A large percentage of the work on the ethics of genetically modified (GM) food has focused on its potentially harmful effects on human health and on the rights of consumers to have their food labeled. A smaller, but still significant percentage has focused instead on the potential environmental impacts of genetically modified food. For example, some authors argue that genetically modified food could lead to a loss of genetic diversity within a particular food crop, leaving that food crop vulnerable to (...) extinction (see, e.g., Lappé and Baily 2002). Another type of argument focuses on potential side effects of genetically modified food. Perhaps there will be gene flow from genetically engineered herbicide resistant crops to a weedy relative, producing a “superweed” that could lead to a “bioinvasion” that takes over an entire ecosystem (see, e.g., Shiva 2002). And Monarch butterflies made headlines when it appeared that corn that was modified to kill corn borers also killed the butterflies, leading to concerns that other species could suffer harm at the hands of genetically modified food as well. (shrink)
1. Drift and selection can be distinguished conceptually. 2. Selection and drift are physical, biological phenomena. 3. Drift and selection can occur simultaneously in a population. 4. Selection and drift should be characterized as processes (see #1), not outcomes. 5. Distinguishing between selection and drift empirically is difficult, but is (sometimes) not impossible. 6. Selection and drift are population-level causal processes.
The status of population genetics has become hotly debated among biologists and philosophers of biology. Many seem to view population genetics as relatively unchanged since the Modern Synthesis and have argued that subjects such as development were left out of the Synthesis. Some have called for an extended evolutionary synthesis or for recognizing the insignificance of population genetics. Yet others such as Michael Lynch have defended population genetics, declaring "nothing in evolution makes sense except in the light of population genetics" (...) (a twist on Dobzhansky's famous slogan that "nothing in biology makes sense except in the light of evolution"). Missing from this discussion is the use of population genetics to shed light on ecology and vice versa, beginning in the 1940s and continuing until the present day. I highlight some of that history through an overview of traditions such as ecological genetics and population biology, followed by a slightly more in-depth look at a contemporary study of the endangered California Tiger Salamander. I argue that population genetics is a powerful and useful tool that continues to be used and modified, even if it isn't required for all evolutionary explanations or doesn't incorporate all the causal factors of evolution. (shrink)
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. (shrink)
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’’ (2009: p. 360). Without questioning Des- mond 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 (atypically) poor. (shrink)
Sex and sensibility: The role of social selection Content Type Journal Article DOI 10.1007/s11016-010-9464-6 Authors Erika L. Milam, Department of History, University of Maryland, 2115 Francis Scott Key Hall, College Park, MD 20742, USA Roberta L. Millstein, Department of Philosophy, University of California, Davis, One Shields Avenue, Davis, CA 95616, USA Angela Potochnik, Department of Philosophy, University of Cincinnati, P.O. Box 210374, Cincinnati, OH 45221, USA Joan E. Roughgarden, Department of Biology, Stanford University, Stanford, CA 94305-5020, USA Journal Metascience Online (...) ISSN 1467-9981 Print ISSN 0815-0796. (shrink)
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’. (shrink)
A review of _Biology’s First Law: The Tendency for Diversity and Complexity to Increase in Evolutionary Systems_, by Daniel W. McShea and Robert N. Brandon. This review argues that the supposed "Zero-Force Evolutionary Law" (ZFEL) is neither a law nor zero-force.
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. (shrink)
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. (shrink)
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. (shrink)
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. (shrink)
Recently, a number of philosophers of biology (e.g., Matthen and Ariew 2002; Walsh, Lewens, and Ariew 2002; Pigliucci and Kaplan 2006; Walsh 2007) 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 (or lack thereof) 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 (p + q)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 (Beatty 1984; Hodge 1987; Millstein 2002, 2005). (shrink)
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. (shrink)
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 (always) impossible. (shrink)
We live in interesting times. Two well-known biologists — E. O. Wilson and Richard Dawkins — and some of their well-known colleagues, who used to employ broadly similar selection models, now deeply disagree over the role of group selection in the evolution of eusociality (or so we argue). Yet they describe their models as interchangeable. As philosophers of biology, we wonder whether there is substantial (i.e., empirical) disagreement here at all, and, if there is, what is this disagreement about? We (...) argue that a substantial disagreement over the processes that caused eusociality best explains this debate, yet the common practice of using overarching definitions for “group selection” and “kin selection” renders empirical differences difficult to detect. We suggest Michael J. Wade’s use of these terms as a basis for models that reveal different selection processes. Wade’s models predict different outcomes for different processes and thus can be tested. (shrink)
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. (shrink)
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. (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 ; Walsh et al. ). 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 ). 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)
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)
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. (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.
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)
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 is an overview of the philosophy of evolution – past, present, and future. It surveys the following topics: the neutralist/selectionist debate, the adapationist programme and its challenges, sociobiology, contingency, laws of biology, the species category problem, the species taxon problem, the tautology problem, fitness, units of selection.
When philosophers of physics explore the nature of chance, they usually look to quantum mechanics. When philosophers of biology explore the nature of chance, they usually look to microevolutionary phenomena, such as mutation or random drift. What has been largely overlooked is the role of chance in macroevolution. The stochastic models of paleobiology employ conceptions of chance that are similar to those at the microevolutionary level, yet different from the conceptions of chance often associated with quantum mechanics and Laplacean determinism.
Links relating to the history and philosophy of biology, assembled by Roberta L. Millstein: reference works, societies, journals, historians and philosophers of biology with papers online, blogs, other resources in the history and philosophy of biology.
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)
Alexander Rosenberg (1994) claims that the omniscient viewpoint of the evolutionary process would have no need for the concept of random drift. However, his argument fails to take into account all of the processes which are considered to be instances of random drift. A consideration of these processes shows that random drift is not eliminable even given a position of omniscience. Furthermore, Rosenberg must take these processes into account in order to support his claims that evolution is deterministic and that (...) evolutionary biology is an instrumental science. (shrink)