Debates concerning the units and levels of selection have persisted for over fifty years. One major question in this literature is whether units and levels of selection are genuine, in the sense that they are objective features of the world, or merely reflect the interests and goals of an observer. Scientists and philosophers have proposed a range of answers to this question. This Element introduces this literature and proposes a novel contribution. It defends a realist stance and offers a way (...) of delineating genuine levels of selection by invoking the notion of a functional unit. (shrink)

I argue against theories that attempt to reduce scientific representation to similarity or isomorphism. These reductive theories aim to radically naturalize the notion of representation, since they treat scientist's purposes and intentions as non-essential to representation. I distinguish between the means and the constituents of representation, and I argue that similarity and isomorphism are common but not universal means of representation. I then present four other arguments to show that similarity and isomorphism are not the constituents of scientific representation. I (...) finish by looking at the prospects for weakened versions of these theories, and I argue that only those that abandon the aim to radically naturalize scientific representation are likely to be successful. (shrink)

Schaffner’s model of theory reduction has played an important role in philosophy of science and philosophy of biology. Here, the model is found to be problematic because of an internal tension. Indeed, standard antireductionist external criticisms concerning reduction functions and laws in biology do not provide a full picture of the limits of Schaffner’s model. However, despite the internal tension, his model usefully highlights the importance of regulative ideals associated with the search for derivational, and embedding, deductive relations among mathematical (...) structures in theoretical biology. A reconstructed Schaffnerian model could therefore shed light on mathematical theory development in the biological sciences and on the epistemology of mathematical practices more generally. *Received November 2006; revised March 2009. †To contact the author, please write to: Philosophy Department, University of California, Santa Cruz, 1156 High St., Santa Cruz, CA 95064; e‐mail: [email protected]. (shrink)

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)

Methodological reductionists practice ‘wannabe reductionism’. They claim that one should pursue reductionism, but never propose how. I integrate two strains in prior work to do so. Three kinds of activities are pursued as “reductionist”. “Successional reduction” and inter-level mechanistic explanation are legitimate and powerful strategies. Eliminativism is generally ill-conceived. Specific problem-solving heuristics for constructing inter-level mechanistic explanations show why and when they can provide powerful and fruitful tools and insights, but sometimes lead to erroneous results. I show how traditional metaphysical (...) approaches fail to engage how science is done. The methods used do so, and support a pragmatic and non-eliminativist realism. (shrink)

Richard Levins’ distinction between aggregate, composed and evolved systems acquires new significance as we recognize the importance of mechanistic explanation. Criteria for aggregativity provide limiting cases for absence of organization, so through their failure, can provide rich detectors for organizational properties. I explore the use of failures of aggregativity for the analysis of mechanistic systems in diverse contexts. Aggregativity appears theoretically desireable, but we are easily fooled. It may be exaggerated through approximation, conditions of derivation, and extrapolating from some conditions (...) of decomposition illegtimately to others. Evolved systems particularly may require analyses under alternative complementary decompositions. Exploring these conditions helps us to better understand the strengths and limits of reductionistic methods. (shrink)

David Hull's (1988c) model of science as a selection process suffers from a two-fold inability: (a) to ascertain when a lineage of theories has been established; i.e., when theories are descendants of older theories or are novelties, and what counts as a distinct lineage; and (b) to specify what the scientific analogue is of genotype and phenotype. This paper seeks to clarify these issues and to propose an abstract model of theories analogous to particulate genetic structure, in order to reconstruct (...) relationships of descent and identity. (shrink)

In 1988, David Hull presented an evolutionary account of science. This was a direct analogy to evolutionary accounts of biological adaptation, and part of a generalized view of Darwinian selection accounts that he based upon the Universal Darwinism of Richard Dawkins. Criticisms of this view were made by, among others, Kim Sterelny, which led to it gaining only limited acceptance. Some of these criticisms are, I will argue, no longer valid in the light of developments in the formal modeling of (...) evolution, in particular that of Sergey Gavrilets’ work on adaptive landscapes. If we can usefully recast the Hullian view of science as being driven by selection in terms of Gavrilets’ and Kaufmann’s view of there being “giant components” of high-fitness networks through any realistic adaptive landscape, we may now find it useful to ask what the adaptive pressures on science are, and to extend the metaphor into a full analogy. This is in effect to reconcile the Fisherianism of the Dawkins–Hull approach to selection and replicators, with a Wrightean drift account of social constructionist views of science, preserving, it is to be hoped, the valuable aspects of both. (shrink)

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to (...) grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as “replicators” which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be “vehicles” of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units.The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to “organs” of group-level “organisms.” Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals. (shrink)

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to (...) grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as "replicators" which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be "vehicles" of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units. The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to "organs" of group-level "organisms." Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals. (shrink)

Many standard philosophical accounts of scientific practice fail to distinguish between modeling and other types of theory construction. This failure is unfortunate because there are important contrasts among the goals, procedures, and representations employed by modelers and other kinds of theorists. We can see some of these differences intuitively when we reflect on the methods of theorists such as Vito Volterra and Linus Pauling on the one hand, and Charles Darwin and Dimitri Mendeleev on the other. Much of Volterra's and (...) Pauling's work involved modeling; much of Darwin's and Mendeleev's did not. In order to capture this distinction, I consider two examples of theory construction in detail: Volterra's treatment of post-WWI fishery dynamics and Mendeleev's construction of the periodic system. I argue that modeling can be distinguished from other forms of theorizing by the procedures modelers use to represent and to study real-world phenomena: indirect representation and analysis. This differentiation between modelers and non-modelers is one component of the larger project of understanding the practice of modeling, its distinctive features, and the strategies of abstraction and idealization it employs. (shrink)

Despite their best efforts, scientists may be unable to construct models that simultaneously exemplify every theoretical virtue. One explanation for this is the existence of tradeoffs: relationships of attenuation that constrain the extent to which models can have such desirable qualities. In this paper, we characterize three types of tradeoffs theorists may confront. These characterizations are then used to examine the relationships between parameter precision and two types of generality. We show that several of these relationships exhibit tradeoffs and discuss (...) what consequences those tradeoffs have for theoretical practice. (shrink)

Roald Hoffmann and other theorists claim that we ought to use highly idealized chemical models (“qualitative models”) in order to increase our understanding of chemical phenomena, even though other models are available which make more highly accurate predictions. I assess this norm by examining one of the tradeoffs faced by model builders and model users—the tradeoff between precision and generality. After arguing that this tradeoff obtains in many cases, I discuss how the existence of this tradeoff can help us defend (...) Hoffmann's norm for modelling. (shrink)

The Darwinian concept of natural selection was conceived within a set of Newtonian background assumptions about systems dynamics. Mendelian genetics at first did not sit well with the gradualist assumptions of the Darwinian theory. Eventually, however, Mendelism and Darwinism were fused by reformulating natural selection in statistical terms. This reflected a shift to a more probabilistic set of background assumptions based upon Boltzmannian systems dynamics. Recent developments in molecular genetics and paleontology have put pressure on Darwinism once again. Current work (...) on self-organizing systems may provide a stimulus not only for increased problem solving within the Darwinian tradition, especially with respect to origins of life, developmental genetics, phylogenetic pattern, and energy-flow ecology, but for deeper understanding of the very phenomenon of natural selection itself. Since self-organizational phenomena depend deeply on stochastic processes, self-organizational systems dynamics advance the probability revolution. In our view, natural selection is an emergent phenomenon of physical and chemical selection. These developments suggest that natural selection may be grounded in physical law more deeply than is allowed by advocates of the autonomy of biology, while still making it possible to deny, with autonomists, that evolutionary explanations can be modeled in terms of a deductive relationship between laws and cases. We explore the relationship between, chance, self-organization, and selection as sources of order in biological systems in order to make these points. (shrink)

Is a policy-friendly philosophy of science possible? In order to respond this question, I consider a particular instance of contemporary philosophy of science, the semantic view of scientific theories, by placing it in the broader methodological landscape of the integration of philosophy of science into STS (Science and Technology Studies) as a component of the overall contribution of the latter to science policy. In that context, I defend a multi-disciplinary methodological integration of the special discipline composing STS against a reductionist (...) interdisciplinary unification, arguing that if STS wants to contribute to policy advising by constructing narratives of science practice feasible for science policy both in terms of descriptive completeness and intelligibility, then it must avoid the explanatory reductionism tendencies of special disciplines in interdisciplinary contexts. This would favour, at the same time, a relaxation of esoteric language. On this basis, it seems that the semantic view is one right candidate among other approaches in the philosophy of science for facilitating the integration of the methodologically different contributions to STS toward policy objectives. In fact, besides offering a more realistic and descriptively complete picture of science practice with respect to its predecessor in the philosophy of science, namely the syntactic view, the semantic view is also able to capture some aspects of science practice that elude even sociological approaches to STS, thus inviting different perspective on the same subject matter. (shrink)

In his recent bookThe Advancement of Science,Philip Kitcher endorses eliminative induction, or the view that confirmation of hypotheses proceeds by the elimination of alternatives. My intention here is to critically examine Kitcher's eliminativist view of confirmation, and his rejection of the widely held Bayesian position, according to which an hypothesis H is confirmed by evidence E just in case the probability of H conditional on E is greater than the simple unconditional probability of H [i.e. p(H/E) > p(H)]. Here, I (...) will maintain that while there are aspects of the eliminative approach which Bayesianism must incorporate, as has been argued by John Earman, the eliminativism advocated by Kitcher is unsatisfactory. In addition, I will suggest that Kitcher's reasons for rejecting Bayesian confirmation theory are unconvincing and depend on an overly restrictive version of the position and that indeed Bayesianism has resources needed by an eliminativist account of confirmation. (shrink)

Common monogenic genetic diseases, ones that have unexpectedly high frequencies in certain populations, have attracted a great number of conflicting evolutionary explanations. This paper will attempt to explain the mystery of why two particularly extensively studied common genetic diseases, Tay Sachs disease and cystic fibrosis, remain evolutionary mysteries despite decades of research. I review the most commonly cited evolutionary processes used to explain common genetic diseases: reproductive compensation, random genetic drift (in the context of founder effect), and especially heterozygote advantage. (...) The latter process has drawn a particularly large amount of attention, having so successfully explained the elevated frequency of sickle cell anemia in malaria-endemic areas. However, the empirical evidence for heterozygote advantage in other common genetic diseases is quite weak. I introduce and illustrate the significance of a hierarchy of genetic disease phenomena found within the genetic disease explanations, which include the phenomena: single mutation variants of a common genetic disease, single genetic diseases, and classes of diseases with related phenotypic effects. I demonstrate that some of the confusion over the explanations of common genetic diseases can be traced back to confusions over which phenomena are being explained. I proceed to briefly evaluate the existing evidence for two common human genetic diseases: Tay Sachs disease and cystic fibrosis. The above considerations will ultimately shed light on why these diseases’ evolutionary explanations remain so deeply unresolved after so such a great volume of research. (shrink)

The architects of punctuated equilibrium and species selection as well as more recent workers (Vrba) have narrowed the original formulation of species selection and made it dependent upon so-called emergent characters. One criticism of this narrow version is the dearth of emergent characters with a consequent diminution in the robustness of species selection as an important evolutionary process. We argue that monomorphic species characters may at times be the focus of selection and that under these circumstances selection at the organism (...) level is by-passed due to the absence of critical variance. Selection therefore shifts to the species level where variability reemerges in a clade. The absence of critical variance among organisms prevents effect macroevolution from operating. If species-wide properties are important in macroevolutionary processes, as we contend, systematists should pay more attention to their elucidation. (shrink)

I defend a radical interpretation of biological populations—what I call population pluralism—which holds that there are many ways that a particular grouping of individuals can be related such that the grouping satisfies the conditions necessary for those individuals to evolve together. More constraining accounts of biological populations face empirical counter-examples and conceptual difficulties. One of the most intuitive and frequently employed conditions, causal connectivity—itself beset with numerous difficulties—is best construed by considering the relevant causal relations as ‘thick’ causal concepts. I (...) argue that the fine-grained causal relations that could constitute membership in a biological population are huge in number and many are manifested by degree, and thus we can construe population membership as being defined by massively multidimensional constructs, the differences between which are largely arbitrary. I end by showing that positions in two recent debates in theoretical biology depend on a view of biological populations at odds with the pluralism defended here. (shrink)

Genic selectionism holds that all selection can be understood as operating on particular genes. Critics (and conventional biological wisdom) insist that this misrepresents the actual causal structure of selective phenomena at higher levels of biological organization, but cannot convincingly defend this intuition. I argue that the real failing of genic selectionism is pragmatic – it prevents us from adopting the most efficient corpus of causal laws for predicting and intervening in the course of affairs – and I offer a Pragmatic (...) account of causation itself which ultimately bears out the claim that genic selectionism misrepresents the causal structure of selective contexts. (shrink)

First, a brief history is provided of Popper's views on the status of evolutionary biology as a science. The views of some prominent biologists are then canvassed on the matter of falsifiability and its relation to evolutionary biology. Following that, I argue that Popper's programme of falsifiability does indeed exclude evolutionary biology from within the circumference of genuine science, that Popper's programme is fundamentally incoherent, and that the correction of this incoherence results in a greatly expanded and much more realistic (...) concept of what is empirical, resulting in the inclusion of evolutionary biology. Finally, this expanded concept of empirical is applied to two particular problems in evolutionary biology — viz., the species problem and the debate over the theory of punctuated equilibria — and it is argued that both of them are still mainly metaphysical. (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)

We review Evolution and the Levels of Selection by Samir Okasha. This important book provides a cohesive philosophical framework for understanding levels-of-selections problems in biology. Concerning evolutionary transitions, Okasha proposes that three stages characterize the shift from a lower level of selection to a higher one. We discuss the application of Okasha’s three-stage concept to the evolutionary transition from unicellularity to multicellularity in the volvocine green algae. Okasha’s concepts are a provocative step towards a more general understanding of the major (...) evolutionary transitions; however, the application of certain ideas to the volvocine model system is not straightforward. (shrink)

This paper examines the nature of theory structure in biology and considers the implications of those theoretical structures for theory reduction. An account of biological theories as interlevel prototypes embodying causal sequences, and related to each other by strong analogies, is presented, and examples from the neurosciences are provided to illustrate these middle-range theories. I then go on to discuss several modifications of Nagel''s classical model of theory reduction, and indicate at what stages in the development of reductions these models (...) might best apply. Finally I consider several implications of these analyses of theory structure and reduction for disciplinary integration in biology. (shrink)

Plant and animal species are information-processing entities of such complexity, integration, and adaptive competence that it may be scientifically fruitful to consider them intelligent. The possibility arises from the analogy between learning and evolution, and from recent developments in evolutionary science, psychology and cognitive science. Species are now described as spatiotemporally localized individuals in an expanded hierarchy of biological entities. Intentional and cognitive abilities are now ascribed to animal, human, and artificial intelligence systems that process information adaptively, and that manifest (...) problem-solving abilities. The structural and functional similarities between such systems and species are extensive, although they “are usually obscured by population-genetic metaphors that have nonetheless contributed much to our understanding of evolution.In this target article, I use Sewall Wright's notion of the “adaptive landscape” to compare the performances of evolving species to those of intelligent organisms. With regard to their adaptive achievements and the kinds of processes by which they are achieved, biological species compare very favorably to intelligent animals by virtue of interactions between populations and their environments, between ontogeny and phylogeny, and between natural, interdemic, organic, and species selection. Addressing the question of whether species are intelligent could help to refine our ideas about species, evolution, and intelligence, and could open new lines of empirical and theoretical inquiry in many disciplines. (shrink)

Plant and animal species are information-processing entities of such complexity, integration, and adaptive competence that it may be scientifically fruitful to consider them intelligent. The possibility arises from the analogy between learning and evolution, and from recent developments in evolutionary science, psychology and cognitive science. Species are now described as spatiotemporally localized individuals in an expanded hierarchy of biological entities. Intentional and cognitive abilities are now ascribed to animal, human, and artificial intelligence systems that process information adaptively, and that manifest (...) problem-solving abilities. The structural and functional similarities between such systems and species are extensive, although they “are usually obscured by population-genetic metaphors that have nonetheless contributed much to our understanding of evolution.In this target article, I use Sewall Wright's notion of the “adaptive landscape” to compare the performances of evolving species to those of intelligent organisms. With regard to their adaptive achievements and the kinds of processes by which they are achieved, biological species compare very favorably to intelligent animals by virtue of interactions between populations and their environments, between ontogeny and phylogeny, and between natural, interdemic, organic, and species selection. Addressing the question of whether species are intelligent could help to refine our ideas about species, evolution, and intelligence, and could open new lines of empirical and theoretical inquiry in many disciplines. (shrink)

This survey of major developments in North American philosophy of science begins with the mid-1960s consolidation of the disciplinary synthesis of internalist history and philosophy of science (HPS) as a response to criticisms of logical empiricism. These developments are grouped for discussion under the following headings: historical metamethodologies, scientific realisms, philosophies of the special sciences, revivals of empiricism, cognitivist naturalisms, social epistemologies, feminist theories of science, studies of experiment and the disunity of science, and studies of science as practice and (...) culture. A unifying theme of the survey is the relation between historical metamethodologists and scientific realists, which dominated philosophical work in the late 1970s. I argue that many of the alternative cognitive naturalisms, social epistemologies, and feminist theories that have been proposed can be understood as analogues to the differences between metamethodological theories of scientific rationality and realist accounts of successful reference to real causal processes. Recent work on experiment, scientific practice, and the culture of science may, however, challenge the underlying conception of the field according to which realism and historical rationalism (or their descendants) are the important alternatives available, and thus may take philosophy of science in new directions. (shrink)

This article elaborates on McShea and Brandon’s idea that drift is unlike the rest of the evolutionary factors because it is constitutive rather than imposed on the evolutionary process. I show that the way they spelled out this idea renders it inadequate and is the reason why it received some objections. I propose a different way in which their point could be understood, that rests on two general distinctions. The first is a distinction between the underlying mathematical apparatus used to (...) formulate a theory and a concept proposed by that theory. With the aid of a formal reconstruction of a population genetic model, I show that drift belongs to the first category. That is, that drift is constitutive of population genetics in the same sense that multiplication is constitutive in classical mechanics, or that circle is constitutive in Ptolemaic astronomy. The second distinction is between eliminating a concept from a theory and setting its value to zero. I will show that even though drift can be set to zero just like the rest of the evolutionary factors, eliminating drift is much harder than eliminating those other factors, since it would require changing the entire mathematical apparatus of standard population genetic theory. I conclude by drawing some other implications from the proposed formal reconstruction. (shrink)

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)

Which domains of biology do philosophers of biology primarily study? The fact that philosophy of biology has been dominated by an interest for evolutionary biology is widely admitted, but it has not been strictly demonstrated. Here I analyse the topics of all the papers published in Biology & Philosophy, just as the journal celebrates its thirtieth anniversary. I then compare the distribution of biological topics in Biology & Philosophy with that of the scientific journal Proceedings of the National Academy of (...) Science of the USA, focusing on the recent period 2003–2015. This comparison reveals a significant mismatch between the distributions of these topics. I examine plausible explanations for that mismatch. Finally, I argue that many biological topics underrepresented in philosophy of biology raise important philosophical issues and should therefore play a more central role in future philosophy of biology. (shrink)