The causal nature of evolution is one of the central topics in the philosophy of biology. The issue concerns whether equations used in evolutionary genetics point to some causal processes or purely phenomenological patterns. To address this question the present article builds well-defined causal models that underlie standard equations in evolutionary genetics. These models are based on minimal and biologically plausible hypotheses about selection and reproduction, and generate statistics to predict evolutionary changes. The causal reconstruction of the evolutionary principles shows (...) adaptive evolution as a genuine causal process, where fitness and selection are both causes of evolution. 1 Introduction2 The Philosophical Puzzle3 Causal Models4 Causal Foundations of Evolutionary Genetics4.1 Univariate quantitative genetics model4.1.1 The causal graph4.1.2 Structural equations4.1.3 Deriving evolutionary transition functions4.2 One-locus population genetics model5 Evolution as a Causal Process5.1 Selection as a causal process5.2 Causes of evolutionary change6 Conclusion. (shrink)

The causal nature of evolution is one of the central topics in the philosophy of biology. The issue concerns whether equations used in evolutionary genetics point to some causal processes or purely phenomenological patterns. To address this question the present article builds well-defined causal models that underlie standard equations in evolutionary genetics. These models are based on minimal and biologically plausible hypotheses about selection and reproduction, and generate statistics to predict evolutionary changes. The causal reconstruction of the evolutionary principles shows (...) adaptive evolution as a genuine causal process, where fitness and selection are both causes of evolution. 1 Introduction2 The Philosophical Puzzle3 Causal Models4 Causal Foundations of Evolutionary Genetics4.1 Univariate quantitative genetics model4.1.1 The causal graph4.1.2 Structural equations4.1.3 Deriving evolutionary transition functions4.2 One-locus population genetics model5 Evolution as a Causal Process5.1 Selection as a causal process5.2 Causes of evolutionary change6 Conclusion. (shrink)

I argue that a phenotypic trait can be an adaptation to a particular environmental condition, as against others, only if the environmental condition and the phenotype interactively cause fitness. Models of interactive environmental causes of fitness generally require that environments be individuated by explicit representation rather than by measures of environmental quality, although the latter understanding of ‘environment’ is more prominent in the philosophy of biology. Hence, talk of adaptations to some but not other environmental conditions relies on conceptions of (...) ‘environment’ importantly different from that commonly presupposed in philosophy of biology. (shrink)

Darwin's theory of evolution by natural selection provided the first, and only, causal-mechanistic account of the existence of adaptations in nature. As such, it provided the first, and only, scientific alternative to the “argument from design”. That alone would account for its philosophical significance. But the theory also raises other philosophical questions not encountered in the study of the theories of physics. Unfortunately the concept of natural selection is intimately intertwined with the other basic concepts of evolutionary theory—such as the (...) concepts of fitness and adaptation —that are themselves philosophically controversial. Fortunately we can make considerable headway in getting clear on natural selection without solving all of those outstanding problems. (shrink)

The contextual approach is a prominent framework for thinking about group selection. Here, I highlight ambiguity about what the contextual approach is. Then, I discuss problematic entailments the contextual approach has for what processes count as group selection—entailments more troublesome than typically noted. However, Sober and Wilson’s version of the Price approach, which is the main alternative to the contextual approach, is problematic too: it leads to an underappreciated paradox called the vanishing selection problem and thereby generates the wrong qualitative (...) account of whether group selection is occurring in a certain family of cases. In response, I develop an account of group selection that can deal with the counterexamples to both the contextual approach and the Price approach. I then discuss the role that contextual analysis can continue to play in the discussion of individual fitness and metapopulation evolution. (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)

Starting from the early decades of the twentieth century, evolutionary biology began to acquire mathematical overtones. This took place via the development of a set of models in which the Darwinian picture of evolution was shown to be consistent with the laws of heredity discovered by Mendel. The models, which came to be elaborated over the years, define a field of study known as population genetics. Population genetics is generally looked upon as an essential component of modern evolutionary theory. This (...) article deals with a famous dispute between J. B. S. Haldane, one of the founders of population genetics, and Ernst Mayr, a major contributor to the way we understand evolution. The philosophical undercurrents of the dispute remain relevant today. Mayr and Haldane agreed that genetics provided a broad explanatory framework for explaining how evolution took place but differed over the relevance of the mathematical models that sought to underpin that framework. The dispute began with a fundamental issue raised by Mayr in 1959: in terms of understanding evolution, did population genetics contribute anything beyond the obvious? Haldane's response came just before his death in 1964. It contained a spirited defense, not just of population genetics, but also of the motivations that lie behind mathematical modelling in biology. While the difference of opinion persisted and was not glossed over, the two continued to maintain cordial personal relations. (shrink)

I define a concept of causal probability and apply it to questions about the role of probability in evolutionary processes. Causal probability is defined in terms of manipulation of patterns in empirical outcomes by manipulating properties that realize objective probabilities. The concept of causal probability allows us see how probabilities characterized by different interpretations of probability can share a similar causal character, and does so in such way as to allow new inferences about relationships between probabilities realized in different chance (...) setups. I clarify relations between probabilities and properties defined in terms of them, and argue that certain widespread uses of computer simulations in evolutionary biology show that many probabilities relevant to evolutionary outcomes are causal probabilities. This supports the claim that higher-level properties such as biological fitness and processes such as natural selection are causal properties and processes, contrary to what some authors have argued. (shrink)

The idea of organism-environment interaction, at least in its modern form, dates only to the mid-nineteenth century. After sketching the origins of the organism-environment dichotomy in the work of Auguste Comte and Herbert Spencer, I will chart its metaphysical and methodological influence on later scientists and philosophers such as Conwy Lloyd Morgan and John Dewey. In biology and psychology, the environment was seen as a causal agent, highlighting questions of organismic variation and plasticity. In philosophy, organism-environment interaction provided a new (...) foundation for ethics, politics, and scientific inquiry. Thinking about organism-environment interaction became indispensable, for it had restructured our view of the biological and social world. (shrink)

Despite the burgeoning interest in new and more complex accounts of the organism-environment dyad by biologists and philosophers, little attention has been paid in the resulting discussions to the history of these ideas and to their deployment in disciplines outside biology—especially in the social sciences. Even in biology and philosophy, there is a lack of detailed conceptual models of the organism-environment relationship. This volume is designed to fill these lacunae by providing the first multidisciplinary discussion of the topic of organism-environment (...) interaction. It brings together scholars from history, philosophy, psychology, anthropology, medicine, and biology to discuss the common focus of their work: entangled life, or the complex interaction of organisms and environments. (shrink)

A strong case has been made for the role and value of mechanistic reasoning in process-oriented sciences, such as molecular biology and neuroscience. This paper shifts focus to assess the role of mechanistic reasoning in an area where it is neither obvious nor expected: population genetics. Population geneticists abstract away from the causal-mechanical details of individual organisms and, instead, use mathematics to describe population-level, statistical phenomena. This paper, first, develops a framework for the identification of mechanistic reasoning where it is (...) not obvious: mathematical and mechanistic styles of scientific reasoning. Second, it applies this framework to demonstrate that both styles are integrated in modern investigations of evolutionary biology. Characteristic of the former, applied population genetic techniques provide statistical evidence for associations between genotype, phenotype, and fitness. Characteristic of the latter, experimental interventions provide causal-mechanical evidence for associations between the very same relationships, often in the same model organisms. The upshot is a richer perspective of how evolutionary biologists build evidence for hypotheses regarding adaptive evolution and a general framework for assessing the scope of mechanistic reasoning across the sciences. (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)

The Darwinian theory of evolution is itself evolving and this book presents the details of the core of modern Darwinism and its latest developmental directions. The authors present current scientific work addressing theoretical problems and challenges in four sections, beginning with the concepts of evolution theory, its processes of variation, heredity, selection, adaptation and function, and its patterns of character, species, descent and life. The second part of this book scrutinizes Darwinism in the philosophy of science and its usefulness in (...) understanding ecosystems, whilst the third section deals with its application in disciplines beyond the biological sciences, including evolutionary psychology and evolutionary economics, Darwinian morality and phylolinguistics. The final section addresses anti-Darwinism, the creationist view and issues around teaching evolution in secondary schools. The reader learns how current experimental biology is opening important perspectives on the sources of variation, and thus of the very power of natural selection. This work examines numerous examples of the extension of the principle of natural selection and provides the opportunity to critically reflect on a rich theory, on the methodological rigour that presides in its extensions and exportations, and on the necessity to measure its advantages and also its limits. Scholars interested in modern Darwinism and scientific research, its concepts, research programs and controversies will find this book an excellent read, and those considering how Darwinism might evolve, how it can apply to the human sciences and other disciplines beyond its origins will find it particularly valuable. Originally produced in French (Les Mondes Darwiniens), the scope and usefulness of the book have led to the production of this English text, to reach a wider audience. This book is a milestone in the impressive penetration by Francophone scholars into the world of Darwinian science, its historiography and philosophy over the last two decades. Alex Rosenberg, R. Taylor Cole Professor of Philosophy, Duke University Until now this useful and comprehensive handbook has only been available to francophones. Thanks to this invaluable new translation, this collection of insightful and original essays can reach the global audience it deserves. Tim Lewens, University of Cambridge. (shrink)

In this short paper, I argue against what I call the “belonging to” interpretation of group selection in scenarios in which a group’s fitness is defined as the per capita reproductive output of the individuals of the group. According to this interpretation, group selection acts on “belonging to” properties of individuals, i.e. on relational or contextual properties that all the individuals of a group share simply by belonging to that group; thus, if differences in the individuals’ “belonging to” properties cause (...) differences in their fitness, group selection sensu the “belonging to” interpretation is said to be at work. I argue that the main problem with the “belonging to” interpretation is that it confuses evolutionary changes due to differences in environmental quality with evolutionary changes due to selection. In other words, I argue that, in the majority of cases, this interpretation actually takes the “selection” out of the “group selection” notion it aims to interpret: by adopting this perspective, one implicitly commits to explaining the evolutionary change under consideration not by a kind of selection, but by differences in the environmental quality experienced by individual types. (shrink)

La tesis del holismo de la confirmación aparece por primera vez presentada de manera explícita y extensa en los escritos de Duhem. Años después, es defendida por Quine, pero no en base a la discusión de teorías científicas particulares, sino en relación con su posición semántica. Desde entonces las relaciones entre el holismo semántico y el holismo de la contrastación han sido discutidas, aunque generalmente de un modo algo despegado del análisis de teorías específicas y sus relaciones. Pretendo mostrar a (...) partir del análisis de un caso, el de las relaciones entre la genética de poblaciones y la teoría de la selección natural, un sentido en el que el holismo de la confirmación puede ser derivado del holismo semántico. The thesis of the confirmation holism appears for the first time explicitly and extensively in the writings of Duhem. Years later, it is defended by Quine, but not based on the discussion of particular scientific theories, but in relation to its semantic position. Since then, the relations between semantic holism and confirmation holism have been discussed, although generally somewhat detached from the analysis of specific theories and their relations. I intend to show from the analysis of a case, that of the relations between population genetics and the theory of natural selection, a sense in which the holism of confirmation can be derived from semantic holism. (shrink)

Causalism is the thesis that natural selection can cause evolution. A standard argument for causalism involves showing that a hypothetical intervention on some population-level property that is identified with natural selection will result in evolution. In a pair of articles, one of which recently appeared in the pages of this journal, Jun Otsuka has put forward a quite different argument for causalism. Otsuka attempts to show that natural selection can cause evolution by considering a hypothetical intervention on an individual-level property. (...) Specifically, Otsuka identifies natural selection with the causal relationship between a trait and fitness, claims an intervention on the strength of this relationship can cause evolution, then concludes that natural selection can cause evolution. Below I describe why Otsuka’s argument for causalism is unconvincing. Central to my criticism is that Otsuka’s argument works only if one adopts an indefensible account of natural selection, according to which natural selection can occur in the absence of trait or fitness variation. I go on to explain why any attempt to demonstrate the truth of causalism via a hypothetical intervention on an individual-level property would appear to require one to adopt an account of natural selection that is inadequate for the same reason. This in turn means the plausibility of causalism does indeed depend on the plausibility of the claim that population-level properties, which supervene on the properties of the individuals in the population, can be causally efficacious. (shrink)

Es una opinión extendida que la teoría de la selección natural, tal como fue formulada originalmente por Darwin de manera “cualitativa”, ha sido reemplazada por una versión cuantitativa superior, brindada por la genética de poblaciones. En este artículo se discute contra esa tesis, sosteniendo en cambio que ambas teorías son complementarias, no sucesivas. Para ello, se introduce una línea de argumentación novedosa, que toma su inspiración en la crítica al “inductivismo estrecho” de Hempel. Se sostiene que los genetistas de poblaciones (...) serían incapaces de aplicar exitosamente su teoría sin hipótesis “ecológicas” preconcebidas, provenientes de la teoría darwiniana, que permitan particionar a la población en rasgos selectivamente relevantes. Se enfatiza además que la falla en notar este punto se debe a una mala comprensión de la estructura de la teoría darwiniana y de su ley o principio fundamental. A la luz de una mejor reconstrucción de dicho principio, se reexamina en mayor detalle la relación existente entre ambas teorías. (shrink)

Recently, Estes and Arnold claimed to have “solved” the paradox of evolutionary stasis; they claim that stabilizing selection, and only stabilizing selection, can explain the patterns of evolutionary divergence observed over “all timescales.” While Estes and Arnold clearly think that they have identified the processes that produce evolutionary stasis, they have not. Instead, Estes and Arnold identify a particular evolutionary pattern but not the processes that produce that pattern. This mistake is important; the slippage between pattern and process is common (...) in population and quantitative genetics and contributes to a persistent misunderstanding of the nature of explanations in evolutionary biology. †To contact the author, please write to: Philosophy Department, 208 Hovland Hall, Oregon State University, Corvallis, OR 97331‐3902; e‐mail: [email protected]. (shrink)

Brandon claims that to explain adaptation one must specify fitnesses in each selective environment and specify the distribution of individuals across selective environments. Glymour claims, using an example of the adaptive evolution of costly plasticity in a symmetric environment, that there are some predictive or explanatory tasks for which Brandon’s claim is limited. In this paper, I provide necessary conditions for carrying out Brandon’s task, produce a new version of the argument for his claim, and show that Glymour’s reasons for (...) making his claim are problematic. I provide a few interpretations of Glymour’s argument but ultimately raise worries for what I take to be the key premises. (shrink)

This work is a critical consideration of several arguments recently given by Elliott Sober that are aimed at undermining the Laplacean stance on probability in evolutionary theory. The Laplacean contends that the only objective probability an event has is the one assigned to it by a complete description of the relevant microparticles. Sober alleges a formal demonstration that the Laplacean stance on probability in evolutionary theory is inconsistent. But Sober’s argument contains a crucial lacuna, one that likely cannot be repaired (...) to yield the conclusion that Sober draws. He also argues that the Laplacean is committed to inferring semantic facts about probabilities from pragmatic facts about agents’ reasons for using probabilities. But Sober’s arguments against inferring semantic facts from pragmatic constraints would only undermine one basis for Laplaceanism. The Laplacean who formulates her position carefully need not leave herself vulnerable to Sober’s objection to inferring semantics from pragmatics. (shrink)

Glymour (Philos Sci 73:369–389, 2006) claims that classical population genetic models can reliably predict short and medium run population dynamics only given information about future fitnesses those models cannot themselves predict, and that in consequence the causal, ecological models which can predict future fitnesses afford a more foundational description of natural selection than do population genetic models. This paper defends the first claim from objections offered by Gildenhuys (Biol Philos, 2011).

It has become customary in multilevel selection theory to use the same terms to denote both two explanatory goals and two explanatory means. This paper spells out some of the benefits that derive from avoiding this terminological conflation. I argue that keeping explanatory means and goals well apart allows us to see that, contrary to a popular recent idea, Price’s equation and contextual analysis—the statistical methods most extensively used for measuring the effects of certain evolutionary factors on the change in (...) the focal individual trait in multilevel selection scenarios—do not come with built-in notions of group selection and, therefore, the efficacy of these methods at analyzing various kinds of cases does not constitute a basis for deciding how group selection should best be defined. Moreover, contrary to another widely accepted idea, I argue that more than one type of group selection may serve as explanatory means when one’s goal is that of explaining the evolution of individual traits in multilevel selection scenarios and I spell out how this explanatory role should be understood. (shrink)

In a recent article, “Wayward Modeling: Population Genetics and Natural Selection,” Bruce Glymour claims that population genetics is burdened by serious predictive and explanatory inadequacies and that the theory itself is to blame. Because Glymour overlooks a variety of formal modeling techniques in population genetics, his arguments do not quite undermine a major scientific theory. However, his arguments are extremely valuable as they provide definitive proof that those who would deploy classical population genetics over natural systems must do so with (...) careful attention to interactions between individual population members and environmental causes. Glymour’s arguments have deep implications for causation in classical population genetics. (shrink)