The developmental properties of organisms play important roles in the generation of variation necessary for evolutionary change. But how can individual development steer the course of evolution? To answer this question, we introduce developmental channeling as a disposition of individual organisms that shapes their possible developmental trajectories and evolutionary dappling as an evolutionary outcome in which the space of possible organismic forms is dappled—it is only partially filled. We then trace out the implications of the channeling-dappling framework for contemporary debates (...) in the philosophy of evolution, including evolvability, reciprocal causation, and the extended evolutionary synthesis. (shrink)

To a large extent, the evidential base of claims in the philosophy of science has switched from thought experiments to case studies. We argue that abandoning thought experiments was a wrong turn, since they can effectively complement case studies. We make our argument via an analogy with the relationship between experiments and observations within science. Just as experiments and ‘natural’ observations can together evidence claims in science, each mitigating the downsides of the other, so too can thought experiments and case (...) studies be mutually supporting. After presenting the main argument, we look at potential concerns about thought experiments, suggesting that a judiciously applied mixed-methods approach can overcome them. (shrink)

Few debates in environmental philosophy have been more heated than the one over the nature of wilderness. And yet, when one surveys the present scene, one finds that a variety of different conceptions of wilderness are still quite popular – some more so in certain professions than others. In this paper, I look at three popular conceptions of wilderness with an eye toward sussing out the good and the bad them. I look at what I call (1) the folk view (...) of wilderness, (2) Leopold’s conception of wilderness, and (3) the legal conception of wilderness (as found in the Wilderness Act of 1964). In the final part of the paper, I sketch out a sort of spectrum account of wilderness, one that I argue allows us to capture more cases of wilderness and might serve as a useful tool in future conservation efforts. (shrink)

In this paper, I will conduct three interrelated analyses. First, I will develop an analysis of various concepts in the history of biology that used to refer to individual-level phenomena but were then reinterpreted by the Modern Synthesis in terms of populations. Second, I argue that a similar situation can be found in contemporary biological theory. While different approaches reflect on the causal role of developing organisms in evolution, proponents of the Modern Synthesis avoid any substantial change by reinterpreting and (...) explaining individual-level phenomena from a population perspective. Finally, I will approach this debate by advocating the statistical reading of natural selection, which holds that explanations by natural selection are statistical. I will argue that the above historical conceptual reinterpretations belong to a new explanatory strategy developed by the Modern Synthesis based on population thinking. The reinterpretation of concepts at the individual level is part of the explanatory framework of the Modern Synthesis and the empty role of development within this framework. Moreover, the statistical perspective adopted here allows for the integration of two explanatory models: population-statistical and individual-causal. Finally, I will argue that this pluralistic framework can help to define the explanatory scope of the different biological approaches in order to achieve a coherent integration of development into evolution without rejecting population thinking. (shrink)

An influential heuristic for thinking about climate adaptation asserts that “natural” adaptation strategies are the best ones. This heuristic has been roundly criticized but is difficult to dislodge in the absence of an alternative. We introduce a new heuristic that assesses adaptation strategies by looking at their maturity, power, and commitment. Maturity is the extent to which we understand an adaptation strategy’s effects. Power is the size of the effect an adaptation strategy will have. Commitment is the degree to which (...) an adaptation strategy is difficult to test or reverse. (shrink)

In this paper, I introduce the notion of ‘philosophy moves’: prominent tropes featured in contemporary academic philosophy. Moves are more than patterns—they are tools for advancing and enriching philosophical debates. By recognizing these patterns in the philosophical literature, we collect an ensemble of moves for deployment in novel contexts, each with the potential to forge new paths of philosophical investigation through a given topic. The moves featured in this paper are constructive and progressive, with the potential to push past stalemates (...) and open a wider conceptual landscape. Following a presentation of five exemplars taken from the philosophy of biology literature, I show how different moves could be made within a current interdisciplinary debate. Overall, I aim to demonstrate that recognizing these patterns is insightful pedagogically and potentially useful when used as part of one’s philosophical methodology. (shrink)

How all philosophical explanations of human consciousness and the fundamental structure of the cosmos are bizarre—and why that’s a good thing Do we live inside a simulated reality or a pocket universe embedded in a larger structure about which we know virtually nothing? Is consciousness a purely physical matter, or might it require something extra, something nonphysical? According to the philosopher Eric Schwitzgebel, it’s hard to say. In The Weirdness of the World, Schwitzgebel argues that the answers to these fundamental (...) questions lie beyond our powers of comprehension. We can be certain only that the truth—whatever it is—is weird. Philosophy, he proposes, can aim to open—to reveal possibilities we had not previously appreciated—or to close, to narrow down to the one correct theory of the phenomenon in question. Schwitzgebel argues for a philosophy that opens. According to Schwitzgebel’s “Universal Bizarreness” thesis, every possible theory of the relation of mind and cosmos defies common sense. According to his complementary “Universal Dubiety” thesis, no general theory of the relationship between mind and cosmos compels rational belief. Might the United States be a conscious organism—a conscious group mind with approximately the intelligence of a rabbit? Might virtually every action we perform cause virtually every possible type of future event, echoing down through the infinite future of an infinite universe? What, if anything, is it like to be a garden snail? Schwitzgebel makes a persuasive case for the thrill of considering the most bizarre philosophical possibilities. (shrink)

Is the ability to speak a language an acquired skill? Leading proponents of the generative approach to human language—notably Chomsky (2000) and Pinker (2003)—have argued that the thesis that language capacities are skills is hopelessly confused and at odds with a range of empirical evidence, which suggests that human language capacities are grounded in a biologically inherited set of language instincts or a Universal Grammar (UG). In this paper, we argue that resistance to the claim that human language capacities are (...) skills has been fostered by naïve and implausible conceptions of the nature of skilled action. Correcting these misconceptions about the nature of skilled action breathes new life into the old idea that human language competences are particular kinds of skills and does so in a way that is fully consistent with appeals to the central importance of UG, broadly construed. More generally, we argue that by taking human language capacities to be skills, we capture what is right about the generative program while at the same time avoiding its most serious pitfalls. (shrink)

This paper focuses on physiological integration in multicellular systems, a notion often associated with biological individuality, but which has not received enough attention and needs a thorough theoretical treatment. Broadly speaking, physiological integration consists in how different components come together into a cohesive unit in which they are dependent on one another for their existence and activity. This paper argues that physiological integration can be understood by considering how the components of a biological multicellular system are controlled and coordinated in (...) such a way that their activities can contribute to the maintenance of the system. The main implication of this perspective is that different ways of controlling their parts may give rise to multicellular organizations with different degrees of integration. After defining control, this paper analyses how control is realized in two examples of multicellular systems located at different ends of the spectrum of multicellularity: biofilms and animals. It focuses on differences in control ranges, and it argues that a high degree of integration implies control exerted at both medium and long ranges, and that insofar as biofilms lack long-range control (relative to their size) they can be considered as less integrated than other multicellular systems. It then discusses the implication of this account for the debate on physiological individuality and the idea that degrees of physiological integration imply degrees of individuality. (shrink)

Abstract: This paper argues that a central explanatory role for the concept of Umwelt in theoretical biology is to be found in developmental biology, in particular in the effort to understand development as a goal-directed and adaptive process that is controlled by the organism itself. I will reach this conclusion in two (interrelated) ways. The first is purely theoretical and relates to the current scenario in the philosophy of biology. Challenging neo-Darwinism requires a new understanding of the various components involved (...) in natural selection processes. An important prerequisite for the explanation is the ability to understand development in a teleological way. Here, the concept of Umwelt plays a crucial role: if organisms are responsible for generating adaptive variation in specific environments, we need a theory that explains the context-dependent nature of adaptively oriented processes. The Umwelt is thus a central element in determining the goal that an adaptive process pursues. The second path in my analysis also has a historical dimension. I will present Karl Ernst von Baer’s reflections on teleological development and his influence on von Uexküll's thinking. I will present various ideas developed by von Baer, such as the distinction between Ziel and Zweck and the use of musical metaphors, which can help to understand development teleologically and give von Uexküll’s theory a central place in this framework. (shrink)

Millikan’s (1984. Language, Thought, and Other Biological Categories: New Foundations for Realism. MIT Press) selected effects theory of functions states that functions are effects for which the ancestors of a trait were selected for. As the function is an effect a thing’s ancestors produced, only things that are reproductions in some sense can have functions. Against this reproduction requirement, Garson (2019. What Biological Functions Are and Why They Matter. Cambridge University Press) argues that not only processes of differential reproduction but (...) also processes of differential persistence can lead to new functions. Since such “persistence functions” have the same explanatory power as traditional selected effects functions, selected effects theorists should include them in their theory. In this paper, I will defend Millikan’s theory against this argument. I will show that the proponents of the generalized theory have yet to provide a working notion of populations that avoids a liberality problem. Further, I will argue that persistence functions are at best a marginal case of functions due to their restricted explanatory power. (shrink)

Agency is a central concept in the organisational approach to organisms, which accounts for their internal purposiveness. Recent recognition of the active role played by organisms in evolution has led researchers to use this concept in an evolutionary approach. Agency is then considered in terms of ‘unintentional’ choice: agents choose from a given repertoire the behaviour most appropriate to their goal, with this choice influencing evolutionary pathways. This view, while allowing for the evolutionary role of the activity of organisms, presents (...) two pitfalls. First, it restricts organisms’ agency by confining their choice within the bounds of a behavioural repertoire, and assuming their goals are dictated by natural selection. Second, this view, while claiming to eliminate the idea of intentionality, retains its structure: organisms are portrayed as rational entities, persistently pursuing specific goals. This leads us back to a teleological thinking, whose use in evolutionary theory has already been heavily criticised. This paper proposes a conception of biological agency which does not assume goal-directedness but considers agency as inventiveness. An organism will be said to be an agent if it is the triggering cause of behaviours falling outside the known repertoire and whose form can only be explained by the unique relationship between the organism and the environment. If these behaviours have implications in evolution, the agent will be considered an evolutionary agent. The merit of this approach is further validated by evidencing the significant role behavioural innovations play in evolution. Finally, the last section delves into the process of invention by examining animal play. (shrink)

The foundations of biology have been a topic of debate for the past few decades. The traditional perspective of the Modern Synthesis, which portrays organisms as passive entities with limited role in evolutionary theory, is giving way to a new paradigm where organisms are recognized as active agents, actively shaping their own phenotypic traits for adaptive purposes. Within this context, this article raises the question of whether contemporary biological theory is undergoing a cognitive revolution. This inquiry can be approached in (...) two ways: from a theoretical standpoint, exploring the centrality of the cognitive sciences in current theoretical biology; and from a historical perspective, examining the resemblance between the current state of theoretical biology and the Cognitive Revolution of the mid-20th century. Both inquiries yield affirmative answers, though important nuances will be emphasized. The cognitive sciences' explanatory framework is employed to elucidate the agentic characteristics of organisms, establishing a clear parallelism between the Cognitive Revolution and the present state of theoretical biology. (shrink)

The field of the philosophy of biology is flourishing in its aim to evaluate and rethink the view inherited from the previous century ---the Modern Synthesis. Different research areas and theories have come to the fore in the last decades in order to account for different biological phenomena that, in the first instance, fall beyond the explanatory scope of the Modern Synthesis. This thesis is anchored and motivated by this revolt in the philosophy of biology. -/- The central target in (...) this context is the possibility of naturalizing teleology, a classical nightmare for the history of biology itself. This requires, principally, understanding the causes of teleological explanations without assuming an unfashioned backward causation of sorts. As the riddles of teleological explanations are about their temporal dimension, I analyze different temporal scales of biological processes: evolutionary, developmental, and physiological. -/- The first one is the one defended in the context of the Modern Synthesis. As expected, one of the aims of this thesis is to evaluate the adequacy of an evolutionary account of teleological explanation. The scrutiny is negative. Evolutionary explanations in the context of the Modern Synthesis lack the necessary causal roots to naturalize teleology. Concerning the physiological scale, a long tradition pushed up by Kant and the organicist movement in the 20th century allows us to better understand how teleological explanations can be naturalized in physiological process. The key notions in this temporal scale are self-organization and the recursive, looped character of physiological process. While the physiological scale may be suitably accounted by contemporaries views, such as Autonomous Systems Theory, different central teleological phenomena remain unexplained from a purely physiological perspective. In particular, different issues concerning the (adaptive) construction of organism ---such as plasticity, robustness, variation, novelty, inheritance--- deserve an ontogenetic analysis. -/- The principal aim of this thesis is to provide a theory of teleological development that falls beyond the Modern Synthesis' framework and is prompted by different insights from the history of biology. I call it Agential Teleosemantics. It rests on two central pillars. First, that developmental processes, beyond any gene-centered stance, can be understood in informational terms; i.e. developmental processes are about the interaction of developmental resources conveying biological information. The second ingredient is agentivity, namely the idea that development is regulated by an agentive system according to the adaptiveness of the phenotypic outcomes produced. The role of agency in Agential Teleosemantic is equivalent to the role of genes in the Modern Synthesis: it is responsible for explaining the order and the adaptive complexity in the living realm. -/- The second target of this thesis regards the possibility of naturalizing intentional explanations in cognitive science. The central project involved in such an aim is known as teleosemantics. Classical teleosemantics however is etiological: it explains the teleofunctions of representational systems in terms of evolutionary processes. The different disputes in the contemporary philosophy of biology provide two insights to analyze teleosemantics in cognitive science. First, the challenges against the Modern Synthesis must be extended to the evolutionary approach of etiological teleosemantics. Second, as Agential Teleosemantics suggests an alternative source for teleofunctions ---ontogeny, I offer an attempt to integrate Agential Teleosemantics into cognitive science in order to provide an alternative teleosemantic project to understand intentional explanations in cognitive science. (shrink)

Cancer biologists ascribe normal functions to parts of cancer. Normal functions are activities that parts of systems are in some minimal sense supposed to perform. Cancer biologists’ finding normality within the abnormality of cancer pose difficulties for two main approaches to normal function. One approach claims that normal functions are activities that parts are selected for. However, some parts of cancers that have normal functions aren’t selected to perform them. The other approach claims that normal functions are part-activities typical for (...) the system and that contribute to survival/reproduction. However, cancers are too heterogeneous to establish what’s typical across a type. (shrink)

Cancer biology features the ascription of normal functions to parts of cancers. At least some ascriptions of function in cancer biology track local normality of parts within the global abnormality of the aberration to which those parts belong. That is, cancer biologists identify as functions activities that, in some sense, parts of cancers are supposed to perform, despite cancers themselves having no purpose. The present paper provides a theory to accommodate these normal function ascriptions—I call it the Modeling Account of (...) Normal Function (MA). MA comprises two claims. First, normal functions are activities whose performance by the function-bearing part contributes to the self-maintenance of the whole system and, thereby, results in the continued presence of that part. Second, MA holds that models of system-level activities that are (partly) constitutive of self-maintenance are improved by including a representation of the relevant function-bearing part and by making reference to the activity/activities that part performs which contribute(s) to those system-level activities. I contrast MA with two other accounts that seek to explicate the ascription of normal functions in biology, namely, the organizational account and the selected effects account. Both struggle to extend to cancer biology. However, I offer ecumenical readings which allow them to recover some ascriptions of normal function to parts of cancers. So, though I contend that MA excels in this respect, the purpose of this paper is served if it provides materials for bridging the gap between cancer biology, philosophy of cancer, and the literature on function. (shrink)

En los últimos años, algunos autores han venido sosteniendo que la validez de la representación arborescente del patrón filético generado por la evolución está siendo menoscabada por el reconocimiento del impacto evolutivo que tendrían la transferencia genética horizontal, la simbiogénesis, la especiación por hibridación y la introgresión. Esa concepción o representación cladogenética de la evolución sólo nos dejaría ver un aspecto parcial de las relaciones de filiación que conectan a los diferentes linajes de seres vivos; ocultando otro aspecto cuya representación (...) más adecuada sería un reticulado de relaciones entre taxones. Con todo, aún sin poner en tela de juicio la frecuencia y la relevancia explicativa de dichos fenómenos, en este trabajo se intentará mostrar que su admisión y reconocimiento no tiene por qué ir en detrimento de esa concepción cladogenética de la evolución cuya relativización se está pregonando. Por el contrario, la comprensión y la representación de tales fenómenos supone dicha concepción. Palabras clave: Filogénesis, Especiación por hibridación, Introgresión, Simbiogénesis, TGH. (shrink)

In this paper I develop a concept of behavioural ecological individuality. Using findings from a case study which employed qualitative methods, I argue that individuality in behavioural ecology should be defined as phenotypic and ecological uniqueness, a concept that is operationalised in terms of individual differences such as animal personality and individual specialisation. This account make sense of how the term “individuality” is used in relation to intrapopulation variation in behavioural ecology. The concept of behavioural ecological individuality can sometimes be (...) used to identify individuals. It also shapes research agendas and methodological choices in behavioural ecology, leading researchers to account for individuals as sources of variation. Overall, this paper draws attention to a field that has been largely overlooked in philosophical discussions of biological individuality and highlights the importance of individual differences and uniqueness for individuality in behavioural ecology. (shrink)

Can heritability estimates provide causal information? This paper argues for an affirmative answer: since a non-nil heritability estimate satisfies certain characteristic properties of causation (i.e., association, manipulability, and counterfactual dependence), it increases the probability that the relation between genotypic variance and phenotypic variance is (at least partly) causal. Contrary to earlier proposals in the literature, the argument does not assume the correctness of any particular conception of the nature of causation, rather focusing on properties that are characteristic of causal relationships. (...) The argument is defended against Lewontin's (1974) locality objection and Kaplan and Turkheimer's (2021) recent critique of Genome-Wide Association Studies (GWAS). (shrink)

This paper introduces the reconstitutor as a comprehensive unit of heredity within the context of evolutionary research. A reconstitutor is the structure resulting from a set of relationships between different elements or processes that are actively involved in the recreation of a specific phenotypic variant in each generation regardless of the biomolecular basis of the elements or whether they stand in a continuous line of ancestry. Firstly, we justify the necessity of introducing the reconstitutor by showing the limitations of other (...) evolutionary conceptions of the unit of heredity, such as the replicator, the reproducer, and the Darwinian individual. We argue that these conceptions are based on the requirement of lineage formation, which we argue to be unnecessary for the existence of evolutionary heredity. In the second part, we introduce the reconstitutor, which we base on the concept of Stability of Traits, and illustrate how it covers cases of hereditary phenomena not covered by the previous accounts. Secondly, we illustrate how the reconstitutor could serve as a platform to rethink ecological inheritance and other forms of inheritance that have been recently introduced under the song/singer model of evolution. (shrink)

The Price equation is currently considered one of the fundamental equations —or even the fundamental equation— of evolution. In this article, we explore the role of this equation within cultural evolutionary theory. More specifically, we use it to account for the explanatory power and the theoretical structure of a certain generalised version of dual-inheritance theory. First, we argue that, in spite of not having a definite empirical content, the Price equation offers a suitable formalisation of the processes of cultural evolution, (...) and provides a powerful heuristic device for discovering the actual causes of cultural change and accumulation. Second, we argue that, as a consequence of this, a certain version of the Price equation is the fundamental law of cultural evolutionary theory. In order to support this claim, we sketch the ideal structure of dual-inheritance theory and we stress the unificatory role that the Price equation plays in it. (shrink)

Philosophers who study the problem of biological function often begin their deliberations by reflecting on the functions of parts of animals, or the behavior of animals. Applying theories of biological function to unconventional or borderline cases can help us to better evaluate and refine those theories. This is the case when we consider whether parts of transposable elements —bits of “selfish” DNA that move about within a host genome—have functions of their own, that is, whether the parts of TEs have (...) the function of helping the TE move about within the genome. Here I argue that whether or not the parts of TEs have functions depends crucially on whether collections of TEs form “populations,” by which I mean, here, a group of individuals of the same type that impact one another’s chances of persistence or multiplication, by impacting one another’s access to a shared resource. I think there is suggestive, but not conclusive, evidence that some TEs have functions of their own. Considering the problem of TE functionality, then, has value both for philosophy and for biology. (shrink)

This thesis focuses on models of conceptual change in science and philosophy. In particular, I developed a new bootstrapping methodology for studying conceptual change, centered around the formalization of several popular models of conceptual change and the collective assessment of their improved formal versions via nine evaluative dimensions. Among the models of conceptual change treated in the thesis are Carnap’s explication, Lakatos’ concept-stretching, Toulmin’s conceptual populations, Waismann’s open texture, Mark Wilson’s patches and facades, Sneed’s structuralism, and Paul Thagard’s conceptual revolutions. (...) -/- In order to analyze and compare the conception of conceptual change provided by these different models, I rely on several historical reconstructions of episodes of scientific conceptual change. The historical episodes of scientific change that figure in this work include the emergence of the morphological concept of fish in biological taxonomies, the development of scientific conceptions of temperature, the Church-Turing thesis and related axiomatizations of effective calculability, the history of the concept of polyhedron in 17th and 18th century mathematics, Hamilton’s invention of the quaternions, the history of the pre-abstract group concepts in 18th and 19th century mathematics, the expansion of Newtonian mechanics to viscous fluids forces phenomena, and the chemical revolution. I will also present five different formal and informal improvements of four specific models of conceptual change. I will first present two different improvements of Carnapian explication, a formal and an informal one. My informal improvement of Carnapian explication will consist of a more fine-grained version of the procedure that adds an intermediate, third step to the two steps of Carnapian explication. I will show how this novel three-step version of explication is more suitable than its traditional two-step relative to handle complex cases of explications. My second, formal improvement of Carnapian explication will be a full explication of the concept of explication itself within the theory of conceptual spaces. By virtue of this formal improvement, the whole procedure of explication together with its application procedures and its pragmatic desiderata will be reconceptualized as a precise procedure involving topological and geometrical constraints inside the theory of conceptual spaces. My third improved model of conceptual change will consist of a formal explication of Darwinian models of conceptual change that will make vast use of Godfrey-Smith’s population-based Darwinism for targeting explicitly mathematical conceptual change. My fourth improvement will be dedicated instead to Wilson’s indeterminate model of conceptual change. I will show how Wilson’s very informal framework can be explicated within a modified version of the structuralist model-theoretic reconstructions of scientific theories. Finally, the fifth improved model of conceptual change will be a belief-revision-like logical framework that reconstructs Thagard’s model of conceptual revolution as specific revision and contraction operations that work on conceptual structures. -/- At the end of this work, a general conception of conceptual change in science and philosophy emerges, thanks to the combined action of the three layers of my methodology. This conception takes conceptual change to be a multi-faceted phenomenon centered around the dynamics of groups of concepts. According to this conception, concepts are best reconstructed as plastic and inter-subjective entities equipped with a non-trivial internal structure and subject to a certain degree of localized holism. Furthermore, conceptual dynamics can be judged from a weakly normative perspective, bound to be dependent on shared values and goals. Conceptual change is then best understood, according to this conception, as a ubiquitous phenomenon underlying all of our intellectual activities, from science to ordinary linguistic practices. As such, conceptual change does not pose any particular problem to value-laden notions of scientific progress, objectivity, and realism. At the same time, this conception prompts all our concept-driven intellectual activities, including philosophical and metaphilosophical reflections, to take into serious consideration the phenomenon of conceptual change. An important consequence of this conception, and of the analysis that generated it, is in fact that an adequate understanding of the dynamics of philosophical concepts is a prerequisite for analytic philosophy to develop a realistic and non-idealized depiction of itself and its activities. (shrink)

This paper isolates a hard, long-standing species problem: developing a comprehensive and exacting theory about the constitutive conditions of the species category, one that is accurate for most of the living world, and which vindicates the widespread view that the species category is of more theoretical import than categories such as genus, sub-species, paradivision, and stirp. The paper then uncovers flaws in several views that imply we have either already solved that hard species problem or dissolved it altogether – so-called (...) We Are Done views. In doing so the paper offers new criticisms of the general lineage species concept (GLSC), evolutionary species concept (EvSC), biological species concept (BSC), other similar concepts, Ereshefsky’s eliminative pluralism about the species category, and both Mishler’s pessimism and Wilkins’ phenomenalism about that category. Opposed to We Are Done views, the paper argues for a Revving Up view, on which we are nearly ready to begin the hard species problem in earnest. To help work towards these conclusions, the paper begins with an outline of a new kind of view of species (Barker 2019a), which proposes they are feedback systems of a mathematically specifiable and empirically testable sort. (shrink)

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 (...) mean thus proves to be a predictively efficacious measure of fitness in examples featuring discrete generations and within- or between-generation variance in offspring output. Unfortunately, this advance has subsequently led some to conclude that the arithmetic mean is never a good measure of fitness and that the geometric mean should accordingly be the default measure of fitness. We show not only that the arithmetic mean is a perfectly reasonable measure of fitness so long as one is clear about what it refers to, but also that it functions as a more general measure when properly interpreted. It must suffice as a measure of fitness in any case where the geometric mean has been effectively deployed as a measure. We conclude with a discussion about why the mathematical equivalence we highlight cannot be dismissed as merely of mathematical interest. (shrink)

The levels of selection debate is generally taken to be a debate about how natural selection can occur at the various levels of biological organization. In this paper, we argue that questions about levels of selection should be analyzed separately from questions about levels of organization. In the deflationary proposal we defend, all that is necessary for multilevel selection is that there are cases in which particles are nested in collectives, and that both the collectives and the particles that compose (...) them each separately undergo natural selection. We argue that adopting this deflationary account helps to disentangle the levels of selection and the levels of organization, and thereby contributes to advancing the levels of selection debate. (shrink)

This essay offers some reflections on Samir Okasha’s new monograph Agents and Goals in Evolution, his style of doing philosophy, and the broader philosophy of nature project of trying to make sense of agency and rationality as natural phenomena.

Darwin provided us with a powerful theoretical framework to explain the evolution of living systems. Natural selection alone, however, has sometimes been seen as insufficient to explain the emergence of new levels of selection. The problem is one of “circularity” for evolutionary explanations: how to explain the origins of Darwinian properties without already invoking their presence at the level they emerge. That is, how does evolution by natural selection commence in the first place? Recent results in experimental evolution suggest a (...) way forward: Paul Rainey and his collaborators have shown that Darwinian properties could be exogenously imposed via what they call “ecological scaffolding.” This could solve the “black box” dilemma faced by Darwinian explanations of new levels of organization. Yet, despite “scaffolding” recently becoming a popular term in the study of cognition, culture, and evolution, the concept has suffered from vagueness and ambiguity. This article aims to show that scaffolding can be turned into a proper scientific concept able to do explanatory work within the context of the major evolutionary transitions. Doing so will allow us to once again extend the scope of the Darwinian model of evolution by natural selection. (shrink)

This MPhil dissertation presents a novel account of teleological explanations in biology. I outline the “shorthand approach” to such explanations, on which they are taken to convey implicit evolutionary explanations. “Selected effects” accounts of teleological explanation dominate recent literature, but they struggle to accommodate teleological explanations of complex traits built through cumulative selection. I articulate the general notion of a landscape explanation, which, applied to biology, explains the evolution of complex features in a population by citing salient features of the (...) population’s fitness landscape. I show that such explanations lend themselves to a teleological shorthand. I close with remarks concerning when a teleological explanation of a trait is legitimate, and why teleological language strikes us as appropriate when we give evolutionary explanations. (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)

Due to its high degree of complexity and its historical nature, evolutionary biology has been traditionally portrayed as a messy science. According to the supporters of such a view, evolutionary biology would be unable to formulate laws and robust theories, instead just delivering coherent narratives and local models. In this article, our aim is to challenge this view by showing how the Price equation can work as the core of a general theoretical framework for evolutionary phenomena. To support this claim, (...) we outline some unnoticed structural similarities between physical theories and evolutionary biology. More specifically, we shall argue that the Price equation, in the same way as fundamental formalisms in physics, can serve as a heuristic principle to formulate and systematise different theories and models in evolutionary biology. (shrink)

Life on Earth descends from a common ancestor. However, it is likely that there are other instances of life in the universe. If so, each abiogenesis event will have given rise to an independently originated life clade, of which Earth-life is an example. In this paper, I argue that the set of all IOLCs in the universe forms a Darwinian population subject to natural selection, with more widely dispersed IOLCs being less likely to face extinction. As a result, we should (...) expect that, over time, more planets will become inhabited by fewer, more successful IOLCs. (shrink)

The goal of this essay is to put forward an original theory of artifact function, which takes on board the results of the debate on the notion of biological function and also accommodates the distinctive aspects of artifacts. More precisely, the paper develops and defends the Dual-Aspect Theory, which is a monist account according to which an artifact’s function depends on intentional and reproductive aspects. It is argued that this approach meets a set of theoretical and meta-theoretical desiderata and is (...) superior to alternative views. (shrink)

Reichenbach’s early solution to the scientific problem of how abstract mathematical representations can successfully express real phenomena is rooted in his view of coordination. In this paper, I claim that a Reichenbach-inspired, ‘layered’ view of coordination provides us with an effective tool to systematically analyse some epistemic and conceptual intricacies resulting from a widespread theorising strategy in evolutionary biology, recently discussed by Okasha (2018) as ‘endogenization’. First, I argue that endogenization is a form of extension of natural selection theory that (...) comprises three stages: quasi-axiomatisation, functional extension, and semantic extension. Then, I argue that the functional extension of one core principle of natural selection theory, namely, the principle of heritability, requires the semantic extension of the concept of inheritance. This is because the semantic extension of ‘inheritance’ is necessary to establish a novel form of coordination between the principle of heritability and the extended domain of phenomena that it is supposed to represent. Finally, I suggest that – despite the current lack of consensus on the right semantic extension of ‘inheritance’ – we can fruitfully understand the reconceptualization of ‘inheritance’ provided by niche construction theorists as the result of a novel form of coordination. (shrink)

The vast literature on negative treatment of outgroups and favoritism toward ingroups provides many local insights but is largely fragmented, lacking an overarching framework that might provide a unified overview and guide conceptual integration. As a result, it remains unclear where different local perspectives conflict, how they may reinforce one another, and where they leave gaps in our knowledge of the phenomena. Our aim is to start constructing a framework to help remedy this situation. We first identify a few key (...) ideas for creating a theoretical roadmap for this complex territory, namely the principles of etiological functionalism and the dual inheritance theory of human evolution. We show how a “molecular” approach to emotions fits into this picture, and use it to illuminate emotions that shape intergroup relations. Finally, we weave the pieces together into the beginnings of a systematic taxonomy of the emotions involved in social interactions, both hostile and friendly. While it is but a start, we have developed the argument in a way that illustrates how the foundational principles of our proposed framework can be extended to accommodate further cases. (shrink)

The goal of this essay is to assess the Selected-Effects Etiological Theory of biological function, according to which a trait has a function F if and only if it has been selected for F. First, I argue that this approach should be understood as describing the paradigm case of functions, rather than as establishing necessary and sufficient conditions for function possession. I contend that, interpreted in this way, the selected-effects approach can explain two central properties of functions and can satisfactorily (...) address some recent counterexamples. This reading, however, shows that there is only a partial overlap between biological functions and selected effects, so the former cannot be reduced to the latter. Finally, I maintain that this result is still compatible with a naturalistic theory of function that appeals to some evolutionary process. (shrink)

The dominant view today on evolutionary progress is that it has been thoroughly debunked. Even value-neutral progress concepts are seen to lack important theoretical underpinnings: natural selection provides no rationale for progress, and natural selection need not even be invoked to explain large-scale evolutionary trends. In this paper I challenge this view by analysing how natural selection acts in heterogeneous environments. This not only undermines key debunking arguments, but also provides a selectionist rationale for a pattern of “evolutionary unfolding”, where (...) life radiates across an increased range of exploitation of environmental heterogeneity. (shrink)

While the struggle for life played an important role in the process of natural selection as it was conceived by Darwin, natural selection is commonly characterized today as a process which does not necessarily involve struggle. Nevertheless, there have been some attempts to show the importance of struggle to the process of natural selection. The present paper aims to continue these attempts and clarify the precise evolutionary role of struggle. The paper focuses on a recent dispute regarding the role of (...) struggle between Peter Godfrey-Smith and Tim Lewens. Godfrey-Smith argues that struggle makes populations into "paradigm cases of Darwinian populations", meaning that struggle leads natural selection to produce adaptations. Lewens, in contrast, argues that Godfrey-Smith's argument fails to show that struggle plays such a role. I argue that Godfrey-Smith's argument that struggle leads selection to produce adaptations can be understood in two different ways. Lewens interprets Godfrey-Smith's argument in one of these ways, and rightly argues that thus understood, this argument fails to show that struggle leads selection to produce adaptations. However, I show that understood in a second way, Godfrey-Smith's argument does show that struggle plays such a role. Still, I show that there is an additional problem with Godfrey-Smith's argument that struggle makes populations into "paradigm cases of Darwinian populations". It does not capture the precise role played by struggle in the relation between selection and adaptation. Thus, the present paper clarifies the unique role that struggle plays in the relation between selection and adaptation in evolutionary theory. (shrink)

An understanding of the factors behind the evolution of multicellularity is one of today’s frontiers in evolutionary biology. This is because multicellular organisms are made of one subset of cells with the capacity to transmit genes to the next generation and another subset responsible for maintaining the functionality of the organism, but incapable of transmitting genes to the next generation. The question arises: why do somatic cells sacrifice their lives for the sake of germline cells? How is germ/soma separation maintained? (...) One conventional answer refers to inclusive fitness theory, according to which somatic cells sacrifice themselves altruistically, because in so doing they enhance the transmission of their genes by virtue of their genetic relatedness to germline cells. In the present article we will argue that this explanation ignores the key role of policing mechanisms in maintaining the germ/soma divide. Based on the pervasiveness of the latter, we argue that the role of altruistic mechanisms in the evolution of multicellularity is limited and that our understanding of this evolution must be enriched through the consideration of coercion mechanisms. (shrink)

There is a certain metaphor that has enjoyed tremendous longevity in the evolution of ageing literature. According to this metaphor, nature has a certain goal or purpose, the perpetuation of the species, or, alternatively, the reproductive success of the individual. In relation to this goal, the individual organism has a function, job, or task, namely, to breed and, in some species, to raise its brood to maturity. On this picture, those who cannot, or can no longer, reproduce are somehow invisible (...) to, or even dispensable to, the evolutionary process. Here, I argue that the metaphor should be discarded, not on the grounds that it is a metaphor, but on the grounds that this particular metaphor distorts our understanding of the evolution of ageing. One reason the metaphor is problematic is that it frames senescence and death as nature’s verdict on the value of older individuals. Instead, we should explore a different metaphor: the lengthy post-reproductive period in humans and some other animals is not an accident of culture, but designed by nature for the purpose of supporting and guiding younger generations. On this alternate picture, different stages of life have their own evolutionary rationales, their distinctive design features, their special mandates. (shrink)

Biological individuality is a notoriously thorny topic for biologists and philosophers of biology. In this paper we argue that biological individuality presents multiple, interconnected questions for biologists and philosophers that together form a problem agenda. Using a case study of an interdisciplinary research group in ecology, behavioral and evolutionary biology, we claim that a debate on biological individuality that seeks to account for diverse practices in the biological sciences should be broadened to include and give prominence to questions about uniqueness (...) and temporality. We show that broadening the problem agenda of biological individuality draws attention to underrecognized philosophical issues and discussions and thereby organizes and enriches the existing debate. (shrink)

The received view holds that there is a significant divide between full-blown representational states and so called ‘detectors’, which are mechanisms set off by specific stimuli that trigger a particular effect. The main goal of this paper is to defend the idea that many detectors are genuine representations, a view that I call ‘Strong Liberal Representationalism’. More precisely, I argue that ascribing semantic properties to them contributes to an explanation of behavior, guides research in useful ways and can accommodate misrepresentation.

High-fidelity cultural transmission, rather than brute intelligence, is the secret of our species’ success, or so many cultural evolutionists claim. It has been selected because it ensures the spread, stability and longevity of beneficial cultural traditions, and it supports cumulative cultural change. To play these roles, however, fidelity must be a causally-efficient property of cultural transmission. This is where the grain problem comes in and challenges the explanatory potency of fidelity. Assessing the degree of fidelity of any episode or mechanism (...) of cultural transmission always depends upon an investigator’s choice of grain of description at which cultural traditions are being studied. The fidelity of cultural transmission then appears to be relative to the granularity at which one approaches cultural variation, and since there is a multiplicity of grains of description by which the same tradition can be studied, there results a multiplicity of measures of fidelity for a same event or mechanism of cultural transmission. If this is correct, because fidelity is always relative to the grain of description dictated by the local and specific research interests of the investigator, then there seems to be no fact of the matter as to whether cultural transmission is faithful or not, independently from a researcher’s framework of analysis. The aims of this paper are to offer a conceptual clarification of the grain problem in cultural evolution, to assess its causes, to unpack its epistemological implications, and to examine its reach and consequences for a science of cultural evolution. (shrink)

Life is a self-maintaining process based on metabolism. Something is said to be alive when it exhibits organization and is actively involved in its own continued existence through carrying out metabolic processes. A life is a spatio-temporally restricted event, which continues while the life processes are occurring in a particular chunk of matter (or, arguably, when they are temporally suspended, but can be restarted at any moment), even though there is continuous replacement of parts. Life is organized in discrete packages, (...) particular cells and multicellular organisms with differing degrees of individuality. Biological species, too, have been shown to be individuals, and not classes, as these collections of organisms are spatio-temporally localized, restricted, continuous, and somewhat cohesive entities, with a definite beginning and end. Assuming that all life on Earth has a common origin, all living organisms, cells, and tissues descending from this origin exhibit continuity of the life processes at the cellular level, as well as many of the features that define the individual character of species: spatio-temporal localization and restriction, continuity, historicity, and cohesiveness. Therefore, life on Earth is an ontological individual. Independent origins of life will have produced other such individuals. These provisionally called ‘life-individuals’ constitute a category of organization of life which has seldom been recognized. The discovery of at least one independent life-individual would go a long way toward the project of the universality of biology. (shrink)

Contrary to the common-sense view and positivist aspirations, scientific concepts are often imprecise. Many of these concepts are ambiguous, vague, or have an under-specified meaning. In this paper, I discuss how imprecise concepts promote integration in biology and thus benefit science. Previous discussions of this issue focus on the concepts of molecular gene and evolutionary novelty. The concept of molecular gene helps biologists integrate explanatory practices, while the notion of evolutionary novelty helps them integrate research questions into an interdisciplinary problem (...) New directions in the philosophy of science, Springer, Dordrecht, 2014). In what follows, I compare molecular gene and evolutionary novelty to another imprecise concept, namely biological lineage. This concept promotes two other types of scientific integration: it helps biologists integrate theoretical principles and methodologies into different areas of biology. The concept of biological lineage facilitates these types of integration because it is broad and under-specified in ways that the concepts of molecular gene and evolutionary novelty are not. Hence, I use the concept of biological lineage as a case study to reveal types of integration that have been overlooked by philosophers. This case study also shows that even very imprecise concepts can be beneficial to scientific practice. (shrink)

Usually evolutionary economists equate evolutionary theory with modern Darwinism. However the rise of evolutionary developmental biology (Evo-Devo) puts into question the monopoly of Darwinism in evolutionary biology. The major divergences between the two paradigms in evolutionary biology are drawn in the analysis of three trade-offs: population vs. typological thinking, creative role of natural selection vs. internal (inherent) change, and microevolution vs. macroevolution. It is argued here that the Evo-Devo breakthrough helps to better understand the limits to Darwinism in the social (...) realm and design the contours of an alternative paradigm in evolutionary economics privileging structural macroevolution and what Schumpeter called “change from within”. (shrink)

Philosophy of immunology is a subfield of philosophy of biology dealing with ontological and epistemological issues related to the studies of the immune system. While speculative investigations and abstract analyses have always been part of immune theorizing, until recently philosophers have largely ignored immunology. Yet the implications for understanding the philosophical basis of organismal functions framed by immunity offer new perspectives on fundamental questions of biology and medicine. Developed in the context of history of medicine, theoretical biology, and medical anthropology, (...) philosophy of immunology differs from these related branches of study in its focus on traditional philosophical questions concerning identity, individuality, ecology, cognition, scientific methodology and theory construction. This broad agenda derives from immunology’s multifaceted research program that has developed from its initial clinical challenges of host defense, transplantation, autoimmunity, tumor immunology, and allergy. In addition to these well-established research areas, immunity is now understood to play a central role in other physiological functions, development, ecology, and evolutionary mechanics. Holding together these diverse domains of inquiry lie philosophical commitments oriented by organismal identity. In this regard, pertinent issues are raised concerning cognition (organization of immune perception and information processing), the character of individuality (framed by the ecological context of immune-mediated assimilation and rejection), and the dynamics of complex systems (understood as holistic systems biology). Indeed, immunology, in the context of cognitive science, evolutionary biology, environmental sciences, and development provides multi-focal perspectives for philosophy of science. (shrink)

This review essay reflects on recent discussions in evolutionary biology and philosophy of science on the central causes of evolution and the structure of causal explanations in evolutionary theory. In this debate, it has been argued that our view of evolutionary causation should be rethought by including more seriously developmental causes and causes of the individual acting organism. I use Tobias Uller’s and Kevin Laland’s volume Evolutionary Causation as well as recent reviews of it as a starting point to reflect (...) on the causal role of agency, individuality, and the environment in evolution. In addition, I critically discuss classical philosophical frameworks of theory change used in this debate to understand changing views of evolutionary causation. (shrink)

Although the definition of ‘signal’ has been controversial for some time within the life sciences, current approaches seem to be converging toward a common analysis. This powerful framework can satisfactorily accommodate many cases of signaling and captures some of its main features. This paper argues, however, that there is a central feature of signals that so far has been largely overlooked: its special causal role. More precisely, I argue that a distinctive feature of signals is that they are minimal causes. (...) I explain this notion, suggest some strategies for identifying its instances and defend its relevance by means of conceptual and empirical considerations. (shrink)