The meaning of niche and competitive ability have long been surrounded by controversy. The reason for this stems from the obscure relationship that exists between these terms. This extends from the views of Darwin through Eltonian tradition to current views in which the meaning of competitive ability is implicitly infused into the paradigm of niche. Distinct operational definitions for niche and competitive ability are therefore established with special reference to plants. It is proposed that potential niche refer explicitly to a (...) theoretical hyperspace of places where a species would leave descendents if all biotic interactions were precluded, and that competitive ability refer to the relative capacity to leave descendents in a particular place in the face of restrictions imposed by competitive interaction. This leads to a qualitative comprehensive theory for coexistence which may be extended to any type of biotic interaction. Niche and competitive ability are both determined by the biological attributes of a species and may be independently adjusted in a population by natural selection in contexts of competition. Species coexistence in nature may therefore be a consequence of alternative evolutionary mechanisms which may operate to various degrees in concert: (1) natural selection leading to niche differentiation; (2) an ongoing process of reciprocal selection (coevolution) which maintains an approximate balance in relative competitive abilities for contested resources. (shrink)

The so-called "adaptationism" of mainstream evolutionary biology has been criticized from a variety of sources. One, which has received relatively little philosophical attention, is developmental biology. Developmental constraints are said to be neglected by adaptationists. This paper explores the divergent methodological and explanatory interests that separate mainstream evolutionary biology from its embryological and developmental critics. It will focus on the concept of constraint itself; even this central concept is understood differently by the two sides of the dispute.

Historians of Biology have divided nineteenth century naturalists into two basic camps, Functionalists and Structuralists. This division is supposed to demarcate the alternative causal presuppositions working beneath research programs. If one is functionally oriented, then organic form will be contingent upon the causal powers of the environment. If structurally oriented, one argues for nonfunctional mechanisms (e.g., internal laws of growth) to account for organic form.Traditionally, Darwin has been grouped with the functionalists because natural selection (an adaptational mechanism) plays the prominent (...) role in shaping organic form. In this paper, I sketch the dichotomy of functionalism versus structuralism and then argue that Darwin cannot be characterized adequately with this dichotomy. I argue that Darwin can incorporate both causal stories because he makes two important modifications to the traditional metaphysical presuppositions. I then offer some brief reflections on the import of Darwin's causal pluralism for the Philosophy of Science. (shrink)

Machine generated contents note: -- 1. Quantum Mechanics as a General Framework -- 2. Classical and Quantum Information and Entropy -- 3. The Brain: An Outlook -- 4. Vision -- 5. Dealing with Target's Motion and Our Own Movement -- 6. Complexity: A Necessary Condition -- 7. General Features of Life -- 8. The Organism as a Semiotic and Cybernetic System -- 9. Phylogeny -- 10. Ontogeny -- 11. Epigeny -- 12. Representational Semiotics -- 13. The Brain as an Information-Control (...) System -- 14. Decisional, Emotional and Cognitive Systems -- 15. Behavior -- 16. Learning -- 17. Memory -- 18. The Basic Symbolic Systems -- 19. What Symbols Are -- 20. Intentionality and Conceptualization -- 21. Consciousness -- 22. Development and Culture -- 23. Language -- 24. Mind and Brain (Body) -- 25. Final Philosophical Remarks. (shrink)

Possible effects of interaction (cross-talk) between signaling pathways is studied in a system of Reaction–Diffusion (RD) equations. Furthermore, the relevance of spontaneous neurite symmetry breaking and Turing instability has been examined through numerical simulations. The interaction between Retinoic Acid (RA) and Notch signaling pathways is considered as a perturbation to RD system of axon-forming potential for N2a neuroblastoma cells. The present work suggests that large increases to the level of RA–Notch interaction can possibly have substantial impacts on neurite outgrowth and (...) on the process of axon formation. This can be observed by the numerical study of the homogeneous system showing that in the absence of RA–Notch interaction the unperturbed homogeneous system may exhibit different saddle-node bifurcations that are robust under small perturbations by low levels of RA–Notch interactions, while large increases in the level of RA–Notch interaction result in a number of transitions of saddle-node bifurcations into Hopf bifurcations. It is speculated that near a Hopf bifurcation, the regulations between the positive and negative feedbacks change in such a way that spontaneous symmetry breaking takes place only when transport of activated Notch protein takes place at a faster rate. (shrink)

The meaning of the concept of fusion is discussed in relation with the works of Payer and those of Van Tieghem. It is pointed out that there is a difference, at the theoretical level, between the concept of fusion congénitale as defined by Payer and the concept of concrescence congénitale formulated by Van Tieghem. The former is inobservable by definition, while the latter deals with intercalary growth. For Van Tieghem, anatomy can prove the existence of fusion, even if we do (...) not see it during ontogenesis.We distinguish three complementary methods for explaining the unions of organs: ontogenetic, typological and phylogenetic. We have attempted, not so much to defend one or other of these methods, as to show that they often invoke very different interpretations for the same morphological phenomena. It is probable that only an analysis of the writings of 19th century botanists will clarify the concept of fusion and more generally the epistemology of plant morphology. (shrink)

This paper is about a general methodology for pattern transformation. Patterns are network representations of the relations among structures and functions within an organism. Transformation refers to any realistic or abstract transformation relevant to biology, e.g. ontogeny, evolution and phenotypic clines. The main aim of the paper is a methodology for analyzing the range of effects on a pattern due to perturbing one or more of its structures and/or functions (transformation morphology). Concepts relevant to such an analysis of pattern transformation (...) are reviewed and several new ones introduced: pattern unit; direct and indirect functional demands; compatibility and trade-off; integrating, adding and decoupling; functional effectiveness; spatial, profile and other architectonic constraints; domains of structure-function relations; goal and process adaptability; multiple pathways. The paper is written from the the perspective of architectonic morphology, viz. functional morphology focusing on the relation between anatomical coherence and the compatibility of functions. The advantages and disadvantages of inductive and deductive approaches are discussed. (shrink)

Alexander Rosenberg recently claimed (1997) that developmental biology is currently being reduced to molecular biology. cite several concrete biological examples that are intended to impugn Rosenberg's claim. I first argue that although Laubichler and Wagner's examples would refute a very strong reductionism, a more moderate reductionism would escape their attacks. Next, taking my cue from the antireductionist's perennial stress on the importance of spatial organization, I describe one form an empirical finding that refutes this moderate reductionism would take. Finally, I (...) point out an actual example, anterior-posterior axis determination in the chick, that challenges the reductionist's belief that all developmental regularities can be explained by molecular biology. In short, I argue that Rosenberg's position can be saved from Laubichler and Wagner's criticisms and putative counter-examples, but it would not survive a different kind of counter-example. (shrink)

Philosophical discussion of molecular and developmental biology began in the late 1960s with the use of genetics as a test case for models of theory reduction. With this exception, the theory of natural selection remained the main focus of philosophy of biology until the late 1970s. It was controversies in evolutionary theory over punctuated equilibrium and adaptationism that first led philosophers to examine the concept of developmental constraint. Developmental biology also gained in prominence in the 1980s as part of a (...) broader interest in the new sciences of self-organization and complexity. The current literature in the philosophy of molecular and developmental biology has grown out of these earlier discussions under the influence of twenty years of rapid and exciting growth of empirical knowledge. Philosophers have examined the concepts of genetic information and genetic program, competing definitions of the gene itself and competing accounts of the role of the gene as a developmental cause. The debate over the relationship between development and evolution has been enriched by theories and results from the new field of 'evolutionary developmental biology'. Future developments seem likely to include an exchange of ideas with the philosophy of psychology, where debates over the concept of innateness have created an interest in genetics and development. (shrink)

The aim of this paper is to evaluate the level of gender bias in Aristotle's Generation of Animals while exercising due care in the analysis of its arguments. I argue that while the GA theory is clearly sexist, the traditional interpretation fails to diagnose the problem correctly. The traditional interpretation focuses on three main sources of evidence: (1) Aristotle's claim that the female is, as it were, a "disabled" (πεπηρωμɛνo;ν) male; (2) the claim at GA IV.3, 767b6-8 that females are (...) a departure from the kind; and (3) Aristotle's supposed claim at GA IV.3, 768a21-8 that the most ideal outcome of reproduction is a male offspring that perfectly resembles its father. I argue that each of these passages has either been misunderstood or misrepresented by commentators. In none of these places is Aristotle suggesting that females are imperfect members of the species or that they result from the failure to achieve some teleological goal. I defend the view that the GA does not see reproduction as occurring for the sake of producing males; rather, what sex an embryo happens to become is determined entirely by non-teleological forces operating through material necessity. This interpretation is consistent with Aristotle's view in GA II.5 that females have the same soul as the male (741a7) as well as the argument in Metaphysics X.9 that sexual difference is not part of the species form but is an affection (παθoç) arising from the matter (1058b21-4). While the traditional interpretation has tended to exaggerate the level of sexism in Aristotle's developmental biology, the GA is by no means free of gender bias as some recent scholarship has claimed. In the final section of the paper I point to one passage where Aristotle clearly falls back on sexist assumptions in order to answer the difficult question, "Why are animals divided into sexes?". I argue that this passage in particular poses a serious challenge to anyone attempting to absolve Aristotle's developmental biology of the charge of sexism. (shrink)

This paper argues in defense of theanti-reductionist consensus in the philosophy ofbiology. More specifically, it takes issues with AlexRosenberg's recent challenge of this position. Weargue that the results of modern developmentalgenetics rather than eliminating the need forfunctional kinds in explanations of developmentactually reinforce their importance.

One foundational question in contemporarybiology is how to `rejoin evolution anddevelopment. The emerging research program(evolutionary developmental biology or`evo-devo) requires a meshing of disciplines,concepts, and explanations that have beendeveloped largely in independence over the pastcentury. In the attempt to comprehend thepresent separation between evolution anddevelopment much attention has been paid to thesplit between genetics and embryology in theearly part of the 20th century with itscodification in the exclusion of embryologyfrom the Modern Synthesis. This encourages acharacterization of evolutionary developmentalbiology as the marriage (...) of evolutionary theoryand embryology via developmental genetics. Butthere remains a largely untold story about thesignificance of morphology and comparativeanatomy (also minimized in the ModernSynthesis). Functional and evolutionarymorphology are critical for understanding thedevelopment of a concept central toevolutionary developmental biology,evolutionary innovation. Highlighting thediscipline of morphology and the concepts ofinnovation and novelty provides an alternativeway of conceptualizing the `evo and the `devoto be synthesized. (shrink)

In this article I consider the challenges for exporting causal knowledge raised by complex biological systems. In particular, James Woodward’s interventionist approach to causality identified three types of stability in causal explanation: invariance, modularity, and insensitivity. I consider an example of robust degeneracy in genetic regulatory networks and knockout experimental practice to pose methodological and conceptual questions for our understanding of causal explanation in biology. †To contact the author, please write to: Department of History and Philosophy of Science, University of (...) Pittsburgh, 1017 Cathedral of Learning, Pittsburgh, PA 15260; e‐mail: smitchel@pitt.edu. (shrink)

The idea that gravity affects dorso-ventral polarization in anouran development contrasts with the theories of self-organization through reaction-diffusion processes. As a result of a literature study we discuss the role of gravity in embryological axis formation and speculate on an influence of gravity on tissue compartmentalization. The involvement of compartmentalization in tissue homeostasis is discussed in the light of the recent progress in mammalian cell culture studies.

Lorenz proposed in his (1935) articulation of a theory of behavioral instincts that the objective of ethology is to distinguish behaviors that are “innate” from behaviors that are “learned” (or “acquired”). Lorenz’s motive was to open the investigation of certain “adaptive” behaviors to evolutionary theorizing. Accordingly, since innate behaviors are “genetic”, they are open to such investigation. By Lorenz’s light an innate/acquired or learned dichotomy rested on a familiar Darwinian distinction between genes and environments. Ever since Lorenz, ascriptions of innateness (...) have become widespread in the cognitive, behavioral, and biological sciences. The trend continues despite decades of strong arguments that show, in particular, the dichotomy that Lorenz invoked in his theory of behavioral instincts is literally false: no biological trait is the product of genes alone. Some critics suggest that the failure of Lorenz’s account shows that innateness is not well-defined in biology and the practice of ascribing innateness to various biological traits should be dropped from respectable science. Elsewhere (Ariew 1996) I argued that despite the arguments of critics, there really is a biological phenomenon underlying the concept of innateness. On my view, innateness is best understood in terms of C.H. Waddington’s concept of “canalization”, i.e. the degree to which a trait is innate is the degree to which its developmental outcome is canalized. The degree to which a developmental outcome is canalized is the degree to which the developmental process is bound to produce a particular endstate despite environmental fluctuations both in the development’s initial state and during the course of development. The canalization account differs in many ways to the traditional ways that ethologists such as Konrad Lorenz originally understood the concept of innateness. Most importantly, on the canalization account the distinction between innate and acquired is not a dichotomy, as Konrad Lorenz had it, but rather a matter of degree difference that lies along a spectrum with highly canalized development outcomes on the one end and highly environmentally sensitive development outcomes on the other end. Nevertheless, I justified the canalization account on the basis of a set of desiderata or criteria that I suggested falls-out of what seemed uncontroversial about Lorenz’s account of innateness (briefly): innateness is a property of a developing individual, innateness denotes environmental stability, and innate-ascriptions are useful in certain natural selection explanations (more below). From that same set of desiderata I argued (in my 1996) that neither the concept of heritability nor of norms of reactions—two concepts from population genetics—suffice to ground innateness. In this essay, I wish to provide further support of the canalization account in two ways. First, I wish to better motivate the desiderata by revisiting a debate between Konrad Lorenz and Daniel Lehrman over the meaning and explanatory usefulness of innate ascriptions in ethology. Second, I wish to compare my canalization account of innateness with accounts proposed by contemporary philosophers, one by Stephen Stich (1975), another by Elliott Sober (forthcoming), and a third by William Wimsatt (1986). (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)

By linking the concepts of homology and morphological organization to evolvability, this paper attempts to 1) bridge the gap between developmental and phylogenetic approaches to homology and to 2) show that developmental constraints and natural selection are compatible and in fact complementary. I conceive of a homologue as a unit of morphological evolvability, i.e., as a part of an organism that can exhibit heritable phenotypic variation independently of the organism’s other homologues. An account of homology therefore consists in explaining how (...) an organism’s developmental constitution results in different homologues/characters as units that can evolve independently of each other. The explanans of an account of homology is developmental, yet the very explanandum is an evolutionary phenomenon: evolvability in a character-by-character fashion, which manifests itself in phylogenetic patterns as recognized by phylogenetic approaches to homology. While developmental constraints and selection have often been viewed as antagonistic forces, I argue that both are complementary as they concern different parts of the evolutionary process. Developmental constraints, conceived of as the presence of the same set of homologues across phenotypic change, pertain to how heritable variation can be generated in the first place (evolvability), while natural selection operates subsequently on the produced variation. (shrink)

Ethical behavior — the conscious attempt to act in accordance with an individually-owned morality — is the product of an advanced stage of the maturing process. Three models of ethical growth derived from research in human development are applied to issues of business ethics.

Background: One of the all-time questions in evolutionary biology regards the evolution of organismal shapes, and in particular why certain forms appear repeatedly in the history of life, others only seldom and still others not at all. Recent research in this field has deployed the conceptual framework of constraints and natural selection as measured by quantitative genetic methods. -/- Scope: In this paper I argue that quantitative genetics can by necessity only provide us with useful statistical sum- maries that may (...) lead researchers to formulate testable causal hypotheses, but that any inferential attempt beyond this is unreasonable. Instead, I suggest that thinking in terms of coordinates in phenotypic spaces, and approaching the problem using a variety of empirical methods (seeking a consilience of evidence), is more likely to lead to solid inferences regarding the causal basis of the historical patterns that make up most of the data available on phenotypic evolution. (shrink)

Background: One of the all-time questions in evolutionary biology regards the evolution of organismal shapes, and in particular why certain forms appear repeatedly in the history of life, others only seldom and still others not at all. Recent research in this field has deployed the conceptual framework of constraints and natural selection as measured by quantitative genetic methods. Scope: In this paper I argue that quantitative genetics can by necessity only provide us with useful statistical sum- maries that may lead (...) researchers to formulate testable causal hypotheses, but that any inferential attempt beyond this is unreasonable. Instead, I suggest that thinking in terms of coordinates in phenotypic spaces, and approaching the problem using a variety of empirical methods (seeking a consilience of evidence), is more likely to lead to solid inferences regarding the causal basis of the historical patterns that make up most of the data available on phenotypic evolution. (shrink)

One current version of the internalism/externalism debate in evolutionary theory focuses on the relative importance of developmental constraints in evolutionary explanation. The received view of developmental constraints sees them as an internalist concept that tend to be shared across related species as opposed to selective pressures that are not. Thus, to the extent that constraints can explain anything, they can better explain similarity across species, while natural selection is better able to explain their differences. I challenge both of these aspects (...) of the received view and propose a hierarchical view of constraints. (shrink)

The highly ordered neuronal projections from the retina to the tectum mesencephali (optic tectum) in several vertebrate groups have been intensively studied. Several hypotheses so far have been proposed, suggesting mechanisms to explain the topographical and biochemical specificity of the retinotectal projections during ontogeny. In the present paper we compare the main hypotheses of retinotectal development with respect to the nature of specificity envisaged, the activity-dependence versus inheritance criterium and the strategy of argument, in casu the descriptive versus interferential type (...) of argument. Matching of the current developmental hypotheses and mechanisms for elimination or persistance of contralateral, respectively ipsilateral connections, is attempted with respect to the known neuroanatomical connectivity data in the amphibian optic tectum and with respect to the symmetry relations between ipsilateral and contralateral connection patterns described in amphibians and other vertebrate groups. Local mechanisms influencing the survival potential of synaptic contacts between retinal afferents and tectal neurons are probably essential for generating global symmetry patterns. Finally, a global topological approach is discussed with respect to its applicability in the amphibian retinotectal projection system. Basic assumptions of topological modeling appear to rely on anatomical and functional properties of the visual system like left-right symmetry, dichotomy, (absence of) convergence and segregation of fibers and neurons into columnar information units. It is concluded however, that more neuroanatomical, physiological and ultrastructural data are needed to establish a formalized operation model of the amphibian retinotectal system. (shrink)

Experiential factors such as long-term deliberate practice are powerful and necessary conditions for outstanding achievement. Nevertheless, to be able to reject the role of biology based individual differences (including genetic ones) in the manifestation of talent requires designs that expose heterogeneous samples to so-called testing-the-limits conditions, allowing asymptotic levels of performance to be analyzed comparatively. When such research has been conducted, as in the field of lifespan cognition, individual differences, including biology based ones, come to the fore and demonstrate that (...) the orchestration of excellence requires joint attention to genetic–biological and experiential factors. (shrink)

There is an argument that has recently been deployed in favor of thinking that the mind is mostly (or even exclusively) composed of cognitive modules; an argument that draws from some ideas and concepts of evolutionary and of developmental biology. In a nutshell, the argument concludes that a mind that is massively composed of cognitive mechanisms that are cognitively modular (henceforth, c-modular) is more evolvable than a mind that is not c-modular (or that is scarcely c-modular), since a cognitive mechanism (...) that is c-modular is likely to be biologically modular (henceforth, b-modular), and b-modular characters are more evolvable (e.g., Sperber 2002, Carruthers 2005). In evolutionary biology, the evolvability of a character in an organism is understood as the “organism’s capacity to facilitate the generation of non-lethal selectable phenotypic variation from random mutation” with respect to that character. Here I will argue that the notion of cognitive modularity needed to make this argument plausible will have to be understood in terms of the biological notion of variational independence; that is, it will have to be understood in such a way that a cognitive feature is c-modular only if few or no other morphological changes (cognitive and not) are significantly correlated with variations of that feature arising in members of the relevant population. I will also argue that all –except for (possibly) one—of the connotations contained in a cluster of notions of cognitive modularity widely accepted in some of the mainstream currents of thought in classical cognitive science, are simply irrelevant to the argument. In order to argue for this, I will have to examine the question as to whether there are any strong theoretical connections between (1) those connotations and (2) notions of modularity accepted in biology, specially in evolutionary and in developmental biology, that are thought to be most relevant to arguments to the effect that biological modularity enhances evolvability. (shrink)

August Weismann is famous for having argued against the inheritance of acquired characters. However, an analysis of his work indicates that Weismann always held that changes in external conditions, acting during development, were the necessary causes of variation in the hereditary material. For much of his career he held that acquired germ-plasm variation was inherited. An irony, which is in tension with much of the standard twentieth-century history of biology, thus exists – Weismann was not a Weismannian. I distinguish three (...) claims regarding the germ-plasm: (1) its continuity, (2) its morphological sequestration, and (3) its variational sequestration. With respect to changes in Weismann’s views on the cause of variation, I divide his career into four stages. For each stage I analyze his beliefs on the relative importance of changes in external conditions and sexual reproduction as causes of variation in the hereditary material. Weismann believed, and Weismannism denies, that variation, heredity, and development were deeply intertwined processes. This article is part of a larger project comparing commitments regarding variation during the latter half of the nineteenth century. (shrink)

This article began as a review of a conference, organized by Gerhard Schlosser, entitled “Modularity in Development and Evolution.” The conference was held at, and sponsored by, the Hanse Wissenschaftskolleg in Delmenhorst, Germany in May, 2000. The article subsequently metamorphosed into a literature and concept review as well as an analysis of the differences in current perspectives on modularity. Consequently, I refer to general aspects of the conference but do not review particular presentations. I divide modules into three kinds: structural, (...) developmental, and physiological. Every module fulfills none, one, or multiple functional roles. Two further orthogonal distinctions are important in this context: module-kinds versus module-variants-of-a-kind and reproducer versus nonreproducer modules. I review criteria for individuation of modules and mechanisms for the phylogenetic origin of modularity. I discuss conceptual and methodological differences between developmental and evolutionary biologists, in particular the difference between integration and competition perspectives on individualization and modular behavior. The variety in views regarding modularity presents challenges that require resolution in order to attain a comprehensive, rather than a piecemeal and fragmentary, evolutionary developmental biology. (shrink)

We show how a simple nonlinear dynamical system (the discrete quadratic iteration on the unit segment) can be the basis for modelling the embryogenesis process. Such an approach, even though being crude, can nevertheless prove to be useful when looking with the two main involved processes:i) on one hand the cell proliferation under successive divisions ii) on the other hand, the differentiation between cell lineages. We illustrate this new approach in the case of Caenorhabditis elegans by looking at the early (...) stages of embryogenesis, up to several hundreds of cells (lima bean larval stage). We show how the many results that have been obtained by several groups can be interpreted in terms of values for the parameters controlling the dynamical system. Furthermore, we can extend the model to the cases of genetic mutations. More precisely the teratogenetic and lethal effects are associated with abnormal variation of the control parameters with time. (shrink)

: This paper juxtaposes Deleuze's notion of the virtual alongside Oyama's notion of a developmental system in order to explore the promises and perils of thinking bodily identity as indeterminate at a time when new technologies render bodily ambiguity increasingly productive of both economic profit and power relations.

Freedom is first apprehended as the pursuit of an activity which implies the choice to defend a thesis among other possible ones. This translation of the problem of freedom in an articulate language presupposes a complex nervous system and sensory apparatuses which we take for granted. In this study, I try to explore the undergrounds of the problem of freedom along with the suggestion that the notion of coding could enable one to bridge nature and the mind. When organisms invent, (...) are they doing it in a spontaneous manner, inscribing in their hereditary and mnemonic instructions a stochastic contrivance of random accidents, or are they attempting to select among a limited number of schemes endowed with some optimality of functioning? If we consider them as submitted to physical forces, it is to the extent that we make them part of a strategy to extend the "laws" of a nature understood to respond passively. I suggest in this study that the epistemological understanding must regionalize itself and admit a hierarchy of dispositions in relation to the phenomenon of selection. I end by suggesting the pursuit of affirmation instead of negation, as this alone contains the requirement of integration to the knowing subject as well as the form in its act of understanding, without giving it a spontaneist position. (shrink)

Some central ideas associated with developmental systems theory (DST) are outlined for non-specialists. These ideas concern the nature of biological development, the alleged distinction between "genetic" and "environmental" traits, the relations between organism and environment, and evolutionary processes. I also discuss some criticisms of the DST approach.

Developmental systems theory (DST) is a general theoretical perspective on development, heredity and evolution. It is intended to facilitate the study of interactions between the many factors that influence development without reviving `dichotomous' debates over nature or nurture, gene or environment, biology or culture. Several recent papers have addressed the relationship between DST and the thriving new discipline of evolutionary developmental biology (EDB). The contributions to this literature by evolutionary developmental biologists contain three important misunderstandings of DST.

Will a synthesis of developmental and evolutionary biology require a focus on the role of nongenetic resources in evolution? Nongenetic variation may exist but be hidden because the phenotypes are stable (developmentally canalized) under certain background conditions. In this case, those differences may come to play important roles in evolution when background conditions change. If this is so, then a focus on the way that developmental resources are made reliable, and the ways in which reliability fails, may prove to be (...) of crucial importance to linking developmental and evolutionary biology. †To contact the author, please write to: 208 Hovland Hall, Philosophy Department, Oregon State University, Corvallis, OR 97331‐3902; e‐mail: jonathan.kaplan@oregonstate.edu. (shrink)

Recent and not so recent advances in our molecular understanding of the genome make the once prevalent view of the genome as a passive container of genetic information (i.e., genes) untenable, and emphasize the importance of the internal organization and re-organization dynamics of the genome for both development and evolution. While this conclusion is by now well accepted, the construction of a comprehensive conceptual framework for studying the genome as a dynamic system, capable of self-organization and adaptive behavior is still (...) underway. This work deals with the effect of such a conceptual shift on evolutionary thought. Specifically, I try to articulate the conceptual commitments and obligations of views that privilege explanatorily or causally the genome, its dynamics and mechanisms, over genes. I refer to this class of views as belonging to ‘the genome perspective’. (shrink)

In reworking a variety of biological concepts, Developmental Systems Theory (DST) has made frequent use of parity of reasoning. We have done this to show, for instance, that factors that have similar sorts of impact on a developing organism tend nevertheless to be invested with quite different causal importance. We have made similar arguments about evolutionary processes. Together, these analyses have allowed DST not only to cut through some age-old muddles about the nature of development, but also to effect a (...) long-delayed reintegration of development into evolutionary theory. Our penchant for causal symmetry, however (or 'causal democracy', as it has recently been termed), has sometimes been misunderstood. This paper shows that causal symmetry is neither a platitude about multiple influences nor a denial of useful distinctions, but a powerful way of exposing hidden assumptions and opening up traditional formulations to fruitful change. (shrink)

The Modern Synthesis of Darwinism and genetics regards non-genetic factors as merely constraints on the genetic variations that result in the characteristics of organisms. Even though the environment (including social interactions and culture) is as necessary as genes in terms of selection and inheritance, it does not contain the information that controls the development of the traits. S. Oyama’s account of the Parity Thesis, however, states that one cannot conceivably distinguish in a meaningful way between nature-based (i.e., gene-based) and nurture-based (...) (i.e., environment-based) characteristics in development because the information necessary for the resulting characteristics is contained at both levels. Oyama and others argue that the Parity Thesis has far-reaching implications for developmental psychology, in that both nativist and interactionist developmental accounts of motor, cognitive, affective, social, and linguistic capacities that presuppose a substantial nature/nurture dichotomy are inadequate. After considering these arguments, we conclude that either Oyama’s version of the Parity Thesis does not differ from the version advocated by liberal interactionists, or it renders precarious any analysis involving abilities present at birth (despite her claim to the contrary). More importantly, developmental psychologists need not discard the distinction between innate characteristics present at birth and those acquired by learning, even if they abandon genocentrism. Furthermore, we suggest a way nativists can disentangle the concept of maturation from a genocentric view of biological nature. More specifically, we suggest they can invoke the maturational segment of the developmental process (which involves genetic, epigenetic and environmental causes) that results in the biological “machinery” (e.g. language acquisition device) which is necessary for learning as a subsequent segment of the developmental process. (shrink)

The concept of innateness is used to make inferences between various better-understood properties, like developmental canalization, evolutionary adaptation, heritability, species-typicality, and so on (‘innateness-related properties’). This paper uses a recently-developed account of the representational content carried by inheritance systems like the genome to explain why innateness-related properties cluster together, especially in non-human organisms. Although inferences between innateness-related properties are deductively invalid, and lead to false conclusions in many actual cases, where some aspect of a phenotypic trait develops in reliance on (...) a genetic representation it will tend, better than chance, to have many of the innateness-related properties. The account also shows why inferences between innateness-related properties sometimes fail and argues that such inferences are especially misleading when applied to human psychology and behaviour because human psychological development is especially reliant on non-genetic inherited representations. (shrink)

Developmental Systems Theory (DST) emphasises the importance of non-genetic factors in development and their relevance to evolution. A common, deflationary reaction is that it has long been appreciated that non-genetic factors are causally indispensable. This paper argues that DST can be reformulated to make a more substantive claim: that the special role played by genes is also played by some (but not all) non-genetic resources. That special role is to transmit inherited representations, in the sense of Shea (2007: Biology and (...) Philosophy, 22, 313-331). Formulating DST as the claim that there are non-genetic inherited representations turns it into a striking, empirically-testable hypothesis, driving the sort of investigations that are only now beginning to appear in the scientific literature. DST’s characteristic rejection of a gene vs. environment dichotomy is preserved, but without dissolving all potentially explanatory distinctions into an interactionist causal soup, as some have alleged. (shrink)

Lorenz proposed in his (1935) articulation of a theory of behavioral instincts that the objective of ethology is to distinguish behaviors that are “innate” from behaviors that are “learned” (or “acquired”). Lorenz’s motive was to open the investigation of certain “adaptive” behaviors to evolutionary theorizing. Accordingly, since innate behaviors are “genetic”, they are open to such investigation. By Lorenz’s light an innate/acquired or learned dichotomy rested on a familiar Darwinian distinction between genes and environments. Ever since Lorenz, ascriptions of innateness (...) have become widespread in the cognitive, behavioral, and biological sciences. The trend continues despite decades of strong arguments that show, in particular, the dichotomy that Lorenz invoked in his theory of behavioral instincts is literally false: no biological trait is the product of genes alone. Some critics suggest that the failure of Lorenz’s account shows that innateness is not well-defined in biology and the practice of ascribing innateness to various biological traits should be dropped from respectable science. Elsewhere (Ariew 1996) I argued that despite the arguments of critics, there really is a biological phenomenon underlying the concept of innateness. On my view, innateness is best understood in terms of C.H. Waddington’s concept of “canalization”, i.e. the degree to which a trait is innate is the degree to which its developmental outcome is canalized. The degree to which a developmental outcome is canalized is the degree to which the developmental process is bound to produce a particular endstate despite environmental fluctuations both in the development’s initial state and during the course of development. The canalization account differs in many ways to the traditional ways that ethologists such as Konrad Lorenz originally understood the concept of innateness. Most importantly, on the canalization account the distinction between innate and acquired is not a dichotomy, as Konrad Lorenz had it, but rather a matter of degree difference that lies along a spectrum with highly canalized development outcomes on the one end and highly environmentally sensitive development outcomes on the other end. Nevertheless, I justified the canalization account on the basis of a set of desiderata or criteria that I suggested falls-out of what seemed uncontroversial about Lorenz’s account of innateness (briefly): innateness is a property of a developing individual, innateness denotes environmental stability, and innate-ascriptions are useful in certain natural selection explanations (more below). From that same set of desiderata I argued (in my 1996) that neither the concept of heritability nor of norms of reactions—two concepts from population genetics—suffice to ground innateness. In this essay, I wish to provide further support of the canalization account in two ways. First, I wish to better motivate the desiderata by revisiting a debate between Konrad Lorenz and Daniel Lehrman over the meaning and explanatory usefulness of innate ascriptions in ethology. Second, I wish to compare my canalization account of innateness with accounts proposed by contemporary philosophers, one by Stephen Stich (1975), another by Elliott Sober (forthcoming), and a third by William Wimsatt (1986). (shrink)

I argue that too much attention has been paid to the Baldwin effect. George Gaylord Simpson was probably right when he said that the effect is theoretically possible and may have actually occurred but that this has no major implications for evolutionary theory. The Baldwin effect is not even central to Baldwin’s own account of ‘social heredity’ and biology-culture co-evolution, an account that in important respects resembles the modern ideas of epigenetic inheritance and niche-construction.

Will a synthesis of developmental and evolutionary biology require a focus on the role of nongenetic resources in evolution? Nongenetic variation may exist but be hidden because the phenotypes are stable (developmentally canalized) under certain background conditions. In this case, those differences may come to play important roles in evolution when background conditions change. If this is so, then a focus on the way that developmental resources are made reliable, and the ways in which reliability fails, may prove to be (...) of crucial importance to linking developmental and evolutionary biology. †To contact the author, please write to: 208 Hovland Hall, Philosophy Department, Oregon State University, Corvallis, OR 97331‐3902; e‐mail: jonathan.kaplan@oregonstate.edu. (shrink)

Organisms inherit various kinds of developmental information and cues from their parents. The study of inheritance systems is aimed at identifying and classifying the various mechanisms and processes of heredity, the types of hereditary information that is passed on by each, the functional interaction between the different systems, and the evolutionary consequences of these properties. We present the discussion of inheritance systems in the context of several debates. First, between proponents of monism about heredity (gene-centric views), holism about heredity (Developmental (...) Systems Theory), and those stressing the role of multiple systems of inheritance. Second, between those analyzing inheritance solely in terms of replication and transmission, and views that stress the multi-generation reproduction of phenotypic traits. A third debate is concerned with different criteria that have been proposed for identifying and delimiting inheritance systems. A fourth controversy revolves around the significance of the “Lamarckian” aspects of some of the inheritance systems that have been identified, such as epigenetic inheritance and behavioral inheritance, that allow the transmission of environmentally induced characters (i.e., “soft inheritance”). (shrink)

Recent and not so recent advances in our molecular understanding of the genome make the once prevalent view of the genome as a passive container of genetic information (i.e., genes) untenable, and emphasize the importance of the internal organization and re-organization dynamics of the genome for both development and evolution. While this conclusion is by now well accepted, the construction of a comprehensive conceptual framework for studying the genome as a dynamic system, capable of self-organization and adaptive behavior is still (...) underway. This work deals with the effect of such a conceptual shift on evolutionary thought. Specifically, I try to articulate the conceptual commitments and obligations of views that privilege explanatorily or causally the genome, its dynamics and mechanisms, over genes. I refer to this class of views as belonging to ‘the genome perspective’. (shrink)

Many natural and biological phenomena can be depicted as networks. Theoretical and empirical analyses of networks have become prevalent. I discuss theoretical biases involved in the delineation of biological networks. The network perspective is shown to dissolve the distinction between regulatory architecture and regulatory state, consistent with the theoretical impossibility of distinguishing a priori between “program” and “data”. The evolutionary significance of the dynamics of trans-generational and inter-organism regulatory networks is explored and implications are presented for understanding the evolution of (...) the biological categories development-heredity; plasticity-evolvability; and epigenetic-genetic. (shrink)

The dichotomy between Nature and Nurture, which has been dismantled within the framework of development, remains embodied in the notions of plasticity and evolvability. We argue that plasticity and evolvability, like development and heredity, are neither dichotomous nor distinct: the very same mechanisms may be involved in both, and the research perspective chosen depends to a large extent on the type of problem being explored and the kinds of questions being asked. Epigenetic inheritance leads to transgenerationally extended plasticity, and developmentally-induced (...) heritable epigenetic variations provide additional foci for selection that can lead to evolutionary change. Moreover, hereditary innovations may result from developmentally induced large-scale genomic repatterning events, which are akin to Goldschmidtian “systemic mutations”. The epigenetic mechanisms involved in repatterning can be activated by both environmental and genomic stress, and lead to phylogenetic as well as ontogenetic changes. Hence, the effects and the mechanisms of plasticity directly contribute to evolvability. (shrink)

In this article I challenge the widely held assumption that human culture is inherited by means of social learning. First, I address the distinction between “social” learning and “individual” learning. I argue that most cultural ideas are not acquired by one form of learning or the other, but from a hybrid of both. Second, I discuss how individual learning can interact with niche construction. I argue that these processes collectively provide a non-social route for learned ideas to be inherited and (...) cumulatively modified. I conclude that human culture is not inherited by social learning alone; the capacities to learn from and modify our environments also play a significant role. (shrink)

August Weismann is famous for having argued against the inheritance of acquired characters. However, an analysis of his work indicates that Weismann always held that changes in external conditions, acting during development, were the necessary causes of variation in the hereditary material. For much of his career he held that acquired germ-plasm variation was inherited. An irony, which is in tension with much of the standard twentieth-century history of biology, thus exists – Weismann was not a Weismannian. I distinguish three (...) claims regarding the germ-plasm: (1) its continuity, (2) its morphological sequestration, and (3) its variational sequestration. With respect to changes in Weismann’s views on the cause of variation, I divide his career into four stages. For each stage I analyze his beliefs on the relative importance of changes in external conditions and sexual reproduction as causes of variation in the hereditary material. Weismann believed, and Weismannism denies, that variation, heredity, and development were deeply intertwined processes. This article is part of a larger project comparing commitments regarding variation during the latter half of the nineteenth century. (shrink)

Darwin’s ideas on variation, heredity, and development differ significantly from twentieth-century views. First, Darwin held that environmental changes, acting either on the reproductive organs or the body, were necessary to generate variation. Second, heredity was a developmental, not a transmissional, process; variation was a change in the developmental process of change. An analysis of Darwin’s elaboration and modification of these two positions from his early notebooks (1836–1844) to the last edition of the /Variation of Animals and Plants Under Domestication/ (1875) (...) complements previous Darwin scholarship on these issues. Included in this analysis is a description of the way Darwin employed the distinction between transmission and development, as well as the conceptual relationship he saw between heredity and variation. This paper is part of a larger project comparing commitments regarding variation during the latter half of the nineteenth century. (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)

The paper works towards an account of explanatory integration in biology, using as a case study explanations of the evolutionary origin of novelties-a problem requiring the integration of several biological fields and approaches. In contrast to the idea that fields studying lower level phenomena are always more fundamental in explanations, I argue that the particular combination of disciplines and theoretical approaches needed to address a complex biological problem and which among them is explanatorily more fundamental varies with the problem pursued. (...) Solving a complex problem need not require theoretical unification or the stable synthesis of different biological fields, as items of knowledge from traditional disciplines can be related solely for the purposes of a specific problem. Apart from the development of genuine interfield theories, successful integration can be effected by smaller epistemic units (concepts, methods, explanations) being linked. Unification or integration is not an aim in itself, but needed for the aim of solving a particular scientific problem, where the problem's nature determines the kind of intellectual integration required. (shrink)

By linking the concepts of homology and morphological organization to evolvability, this paper attempts to 1) bridge the gap between developmental and phylogenetic approaches to homology and to 2) show that developmental constraints and natural selection are compatible and in fact complementary. I conceive of a homologue as a unit of morphological evolvability, i.e., as a part of an organism that can exhibit heritable phenotypic variation independently of the organism’s other homologues. An account of homology therefore consists in explaining how (...) an organism’s developmental constitution results in different homologues/characters as units that can evolve independently of each other. The explanans of an account of homology is developmental, yet the very explanandum is an evolutionary phenomenon: evolvability in a character-by-character fashion, which manifests itself in phylogenetic patterns as recognized by phylogenetic approaches to homology. While developmental constraints and selection have often been viewed as antagonistic forces, I argue that both are complementary as they concern different parts of the evolutionary process. Developmental constraints, conceived of as the presence of the same set of homologues across phenotypic change, pertain to how heritable variation can be generated in the first place (evolvability), while natural selection operates subsequently on the produced variation. (shrink)

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

The evolutionary embryologist Gavin Rylands de Beer can be viewed as one of the forerunners of modern evolutionary developmental biology in that he posed crucial questions and proposed relevant answers about the causal relationship between ontogeny and phylogeny. In his developmental approach to the phylogenetic phenomenon of homology, he emphasized that homology of morphological structures is to be identified neither with the sameness of the underlying developmental processes nor with the homology of the genes that are in involved in the (...) development of the structures. De Beer’s work on developmental evolution focused on the notion of heterochrony, arguing that paedomorphosis increases morphological evolvability and is thereby an important mode of evolution that accounts for the origin of many taxa, including higher taxa. (shrink)

The present paper analyzes the use and understanding of the homology concept across different biological disciplines. It is argued that in its history, the homology concept underwent a sort of adaptive radiation. Once it migrated from comparative anatomy into new biological fields, the homology concept changed in accordance with the theoretical aims and interests of these disciplines. The paper gives a case study of the theoretical role that homology plays in comparative and evolutionary biology, in molecular biology, and in evolutionary (...) developmental biology. It is shown that the concept or variant of homology preferred by a particular biological field is used to bring about items of biological knowledge that are characteristic for this field. A particular branch of biology uses its homology concept to pursue its specific theoretical goals. (shrink)

Exploring whether clades can reproduce leads to new perspectives on general accounts of biological development and individuation. Here we apply James Griesemer's general account of reproduction to clades. Griesemer's account of reproduction includes a requirement for development, raising the question of whether clades may bemeaningfully said to develop. We offer two illustrative examples of what clade development might look like, though evaluating these examples proves difficult due to the paucity of general accounts of development. This difficulty, however, is instructive about (...) what a general account of development should look like and how it may usefully be applied to research problems (further suggesting a means for evaluating general accounts of development). Reproduction also requires individuation of parent and offspring. We argue that there is no special problem of individuating older and younger clades. The vagaries involved with determining when clades begin, mature, and end are precisely the same as those that arise when the same questions are asked of cells, organisms, or species. Though the question of clade reproduction and selection may still be open, the process of discovery presents new insights into old problems. (shrink)

Evolutionary developmental biology (Evo-Devo) is a new and rapidly developing field of biology which focuses on questions in the intersection of evolution and development and has been seen by many as a potential synthesis of these two fields. This synthesis is the topic of the books reviewed here. Integrating Evolution and Development (edited by Roger Sansom and Robert Brandon), is a collection of papers on conceptual issues in Evo-Devo, while From Embryology to Evo-Devo (edited by Manfred Laubichler and Jane Maienschein) (...) is a history of the problem of the relations between ontogeny and phylogeny. (shrink)

This volume joins a growing list of books, monographs, and proceedings from scientific meetings that attempt to consolidate the wide spectrum of approaches emphasizing the role of development in evolution into a coherent and productive synthesis, often called evo-devo. Evo-devo is seen as a replacement or amendment of the modern synthesis that has dominated the field of evolution since the 1940s and which, as even its architects confessed, was fundamentally incomplete because development remained outside its theoretical framework (Mayr and Provine (...) 1980).As the volume attests, there is now a strong feeling that the time is ripe for the onsolidation of evo-devo, and that the field is mature enough so that mapping the theoretical terrain and experimental approaches is both feasible and scientifically productive. Now is an appropriate time to try to weave the strands of reasoning leading to the developmental perspective and offer a synthesis. (shrink)

Will a synthesis of developmental and evolutionary biology require a focus on the role of nongenetic resources in evolution? Nongenetic variation may exist but be hidden because the phenotypes are stable (developmentally canalized) under certain background conditions. In this case, those differences may come to play important roles in evolution when background conditions change. If this is so, then a focus on the way that developmental resources are made reliable, and the ways in which reliability fails, may prove to be (...) of crucial importance to linking developmental and evolutionary biology. †To contact the author, please write to: 208 Hovland Hall, Philosophy Department, Oregon State University, Corvallis, OR 97331‐3902; e‐mail: jonathan.kaplan@oregonstate.edu. (shrink)

Organisms inherit various kinds of developmental information and cues from their parents. The study of inheritance systems is aimed at identifying and classifying the various mechanisms and processes of heredity, the types of hereditary information that is passed on by each, the functional interaction between the different systems, and the evolutionary consequences of these properties. We present the discussion of inheritance systems in the context of several debates. First, between proponents of monism about heredity (gene-centric views), holism about heredity (Developmental (...) Systems Theory), and those stressing the role of multiple systems of inheritance. Second, between those analyzing inheritance solely in terms of replication and transmission, and views that stress the multi-generation reproduction of phenotypic traits. A third debate is concerned with different criteria that have been proposed for identifying and delimiting inheritance systems. A fourth controversy revolves around the significance of the “Lamarckian” aspects of some of the inheritance systems that have been identified, such as epigenetic inheritance and behavioral inheritance, that allow the transmission of environmentally induced characters (i.e., “soft inheritance”). (shrink)

Many natural and biological phenomena can be depicted as networks. Theoretical and empirical analyses of networks have become prevalent. I discuss theoretical biases involved in the delineation of biological networks. The network perspective is shown to dissolve the distinction between regulatory architecture and regulatory state, consistent with the theoretical impossibility of distinguishing a priori between “program” and “data”. The evolutionary significance of the dynamics of trans-generational and inter-organism regulatory networks is explored and implications are presented for understanding the evolution of (...) the biological categories development-heredity; plasticity-evolvability; and epigenetic-genetic. (shrink)

This paper is about mechanisms and models, and how they interact. In part, it is a response to recent discussion in philosophy of biology regarding whether natural selection is a mechanism. We suggest that this debate is indicative of a more general problem that occurs when scientists produce mechanistic models of populations and their behaviour. We can make sense of claims that there are mechanisms that drive population-level phenomena such as macroeconomics, natural selection, ecology, and epidemiology. But talk of mechanisms (...) and mechanistic explanation evokes objects with well-defined and localisable parts which interact in discrete ways, while models of populations include parts and interactions that are neither local nor discrete in any actual populations. This apparent tension can be resolved by carefully distinguishing between the properties of a model and those of the system it represents. To this end, we provide an analysis that recognises the flexible relationship between a mechanistic model and its target system. In turn, this reveals a surprising feature of mechanistic representation and explanation: it can occur even when there is a mismatch between the mechanism of the model and that of its target. Our analysis reframes the debate, providing an alternative way to interpret scientists’ mechanism-talk , which initially motivated the issue. We suggest that the relevant question is not whether any population-level phenomenon such as natural selection is a mechanism, but whether it can be usefully modelled as though it were a particular type of mechanism. (shrink)

One important aspect of biological explanation is detailed causal modeling of particular phenomena in limited experimental background conditions. Recognising this allows a new avenue for intertheoretic reduction to be seen. Reductions in biology are possible, when one fully recognises that a sufficient condition for a reduction in biology is a molecular model of 1) only the demonstrated causal parameters of a biological model and 2) only within a replicable experimental background. These intertheoretic identifications –which are ubiquitous in biology and form (...) the basis of ruthless reductions (Bickle 2003)- are criticised as merely “local” (Sullivan 2009) or “fragmentary” (Schaffner 2006). However, in an instructive case, a biological model is preserved in molecular terms, and a complex biological phenomenon has been successfully reduced. In doing this the molecular model remains valid in a broader range of background conditions and meaningfully unites disparate biological phenomena. (shrink)

Recent work on the heat-shock protein Hsp90 by Rutherford and Lindquist (1998) has been included among the pieces of evidence taken to show the essential role of developmental processes in evolution; Hsp90 acts as a buffer against phenotypic variation, allowing genotypic variation to build. When the buffering capacity of Hsp90 is altered (e.g., in nature, by mutation or environmental stress), the genetic variation is "revealed," manifesting itself as phenotypic variation. This phenomenon raises questions about the genetic variation before and after (...) what I will call a "revelation event": Is it neutral, nearly neutral, or non-neutral (i.e., strongly deleterious or strongly advantageous)? Moreover, what kinds of evolutionary processes do we take to be at work? Rutherford and Lindquist (1998) focus on the implications of non-neutral variation and selection. Later work by Queitsch, Sangster, and Lindquist (2002) and Sangster, Lindquist, and Queitsch (2004) raises the possibility that Hsp90 buffering may play the role that was played by drift in Sewall Wright's shifting balance model, permitting transition from one adaptive peak to another. However, Ohta (2002) suggests that much of this variation may be nearly neutral, which in turn, would imply a strong role for drift as well as selection. The primary goal of this paper is to illuminate the alternative scenarios and the processes operating in each. At the end, I raise the possibility of a synthesis between evo-devo and nearly neutral evolution. (shrink)

Evolutionary developmental biology (evo-devo) offers both an account of developmental processes and also new integrative frameworks for analyzing interactions between development and evolution. Biologists and philosophers are keen on evo-devo in part because it appears to offer a comfort zone between, on the one hand, what some take to be the relative inability of mainstream evolutionary biology to integrate a developmental perspective; and, on the other hand, what some take to be more intractable syntheses of development and evolution. In this (...) article, I outline core concerns of evo-devo, distinguish theoretical and practical variants, and counter Sterelny's recent argument that evo-devo's attention to development, while important, offers no significant challenge to evolutionary theory as we know it. (shrink)

Mesoudi et al.'s new synthesis for cultural evolution closely parallels the evolutionary synthesis of Neo-Darwinism. It too draws inspiration from population genetics, recruits other fields, and, unfortunately, also ignores development. Enculturation involves many serially acquired skills and dependencies that allow us to build a rich cumulative culture. The newer synthesis, evolutionary developmental biology, provides a key tool, generative entrenchment, to analyze them. (Published Online November 9 2006).

Evolutionary Developmental Biology (Evo-Devo) is philosophically fascinating because of its plurality of scientific “cultures” of practice and theory that continue making progress towards a better understanding of complex biological reality. In this chapter, through an examination of a variety of the scientific cultures pertinent to Evo-Devo, I show that Evo-Devo can be usefully understood as a /trading zone/ (Galison 1997). That is, a variety of disciplines, styles, and paradigms negotiate heavily with each other in the domain of Evo-Devo. I am (...) concerned with the differences, the interactions, and the relative openness and flexibility of these cultures. When are the cultures acting—individually or collectively—in ways that further research, empirically, theoretically, and ethically? When do they become imperialistic, in the sense of excluding and subordinating other cultures? I wish to develop a critical /assumption archeology/ (my term, following Michel Foucault, Ian Hacking, and Michael Friedman), which explores some of the key presuppositions standing behind or under or within each of these cultures. These assumptions ground the concepts, methods, and models of each culture. The goal of this chapter is to identify six cultures of Evo-Devo (three styles and three paradigms), and provide an initial archeology of their internal structure, and mutual relations, through the concept of trading zone. My main excavation site is Bonner (1982), founding text of Evo-Devo and product of the 1981 Dahlem “Evolution and Development” workshop, on which this 2011 anthology (and workshop) was also based. (shrink)

This article began as a review of a conference, organized by Gerhard Schlosser, entitled “Modularity in Development and Evolution.” The conference was held at, and sponsored by, the Hanse Wissenschaftskolleg in Delmenhorst, Germany in May, 2000. The article subsequently metamorphosed into a literature and concept review as well as an analysis of the differences in current perspectives on modularity. Consequently, I refer to general aspects of the conference but do not review particular presentations. I divide modules into three kinds: structural, (...) developmental, and physiological. Every module fulfills none, one, or multiple functional roles. Two further orthogonal distinctions are important in this context: module-kinds versus module-variants-of-a-kind and reproducer versus nonreproducer modules. I review criteria for individuation of modules and mechanisms for the phylogenetic origin of modularity. I discuss conceptual and methodological differences between developmental and evolutionary biologists, in particular the difference between integration and competition perspectives on individualization and modular behavior. The variety in views regarding modularity presents challenges that require resolution in order to attain a comprehensive, rather than a piecemeal and fragmentary, evolutionary developmental biology. (shrink)

As Paul Griffiths [2002] puts it, “innateness” is associated with different clusters of related ideas where each cluster depends on different historical, cultural and intellectual contexts. In psychology innateness is typically opposed to learning while the biological opposite of innate is ‘acquired’. ‘Acquired’ and ‘learned’ have different extensions. Learning is one way to acquire a character but there are others. Cuts and scratches are unlearned yet acquired; if we could acquire languages by popping a pill, then languages would be unlearned (...) yet acquired according to the wide biological application of the term [Sober, 1998]. Further, in psychology and philosophy innateness is often associated with both “universality” (or species-specificity), and, relatedely, innate traits are often thought to be “fixed” or “unmodifiable”. But, biologists recognize a range of developmental patterns that a specific trait may take. Some are universal, but others are not, as in the case of innate diseases. Some are “fixed” in the sense that once we develop them we have them for the rest of our lives; some innate diseases are like this, but others, are modifiable. Sober [1998] cites a case of an Egyptian vulture that when first confronted with an ostrich egg and a stone, will break the egg with the stone, but if the vulture repeatedly comes to find broken eggs to be empty, it will eventually stop breaking eggs. These examples lend support to Griffiths’s thesis, since the concept of innateness in psychology appears to be in several ways distinct from the concept of innateness in biology. (shrink)

Lorenz proposed in his (1935) articulation of a theory of behavioral instincts that the objective of ethology is to distinguish behaviors that are “innate” from behaviors that are “learned” (or “acquired”). Lorenz’s motive was to open the investigation of certain “adaptive” behaviors to evolutionary theorizing. Accordingly, since innate behaviors are “genetic”, they are open to such investigation. By Lorenz’s light an innate/acquired or learned dichotomy rested on a familiar Darwinian distinction between genes and environments. Ever since Lorenz, ascriptions of innateness (...) have become widespread in the cognitive, behavioral, and biological sciences. The trend continues despite decades of strong arguments that show, in particular, the dichotomy that Lorenz invoked in his theory of behavioral instincts is literally false: no biological trait is the product of genes alone. Some critics suggest that the failure of Lorenz’s account shows that innateness is not well-defined in biology and the practice of ascribing innateness to various biological traits should be dropped from respectable science. Elsewhere (Ariew 1996) I argued that despite the arguments of critics, there really is a biological phenomenon underlying the concept of innateness. On my view, innateness is best understood in terms of C.H. Waddington’s concept of “canalization”, i.e. the degree to which a trait is innate is the degree to which its developmental outcome is canalized. The degree to which a developmental outcome is canalized is the degree to which the developmental process is bound to produce a particular endstate despite environmental fluctuations both in the development’s initial state and during the course of development. The canalization account differs in many ways to the traditional ways that ethologists such as Konrad Lorenz originally understood the concept of innateness. Most importantly, on the canalization account the distinction between innate and acquired is not a dichotomy, as Konrad Lorenz had it, but rather a matter of degree difference that lies along a spectrum with highly canalized development outcomes on the one end and highly environmentally sensitive development outcomes on the other end. Nevertheless, I justified the canalization account on the basis of a set of desiderata or criteria that I suggested falls-out of what seemed uncontroversial about Lorenz’s account of innateness (briefly): innateness is a property of a developing individual, innateness denotes environmental stability, and innate-ascriptions are useful in certain natural selection explanations (more below). From that same set of desiderata I argued (in my 1996) that neither the concept of heritability nor of norms of reactions—two concepts from population genetics—suffice to ground innateness. In this essay, I wish to provide further support of the canalization account in two ways. First, I wish to better motivate the desiderata by revisiting a debate between Konrad Lorenz and Daniel Lehrman over the meaning and explanatory usefulness of innate ascriptions in ethology. Second, I wish to compare my canalization account of innateness with accounts proposed by contemporary philosophers, one by Stephen Stich (1975), another by Elliott Sober (forthcoming), and a third by William Wimsatt (1986). (shrink)

Cognitive scientists often employ the notion of innateness without defining it. The issue is, how is innateness defined in biology? Some critics contend that innateness is not a legitimate concept in biology. In this paper I will argue that it is. However, neither the concept of high heritability nor the concept of flat norm of reaction (two popular accounts in the biology literature) define innateness. An adequate account is found in developmental biology. I propose that innateness is best defined in (...) terms of C. H. Waddington's concept of canalization. (shrink)

The question whether ethical behavior is biologically determined may refer either to the capacity for ethics (i.e., the proclivity to judge human actions as either right or wrong), or to the moral norms accepted by human beings for guiding their actions. I herein propose: (1) that the capacity for ethics is a necessary attribute of human nature; and (2) that moral norms are products of cultural evolution, not of biological evolution. Humans exhibit ethical behavior by nature because their biological makeup (...) determines the presence of three necessary conditions for ethical behavior: (i) the ability to anticipate the consequences of one’s own actions; (ii) the ability to make value judgments; and (iii) the ability to choose between alternative courses of action. Ethical behavior came about in evolution not because it is adaptive in itself, but as a necessary consequence of man’s eminent intellectual abilities, which are an attribute directly promoted by natural selection. That is, morally evolved as an exaptation, not as an adaptation. Since Darwin’s time there have been evolutionists proposing that the norms of morality are derived from biological evolution. Sociobiologists represent the most recent and most subtle version of that proposal. The sociobiologists' argument is that human ethical norms are sociocultural correlates of behaviors fostered by biological evolution. I argue that such proposals are misguided and do not escape the naturalistic fallacy. The isomorphism between the behaviors promoted by natural selection and those sanctioned by moral norms exist only with respect to the consequences of the behaviors; the underlying causations are completely disparate. (shrink)

The hunt for a biologically respectable definition for the folk concept of innateness is still on. I defend Ariew’s Canalization account of innateness against the criticisms of Griffiths and Machery, but highlight the remaining flaws in this proposal. I develop a new analysis based on the notion of environmental induction. A trait is innate, I argue, iff it is not environmentally induced. I augment this definition with a novel analysis of environmental induction that draws on the contrastive nature of causal (...) explanation. Whether a trait is environmentally induced, I argue, depends on a context sensitive contrast class. I argue that a “Noninduction” analysis of innateness allows the concept an explanatory role in biology. I show how my proposal co-opts the successes of the Canalization account whilst avoiding its pitfalls, and I account for why biologists associate a range of disparate properties with innateness. (shrink)

This is the first volume of a projected three-volume set on the subject of innateness. The extent to which the mind is innate is one of the central questions in the human sciences, with important implications for many surrounding debates. By bringing together the top nativist scholars in philosophy, psychology, and allied disciplines these volumes provide a comprehensive assessment of nativist thought and a definitive reference point for future nativist inquiry. The Innate Mind: Structure and Content, concerns the fundamental architecture (...) of the mind, addressing such question as: What capacities, processes, representations, biases, and connections are innate? How do these innate elements feed into a story about the development of our mature cognitive capacities, and which of them are shared with other members of the animal kingdom? The editors have provided an introduction giving some of the background to debates about innateness and introducing each of the subsequent essays, as well as a consolidated bibliography that will be a valuable reference resource for all those interested in this area. The volume will be of great importance to all researchers and students interested in the fundamental nature and powers of the human mind. Together, the three volumes in the series will provide the most intensive and richly cross-disciplinary investigation of nativism ever undertaken. They point the way toward a synthesis of nativist work that promises to provide a new understanding of our minds and their place in the natural order. (shrink)

The notion of innateness is widely used, particularly in philosophy of mind, cognitive science and linguistics. Despite this popularity, it remains a controversial idea. This is partly because of the variety of ways in which it can be explicated and partly because it appears to embody the suggestion that we can determine the relative causal contributions of genes and environment in the development of biological individuals. As these causes are not independent, the claim is metaphysically suspect. This paper argues that (...) there is a plausible reconstruction of the notion of innateness. This involves defining it sufficiently broadly to cover most of the current usages as well as making it an informational rather than a causal property. This has two consequences. Firstly, innateness becomes a matter of degree. Secondly, we have to abandon the idea, originally proposed by ethologists, that innate traits are necessarily the products of genetic information. (shrink)

The capacity to engage with art is a human universal present in all cultures and just about every individual human. This indicates that this capacity is evolved. In this Critical Notice of Denis Dutton's The Art Instinct, I discuss various evolutionary scenarios and their consequences. Dutton and I both reject the "spandrel" approach that originates from the work of Gould and Lewontin. Dutton proposes, following work of Geoffrey Miller, that art is sexually selected--that art-production is a sign of a fit (...) genome in males. I argue that while assortative mating may well have had a role in the evolution of "the art instinct", group selection is a better explanation. I also take issue with Dutton's "cluster concept" approach to defining art, and argue that it is a universal and essential characteristic of art that it is appreciated both for what it expresses and for the way that it expresses. It thus requires a reflexive capacity that is not operative in the appreciation of sport spectacles and pornography. (shrink)

This article is inspired by recent psychological studies confirming that a child is not born a blank slate but has important innate capabilities. An important part of the ``learning'' required to deal with the three dimensional world of objects, processes, and other beings was done by evolution. Each child need not do this learning itself.

S. Oyama’s prominent account of the Parity Thesis states that one cannot distinguish in a meaningful way between nature-based (i.e. gene-based) and nurture-based (i.e. environment-based) characteristics in development because the information necessary for the resulting characteristics is contained at both levels. Oyama as well as P. E. Griffiths and K. Stotz argue that the Parity Thesis has far-reaching implications for developmental psychology in that both nativist and interactionist developmental accounts of psychological capacities that presuppose a substantial nature/nurture dichotomy are inadequate. (...) We argue that well-motivated abandoning of the nature/nurture dichotomy, as advocated in converging versions of the Parity Thesis in biology, does not necessarily entail abandoning the distinction between biologically given abilities necessary for the development of higher psychological capacities and the learning process they enable. Thus, contrary to the claims of the aforementioned authors, developmental psychologists need not discard a substantial distinction between innate (biologically given) characteristics and those acquired by learning, even if they accept the Parity Thesis. We suggest a two-stage account of development: the first stage is maturational and involves interaction of genetic, epigenetic and environmental causes, resulting in the endogenous biological ‘machinery’ (e.g. language acquisition device), responsible for learning in the subsequent stage of the developmental process by determining the organism’s responses to the environment. This account retains the crux of nativism (the endogenous biological structure determines the way the organism learns/responds to an environment) whilst adopting the developmentalist view of biology by characterizing environments as distinctly different in terms of structure and function in two developmental stages. (shrink)

The discovery that genes have something to do with behavior came as a shock in an era in which people thought that the mind of a newborn was a blank slate and that anyone could do anything if only they strove hard enough. And it continues to set off alarm bells. Many people worry about a Brave New World in which parents or governments will try to re-engineer human nature. Others see genes as a threat to free will and personal (...) responsibility, citing headlines such as “Man’s genes made him kill, his lawyers claim.” Behavioral geneticists are sometimes picketed, censored, or compared to Nazis. With increasing knowledge of how the genome works, many beliefs about ourselves will indeed have to be rethought. But the worst fears of the genophobes are misplaced. It is easy to exaggerate the significance of behavioral genetics for our lives. (shrink)

First I should clarify my thesis. When I say the mind starts off as a blank slate, I’m saying that it’s devoid of substantive concepts or ideas, that is non-logical concepts or ideas. Some examples of substantive concepts are: the concept of a cat, the concept of a quark, the concept of being square, and the concept of heaviness.

The concept of innateness is used to make inferences between various better-understood properties, like developmental canalization, evolutionary adaptation, heritability, species-typicality, and so on (‘innateness-related properties’). This paper uses a recently-developed account of the representational content carried by inheritance systems like the genome to explain why innateness-related properties cluster together, especially in non-human organisms. Although inferences between innateness-related properties are deductively invalid, and lead to false conclusions in many actual cases, where some aspect of a phenotypic trait develops in reliance on (...) a genetic representation it will tend, better than chance, to have many of the innateness-related properties. The account also shows why inferences between innateness-related properties sometimes fail and argues that such inferences are especially misleading when applied to human psychology and behaviour because human psychological development is especially reliant on non-genetic inherited representations. (shrink)

Recent theoretical work has identified a tightly-constrained sense in which genes carry representational content. Representational properties of the genome are founded in the transmission of DNA over phylogenetic time and its role in natural selection. However, genetic representation is not just relevant to questions of selection and evolution. This paper goes beyond existing treatments and argues for the heterodox view that information generated by a process of selection over phylogenetic time can be read in ontogenetic time, in the course of (...) individual development. Recent results in evolutionary biology, drawn both from modelling work, and from experimental and observational data, support a role for genetic representation in explaining individual ontogeny: both genetic representations and environmental information are read by the mechanisms of development, in an individual, so as to lead to adaptive phenotypes. Furthermore, in some cases there appears to have been selection between individuals that rely to different degrees on the two sources of information. Thus, the theory of representation in inheritance systems like the genome is much more than just a coherent reconstruction of information talk in biology. Genetic representation is a property with considerable explanatory utility. (shrink)

What Science Offers the Humanities examines some of the deep problems facing current approaches to the study of culture. It focuses especially on the excesses of postmodernism, but also acknowledges serious problems with postmodernism's harshest critics. In short, Edward Slingerland argues that in order for the humanities to progress, its scholars need to take seriously contributions from the natural sciences—and particular research on human cognition—which demonstrate that any separation of the mind and the body is entirely untenable. The author provides (...) suggestions for how humanists might begin to utilize these scientific discoveries without conceding that science has the last word on morality, religion, art, and literature. Calling into question such deeply entrenched dogmas as the "blank slate" theory of nature, strong social constructivism, and the ideal of disembodied reason, What Science Offers the Humanities replaces the human-sciences divide with a more integrated approach to the study of culture. (shrink)

Human, cognition, interaction, and culture are thoroughly intertwined. Without cognition and interaction, there would be no culture. Without culture, cognition and interaction would be very different affairs, as they are among other social species. The effect of culture on mental life has always been a main concern of the social sciences and, after a long period of almost total neglect, it is more and more taken into consideration in cognitive psychology. The effect of cognition, and in particular of the ability (...) to attribute mental states to others, on interaction has become a important topic of investigation in developmental and social psychology. Little attention, on the other hand, is paid to the effect of cognition on culture. All would agree, of course that the human mind is what makes human culture possible. Quite generally, however, the mind is seen as a mere enabler of culture, a pure opportunity with no constraints attached, nothing that might contribute to shaping, or at least to biasing cultural contents. Against this neglect of cognition, I will argue that understanding the mind is doubly important to the study of culture. Psychological considerations are crucial both to a proper characterization of what is cultural and to a proper explanation of cultural phenomena. Most anthropologists may be too savvy to still talk of the mind as a “blank slate”, but they too often meet any discussion of psychological factors in culture with a blank stare. Most evolutionary theorists who see cultural evolution as an analogue of biological evolution by natural selection postulate rather than investigate the few particulars of the human mind.. (shrink)