Organisms change their shape and behavior during ontogenesis in response to incentives—what biologists call “phenotypic plasticity” or what is called here more specifically “behavioral plasticity.” Such plasticity is usually in the direction of enhancing welfare or fitness. In light of basic concepts in economics, such behavioral plasticity is nothing but rationality. Such rationality is not limited to organisms with neural systems. It also characterizes brainless organisms such as plants, fungi, and unicellular organisms. The gist of the article is the distinction (...) between rationality and intelligence. Whereas rationality is ubiquitous in all organisms, intelligence varies in degrees depending on division of labor that may involve the evolution of a neural system. This article aims to defend the universality of rationality—the Organismus oeconomicus hypothesis. It argues that neither the notion of bounded rationality of behavioral economics nor Herbert Simon’s “procedural rationality” can ultimately undermine the Organismus oeconomicus hypothesis. (shrink)

In this introduction, we give a brief overview of the main concepts of modularity that have been offered in recent literature. After this, we turn to a summary of the papers collected in this volume. Our primary aim is to explain how the modularity of emotion question relates to traditional debates in emotion theory.

The growing interest and major advances of the last decades in evolutionary developmental biology (EvoDevo) have led to the recognition of the incompleteness of the Modern Synthesis of evolutionary theory. Here we discuss how paleontology makes significant contributions to integrate evolution and development. First, extinct organisms often inform us about developmental processes by showing a combination of features unrecorded in living species. We illustrate this point using the vertebrate fossil record and studies relating bone ossification to life history traits. Second, (...) we discuss exceptionally preserved fossils that document rare ontogenetic sequences and illustrate this case with the patterns of heterochrony observed in Cambrian crustacean larvae preserved three-dimensionally. Third, most fossils potentially document the evolutionary patterns of allometry and modularity, as well as some of the (paleo)ecological factors that had influenced them. The temporal persistence of adaptive patterns in rodent evolution serves to address the importance of ecological constraints in evolution. Fourth, we discuss how the macroevolutionary patterns observed in the tetrapod limb, in the mammal molar proportions, and in the molluscan shell provide independent tests of the validity of morphogenetic models proposed on living species. Reciprocally, these macroevolutionary patterns often act as a source of inspiration to investigate the underlying rules of development, because, at the end, they are the patterns that the neo-Darwinian theory was unable to account for. (shrink)

In a macroevolutionary timescale, evolvability itself evolves. Lineages are sorted based on their ability to generate adaptive novelties, which leads to the optimization of their genotype-phenotype map. The system of translation of genetic or epigenetic changes to the phenotype may reach significant horizontal and vertical complexity, and may even exhibit certain aspects of learning behaviour. This continuously evolving semiotic system probably enables the origin of complex yet functional and internally compatible adaptations. However, it also has a second, “darker”, side. As (...) was pointed out by several authors, the same process gradually reduces the probability of the origination of significant evolutionary novelties. In a similar way to the evolution of societies, teachings, or languages, in which the growing number of internal linkages gradually solidifies their overall structure and the structure or interpretation of their constitutive elements, the evolutionary potential of lineages decreases during biological evolution. Possible adaptations become limited to small “peripheral” modifications. According to the Frozen Evolution theory, some of the proximate causes of this “macroevolutionary freezing” are more pronounced or present exclusively in sexual lineages. Sorting based on the highest evolvability probably leads to the establishment of certain structural features of complex organisms, e.g. the modular character of their development and morphology. However, modules also “macroevolutionary freeze” whereas the hypothetical “thawing” of modules or their novel adaptive combinations becomes rarer and rarer. Some possible ways out of this dead end include the rearrangement of individual development, e.g. neoteny, radical simplification, i.e. sacculinization, and transition to a higher level of organization, e.g. symbiosis or symbiogenesis. The evolution of evolvability is essentially a biosemiotic process situated at the intersection of the genocentric modern synthesis and the evo-devo-centric extended synthesis. Therefore, evolvability may eventually connect these three not necessarily contradictory approaches. (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)

What is ‘counterintuitive’? There is general agreement that it refers to a violation of previously held knowledge, but the precise definition seems to vary with every author and study. The aim of this paper is to deconstruct the notion of ‘counterintuitive’ and provide a more philosophically rigorous definition congruent with the history of psychology, recent experimental work in ‘minimally counterintuitive’ concepts, the science vs. religion debate, and the developmental and evolutionary background of human beings. We conclude that previous definitions of (...) counterintuitiveness have been flawed and did not resolve the conflict between a believer’s conception of the supernatural entity (an atypical “real kind”) and the non-believer’s conception (empty name/fictional). Furthermore, too much emphasis has been placed on the universality and (presumed) innateness of intuitive concepts (and hence the criteria for what is counterintuitive)—and far too little attention paid to learning and expertise. We argue that many putatively universal concepts are not innate, but mostly learned and defeasible—part of a religious believer’s repertoire of expert knowledge. Nonetheless, the results from empirical studies about the memorability of counterintuitive concepts have been convincing and it is difficult to improve on existing designs and methodologies. However, future studies in counterintuitive concepts need to embed their work in research about context effects, typicality, the psychology of learning and expertise (for example, the formation of expert templates and range defaults), with more attention to the sources of knowledge (direct and indirect knowledge) and a better idea of what ‘default’ knowledge really is. (shrink)

More than one researcher is currently proposing that the notion of information become an important element for defining living systems as well as for explaining conditions that make their origins possible. During the pre-biotic era, the type of compounds encountered would mainly have been very simple in nature and might have been immersed in the natural dynamic of the physical world and in processes of self-organization. It is furthermore quite possible that they formed a relationship between and among certain types (...) of processes that here we are specifically proposing as central for the emergence of cell organization. Consequently, an important initial step towards constructing a theory of biological information is to ask ourselves the question: how do biological systems process information? In this way, we will be contributing to the proposals of this paper where we seek to identify general principles that govern biological computing and that deal with biosemiotic approaches as they are defended in naturalistic normative terms. (shrink)

Morphological EvoDevo is a field of biological inquiry in which explicit relations between evolutionary patterns and growth or morphogenetic processes are made. Historically, morphological EvoDevo results from the coming together of several traditions, notably Naturphilosophie, embryology, the study of heterochrony, and developmental constraints. A special feature binding different approaches to morphological EvoDevo is the use of formalisms and mathematical models. Here we will introduce anatomical network analysis, a new approach centered on connectivity patterns formed by anatomical parts, with its own (...) concepts and tools specifically designed for the study of morphological EvoDevo questions. Riedl’s concept of burden is tightly related to the use of anatomical networks, providing a nexus between the evolutionary patterns and the structural constraints that shape them. (shrink)

Evolutionary systems biology is an emerging hybrid approach that integrates methods, models, and data from evolutionary and systems biology. Drawing on themes that arose at a cross-disciplinary meeting on ESB in 2013, we discuss in detail some of the explanatory friction that arises in the interaction between evolutionary and systems biology. These tensions appear because of different modeling approaches, diverse explanatory aims and strategies, and divergent views about the scope of the evolutionary synthesis. We locate these discussions in the context (...) of long-running philosophical deliberations on explanation, modeling, and theoretical synthesis. We show how many of the issues central to ESB’s progress can be understood as general philosophical problems. The benefits of addressing these philosophical issues feed back into philosophy too, because ESB provides excellent examples of scientific practice for the development of philosophy of science and philosophy of biology. (shrink)

Rudolf Leuckart's 1851 pamphlet Ueber den Polymorphismus der Individuen (On the polymorphism of individuals) stood at the heart of naturalists' discussions on biological individuals, parts and wholes in mid-nineteenth-century Britain and Europe. Our analysis, which accompanies the first translation of this pamphlet into English, situates Leuckart's contribution to these discussions in two ways. First, we present it as part of a complex conceptual knot involving not only individuality and the understanding of compound organisms, but also the alternation of generations, the (...) division of labor in nature, and the possibility of finding general laws of the organic world. Leuckart's pamphlet is important as a novel attempt to give order to the strands of this knot. It also solved a set of key biological problems in a way that avoided some of the drawbacks of an earlier teleological tradition. Second, we situate the pamphlet within a longer trajectory of inquiry into part-whole relations in biology from the mid-eighteenth century to the present. We argue that biological individuality, along with the problem-complexes with which it engaged, was as central a problem to naturalists before 1859 as evolution, and that Leuckart's contributions to it left a long legacy that persisted well into the twentieth century. As biologists' interests in part-whole relations are once again on the upswing, the longue durée of this problem merits renewed consideration. (shrink)

Rudolf Leuckart’s 1851 pamphlet Ueber den Polymorphismus der Individuen stood at the heart of naturalists’ discussions on biological individuals, parts and wholes in mid-nineteenth-century Britain and Europe. Our analysis, which accompanies the first translation of this pamphlet into English, situates Leuckart’s contribution to these discussions in two ways. First, we present it as part of a complex conceptual knot involving not only individuality and the understanding of compound organisms, but also the alternation of generations, the division of labor in nature, (...) and the possibility of finding general laws of the organic world. Leuckart’s pamphlet is important as a novel attempt to give order to the strands of this knot. It also solved a set of key biological problems in a way that avoided some of the drawbacks of an earlier teleological tradition. Second, we situate the pamphlet within a longer trajectory of inquiry into part-whole relations in biology from the mid-eighteenth century to the present. We argue that biological individuality, along with the problem-complexes with which it engaged, was as central a problem to naturalists before 1859 as evolution, and that Leuckart’s contributions to it left a long legacy that persisted well into the twentieth century. As biologists’ interests in part-whole relations are once again on the upswing, the longue durée of this problem merits renewed consideration. (shrink)

No comprehensive theory of development is available yet. Traditionally, we regard the development of animals as a sequence of changes through which an adult multicellular animal is produced, starting from a single cell which is usually a fertilized egg, through increasingly complex stages. However, many phenomena that would not qualify as developmental according to these criteria would nevertheless qualify as developmental in that they imply nontrivial (e.g., non degenerative) changes of form, and/or substantial changes in gene expression. A broad, comparative (...) approach is badly needed. In the Cnidaria, for example, even the boundary between generations is problematic. Describing their life cycle in terms of metagenesis (alternation between polyp generation and medusa generation) or in terms of metamorphosis (polyp as larva or juvenile) are matters of semantics more than biology. The life cycle of other metazoans, described in textbooks in terms of larva-to-adult metamorphosis, is hardly different from a typical metagenetic life cycle of cnidarians. This applies to holometabolous insects and to marine invertebrates like sea urchins, where most of the larval cells are discarded at metamorphosis. The uncertain temporal and spatial boundaries of individual development are also shown by the widespread lack of a strict correspondence between adult and mature. A comprehensive theory of development should start with a zero principle of “developmental inertia,” corresponding to an indeterminate local self-perpetuation of cell-level dynamics. Indeterminate growth, scale-invariance, segmentation, and regeneration provide examples of developmental dynamics close to that. (shrink)

In this paper, we argue that several recent ‘wide’ perspectives on cognition (embodied, embedded, extended, enactive, and distributed) are only partially relevant to the study of cognition. While these wide accounts override traditional methodological individualism, the study of cognition has already progressed beyond these proposed perspectives towards building integrated explanations of the mechanisms involved, including not only internal submechanisms but also interactions with others, groups, cognitive artifacts, and their environment. The claim is substantiated with reference to recent developments in the (...) study of “mindreading” and debates on emotions. We claim that the current practice in cognitive (neuro)science has undergone, in effect, a silent mechanistic revolution, and has turned from initial binary oppositions and abstract proposals towards the integration of wide perspectives with the rest of the cognitive (neuro)sciences. (shrink)

If all origins of life or of any new grade, level, or major transition as such begin with “competitive development”—with juveniles rather than adults, and multiple individuals rather than a single one—then the evolution of progeneration and of replication always requires an explanation. This article proposes that principles of evolutionary ecology such as density-dependence can be used to explain three kinds of developmental repetitions, viz., sequences of inductive and niche-constructing interactions between the ecological environment and population members, which take place (...) in such a way that the sequence is repeated: individual or somatic repetitions, discussed in Section 1, demographic or progenerative repetitions , discussed in Section 2, and replicative repetitions , discussed in Section 3. This results in a statement of the evolutionary process that includes rather than excludes development and ecology, but one which requires some additions to Van Valen’s aphorism that evolution is the control of development by ecology. (shrink)

There is a trend within philosophy of biology to concentrate on questions that are strongly related to particular biological research programs rather than on the general scope of the field and its relation to other sciences. Projects of the latter kind, of course, are followed as well but will not be the topic of this review. Shifting the focus to particular research programs reflects philosophers’ increased interest in knowledge of, and contribution to, actual biological research, which is organized in such (...) programs. It is accompanied by the increasing enthusiasm of biologists to involve philosophers in the conceptual work of theoretical biology. I concentrate on the philosophies of four biological research programs, three of which are devoted to evolutionary biology, which is still the main field of interest among philosophers of biology: adaptationism, EvoDevo, and the developmental systems approach. In addition, a short sketch of the newly emerging philosophy of systems biology is given. Several lines of philosophical inquiry can be found in all of the fields considered here: philosophy contributes to the conceptual development of biological research programs, it analyzes structures and delineations of particular research programs, and sometimes it is involved in a comparative assessment of biological programs as well. Philosophical projects that start from the level of a particular research program may give rise to a bottom-up perspective on biology and allow for an integrative view of biological research. This may open up the opportunity to tackle “larger” questions again in an altered and fruitful manner. (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)

First the ‘Weismann barrier’ and later on Francis Crick’s ‘central dogma’ of molecular biology nourished the gene-centric paradigm of life, i.e., the conception of the gene/genome as a ‘central source’ from which hereditary specificity unidirectionally flows or radiates into cellular biochemistry and development. Today, due to advances in molecular genetics and epigenetics, such as the discovery of complex post-genomic and epigenetic processes in which genes are causally integrated, many theorists argue that a gene-centric conception of the organism has become problematic. (...) Here, we first explore the causal implications of the following two central dogma-related issues: (1) widespread reverse transcription—arguing for an extension from ‘DNA-genome’ to RNA-encompassing ‘NA-genome’ and, thus, from traditional DNA-centrism to a broader ‘NA-centrism’; and (2) the absence of a mechanism of reverse translation—arguing for the ‘structural primacy’ of NA-sequence over protein in cellular biochemistry. Secondly, we explore whether this latter conclusion can be extended to a ‘functional primacy’ of NA-sequence over protein in cellular biochemistry, which would imply a limited kind of ‘gene/NA-centrism’ confined to the subcellular level of NA/protein-based biochemistry. Finally, we explore the conditions—and their (non)fulfilment—for a more generalised form of gene-centrism extendable to higher levels of biological organisation. We conclude that the higher we go in the biological hierarchy, the more dubious gene-centric claims become. (shrink)

This is the second of two articles in which I reflect on “generalized Darwinism” as currently discussed in evolutionary economics. In the companion article (Callebaut, Biol Theory 6. doi: 10.1007/s13752-013-0086-2, 2011, this issue) I approached evolutionary economics from the naturalistic perspectives of evolutionary epistemology and the philosophy of biology, contrasted evolutionary economists’ cautious generalizations of Darwinism with “imperialistic” proposals to unify the behavioral sciences, and discussed the continued resistance to biological ideas in the social sciences. Here I assess Generalized Darwinism (...) as propounded by Geoffrey Hodgson, Thorbjørn Knudsen, and others, concentrating on the roles of theory and model building in science (and the roles of analogy and metaphor therein), generative replication, and the relation between selection and self-organization. I then point to advances in current biology that promise to be more fruitful as sources of inspiration for evolutionary economics than the project to generalize Darwinism in its current, “hardened Modern Synthesis” form; and I draw some conclusions. (shrink)

There are many things that philosophy of biology might be. But, given the existence of a professional philosophy of biology that is arguably a progressive research program and, as such, unrivaled, it makes sense to define philosophy of biology more narrowly than the totality of intersecting concerns biologists and philosophers (let alone other scholars) might have. The reasons for the success of the “new” philosophy of biology remain poorly understood. I reflect on what Dutch and Flemish, and, more generally, European (...) philosophers of biology could do to improve the situation of their discipline locally, regionally, and internationally, paying particular attention to the lessons to be learned from the “Science Wars.”. (shrink)

Complexity in our universe, Herbert Simon once noted, generally takes a hierarchical, nearly decomposable form. If our purpose as biologists is to "carve Nature at the joints," then the quantitative biologist's pattern questions must embody some tentative claim of where the explanatory joints are—only after meaningful qualifications can notions of variance and covariance make sense. In morphometrics, specimens and variables alike can be "carved at the joints," with a correspondingly great gain in explanatory power in both versions. Simon's advice is (...) that the competent biologist's measurements should lie entirely within a single organismal component or else deal entirely with one of the joints. In either context, our best contemporary rhetorics of explanation in biology may resemble morphometrics in their frank combination of carefully supervised parallel quantifications that, taken together, result in new qualifications, leading in turn to new quantifications, and so on. In short, the relation between qualitative and quantitative in the organismal biological sciences is not an opposition but a complementarity, and the modern biometrical statistics of organismal form may be a particularly apposite praxis for exploring it. (shrink)

Evolutionary developmental biologists categorize many different kinds of things, from ontogenetic stages to modules of gene activity. The process of categorization—the establishment of “kinds”—is an implicit part of describing the natural world in consistent, useful ways, and has an essentially practical rather than philosophical basis. Kinds commonly serve one of three purposes: they may function (1) as practical tools for communication; (2) to support prediction and generalization; or (3) as a basis for theoretical discussions. Beyond the minimal requirement that classifications (...) reflect biological reality, what sorts of kinds or classification will be useful in advancing a research program depends on the epistemological context. Thus, the important meaning of “natural” in “natural kinds” is not “natural with respect to nature,” but “natural with respect to the question.” This conclusion arises from the recognition that the proper role of concepts (e.g. natural kind, module, homology, model) is not to answer biological questions, but rather to help frame them. From a scientist’s perspective, arguing about the wording (or existence) of a single definition of “natural” or “kind” is beside the point: we get more work done by letting the question at hand determine what kinds of kinds are natural, on the basis of their ability to help answer it. We should be content to let “natural kinds” remain vague, multivalent, and–therefore broadly useful. For a philosopher like Hacking, the diverse, disparate, and ultimately incommensurable uses of the term “natural kind” have diluted its value so far that it loses all meaning. For practicing scientists, however, defining useful kinds in the context of particular questions and systems remains a productive epistemological strategy. (shrink)

Recent advances in genetics and neurobiology have greatly increased the degree of variation that one finds in what is taken to provide the biological foundations of our species-specific linguistic capacities. In particular, this variation seems to cast doubt on the purportedly homogeneous nature of the language faculty traditionally captured by the concept of “Universal Grammar.” In this article we discuss what this new source of diversity reveals about the biological reality underlying Universal Grammar. Our discussion leads us to support (1) (...) certain hypotheses advanced in evolutionary developmental biology that argue for the existence of robust biological mechanisms capable of canalizing variation at different levels, and (2) a bottom-up perspective on comparative cognition. We conclude by sketching future directions for what we call “comparative biolinguistics,” specifying which experimental directions may help us succeed in this new research avenue. (shrink)

Archaeologists have proposed that behavioral knowledge of a tool can be conceptualized as a “recipe”—a unit of cultural transmission that combines the preparation of raw materials, construction, and use of the tool, and contingency plans for repair and maintenance. This parallels theories in cognitive psychology that behavioral knowledge is hierarchically structured—sequences of actions are divided into higher level, partially independent subunits. Here we use an agent-based simulation model to explore the costs and benefits of hierarchical learning relative to holistic learning, (...) where entire behavioral sequences are learned in an all-or-nothing fashion, and diffusionist learning, where actions are completely independent. Hierarchical learning is favored under the reasonable assumptions that learning is associated with some degree of both error and cost, and that behavior can be grouped into subunits that repeat in one or more tool recipes. These general predictions can be tested in the archaeological and ethnographic record. Recent advances in evolutionary developmental biology have revealed a number of parallels between the hierarchically structured, recipe-like organization of behavioral knowledge that we examine here and the manner in which biological organisms develop. (shrink)

The phylogenetic trees reconstructed from molecular data have led to the discovery that all living creatures belong to three primary kingdoms, or domains, because there are three types of cells in nature. The primary kingdoms are referred to as Archaea, Bacteria, and Eukaya, and their first representatives were the first modern cells that appeared on Earth. All known cells, on the other hand, contain a virtually universal genetic code, and this implies that the code evolved in a population of primitive (...) systems that preceded the first modern cells and is collectively known as the common ancestor of all life. This gives us the problem of understanding how the descendants of the common ancestor gave origin to the first modern cells. In this article it is argued that the appearance of the genetic code allowed the ancestral systems to translate genes into specific proteins, but their behavior was still ambiguous because they were unable to produce specific responses to the signals from the environment. To that purpose they needed to evolve signal processing codes, and here it is proposed that the development of these codes was a crucial step in the evolution of the first modern cells. (shrink)

An important strategy in the discovery of biological mechanisms involves the piecing together of experimental results from interventions. However, if mechanisms are investigated by means of ideal interventions, as defined by James Woodward and others, then the kind of information revealed is insufficient to discriminate between modular and non-modular causal contributions. Ideal interventions suffice for constructing webs of causal dependencies that can be used to make some predictions about experimental outcomes, but tell us little about how causally relevant factors are (...) organized together and how they interact with each other in order to produce a phenomenon. I argue that lab research relies on more elaborated types of interventions targeting in a controlled fashion multiple variables at the same time in order to probe the temporal organization of causally-relevant factors along distinct causal pathways and to test for non-modular interaction effects, thus providing crucial spatial-temporal constraints guiding the formulation of more detailed mechanistic explanations. (shrink)

The purpose of this article is to update and defend syntax-based gene concepts. I show how syntax-based concepts can and have been extended to accommodate complex cases of processing and gene expression regulation. In response to difficult cases and causal parity objections, I argue that a syntax-based approach fleshes out a deflationary concept defining genes as genomic sequences and organizational features of the genome contributing to a phenotype. These organizational features are an important part of accepted molecular explanations, provide the (...) theoretical basis for a large number of experimental techniques and practical applications, and play a crucial role in in annotating the genome, deriving predictions and constructing bioinformatics models. (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)

The advent of contemporary evolutionary theory ushered in the eventual decline of Aristotelian Essentialism (Æ) – for it is widely assumed that essence does not, and cannot have any proper place in the age of evolution. This paper argues that this assumption is a mistake: if Æ can be suitably evolved, it need not face extinction. In it, I claim that if that theory’s fundamental ontology consists of dispositional properties, and if its characteristic metaphysical machinery is interpreted within the framework (...) of contemporary evolutionary developmental biology, an evolved essentialism is available. The reformulated theory of Æ offered in this paper not only fails to fall prey to the typical collection of criticisms, but is also independently both theoretically and empirically plausible. The paper contends that, properly understood, essence belongs in the age of evolution. (shrink)

Evolutionary developmental biology (or developmental evolution) is in the middle stages of its “development.” Its early ontogeny cannot be traced back to fertilization but pivotal developmental events included Gould’s (1977) treatment of heterochrony, Riedl’s (1978) analysis of “burden”, the Dahlem conference of 1981, a British Society of Developmental Biologists Symposium, as well as books that incorporated developmental genetics into older comparative themes. A major inductive process began with the discovery of widespread phylogenetic conservation in homeobox-containing genes. One interpretation of these (...) early developmental dynamics holds that an increasingly prominent problem agenda (e.g., how do novel structures and functions originate?), inherited from long-standing research programs in comparative embryology, morphology, and paleontology, was handed a powerful set of experimental tools from the lineage of experimental embryology. This allowed for an experimental probing of development and its evolution in an unprecedented fashion. Although the relationship between evolution and development has exhibited a protracted and complex history, we are still in the process of comprehending these early stages of “modern” EvoDevo’s ontogeny. (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)

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)

According to conventional wisdom, local gauge symmetry is not a symmetry of nature, but an artifact of how our theories represent nature. But a study of the so-called theta-vacuum appears to refute this view. The ground state of a quantized non-Abelian Yang-Mills gauge theory is characterized by a real-valued, dimensionless parameter theta—a fundamental new constant of nature. The structure of this vacuum state is often said to arise from a degeneracy of the vacuum of the corresponding classical theory, which degeneracy (...) allegedly arises from the fact that “large” (but not “small”) local gauge transformations connect physically distinct states of zero field energy. If that is right, then some local gauge transformations do generate empirical symmetries. In defending conventional wisdom against this challenge I hope to clarify the meaning of empirical symmetry while deepening our understanding of gauge transformations. I distinguish empirical from theoretical symmetries. Using Galileo’s ship and Faraday’s cube as illustrations, I say when an empirical symmetry is implied by a theoretical symmetry. I explain how the theta-vacuum arises, and how “large” gauge transformations differ from “small” ones. I then present two analogies from elementary quantum mechanics. By applying my analysis of the relation between empirical and theoretical symmetries, I show which analogy faithfully portrays the character of the vacuum state of a classical non-Abelian Yang-Mills gauge theory. The upshot is that “large” as well as “small” gauge transformations are purely formal symmetries of non-Abelian Yang-Mills gauge theories, whether classical or quantized. It is still worth distinguishing between these kinds of symmetries. An analysis of gauge within the constrained-Hamiltonian formalism yields the result that “large” gauge transformations should not be classified as gauge transformations; indeed, nor should “global” gauge transformations. In a theory in which boundary conditions are modeled dynamically, “global” gauge transformations may be associated with physical symmetries, corresponding to translations of these extra dynamical variables. Such translations are symmetries if and only if charge is conserved. But it is hard to argue that these symmetries are empirical, and in any case they do not correspond to any constant phase change in a quantum state. (shrink)

Biochemists often adopt what may be called the “Strategy of Decomposition” for the causal discovery of biochemical pathway dynamic behaviours. This involves decomposing a pathway into a set of isolated parts, which are then analysed separately. It is assumed that knowledge gained of the isolated parts can then be used to explain the dynamic behaviours of the whole pathway. My thesis addresses the extent to which use of the Strategy of Decomposition is warranted. I evaluate two challenges contained in Bechtel (...) and Richardson’s Discovering Complexity. The first challenge is that pathways lack the ‘modular’ structure assumed in the Strategy of Decomposition. Bechtel and Richardson take biochemists to use a concept of modularity called ‘near decomposability’. The second challenge is that pathways have ‘Pathway Emergent’ behaviours. I reject both challenges. I show that near decomposability is the wrong type of modularity to apply to pathways, and that the occurrence of Pathway Emergence has not been established. I argue that an underlying problem with Bechtel and Richardson’s analyses is that they overstate the consequences of feedback and nonlinearity for the Strategy of Decomposition. Instead, the analysis of pathway modularity and emergence needs to be centered on the context-sensitivity of pathways’ ‘local causal laws’. I identify that the type of modularity assumed in the Strategy of Decomposition is ‘causal law modularity’, which requires the invariance of local causal laws. I also identify a necessary condition for Pathway Emergence: a pathway must manifest at least one local causal law that is not manifested by its isolated parts. I argue that the use of the Strategy of Decomposition may often be unwarranted. This is because the local causal laws of pathways are highly context-sensitive, and pathways might often not be causal law modular. This context-sensitivity also leaves open the possibility of Pathway Emergence. (shrink)

How can we explain the intentional nature of an expert’s actions, performed without immediate and conscious control, relying instead on automatic cognitive processes? How can we account for the differences and similarities with a novice’s performance of the same actions? Can a naturalist explanation of intentional expert action be in line with a philosophical concept of intentional action? Answering these and related questions in a positive sense, this dissertation develops a three-step argument. Part I considers different methods of explanations in (...) cognitive neuroscience (Bennett & Hacker’s philosophical, conceptual analysis; Marr’s three levels of explanation; Neural Correlates of Consciousness research; mechanistic explanation), defending ‘mechanistic explanation’ as a method that provides the necessary tools for integrating interdisciplinary insights into human action. Furthermore, a dynamic, explanatory mechanism allows the assessment of the impact of learning and development on expert action in a valuable way that other methods don’t. Part II continues by scrutinizing several cognitive neuroscientific theories of learning and development (neuroconstructivism; dual-processing theories; simulation theory; extended mind/cognition hypothesis), arguing for the complex interactions between different types of processing and different action representations involved in expert action performances. Moreover, according to our discussion of a particular ‘simulation theory’ these interactions can be influenced in several ways with the use of language, allowing an agent to configure a specific action representation for performance at a later stage. The results of Parts I and II are then applied in Part III to a parallel discussion of philosophical analyses of intentional action (discussing i.a. Frankfurt, Bratman, Pacherie and Ricoeur) and cognitive neuroscientific insights in it. Both approaches are found to converge in emphasizing the importance for an expert to develop stable patterns of actions that comply maximally not only with his intentions, but also with his motor expertise and with situational conditions. Consequently, his actions – automatic, or not – rely on this ‘sculpted space of actions’. (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)

In this paper, I suggest that the notion of module explicitly defined by Peter Carruthers in The Architecture of The Mind (Carruthers 2006) is not really In use in the book. Instead, a more robust notion seems to be actually in play. The more robust notion, albeit implicitly assumed, seems to be far more useful for making claims about the modularity of mind. Otherwise, the claims would become trivial. This robust notion will be reconstructed and improved upon by putting it (...) into a more general framework of mental architecture. I defend the view that modules are the outcome of structural rather than functional decomposition and that they should be conceived as near decomposable systems. (shrink)