Despite the traditional focus on metaphysical issues in discussions of natural kinds in biology, epistemological considerations are at least as important. By revisiting the debate as to whether taxa are kinds or individuals, I argue that both accounts are metaphysically compatible, but that one or the other approach can be pragmatically preferable depending on the epistemic context. Recent objections against construing species as homeostatic property cluster kinds are also addressed. The second part of the paper broadens the perspective by considering (...) homologues as another example of natural kinds, comparing them with analogues as functionally defined kinds. Given that there are various types of natural kinds, I discuss the different theoretical purposes served by diverse kind concepts, suggesting that there is no clear-cut distinction between natural kinds and other kinds, such as functional kinds. Rather than attempting to offer a unique metaphysical account of ‘natural’ kind, a more fruitful approach consists in the epistemological study of how different natural kind concepts are employed in scientific reasoning. (shrink)

Aristotle’s influence on D’Arcy Thompson was praised by Thompson himself and has been recognized by others in various respects, including the aesthetic and normative dimensions of biology, and the multicausal explanation of living forms. This article focuses on the relatedness of organic forms, one of the core problems addressed by both Aristotle’s History of Animals (HA), and the renowned chapter of Thompson’s On Growth and Form (G&F), “On the Theory of Transformations, or the Comparison of Related Forms.” We contend that, (...) far from being an incidental inspiration stemming from Thompson’s classicist background, his translation of HA played a pivotal role in developing his theory of transformations. Furthermore, we argue that Thompson’s interpretation of the Aristotelian method of comparison challenges the prevailing view of Aristotle as the founder of “typological essentialism,” and is a key episode in the revision of this narrative. Thompson understood that the method Aristotle used in HA to compare animal forms is better comprehended as a “method of transformations,” leading to a morphological arrangement of animal diversity, as opposed to a taxonomical classification. Finally, we examine how this approach to the relatedness of forms lay the foundation for a causal understanding of parts and their interconnections. Although Aristotle and Thompson emphasized distinct types of causes, we contend that they both differ in a fundamental sense from the one introduced by Darwin’s theory of natural selection, which was formulated as a solution to the species problem rather than the form problem. We conclude that Thompson’s interpretation of Aristotle’s approach to form comparison has not only impacted contemporary scholarship on Aristotle’s biology, but revitalized a perspective that has regained significance due to the resurgence of the problem of form in evo-devo. (shrink)

Molecular methods have revolutionised virtually every area of biology, and metazoan phylogenetics is no exception: molecular phylogenies, molecular clocks, comparative phylogenomics, and developmental genetics have generated a plethora of molecular data spanning numerous taxa and collectively transformed our understanding of the evolutionary history of animals, often corroborating but at times opposing results of more traditional approaches. Moreover, the diversity of methods and models within molecular phylogenetics has resulted in significant disagreement among molecular phylogenies as well as between these and earlier (...) phylogenies. How should this broad and multifaceted problem be tackled? I argue that the answer lies in integrating evidence to infer the best evolutionary scenario. I begin with an overview of recent development in early metazoan phylogenetics, followed by a discussion of key conceptual issues in phylogenetics revolving around phylogenetic evidence, theory, methodology, and interrelations thereof. I then argue that the integration of different kinds of evidence is necessary for arriving at the best evolutionary scenario rather than merely the best-fitting cladogram. Finally, I discuss the prospects of this view in stimulating interdisciplinary cross-talk in early metazoan research and beyond, and challenges that need to be overcome. (shrink)

This paper attempts to reconcile the conceptual conflict between typological and population thinking to provide a philosophical foundation for extended evolutionary synthesis. Typological thinking has been considered a pre-Darwinian, essentialist dogma incompatible with population thinking, which is the core notion of Darwinism. More recent philosophical and historical studies suggest that a non-essentialist form of typology has some advantages in the study of evolutionary biology. However, even if we adopt such an epistemological interpretation of typological thinking, there still remains an epistemological (...) and methodological conflict between these two styles of thinking. How can we relate typological thinking with population thinking in pursuit of more integrated or interconnected research into evolutionary biology? I propose that homology thinking, which is another style of thinking that recognizes homologous characters, provides a common basis for typological representations of character states and for character dynamics in an evolving population. Good examples of this bridging role are found in teratology and breeding, where variation and novelty are recognized in developmental and morphological traits, gene expression patterns, and so on. Essentialism-free, dynamic views of homology have great potential to reconcile typological and population thinking and to set the stage for the EES. (shrink)

Although there is no in-principle impediment to an EvoDevo of behavior, such an endeavor is not as straightforward as one might think; many of the key terms and concepts used in EvoDevo are tailored to suit its traditional focus on morphology, and are consequently difficult to apply to behavior. In this light, the application of the EvoDevo conceptual toolkit to the behavioral domain requires the establishment of a set of tractable concepts that are readily applicable to behavioral characters. Here, I (...) begin the type of theoretical work that needs to be undertaken in order to achieve this, focusing in particular on the key concept of “novelty.” Building on existing criteria used for the identification of behavioral homology from behavioral ecology, I develop a set of operational criteria for identifying novelty in the behavioral domain. These criteria provide a conceptual foundation for the study of novelty in behavioral traits. (shrink)

En este artículo argumento que para la Síntesis extendida resulta imperioso reemplazarlos modelos causales tradicionales en biología, el de causas próximas— últimas y el de lacausalidad lineal ascendente, debido a su incapacidad para capturar la causalidad biológicacompleja que describen la mayoría de áreas de investigación en la actualidad. Estas nuevastendencias muestran la necesidad de reconceptualizar la causalidad y así permitir la construcciónde modelos multinivel que conecten bidireccionalmente diversos niveles de organizacióny diferentes escalas de tiempo. La causalidad multinivel es fundamental en (...) losprocesos de organización natural, por lo que, argumento, la extensión de la síntesis debeubicarla en su base. (shrink)

This volume provides a broad overview of issues in the philosophy of behavioral biology, covering four main themes: genetic, developmental, evolutionary, and neurobiological explanations of behavior. It is both interdisciplinary and empirically informed in its approach, addressing philosophical issues that arise from recent scientific findings in biological research on human and non-human animal behavior. Accordingly, it includes papers by professional philosophers and philosophers of science, as well as practicing scientists. Much of the work in this volume builds on presentations given (...) at the international conference, “Biological Explanations of Behavior: Philosophical Perspectives”, held in 2008 at the Leibniz Universität Hannover in Germany. The volume is intended to be of interest to a broad range of audiences, which includes philosophers (e.g., philosophers of mind, philosophers of biology, and metaethicists), as well as practicing scientists, such as biologists or psychologists whose interests relate to biological explanations of behavior. (shrink)

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

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)

Researchers in the cognitive sciences often seek neural correlates of psychological constructs. In this paper, I argue that even when these correlates are discovered, they do not always lead to reductive outcomes. To this end, I examine the psychological construct of a critical period and briefly describe research identifying its neural correlates. Although the critical period is correlated with certain neural mechanisms, this does not imply that there is a reductionist relationship between this psychological construct and its neural correlates. Instead, (...) this case study suggests that there may be many-to-many psychological-neural mappings, not just one-to-one or even one-to-many relations between psychological kinds and types of neural mechanisms. (shrink)

The traditional practice of establishing morphological types and investigating morphological organization has found new support from evolutionary developmental biology (evo-devo), especially with respect to the notion of body plans. Despite recurring claims that typology is at odds with evolutionary thinking, evo-devo offers mechanistic explanations of the evolutionary origin, transformation, and evolvability of morphological organization. In parallel, philosophers have developed non-essentialist conceptions of natural kinds that permit kinds to exhibit variation and undergo change. This not only facilitates a construal of species (...) and higher taxa as natural kinds, but also broadens our perspective on the diversity of kinds found in biology. There are many different natural kinds relevant to the investigative and explanatory aims of evo-devo, including homologues and developmental modules. (shrink)

I will begin by proposing a taxonomy of taxonomic positions regarding the mind–brain: localism, globalism, revisionism, and contextualism, and will go on to focus on the last position. Although some versions of contextualism have been defended by various researchers, they largely limit themselves to a version of neural contextualism: different brain regions perform different functions in different neural contexts. I will defend what I call “environmental-etiological contextualism,” according to which the psychological functions carried out by various neural regions can only (...) be identified and individuated against an environmental context or with reference to a causal history. While this idea may seem innocuous enough, it has important implications for a structure-to-function mapping in the mind and brain sciences. It entails that the same neural structures can subserve different psychological functions in different contexts, leading to crosscutting psycho-neural mappings. I will try to illustrate how this can occur with reference to recent research on episodic memory. (shrink)

The biological sciences have always proven a fertile ground for philosophical analysis, one from which has grown a rich tradition stemming from Aristotle and flowering with Darwin. And although contemporary philosophy is increasingly becoming conceptually entwined with the study of the empirical sciences with the data of the latter now being regularly utilised in the establishment and defence of the frameworks of the former, a practice especially prominent in the philosophy of physics, the development of that tradition hasn’t received the (...) wider attention it so thoroughly deserves. This review will briefly introduce some recent significant topics of debate within the philosophy of biology, focusing on those whose metaphysical themes (in everything from composition to causation) are likely to be of wide-reaching, cross-disciplinary interest. (shrink)

Evolutionary developmental biology represents a paradigm shift in the understanding of the ontogenesis and evolutionary progression of the denizens of the natural world. Given the empirical successes of the evo-devo framework, and its now widespread acceptance, a timely and important task for the philosophy of biology is to critically discern the ontological commitments of that framework and assess whether and to what extent our current metaphysical models are able to accommodate them. In this paper, I argue that one particular model (...) is a natural fit: an ontology of dispositional properties coherently and adequately captures the crucial casual-cum-explanatory role that the fundamental elements of evo-devo play within that framework. (shrink)

Though the realm of biology has long been under the philosophical rule of the mechanistic magisterium, recent years have seen a surprisingly steady rise in the usurping prowess of process ontology. According to its proponents, theoretical advances in the contemporary science of evo-devo have afforded that ontology a particularly powerful claim to the throne: in that increasingly empirically confirmed discipline, emergently autonomous, higher-order entities are the reigning explanantia. If we are to accept the election of evo-devo as our best conceptualisation (...) of the biological realm with metaphysical rigour, must we depose our mechanistic ontology for failing to properly “carve at the joints” of organisms? In this paper, I challenge the legitimacy of that claim: not only can the theoretical benefits offered by a process ontology be had without it, they cannot be sufficiently grounded without the metaphysical underpinning of the very mechanisms which processes purport to replace. The biological realm, I argue, remains one best understood as under the governance of mechanistic principles. (shrink)

Evolutionary developmental biology (Evo-devo) is a vibrant area of contemporary life science that should be (and is) increasingly incorporated into teaching curricula. Although the inclusion of this content is important for biological pedagogy at multiple levels of instruction, there are also philosophical lessons that can be drawn from the scientific practices found in Evo-devo. One feature of particular significance is the interdisciplinary nature of Evo-devo investigations and their resulting explanations. Instead of a single disciplinary approach being the most explanatory or (...) fundamental, different methodologies from biological disciplines must be synthesized to generate empirically adequate explanations. Thus, Evo-devo points toward a non-reductionist epistemology in biology. I review three areas where these synthetic efforts become manifest as a result of Evo-devo’s practices (form versus function reasoning styles; problem-structured investigations; idealizations related to studying model organisms), and then sketch some possible applications to teaching biology. These philosophical considerations provide resources for life science educators to address (and challenge) key aspects of the National Science Education Standards and Benchmarks for Scientific Literacy. (shrink)

The goal of this paper is to encourage a reconfiguration of the discussion about typology in biology away from the metaphysics of essentialism and toward the epistemology of classifying natural phenomena for the purposes of empirical inquiry. First, I briefly review arguments concerning ‘typological thinking’, essentialism, species, and natural kinds, highlighting their predominantly metaphysical nature. Second, I use a distinction between the aims, strategies, and tactics of science to suggest how a shift from metaphysics to epistemology might be accomplished. Typological (...) thinking can be understood as a scientific tactic that involves representing natural phenomena using idealizations and approximations, which facilitates explanation, investigation, and theorizing via abstraction and generalization. Third, a variety of typologies from different areas of biology are introduced to emphasize the diversity of this representational reasoning. One particular example is used to examine how there can be epistemological conflict between typology and evolutionary analysis. This demonstrates that alternative strategies of typological thinking arise due to the divergent explanatory goals of researchers working in different disciplines with disparate methodologies. I conclude with several research questions that emerge from an epistemological reconfiguration of typology. (shrink)

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

Advances in developmental genetics and evo-devo in the last several decades have enabled the growth of novel developmental approaches to the classic theme of homology. These approaches depart from the more standard phylogenetic view by contending that homology between morphological characters depends on developmental-genetic individuation and explanation. This article provides a systematic re-examination of the relationship between developmental and phylogenetic homology in light of current evidence from developmental and evolutionary genetics and genomics. I present a qualitative model of the processes (...) that cause de-coupling of morphological and molecular evolution by developmental system drift (DSD), and hypothesize that DSD is a widespread and regular stochastic process that is predictable from parameters such as population size, pleiotropy, and regulatory redundancy. These theoretical findings support an integrative approach in which models of DSD aid in determining when developmental-genetic explanations of homology apply and when other explanations such as stabilizing selection are needed. I argue that a phylogenetic monism about the definition and criteria of homology supports the integration of different explanations into a theory of homology better than pluralism does. (shrink)

What is the relationship between evolutionary contingency and diversity? The evolutionary contingency thesis emphasizes dependency relations and chance as the hallmarks of evolution. While contingency can be destructive of, for example, the fragile and complex dynamics in an ecosystem, I will mainly focus on the productive or causal aspect of contingency for a particular sort of diversity. There are many sorts of diversities: Gould is most famous for his diversity-to-decimation model, which includes disparate body plans distinguishing different phyla. However, structural (...) diversity construed more broadly spans scales, such as organization in and among cells, structural arrangements and biomechanics on various scales, and even the profile of ancestor-descendent relationships or community structure of interactions within ecosystems. By focusing on stochastic processes in contingent evolution, I argue that contingency causes structural diversity. Specifically, I focus on the plurality of structural types of cells, genetic codes, and phyla diversity as case studies. (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)

Homology is a fundamental concept in biology. However, the metaphysical status of homology, especially whether a homolog is a part of an individual or a member of a natural kind, is still a matter of intense debate. The proponents of the individuality view of homology criticize the natural kind view of homology by pointing out that homologs are subject to evolutionary transformation, and natural kinds do not change in the evolutionary process. Conversely, some proponents of the natural kind view of (...) homology argue that a homolog can be construed both as a part of an individual and a member of a natural kind. They adopt the Homeostatic Property Cluster theory of natural kinds, and the theory seems to strongly support their construal. Note that this construal implies the acceptance of essentialism. However, looking back on the history of the concept of homology, we should not overlook the fact that the individuality view was proposed to reject the essentialist interpretation of homology. Moreover, the essentialist notions of natural kinds can, in our view, mislead biologists about the phenomena of homology. Consequently, we need a non-essentialist view of homology, which we name the “persistently reproducible module” view. This view highlights both the individual-like and kind-like aspects of homologs while stripping down both essentialist and anti-essentialist interpretations of homology. In this article, we articulate the PRM view of homology and explain why it is recommended over the other two views. (shrink)

In this introductory article to the special issue on Multi-level semiosis we attempt to stage the background for qualifying the notion of “multi-levelness” when considering communication processes and semiosis in all life forms, i.e. from the cellular to the organismic level. While structures are organized hierarchically, communication processes require a kind of processual organization that may be better described as being heterarchical. Theoretically, the challenge arises in the temporal domain, that is, in the developmental and evolutionary dimension of dynamic semiotic (...) processes. We discuss the importance of this fundamental difference in order to explain how levels, domains and orders of magnitude, on the one hand, and synchronic and diachronic processes, on the other, contribute to the overall organization of every living being. To account for such multi-level organization, semiotic freedom is assumed to be a scalar property that endows living systems at different levels and domains with the capacity to ponder selectively the overall structural coherence and functional compatibility of their heterarchical processing, which is increasingly less conditioned by the underlying molecular determinism. (shrink)

The 1981 Dahlem conference was a catalyst for contemporary evolutionary developmental biology (Evo-devo). This introductory chapter rehearses some of the details of the history surrounding the original conference and its associated edited volume, explicates the philosophical problem of conceptual change that provided the rationale for a workshop devoted to evaluating the epistemic revisions and transformations that occurred in the interim, explores conceptual change with respect to the concept of evolutionary novelty, and highlights some of the themes and patterns in the (...) different contributions to the present volume, Conceptual Change in Biology: Scientific and Philosophical Perspectives on Evolution and Development. (shrink)

The aim of this article is to detail some reservations against the beliefs, claims, or presuppositions that current essentialist natural kind concepts (including homeostatic property cluster kinds) model grouping practices in the life sciences accurately and generally. Such concepts fit reasoning into particular preconceived epistemic and semantic patterns. The ability of these patterns to fit scientific practice is often argued in support of homeostatic property cluster accounts, yet there are reasons to think that in the life sciences kind concepts exhibit (...) a diversity of grouping practices that are flattened out by conceptualizing them as natural kinds. Instead this article argues that the process of understanding grouping practices needs to start from a more neutral position independent of any ontological account. Following Love (Acta Biotheor 57:51–75, 2009) this paper suggests that typical natural kind concepts should be broached in the first place as grouping strategies that use a variety of semantic and epistemic tactics to apply group-bound information to tasks of explanation and understanding. (shrink)

A number of areas of biology raise questions about what is of value in the natural environment and how we ought to behave towards it: conservation biology, environmental science, and ecology, to name a few. Based on my experience teaching students from these and similar majors, I argue that the field of environmental ethics has much to teach these students. They come to me with pent-up questions and a feeling that more is needed to fully engage in their subjects, and (...) I believe some exposure to environmental ethics can help focus their interests and goals. I identify three primary areas in which environmental ethics can con- tribute to their education. The first is an examination of who (or what) should be considered to be part of our moral community (i.e., the community to whom we owe direct duties). Is it humans only? Or does it include all sentient life? Or all life? Or ecosystems considered holistically? Often, readings implicitly assume one or more of these answers; the goal is to make the student more sensitive to these implicit claims and to get them to think about the different reasons that support them. The second area, related to the first, is the application of the different answers concerning the extent of the ethical community to real environmental issues and problems. Students need to be aware of how the different answers concerning the moral community can imply conflicting answers for how we should act in certain cases and to think about ways to move toward conflict resolution. The third area in which environmental ethics can contribute is a more conceptual one, focusing on central concepts such as biodiversity, sustainability, species, and ecosystems. Exploring and evaluating various meanings of these terms will make students more reflective and thoughtful citizens and biologists, sensitive to the implications that different conceptual choices make. (shrink)

The concept of developmental constraint was at the heart of developmental approaches to evolution of the 1980s. While this idea was widely used to criticize neo-Darwinian evolutionary theory, critique does not yield an alternative framework that offers evolutionary explanations. In current Evo-devo the concept of constraint is of minor importance, whereas notions as evolvability are at the center of attention. The latter clearly defines an explanatory agenda for evolutionary research, so that one could view the historical shift from ‘developmental constraint’ (...) towards ‘evolvability’ as the move from a concept that is a mere tool of criticism to a concept that establishes a positive explanatory project. However, by taking a look at how the concept of constraint was employed in the 1980s, I argue that developmental constraint was not just seen as restricting possibilities (‘constraining’), but also as facilitating morphological change in several ways. Accounting for macroevolutionary transformation and the origin of novel form was an aim of these developmental approaches to evolution. Thus, the concept of developmental constraint was part of a positive explanatory agenda long before the advent of Evo-devo as a genuine scientific discipline. In the 1980s, despite the lack of a clear disciplinary identity, this concept coordinated research among paleontologists, morphologists, and developmentally inclined evolutionary biologists. I discuss the different functions that scientific concepts can have, highlighting that instead of classifying or explaining natural phenomena, concepts such as ‘developmental constraint’ and ‘evolvability’ are more important in setting explanatory agendas so as to provide intellectual coherence to scientific approaches. The essay concludes with a puzzle about how to conceptually distinguish evolvability and selection. (shrink)

Philosophers of science have often favoured reductive approaches to how-possibly explanation. This article identifies three alternative conceptions making how-possibly explanation an interesting phenomenon in its own right. The first variety approaches “how possibly X?” by showing that X is not epistemically impossible. This can sometimes be achieved by removing misunderstandings concerning the implications of one’s current belief system but involves characteristically a modification of this belief system so that acceptance of X does not result in contradiction. The second variety offers (...) a potential how-explanation of X. It is usually followed by a range of further potential how-explanations of the same phenomenon. In recent literature the factual claims implied by the second variety have been downplayed whereas the heuristic role of mapping the space of conceptual possibilities has been emphasized. I will focus especially on this truth-bracketing sense of potentiality when looking closer at the second variety in the paper. The third variety has attracted less interest. It presents a partial how-explanation of X. Typically it aims to establish the existence of a mechanism by which X could be and was generated. The third conception stands out as the natural alternative for the advocate of ontic how-possibly explanations. This article transfers Salmon’s view that explanation-concepts can be broadly divided into epistemic, modal, and ontic to the context of how-possibly explanations. Moreover, it is argued that each of the three above-mentioned varieties of how-possibly explanation occurs in science. To recognize this may be especially relevant for philosophers. We are often misled by the promises of various why-explanation accounts, and seem to have forgotten nearly everything about the diversity of how-possibly explanations. (shrink)

The Darwinian theory of evolution is itself evolving and this book presents the details of the core of modern Darwinism and its latest developmental directions. The authors present current scientific work addressing theoretical problems and challenges in four sections, beginning with the concepts of evolution theory, its processes of variation, heredity, selection, adaptation and function, and its patterns of character, species, descent and life. The second part of this book scrutinizes Darwinism in the philosophy of science and its usefulness in (...) understanding ecosystems, whilst the third section deals with its application in disciplines beyond the biological sciences, including evolutionary psychology and evolutionary economics, Darwinian morality and phylolinguistics. The final section addresses anti-Darwinism, the creationist view and issues around teaching evolution in secondary schools. The reader learns how current experimental biology is opening important perspectives on the sources of variation, and thus of the very power of natural selection. This work examines numerous examples of the extension of the principle of natural selection and provides the opportunity to critically reflect on a rich theory, on the methodological rigour that presides in its extensions and exportations, and on the necessity to measure its advantages and also its limits. Scholars interested in modern Darwinism and scientific research, its concepts, research programs and controversies will find this book an excellent read, and those considering how Darwinism might evolve, how it can apply to the human sciences and other disciplines beyond its origins will find it particularly valuable. Originally produced in French (Les Mondes Darwiniens), the scope and usefulness of the book have led to the production of this English text, to reach a wider audience. This book is a milestone in the impressive penetration by Francophone scholars into the world of Darwinian science, its historiography and philosophy over the last two decades. Alex Rosenberg, R. Taylor Cole Professor of Philosophy, Duke University Until now this useful and comprehensive handbook has only been available to francophones. Thanks to this invaluable new translation, this collection of insightful and original essays can reach the global audience it deserves. Tim Lewens, University of Cambridge. (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)

Over the past decade, it has been discovered that disparate aspects of morphology – often of distantly related groups of organisms – are regulated by the same genetic regulatory mechanisms. Those discoveries provide a new perspective on morphological evolutionary change. A conceptual framework for exploring these research findings is termed ‘deep homology’. A comparative framework for morphological relations of homology is provided that distinguishes analogy, homoplasy, plesiomorphy and synapomorphy. Four examples – three from plants and one from animals – demonstrate (...) that homologous developmental mechanisms can regulate a range of morphological relations including analogy, homoplasy and examples of uncertain homology. Deep homology is part of a much wider range of phenomena in which biological (genes, regulatory mechanisms, morphological traits) and phylogenetic levels of homology can both be disassociated. Therefore, to understand homology, precise, comparative, independent statements of both biological and phylogenetic levels of homology are necessary. (shrink)

Evolutionary developmental biology (evo-devo) is considered a ‘mechanistic science,’ in that it causally explains morphological evolution in terms of changes in developmental mechanisms. Evo-devo is also an interdisciplinary and integrative approach, as its explanations use contributions from many fields and pertain to different levels of organismal organization. Philosophical accounts of mechanistic explanation are currently highly prominent, and have been particularly able to capture the integrative nature of multifield and multilevel explanations. However, I argue that evo-devo demonstrates the need for a (...) broadened philosophical conception of mechanisms and mechanistic explanation. Mechanistic explanation (in terms of the qualitative interactions of the structural parts of a whole) has been developed as an alternative to the traditional idea of explanation as derivation from laws or quantitative principles. Against the picture promoted by Carl Craver, that mathematical models describe but usually do not explain, my discussion of cases from the strand of evo-devo which is concerned with developmental processes points to qualitative phenomena where quantitative mathematical models are an indispensable part of the explanation. While philosophical accounts have focused on the actual organization and operation of mechanisms, properties of developmental mechanisms that are about how a mechanism reacts to modifications are of major evolutionary significance, including robustness, phenotypic plasticity, and modularity. A philosophical conception of mechanisms is needed that takes into account quantitative changes, transient entities and the generation of novel types of entities, feedback loops and complex interaction networks, emergent properties, and, in particular, functional-dynamical aspects of mechanisms, including functional (as opposed to structural) organization and distributed, system-wide phenomena. I conclude with general remarks on philosophical accounts of explanation. (shrink)

Taxa and homologues can in our view be construed both as kinds and as individuals. However, the conceptualization of taxa as natural kinds in the sense of homeostatic property cluster kinds has been criticized by some systematists, as it seems that even such kinds cannot evolve due to their being homeostatic. We reply by arguing that the treatment of transformational and taxic homologies, respectively, as dynamic and static aspects of the same homeostatic property cluster kind represents a good perspective for (...) supporting the conceptualization of taxa as kinds. The focus on a phenomenon of homology based on causal processes (e.g., connectivity, activity-function, genetics, inheritance, and modularity) and implying relationship with modification yields a notion of natural kinds conforming to the phylogenetic-evolutionary framework. Nevertheless, homeostatic property cluster kinds in taxonomic and evolutionary practice must be rooted in the primacy of epistemological classification (homology as observational properties) over metaphysical generalization (series of transformation and common ancestry as unobservational processes). The perspective of individuating characters exclusively by historical-transformational independence instead of their developmental, structural, and functional independence fails to yield a sufficient practical interplay between theory and observation. Purely ontological and ostensional perspectives in evolution and phylogeny (e.g., an ideographic character concept and PhyloCode’s ‘individualism’ of clades) may be pragmatically contested in the case of urgent issues in biodiversity research, conservation, and systematics. (shrink)

It has been claimed that the intentional stance is necessary to individuate behavioral traits. This thesis, while clearly false, points to two interesting sets of problems concerning biological explanations of behavior: The first is a general in the philosophy of science: the theory-ladenness of observation. The second problem concerns the principles of trait individuation, which is a general problem in philosophy of biology. After discussing some alternatives, I show that one way of individuating the behavioral traits of an organism is (...) by a special use of the concept of biological function, as understood in an enriched causal role (not selected effect) sense. On this view, a behavioral trait is essentially a special kind of regularity, namely a regularity that is produced by some regulatory mechanism. Regulatory mechanisms always require goal states, which can only be provided by functional considerations. As an example from actual (as opposed to folk) science, I examine the case of social behavior in nematodes. I show that the attempt to explain this phenomenon actually transformed it. This supports the view that scientific explanation does not explain an explanandum phenomenon that is given prior to the explanation; rather, the explanandum is changed by the explanation. This means that there could be a plurality of stances that have some heuristic value initially, but which will be abandoned in favor of a functional characterization eventually. (shrink)

There have been repeated attempts in the history of comparative biology to provide a mechanistic account of morphological homology. However, it is well-established that homologues can develop from diverse sets of developmental causes, appearing not to share any core causal architecture that underwrites character identity. We address this challenge with a new conceptual model of Character Identity Mechanisms. ChIMs are cohesive mechanisms with a recognizable causal profile that allows them to be traced through evolution as homologues despite having a diverse (...) etiological organization. Our model hypothesizes that anatomical units at different levels of organization—cell types, tissues, and organs—have level-specific ChIMs with different conserved parts, activities, and organization. Relying on a methodology of conceptual engineering, we show how the ChIM concept advances our understanding of the developmental basis of morphological characters, while forging an important link between comparative and mechanistic biology. (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)

The Mind-Brain Identity Theory lived a short life as a respectable philosophical position in the late 1950s, until Hilary Putnam developed his famous argument on the multiple realizability of mental states. The argument was, and still is, taken as the definitive demonstration of the falsity of Identity Theory and the foundation on which contemporary functionalist computational cognitive science was to be grounded. In this paper, in the wake of some contemporary philosophers, we reopen the case for Identity Theory and offer (...) a solution to the problem of multiple realizabilty. The solution is based on the necessity, at the time of establishing identity relations, of appealing to the notions of “homology” and “analogy” developed in the nineteenth century by Richard Owen. We also suggest that these notions are useful in order to correct certain shortcomings of some recent attempts at rebutting the Multiple Realizability argument. (shrink)

Many of the current comparisons of taxic phylogenetic and biological homology in the context of morphology focus on what are seen as categorical distinctions between the two concepts. The first, it is claimed, identifies historical patterns of conservation and variation relating taxa; the second provides a causal framework for the explanation of this conservation and variation. This leads to the conclusion that the two need not be placed in conflict and are in fact compatible, having non-competing epistemic purposes or mapping (...) the same extensions in the form of monophyletic groupings (see Roth, The biological basis of homology 1–26, 1988; Sluys, J Zool Syst Evol Res 34:145–152, 1996; Abouheif, Trends Ecol Evol 12:405–408, 1997; Brigandt, J Exp Zool 299:9–17, 2003, Biol Philos 22:709–725, 2007; Assis and Brigandt, Evol Biol 36:248–255, 2009). This article argues that moves in this direction miss the essential disagreement between these concepts as they have been developed in the context of the debate concerning the best concept for evolutionary investigation. We should rather see these concepts employing a common fundamental methodological approach to homology, but disagreeing about how to apply the methodology effectively. Both concepts employ class reasoning, which pursues homologies as units of generalization—more precisely, as sources of reliable and relevant group-bound information in the form of shared underlying causes. The dispute can be better understood by two poles that structure such reasoning: the need for a reliable basis for projections about the causal history of shared structures, and the desire to identify homologous characters with more informative and specific causal information relevant to generalizing about evolutionary processes. Judgments in favor of one or the other in turn have affected the scope or extension of these competing homology concepts. (shrink)

I develop an account of homology and homoplasy drawing on their use in biological inference and explanation. Biologists call on homology and homoplasy to infer character states, support adaptationist explanations, identify evolutionary novelties and hypothesize phylogenetic relationships. In these contexts, the concepts must be understood phylogenetically and kept separate: as they play divergent roles, overlap between the two ought to be avoided. I use these considerations to criticize an otherwise attractive view defended by Gould, Hall, and Ramsey & Peterson. By (...) this view, homology and homoplasy can only be delineated qua some level of description, and some homoplasies (parallelisms) are counted as homologous. I develop an account which retains the first, but rejects the second, aspect of that view. I then characterize parallelisms and convergences in terms of their causal role. By the Strict Continuity account, homology and homoplasy are defined phylogenetically and without overlaps, meeting my restriction. Convergence and parallelisms are defined as two types of homoplasy: convergent homoplasies are largely constrained by external factors, while parallelisms are due to internal constraints. (shrink)

The scientific status of evolutionary theory seems to be more or less perennially under question. I am not referring here (just) to the silliness of young Earth creation- ism (Pigliucci 2002; Boudry and Braeckman 2010), or even of the barely more intel- lectually sophisticated so-called Intelligent Design theory (Recker 2010; Brigandt this volume), but rather to discussions among scientists and philosophers of science concerning the epistemic status of evolutionary theory (Sober 2010). As we shall see in what follows, this debate (...) has a long history, dating all the way back to Darwin, and it is in great part rooted in the fundamental dichotomy between what French biologist and Nobel laureate Jacques Monod (1971) called chance and necessity—i.e., the inevitable and inextricable interplay of deterministic and stochastic mechanisms operating during the course of evolution. (shrink)

Against the neo-Darwinian assumption that genetic factors are the principal source of variation upon which natural selection operates, a phenotype-first hypothesis strikes us as revolutionary because development would seem to constitute an independent source of variability. Richard Watson and his co-authors have argued that developmental memory constitutes one such variety of phenotypic variability. While this version of the phenotype-first hypothesis is especially well-suited for the late metazoan context, where animals have a sufficient history of selection from which to draw, appeals (...) to developmental memory seem less plausible in the evolutionary context of the early metazoans. I provide an interpretation of Stuart Newman’s account of deep metazoan phylogenesis that suggests that spandrels are, in addition to developmental memory, an important reservoir of phenotypic variability. I conclude by arguing that Gerd Müller’s “side-effect hypothesis” is an illuminating generalization of the proposed non-Watsonian version of the phenotype-first hypothesis. (shrink)

Il concetto di omologia riveste un ruolo centrale nella biologia teoretica contemporanea perché attorno ad esso e per suo tramite trovano articolazione numerosi progetti di ricerca miranti a ripensare la centralità dell’organismo, i suoi percorsi evolutivi e il significato della sua interazione con la realtà ambientale. La rete di omologie che tesse l’organismo permette di strutturarne e canalizzarne le innovazioni morfologiche e funzionali. Appunto questa componente sistemica rende dunque l’omologia un concetto fondamentale per una considerazione dinamica della forma vivente, come (...) quella cui mira un’estetica che non intenda limitarsi a indagare sull’atteggiamento estetico e sulle produzioni artistiche, ma sappia tematizzare in senso più comprensivo la questione (eminentemente propria della tradizione morfologica) dell’unità dell’agire e del percepire come configurazione estetica del vivente.Il presente studio propone una ricognizione di alcuni dei principali momenti fondativi del concetto di omologia, a partire dalle ricerche botaniche di Goethe e dal significato che esse rivestono nel progetto morfologico goethiano, per proseguire con l’elaborazione del concetto di “elemento anatomico” da parte di Geoffroy Saint-Hilaire e la sua problematica rilettura da parte di Arthur Schopenhauer, conducendo sino alla fondazione del concetto moderno di omologia con Richard Owen, alla sua messa in relazione col concetto di analogia e all’elaborazione del concetto di “archetipo del vertebrato”. In tal modo, il presente studio mira a individuare alcuni motivi salienti di attualità del concetto e delle potenzialità teoriche che in esso trovano espressione, fra prospettiva morfologica e attenzione ai vincoli materiali della forma. (shrink)

While philosophers tend to consider a single type of causal history, biologists distinguish between two kinds of causal history: evolutionary history and developmental history. This essay studies the peculiarity of development as a criterion for the individuation of biological traits and its relation to form, function, and evolution. By focusing on examples involving serial homologies and genetic reprogramming, we argue that morphology (form) and function, even when supplemented with evolutionary history, are sometimes insufficient to individuate traits. Developmental mechanisms bring in (...) a novel aspect to the business of classification—identity of process-type—according to which entities are type-identical across individuals and natural kinds in virtue of the fact that they form and develop through similar processes. These considerations bear important metaphysical implications and have potential applications in several areas of philosophy. (shrink)

Evo-Devo exhibits a plurality of scientific “cultures” of practice and theory. When are the cultures acting—individually or collectively—in ways that actually move research forward, empirically, theoretically, and ethically? When do they become imperialistic, in the sense of excluding and subordinating other cultures? This chapter identifies six cultures – three /styles/ (mathematical modeling, mechanism, and history) and three /paradigms/ (adaptationism, structuralism, and cladism). The key assumptions standing behind, under, or within each of these cultures are explored. Characterizing the internal structure of (...) the cultures is necessary for understanding how they collaborate or compete, and how they are fragmented or integrated, in the rich interdisciplinary /trading zone/ (Galison 1997) of Evo-Devo. Evo-Devo is an important example of how science can progress through a radical plurality of perspectives and cultures. (shrink)

I saggi raccolti in questo fascicolo mirano a comprendere la polarità omologia/analogia e ad analizzarne gli usi nei diversi ambiti concettuali in cui essa ricorre nel dibattito contemporaneo in filosofia, nelle scienze della vita, nei discorsi sulle arti. La storia di questa coppia concettuale mostra la capacità pervasiva di costruire relazioni tra i diversi oggetti della conoscenza e, cosa ancor più interessante, la possibilità di raggiungere una sintesi concettuale fra approcci metodologic...

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)