While niche construction theory locates animal artefacts in their constructors’ environment, hence treating them as capable of exerting selective pressure on both the constructors and their descendants, the extended phenotype concept assimilates artefacts with their constructors’ genes. Analogous contrasts apply in the case of endoparasite and brood parasite genes influencing host behaviour. The explanatory power of these competing approaches are assessed by re-examining the core chapters of Richard Dawkins’ _The Extended Phenotype_. Because animal artefacts have multiple evolutionary consequences for their (...) constructors, the extra-body effects of a gene seemingly include feedback effects on multiple other genes, a result which is more consistent with niche construction theory than with selfish gene theory. In the case of endoparasite genes influencing host behaviour, Dawkins’ argument leaves out what appears to be the key explanatory component, namely the role of the host’s own bodily systems in making it possible for such genes to exist. For action at a distance, it is unclear whether the key genes have extended effects because they sit in the body of the manipulating organism, or alternatively do not have such effects because they sit in the body of its victim. It is argued that niche construction theory offers a superior explanation in all three cases, regardless of whether the extended phenotype concept is interpreted in selfish gene or selfish organism terms. (shrink)

In his 2009 monograph, Darwinian Populations and Natural Selection, Peter Godfrey-Smith accuses biologists of demonstrating ‘Darwinian Paranoia’ when they engage in what he dubs ‘agential thinking’. But as Daniel Dennett points out, he offers neither an illuminating set of examples nor an extended argument for this assertion, deeming it to be a brilliant propaganda stroke against what is actually a useful way of thinking. Compared to the dangers of teleological thinking in biology, the dangers of agential thinking have unfortunately rarely (...) been discussed. Drawing on recent work by Samir Okasha, I attempt to remedy this omission, through analyzing the nature of agential thinking, and providing a philosophical treatment of the unexamined dangers in this peculiar, yet tempting way of thinking. (shrink)

At the core of anthropomorphism lies a false-positive cognitive bias to over-attribute the pattern of the human body and/or mind. Anthropomorphism is independently discussed in various disciplines, is presumed to have deep biological roots, but its cognitive bases are rarely explored in an integrative way. I present an inclusive, multifaceted interdisciplinary approach to refine the psychological bases of mental anthropomorphism. I have integrated 13 conceptual dissections of folk finalistic reasoning into four psychological inference systems (physical, design, basic-goal and belief stances); (...) the latter three are truly teleological and thus prone to anthropomorphisms. I then have empirically integrated the cross-disciplinary genetic, neural, cognitive, psychiatric, developmental, comparative and evolutionary/adaptive evidence that converge supporting the nature of the distinct stances. The over-reactive calibration of the three teleological systems is framed as an evolved design feature to avoid harmful ancestral contexts. Nowadays, these stances easily engage with scientific reasoning about bio-evolutionary matters with negative and positive consequences. Design, basic-goal, and belief stances benefit biology by providing cognitive foundations, expressing a high-powered explanatory system, promoting functional generalization, fostering new research questions and discoveries, enabling metaphorical/analogical thinking and explaining didactically with brevity. Hence, it is neither feasible nor advantageous to completely eliminate teleology from biology. Instead, we should engage with the eight classes of problems in bio-philosophy and bio-education that relate to the three stances: types of anthropomorphism, variety of misunderstandings, misleading appeal, legitimacy controversy, gateway to mysticism, total prohibition and its backfire effect. Recognizing the distinction among design, basic-goal, and belief stances helps to elucidate much of the logic underlying these issues, so that it enables a much more detailed taxonomy of anthropomorphisms, and organizes the various misunderstandings about evolution by natural selection. It also offers a solid psychological grounding for anchoring definitions and terminology. This tripartite framework also shed some light on how to better deal with the over-reactive stances in bio-education, by organizing previous pedagogical strategies and by suggesting new possibilities to be tested. Therefore, this framework constitutes a promising approach to advance the debate regarding the psychological underpinnings of anthropomorphisms and to further support regulating and clarifying teleology and anthropomorphism in biology. (shrink)

The concept of biological inheritance has recently been extended so as to integrate, among other elements, parts of organisms’ environments. The literature refers to the trans-generational reconstruction of these parts in terms of environmental or ecological inheritance. This article’s main objective is to clarify the different meanings of "environmental inheritance," to underline so far unnoticed theoretical difficulties associated to this polysemous notion and to consequently argue that inheritance, even when extended, should be theoretically distinguished from trans-generational environmental stability. After disentangling (...) the different meanings of environmental inheritance, I underline that studies dealing with this concept place themselves in the wake of earlier contributions about biological environment and elaborate on the role of organisms in the determination of their relevant developmental and selective surroundings. This leads me to question the legitimacy of the category shift operated by niche inheritance proponents—from environment to inheritance—and to explain why the very concept of inherited environment shows important and so far unnoticed theoretical limitations. In this context, I assert the necessity to distinguish two related but different research programs: the construction of a finer-grained theory of environment and the elaboration of an extended theory of inheritance. (shrink)

The concept of biological inheritance has recently been extended so as to integrate, among other elements, parts of organisms’ environments. The literature refers to the trans-generational reconstruction of these parts in terms of environmental or ecological inheritance. This article’s main objective is to clarify the different meanings of "environmental inheritance," to underline so far unnoticed theoretical difficulties associated to this polysemous notion and to consequently argue that inheritance, even when extended, should be theoretically distinguished from trans-generational environmental stability. After disentangling (...) the different meanings of environmental inheritance, I underline that studies dealing with this concept place themselves in the wake of earlier contributions about biological environment and elaborate on the role of organisms in the determination of their relevant developmental and selective surroundings. This leads me to question the legitimacy of the category shift operated by niche inheritance proponents—from environment to inheritance—and to explain why the very concept of inherited environment shows important and so far unnoticed theoretical limitations. In this context, I assert the necessity to distinguish two related but different research programs: the construction of a finer-grained theory of environment and the elaboration of an extended theory of inheritance. (shrink)

Both Aristotle and Ernst Mayr present theories of dual explanation in biology, with proximal, clearly physical explanations and more distal, biology-specific explanations. Aristotle’s presentation of final cause explanations in Posterior Analytics relates final causes to the necessary material, formal, and efficient causes that mediate them. Johnson and Leunissen demonstrate the problematic nature of historical and recent interpretations and open the door for a new interpretation consistent with modern evolutionary theory. Mayr’s differentiation of proximate and ultimate/evolutionary causes provides a key to (...) appropriating Aristotle for modern use, if care is taken to differentiate Mayr’s proximate/distal and mechanical/evolutionary distinctions from his intentional/non-intentional distinction. Building on Mayr, and noting his appropriation of ancient and recent commentary, a strong case can be made for nested explanation, wherein evolutionary explanations are instantiated in systems of mechanical explanations. Biological concepts of organism, gene, function, etc., are etiologically prior to mechanical explanations of function, but temporally posterior to mechanical explanations of historical adaptation. Such an analysis sheds light on recent arguments within evolutionary biology while highlighting the importance of epistemology in contemporary science. (shrink)

The primary challenge for generating a useful scientific definition of life comes from competing concepts of biological activity and our failure to make them explicit in our models. I set forth a three-part scheme for characterizing definitions of life, identifying a binary , a range , and a preference . The three components together form a proper activity in biology . To be clear, I am not proposing that proper activity be adopted as the best definition of life or even (...) as a desirable definition for life. Instead, I am arguing that some notion of proper activity already exists within common scientific definitions. By making the implicit elements explicit, the notion can be analyzed to see whether it is useful and appropriate in the context of the biological sciences. (shrink)

A high heritability estimate usually corresponds to a situation in which trait variation is largely caused by genetic variation. However, in some cases of gene-environment covariance, causal intuitions about the sources of trait difference can vary, leading experts to disagree as to how the heritability estimate should be interpreted. We argue that the source of contention for these cases is an inconsistency in the interpretation of the concepts ‘genotype’, ‘phenotype’, and ‘environment’. We propose an interpretation of these terms under which (...) trait variance initially caused by genetic variance is subsumed into a heritability for all cases of gene-environment covariance. (shrink)

Advocates of an ‘extended evolutionary synthesis’ have claimed that standard evolutionary theory fails to accommodate epigenetic inheritance. The opponents of the extended synthesis argue that the evidence for epigenetic inheritance causing adaptive evolution in nature is insufficient. We suggest that the ambiguity surrounding the conception of the gene represents a background semantic issue in the debate. Starting from Haig’s gene-selectionist framework and Griffiths and Neumann-Held’s notion of the evolutionary gene, we define senses of ‘gene’, ‘environment’, and ‘phenotype’ in a way (...) that makes them consistent with gene-centric evolutionary theory. We argue that the evolutionary gene, when being materialized, need not be restricted to nucleic acids but can encompass other heritable units such as epialleles. If the evolutionary gene is understood more broadly, and the notions of environment and phenotype are defined accordingly, current evolutionary theory does not require a major conceptual change in order to incorporate the mechanisms of epigenetic inheritance. _1_ Introduction _2_ The Gene-centric Evolutionary Theory and the ‘Evolutionary Gene’ _2.1_ The evolutionary gene _2.2_ Genes, phenotypes, and environments _3_ Epigenetic Inheritance and the Gene-Centred Framework _3.1_ Treating the gene as the sole heritable material? _3.2_ Epigenetics and phenotypic plasticity _4_ Conclusion. (shrink)

In the Sleeping Beauty problem, Beauty is woken once if a coin lands heads or twice if the coin lands tails but promptly forgets each waking on returning to sleep. Philosophers have divided over whether her waking credence in heads should be a half or a third. Beauty has centered beliefs about her world and about her location in that world. When given new information about her location she should update her worldly beliefs before updating her locative beliefs. When she (...) conditionalizes in this way, her credence in heads is a half before and after being told it is Monday. In applications of Dutch Book arguments to the Sleeping Beauty problem, the probability of a particular outcome has often been confounded with consequences of that outcome. Heads and tails are equally likely but twice as much is at stake if the coin falls tails because Beauty is fated to make the same choice twice. As a consequence, the possibility of tails should be given twice the weight of the possibility of heads when deciding whether to bet on heads even though heads and tails are equally likely. (shrink)

Organisms and their genomes are mosaics of features of different evolutionary age. Older features are maintained by ‘negative’ selection and comprise part of the selective environment that has shaped the evolution of newer features by ‘positive’ selection. Body plans and body parts are among the most conservative elements of the environment in which genetic differences are selected. By this process, well-trodden paths of development constrain and direct paths of evolutionary change. Structuralism and adaptationism are both vindicated. Form plays a selective (...) role in the molding of form. (shrink)

Concepts of cause, choice, and information are closely related. A cause is a choice that can be held responsible. It is a difference that makes a difference. Information about past causes and their effects is a valuable commodity because it can be used to guide future choices. Information about criteria of choice is generated by choosing a subset from an ensemble for ‘reasons’ and has meaning for an interpreter when it is used to achieve an end. Natural selection evolves interpreters (...) with ends. Surviving genes embody a textual record of past choices that had favorable outcomes. Consultation of these archives guides current choices. Purposive choice is well-informed difference making. (shrink)

Organisms can be treated as optimizers when there is consensus among their genes about what is best to be done, but genomic consensus is often lacking, especially in interactions among kin because kin share some genes but not others. Grafen adopts a majoritarian perspective in which an individual’s interests are identified with the interests of the largest coreplicon of its genome, but genomic imprinting and recombination factionalize the genome so that no faction may predominate in some interactions among kin. Once (...) intragenomic conflicts are recognized, the individual organism can be conceptualized as an arbiter among competing interests within a collective. Organismal adaptation can be recognized without phenotypes being optimized. (shrink)

Although classical evolutionary theory, i.e., population genetics and the Modern Synthesis, was already implicitly ‘gene-centred’, the organism was, in practice, still generally regarded as the individual unit of which a population is composed. The gene-centred approach to evolution only reached a logical conclusion with the advent of the gene-selectionist or gene’s eye view in the 1960s and 1970s. Whereas classical evolutionary theory can only work with (genotypically represented) fitness differences between individual organisms, gene-selectionism is capable of working with fitness differences (...) among genes within the same organism and genome. Here, we explore the explanatory potential of ‘intra-organismic’ and ‘intra-genomic’ gene-selectionism, i.e., of a behavioural-ecological ‘gene’s eye view’ on genetic, genomic and organismal evolution. First, we give a general outline of the framework and how it complements the—to some extent—still ‘organism-centred’ approach of classical evolutionary theory. Secondly, we give a more in-depth assessment of its explanatory potential for biological evolution, i.e., for Darwin’s ‘common descent with modification’ or, more specifically, for ‘historical continuity or homology with modular evolutionary change’ as it has been studied by evolutionary developmental biology (evo-devo) during the last few decades. In contrast with classical evolutionary theory, evo-devo focuses on ‘within-organism’ developmental processes. Given the capacity of gene-selectionism to adopt an intra-organismal gene’s eye view, we outline the relevance of the latter model for evo-devo. Overall, we aim for the conceptual integration between the gene’s eye view on the one hand, and more organism-centred evolutionary models (both classical evolutionary theory and evo-devo) on the other. (shrink)

Although classical evolutionary theory, i.e., population genetics and the Modern Synthesis, was already implicitly ‘gene-centred’, the organism was, in practice, still generally regarded as the individual unit of which a population is composed. The gene-centred approach to evolution only reached a logical conclusion with the advent of the gene-selectionist or gene’s eye view in the 1960s and 1970s. Whereas classical evolutionary theory can only work with fitness differences between individual organisms, gene-selectionism is capable of working with fitness differences among genes (...) within the same organism and genome. Here, we explore the explanatory potential of ‘intra-organismic’ and ‘intra-genomic’ gene-selectionism, i.e., of a behavioural-ecological ‘gene’s eye view’ on genetic, genomic and organismal evolution. First, we give a general outline of the framework and how it complements the—to some extent—still ‘organism-centred’ approach of classical evolutionary theory. Secondly, we give a more in-depth assessment of its explanatory potential for biological evolution, i.e., for Darwin’s ‘common descent with modification’ or, more specifically, for ‘historical continuity or homology with modular evolutionary change’ as it has been studied by evolutionary developmental biology during the last few decades. In contrast with classical evolutionary theory, evo-devo focuses on ‘within-organism’ developmental processes. Given the capacity of gene-selectionism to adopt an intra-organismal gene’s eye view, we outline the relevance of the latter model for evo-devo. Overall, we aim for the conceptual integration between the gene’s eye view on the one hand, and more organism-centred evolutionary models on the other. (shrink)

I argue that Grafen’s formal darwinism project could profitably incorporate a gene’s-eye view, as informed by the major transitions framework. In this, instead of the individual being assumed to maximise its inclusive fitness, genes are assumed to maximise their inclusive fitness. Maximisation of fitness at the individual level is not a straightforward concept because the major transitions framework shows that there are several kinds of biological individual. In addition, individuals have a definable fitness, exhibit individual-level adaptations and arise in a (...) major transition, only to the extent that the inclusive-fitness interests of genes within them coincide. Therefore, as others have suggested, the fundamental level at which fitness is maximised is the gene level. Previous reconciliations of the concepts of gene-level fitness and individual-level fitness implicitly recognise this point. Adaptations always maximise the fitness of their causative genes, but may be simple or complex. Simple adaptations may be controlled by single genes and be maladaptive at higher levels, whereas complex adaptations are controlled by multiple genes and rely on those genes having coinciding fitness interests at a higher level, for a given trait. (shrink)

In this paper I critically evaluate Reisman and Forber’s :1113–1123, 2005) arguments that drift and natural selection are population-level causes of evolution based on what they call the manipulation condition. Although I agree that this condition is an important step for identifying causes for evolutionary change, it is insufficient. Following Woodward, I argue that the invariance of a relationship is another crucial parameter to take into consideration for causal explanations. Starting from Reisman and Forber’s example on drift and after having (...) briefly presented the criterion of invariance, I show that once both the manipulation condition and the criterion of invariance are taken into account, drift, in this example, should better be understood as an individual-level rather than a population-level cause. Later, I concede that it is legitimate to interpret natural selection and drift as population-level causes when they rely on genuinely indeterministic events and some cases of frequency-dependent selection. (shrink)

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...

Altruism is one of the most studied topics in theoretical evolutionary biology. The debate surrounding the evolution of altruism has generally focused on the conditions under which altruism can evolve and whether it is better explained by kin selection or multilevel selection. This debate has occupied the forefront of the stage and left behind a number of equally important questions. One of them, which is the subject of this article, is whether the word “selection” in “kin selection” and “multilevel selection” (...) necessarily refers to “evolution by natural selection.” I show, using a simple individual-centered model, that once clear conditions for natural selection and altruism are specified, one can distinguish two kinds of evolution of altruism, only one of which corresponds to the evolution of altruism by natural selection, the other resulting from other evolutionary processes. (shrink)

Sociological, economic and evolutionary paradigms of human agency have often seen social agents either as the rational controllers of their fate or as marionettes on the strings of historical, functional or adaptive necessity. They found it therefore difficult to account for the variability, intentionality and creativity of human behaviour and for its frequently redundant or harmful results. This paper argues that human agency is a product of evolution, but that genetic variation and inheritance can only provide a limited explanation of (...) its complex nature. The primary evolutionary problem which human agents face while they are alive is not to adapt to stable environments, but to respond flexibly and creatively to a contingent, uncertain world. Variation and selection therefore take two connected but distinct forms, one external, genetic, and inherited across generations, the other internal and cognitive, and operating during the lifetime of individuals. An examination of this lived part of evolution provides a better understanding of key properties of agency. (shrink)

Causation has multiple distinct meanings in genetics. One reason for this is meaning slippage between two concepts of the gene: Mendelian and molecular. Another reason is that a variety of genetic methods address different kinds of causal relationships. Some genetic studies address causes of traits in individuals, which can only be assessed when single genes follow predictable inheritance patterns that reliably cause a trait. A second sense concerns the causes of trait differences within a population. Whereas some single genes can (...) be said to cause population-level differences, most often these claims concern the effects of many genes. Polygenic traits can be understood using heritability estimates, which estimate the relative influences of genetic and environmental differences to trait differences within a population. Attempts to understand the molecular mechanisms underlying polygenic traits have been developed, although causal inference based on these results remains controversial. Genetic variation has also recently been leveraged as a randomizing factor to identify environmental causes of trait differences. This technique—Mendelian randomization—offers some solutions to traditional epidemiological challenges, although it is limited to the study of environments with known genetic influences. (shrink)