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  1. Marshall Abrams (2009). Fitness “Kinematics”: Biological Function, Altruism, and Organism–Environment Development. Biology and Philosophy 24 (4):487-504.
    It’s recently been argued that biological fitness can’t change over the course of an organism’s life as a result of organisms’ behaviors. However, some characterizations of biological function and biological altruism tacitly or explicitly assume that an effect of a trait can change an organism’s fitness. In the first part of the paper, I explain that the core idea of changing fitness can be understood in terms of conditional probabilities defined over sequences of events in an organism’s life. The result (...)
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  2. Marshall Abrams (2009). The Unity of Fitness. Philosophy of Science 76 (5):750-761.
    It has been argued that biological fitness cannot be defined as expected number of offspring in all contexts. Some authors argue that fitness therefore merely satisfies a common schema or that no unified mathematical characterization of fitness is possible. I argue that comparative fitness must be relativized to an evolutionary effect; thus relativized, fitness can be given a unitary mathematical characterization in terms of probabilities of producing offspring and other effects. Such fitnesses will sometimes be defined in terms of probabilities (...)
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  3. Marshall Abrams (2009). What Determines Biological Fitness? The Problem of the Reference Environment. Synthese 166 (1):21 - 40.
    Organisms' environments are thought to play a fundamental role in determining their fitness and hence in natural selection. Existing intuitive conceptions of environment are sufficient for biological practice. I argue, however, that attempts to produce a general characterization of fitness and natural selection are incomplete without the help of general conceptions of what conditions are included in the environment. Thus there is a "problem of the reference environment"—more particularly, problems of specifying principles which pick out those environmental conditions which determine (...)
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  4. Marshall Abrams (2007). Fitness and Propensity's Annulment? Biology and Philosophy 22 (1):115-130.
    Recent debate on the nature of probabilities in evolutionary biology has focused largely on the propensity interpretation of fitness (PIF), which defines fitness in terms of a conception of probability known as “propensity”. However, proponents of this conception of fitness have misconceived the role of probability in the constitution of fitness. First, discussions of probability and fitness have almost always focused on organism effect probability, the probability that an organism and its environment cause effects. I argue that much of the (...)
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  5. Andre Ariew (2009). What Fitness Can't Be. Erkenntnis 71 (3):289 - 301.
    Recently advocates of the propensity interpretation of fitness have turned critics. To accommodate examples from the population genetics literature they conclude that fitness is better defined broadly as a family of propensities rather than the propensity to contribute descendants to some future generation. We argue that the propensity theorists have misunderstood the deeper ramifications of the examples they cite. These examples demonstrate why there are factors outside of propensities that determine fitness. We go on to argue for the more general (...)
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  6. André Ariew & R. C. Lewontin (2004). The Confusions of Fitness. British Journal for the Philosophy of Science 55 (2):347-363.
    The central point of this essay is to demonstrate the incommensurability of ‘Darwinian fitness’ with the numeric values associated with reproductive rates used in population genetics. While sometimes both are called ‘fitness’, they are distinct concepts coming from distinct explanatory schemes. Further, we try to outline a possible answer to the following question: from the natural properties of organisms and a knowledge of their environment, can we construct an algorithm for a particular kind of organismic life-history pattern that itself will (...)
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  7. Jonathan Birch (2014). Gene Mobility and the Concept of Relatedness. Biology and Philosophy 29 (4):445-476.
    Cooperation is rife in the microbial world, yet our best current theories of the evolution of cooperation were developed with multicellular animals in mind. Hamilton’s theory of inclusive fitness is an important case in point: applying the theory in a microbial setting is far from straightforward, as social evolution in microbes has a number of distinctive features that the theory was never intended to capture. In this article, I focus on the conceptual challenges posed by the project of extending Hamilton’s (...)
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  8. Jonathan Birch (2014). Has Grafen Formalized Darwin? Biology and Philosophy 29 (2):175-180.
    One key aim of Grafen’s Formal Darwinism project is to formalize ‘modern biology’s understanding and updating of Darwin’s central argument’. In this commentary, I consider whether Grafen has succeeded in this aim.
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  9. Jonathan Birch (2013). Samir Okasha and Ken Binmore (Eds) Evolution and Rationality: Decisions, Cooperation, and Strategic Behaviour. [REVIEW] British Journal for the Philosophy of Science 64 (3):669-673.
  10. Marion Blute (2001). A Single-Process Learning Theory. Behavioral and Brain Sciences 24 (3):529-531.
    Many analogies exist between the process of evolution by natural selection and of learning by reinforcement and punishment. A full extension of the evolutionary analogy to learning to include analogues of the fitness, genotype, development, environmental influences, and phenotype concepts makes possible a single theory of the learning process able to encompass all of the elementary procedures known to yield learning.
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  11. Frédéric Bouchard (2011). Darwinism Without Populations: A More Inclusive Understanding of the “Survival of the Fittest”. Studies in History and Philosophy of Science Part C 42 (1):106-114.
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  12. Frédéric Bouchard (2008). Causal Processes, Fitness, and the Differential Persistence of Lineages. Philosophy of Science 75 (5):560-570.
    Ecological fitness has been suggested to provide a unifying definition of fitness. However, a metric for this notion of fitness was in most cases unavailable except by proxy with differential reproductive success. In this article, I show how differential persistence of lineages can be used as a way to assess ecological fitness. This view is inspired by a better understanding of the evolution of some clonal plants, colonial organisms, and ecosystems. Differential persistence shows the limitation of an ensemblist noncausal understanding (...)
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  13. Frédéric Bouchard & Alex Rosenberg (2004). Fitness, Probability and the Principles of Natural Selection. British Journal for the Philosophy of Science 55 (4):693-712.
    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 (...)
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  14. Henry Byerly (1986). Fitness as a Function. PSA: Proceedings of the Biennial Meeting of the Philosophy of Science Association 1986:494 - 501.
    Fitness in the sense of actual rate of increase of genotypes, commonly used in population genetics, is contrasted with fitness in the ordinary sense (and Darwin's) of adaptedness of organisms. Fitness as actual reproductive success is interpreted as a function of variables representing intrinsic adaptive capacities and environmental properties. Adaptive capacities causally contribute to fitness as actual reproductive success which in turn, as relative increase of genotypes, determines evolutionary change. The propensity interpretation of fitness is shown not to play a (...)
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  15. Henry C. Byerly & Richard E. Michod (1991). Fitness and Evolutionary Explanation. [REVIEW] Biology and Philosophy 6 (1):45-53.
    Recent philosophical discussions have failed to clarify the roles of the concept fitness in evolutionary theory. Neither the propensity interpretation of fitness nor the construal of fitness as a primitive theoretical term succeed in explicating the empirical content and explanatory power of the theory of natural selection. By appealing to the structure of simple mathematical models of natural selection, we separate out different contrasts which have tended to confuse discussions of fitness: the distinction between what fitness is defined as versus (...)
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  16. Gregory Cooper (1988). Fitness and Explanation. PSA: Proceedings of the Biennial Meeting of the Philosophy of Science Association 1988:207 - 215.
    Although consensus appears to be on the horizon, the foundations of the theory of natural selection remain a matter of controversy. This paper looks at two recent challenges to the emerging "received view" of this theory. It argues that different views of the nature of scientific explanation are playing a pivotal role in the debates. Do explanations in biology fit the covering-law paradigm? What are the explanatory laws of biology like? Until agreement is reached on these fundamental questions, there is (...)
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  17. A. Brito Da Cunha (1991). Commentary on the Paper by HC Byerly and RE Michod,“Fitness and Evolutionary Explanation”. Biology and Philosophy 6 (1):23-27.
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  18. Klaus Dietz (2005). Darwinian Fitness, Evolutionary Entropy and Directionality Theory. Bioessays 27 (11):1097-1101.
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  19. Iris Fry (1996). On the Biological Significance of the Properties of Matter: L. J. Henderson's Theory of the Fitness of the Environment. [REVIEW] Journal of the History of Biology 29 (2):155 - 196.
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  20. Lasse Gerrits & Peter Marks (forthcoming). The Evolution of Wright's (1932) Adaptive Field to Contemporary Interpretations and Uses of Fitness Landscapes in the Social Sciences. Biology and Philosophy:1-21.
    The concepts of adaptation and fitness have such an appeal that they have been used in other scientific domains, including the social sciences. One particular aspect of this theory transfer concerns the so-called fitness landscape models. At first sight, fitness landscapes visualize how an agent, of any kind, relates to its environment, how its position is conditional because of the mutual interaction with other agents, and the potential routes towards improved fitness. The allure of fitness landscapes is first and foremost (...)
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  21. Herbert Gintis (2014). Inclusive Fitness and the Sociobiology of the Genome. Biology and Philosophy 29 (4):477-515.
    Inclusive fitness theory provides conditions for the evolutionary success of a gene. These conditions ensure that the gene is selfish in the sense of Dawkins (The selfish gene, Oxford University Press, Oxford, 1976): genes do not and cannot sacrifice their own fitness on behalf of the reproductive population. Therefore, while natural selection explains the appearance of design in the living world (Dawkins in The blind watchmaker: why the evidence of evolution reveals a universe without design, W. W. Norton, New York, (...)
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  22. Peter Godfrey-Smith (1992). Additivity and the Units of Selection. PSA: Proceedings of the Biennial Meeting of the Philosophy of Science Association 1992:315 - 328.
    "Additive variance in fitness" is an important concept in the formal apparatus of population genetics. Wimsatt and Lloyd have argued that this concept can also be used to decide the "unit of selection" in an evolutionary process. The paper argues that the proposed criteria of Wimsatt and Lloyd are ambiguous, and several interpretations of their views are presented. It is argued that none of these interpretations provide acceptable criteria for deciding units of selection. The reason is that additive variance in (...)
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  23. Paul E. Griffiths (1993). Functional Analysis and Proper Functions. British Journal for the Philosophy of Science 44 (3):409-422.
    The etiological approach to ‘proper functions’ in biology can be strengthened by relating it to Robert Cummins' general treatment of function ascription. The proper functions of a biological trait are the functions it is assigned in a Cummins-style functional explanation of the fitness of ancestors. These functions figure in selective explanations of the trait. It is also argued that some recent etiological theories include inaccurate accounts of selective explanation in biology. Finally, a generalization of the notion of selective explanation allows (...)
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  24. Jonathan Kaplan (2008). The End of the Adaptive Landscape Metaphor? Biology and Philosophy 23 (5):625-638.
    The concepts of adaptive/fitness landscapes and adaptive peaks are a central part of much of contemporary evolutionary biology; the concepts are introduced in introductory texts, developed in more detail in graduate-level treatments, and are used extensively in papers published in the major journals in the field. The appeal of visualizing the process of evolution in terms of the movement of populations on such landscapes is very strong; as one becomes familiar with the metaphor, one often develops the feeling that it (...)
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  25. David Keane (2010). Survival of the Fairest? Evolution and the Geneticization of Rights. Oxford Journal of Legal Studies 30 (3):467-494.
    The process of evolution is largely absent from philosophical legal literature, to the extent that the possibility of a genetic origin of rights has not been explored. This is striking given that human rights theory stems from natural law and natural rights, which seems to imply a potential link with natural selection. Furthermore, the concept of nature has played a significant role in the philosophical foundations of international legal norms of rights and responsibilities. On the surface it may seem desirable (...)
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  26. Sa Kleiner (1991). Fitness and Evolutionary Explanation-Comment. Biology and Philosophy 6 (1):29-32.
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  27. Costas B. Krimbas (2004). On Fitness. Biology and Philosophy 19 (2):185-203.
    The concept of fitness, central to population genetics and to the synthetic theory of evolution, is discussed. After a historical introduction on the origin of this concept, the current meaning of it in population genetics is examined: a cause of the selective process and its quantification. Several difficulties arise for its exact definition. Three adequacy criteria for such a definition are formulated. It is shown that it is impossible to formulate an adequate definition of fitness respecting these criteria. The propensity (...)
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  28. Maria Kronfeldner (2010). Wenn Philosophen Auf Biologen Treffen: Über Die Arbeit Am Begriff Im Dienste der Interdisziplinarität. Briefe Zur Interdisziplinarität 6:7-16.
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  29. Nick Lane (2011). Mitonuclear Match: Optimizing Fitness and Fertility Over Generations Drives Ageing Within Generations. Bioessays 33 (11):860-869.
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  30. Laurent Lehmann & François Rousset (2014). Fitness, Inclusive Fitness, and Optimization. Biology and Philosophy 29 (2):181-195.
    Individual-as-maximizing agent analogies result in a simple understanding of the functioning of the biological world. Identifying the conditions under which individuals can be regarded as fitness maximizing agents is thus of considerable interest to biologists. Here, we compare different concepts of fitness maximization, and discuss within a single framework the relationship between Hamilton’s (J Theor Biol 7:1–16, 1964) model of social interactions, Grafen’s (J Evol Biol 20:1243–1254, 2007a) formal Darwinism project, and the idea of evolutionary stable strategies. We distinguish cases (...)
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  31. Jg Lennox (1991). Fitness and Evolutionary Explanation-Comment. Biology and Philosophy 6 (1):33-37.
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  32. Elisabeth A. Lloyd (2006). Response to Puts and Dawood's 'The Evolution of Female Orgasm: Adaptation or Byproduct?'--Been There. Twin Studies and Human Genetics 9 (4).
  33. Elisabeth A. Lloyd (2001). Science Gone Astray: Evolution and Rape. [REVIEW] Michigan Law Review 99 (6):1536-1559.
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  34. James Maclaurin, Fitness: Philosophical Problems. Encyclopedia of Life Sciences.
    A philosophical discussion of conceptual and theoretical issues raised by the scientific use of the term ‘fitness’ to describe a property of evolving systems.
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  35. Mohan Matthen & André Ariew (2002). Two Ways of Thinking About Fitness and Natural Selection. Journal of Philosophy 99 (2):55-83.
    How do fitness and natural selection relate to other evolutionary factors like architectural constraint, mode of reproduction, and drift? In one way of thinking, drawn from Newtonian dynamics, fitness is one force driving evolutionary change and added to other factors. In another, drawn from statistical thermodynamics, it is a statistical trend that manifests itself in natural selection histories. It is argued that the first model is incoherent, the second appropriate; a hierarchical realization model is proposed as a basis for a (...)
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  36. Richard E. Michod (2005). On the Transfer of Fitness From the Cell to the Multicellular Organism. Biology and Philosophy 20 (5):967-987.
    The fitness of any evolutionary unit can be understood in terms of its two basic components: fecundity (reproduction) and viability (survival). Trade-offs between these fitness components drive the evolution of life-history traits in extant multicellular organisms. We argue that these trade-offs gain special significance during the transition from unicellular to multicellular life. In particular, the evolution of germ–soma specialization and the emergence of individuality at the cell group (or organism) level are also consequences of trade-offs between the two basic fitness (...)
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  37. Susan K. Mills & John H. Beatty (1979). The Propensity Interpretation of Fitness. Philosophy of Science 46 (2):263-286.
    The concept of "fitness" is a notion of central importance to evolutionary theory. Yet the interpretation of this concept and its role in explanations of evolutionary phenomena have remained obscure. We provide a propensity interpretation of fitness, which we argue captures the intended reference of this term as it is used by evolutionary theorists. Using the propensity interpretation of fitness, we provide a Hempelian reconstruction of explanations of evolutionary phenomena, and we show why charges of circularity which have been levelled (...)
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  38. Roberta L. Millstein (forthcoming). Probability in Biology: The Case of Fitness. In A. Hájek & C. R. Hitchcock (eds.), The Oxford Handbook of Probability and Philosophy. Oxford University Press.
    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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  39. D. E. Montoya, D. A. Peck, N. L. Montoya & C. P. Montoya (2009). A Transdisciplinary Perspective Concerning the Origin of the Species: The Migratory Theory of Genetic Fitness. World Futures 65 (3):166 – 175.
    Although the Neo-Darwin Theory of Evolution is one of the most celebrated theories in science, nonetheless it has received many criticisms. These criticisms are documented and a new transdisciplinary theory of origin is introduced. Darwin's original argument was that natural selection, through heritable changes, changed simple organisms over time. These heritable changes are responsible for the complex plethora of life seen around us today. Darwin's original theory, however, was deconstructed after the (...)
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  40. Bence Nanay (2011). Replication Without Replicators. Synthese 179 (455):477.
    According to a once influential view of selection, it consists of repeated cycles of replication and interaction. It has been argued that this view is wrong: replication is not necessary for evolution by natural selection. I analyze the nine most influential arguments for this claim and defend the replication–interaction conception of selection against these objections. In order to do so, however, the replication–interaction conception of selection needs to be modified significantly. My proposal is that replication is not the copying of (...)
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  41. Bence Nanay (2002). The Return of the Replicator: What is Philosophically Significant in a General Account of Replication and Selection? [REVIEW] Biology and Philosophy 17 (1):109-121.
    The aim of this paper is to outline a typologyof selection processes, and show that differentsub-categories have different explanatorypower. The basis of this typology of selectionprocesses is argued to be the difference ofreplication processes involved in them. Inorder to show this, I argue that: 1.Replication is necessary for selection and 2.Different types of replication lead todifferent types of selection. Finally, it isargued that this typology is philosophicallysignificant, since it contrasts cases ofselection (on the basis of the replicationprocesses involved in them) (...)
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  42. Alfred Nordmann (1990). Persistent Propensities: Portrait of a Familiar Controversy. [REVIEW] Biology and Philosophy 5 (4):379-399.
    Susan Mills and John Beatty's propensity interpretation of fitness encountered very different philosophical criticisms by Alexander Rosenberg and Kenneth Waters. These criticisms and the rejoinders to them are both predictable and important. They are predictable as raisingkinds of issues typically associated with disposition concepts (this is established through a systematic review of the problems generated by Carnap's dispositional interpretation of all scientific terms). They are important as referring the resolution of these issues to the development of evolutionary biology. This historical (...)
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  43. Samir Okasha (2009). Individuals, Groups, Fitness and Utility: Multi-Level Selection Meets Social Choice Theory. Biology and Philosophy 24 (5):561-584.
    In models of multi-level selection, the property of Darwinian fitness is attributed to entities at more than one level of the biological hierarchy, e.g. individuals and groups. However, the relation between individual and group fitness is a controversial matter. Theorists disagree about whether group fitness should always, or ever, be defined as total (or average) individual fitness. This paper tries to shed light on the issue by drawing on work in social choice theory, and pursuing an analogy between fitness and (...)
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  44. Samir Okasha & Cédric Paternotte (2014). Adaptation, Fitness and the Selection-Optimality Links. Biology and Philosophy 29 (2):225-232.
    We critically examine a number of aspects of Grafen’s ‘formal Darwinism’ project. We argue that Grafen’s ‘selection-optimality’ links do not quite succeed in vindicating the working assumption made by behavioural ecologists and others—that selection will lead organisms to exhibit adaptive behaviour—since these links hold true even in the presence of strong genetic and developmental constraints. However we suggest that the selection-optimality links can profitably be viewed as constituting an axiomatic theory of fitness. Finally, we compare Grafen’s project with Fisher’s ‘fundamental (...)
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  45. J. G. Ollason (1991). What is This Stuff Called Fitness? Biology and Philosophy 6 (1):81-92.
    This paper considers a variety of attempts to define fitness in such a way as to defend the theory of evolution by natural selection from the criticism that it is a circular argument. Each of the definitions is shown to be inconsistent with the others. The paper argues that the environment in which an animal evolves can be defined only with respect to the properties of the phenotype of the animal and that it is therefore not illuminating to try to (...)
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  46. Jun Otsuka, Trin Turner, Colin Allen & Elisabeth Lloyd (2011). Why the Causal View of Fitness Survives. Philosophy of Science 78 (2):209-224.
    We critically examine Denis Walsh’s latest attack on the causalist view of fitness. Relying on Judea Pearl’s Sure-Thing Principle and geneticist John Gillespie’s model for fitness, Walsh has argued that the causal interpretation of fitness results in a reductio. We show that his conclusion only follows from misuse of the models, that is, (1) the disregard of the real biological bearing of the population-size parameter in Gillespie’s model and (2) the confusion of the distinction between ordinary probability and Pearl’s causal (...)
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  47. Cedric Paternotte (forthcoming). Shared Adaptiveness is Not Group Adaptation - Commentary of E. Van der Vliert's 'Climato-Economic Habitats Support Patterns of Human Needs, Stresses, and Freedoms'. Behavioral and Brain Sciences.
    Climate stresses and monetary resources seem to lead to different collective adaptations. However, the reference to adaptation and to ambiguous collective dimensions appears premature; populations may entertain nothing more than shared adaptiveness. At this point, the intricacy of the underlying evolutionary processes (cultural selection, fitness-utility decoupling) very much obscures any diagnosis based on correlations.
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  48. Charles H. Pence & Grant Ramsey (2013). A New Foundation for the Propensity Interpretation of Fitness. British Journal for the Philosophy of Science 64 (4):851-881.
    The propensity interpretation of fitness (PIF) is commonly taken to be subject to a set of simple counterexamples. We argue that three of the most important of these are not counterexamples to the PIF itself, but only to the traditional mathematical model of this propensity: fitness as expected number of offspring. They fail to demonstrate that a new mathematical model of the PIF could not succeed where this older model fails. We then propose a new formalization of the PIF that (...)
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  49. Beth Preston (2009). Biological and Cultural Proper Functions in Comparative Perspective. In Ulrich Krohs & Peter Kroes (eds.), Functions in Biological and Artificial Worlds: Comparative Philosophical Perspectives. Mit Press.
    Both biological traits and artifacts have proper functions. But accounts of proper function are typically based on the biological case. So adapting these accounts to the artifact case requires finding cultural analogues of biological concepts. This can go wrong in two ways. The biological concepts may not pick out either biological or cultural proper functions correctly; or they may have no cultural analogues. I argue that things have gone wrong in the first way with regard to selection and in the (...)
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  50. Grant Ramsey (2013). Can Fitness Differences Be a Cause of Evolution? Philosophy and Theory in Biology 5 (20130604).
    Biological fitness is a foundational concept in the theory of natural selection. Natural selection is often defined in terms of fitness differences as “any consistent difference in fitness (i.e., survival and reproduction) among phenotypically different biological entities” (Futuyma 1998, 349). And in Lewontin’s (1970) classic articulation of the theory of natural selection, he lists fitness differences as one of the necessary conditions for evolution by natural selection to occur. Despite this foundational position of fitness, there remains much debate over the (...)
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