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)

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.

Scientific conflicts often stem from differences in the conceptual framework through which scientists view and understand their own field. In this chapter, I analyze the ontological and methodological assumptions of three traditions in evolutionary biology, namely, Ernst Mayr’s population thinking, the gene-centered view of the Modern Syn thesis, and the Extended Evolutionary Synthesis. Each of these frameworks presupposes a different account of "evolutionary causes," and this discrepancy prevents mutual understanding and objective evaluation in the recent contention surrounding the EES. From (...) this perspective, the chapter characterizes the EES research program as an attempt to introduce causal structures beyond genes as additional units of evolution, and compares its research methodology and objectives with those of the traditional MS framework. (shrink)

I argue that Darwinian evolutionary theory has a rhetorical dimension and that rhetorical criticism plays a role in how evolutionary science acquires knowledge. I define what I mean by rhetoric by considering Darwin’s Origin. I use the Modern Evolutionary Synthesis to show how rhetoric conceived as situated and addressed argumentation enters into evolutionary theorizing. Finally, I argue that rhetorical criticism helps judge the success, limits, and failures of these theories.

W&S correctly ask if groups can be like individuals in the harmony and cooperation of their parts, but in their answer, they ignore the importance of the difference between genetically related and unrelated components, and also misconstrue the import of the Hutterites.

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 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..

Feminist philosophers have discussed the prospects for assessing values empirically, particularly given the ongoing threat of sexism and other oppressive values influencing science and society. Some advocates of such tests now champion a “values as evidence” approach, and they criticize Helen Longino’s contextual empiricism for not holding values to the same level of empirical scrutiny as other claims. In this paper, we defend contextual empiricism by arguing that many of these criticisms are based on mischaracterizations of Longino’s position, overstatements of (...) certain claims, and false dichotomies. Her contextual empiricism not only allows for the empirical support and disconfirmation of values, but Longino explicitly discusses when values can be empirically adjudicated and emphasizes the crucial role of the community for standards of evidence. We support contextual empiricism and elaborate a less direct account of “values as heuristics” by reviewing Longino’s theory of evidence and then using a case study from Elisabeth Lloyd on the biology of female orgasm, demonstrating the disconfirmation of androcentric values in evolutionary science. Within Longino’s and Lloyd’s contextual empiricism, values do not get treated as empirical evidence to be directly assessed by individuals, but rather values are heuristic tools to build models whose use can be validated or invalidated by communities based on their empirical fruitfulness in the logic and pragmatics of research questions in specific historical and cultural contexts. (shrink)

Philosophical analysis of ecological theories has lagged behind the study of evolutionary theory. The semantic conception of scientific theories, which has been employed successfully in the analysis of evolutionary theory, is adopted here to analyse ecological theory. Two general problems in ecology are discussed. One arises from the continued use of covering law models in ecology, and the other concerns the applicability of ecological theory in conservation biology. The semantic conception of ecological theories is used to resolve these problems.

We attempt a conclusive resolution of the debate over whether the principle of natural selection (PNS), especially conceived as the `principle' of the `survival of the fittest', is a tautology. This debate has been largely ignored for the past 15 years but not, we think, because it has actually been settled. We begin by describing the tautology objection, and situating the problem in the philosophical and biology literature. We then demonstrate the inadequacy of six prima facie plausible reasons for believing (...) that the tautology debate has been satisfactorily resolved (the PNS is strictly a methodological principle; scientific theories can contain tautologies; the scope of the PNS has been reduced; theories should be understood as models and not exceptionless laws; the widespread acceptance of the propensity interpretation of fitness; and the abandonment of operationalism and verificationism). We proceed to a detailed discussion of Brandon's law (D) describing the PNS, and show that law (D) seriously misrepresents the structure of evolution by natural selection. In the final sections, we provide and defend a novel reinterpretation of the structure of the principle (or, we prefer, model) of evolution by natural selection. (shrink)

Although “theory” has been the prevalent unit of analysis in the meta-study of science throughout most of the twentieth century, the concept remains elusive. I further explore the leitmotiv of several authors in this issue: that we should deal with theorizing (rather than theory) in biology as a cognitive activity that is to be investigated naturalistically. I first contrast how philosophers and biologists have tended to think about theory in the last century or so, and consider recent calls to upgrade (...) the role of theory in the life sciences against the background of the recent “data deluge” in molecular biology, systems biology, etc. I then review thinking about theory in biology in relation to physical theory as a positive or negative exemplar. I conclude by discussing various aspects of a positive program for “naturalizing theorizing.”. (shrink)

The main goal of this book, The Structure and Confirmation of Evolutionary Theory, “is to introduce, develop, and demonstrate the usefulness of a precise analysis of the structure of evolutionary theory”, an analysis that treats evolutionary theory as built up out of complexly interrelated mathematical models. The book has a second goal as well, namely, “to offer … evidence of the appropriateness and utility of the semantic view of scientific theories”. The appropriate audience for the book is not obvious. It (...) is well suited for graduate courses in philosophy of biology, though it demands substantial knowledge of both disciplines. Lloyd has attempted to make it accessible to wider biological and philosophical audiences by minimizing the technical treatment of mathematical models, but I doubt that she has succeeded in making the book accessible to those unfamiliar with the models of population genetics or to biologists previously unversed in the philosophy of science. (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)

Richard Lewontin's (1970) early work on the units of selection initiated the conceptual and theoretical investigations that have led to the hierarchical perspective on selection that has reached near consensus status today. This paper explores other aspects of his work, work on what he termed continuity and quasi-independence, that connect to contemporary explorations of modularity in development and evolution. I characterize such modules and argue that they are the true units of selection in that they are what evolution by natural (...) selection individuates, selects among, and transforms. (shrink)

Genic selectionists (Williams 1966; Dawkins 1976) defend the view that genes are the (unique) units of selection and that all evolutionary events can be adequately represented at the genic level. Pluralistic genic selectionists (Sterelny and Kitcher 1988; Waters 1991; Dawkins 1982) defend the weaker view that in many cases there are multiple equally adequate accounts of evolutionary events, but that always among the set of equally adequate representations will be one at the genic level. We describe a range of cases (...) all involving stable equilibria actively maintained by selection. In these cases genotypic models correctly show that selection is active at the equilibrium point. In contrast, the genic models have selection disappearing at equilibrium. For deterministic models this difference makes no difference. However, once drift is added in, the two sets of models diverge in their predicted evolutionary trajectories. Thus, contrary to received wisdom on this matter, the two sets of models are not empirically equivalent. Moreover, the genic models get the facts wrong. (shrink)

Sober and Elgin defend the claim that there are a priori causal laws in biology. Lange and Rosenberg take issue with this on Humean grounds, among others. I will argue that Sober and Elgin don’t go far enough – there are a priori causal laws in many sciences. Furthermore, I will argue that this thesis is compatible with a Humean metaphysics and an empiricist epistemology.

The formalism used to describe evolutionary change in a multilevel setting can be used equally to re-describe the situation as one where all the selection occurs at the individual level. Thus, whether multilevel or individual-level selection occurs seems to be a matter of convention rather than fact. Yet, group selection is regarded by some as an important concept with factual rather than conventional elements. I flesh out an alternative position that regards groups as a target of selection in a way (...) that is not merely definitional fiat and provide a theoretical basis for this position. (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)

Yes, fitness is the central concept of evolutionary biology, but it is an elusive concept. Almost everyone who looks at it seriously comes out in a different place.

We argue that a fashionable interpretation of the theory of natural selection as a claim exclusively about populations is mistaken. The interpretation rests on adopting an analysis of fitness as a probabilistic propensity which cannot be substantiated, draws parallels with thermodynamics which are without foundations, and fails to do justice to the fundamental distinction between drift and selection. This distinction requires a notion of fitness as a pairwise comparison between individuals taken two at a time, and so vitiates the interpretation (...) of the theory as one about populations exclusively. (shrink)

This article proposes two conditions to assess whether an entity at a level of description is a unit of selection qua interactor. These two conditions make it possible to (1) distinguish biologically relevant entities from arbitrary ones and (2) distinguish units that can _potentially_ enter a selection process from those that have already done so. I show that the classical approaches used in the literature on units and levels of selection do not fare well with respect to either or both (...) of these desiderata. (shrink)

David Lack of Oxford University and V. C. Wynne- Edwards of Aberdeen University were renowned ornithologists with contrasting views of the modern synthesis which deeply influenced their interpretation and explanation of bird behavior. In the 1950's and 60's Lack became the chief advocate of neo-Darwinism with respect to avian ecology, while Wynne- Edwards developed his theory of group selection. Lack 's position was consistent with the developing focus on individual level adaptation, which was a core concept of the modern synthesis. (...) Alternatively, Wynne- Edwards viewed the emphasis on populations as the most important development provided by the modern synthesis. In this paper, I present the development of these two positions and trace their roots in the literature of the synthesis. Through an analysis of Lack 's 1966 critique of Wynne- Edwards I conclude that Wynne- Edwards was, in many ways, justified in his pursuit of group level explanations. (shrink)

We offer to extend structural realism into the field of mind and brain studies. The naturalised metaphysics of structural realism has been defined in terms of unification of sciences. The unification program has been carried out nicely in fields of Quantum Mechanics and General Relativity. But for the structural realist metaphysics to receive a recommendation, the unification program needs to be extended to the fields of especial sciences. Our aim in the paper is twofold. On the one hand, we present (...) a structural realist theory of the brain that is well-supported by the best theories of computational neurosciences. On the other hand, and at the same time, we provide further reason for being optimistic about the structural realism metaphysical claim which is based on the idea of unification in sciences. (shrink)

Philosophy of science is positioned to make distinctive contributions to cognitive science by providing perspective on its conceptual foundations and by advancing normative recommendations. The philosophy of science I embrace is naturalistic in that it is grounded in the study of actual science. Focusing on explanation, I describe the recent development of a mechanistic philosophy of science from which I draw three normative consequences for cognitive science. First, insofar as cognitive mechanisms are information-processing mechanisms, cognitive science needs an account of (...) how the representations invoked in cognitive mechanisms carry information about contents, and I suggest that control theory offers the needed perspective on the relation of representations to contents. Second, I argue that cognitive science requires, but is still in search of, a catalog of cognitive operations that researchers can draw upon in explaining cognitive mechanisms. Last, I provide a new perspective on the relation of cognitive science to brain sciences, one which embraces both reductive research on neural components that figure in cognitive mechanisms and a concern with recomposing higher-level mechanisms from their components and situating them in their environments. (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)

Two strategies for using a model as “null” are distinguished. Null modeling evaluates whether a process is causally responsible for a pattern by testing it against a null model. Baseline modeling measures the relative significance of various processes responsible for a pattern by detecting deviations from a baseline model. When these strategies are conflated, models are illegitimately privileged as accepted until rejected. I illustrate this using the neutral theory of ecology and draw general lessons from this case. First, scientists cannot (...) draw certain conclusions using null modeling. Second, these conclusions follow using baseline modeling, but doing so requires more evidence. (shrink)

This paper examines the empirical aspect of the debates over group selection. I argue that specific narrow readings of the notion of ‘group’ result in a deficient experimentation of the group selection process. Criteria for empirical testability are presented and used to reexamine two well-known experiments of group selection. I argue that the former holds a narrow image of ‘group’ that does not distinguish group selection from selection at other levels; while the latter holds a multifaceted image of ‘group’ that (...) offers a way to achieve substantive progress in the debate. However, most supporters and opponent of group selection maintain a narrow and restricted image of ‘group’, which implicitly undermines informative details regarding population dynamics and population structure. These details may improve our understanding of the evolution of altruism and the evolution of sociality. To conclude, adopting a wide-ranging meaning of ‘group’ may empirically advance the group selection debates. (shrink)

These preprints were automatically compiled into a PDF from the collection of papers deposited in PhilSci-Archive in conjunction with the PSA 2018.

This is the most comprehensive book ever published on philosophical methodology. A team of thirty-eight of the world's leading philosophers present original essays on various aspects of how philosophy should be and is done. The first part is devoted to broad traditions and approaches to philosophical methodology. The entries in the second part address topics in philosophical methodology, such as intuitions, conceptual analysis, and transcendental arguments. The third part of the book is devoted to essays about the interconnections between philosophy (...) and neighbouring fields, including those of mathematics, psychology, literature and film, and neuroscience. (shrink)

This paper considers recent heated debates led by Jerry A. Coyne andMichael J. Wade on issues stemming from the 1929–1962 R.A. Fisher-Sewall Wrightcontroversy in population genetics. William B. Provine once remarked that theFisher-Wright controversy is central, fundamental, and very influential.Indeed,it is also persistent. The argumentative structure of therecent (1997–2000) debates is analyzed with the aim of eliminating a logicalconflict in them, viz., that the two sides in the debates havedifferent aims and that, as such, they are talking past each other. (...) Given aphilosophical analysis of the argumentative structure of the debates,suggestions supportive of Wade's work on the debate are made that areaimed, modestly, at putting the persistent Fisher-Wright controversy on thecourse to resolution. (shrink)

The semantic view of theories is normally considered to be an ac-count of theories congenial to Scientific Realism. Recently, it has been argued that Ontic Structural Realism could be fruitfully applied, in combination with the semantic view, to some of the philosophical issues peculiarly related to bi-ology. Given the central role that models have in the semantic view, and the relevance that mathematics has in the definition of the concept of model, the fo-cus will be on population genetics, which is (...) one of the most mathematized areas in biology. We will analyse some of the difficulties which arise when trying to use Ontic Structural Realism to account for evolutionary biology. (shrink)