature of counterfactual-probability is the role it plays in conveying important counterfactual information in explanations. This distinguishes counterfactual probability from chance as a second concept of objective probability. (shrink)

It’s recently been argued that biological fitness can’t change over the course of an organism’s life as a result of organisms’ behaviors. However, some characterizations of biological function and biological altruism tacitly or explicitly assume that an effect of a trait can change an organism’s fitness. In the first part of the paper, I explain that the core idea of changing fitness can be understood in terms of conditional probabilities defined over sequences of events in an organism’s life. The result (...) is a notion of “conditional fitness” which is static but which captures intuitions about apparent behavioral effects on fitness. The second part of the paper investigates the possibility of providing a systematic foundation for conditional fitness in terms of spaces of sequences of states of an organism and its environment. I argue that the resulting “organism–environment history conception” helps unify diverse biological perspectives, and may provide part of a metaphysics of natural selection. (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)

Leslie Graves, Barbara Horan and Alex Rosenberg (1999) have argued that the process of evolution is really deterministic, so we should be instrumentalists about our probabilistic evolutionary theory. I criticize the consistency of their view. I argue that because they are realists towards multiple theories (quantum mechanics and macrophysics) their arguments against realism for another scientific theory fail. The main point of this paper is critical, but in order to set up this criticism I explore the ramifications of realism for (...) multiple theories. Finally, I offer a brief metaphysical justification for realism about multiple theories. This view justifies realism for evolutionary theory, which has been defended by Robert Brandon and Scott Carson (1996). (shrink)

In "The Indeterministic Character of Evolutionary Theory: No 'Hidden Variables Proof' But No Room for Determinism Either," Brandon and Carson (1996) argue that evolutionary theory is statistical because the processes it describes are fundamentally statistical. In "Is Indeterminism the Source of the Statistical Character of Evolutionary Theory?" Graves, Horan, and Rosenberg (1999) argue in reply that the processes of evolutionary biology are fundamentally deterministic and that the statistical character of evolutionary theory is explained by epistemological rather than ontological considerations. In (...) this paper I focus on the topic of mutation. By focusing on some of the theory and research on this topic from early to late, I show how quantum indeterminism hooks up to point mutations (via tautomeric shifts, proton tunneling, and aqueous thermal motion). I conclude with a few thoughts on some of the wider implications of this topic. (shrink)

ABSTRACTIn Darwinian Population and Natural Selection, Peter Godfrey-Smith brought the topic of natural selection back to the forefront of philosophy of biology, highlighting different issues surro...

One finds intertwined with ideas at the core of evolutionary theory claims about frequencies in counterfactual and infinitely large populations of organisms, as well as in sets of populations of organisms. One also finds claims about frequencies in counterfactual and infinitely large populations—of events—at the core of an answer to a question concerning the foundations of evolutionary theory. The question is this: To what do the numerical probabilities found throughout evolutionary theory correspond? The answer in question says that evolutionary probabilities (...) are “hypothetical frequencies” (including what are sometimes called “long-run frequencies” and “long-run propensities”). In this paper, I review two arguments against hypothetical frequencies. The arguments have implications for the interpretation of evolutionary probabilities, but more importantly, they seem to raise problems for biologists’ claims about frequencies in counterfactual or infinite populations of organisms and sets of populations of organisms. I argue that when properly understood, claims about frequencies in large and infinite populations of organisms and sets of populations are not threatened by the arguments. Seeing why gives us a clearer understanding of the nature of counterfactual and infinite population claims and probability in evolutionary theory. (shrink)

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)

Recently, philosophers of biology have debated the status of the evolutionary process: is it deterministic or indeterministic? I argue that there is insufficient reason to favor one side of the debate over the other, and that a more philosophically defensible position argues neither for the determinacy nor for the indeterminacy of the evolutionary process. In other words, I maintain that the appropriate stand to take towards the question of the determinism of the evolutionary process is agnosticism. I then suggest that (...) an examination of the phenomenon of developmental noise might yield a solution to the problem. (shrink)

This volume focuses on various questions concerning the interpretation of probability and probabilistic reasoning in biology and physics. It is inspired by the idea that philosophers of biology and philosophers of physics who work on the foundations of their disciplines encounter similar questions and problems concerning the role and application of probability, and that interaction between the two communities will be both interesting and fruitful. In this introduction we present the background to the main questions that the volume focuses on (...) and summarize the highlights of the individual contributions. (shrink)

This paper deals with the dependence of directionality in the course of events-or our claims concerning such directionality-on the modes of description we use in speaking of the events in question. I argue that criteria of similarity and individuation play a crucial role in assessments of directionality. This is an extension of Davidson's claim regarding the difference between causal and explanatory contexts. The argument is based on a characterisation of notions of necessity and contingency that differ from their modal logic (...) counterparts on the one hand, and from causality and chance on the other. I show that some types of directionality are perfectly compatible with both determinism and indeterminism at the microscopic level, and that there is no likelihood of, or advantage to, reducing such directionality to other laws or causal processes. (shrink)

Recent discussion of the statistical character of evolutionary theory has centered around two positions: Determinism combined with the claim that the statistical character is eliminable, a subjective interpretation of probability, and instrumentalism; Indeterminism combined with the claim that the statistical character is ineliminable, a propensity interpretation of probability, and realism. I point out some internal problems in these positions and show that the relationship between determinism, eliminability, realism, and the interpretation of probability is more complex than previously assumed in this (...) debate. Furthermore, I take some initial steps towards a more adequate account of the statistical character of evolutionary theory. (shrink)