Roberta L. Millstein

I argue that the propensity interpretation of fitness, properly understood, not only solves the explanatory circularity problem and the mismatch problem, but can also withstand the Pandora’s box full of problems that have been thrown at it. Fitness is the propensity (i.e., probabilistic ability, based on heritable physical traits) for organisms or types of organisms to survive and reproduce in particular environments and in particular populations for a specified number of generations; if greater than one generation, “reproduction” includes descendants of descendants. Fitness values can be described in terms of distributions of propensities to produce varying number of offspring and can be modeled for any number of generations using computer simulations, thus providing both predictive power and a means for comparing the fitness of different phenotypes. Fitness is a causal concept, most notably at the population level, where fitness differences are causally responsible for differences in reproductive success. Relative fitness is ultimately what matters for natural selection.

The “land community” (or “biotic community”) that features centrally in Aldo Leopold’s Land Ethic has typically been equated with the concept of “ecosystem.” Moreover, some have challenged this central Leopoldean concept given the multitude of meanings of the term “ecosystem” and the changes the term has undergone since Leopold’s time (see, e.g., Shrader-Frechette 1996). Even one of Leopold’s primary defenders, J. Baird Callicott, asserts that there are difficulties in identifying the boundaries of ecosystems and suggests that we recognize that their boundaries are determined by scientific questions ecologists pose (Callicott 2013). I argue that we need to rethink Leopold’s concept of land community in the following ways. First, we should recognize that Leopold’s views are not identical to those of his contemporaries (e.g., Clements, Elton), although they resemble those of some subsequent ecologists, including some of our contemporaries (e.g., O’Neill 2001, Post et al. 2007, Hastings and Gross 2012). Second, the land community concept does not map cleanly onto the concept of “ecosystem”; it also incorporates elements of the “community” concept in community ecology by emphasizing the interactions between organisms and not just the matter/energy flow of the ecosystem concept. Third, the boundary question can be illuminated by considering some of the recent literature on the nature of biological individuals (in particular, Odenbaugh 2007; Hamilton, Smith, and Haber 2009; Millstein 2009), focusing on concentrations of causal relations as determinative of the boundaries of the land community qua individual. There are challenges to be worked out, particularly when the interactions of community members do not map cleanly onto matter/energy flows, but I argue that these challenges can be resolved. The result is a defensible land community concept that is ontologically robust enough to be a locus of moral obligation while being consistent with contemporary ecological theory and practice.

In Darwin’s Sacred Cause, Adrian Desmond and James Moore contend that “Darwin would put his utmost into sexual selection because the subject intrigued him, no doubt, but also for a deeper reason: the theory vindicated his lifelong commitment to human brotherhood”. Without questioning Desmond and Moore’s evidence, I will raise some puzzles for their view. I will show that attention to the structure of Darwin’s arguments in the Descent of Man shows that they are far from straightforward. As Desmond and Moore note, Darwin seems to have intended sexual selection in non-human animals to serve as evidence for sexual selection in humans. However, Darwin’s account of sexual selection in humans was different from the canonical cases that Darwin described at great length. If explaining the origin of human races was the main reason for introducing sexual selection, and if sexual selection was a key piece of Darwin’s anti-slavery arguments, then it is puzzling why Darwin would have spent so much time discussing cases that did not really support his argument for the origin of human races, and it is also puzzling that his argument for the origin of human races would be so poor.

This paper aims to illustrate one of the primary goals of the philosophy of biology⎯namely, the examination of central concepts in biological theory and practice⎯through an analysis of the concepts of population and metapopulation in evolutionary biology and ecology. I will first provide a brief background for my analysis, followed by a characterization of my proposed concepts: the causal interactionist concepts of population and metapopulation. I will then illustrate how the concepts apply to six cases that differ in their population structure; this analysis will also serve to flesh out and defend the concepts a bit more. Finally, I will respond to some possible questions that my analysis may have raised and then conclude briefly.

Genetic drift (variously called “random drift”, “random genetic drift”, or sometimes just “drift”) has been a source of ongoing controversy within the philosophy of biology and evolutionary biology communities, to the extent that even the question of what drift is has become controversial. There seems to be agreement that drift is a chance (or probabilistic or statistical) element within population genetics and within evolutionary biology more generally, and that the term “random” isn’t invoking indeterminism or any technical mathematical meaning, but that’s about where agreement ends. Yet genetic drift models are a staple topic in population genetics textbooks and research, with genetic drift described as one of the main factors of evolution alongside selection, mutation, and migration. Some claim that genetic drift has played a major role in evolution (particularly molecular evolution), while others claim it to be minor. This article examines these and other controversies. In order to break through the logjam of competing definitions of drift, this entry begins with a brief history of the concept, before examining various philosophical claims about the proper characterization of drift and whether it can be distinguished from natural selection; the relation of drift to debates over statisticalism; whether drift can be detected empirically and if so, how; and the proper understanding of drift as a model and as a (purported) law.

Recently, a number of philosophers of biology have endorsed views about random drift that, we will argue, rest on an implicit assumption that the meaning of concepts such as drift can be understood through an examination of the mathematical models in which drift appears. They also seem to implicitly assume that ontological questions about the causality of terms appearing in the models can be gleaned from the models alone. We will question these general assumptions by showing how the same equation — the simple 2 = p2 + 2pq + q2 — can be given radically different interpretations, one of which is a physical, causal process and one of which is not. This shows that mathematical models on their own yield neither interpretations nor ontological conclusions. Instead, we argue that these issues can only be resolved by considering the phenomena that the models were originally designed to represent and the phenomena to which the models are currently applied. When one does take those factors into account, starting with the motivation for Sewall Wright’s and R.A. Fisher’s early drift models and ending with contemporary applications, a very different picture of the concept of drift emerges. On this view, drift is a term for a set of physical processes, namely, indiscriminate sampling processes

Biologists studying ecology and evolution use the term “population” in many different ways. Yet little philosophical analysis of the concept has been done, either by biologists or philosophers, in contrast to the voluminous literature on the concept of “species.” This is in spite of the fact that “population” is arguably a far more central concept in ecological and evolutionary studies than “species” is. The fact that such a central concept has been employed in so many different ways is potentially problematic for the reason that inconsistent usages (especially when the usage has not been made explicit) might lead to false controversies in which disputants are simply talking past one another. However, the inconsistent usages are not the only, or even the most important reason to examine the concept. If any set of organisms is legitimately called a “population,” selection and drift processes become purely arbitrary, too. Moreover, key ecological variables, such as abundance and distribution, depend on a nonarbitrary way of identifying populations. I sketch the beginnings of a population concept, drawing inspiration from the Ghiselin-Hull individuality thesis, and show why some alternative approaches are nonstarters.

Recent work on the heat-shock protein Hsp90 by Rutherford and Lindquist (1998) has been included among the pieces of evidence taken to show the essential role of developmental processes in evolution; Hsp90 acts as a buffer against phenotypic variation, allowing genotypic variation to build. When the buffering capacity of Hsp90 is altered (e.g., in nature, by mutation or environmental stress), the genetic variation is "revealed," manifesting itself as phenotypic variation. This phenomenon raises questions about the genetic variation before and after what I will call a "revelation event": Is it neutral, nearly neutral, or non-neutral (i.e., strongly deleterious or strongly advantageous)? Moreover, what kinds of evolutionary processes do we take to be at work? Rutherford and Lindquist (1998) focus on the implications of non-neutral variation and selection. Later work by Queitsch, Sangster, and Lindquist (2002) and Sangster, Lindquist, and Queitsch (2004) raises the possibility that Hsp90 buffering may play the role that was played by drift in Sewall Wright's shifting balance model, permitting transition from one adaptive peak to another. However, Ohta (2002) suggests that much of this variation may be nearly neutral, which in turn, would imply a strong role for drift as well as selection. The primary goal of this paper is to illuminate the alternative scenarios and the processes operating in each. At the end, I raise the possibility of a synthesis between evo-devo and nearly neutral evolution.

Recently, much philosophical discussion has centered on the best way to characterize the concepts of random drift and natural selection, and, in particular, whether selection and drift can be conceptually distinguished (Beatty, 1984; Brandon, 2005; Hodge, 1983, 1987; Millstein, 2002, 2005; Pfeifer, 2005; Shanahan, 1992; Stephens, 2004). These authors all contend, to a greater or lesser degree, that their concepts make sense of biological practice. So it should be instructive to see how the concepts of drift and selection were distinguished by the disputants in a high-profile debate; debates such as these often force biologists to take a more philosophical turn, discussing the concepts at issue in greater detail than usual. Moreover, it is important to consider a debate where the disputants are actually trying to apply the models of population genetics to natural populations; only then can their proper interpretations become fully apparent. (Indeed, I contend that some of the philosophical confusion has arisen because authors have considered only the models themselves, and not the phenomena that the models are attempting to represent). A prime candidate for just such a case study is what Provine (1986) has termed “The Great Snail Debate,” that is, the debates over the highly polymorphic land snails Cepaea nemoralis and C. hortensis in the 1950s and early 1960s. These studies represent one of the best, if not the best, of the early attempts to demonstrate drift in natural populations.

As a number of biologists and philosophers have emphasized, ‘chance’ has multiple meanings in evolutionary biology. Seven have been identified. I will argue that there is a unified concept of chance underlying these seven, which I call the UCC (Unified Chance Concept). I will argue that each is characterized by which causes are consid- ered, ignored, or prohibited. Thus, chance in evolutionary biology can only be under- stood through understanding the causes at work. The UCC aids in comparing the different concepts and allows us to characterize our concepts of chance in probabilistic terms, i.e. provides a way to translate between ‘chance’ and ‘probability’.

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.

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

The neutral and nearly neutral theories of molecular evolution are sometimes characterized as theories about drift alone, where drift is described solely as an outcome, rather than a process. We argue, however, that both selection and drift, as causal processes, are integral parts of both theories. However, the nearly neutral theory explicitly recognizes alleles and/or molecular substitutions that, while engaging in weakly selected causal processes, exhibit outcomes thought to be characteristic of random drift. A narrow focus on outcomes obscures the significant role of weakly selected causal processes in the nearly neutral theory.

Given the role of values in the deployment of GMOs, given the lack of mandatory and long-term testing of GMOs with outside oversight, and given the demonstrated environmental harms, it is not anti-science to want to GMOs labelled as GMOs. People who would like to avoid GMOs, whether out of concerns for potential health harms or concerns over actual environmental harms, are not being allowed to judge the risks and make choices for themselves and their families. For these reasons—so that people can follow their reasonably held values—we ought to label GMOs as GMOs.

This paper explores whether natural selection, a putative evolutionary mechanism, and a main one at that, can be characterized on either of the two dominant conceptions of mechanism, due to Glennan and the team of Machamer, Darden, and Craver, that constitute the “new mechanistic philosophy.” The results of the analysis are that neither of the dominant conceptions of mechanism adequately captures natural selection. Nevertheless, the new mechanistic philosophy possesses the resources for an understanding of natural selection under the rubric.

This volume addresses fundamental issues in the philosophy of science in the context of two most intriguing fields: biology and economics. Written by authorities and experts in the philosophy of biology and economics, Mechanism and Causality in Biology and Economics provides a structured study of the concepts of mechanism and causality in these disciplines and draws careful juxtapositions between philosophical apparatus and scientific practice. By exploring the issues that are most salient to the contemporary philosophies of biology and economics and by presenting comparative analyses, the book serves as a platform not only for gaining mutual understanding between scientists and philosophers of the life sciences and those of the social sciences, but also for sharing interdisciplinary research that combines both philosophical concepts in both fields. The book begins by defining the concepts of mechanism and causality in biology and economics, respectively. The second and third parts investigate philosophical perspectives of various causal and mechanistic issues in scientific practice in the two fields. These two sections include chapters on causal issues in the theory of evolution; experiments and scientific discovery; representation of causal relations and mechanism by models in economics. The concluding section presents interdisciplinary studies of various topics concerning extrapolation of life sciences and social sciences, including chapters on the philosophical investigation of conjoining biological and economic analyses with, respectively, demography, medicine and sociology.

Population genetics attempts to measure the influence of the causes of evolution, viz., mutation, migration, natural selection, and random genetic drift, by understanding the way those causes change the genetics of populations. But how does it accomplish this goal? After a short introduction, we begin in section (2) with a brief historical outline of the origins of population genetics. In section (3), we sketch the model theoretic structure of population genetics, providing the flavor of the ways in which population genetics theory might be understood as incorporating causes. In sections (4) and (5) we discuss two specific problems concerning the relationship between population genetics and evolutionary causes, viz., the problem of conceptually distinguishing natural selection from random genetic drift, and the problem of interpreting fitness. In section (6), we briefly discuss the methodology and key epistemological problems faced by population geneticists in uncovering the causes of evolution. Section (7) of the essay contains concluding remarks.

In “‘Population’ is Not a Natural Kind of Kinds,” Jacob Stegenga argues against the claim that the concept of “population” is a natural kind and in favor of conceptual pluralism, ostensibly in response to two papers of mine (Millstein 2009, 2010). Pluralism is often an attractive position in the philosophy of science. It certainly is a live possibility for the concept of population in ecology and evolutionary biology, and I welcome the opportunity to discuss the topic further. However, I argue that the case for conceptual pluralism has not yet been made. In what follows, I first clarify the issues at stake before taking up the topic of conceptual pluralism and responding to Stegenga’s criticisms of the causal interactionist population concept.

Many philosophers have become familiar with Leopold’s land ethic through the writings of J. Baird Callicott, who claims that Leopold bases his land ethic on a ‘protosociobiological’ argument that Darwin gives in the Descent of Man. On this view, which has become the canonical interpretation, Leopold’s land ethic is based on extending our moral sentiments to ecosystems. I argue that the evidence weighs in favor of an alternative interpretation of Leopold; his reference to Darwin does not refer to the Descent, but rather to the Origin of Species, where Darwin discusses the interdependencies between organisms in the struggle for existence.

A number of areas of biology raise questions about what is of value in the natural environment and how we ought to behave towards it: conservation biology, environmental science, and ecology, to name a few. Based on my experience teaching students from these and similar majors, I argue that the field of environmental ethics has much to teach these students. They come to me with pent-up questions and a feeling that more is needed to fully engage in their subjects, and I believe some exposure to environmental ethics can help focus their interests and goals. I identify three primary areas in which environmental ethics can con- tribute to their education. The first is an examination of who (or what) should be considered to be part of our moral community (i.e., the community to whom we owe direct duties). Is it humans only? Or does it include all sentient life? Or all life? Or ecosystems considered holistically? Often, readings implicitly assume one or more of these answers; the goal is to make the student more sensitive to these implicit claims and to get them to think about the different reasons that support them. The second area, related to the first, is the application of the different answers concerning the extent of the ethical community to real environmental issues and problems. Students need to be aware of how the different answers concerning the moral community can imply conflicting answers for how we should act in certain cases and to think about ways to move toward conflict resolution. The third area in which environmental ethics can contribute is a more conceptual one, focusing on central concepts such as biodiversity, sustainability, species, and ecosystems. Exploring and evaluating various meanings of these terms will make students more reflective and thoughtful citizens and biologists, sensitive to the implications that different conceptual choices make.

I respond to Brandon's (2005) criticisms of my earlier (2002) essay. I argue that (1) biologists are inconsistent in their use of the terms 'selection' and 'drift' -- vacillating between 'process' and 'outcome' -- but that the process-oriented definitions I defend make better sense of the neutralist/selectionist debate; (2) Brandon's purported demonstration that there is no qualitative difference between drift and selection as processes begs the question against my account; and (3) biologists (e.g., Kimura) have argued for genuinely neutral variants. Whether any such variants actually exist is an empirical question. However, the philosophical question at hand is conceptual, not empirical.