Two controversies exist regarding the appropriate characterization of hierarchical and adaptive evolution in natural populations. In biology, there is the Wright-Fisher controversy over the relative roles of random genetic drift, natural selection, population structure, and interdemic selection in adaptive evolution begun by Sewall Wright and Ronald Aylmer Fisher. There is also the Units of Selection debate, spanning both the biological and the philosophical literature and including the impassioned group-selection debate. Why do these two discourses exist separately, and interact relatively little? (...) We postulate that the reason for this schism can be found in the differing focus of each controversy, a deep difference itself determined by distinct general styles of scientific research guiding each discourse. That is, the Wright-Fisher debate focuses on adaptive process, and tends to be instructed by the mathematical modeling style, while the focus of the Units of Selection controversy is adaptive product, and is typically guided by the function style. The differences between the two discourses can be usefully tracked by examining their interpretations of two contested strategies for theorizing hierarchical selection: horizontal and vertical averaging. (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)

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)

The object of this paper is to reply to Morrison's ([2000]) claim that while ‘structural unity’ was achieved at the level of the mathematical models of population genetics in the early synthesis, there was explanatory disunity. I argue to the contrary, that the early synthesis effected by the founders of theoretical population genetics was unifying and explanatory both. Defending this requires a reconsideration of Morrison's notion of explanation. In Morrison's view, all and only answers to ‘why’ questions which include the (...) ‘cause or mechanism’ for some phenomenon count as explanatory. In my view, mathematical demonstrations that answer ‘how possibly’ and ‘why necessarily’ questions may also count as explanatory. The authors of the synthesis explained how evolution was possible on a Mendelian system of inheritance, answered skepticism about the sufficiency of selection, and thus explained why and how a Darwinian research program was warranted. While today we take many of these claims as obvious, they required argument, and part of the explanatory work of the formal sciences is providing such arguments. Surely, Fisher and Wright had competing views as to the optimal means of generating adaptation. Nevertheless, they had common opponents and a common unifying and explanatory goal that their mathematical demonstrations served. Introduction: Morrison's challenge Fisher v. Wright revisited The early synthesis Conclusion: unification and explanation reconciled. (shrink)

Speciation is an aspect of evolutionary biology that has received little philosophical attention apart from articles mainly by biologists such as Mayr (1988). The role of speciation as a terminus a quo for the individuality of species or in the context of punctuated equilibrium theory has been discussed, but not the nature of speciation events themselves. It is the task of this paper to attempt to bring speciation events into some kind of general scheme, based primarily upon the work of (...) Sergey Gavrilets on adaptive landscapes, using migration rate, or gene flow, as the primary scale, and concluding that adaptive and drift explanations are complementary rather than competing. I propose a distinction between intrinsic and extrinsic selection, and the notion of reproductive reach and argue that speciation modes should be discriminated in terms of gene flow, the nature of selection maintaining reproductive reach, and whether the predominant cause is selective or stochastic. I also suggest that the notion of an adaptive “quasispecies” for asexual species is the primitive notion of species, and that members of reproductively coherent sexual species are additionally coadapted to their mating partners. (shrink)

This paper outlines a critique of the use of the genetic variance–covariance matrix (G), one of the central concepts in the modern study of natural selection and evolution. Specifically, I argue that for both conceptual and empirical reasons, studies of G cannot be used to elucidate so-called constraints on natural selection, nor can they be employed to detect or to measure past selection in natural populations – contrary to what assumed by most practicing biologists. I suggest that the search for (...) a general solution to the difficult problem of identifying causal structures given observed correlation’s has led evolutionary quantitative geneticists to substitute statistical modeling for the more difficult, but much more valuable, job of teasing apart the many possible causes underlying the action of natural selection. Hence, the entire evolutionary quantitative genetics research program may be in need of a fundamental reconsideration of its goals and how they correspond to the array of mathematical and experimental techniques normally employed by its practitioners. (shrink)

Recent discussion of mechanism has suggested new approaches to several issues in the philosophy of science, including theory structure, causal explanation, and reductionism. Here, I apply what I take to be the fruits of the 'new mechanical philosophy' to an analysis of a contemporary debate in evolutionary biology about the role of natural selection in speciation. Traditional accounts of that debate focus on the geographic context of genetic divergence--namely, whether divergence in the absence of geographic isolation is possible (or significant). (...) Those accounts are at best incomplete, I argue, because they ignore the mechanisms producing divergence and miss what is at stake in the biological debate. I argue that the biological debate instead concerns the scope of particular speciation mechanisms which assign different roles to natural selection at various stages of divergence. The upshot is a new interpretation of the crux of that debate--namely, whether divergence with gene flow is possible (or significant) and whether the isolating mechanisms producing it are adaptive. (shrink)

Niche construction is the process of organisms changing themselves or their environment—or their relationship with their environment—in ways that affect the evolutionary trajectory of their population. These evolutionary trajectory changes are traditionally understood to be triggered by changes in selection pressures. Niche construction thus necessarily involves organisms altering selection pressures. In this paper, we argue that changes in selection pressures is not the only way organisms can influence the evolutionary futures of their population. We propose that organisms can also affect (...) drift probabilities, and that such changes should be considered niche construction. Drift probabilities can be modulated by altering population size or by affecting driftability (individual variance in possible reproductive outcomes). We consider both and provide examples of how niche construction can stabilize, increase, or dampen drift probabilities. Finally, we revisit and broaden the traditional definition of niche construction. We hold that organismic activities that modify drift probabilities should count as niche construction, even if selection pressures remain unaltered. (shrink)

Empirical adaptationism is often said to be an empirical claim about nature, which concerns the overall relative causal importance of natural selection in evolution compared with other evolutionary factors. Philosophers and biologists who have tried to clarify the meaning of empirical adaptationism usually share, explicitly or implicitly, two assumptions: (1) Empirical adaptationism is an empirical claim that is scientifically testable; (2) testing empirical adaptationism is scientifically valuable. In this article, I challenge these two assumptions and argue that both are unwarranted (...) given how empirical adaptationism is currently formulated. I identify a series of conceptual and methodological difficulties that makes testing empirical adaptationism in a biologically non-arbitrary way virtually impossible. Moreover, I show that those in favor of testing empirical adaptationism have yet to demonstrate the distinctive value and necessity of conducing such a test. My analysis of the case of empirical adaptationism also provides reasons for scientists to reconsider the value and necessity of engaging in scientific debates involving the notion of overall relative causal importance. (shrink)

Recent discussion of mechanism has suggested new approaches to several issues in the philosophy of science, including theory structure, causal explanation, and reductionism. Here, I apply what I take to be the fruits of the Ônew mechanical philosophyÕ to an analysis of a contemporary debate in evolutionary biology about the role of natural selection in speciation. Traditional accounts of that debate focus on the geographic context of genetic divergence— namely, whether divergence in the absence of geographic isolation is possible (or (...) significant). Those accounts are at best incomplete, I argue, because they ignore the mechanisms producing divergence and miss what is at stake in the biological debate. I argue that the biological debate instead concerns the scope of particular speciation mechanisms which assign different roles to natural selection at various stages of divergence. The upshot is a new interpretation of the crux of that debate—namely, whether divergence with gene flow is possible (or significant) and whether the isolating mechanisms producing it are adaptive. Ó 2005 Elsevier Ltd. All rights reserved. (shrink)

This essay examines the origin of genotype-environment interaction, or G×E. "Origin" and not "the origin" because the thesis is that there were actually two distinct concepts of G×E at this beginning: a biometric concept, or \[G \times E_B\], and a developmental concept, or \[G \times E_D \]. R. A. Fisher, one of the founders of population genetics and the creator of the statistical analysis of variance, introduced the biometric concept as he attempted to resolve one of the main problems in (...) the biometric tradition of biology - partitioning the relative contributions of nature and nurture responsible for variation in a population. Lancelot Hogben, an experimental embryologist and also a statistician, introduced the developmental concept as he attempted to resolve one of the main problems in the developmental tradition of biology - determining the role that developmental relationships between genotype and environment played in the generation of variation. To argue for this thesis, I outline Fisher and Hogben's separate routes to their respective concepts of G × E; then these separate interpretations of G × E are drawn on to explicate a debate between Fisher and Hogben over the importance of G × E, the first installment of a persistent controversy. Finally, Fisher's \[G \times E_B\] and Hogben's \[G \times E_D \] are traced beyond their own work into mid-2Oth century population and developmental genetics, and then into the infamous IQ Controversy of the 1970s. (shrink)

The relatively new field of evolutionary developmental biology continues to attract considerable attention from biologists, philosophers, and historians, in part, because work in this field demonstrates that important changes are underway within biology. Though studies of development and evolution were closely connected during the 19th century, continued work in genetics fostered a general split between the two during the first decades of the twentieth century (e.g., Allen 1978; Gilbert 1978; Mayr and Provine 1980; Gilbert, Opitz and..

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. (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)

Scientific disputes about how often different processes or patterns occur are relative frequency controversies. These controversies occur across the sciences. In some areas—especially biology—they are even the dominant mode of dispute. Yet they depart from the standard picture of what a scientific controversy is like. In fact, standard philosophical accounts of scientific controversies suggest that relative frequency controversies are irrational or lacking in epistemic value. This is because standard philosophical accounts of scientific controversies often assume that in order to be (...) rational, a scientific controversy must reach a resolution and be about a scientifically interesting question. Relative frequency controversies rarely reach a resolution, however, and some scientists and philosophers are skeptical that these controversies center on scientifically interesting questions. In this paper, I provide a novel account of the epistemic contribution that relative frequency controversies make to science. I show that these controversies are rational in the sense of furthering the epistemic aims of the scientific communities in which they occur. They do this despite rarely reaching a resolution, and independent of whether the controversies are about scientifically interesting questions. This means that assumptions and about what is required for a controversy to be rational are wrong. Controversies do not need to reach a resolution in order to be rational. And they do not need to be about anything scientifically interesting in order to make valuable epistemic contributions to science. (shrink)

Confirmation in evolutionary biology depends on what biologists take to be the genuine rivals. Investigating what constrains the scope of biological possibility provides part of the story: explaining how possible helps determine what counts as a genuine rival and thus informs confirmation. To clarify the criteria for genuine rivalry I distinguish between global and local constraints on biological possibility, and offer an account of how-possibly explanation. To sharpen the connection between confirmation and explaining how possible I discuss the view that (...) formal inquiry can provide a kind of confirmation-theoretic support for evolutionary models, and offer an example of how-possibly explanation interacting with testing practice. (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)

: Where there are cases of underdetermination in scientific controversies, such as the case of the molecular clock, scientists may direct the course and terms of dispute by playing off the multidimensional framework of theory evaluation. This is because assessment strategies themselves are underdetermined. Within the framework of assessment, there are a variety of trade-offs between different strategies as well as shifting emphases as specific strategies are given more or less weight in assessment situations. When a strategy is underdetermined, scientists (...) can change the dynamics of a controversy by making assessments using different combinations of evaluation strategies and/or weighting whatever strategies are in play in different ways. Following an underdetermination strategy does not end or resolve a scientific dispute. Consequently, manipulating underdetermination is a feature of controversy dynamics and not controversy closure. (shrink)

: This paper explores the calibration of laboratory models in population genetics as an experimental strategy for justifying experimental results and claims based upon them following Franklin (1986, 1990) and Rudge (1996, 1998). The analysis provided undermines Coyne et al.'s (1997) critique of Wade and Goodnight's (1991) experimental study of Wright's (1931, 1932) Shifting Balance Theory. The essay concludes by further demonstrating how this analysis bears on Diamond's (1986) claims regarding the weakness of laboratory experiments as evidence, and further how (...) the calibration strategy fits within Lloyd's (1987, 1988) account of the confirmation of ecological and evolutionary models. (shrink)

The concept of dominance poses several dilemmas. First, while entrenched in genetics education, the metaphor of dominance promotes several misconceptions and misleading cultural perspectives. Second, the metaphors of power, prevalence and competition extend into science, shaping assumptions and default concepts. Third, because genetic causality is complex, the simplified concepts of dominance found in practice are highly contingent or inconsistent. The conceptual problems are illustrated in the history of studies on the evolution of dominance. Conceptual clarity may be fostered, I claim, (...) by viewing diploid organisms as diphenic and by framing genetic causality modestly through individual alleles and their corresponding haplophenotypes. (shrink)