This paper uses formal Darwinism as elaborated by Alan Grafen to articulate an explanatory pluralism that casts light upon two strands of controversies running across evolutionary biology, viz., the place of organisms versus genes, and the role of adaptation. Formal Darwinism shows that natural selection can be viewed either physics-style, as a dynamics of alleles, or in the style of economics as an optimizing process. After presenting such pluralism, I argue first that whereas population genetics does not support optimization, optimality (...) can still be taken as a default hypothesis when modeling evolutionary processes; and second, that organisms have an explanatory role in evolutionary theory, since they are involved in the economic perspective of optimization. Finally, in order to ask whether the Modern Synthesis can indeed provide a theory of organisms, I apply a Kantian-inspired theoretical view of organisms (underlying much developmental modeling), according to which they are both designed entities and subjects of intrinsic circular processes involving the whole organism and its parts. I first show that the design aspect is accountable for in terms of the Modern Synthesis understood in the formal Darwinism framework. I then question whether the latter aspect of organisms can also be ultimately captured in the same framework, and to this purpose devise an empirical test relying on an assessment of the relative weight of genetic elements in developmental and functional gene regulatory networks. (shrink)

Besides mechanistic explanations of phenomena, which have been seriously investigated in the last decade, biology and ecology also include explanations that pinpoint specific mathematical properties as explanatory of the explanandum under focus. Among these structural explanations, one finds topological explanations, and recent science pervasively relies on them. This reliance is especially due to the necessity to model large sets of data with no practical possibility to track the proper activities of all the numerous entities. The paper first defines topological explanations (...) and then explains why topological explanations and mechanisms are different in principle. Then it shows that they are pervasive both in the study of networks—whose importance has been increasingly acknowledged at each level of the biological hierarchy—and in contexts where the notion of selective neutrality is crucial; this allows me to capture the difference between mechanisms and topological explanations in terms of practical modelling practices. The rest of the paper investigates how in practice mechanisms and topologies are combined. They may be articulated in theoretical structures and explanatory strategies, first through a relation of constraint, second in interlevel theories, or they may condition each other. Finally, I explore how a particular model can integrate mechanistic informations, by focusing on the recent practice of merging networks in ecology and its consequences upon multiscale modelling. (shrink)

Although it may seem like a truism to assert that biology is the science that studies organisms, during the second half of the twentieth century the organism category disappeared from biological theory. Over the past decade, however, biology has begun to witness the return of the organism as a fundamental explanatory concept. There are three major causes: (a) the realization that the Modern Synthesis does not provide a fully satisfactory understanding of evolution; (b) the growing awareness of the limits of (...) reductionism in molecular biology; and (c) the renewed interest in the nature of life as a genuine scientific problem. This essay examines these recent developments and considers the new epistemological roles being played by the organism in each of them. It also reflects on what the present resurgence of the organism means for the philosophy of biology. (shrink)

The category of ‘organism’ has an ambiguous status: is it scientific or is it philosophical? Or, if one looks at it from within the relatively recent field or sub-field of philosophy of biology, is it a central, or at least legitimate category therein, or should it be dispensed with? In any case, it has long served as a kind of scientific “bolstering” for a philosophical train of argument which seeks to refute the “mechanistic” or “reductionist” trend, which has been perceived (...) as dominant since the 17th century, whether in the case of Stahlian animism, Leibnizian monadology, the neo-vitalism of Hans Driesch, or, lastly, of the “phenomenology of organic life” in the 20th century, with authors such as Kurt Goldstein, Maurice Merleau-Ponty, and Georges Canguilhem. In this paper I try to reconstruct some of the main interpretive ‘stages’ or ‘layers’ of the concept of organism in order to critically evaluate it. How might ‘organism’ be a useful concept if one rules out the excesses of ‘organismic’ biology and metaphysics? Varieties of instrumentalism and what I call the ‘projective’ concept of organism are appealing, but perhaps ultimately unsatisfying. (shrink)

Well prior to the invention of the term ‘biology’ in the early 1800s by Lamarck and Treviranus, and also prior to the appearance of terms such as ‘organism’ under the pen of Leibniz in the early 1700s, the question of ‘Life’, that is, the status of living organisms within the broader physico-mechanical universe, agitated different corners of the European intellectual scene. From modern Epicureanism to medical Newtonianism, from Stahlian animism to the discourse on the ‘animal economy’ in vitalist medicine, models (...) of living being were constructed in opposition to ‘merely anatomical’, structural, mechanical models. It is therefore curious to turn to the ‘passion play’ of the Scientific Revolution – whether in its early, canonical definitions or its more recent, hybridized, reconstructed and expanded versions: from Koyré to Biagioli, from Merton to Shapin – and find there a conspicuous absence of worry over what status to grant living beings in a newly physicalized universe. Neither Harvey, nor Boyle, nor Locke (to name some likely candidates, the latter having studied with Willis and collaborated with Sydenham) ever ask what makes organisms unique, or conversely, what does not. In this paper I seek to establish how ‘Life’ became a source of contention in early modern thought, and how the Scientific Revolution missed the controversy. (shrink)

Explanatory Externalism states that the only adaptive force in evolution is natural selection. Explanatory Externalism is a central thesis of the Modern Evolutionary Synthesis. The etiological theory of natural selected-effect functions also advocates Explanatory Externalism. According to this theory, natural selection is the process responsible for determining the proper natural functions of traits. However, I will point out several challenges to Explanatory Externalism that are proposed primarily by developmental biology and its various subfields. Based on these challenges, this paper will (...) argue why biological functions cannot be fully explained by the selected-effect theory, i.e., that a theory of biological functions must adopt some kind of Explanatory Internalism. At the end of this paper, I will discuss whether or not Explanatory Internalism entails a pluralistic view of biological functions, and how this discussion is related to the different explanations of fit and diversity found in evolutionary biology. (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)

Abstract: The use of evolutionary explanations to account for proximate phenomena has been labeled by various authors as an explanatory error, the so-called phylogeny fallacy. In this paper, this fallacy will be analyzed in the context of teleosemantics. I will discuss whether teleosemantics projects that rely on the Selected-Effect Theory of Functions (i.e., mainstream teleosemantics) generally commit the fallacy. To frame the discussion, I will present two desiderata that, as argued here, every teleosemantic project must fulfill. The actuality desideratum, motivated (...) by Brentano’s problem, demands for a proximate analysis of the properties that make a system intentional. The historical desideratum, on the other hand, says that a teleological view of intentionality requires a historical dimension in order to understand the causal basis of teleological explanations. While I give some reasons why mainstream teleosemantics might be implicated in the phylogeny fallacy, my main argument aims to highlight an explanatory gap in mainstream teleosemantics: the population perspective on teleofunctions is inadequate to explain the proximate properties of intentional systems. In other words, the account of the historical desideratum in population-evolutionary terms prevents an adequate solution to the actuality desideratum. This is related to two central goals of teleosemantics: The goal of explaining content determination by recourse to natural selection processes makes it impossible to solve another goal of teleosemantics, the so-called problem of representational status. I analyze this situation and argue that inherited information (sometimes interpreted in a teleosemantic sense) plays the role of closing the gap. This leads teleosemantics into a complex situation: in order to close the gap, it must commit the fallacy. At the end of the article, I propose an alternative to fulfill both desiderata, motivated by an evo-devo perspective. I state that lineage explanations have the property of providing historical explanations based on proximate causes. (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)

We characterize Hegel’s stance on biological purposiveness as consisting in a twofold move, which conceives organisms as intrinsically purposive natural systems and focuses on their behavioral and cognitive abilities. We submit that a Hegelian stance is at play in enactivism, the branch of the contemporary theory of biological autonomy devoted to the study of cognition and the mind. What is at stake in the Hegelian stance is the elaboration of a naturalized, although non-reductive, understanding of natural purposiveness.

This paper investigates Hegel’s account of the animal organism as it is presented in the Philosophy of Nature, with a special focus on its normative implications. I argue that the notion of “organisation” is fundamental to Hegel’s theory of animal normativity. The paper starts by showing how a Hegelian approach takes up the scientific image of organism and assigns a basic explanatory role to the notion of “organisation” in its understanding living beings. Moving from this premise, the paper turns to (...) the group of accounts in contemporary theoretical biology known as “organisational accounts”, which offer a widely debated strategy for naturalizing teleology and normativity in organisms. As recent scholarship recognizes, these accounts explicitly rely on insights from Kant and Post-Kantianism. I make the historical and conceptual argument that Hegel’s view of the organism shares several basic commitments with OAs, especially regarding the notion of “organisational closure”. I assess the account of normativity that such accounts advance and its implications for how we approach Hegel. Finally, I argue that the notion of “organisation” is more fundamental to Hegel’s theory of animal normativity than the Aristotelian notion of “Gattung” or “species”, which by contrast appears derivative – at least in the Philosophy of Nature and the Lectures – and does not play the central role in his account maintained by some scholars. (shrink)

In this paper, we analyze the debate between the Modern Synthesis and the Extended Evolutionary Synthesis in light of the concept of incommensurability developed by Thomas Kuhn. In order to do so, first we briefly present both the Modern Synthesis and the Extended Evolutionary Synthesis. Then, we clarify the meaning and interpretations of incommensurability throughout Kuhn’s works, concluding that the version of this concept deployed in The Structure of Scientific Revolutions is the best suited to the analysis of scientific disputes. (...) After discussing incommensurability in Kuhn’s works, we address the question of whether the Modern Synthesis and the Extended Evolutionary Synthesis can be considered semantically, methodologically, and ontologically incommensurable, concluding that they can. Finally, we discuss three problems that arise from such a conclusion: firstly, what are the consequences of incommensurability; secondly, which mode of scientific change better explains this current dispute in evolutionary biology; and thirdly, whether rational theory comparison is possible given incommensurability. We suggest that the main consequence of incommensurability is profound disagreement, that the kind of scientific change that better explains the current dispute between the MS and the EES may be scientific specialization, and that incommensurability does not preclude rational theory comparison. (shrink)

Both biosemiotics and evolutionary epistemology are concerned with how knowledge evolves. (Applied) Evolutionary Epistemology thereby focuses on identifying the units, levels, and mechanisms or processes that underlie the evolutionary development of knowing and knowledge, while biosemiotics places emphasis on the study of how signs underlie the development of meaning. We compare the two schools of thought and analyze how in delineating their research program, biosemiotics runs into several problems that are overcome by evolutionary epistemologists. For one, by emphasizing signs, biosemiotics (...) needs to delineate a semiotic threshold, which is a problem not encountered by evolutionary epistemologists. Instead, the latter recognizes that all organisms are knowers that evolve knowledge, which they recognize to extend toward phenomena produced by organisms such as behavior, cognition, language, culture, science, and technology. Secondly, biosemiotics attempts at continuing adaptationist notions on how organisms relate to their environment, while especially Applied Evolutionary Epistemology comes to redefine the nature of the organism–environment relationship in such a way that it recognizes the spatiotemporal boundedness of existence, which in turn makes adaptationist accounts obsolete. (shrink)

According to neo-Aristotelian ethical naturalism, moral goodness is an instance of natural goodness, a kind of normativity supposedly already present in nature in the biological realm of non-human living things. Proponents of this view appeal to Michael Thompson’s conception of a life-form--the form of a living organism--to give an account of natural goodness. However, although neo-Aristotelians call themselves naturalists, they hardly ever consult the science of biology to defend their commitments regarding biological organisms. This has led many critics to argue (...) that the neo-Aristotelian account of natural normativity is out of touch with the findings of modern evolutionary biology. One line of response to this objection, presented by John Hacker-Wright and Micah Lott, claims that the neo-Aristotelian concept of a living organism has to be presupposed in evolutionary biology as long as organisms are the subjects of evolutionary explanation. In this paper, I examine this response by tracing the concept of organism in modern evolutionary biology. I first argue that the Modern Synthesis theory of evolution, which understands evolution as change in gene frequencies within a population, does not presuppose the relevant concept of organism. I then explore an alternative view of evolution that has emerged in the past twenty years from advances in evolutionary developmental biology. I argue that this so called ‘evo-devo’ approach makes room for an explanatory concept of organism that can be reconciled with the neo-Aristotelian view. Moreover, I argue that although the explanatory role of the concept of organism in evolutionary biology is still contentious, the well-established role of this concept in developmental biology can be used to defend the biological commitments of neo-Aristotelian naturalism. (shrink)

In the theory and history of ecology, Frederic Clements’s theory of plant communities is usually presented as the historical prototype and a paradigmatic example of synecological organicism, characterised by the assumption that ecological communities are functionally integrated units of mutually dependent species. In this paper, I will object to this standard interpretation of Clements’s theory. Undoubtedly, Clements compares plant communities with organisms and calls them “complex organisms” and “superorganisms”. Further, he can indeed be regarded as a proponent of ecological organicism—provided (...) that one defines ecological organicism as the interpretation of synecological units according to the model of the individual organism. However, Clements’s theory does not include the assumption that mutual dependence is a principle of the organisation of plant communities. Rather, he interprets plant communities as top-down control-hierarchical entities, in which subordinate species depend on dominant species—but not the other way around. Therefore, his theory represents what may be called ‘control-hierarchical organicism’ as against ‘mutualistic organicism’. The erroneous attribution to Clements of ‘mutualistic organicism’ might be due to an unawareness of the existence of different concepts of the organism. This unawareness results in the projection on Clements’s theory of a seemingly self-evident mutualistic concept of organism that Clements himself did not use as a basis for his theory of plant communities. (shrink)

Given the central place organisms occupy in Kant’s account of living nature, it might seem unlikely that his claims about biological wholes could be relevant to current debates over the problem of biological individuality. These debates acknowledge the multiple realizability of biological individuality in vastly different forms, including parts of organisms and complex groups of organisms at various levels of the biological hierarchy, sparking much controversy in attempts to characterize a biological individual. I argue that, far from being irrelevant to (...) this controversy, Kant’s account provides a key insight for addressing the multiple realizability problem. I show how the reciprocal causality between a self-organizing whole and its parts, which Kant thinks characterizes a natural end, is not limited to organisms but is exhibited by numerous types of beings in living nature. Self-organizing wholes of various kinds, and at various biological levels, may count as biological individuals, depending on the degree to which their functionally integrated parts are represented by reflective judgement as a natural end. (shrink)

We outline an alternative to both scientific and liberal naturalism which attempts to reconcile Sellars’ apparently conflicting commitments to the scientific accountability of human nature and the autonomy of the space of reasons. Scientific naturalism holds that agency and associated concepts are a mechanical product of the realm of laws, while liberal naturalism contends that the autonomy of the space of reason requires that we leave nature behind. The third way we present follows in the footsteps of German Idealism, which (...) attempted to overcome the Kantian chasm between nature and agency, and is thus dubbed ‘post-Kantian.’ We point to an overlooked group of scholars in the naturalism debate who, along with recent work in biology and cognitive science, offer a path to overcome the reductive tendencies of empiricism while avoiding the dichotomy of logical spaces. We then bring together these different streams of research, by foregrounding and expanding on what they share: the idea of organisms as living agents and that of a continuity without identity between life and mind. We qualify this as a bottom-up transformative approach to rational agency, which grounds cognition in the intrinsically purposive nature of organisms, while emphasizing the distinction between biological agency and full-fledged mindedness. (shrink)

Canguilhem criticized the concept of “public health”: health and disease are concepts that only apply to individuals, taken as organic totalities. Their extension to a different level of organization is purely metaphorical. The importance assumed by epidemiology in the construction of our knowledge of the normal and the pathological does, however, call for reflection on the role and the status of the population level of organization in our approach to health phenomena. The entanglement of the biological and the social in (...) human life and in contemporary societies justifies this level of analysis for better understanding the complexity and the interaction of health determinants both at the level of individuals and their interactions and at that of the population. But is this population level just a useful level of analysis that makes it possible to bring to light the social determinants of health at the individual level, or does it rest instead on characteristics of the population that are irreducible to individual characteristics, but which are nevertheless important for understanding and taking action with respect to both population and individual health? Defending this second alternative, I show how the epidemiological point of view, and in particular that of social epidemiology, leads us to rethink the possibility of a concept of “population health” that is not reducible to the sum of individual instances of health. (shrink)

This paper uses the framework of Formal Darwinism (FD) to evaluate organism-centric critiques of the Modern Synthesis (MS). The first section argues that the FD project reconciles two kinds of selective explanations in biology. Thus it is not correct to say that the MS neglects organisms—instead, it explains organisms’ design, as argued in the second section. In the third section I employ a concept of the organism derived from Kant that has two aspects: the parts presupposing the whole, and the (...) productivity of the parts in relation to the whole. The first aspect corresponds to the “design” that FD explains, whereas the second aspect is something about which the MS is largely silent. (shrink)

The relationship between statistical mechanics and population genetics has a long history. Both take advantage of statistics to address the behavior of large groups of entities. The main objective of this article is to assess the obstacles population genetics is meeting in its claim to explain biological phenomena from the conceptual apparatus of statistical mechanics according to two recent articles. Several tools available to the latter are missing in the former. Thus, in the absence of an adequate justification of the (...) relevance of the analogy and without the satisfaction of certain basic assumptions, I argue that the explanatory strategy of the analogously based model is seriously compromised. La relation entre mécanique statistique et génétique des populations a une longue histoire. À l’instar de la première, la seconde tire profit des statistiques afin de traiter le comportement des grands ensembles d’entités. L’objectif principal du présent article est d’identifier les obstacles que rencontre la génétique des populations dans sa prétention d’explication des phénomènes biologiques à partir de l’appareillage conceptuel de la mécanique statistique selon deux articles récents. Or, plusieurs outils dont dispose cette dernière manque à la première. Ainsi, en l’absence d’une justification adéquate de la pertinence de l’analogie et sans la satisfaction de certaines hypothèses fondamentales, je soutiens que la stratégie explicative du modèle analogue est sérieusement compromise. (shrink)

Arguments for an extended evolutionary synthesis often center on the concept of “reciprocal causation.” Proponents argue that reciprocal causation is superior to standard models of evolutionary causation for at least two reasons. First, it leads to better scientific models with more predictive power. Second, it more accurately represents the causal structure of the biological world. Simply put, proponents of an extended evolutionary synthesis argue that reciprocal causation is empirically and explanatorily apt relative to competing causal frameworks. In this paper, I (...) present quantitative survey data from faculty members in biology departments at universities across the United States to evaluate this claim. The survey data indicate that a majority of the participants do not agree (i.e., most either disagree or neither agree nor disagree) that the concept of reciprocal causation confers a larger advantage on research practices. However, a majority of the participants agree that the causal framework of the extended evolutionary synthesis more accurately represents the structure of the biological world. These results demonstrate that the explanatory merits of a conceptual framework and its practical utility can come apart in interesting and informative ways. (shrink)

This paper questions the form and prospects of “extended theories” which have been simultaneously and independently advocated both in the philosophy of mind and in the philosophy of biology. It focuses on Extend Mind Theory (EMT) and Developmental Systems Theory (DST). It shows first that the two theories vindicate a parallel extension of received views, the former concerning extending cognition beyond the brain, the latter concerned with extending evolution and development beyond the genes. It also shows that both arguments rely (...) on the demonstration of causal parities, which have been undermined by the classical received view. Then I question whether the argument that there is an illegitimate inference from parities or coupling to constitution claims, which has been objected by Adams and Aizawa in The bounds of cognition, (2008) to EMT, also holds against DST. To this aim, I consider two defenses against DST that are parallel to two defenses against EMT, one about intrinsic content, the other about the difference between what’s in principle possible and what happens in practice. I conclude by claiming that the weaknesses and strengths of both theories are different regarding these two kinds of objections. (shrink)