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)

Para algunos la selección natural se identifica con las diferencias de éxito de distintos organismos en la reproducción diferencial. Si esto fuese así, el principio de Hardy-Weinberg, por permitir determinar con bastante precisión bajo ciertos supuestos que la frecuencia génica en una población no es la esperada, podría ser visto como una versión cuantitativa de la selección natural cualitativa propuesta por Darwin. Es mi intención mostrar, a través del análisis de explicaciones dadas por Darwin, que la selección natural es más (...) que la mera diferencia en el éxito en la reproducción diferencial e investigar algunas relaciones entre la teoría de la selección natural y la genética de poblaciones. (shrink)

Existe un acuerdo relativo en la necesidad de distinguir dos usos del término «fitness»: el ecológico y el de la genética de poblaciones. Algunos consideran que el segundo ha venido a reemplazar al primero. Otros que el fitness ecológico tiene cierta capacidad explicativa de la que el segundo carece. Estos últimos autores han intentado dar respuesta a cómo es que el fitness ecológico se relaciona con las propiedades particulares de los organismos, siendo estas tan heterogéneas. En este trabajo intentaré dar (...) una respuesta más adecuada a esta cuestión, utilizando el marco conceptual de la metateoría estructuralista. -/- AB STRACT There is relative agreement on the need to distinguish two different uses of the term «fitness»: the ecological fitness and the population genetics fitness. Many consider that the latter has come to replace the former. Others think that the ecological fitness has certain explanatory power that the population genetics fitness lacks of. Among the last ones, many have tried to give response to how ecological fitness relates to organism’s properties, especially because of their being so heterogeneous. In this paper I will try to give a better answer to this matter using the conceptual framework of metatheoretical structuralism. (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)

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)

Philosophy can shed light on mathematical modeling and the juxtaposition of modeling and empirical data. This paper explores three philosophical traditions of the structure of scientific theory—Syntactic, Semantic, and Pragmatic—to show that each illuminates mathematical modeling. The Pragmatic View identifies four critical functions of mathematical modeling: (1) unification of both models and data, (2) model fitting to data, (3) mechanism identification accounting for observation, and (4) prediction of future observations. Such facets are explored using a recent exchange between two groups (...) of mathematical modelers in plant biology. Scientific debate can arise from different modeling philosophies. (shrink)

Critics of contemporary metaphysics argue that it attempts to do the hard work of science from the ease of the armchair. Physics, not metaphysics, tells us about the fundamental facts of the world, and empirical psychology is best placed to reveal the content of our concepts about the world. Exploring and understanding the world through metaphysical reflection is obsolete. In this paper, I will show why this critique of metaphysics fails, arguing that metaphysical methods used to make claims about the (...) world are similar to scientific methods used to make claims about the world, but that the subjects of metaphysics are not the subjects of science. Those who argue that metaphysics uses a problematic methodology to make claims about subjects better covered by natural science get the situation exactly the wrong way around: metaphysics has a distinctive subject matter, not a distinctive methodology. The questions metaphysicians address are different from those of scientists, but the methods employed to develop and select theories are similar. In the first section of the paper, I will describe the sort of subject matter that metaphysics tends to engage with. In the second section of the paper, I will show how metaphysical theories are classes of models and discuss the roles of experience, common sense and thought experiments in the construction and evaluation of such models. Finally, in the last section I will discuss the way these methodological points help us to understand the metaphysical project. Getting the right account of the metaphysical method allows us to better understand the relationship between science and metaphysics, to explain why doing metaphysics successfully involves having a range of different theories, to understand the role of thought experiments involving fictional worlds, and to situate metaphysical realism in a scientifically realist context. (shrink)

The complexity and heterogeneity of causes influencing ecology’s domain challenge its capacity to generate a general theory without exceptions, raising the question of whether ecology is capable, even in principle, of achieving the sort of theoretical success enjoyed by physics. Weber has argued that competition theory built around the Competitive Exclusion Principle (especially Tilman’s resource-competition model) offers an example of ecology identifying a law-like causal regularity. However, I suggest that as Weber presents it, the CEP is not yet a causal (...) regularity. Instead, I argue that the scientific understanding in Tilman’s theory takes a different form. The theory explains through a structure I call “channeling explanation” which does not depend on deduction from general laws, but rather builds on constraints and trade-offs represented in state-space. Recognizing this structure supports the more general point that ecology and other so-called special sciences can reveal novel theoretical approaches to philosophy of science when approached with openness to their uniqueness. (shrink)

This essay analyzes and develops recent views about explanation in biology. Philosophers of biology have parted with the received deductive-nomological model of scientific explanation primarily by attempting to capture actual biological theorizing and practice. This includes an endorsement of different kinds of explanation (e.g., mathematical and causal-mechanistic), a joint study of discovery and explanation, and an abandonment of models of theory reduction in favor of accounts of explanatory reduction. Of particular current interest are philosophical accounts of complex explanations that appeal (...) to different levels of organismal organization and use contributions from different biological disciplines. The essay lays out one model that views explanatory integration across different disciplines as being structured by scientific problems. I emphasize the philosophical need to take the explanatory aims pursued by different groups of scientists into account, as explanatory aims determine whether different explanations are competing or complementary and govern the dynamics of scientific practice, including interdisciplinary research. I distinguish different kinds of pluralism that philosophers have endorsed in the context of explanation in biology, and draw several implications for science education, especially the need to teach science as an interdisciplinary and dynamic practice guided by scientific problems and explanatory aims. (shrink)

this requirement for adaptations. Emergent characters are always potential adaptations. Not all selection processes produce adaptations, however. The key issue, in delineating a selection process, is the relationship between a character and fitness. The emergent character approach is more restrictive than alternative schemas that delineate selection..

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)

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)

Selectionist evolutionary theory has often been faulted for not making novel predictions that are surprising, risky, and correct. I argue that it in fact exhibits the theoretical virtue of predictive capacity in addition to two other virtues: explanatory unification and model fitting. Two case studies show the predictive capacity of selectionist evolutionary theory: parallel evolutionary change in E. coli, and the origin of eukaryotic cells through endosymbiosis.

Today's climate models are supported in a couple of ways that receive little attention from philosophers or climate scientists. In addition to standard 'model fit', wherein a model's simulation is compared to observational data, there is an additional type of confirmation available through the variety of instances of model fit. When a model performs well at fitting first one variable and then another, the probability of the model under some standard confirmation function, say, likelihood, goes up more than under each (...) individual case of fit alone. Thus, two instances of fit of distinct variables of a global climate model using distinct data sets considered collectively will provide stronger evidence for a model than either one of the instances considered individually. This has consequences for model robustness. Sets of models that produce robust results will, if their assumptions vary enough and they each are observationally sound, provide reasons to endorse common structures found in those models. Finally, independent empirical support for aspects and assumptions of the model provides an additional confirmational virtue for climate models, contrary to what is implied by some current philosophical writing on this topic. (shrink)

Multiply realizable properties are those whose realizers are physically diverse. It is often argued that theories which contain them are ipso facto irreducible. These arguments assume that physical explanations are restricted to the most specific descriptions possible of physical entities. This assumption is descriptively false, and philosophically unmotivated. I argue that it is a holdover from the late positivist axiomatic view of theories. A semantic view of theories, by contrast, correctly allows scientific explanations to be couched in the most perspicuous, (...) powerful language available. On a semantic view, traditional notions of multiple realizability are thus very hard to motivate. At best, one must abandon either the idea that multiple realizability is an interesting scientific notion, or else admit that multiply realizable properties do not automatically block scientific reductions. (shrink)

The paper, as Part I of a two-part series, argues for a hybrid formulation of the semantic view of scientific theories. For stage-setting, it first reviews the elements of the model theory in mathematical logic (on whose foundation the semantic view rests), the syntactic and the semantic view, and the different notions of models used in the practice of science. The paper then argues for an integration of the notions into the semantic view, and thereby offers a hybrid semantic view, (...) which at once secures the view's logical foundations and enhances its applicability. The dilemma of either losing touch with the practice of science or yielding up the benefits of the model theory is thus avoided. (shrink)

Recent work on the semantic view of scientific theories is highly critical of the position. This paper identifies two common criticisms of the view, describes two popular alternatives for responding to them, and argues those responses do not suffice. Subsequently, it argues that retuning to Patrick Suppes’ interpretation of the position provides the conceptual resources for rehabilitating the semantic view.

This paper is the second of a two-part series on models and theories, the first of which appeared in International Studies in the Philosophy of Science, Vol. 11, No. 2, 1997. It further explores some of themes of the first paper and examines applications, including: the relations between “similarity” and “isomorphism”, and between “model” and “interpretation”, and the notion of structural explanation.

The dangers of character reification for cladistic inference are explored. The identification and analysis of characters always involves theory-laden abstraction—there is no theory-free “view from nowhere.” Given theory-ladenness, and given a real world with actual objects and processes, how can we separate robustly real biological characters from uncritically reified characters? One way to avoid reification is through the employment of objectivity criteria that give us good methods for identifying robust primary homology statements. I identify six such criteria and explore each (...) with examples. Ultimately, it is important to minimize character reification, because poor character analysis leads to dismal cladograms, even when proper phylogenetic analysis is employed. Given the deep and systemic problems associated with character reification, it is ironic that philosophers have focused almost entirely on phylogenetic analysis and neglected character analysis. (shrink)

The fate of optimality modeling is typically linked to that of adaptationism: the two are thought to stand or fall together (Gould and Lewontin, Proc Relig Soc Lond 205:581–598, 1979; Orzack and Sober, Am Nat 143(3):361–380, 1994). I argue here that this is mistaken. The debate over adaptationism has tended to focus on one particular use of optimality models, which I refer to here as their strong use. The strong use of an optimality model involves the claim that selection is (...) the only important influence on the evolutionary outcome in question and is thus linked to adaptationism. However, biologists seldom intend this strong use of optimality models. One common alternative that I term the weak use simply involves the claim that an optimality model accurately represents the role of selection in bringing about the outcome. This and other weaker uses of optimality models insulate the optimality approach from criticisms of adaptationism, and they account for the prominence of optimality modeling (broadly construed) in population biology. The centrality of these uses of optimality models ensures a continuing role for the optimality approach, regardless of the fate of adaptationism. (shrink)

Non-actual model systems discussed in scientific theories are compared to fictions in literature. This comparison may help with the understanding of similarity relations between models and real-world target systems. The ontological problems surrounding fictions in science may be particularly difficult, however. A comparison is also made to ontological problems that arise in the philosophy of mathematics.

In this paper, a modest version of the Semantic View is motivated as both tenable and potentially fruitful for philosophy of science. An analysis is proposed in which the Semantic View is characterized by three main claims. For each of these claims, a distinction is made between stronger and more modest interpretations. It is argued that the criticisms recently leveled against the Semantic View hold only under the stronger interpretations of these claims. However, if one only commits to the modest (...) interpretation for all the claims, then the view obtained, the Modest Semantic View, is tenable and fruitful for the philosophy of science. (shrink)

Most models of generational succession in sexually reproducing populations necessarily move back and forth between genic and genotypic spaces. We show that transitions between and within these spaces are usually hidden by unstated assumptions about processes in these spaces. We also examine a widely endorsed claim regarding the mathematical equivalence of kin-, group-, individual-, and allelic-selection models made by Lee Dugatkin and Kern Reeve. We show that the claimed mathematical equivalence of the models does not hold. *Received January 2007; revised (...) April 2008. †To contact the authors, please write to: Elisabeth Lloyd, Department of History and Philosophy of Science, 130 Goodbody Hall, Indiana University, Bloomington, IN 47405; e-mail: [email protected]; Richard Lewontin, Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138; Marcus Feldman, Department of Biological Sciences, Stanford University, Stanford, CA 94305; e-mail: [email protected]. (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)

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)

That biology provides explanations is not open to doubt. But how it does so must be a vexed question for those who deny that biology embodies laws or other generalizations with the sort of explanatory force that the philosophy of science recognizes. The most common response to this problem has involved redefining law so that those grammatically general statements which biologists invoke in explanations can be counted as laws. But this terminological innovation cannot identify the source of biology's explanatory power. (...) I argue that because biological science is historical, the problem of biological explanation can be assimilated to the parallel problem in the philosophy of history, and that the problem was solved by Carl Hempel. All we need to do is recognize that the only laws that biology—in all its compartments from the molecular onward—has or needs are the laws of natural selection. (shrink)

David Hull's analysis of conceptual change in science, as presentedin his book, Science as a Process (1988), provides a useful framework for understanding one of the scientific controversies in which he actively and constructively intervened, the units of selectiondebates in evolutionary biology. What follows is a brief overview ofthose debates and some reflections on them.

I argue that four of the fundamental claims of those calling themselves `genic pluralists'Philip Kitcher, Kim Sterelny, and Ken Watersare defective. First, they claim that once genic selectionism is recognized, the units of selection problems will be dissolved. Second, Sterelny and Kitcher claim that there are no targets of selection. Third, Sterelny, Kitcher, and Waters claim that they have a concept of genic causation that allows them to give independent genic causal accounts of all selection processes. I argue that each (...) one of these claims is either false or misleading. Moreover, the challenge that arises from the availability of genic causal accounts, namely, the inability to choose on rational grounds among genic and higher-level accounts, is unsupported. (shrink)

Throughout his career David Hull has sought to bring the philosophy of science into closer contact with science and especially with biological science (Hull 1969, 1997b). This effort has taken many forms. Sometimes it has meant ‘either explaining basic biology to philosophers or explaining basic philosophy to biologists’ (Hull 1996, p. 77). The first of these tasks, simple as it sounds, has been responsible for revolutionary changes. It is well known that traditional philosophy of science, modeled as it was on (...) theoretical physics, proved inadequate when philosophers turned their attention to biological science. Biological examples have driven major revisions of accounts of reduction (Hull 1974; Schaffner 1993, Ch. 9), laws of nature (Beatty et al. 1997), theories (Lloyd 1988) and natural kinds (Wilson 1999, Part III). Nor is explaining basic philosophy to biologists a task to be looked down upon. It is useful, not because philosophy has all the answers, but because scientists must think about how to do science, that is doing philosophy of science and scientists frequently reinvent philosophical views with known flaws. Early in his career Hull found biological systematists in the grip of a crude operationalism about scientific concepts and said so in the pages of Systematic Zoology (Hull 1968). For the next thirty years, as biologists debated the nature of species and the correct principles of classification, Hull added a philosophical note at the same congresses and in the same journals (Hull 1970, 1976, 1980, 1997a, 1999). (shrink)

Paul Thompson, John Beatty, and Elisabeth Lloyd argue that attempts to resolve certain conceptual issues within evolutionary biology have failed because of a general adherence to the received view of scientific theories. They maintain that such issues can be clarified and resolved when one adopts a semantic approach to theories. In this paper, I argue that such conceptual issues are just as problematic on a semantic approach. Such issues arise from the complexity involved in providing formal accounts of theoretical laws (...) and scientific explanations. That complexity is due to empirical and pragmatic considerations, not one's adherence to a particular formal approach to theories. This analysis raises a broader question. How can any formal account properly represent the complex nature of empirical phenomena? (shrink)

In the 1960s molecular population geneticists used Monte Carlo experiments to evaluate particular diffusion equation models. In this paper I examine the nature of this comparative evaluation and argue for three claims: first, Monte Carlo experiments are genuine experiments: second, Monte Carlo experiments can provide an important meansfor evaluating the adequacy of highly idealized theoretical models; and, third, the evaluation of the computational adequacy of a diffusion model with Monte Carlo experiments is significantlydifferent from the evaluation of the emperical adequacy (...) of the same diffusion model. (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)

The conflation of two fundamentally distinct issues has generated serious confusion in the philosophical and biological literature concerning the units of selection. The question of how a unit of selection of defined, theoretically, is rarely distinguished from the question of how to determine the empirical accuracy of claims--either specific or general--concerning which unit(s) is undergoing selection processes. In this paper, I begin by refining a definition of the unit of selection, first presented in the philosophical literature by William Wimsatt, which (...) is grounded in the structure of natural selection models. I then explore the implications of this structural definition for empirical evaluation of claims about units of selection. I consider criticisms of this view presented by Elliott Sober--criticisms taken by some (for example, Mayo and Gilinsky 1987) to provide definitive damage to the structuralist account. I shall show that Sober has misinterpreted the structuralist views; he knocks down a straw man in order to motivate his own causal account. Furthermore, I shall argue, Sober's causal account is dependent on the structuralist account that he rejects. I conclude by indicating how the refined structural definition can clarify which sorts of empirical evidence could be brought to bear on a controversial case involving units of selection. (shrink)

I argue that everyday folk-psychological skill might best be explained in terms of the deployment of something like a model, in a specific sense drawn from recent philosophy of science. Theoretical models in this sense do not make definite commitments about the systems they are used to understand; they are employed with a particular kind of flexibility. This analysis is used to dissolve the eliminativism debate of the 1980s, and to transform a number of other questions about the status and (...) role of folk psychology. (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)

Several prominent debates in biology, such as those surrounding adaptationism, group selection, and punctuated equilibrium, have focused on disagreements about the relative importance of a cause in producing a phenomenon of interest. Some philosophers, such as John Beatty have expressed scepticism about the scientific value of engaging in these controversies, and Karen Kovaka has suggested that their value might be limited. In this paper, I challenge that scepticism by giving a novel analysis of relative significance controversies, showing that there are (...) three forms they can take. I argue that these controversies can have significant epistemic upshots, in that they help scientists form predictions about new instances of the phenomenon of interest. Finally, using two historical examples, I show how engaging in these controversies can improve our understanding of causal relationships. (shrink)

The structuralist approach represents the relation between a model and physical system as a relation between two mathematical structures. However, since a physical system is _prima facie_ _not_ a mathematical structure, the structuralist approach seemingly fails to represent the fact that science is about concrete, physical reality. In this paper, I take up this _problem of lost reality_ and suggest how it may be solved in a purely structuralist fashion. I start by briefly introducing both the structuralist approach and the (...) problem of lost reality and discussing the various (non-structuralist) solutions that have been proposed in the literature. Following this, I decompose the problem into the _ontological mismatch_ and _specification_ problems. In response to the former, I present a _metascientific dissolution argument_, according to which the difference in kind between mathematical structures and physical systems poses no deep obstacle to the structuralist approach, and consider some upshots of this argument for our views on representation. By way of conclusion, I argue that the metascientific dissolution argument paves the way for a solution to the specification problem as well. (shrink)

This special issue for the Journal for General Philosophy of Science is devoted to exploring the impact and many ramifications of current research in evolutionary epistemology. Evolutionary epistemology is an inter- and multidisciplinary area of research that can be divided into two ever-inclusive research avenues. One research avenue expands on the EEM program and investigates the epistemology of evolution. The other research avenue builds on the EET program and researches the evolution of epistemology. Since its conception, EE has developed three (...) schools of thought: adaptationist, non-adaptationist, and applied EE. Although diverse in outlook and theoretical background, these research avenues and schools share the same agenda of understanding how knowledge evolves, and how it relates to the world. In this paper, we first explain wherefrom evolutionary epistemological schools of thought developed, and then we highlight current debates in EE by briefly reviewing the papers that form part of this special issue. (shrink)

⦿ In my dissertation I introduce, motivate and take the first steps in the implementation of, the project of naturalising modal metaphysics: the transformation of the field into a chapter of the philosophy of science rather than speculative, autonomous metaphysics. -/- ⦿ In the introduction, I explain the concept of naturalisation that I apply throughout the dissertation, which I argue to be an improvement on Ladyman and Ross' proposal for naturalised metaphysics. I also object to Williamson's proposal that modal metaphysics (...) --- or some view in the area --- is already a quasi-scientific discipline. -/- ⦿ Recently, some philosophers have argued that the notion of metaphysical modality is as ill defined as to be of little theoretical utility. In the second chapter I intend to contribute to such skepticism. First, I observe that each of the proposed marks of the concept, except for factivity, is highly controversial; thus, its logical structure is deeply obscure. With the failure of the "first principles" approach, I examine the paradigmatic intended applications of the concept, and argue that each makes it a device for a very specific and controversial project: a device, therefore, for which a naturalist will find no use for. I conclude that there is no well-defined or theoretically useful notion of objective necessity other than logical or physical necessity, and I suggest that naturalising modal metaphysics can provide more stable methodological foundations. -/- ⦿ In the third chapter I answer a possible objection against the in-principle viability of the project: that the concept of metaphysical modality cannot be understood through the philosophical analysis of any scientific theory, since metaphysical necessity "transcends'' natural necessity, and science only deals with the latter. I argue that the most important arguments for this transcendence thesis fail or face problems that, as of today, remain unsolved. -/- ⦿ Call the idea that science doesn't need modality, "demodalism''. Demodalism is a first step in a naturalistic argument for modal antirealism. In the fourth chapter I examine six versions of demodalism to explain why a family of formalisms, that I call "spaces of possibility'', are (i) used in a quasi-ubiquitous way in mathematised sciences (I provide examples from theoretical computer science to microeconomics), (ii) scientifically interpreted in modal terms, and (iii) used for at least six important tasks: (1) defining laws and theories; (2) defining important concepts from different sciences (I give several examples); (3) making essential classifications; (4) providing different types of explanations; (5) providing the connection between theory and statistics, and (6) understanding the transition between a theory and its successor (as is the case with quantisation). -/- ⦿ In fifth chapter I propose and defend a naturalised modal ontology. This is a realism about modal structure: my realism about constraints. The modal structure of a system are the relationships between its possible states and between its possible states and those of other systems. It is given by the plurality of restrictions to which said system is subject. A constraint is a factor that explains the impossibility of a class of states; I explain this concept further. First, I defend my point of view by rejecting some of its main rivals: constructive empiricism, Humean conventionalism, and wave function realism, as they fail to make sense of quantum chaos. This is because the field requires the notion of objective modal structure, and the mentioned views have trouble explaining the modal facts of quantum dynamics. Then, I argue that constraint realism supersedes these views in the context of Bohm's standard theory and mechanics, and underpins the study of quantum chaos. Finally, I consider and reject two possible problems for my point of view. -/- ⦿ A central concern of modal metaphysicians has been to understand the logical system that best characterises necessity. In the sixth chapter I intend to recover the logical project applied to my naturalistic modal metaphysics. Scientists and philosophers of science accept different degrees of physical necessity, ranging from purely mathematically necessary facts that restrict physical behaviour, to kinetic principles, to particular dynamical constraints. I argue that this motivates a multimodal approach to modal logic, and that the time dependence of dynamics motivates a logic of historical necessity. I propose multimodal propositional (classical) logics for Bohmian mechanics and the Everettian theory of many divergent worlds, and I close with a criticism of Williamson's approach to the logic of state spaces of dynamic systems. (shrink)

Three metascientific concepts that have been object of philosophical analysis are the concepts oflaw, model and theory. The aim ofthis article is to present the explication of these concepts, and of their relationships, made within the framework of Sneedean or Metatheoretical Structuralism (Balzer et al. 1987), and of their application to a case from the realm of biology: Population Dynamics. The analysis carried out will make it possible to support, contrary to what some philosophers of science in general and of (...) biology in particular hold, the following claims: a) there are "laws" in biological sciences, b) many of the heterogeneous and different "models" of biology can be accommodated under some "theory", and c) this is exactly what confers great unifying power to biological theories. (shrink)

Two metascientific concepts that have been ― and still are ― object of philosophical analysis are the concepts of model and theory. But while the concept of scientific theory was one of the concepts to which philosophers of science devoted most attention during the 20th century, it is only in recent decades that the concept of scientific model has come to occupy a central position in philosophical reflection. However, it has done so in such a way that, at present, as (...) Jim Bogen states in the back cover of the book Scientific Models in the Philosophy of Science, by Daniela Bailer-Jones, “[t]he standard philosophical literature on the role of models in scientific reasoning is voluminous, disorganized, and confusing”. In spite of this, one of the axes that would allow us to organize at least part of this literature, and with which Bailer-Jones’ book closes, is that which is identified as one of the “contemporary philosophical issues: how theories and models relate to each other” (Bailer-Jones 2009, p. 208).That is why, in this introduction to the special issues of Metatheoria devoted to the topic of “Models and Theories in Biology”, we will present the main advances that have been made in the philosophical analysis of the concepts of model and theory in general and in biology in particular, and we will also do the same with the answers that have been given to the problem of "how theories and models relate to each other”. (shrink)

Neutral Theory is controversial in ecology. Ecologists and philosophers have diagnosed the source of the controversy as: its false assumption that individuals in different species within the same trophic level are ecologically equivalent, its conflict with Competition Theory and the adaptation of species, its role as a null hypothesis, and as a Lakatosian research programme. In this paper, I show why we should instead understand the conflict at the level of research programs which involve more than theory. The Neutralist and (...) Competitionist research programs borrow and construct theories, models, and experiments for various aims and given their home ecological systems. I present a holistic and pragmatic view of the controversy that foregrounds the interrelation between many kinds of practices and decisions in ecological research. (shrink)

The units or levels of selection debate concerns the question of what kind of biological systems are stable enough that part of their evolution is a result of the process of natural selection acting at their level. Traditionally, the debate has concerned at least two different, though related, questions: the question of the level at which interaction with the environment occurs, and the question of the level at which reproduction occurs. In recent years, biologists and philosophers have discussed a new (...) aspect of this debate, namely the possibility that certain multi-species consortia formed by a host and its microbiome may act as a unit of selection. This thesis, however, has not been without criticism, as it is doubtful that such consortia could meet the conditions required to achieve the degree of stability that would allow them to experience natural selection. The purpose of this paper is to systematically examine such criticisms and to defend the thesis that the holobiont/hologenome can act as a genuine level of selection both in the form of an interactor and in the form of a reproducer. To do so, it will be argued that the microbiome should be characterized in functional rather than taxonomic terms. (shrink)

The hologenome concept of evolution is a hypothesis about the evolution of animals and plants. It asserts that the evolution of animals and plants was partially triggered by their interactions with their symbiotic microbiomes. In that vein, the hologenome concept posits that the holobiont (animal host + symbionts of the microbiome) is a unit of selection. -/- The hologenome concept has been severely criticized on the basis that selection on holobionts would only be possible if there were a tight transgenerational (...) host-genotype-to-symbiont-genotype connection. As our current evidence suggests that this is not the case for most of the symbiont species that compose the microbiome of animals and plants, the opportunity for holobiont selection is very low in relation to the opportunity for selection on each of the species that compose the host microbiome. Therefore, holobiont selection will always be disrupted ‘from below’, by selection on each of the species that compose the microbiome. -/- This thesis constitutes a conceptual effort to defend philosophically the hologenome concept. I argue that the criticism according to which holobiont selection requires tight transgenerational host-genotype-to-symbiont-genotype connection is grounded on a metaphysical view of the world according to which the biological hierarchy needs to be nested, such that each new level of selection includes every entity from below. Applied to hologenomes, it entails that the hologenome is a collection of genomes, and selection of hologenomes is assumed to entail cospeciation of the host with the species that constitute its microbiome. -/- Against that interpretation, I propose the ‘stability of traits’ account, according to which hologenome evolution is the result of the action of natural selection in a non-nested hierarchical world. In that vein, hologenome evolution does not entail cospeciation, and thus it does not require tight transgenerational host-genotype-to-symbiont-genotype connection. By embracing a multilevel selection perspective, I argue that hologenome evolution results from the simultaneous action of natural selection on each of the lineages that compose the microbiome, and on the assemblage composed by the host genome plus the functional traits of its microbiome. Hologenome selection occurs when the evolution of the traits of the microbiome result from their effects on the fitness of the host, and it can take the form of multilevel selection 1, or multilevel selection 2. In both cases, hologenome selection entails the evolution of microbiome traits, as well as evolution of the host genome, rather than cospeciation of lineages. (shrink)

We address the controversy in the literature concerning the definition of holobionts and the apparent constraints on their evolution using concepts from community population genetics. The genetics of holobionts, consisting of a host and diverse microbial symbionts, has been neglected in many discussions of the topic, and, where it has been discussed, a gene-centric, species-centric view, based in genomic conflict, has been predominant. Because coevolution takes place between traits or genes in two or more species and not, strictly speaking, between (...) species, it may affect some traits but not others in either host or symbiont. Moreover, when interacting species pairs are embedded in a larger community, indirect ecological effects can alter the expected pairwise dynamics. Mode of symbiont transmission and the degree of host inbreeding both affect the extent of microbial mixing across host lineages and thereby the degree to which selection on one trait of either partner affects other aspects of a holobiont phenotype. We discuss several potential defining criteria for holobionts using community genetics and population genetics models, suggesting their application and limitations. Using community genetics models, we show how conflict between genomes can be self-limiting, while cooperation and mutualism tend to be self-accelerating. It is likely that this bias in the evolutionary dynamics of interaction between hosts and symbionts is an important feature of holobionts. This bias in the evolutionary dynamic could contribute to explaining the absence of cheaters from natural mutualisms, although cheaters are predicted by gene-centered conflict theory to cause the evolutionary instability of mutualisms. Additionally, it may help explain the more frequent origin of mutualisms from parasitic than from free-living systems, an evolutionary trajectory opposite to that predicted by genome conflict theory. (shrink)

This PhD dissertation examines the conceptual and theoretical foundations of the most general and most widely used framework for understanding social evolution, W. D. Hamilton's theory of kin selection. While the core idea is intuitive enough (when organisms share genes, they sometimes have an evolutionary incentive to help one another), its apparent simplicity masks a host of conceptual subtleties, and the theory has proved a perennial source of controversy in evolutionary biology. To move towards a resolution of these controversies, we (...) need a careful and rigorous analysis of the philosophical foundations of the theory. My aim in this work is to provide such an analysis. I begin with an examination of the concepts behavioural ecologists employ to describe and classify types of social behaviour. I stress the need to distinguish concepts that are often conflated: for example, we need to distinguish simple cooperation from collaboration in collective tasks, behaviours from strategies, and control from manipulation and coercion. I proceed from here to the formal representation of kin selection via George R. Price’s covariance selection mathematics. I address a number of interpretative issues the Price formalism raises, including the vexed question of whether kin selection theory is ‘formally equivalent’ to multi-level selection theory. In the second half of the dissertation, I assess the uses and limits of Hamilton’s rule for the evolution of social behaviour; I provide a precise statement of the conditions under which the rival neighbour-modulated fitness and inclusive fitness approaches in contemporary kin selection theory are equivalent (and describe cases in which they are not); and I criticize recent formal attempts to establish the controversial claim that kin selection leads to organisms behaving as if maximizing their inclusive fitness. (shrink)

We assess the arguments for recognising functionally integrated multispecies consortia as genuine biological individuals, including cases of so-called ‘holobionts’. We provide two examples in which the same core biochemical processes that sustain life are distributed across a consortium of individuals of different species. Although the same chemistry features in both examples, proponents of the holobiont as unit of evolution would recognize one of the two cases as a multispecies individual whilst they would consider the other as a compelling case of (...) ecological dependence between separate individuals. Some widely used arguments in support of the ‘holobiont’ concept apply equally to both cases, suggesting that those arguments have misidentified what is at stake when seeking to identify a new level of biological individuality. One important aspect of biological individuality is evolutionary individuality. In line with other work on the evolution of individuality, we show that our cases can be distinguished by focusing on the fitness alignment between the partners of the consortia. We conclude that much of the evidence currently presented for the ubiquity and importance of multi-species individuals is simply not to the point, at least unless the issue of biological individuality is firmly divorced from the question of evolutionary individuality. (shrink)