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  1. The Proximate-Ultimate Distinction and the Active Role of the Organism in Evolution.Bendik Hellem Aaby & Grant Ramsey - 2022 - Biology and Philosophy 37 (4):1-20.
    The validity and utility of the proximate-ultimate distinction in biology have recently been under debate. Opponents of the distinction argue that it rules out individual-level organismic processes from evolutionary explanations, thereby leading to an unfounded separation between organismic causation and evolutionary causation. Proponents of the proximate-ultimate distinction, on the other hand, argue that it serves an important epistemological role in forming different kinds of explanation-seeking questions in biology. In this paper we offer an interpretation the proximate-ultimate distinction not only as (...)
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  • Evolution is About Populations, But Its Causes are About Individuals.Pierrick Bourrat - 2019 - Biological Theory 14 (4):254-266.
    There is a tension between, on the one hand, the view that natural selection refers to individual-level causes, and on the other hand, the view that it refers to a population-level cause. In this article, I make the case for the individual-level cause view. I respond to recent claims made by McLoone that the individual-level cause view is inconsistent. I show that if one were to follow his arguments, any causal claim in any context would have to be regarded as (...)
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  • W.F.R. Weldon changes his mind.Charles H. Pence - 2021 - European Journal for Philosophy of Science 11 (3):1-20.
    A recent debate over the causal foundations of evolutionary theory pits those who believe that natural selection causally explains long-term, adaptive population change against those who do not. In this paper, I argue that this debate – far from being an invention of several articles in 2002 – dates from our very first engagements with evolution as a quantified, statistical science. Further, when we analyze that history, we see that a pivotal figure in the early use of statistical methodology in (...)
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  • Is Organismic Fitness at the Basis of Evolutionary Theory?Charles H. Pence & Grant Ramsey - 2015 - Philosophy of Science 82 (5):1081-1091.
    Fitness is a central theoretical concept in evolutionary theory. Despite its importance, much debate has occurred over how to conceptualize and formalize fitness. One point of debate concerns the roles of organismic and trait fitness. In a recent addition to this debate, Elliott Sober argues that trait fitness is the central fitness concept, and that organismic fitness is of little value. In this paper, by contrast, we argue that it is organismic fitness that lies at the bases of both the (...)
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  • Darwin’s empirical claim and the janiform character of fitness proxies.Ulrich Krohs - 2022 - Biology and Philosophy 37 (2):1-23.
    Darwin’s claim about natural selection is reconstructed as an empirical claim about a causal connection leading from the match of the physiology of an individual and its environment to leaving surviving progeny. Variations in this match, Darwin claims, cause differences in the survival of the progeny. Modern concepts of fitness focus the survival side of this chain. Therefore, the assumption that evolutionary theory wants to explain reproductive success in terms of a modern concept of fitness has given rise to the (...)
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  • The Early History of Chance in Evolution.Charles H. Pence - 2015 - Studies in History and Philosophy of Science Part A 50:48-58.
    Work throughout the history and philosophy of biology frequently employs ‘chance’, ‘unpredictability’, ‘probability’, and many similar terms. One common way of understanding how these concepts were introduced in evolution focuses on two central issues: the first use of statistical methods in evolution (Galton), and the first use of the concept of “objective chance” in evolution (Wright). I argue that while this approach has merit, it fails to fully capture interesting philosophical reflections on the role of chance expounded by two of (...)
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  • On Mechanistic Reasoning in Unexpected Places: The Case of Population Genetics.Lucas J. Matthews - 2017 - Biology and Philosophy 32 (6):999-1018.
    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 (...)
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  • Selection in a Complex World: Deriving Causality From Stable Equilibrium.Hugh Desmond - 2018 - Erkenntnis 83 (2):265-286.
    It is an ongoing controversy whether natural selection is a cause of population change, or a mere statistical description of how individual births and deaths accumulate. In this paper I restate the problem in terms of the reference class problem, and propose how the structure of stable equilibrium can provide a solution in continuity with biological practice. Insofar natural selection can be understood as a tendency towards equilibrium, key statisticalist criticisms are avoided. Further, in a modification of the Newtonian-force analogy, (...)
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  • What’s Wrong with Evolutionary Biology?John J. Welch - 2017 - Biology and Philosophy 32 (2):263-279.
    There have been periodic claims that evolutionary biology needs urgent reform, and this article tries to account for the volume and persistence of this discontent. It is argued that a few inescapable properties of the field make it prone to criticisms of predictable kinds, whether or not the criticisms have any merit. For example, the variety of living things and the complexity of evolution make it easy to generate data that seem revolutionary, and lead to disappointment with existing explanatory frameworks. (...)
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  • The arithmetic mean of what? A Cautionary Tale about the Use of the Geometric Mean as a Measure of Fitness.Peter Takacs & Pierrick Bourrat - 2022 - Biology and Philosophy 37 (2):1-22.
    Showing that the arithmetic mean number of offspring for a trait type often fails to be a predictive measure of fitness was a welcome correction to the philosophical literature on fitness. While the higher mathematical moments of a probability-weighted offspring distribution can influence fitness measurement in distinct ways, the geometric mean number of offspring is commonly singled out as the most appropriate measure. For it is well-suited to a compounding process and is sensitive to variance in offspring number. The geometric (...)
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  • The ‘niche’ in niche-based theorizing: much ado about nothing.Samantha Wakil & James Justus - 2022 - Biology and Philosophy 37 (2):1-21.
    The niche is allegedly the conceptual bedrock underpinning the most prominent, and some would say most important, theorizing in ecology. We argue this point of view is more aspirational than veridical. Rather than critically dissect existing definitions of the concept, the supposedly significant work it is thought to have done in ecology is our evaluative target. There is no denying the impressive mathematical sophistication and theoretical ingenuity of the ecological modeling that invokes ‘niche’ terminology. But despite the pervasive labeling, we (...)
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  • A Critical Review of the Statisticalist Debate.Jun Otsuka - 2016 - Biology and Philosophy 31 (4):459-482.
    Over the past decade philosophers of biology have discussed whether evolutionary theory is a causal theory or a phenomenological study of evolution based solely on the statistical features of a population. This article reviews this controversy from three aspects, respectively concerning the assumptions, applications, and explanations of evolutionary theory, with a view to arriving at a definite conclusion in each contention. In so doing I also argue that an implicit methodological assumption shared by both sides of the debate, namely the (...)
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  • Organism-Environment Interactions in Evolutionary Theory.Bendik Hellem Aaby - 2021 - Dissertation, KU Leuven
    This dissertation concerns the active role of the organism in evolutionary theory. In particular, it concerns how our conception of the relationship between organism and environment, and the nature of natural selection, influences the causal and explanatory role of organismic activity and behavior in evolutionary explanations. The overarching aim is to argue that the behaviors and activities of organisms can serve both as the explananda (that which is explained) and the explanantia (that which explains) in evolutionary explanations. I attempt to (...)
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  • Darwinian Rational Expectations.Kobi Finestone - 2022 - Journal of Economic Methodology 29 (2):113-123.
    The rational expectations hypothesis holds that agents should be modeled as not making systematic forecasting errors and has become a central model-building principle of modern economics. The hypot...
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  • Probability and Manipulation: Evolution and Simulation in Applied Population Genetics.Marshall Abrams - 2015 - Erkenntnis 80 (S3):519-549.
    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 (...)
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  • Darwinian Rational Expectations.Kobi Finestone - forthcoming - Tandf: Journal of Economic Methodology:1-11.
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  • The Return of the Organism as a Fundamental Explanatory Concept in Biology.Daniel J. Nicholson - 2014 - Philosophy Compass 9 (5):347-359.
    Although it may seem like a truism to assert that biology is the science that studies organisms, during the second half of the twentieth century the organism category disappeared from biological theory. Over the past decade, however, biology has begun to witness the return of the organism as a fundamental explanatory concept. There are three major causes: (a) the realization that the Modern Synthesis does not provide a fully satisfactory understanding of evolution; (b) the growing awareness of the limits of (...)
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  • The Complex Nexus of Evolutionary Fitness.Mauricio Suárez - 2022 - European Journal for Philosophy of Science 12 (1):1-26.
    The propensity nature of evolutionary fitness has long been appreciated and is nowadays amply discussed. The discussion has, however, on occasion followed long standing conflations in the philosophy of probability literature between propensities, probabilities, and frequencies. In this paper, I apply a more recent conception of propensities in modelling practice to some of the key issues, regarding the mathematical representation of fitness and how it may be regarded as explanatory. The ensuing complex nexus of fitness emphasises the distinction between biological (...)
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  • The Idea of Mismatch in Evolutionary Medicine.Pierrick Bourrat & Paul Edmund Griffiths - forthcoming - British Journal for the Philosophy of Science.
    Mismatch is a prominent concept in evolutionary medicine and a number of philosophers have published analyses of this concept. The word ‘mismatch’ has been used in a diversity of ways across a range of sciences, leading these authors to regard it as a vague concept in need of philosophical clarification. Here, in contrast, we concentrate on the use of mismatch in modelling and experimentation in evolutionary medicine. This reveals a rigorous theory of mismatch within which the term ‘mismatch’ is indeed (...)
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  • Fitness: Philosophical Problems.Grant Ramsey & Charles Pence - 2013 - eLS.
    Fitness plays many roles throughout evolutionary theory, from a measure of populations in the wild to a central element in abstract theoretical presentations of natural selection. It has thus been the subject of an extensive philosophical literature, which has primarily centered on the way to understand the relationship between fitness values and reproductive outcomes. If fitness is a probabilistic or statistical quantity, how is it to be defined in general theoretical contexts? How can it be measured? Can a single conceptual (...)
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  • Levels of Selection Are Artefacts of Different Fitness Temporal Measures.Pierrick Bourrat - 2015 - Ratio 28 (1):40-50.
    In this paper I argue against the claim, recently put forward by some philosophers of biology and evolutionary biologists, that there can be two or more ontologically distinct levels of selection. I show by comparing the fitness of individuals with that of collectives of individuals in the same environment and over the same period of time – as required to decide if one or more levels of selection is acting in a population – that the selection of collectives is a (...)
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  • Taming Fitness: Organism‐Environment Interdependencies Preclude Long‐Term Fitness Forecasting.Guilhem Doulcier, Peter Takacs & Pierrick Bourrat - 2021 - Bioessays 43 (1):2000157.
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  • Framing the Epistemic Schism of Statistical Mechanics.Javier Anta - 2021 - Proceedings of the X Conference of the Spanish Society of Logic, Methodology and Philosophy of Science.
    In this talk I present the main results from Anta (2021), namely, that the theoretical division between Boltzmannian and Gibbsian statistical mechanics should be understood as a separation in the epistemic capabilities of this physical discipline. In particular, while from the Boltzmannian framework one can generate powerful explanations of thermal processes by appealing to their microdynamics, from the Gibbsian framework one can predict observable values in a computationally effective way. Finally, I argue that this statistical mechanical schism contradicts the Hempelian (...)
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  • Explaining Drift From a Deterministic Setting.Pierrick Bourrat - 2017 - Biological Theory 12 (1):27-38.
    Drift is often characterized in statistical terms. Yet such a purely statistical characterization is ambiguous for it can accept multiple physical interpretations. Because of this ambiguity it is important to distinguish what sorts of processes can lead to this statistical phenomenon. After presenting a physical interpretation of drift originating from the most popular interpretation of fitness, namely the propensity interpretation, I propose a different one starting from an analysis of the concept of drift made by Godfrey-Smith. Further on, I show (...)
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  • Fitness: static or dynamic?Peter Takacs & Pierrick Bourrat - 2021 - European Journal for Philosophy of Science 11 (4):1-20.
    The most consistent definition of fitness makes it a static property of organisms. However, this is not how fitness is used in many evolutionary models. In those models, fitness is permitted to vary with an organism’s circumstances. According to this second conception, fitness is dynamic. There is consequently tension between these two conceptions of fitness. One recently proposed solution suggests resorting to conditional properties. We argue, however, that this solution is unsatisfactory. Using a very simple model, we show that it (...)
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  • Naturalización de la Metafísica Modal.Carlos Romero - 2021 - Dissertation, National Autonomous University of Mexico
    ⦿ In my dissertation I introduce, motivate and take the first steps in the realization 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 --- (...)
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  • Roles of Mitonuclear Ecology and Sex in Conceptualizing Evolutionary Fitness.Elay Shech & Kyle B. Heine - 2021 - Biology and Philosophy 36 (3):1-20.
    We look to mitonuclear ecology and the phenomenon of Mother’s Curse to argue that the sex of parents and offspring among populations of eukaryotic organisms, as well as the mitochondrial genome, ought to be taken into account in the conceptualization of evolutionary fitness. Subsequently, we show how characterizations of fitness considered by philosophers that do not take sex and the mitochondrial genome into account may suffer. Last, we reflect on the debate regarding the fundamentality of trait versus organism fitness and (...)
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  • Is Cultural Fitness Hopelessly Confused?Grant Ramsey & Andreas De Block - 2017 - British Journal for the Philosophy of Science 68 (2).
    Fitness is a central concept in evolutionary theory. Just as it is central to biological evolution, so, it seems, it should be central to cultural evolutionary theory. But importing the biological fitness concept to CET is no straightforward task—there are many features unique to cultural evolution that make this difficult. This has led some theorists to argue that there are fundamental problems with cultural fitness that render it hopelessly confused. In this essay, we defend the coherency of cultural fitness against (...)
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  • Is Genetic Drift a Force?Charles H. Pence - manuscript
    One hotly debated philosophical question in the analysis of evolutionary theory concerns whether or not evolution and the various factors which constitute it may profitably be considered as analogous to “forces” in the traditional, Newtonian sense. Several compelling arguments assert that the force picture is incoherent, due to the peculiar nature of genetic drift. I consider two of those arguments here – that drift lacks a predictable direction, and that drift is constitutive of evolutionary systems – and show that they (...)
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  • Trait Fitness is Not a Propensity, but Fitness Variation Is.Elliott Sober - 2013 - Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences 44 (3):336-341.
    The propensity interpretation of fitness draws on the propensity interpretation of probability, but advocates of the former have not attended sufficiently to problems with the latter. The causal power of C to bring about E is not well-represented by the conditional probability Pr. Since the viability fitness of trait T is the conditional probability Pr, the viability fitness of the trait does not represent the degree to which having the trait causally promotes surviving. The same point holds for fertility fitness. (...)
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  • Optimality Models and the Propensity Interpretation of Fitness.Ariel Jonathan Roffé & Santiago Ginnobili - 2020 - Acta Biotheoretica 68 (3):367-385.
    The propensity account of fitness intends to solve the classical tautologicity issue by identifying fitness with a disposition, the ability to survive and reproduce. As proponents recognized early on, this account requires operational independence from actual reproductive success to avoid circularity and vacuousness charges. They suggested that operational independence is achieved by measuring fitness values through optimality models. Our goal in this article is to develop this suggestion. We show that one plausible procedure by which these independent operationalizations could be (...)
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  • Probability in Biology: The Case of Fitness.Roberta L. Millstein - 2016 - In A. Hájek & C. R. Hitchcock (eds.), The Oxford Handbook of Probability and Philosophy. Oxford University Press. pp. 601-622.
    I argue that the propensity interpretation of fitness, properly understood, not only solves the explanatory circularity problem and the mismatch problem, but can also withstand the Pandora’s box full of problems that have been thrown at it. Fitness is the propensity (i.e., probabilistic ability, based on heritable physical traits) for organisms or types of organisms to survive and reproduce in particular environments and in particular populations for a specified number of generations; if greater than one generation, “reproduction” includes descendants of (...)
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  • In What Sense Can There Be Evolution by Natural Selection Without Perfect Inheritance?Pierrick Bourrat - 2019 - International Studies in the Philosophy of Science 32 (1):13-31.
    ABSTRACTIn Darwinian Population and Natural Selection, Peter Godfrey-Smith brought the topic of natural selection back to the forefront of philosophy of biology, highlighting different issues surro...
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  • The Causal Structure of Evolutionary Theory.Grant Ramsey - 2016 - Australasian Journal of Philosophy 94 (3):421-434.
    ABSTRACTOne contentious debate in the philosophy of biology is that between the statisticalists and causalists. The former understand core evolutionary concepts like fitness and selection to be mere statistical summaries of underlying causal processes. In this view, evolutionary changes cannot be causally explained by selection or fitness. The causalist side, on the other hand, holds that populations can change in response to selection—one can cite fitness differences or driftability in causal explanations of evolutionary change. But, on the causalist side, it (...)
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  • Musing on Means: Fitness, Expectation, and the Principles of Natural Selection.Bengt Autzen - 2020 - British Journal for the Philosophy of Science 71 (1):373-389.
    How to measure fitness in the theory of natural selection? A fitness measure that has been proposed in both the biological and the philosophical literature is the expected relative reproductive success. The aim of this article is to examine the relationship between expected relative reproductive success and future actual evolutionary success. Doing so will not only clarify the use of expected relative reproductive success as a fitness measure but also shed light on the role of fitness in the theory of (...)
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  • Locating Uncertainty in Stochastic Evolutionary Models: Divergence Time Estimation.Charles H. Pence - 2019 - Biology and Philosophy 34 (2):21.
    Philosophers of biology have worked extensively on how we ought best to interpret the probabilities which arise throughout evolutionary theory. In spite of this substantial work, however, much of the debate has remained persistently intractable. I offer the example of Bayesian models of divergence time estimation as a case study in how we might bring further resources from the biological literature to bear on these debates. These models offer us an example in which a number of different sources of uncertainty (...)
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  • Natural Selection and the Reference Grain Problem.Pierrick Bourrat - 2020 - Studies in History and Philosophy of Science Part A 80:1-8.
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  • Driftability.Grant Ramsey - 2013 - Synthese 190 (17):3909-3928.
    In this paper, I argue (contra some recent philosophical work) that an objective distinction between natural selection and drift can be drawn. I draw this distinction by conceiving of drift, in the most fundamental sense, as an individual-level phenomenon. This goes against some other attempts to distinguish selection from drift, which have argued either that drift is a population-level process or that it is a population-level product. Instead of identifying drift with population-level features, the account introduced here can explain these (...)
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  • Toward a Propensity Interpretation of Stochastic Mechanism for the Life Sciences.Lane DesAutels - 2015 - Synthese 192 (9):2921-2953.
    In what follows, I suggest that it makes good sense to think of the truth of the probabilistic generalizations made in the life sciences as metaphysically grounded in stochastic mechanisms in the world. To further understand these stochastic mechanisms, I take the general characterization of mechanism offered by MDC :1–25, 2000) and explore how it fits with several of the going philosophical accounts of chance: subjectivism, frequentism, Lewisian best-systems, and propensity. I argue that neither subjectivism, frequentism, nor a best-system-style interpretation (...)
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