Ernst Mayr’s classical work on the nature of causation in biology has had a huge influence on biologists as well as philosophers. Although his distinction between proximate and ultimate causation recently came under criticism from those who emphasize the role of development in evolutionary processes, the formal relationship between these two notions remains elusive. Using causal graph theory, this paper offers a unified framework to systematically translate a given “proximate” causal structure into an “ultimate” evolutionary response, and illustrates evolutionary implications (...) of various kinds of causal mechanisms including epigenetic inheritance, maternal effects, and niche construction. These results not only reveal the essential interplay between proximate and ultimate causation in the study of evolution, but also provide a formal method to evaluate or discover non-standard or yet unknown evolutionary phenomena. (shrink)

In this article we argue that the classical—linear and bottom-up directed—models of causation in biology, and the ‘‘proximate/ultimate’’ dichotomy, are inappropriate to capture the complexity inherent to biological processes. We introduce a new notion of ‘‘multilevel causation’’ where old dichotomies such as proximate/ultimate and bottom-up/ top-down are reinterpreted within a multilevel, web-like, approach. In briefly reviewing some recent work on complexity, EvoDevo, carcinogenesis, autocatalysis, comparative genomics, animal regeneration, phenotypic plasticity, and niche construction, we will argue that such reinterpretation is a (...) necessary step for the advancement of the ‘‘Extended Synthesis.’’. (shrink)

Niche construction theory (NCT) is distinctive for being explicit in recognizing environmental modification by organisms—niche construction—and its legacy—ecological inheritance—to be evolutionary processes in their own right. Humans are widely regarded as champion niche constructors, largely as a direct result of our capacity for the cultural transmission of knowledge and its expression in human behavior, engineering, and technology. This raises the question of how human ecological inheritance relates to human cultural inheritance. If NCT is to provide a conceptual framework for the (...) human sciences, then it is important that the relationship between these two legacies is clear. We suggest that cultural processes and cultural inheritance can be viewed as the primary means by which humans engage in the universal process of niche construction. (shrink)

Using the context of controversies surrounding evolutionary developmental biology (EvoDevo) and the possibility of an Extended Evolutionary Synthesis, I provide an account of theory structure as idealized theory presentations that are always incomplete (partial) and shaped by their conceptual content (material rather than formal organization). These two characteristics are salient because the goals that organize and regulate scientific practice, including the activity of using a theory, are heterogeneous. This means that the same theory can be structured differently, in part because (...) theory presentations (as idealizations) intentionally depart from different features known to be present in a theory. Since there are diverse and potentially incompatible theory structures derived from heterogeneous goals found in scientific practices, a question arises about the absence of a unifying theory structure in the background. The notion of a “theory façade” offers a fruitful perspective on this potentially unsettling result. (shrink)

The essentialism story is a version of the history of biological classification that was fabricated between 1953 and 1968 by Ernst Mayr, who combined contributions from Arthur Cain and David Hull with his own grudge against Plato. It portrays pre-Darwinian taxonomists as caught in the grip of an ancient philosophy called essentialism, from which they were not released until Charles Darwin's 1859 Origin of Species. Mayr's motive was to promote the Modern Synthesis in opposition to the typology of idealist morphologists; (...) demonizing Plato served this end. Arthur Cain's picture of Linnaeus as a follower of 'Aristotelian' (scholastic) logic was woven into the story, along with David Hull's application of Karl Popper's term, 'essentialism', which Mayr accepted in 1968 as a synonym for what he had called 'typological thinking'. Although Mayr also pointed out the importance of empiricism in the history of taxonomy, the essentialism story still dominates the secondary literature. The history of the first telling of the essentialism story exposes its scant basis in fact. (shrink)

Like Laland et al., I think Mayr’s distinction is problematic, but I identify a further problem with it. I argue that Mayr’s distinction is a false dichotomy, and obscures an important question about evolutionary change. I show how this question, once revealed, sheds light on some debates in evo-devo that Laland et al.’s analysis cannot, and suggest that it provides a different view about how future integration between biological disciplines might proceed.

In 1961, Ernst Mayr published a highly influential article on the nature of causation in biology, in which he distinguished between proximate and ultimate causes. Mayr argued that proximate causes (e.g. physiological factors) and ultimate causes (e.g. natural selection) addressed distinct ‘how’ and ‘why’ questions and were not competing alternatives. That distinction retains explanatory value today. However, the adoption of Mayr’s heuristic led to the widespread belief that ontogenetic processes are irrelevant to evolutionary questions, a belief that has (1) hindered (...) progress within evolutionary biology, (2) forged divisions between evolutionary biology and adjacent disciplines and (3) obstructed several contemporary debates in biology. Here we expand on our earlier (Laland et al. in Science 334:1512–1516, 2011) argument that Mayr’s dichotomous formulation has now run its useful course, and that evolutionary biology would be better served by a concept of reciprocal causation, in which causation is perceived to cycle through biological systems recursively. We further suggest that a newer evolutionary synthesis is unlikely to emerge without this change in thinking about causation. (shrink)

This essay is concerned with concentrations of entities, which play an important—albeit often overlooked—role in scientific explanation. First, I discuss an example from molecular biology to show that concentrations can play an irreducible causal role. Second, I provide a preliminary philosophical analysis of this causal role, suggesting some implications for extant theories of causation. I conclude by introducing the concept of causation by concentration, a form of statistical causation whose widespread presence throughout the sciences has been unduly neglected and which (...) deserves to be studied in more depth. 1 Introduction2 Solving Lillie's Paradox: Lysogenic Induction in Phage λ3 Repressor Concentration and the Tuning of the Switch4 Concentration and Causality5 Preemption in Concentrations: Analysis and Implications6 Causation by Concentration: General Definition, Refinements, and Further Applications. (shrink)

In this paper, using a multilevel approach, we defend the positive role of natural selection in the generation of organismal form. Despite the currently widespread opinion that natural selection only plays a negative role in the evolution of form, we argue, in contrast, that the Darwinian factor is a crucial (but not exclusive) factor in morphological organization. Analyzing some classic arguments, we propose incorporating the notion of ‘downward causation’ into the concept of ‘natural selection.’ In our opinion, this kind of (...) causation is fundamental to the operation of selection as a creative evolutionary process. (shrink)

Griffiths and Machery (2008) argue that innateness is a ?folk biological? notion, which, as such, has no useful reconstruction in contemporary biology. If this is so, not only is it wrong to identify the vernacular notion with the precise theoretical concept of canalization, but worse, it would appear that many of the putative scientific claims for particular competences and capacities being innate are simply misplaced. The present paper challenges the core substantive claim of Griffiths and Machery's position, namely, that innateness (...) understood on canalization lines as environment-independent development (somehow and to some degree) is a confused, outmoded notion. It will be contended that the modality-independence of language offers a prima facie case against Griffiths and Machery's general position. (shrink)

This article reviews the recent reissuing of Richard Owen’s On the Nature of Limbs and its three novel, introductory essays. These essays make Owen’s 1849 text very accessible by discussing the historical context of his work and explaining how Owen’s ideas relate to his larger intellectual framework. In addition to the ways in which the essays point to Owen’s relevance for contemporary biology, I discuss how Owen’s unity of type theory and his homology claims about fins and limbs compare with (...) modern views. While the phenomena studied by Owen are nowadays of major interest to evolutionary developmental biology, research in evo-devo has largely shifted from homology (which was Owen’s concern) towards evolutionary novelty, e.g., accounting for fins as a novelty. Still, I argue that questions about homology are important and raise challenges even for explanations of novelty. (shrink)

Among many properties distinguishing emergence, such as novelty, irreducibility and unpredictability, computational accounts of emergence in terms of computational incompressibility aim first at making sense of such unpredictability. Those accounts prove to be more objective than usual accounts in terms of levels of mereology, which often face objections of being too epistemic. The present paper defends computational accounts against some objections, and develops what such notions bring to the usual idea of unpredictability. I distinguish the objective unpredictability, compatible with determinism (...) and entailed by emergence, and various possibilities of predictability at emergent levels. This makes sense of practices common in complex systems studies that forge qualitative predictions on the basis of comparisons of simulations with multiple values of parameters. I consider robustness analysis as a way to ensure the ontological character of computational emergence. Finally, I focus on the property of novelty, as it is displayed by biological evolution, and ask whether computer simulations of evolution can produce the same kind of emergence as the open-ended evolution attested in Phanerozoic records. (shrink)

Selectionist evolutionary theory has often been faulted for not making novel predictions that are surprising, risky, and correct. I argue that it in fact exhibits the theoretical virtue of predictive capacity in addition to two other virtues: explanatory unification and model fitting. Two case studies show the predictive capacity of selectionist evolutionary theory: parallel evolutionary change in E. coli, and the origin of eukaryotic cells through endosymbiosis.

According to Aristotelian essentialism, the nature of an organism is constituted of a particular goal-directed disposition to produce an organism typical of its kind. This paper argues—against the prevailing orthodoxy—that essentialism of this sort is indispensable to evolutionary biology. The most powerful anti-essentialist arguments purport to show that the natures of organisms play no explanatory role in modern synthesis biology. I argue that recent evolutionary developmental biology provides compelling evidence to the contrary. Developmental biology shows that one must appeal to (...) the capacities of organisms to explain what makes adaptive evolution adaptive. Moreover, the specific capacities in question are precisely those that, according to Aristotle, constitute the nature of an organism. Essentialism 1.1 Aristotelian biological kinds Evolutionary anti-essentialism 2.1 Taxonomic anti-essentialism 2.2 Explanatory anti-essentialism Adaptation 3.1 Stability 3.2 Mutability 3.3 Phenotypic plasticity and adaptive evolution The natures of organisms Conclusion. (shrink)

If the question ``What is a gene?'' proves to be worth asking it must be able to elicit an answer which both recognizes and address the reasons why the concept of the gene ever seemed to be something worth getting excited about in the first place as well analyzing and evaluating the latest develops in the molecular biology of DNA. Each of the preceding papers fails to do one of these and sufferrs the consequences. Where Rolston responds to the apparent (...) failure of molecular biology to make good on the desideratum of the classical gene by veering off into fanciful talk about ``cybernetic genes,'' Griffiths and Stotz lose themselves in the molecular fine print and forget to ask themselves why ``genes'' should be of any special interst anyway. (shrink)

Essentialism is widely regarded as a mistaken view of biological kinds, such as species. After recounting why (sections 2-3), we provide a brief survey of the chief responses to the “death of essentialism” in the philosophy of biology (section 4). We then develop one of these responses, the claim that biological kinds are homeostatic property clusters (sections 5-6) illustrating this view with several novel examples (section 7). Although this view was first expressed 20 years ago, and has received recent discussion (...) and critique, it remains underdeveloped and is often misrepresented by its critics (section 8). (shrink)

Editors' introduction to the special issue on homology (Biology and Philosophy Vol. 22, Issue 5, 2007).

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)

The validity and utility of the proximate-ultimate distinction in biology have recently been under debate. Opponents of the distinction argue that it rules out individual-level organismic processes from evolutionary explanations, thereby leading to an unfounded separation between organismic causation and evolutionary causation. Proponents of the proximate-ultimate distinction, on the other hand, argue that it serves an important epistemological role in forming different kinds of explanation-seeking questions in biology. In this paper we offer an interpretation the proximate-ultimate distinction not only as (...) a means of forming explanation-seeking questions, but also as a distinction that can help highlight the way in which individual-level organismic processes can be evolutionary causes. We do this by interpreting the distinction between proximate and ultimate causes as a distinction between structuring and triggering causes. (shrink)

This dissertation concerns the active role of the organism in evolutionary theory. In particular, it concerns how our conception of the relationship between organism and environment, and the nature of natural selection, influences the causal and explanatory role of organismic activity and behavior in evolutionary explanations. The overarching aim is to argue that the behaviors and activities of organisms can serve both as the explananda (that which is explained) and the explanantia (that which explains) in evolutionary explanations. I attempt to (...) achieve this aim by offering three central arguments. First, that the organism-environment relation is the ontologically basic unit of biology. A common way of conceiving the relationship between organism and environment is as a duality—as two causally independent systems that overlap through interaction. I think this is a mistake. There cannot be organisms without environments and vice versa. They are codependent and codetermined. Second, that natural selection is an ecological process. By this I mean that natural selection acts primarily on organism-environment interactions. It is only in virtue of organisms interacting with their environments that there can be fitness differences amongst individual organisms in a population. The ecological approach to natural selection also entails that the nature of the system(s) of inheritance involved in the reoccurrence of favorable organism-environment interactions is, in principle, immaterial to the process of selection. It only matters for selection that organism-environment interactions actually do reoccur in subsequent generations. Third, that niche construction theory—the most well-developed theoretical and conceptual framework for studying how the activities of organisms influences evolutionary dynamics—can be seen as fully a compatible and integrated part of evolutionary theory. Together, these three arguments constitute an overarching argument. Namely that we both can, and should, take organismic activity and behavior to be crucial explanatory elements in evolutionary theory. Throughout the dissertation I also show how these three arguments have consequences for other debates in the philosophy of evolutionary theory, such as the nature of evolutionary processes, the structure of selection-based explanations, the validity and utility of the proximate-ultimate distinction, and the nature of teleology in evolutionary systems. (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)

Biology has improved a lot over the centuries. The success of developmental biology is huge in every field of physiology, anatomy, paleontology, embryology. The term homology has not been used in biology for a long time. The need for homology in biology is understandable. I have discussed in this paper the necessity of homology in the field of biology and argued that it is unreasonable to refer to homology as evidence of Darwinism, both philosophically and biologically. This paper evaluates homology (...) in the light of the philosophy of biology as evidence for evolution. (shrink)

En este artículo analizaré algunos puntos de convergencia entre la filosofía transhumanista y la crítica filosófica contemporánea al antropocentrismo. Para ello distinguiré entre un tipo de programa normativo fuerte y uno débil en el transhumanismo. El argumento central reside en que la normatividad fuerte en el discurso transhumanista, al suponer que es posible seleccionar exclusivamente las características “propiamente humanas” para ser conservadas en el estadio post-humano, se basa en una concepción pre-darwiniana de la especie humana, lo cual posee implicancias tanto (...) epistemológicas como éticas. (shrink)

Historiographical accounts typically place the formulation of the first embryological theory of the evolutionary origin of vertebrates after the publication of Darwin’s On the Origin of Species. However, the French naturalist Étienne Geoffroy Saint-Hilaire developed an embryological evolutionary model in the 1820s that followed the Lamarckian theory. Geoffroy was the first to establish a direct embryological relationship between vertebrates and invertebrates. This idea was not forgotten, and the embryologists Anton Dohrn and Carl Semper subsequently updated it in their annelid theory (...) as part of a debate about the origin of vertebrates that occurred during the latter part of the nineteenth century. This paper reviews the traditional historiography, analyzing and integrating Geoffroy’s model into the current body of ideas. (shrink)

Comparative biology builds up systematic knowledge of the diversity of life, across evolutionary lineages and levels of organization, starting with evidence from a sparse sample of model organisms. In developmental biology, a key obstacle to the growth of comparative approaches is that the concept of homology is not very well defined for levels of organization that are intermediate between individual genes and morphological characters. In this paper, we investigate what it means for ontogenetic processes to be homologous, focusing specifically on (...) the examples of insect segmentation and vertebrate somitogenesis. These processes can be homologous without homology of the underlying genes or gene networks, since the latter can diverge over evolutionary time, while the dynamics of the process remain the same. Ontogenetic processes like these therefore constitute a dissociable level and distinctive unit of comparison requiring their own specific criteria of homology. In addition, such processes are typically complex and nonlinear, such that their rigorous description and comparison not only requires observation and experimentation, but also dynamical modeling. We propose six criteria of process homology, combining recognized indicators with novel ones derived from dynamical systems modeling. We show how these criteria apply to animal segmentation and other ontogenetic processes. We conclude by situating our proposed dynamical framework for homology of process in relation to similar research programs, such as process structuralism and developmental approaches to morphological homology. (shrink)

A given explanatory theory T falls into circular reasoning if the only way to determine its explanandum is through the application of T. To find an underlying theory T′ that determines T′s explanandum helps us save T from this accusation of circularity. We follow the structuralist view of theories in presenting and dealing with this issue, by applying it to particular theories. More specifically, we focus on the relationship between the Darwinian theory of common ancestry and the determination of homologies.

En este trabajo discutimos la extensión de la influencia que el pensamiento de Richard Owen tuvo sobre el de Charles Darwin. Además, se intentará mostrar lo heterogéneo de tal influencia, que va desde teorías específicas a giros retóricos. Esta influencia es en muchos casos subestimada, dando la sensación de que la novedad darwiniana consistió únicamente en mirar con ojos desprejuiciados lo que los otros no habían visto. Esta visión resulta injusta con Owen, y también con el esfuerzo conceptual llevado adelante (...) por Darwin con las piezas brindadas por sus precursores. Finalmente, este es un caso interesante para entender el tipo de novedad aportada por las revoluciones científicas y el modo sofisticado en que tal novedad se sustenta sobre el trabajo de los enfoques previos. (shrink)

The standard perception of the dichotomy between population thinking and essentialism (typological thinking) in evolutionary economics descends from the golden age of the neo-Darwinian Synthesis. Over the last few decades the received view on population thinking has been seriously challenged in biology and its philosophy. First, the strong version of population thinking that banishes essentialism witnessed important tensions stemming from the ontological status of species. These tensions have been amplified by the demise of positivism and the rise of a new (...) essentialism in philosophy of science. Second, the soft version that transforms the opposition between population thinking and essentialism to the dichotomy between ultimate and proximate causation has led to contradictory interpretations regarding the locus of ultimate causes. Taking stock of the previous discussion the paper addresses the limits to population thinking in the socio-economic realm. The upshot is that without denying the important achievements made by the application of population thinking in sub-disciplines like industrial dynamics and economic anthropology, the idea to generalize these applications into the whole socio-economic realm is problematic. The aforementioned achievements cannot come to grips with the structural aspects of capitalism, its different periods (e.g. the contemporary finance-led capitalism) and its geographical varieties. The resulting gap points to the importance of structural analysis (essentialism) and evolutionary political economy. The latter is distinguished from the rest of evolutionary economics by its project to go beyond the surface of economic phenomena and to critically analyze their underlying social structures. (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)

The theory of concepts advanced in the dissertation aims at accounting for a) how a concept makes successful practice possible, and b) how a scientific concept can be subject to rational change in the course of history. Traditional accounts in the philosophy of science have usually studied concepts in terms only of their reference; their concern is to establish a stability of reference in order to address the incommensurability problem. My discussion, in contrast, suggests that each scientific concept consists of (...) three components of content: 1) reference, 2) inferential role, and 3) the epistemic goal pursued with the concept's use. I argue that in the course of history a concept can change in any of these three components, and that change in one component—including change of reference—can be accounted for as being rational relative to other components, in particular a concept's epistemic goal.This semantic framework is applied to two cases from the history of biology: the homology concept as used in 19th and 20th century biology, and the gene concept as used in different parts of the 20th century. The homology case study argues that the advent of Darwinian evolutionary theory, despite introducing a new definition of homology, did not bring about a new homology concept (distinct from the pre-Darwinian concept) in the 19th century. Nowadays, however, distinct homology concepts are used in systematics/evolutionary biology, in evolutionary developmental biology, and in molecular biology. The emergence of these different homology concepts is explained as occurring in a rational fashion. The gene case study argues that conceptual progress occurred with the transition from the classical to the molecular gene concept, despite a change in reference. In the last two decades, change occurred internal to the molecular gene concept, so that nowadays this concept's usage and reference varies from context to context. I argue that this situation emerged rationally and that the current variation in usage and reference is conducive to biological practice.The dissertation uses ideas and methodological tools from the philosophy of mind and language, the philosophy of science, the history of science, and the psychology of concepts. (shrink)

ZusammenfassungDer Artikel führt den Begriff der epistemischen Konkurrenz ein. Im Gegensatz zu „wissenschaftliche Kontroverse“ beschreibt er eine Situation, in der sich zwei Forschungsfelder gegenseitig als mit demselben Bereich von Phänomen befasst wahrnehmen, wobei ihre methodischen Ansätze und theoretischen Erklärungen jedoch so unterschiedlich sind, dass ein offener Konflikt über die Wahrheit oder Falschheit bestimmter Aussagen oder die Genauigkeit in der Anwendung einer Methode nicht stattfindet. Nichtsdestotrotz streben beide Parteien danach, die maßgebliche Erklärung der entsprechenden Phänomene anzubieten. Indem die erweiterte Gemeinschaft der (...) Forschenden oder die Öffentlichkeit einen Ansatz als maßgeblich anerkennt, handeln sie als eine dritte, gewissermaßen neutrale Partei, die einen Preis vergibt, um den die anderen Parteien konkurrieren. Der Artikel beschreibt das Verhältnis von Genetik und Entwicklungsbiologie um 1900 als epistemische Konkurrenz. Diese Forschungsfelder geben unterschiedliche Erklärungen für die Phänomene der organischen Reproduktion. Die Erklärungen unterscheiden sich bezüglich der Formbegriffe und entsprechend hinsichtlich der Begriffe der Vererbung von Aspekten der Form eines Organismus. Zudem basieren die Erklärungen der beiden Felder auf unterschiedlichen Arten kausalen Schließens, die in unterschiedliche experimentelle Ansätze eingebettet sind. Diese Unterschiede können als Instanzen eines allgemeineren Unterschieds zwischen der Tradition der Naturgeschichte, die sich mit Unterschieden zwischen Organismen auseinandersetzt, und der Tradition der Anatomie, die sich mit den Teilen der Organismen beschäftigt, angesehen werden. Dieses Bild der Genetik und Entwicklungsbiologie als Zweige unterschiedlicher fest verankerter Traditionen widerspricht dem Narrativ der Trennung des Begriffs der Vererbung von dem der Entwicklung im Zuge einer Trennung der Disziplinen der Genetik und der Embryologie, das in der Historiographie der Biologie nach wie vor weit verbreitet ist. (shrink)

Research on animal behavior is typically organized according to a combination of two influential frameworks: Ernst Mayr’s distinction between proximate and ultimate causes, and Niko Tinbergen’s “four questions”. My aim is to debunk two common interpretive misconceptions about Mayr’s proximate–ultimate distinction and its relationship to Tinbergen’s four questions, and to offer a new interpretation that avoids both. The first misconception is that the proximate–ultimate distinction maps cleanly onto Tinbergen’s four questions, marking a boundary between Tinbergen’s evolutionary and survival value questions (...) versus developmental and mechanistic questions. The second is that Mayr’s proximate–ultimate distinction is meant to rule out the relevance of proximate causes to evolutionary explanations. I argue that neither is plausible given the text and Mayr’s philosophical aims, namely, to argue that evolutionary biology cannot be reduced to either the physical sciences or to other areas of biology. Through a reconstruction of Mayr’s anti-reductionist argument, I develop an interpretation according to which the proximate–ultimate distinction marks two ways that teleological reasoning can be naturalistically grounded in biology, corresponding to Mayr’s distinction between teleonomic and adapted systems. Mayr distinguishes reduction, which the proximate–ultimate distinction is meant to block, from analysis, through which he allows that proximate causes, causes that are neither proximate nor ultimate, and chance can all contribute to evolutionary explanations. I conclude by suggesting some ways in which the interpretation defended here reframes our understanding of Mayr’s disagreements with some evolutionary-developmental biologists. (shrink)

Research on animal behavior is typically organized according to a combination of two influential frameworks: Ernst Mayr’s distinction between proximate and ultimate causes, and Niko Tinbergen’s “four questions”. My aim is to debunk two common interpretive misconceptions about Mayr’s proximate–ultimate distinction and its relationship to Tinbergen’s four questions, and to offer a new interpretation that avoids both. The first misconception is that the proximate–ultimate distinction maps cleanly onto Tinbergen’s four questions, marking a boundary between Tinbergen’s evolutionary and survival value questions (...) versus developmental and mechanistic questions. The second is that Mayr’s proximate–ultimate distinction is meant to rule out the relevance of proximate causes to evolutionary explanations. I argue that neither is plausible given the text and Mayr’s philosophical aims, namely, to argue that evolutionary biology cannot be reduced to either the physical sciences or to other areas of biology. Through a reconstruction of Mayr’s anti-reductionist argument, I develop an interpretation according to which the proximate–ultimate distinction marks two ways that teleological reasoning can be naturalistically grounded in biology, corresponding to Mayr’s distinction between teleonomic and adapted systems. Mayr distinguishes reduction, which the proximate–ultimate distinction is meant to block, from analysis, through which he allows that proximate causes, causes that are neither proximate nor ultimate, and chance can all contribute to evolutionary explanations. I conclude by suggesting some ways in which the interpretation defended here reframes our understanding of Mayr’s disagreements with some evolutionary-developmental biologists. (shrink)

This book is divided in two parts, the first of which shows how, beyond paleontology and systematics, macroevolutionary theories apply key insights from ecology and biogeography, developmental biology, biophysics, molecular phylogenetics, and even the sociocultural sciences to explain evolution in deep time. In the second part, the phenomenon of macroevolution is examined with the help of real life-history case studies on the evolution of eukaryotic sex, the formation of anatomical form and body-plans, extinction and speciation events of marine invertebrates, hominin (...) evolution and species conservation ethics. The book brings together leading experts, who explain pivotal concepts such as Punctuated Equilibria, Stasis, Developmental Constraints, Adaptive Radiations, Habitat Tracking, Turnovers, (Mass) Extinctions, Species Sorting, Major Transitions, Trends, and Hierarchies – key premises that allow macroevolutionary epistemic frameworks to transcend microevolutionary theories that focus on genetic variation, selection, migration and fitness. Along the way, the contributing authors review ongoing debates and current scientific challenges; detail new and fascinating scientific tools and techniques that allow us to cross the classic borders between disciplines; demonstrate how their theories make it possible to extend the Modern Synthesis; present guidelines on how the macroevolutionary field could be further developed; and provide a rich view of just how it was that life evolved across time and space. In short, this book is a must-read for active scholars and, because the technical aspects are fully explained, it is also accessible for non-specialists. Understanding evolution requires a solid grasp of above-population phenomena. Species are real biological individuals, and abiotic factors impact the future course of evolution. Beyond observation, when the explanation of macroevolution is the goal, we need both evidence and theory that enable us to explain and interpret how life evolves at the grand scale. (shrink)

Throughout the last quarter of the nineteenth century, researchers became increasingly interested in explaining the ways in which mammalian teeth, especially molars, and their complex arrangements of cusps arose along both developmental and evolutionary timescales. By the 1890s, two theories garnered special prominence; the tritubercular theory and the concrescence theory. The tritubercular theory was proposed by Edward Drinker Cope in 1883, and later expanded by Henry Fairfield Osborn in 1888, while the concrescence theory was developed by Carl Röse in 1892. (...) Reviews concerning the evolution of mammalian molar teeth tended to paint the two theories as occupying opposing sides, and debates arose between their main proponents; however, their tenets do not seem logically incompatible. Throughout this paper, I argue that the conflict that arose was due not to the content of the theories, but to a diverse array of commitments Cope, Osborn, and Röse held, which turned into background assumptions within the setting of these theories. This history traces the context in which Cope, Osborn, and Röse developed the tritubercular and concrescence theories, and the ways in which the assumptions that these investigators held influenced their perceptions of their theories. (shrink)

Recent challenges to non-traditional theories of natural kinds demand clarifications and revisions to those theories. Highlights: The semantics of natural kind terms is a special case of a general naturalistic conception of signaling in organisms that explains the epistemic reliability of signaling. Natural kinds and reference are two aspects of the same natural phenomenon. Natural kind definitions are phenomena in nature not linguistic or representational entities; their relation to conceptualized definitions is complex. Reference and truth are special cases of a (...) broader phenomenon of accommodation between aspects of signaling and epistemically relevant causal structures. (shrink)

Contemporary evolutionary biology comprises a plural landscape of multiple co-existent conceptual frameworks and strenuous voices that disagree on the nature and scope of evolutionary theory. Since the mid-eighties, some of these conceptual frameworks have denounced the ontologies of the Modern Synthesis and of the updated Standard Theory of Evolution as unfinished or even flawed. In this paper, we analyze and compare two of those conceptual frameworks, namely Niles Eldredge’s Hierarchy Theory of Evolution (with its extended ontology of evolutionary entities) and (...) the Extended Evolutionary Synthesis (with its proposal of an extended ontology of evolutionary processes), in an attempt to map some epistemic bridges (e.g. compatible views of causation; niche construction) and some conceptual rifts (e.g. extra-genetic inheritance; different perspectives on macroevolution; contrasting standpoints held in the “externalism–internalism” debate) that exist between them. This paper seeks to encourage theoretical, philosophical and historiographical discussions about pluralism or the possible unification of contemporary evolutionary biology. (shrink)

The Extended Evolutionary Synthesis (EES) debate is gaining ground in contemporary evolutionary biology. In parallel, a number of philosophical standpoints have emerged in an attempt to clarify what exactly is represented by the EES. For Massimo Pigliucci, we are in the wake of the newest instantiation of a persisting Kuhnian paradigm; in contrast, Telmo Pievani has contended that the transition to an EES could be best represented as a progressive reformation of a prior Lakatosian scientific research program, with the extension (...) of its Neo-Darwinian core and the addition of a brand-new protective belt of assumptions and auxiliary hypotheses. Here, we argue that those philosophical vantage points are not the only ways to interpret what current proposals to ‘extend’ the Modern Synthesis-derived ‘standard evolutionary theory’ (SET) entail in terms of theoretical change in evolutionary biology. We specifically propose the image of the emergent EES as a vast network of models and interweaved representations that, instantiated in diverse practices, are connected and related in multiple ways. Under that assumption, the EES could be articulated around a paraconsistent network of evolutionary theories (including some elements of the SET), as well as models, practices and representation systems of contemporary evolutionary biology, with edges and nodes that change their position and centrality as a consequence of the co-construction and stabilization of facts and historical discussions revolving around the epistemic goals of this area of the life sciences. We then critically examine the purported structure of the EES—published by Laland and collaborators in 2015—in light of our own network-based proposal. Finally, we consider which epistemic units of Evo-Devo are present or still missing from the EES, in preparation for further analyses of the topic of explanatory integration in this conceptual framework. (shrink)

Der Artikel führt den Begriff der epistemischen Konkurrenz ein. Im Gegensatz zu „wissenschaftliche Kontroverse“ beschreibt er eine Situation, in der sich zwei Forschungsfelder gegenseitig als mit demselben Bereich von Phänomen befasst wahrnehmen, wobei ihre methodischen Ansätze und theoretischen Erklärungen jedoch so unterschiedlich sind, dass ein offener Konflikt über die Wahrheit oder Falschheit bestimmter Aussagen oder die Genauigkeit in der Anwendung einer Methode nicht stattfindet. Nichtsdestotrotz streben beide Parteien danach, die maßgebliche Erklärung der entsprechenden Phänomene anzubieten. Indem die erweiterte Gemeinschaft der (...) Forschenden oder die Öffentlichkeit einen Ansatz als maßgeblich anerkennt, handeln sie als eine dritte, gewissermaßen neutrale Partei, die einen Preis vergibt, um den die anderen Parteien konkurrieren. Der Artikel beschreibt das Verhältnis von Genetik und Entwicklungsbiologie um 1900 als epistemische Konkurrenz. Diese Forschungsfelder geben unterschiedliche Erklärungen für die Phänomene der organischen Reproduktion. Die Erklärungen unterscheiden sich bezüglich der Formbegriffe und entsprechend hinsichtlich der Begriffe der Vererbung von Aspekten der Form eines Organismus. Zudem basieren die Erklärungen der beiden Felder auf unterschiedlichen Arten kausalen Schließens, die in unterschiedliche experimentelle Ansätze eingebettet sind. Diese Unterschiede können als Instanzen eines allgemeineren Unterschieds zwischen der Tradition der Naturgeschichte, die sich mit Unterschieden zwischen Organismen auseinandersetzt, und der Tradition der Anatomie, die sich mit den Teilen der Organismen beschäftigt, angesehen werden. Dieses Bild der Genetik und Entwicklungsbiologie als Zweige unterschiedlicher fest verankerter Traditionen widerspricht dem Narrativ der Trennung des Begriffs der Vererbung von dem der Entwicklung im Zuge einer Trennung der Disziplinen der Genetik und der Embryologie, das in der Historiographie der Biologie nach wie vor weit verbreitet ist. (shrink)

There is long-standing conflict between genealogical and developmental accounts of homology. This paper provides a general framework that shows that these accounts are compatible and clarifies precisely how they are related. According to this framework, understanding homology requires both an abstract genealogical account that unifies the application of the term to all types of characters used in phylogenetic systematics and locally enriched accounts that apply only to specific types of characters. The genealogical account serves this unifying role by relying on (...) abstract notions of ‘descent’ and ‘character’. As a result, it takes for granted the existence of such characters. This requires theoretical justification that is provided by enriched accounts, which incorporate the details by which characters are inherited. These enriched accounts apply to limited domains, providing the needed theoretical justification for recognizing characters within that domain. Though connected to the genealogical account of homology in this way, enriched accounts include phenomena that fall outside the scope of the genealogical account. They therefore overlap, but are not nested within, the genealogical account. Developmental accounts of homology are to be understood as enriched accounts of body part homology. Once they are seen in this light, the conflict with the genealogical account vanishes. It is only by understanding the fine conceptual structure undergirding the many uses of the term ‘homology’ that we can understand how these uses hang together. (shrink)

I propose a new account of homology, according to which homology is a correspondence of developmental mechanisms due to common ancestry, formally defined as an isomorphism of causal graphs over lineages. The semiformal definition highlights the role of homology as a higher-order principle unifying evolutionary models and also provides definite meanings to concepts like constraints, evolvability, and novelty. The novel interpretation of homology suggests a broad perspective that accommodates evolutionary developmental biology and traditional population genetics as distinct but complementary approaches (...) to understand evolution, prompting further empirical and theoretical research. (shrink)

By formulating in 1875 his major theoretical achievement, the “principle of succession of functions”, Dohrn was consciously entering the controversy between Darwin and Mivart. Dohrn’s principle enjoyed the approval of Darwin, but not his enthusiasm. The paper examines the evolution of Darwin’s original idea of ‘conversion of functions’ in the 6th edition of his Origin, following Mivart’s criticism, and contrasts the overtly functionalist interpretation entailed in Dohrn’s formulation with Darwin’s increasing structuralist hesitations as to the origin of evolutionary novelty. A (...) more accurate analysis of Dohrn’s principle, however, appears to corroborate the thesis that Dohrn was equally receptive to Darwin’s argument as to Mivart’s criticism. (shrink)

An important question facing contemporary philosophy of science is whether the natural sciences in terms of their historical records exhibit distinguishing developmental patterns or structures. At least two philosophical stances are possible in answering this question. The first pertains to the plurality of the individual sciences. From this stance, the various sciences are analyzed individually and compared with one another in order to derive potential commonalities, if any, among them. The second stance involves a general philosophy of science in which (...) a thorough theory of the natural sciences is developed. The latter stance strives to account for more than possible commonalities among the sciences but also to provide a broad-spectrum philosophical framework to account for, or to explicate, the nature of science itself and its progress. In this paper, the second stance is taken in which an evolutionary philosophy of science is proposed. To that end, Thomas Kuhn’s evolutionary philosophy of science is initially discussed and critiqued. An evolutionary philosophy of science is then proposed based on a revision of Kuhn’s evolutionary philosophy of science in terms of George Gaylord Simpson’s various tempos and modes for biological evolution. Next, two historical case studies from the biological sciences are reconstructed to illustrate the robustness of the proposed evolutionary philosophy of science for explicating the progress of the natural sciences. A concluding section discusses the proposed evolutionary philosophy of science with respect to providing a broad-spectrum framework or general philosophy of science for understanding the nature and progress of the natural sciences. (shrink)

Although contemporary metaphysics has recently undergone a neo-Aristotelian revival wherein dispositions, or capacities are now commonplace in empirically grounded ontologies, being routinely utilised in theories of causality and modality, a central Aristotelian concept has yet to be given serious attention – the doctrine of hylomorphism. The reason for this is clear: while the Aristotelian ontological distinction between actuality and potentiality has proven to be a fruitful conceptual framework with which to model the operation of the natural world, the distinction between (...) form and matter has yet to similarly earn its keep. In this chapter, I offer a first step toward showing that the hylomorphic framework is up to that task. To do so, I return to the birthplace of that doctrine - the biological realm. Utilising recent advances in developmental biology, I argue that the hylomorphic framework is an empirically adequate and conceptually rich explanatory schema with which to model the nature of organisms. (shrink)