We critically examine a number of aspects of Grafen’s ‘formal Darwinism’ project. We argue that Grafen’s ‘selection-optimality’ links do not quite succeed in vindicating the working assumption made by behavioural ecologists and others—that selection will lead organisms to exhibit adaptive behaviour—since these links hold true even in the presence of strong genetic and developmental constraints. However we suggest that the selection-optimality links can profitably be viewed as constituting an axiomatic theory of fitness. Finally, we (...) compare Grafen’s project with Fisher’s ‘fundamental theorem of natural selection’, and we speculate about whether Grafen’s results can be extended to a game-theoretic setting. (shrink)

Developmental systems theory (DST) is a wholeheartedly epigenetic approach to development, inheritance and evolution. The developmental system of an organism is the entire matrix of resources that are needed to reproduce the life cycle. The range of developmental resources that are properly described as being inherited, and which are subject to natural selection, is far wider than has traditionally been allowed. Evolution acts on this extended set of developmental resources. From a (...) class='Hi'>developmental systems perspective, development does not proceed according to a preformed plan; what is inherited is much more than DNA; and evolution is change not only in gene frequencies, but in entire developmental systems. (shrink)

The presence of various mechanisms of non-genetic inheritance is one of the main problems for current evolutionary theory according to several critics. Sufficient empirical and conceptual reasons exist to take this claim seriously, but there is little consensus on the implications of multiple inheritance systems for evolutionary processes. Here we use the Price Equation as a starting point for a discussion of the differences between four recently proposed categories of inheritance systems; genetic, epigenetic, behavioral and (...) symbolic. Specifically, we address how the components of the Price Equation encompass different non-genetic systems of inheritance in an attempt to clarify how the different systems are conceptually related. We conclude that the four classes of inheritance systems do not form distinct clusters with respect to their effect on the rate and direction of phenotypic change from one generation to the next in the absence or presence of selection. Instead, our analyses suggest that different inheritance systems can share features that are conceptually very similar, but that their implications for adaptive evolution nevertheless differ substantially as a result of differences in their ability to couple selection and inheritance. (shrink)

FOREWORD. Gilbert Gottlieb and the Developmental Point of View. I. INTRODUCTION. 1. Developmental Systems, Nature-Nurture, and the Role of Genes in Behavior and Development: On the Legacy of Gilbert Gottlieb. 2. Normally Occurring Environmental and Behavioral Influences on Gene Activity: From Central Dogma to Probabilistic Epigenesis. II. THEORETICAL FOUNDATIONS FOR THE DEVELOPMENTAL STUDY OF BEHAVIOR AND GENETICS. 3. Historical and Philosophical Perspectives on Behavioral Genetics and Developmental Science. 4. Development and Evolution Revisited. 5. Probabilistic Epigenesis (...) and Modern Behavioral and Neural Genetics. 6. The Roles of Environment, Experience, and Learning in Behavioral Development. 7. Contemporary Ideas in Physics and Biology in Gottlieb’s Psychology. III. EMPIRICAL STUDIES OF BEHAVIORAL DEVELOPMENT AND GENETICS. 8. Behavioral Development during the Mother-Young Interaction in Placental Mammals: The Development of Behavior in the Relationship with the Mother. 9. Amniotic Fluid as an Extended Milieu Interieur. 10. Developmental Effects of Selective Breeding for an Infant Trait. 11. Emergence and Constraint in Novel Behavioral Adaptations. 12. Nonhuman Primate Research Contributions to Understanding Genetic and Environmental Influences on Phenotypic Outcomes across Development. 13. Interactive Contributions of Genes and Early Experience to Behavioural Development: Sensitive Periods and Lateralized Brain and Behaviour. 14. Trans-Generational Epigenetic Inheritance. 15. The Significance of Non-Replication of Gene-Phenotype Associations. 16. Canalization and Malleability Reconsidered: The Developmental Basis of Phenotypic Stability and Variability. IV. APPLICATIONS TO DEVELOPMENT. 17. Gene-Parenting Interplay in the Development of Infant Emotionality. 18. Genetic Research in Psychiatry and Psychology: A Critical Overview. 19. On the Limits of Standard Quantitative Genetic Modeling of Inter-Individual Variation: Extensions, Ergodic Conditions and a New Genetic Factor Model of Intra-Individual Variation. 20. Songs My Mother Taught Me: Gene-Environment Interactions, Brain Development and the Auditory System: Thoughts on Non-Kin Rejection, Part II. 21. Applications of Developmental Systems Theory to Benefit Human Development: On the Contributions of Gilbert Gottlieb to Individuals, Families, and Communities. (shrink)

The themes, problems and challenges of developmental systems theory as described in Cycles of Contingency are discussed. We argue in favor of a robust approach to philosophical and scientific problems of extended heredity and the integration of behavior, development, inheritance, and evolution. Problems with Sterelny's proposal to evaluate inheritance systems using his `Hoyle criteria' are discussed and critically evaluated. Additional support for a developmental systems perspective is sought in evolutionary studies of performance (...) and behavior modulation of fitness. (shrink)

What kind mechanisms one deems central for the evolutionary process deeply influences one's understanding of the nature of organisms, including cognition. Reversely, adopting a certain approach to the nature of life and cognition and the relationship between them or between the organism and its environment should affect one's view of evolutionary theory. This paper explores this reciprocal relationship in more detail. In particular it argues that the view of living and cognitive systems, especially humans, as deeply integrated beings embedded (...) in and transformed by their genetic, epigenetic, behavioral, ecological, socio-cultural and cognitive-symbolic legacies calls for an extended evolutionary synthesis that goes beyond either a theory of genes juxtaposed against a theory of cultural evolution and or even more sophisticated theories of gene-culture coevolution and niche construction. Environments, particularly in the form of developmental environments, do not just select for variation, they also create new variation by influencing development through the reliable transmission of non-genetic but heritable information. This paper stresses particularly views of embodied, embedded, enacted and extended cognition, and their relationship to those aspects of extended inheritance that lie between genetic and cultural inheritance, the still gray area of epigenetic and behavioral inheritance systems that play a role in parental effect. These are the processes that can be regarded as transgenerational developmental plasticity and that I think can most fruitfully contribute to, and be investigated by, developmental psychology. (shrink)

In the past 150 years there have been many attempts to draw parallels between cultural and biological evolution. Most of these attempts were flawed due to lack of knowledge and false ideas about evolution. In recent decades these shortcomings have been cleared away, thus triggering a renewed interest in the subject. This paper offers a critical survey of the main issues and arguments in that discussion. The paper starts with an explication of the Darwinian algorithm of evolution. It is argued (...) that this ‘formula’ is substrate-neutral, which means that biological evolution might not be the only Darwinian process. Other dynamic systems could evolve as well provided that certain conditions are met. In the case of human culture this seems to be the case. The paper then focuses on the notion of niche construction. It is argued that niche construction plays a crucial role in human evolution because it has altered the sources of natural selection and thus the path of evolution. Next two approaches to cultural evolution are discussed: sociobiology and memetics. I will argue that both approaches have flaws because they either underestimate the influence of culture or they stretch analogies too far. Finally two common objections against the idea of cultural evolution are addressed: Lamarckian inheritance and the issue of guided variation. I will argue that although cultural evolution differs from biological evolution in several respects, these discrepancies do not jeopardize the claim that cultural evolution is essentially Darwinian. (shrink)

The neo-Darwinian paradigm, focusing on natural selection of genes responsible for differential adaption, provides the foundation for explaining evolutionary processes. The modern synthesis is broader, however, focusing on organisms rather than on gene transmissions per se. Yet, strands of current biology argue for further supplementation of Darwinian theory, pointing to nonbiotic drivers of evolutionary development, for example, self-organization of physical structures, and the interaction between individual organisms, groups of organisms, and their nonbiotic environments. According to niche construction theory, when (...) organisms and groups develop, they not only adapt to their environments but modify their environments, creating new habitats for later generations. Insofar as ecological niches persist beyond the lifecycle of individual organisms, an ecological inheritance system exists alongside genetic inheritance. Such ecological structures may even facilitate the development of a cultural inheritance system, as we see in humans. The article discusses theological perspectives of such new developments within holistic biology. (shrink)

Darwinism is defined here as an evolving research tradition based upon the concepts of natural selection acting upon heritable variation articulated via background assumptions about systems dynamics. Darwin’s theory of evolution was developed within a context of the background assumptions of Newtonian systems dynamics. The Modern Evolutionary Synthesis, or neo-Darwinism, successfully joined Darwinian selection and Mendelian genetics by developing population genetics informed by background assumptions of Boltzmannian systems dynamics. Currently the Darwinian Research Tradition (...) is changing as it incorporates new information and ideas from molecular biology, paleontology, developmental biology, and systems ecology. This putative expanded and extended synthesis is most perspicuously deployed using background assumptions from complex systems dynamics. Such attempts seek to not only broaden the range of phenomena encompassed by the Darwinian Research Tradition, such as neutral molecular evolution, punctuated equilibrium, as well as developmental biology, and systems ecology more generally, but to also address issues of the emergence of evolutionary novelties as well as of life itself. (shrink)

Recent theories in cognitive science have begun to focus on the active role of organisms in shaping their own environment, and the role of these environmental resources for cognition. Approaches such as situated, embedded, ecological, distributed and particularly extended cognition look beyond ‘what is inside your head’ to the old Gibsonian question of ‘what your head is inside of’ and with which it forms a wider whole—its internal and external cognitive niche. Since these views have been treated as a (...) radical departure from the received view of cognition, their proponents have looked for support to similar extended views within (the philosophy of) biology, most notably the theory of niche construction. This paper argues that there is an even closer and more fruitful parallel with developmental systems theory and developmental niche construction. These ask not ‘what is inside the genes you inherited’, but ‘what the inherited genes are inside of’ and with which they form a wider whole—their internal and external ontogenetic niche, understood as the set of epigenetic, social, ecological, epistemic and symbolic legacies inherited by the organism as necessary developmental resources. To the cognizing agent, the epistemic niche presents itself not just as a partially self-engineered selective niche, as the niche construction paradigm will have it, but even more so as a partially self-engineered ontogenetic niche, a problem-solving resource and scaffold for individual development and learning. This move should be beneficial for coming to grips with our own (including cognitive) nature: what is most distinctive about humans is their developmentally plastic brains immersed into a well-engineered, cumulatively constructed cognitive–developmental niche. (shrink)

This paper evaluates and criticises the developmental systems conception of evolution and develops instead an extension of the gene's eye conception of evolution. We argue (i) Dawkin's attempt to segregate developmental and evolutionary issues about genes is unsatisfactory. On plausible views of development it is arbitrary to single out genes as the units of selection. (ii) The genotype does not carry information about the phenotype in any way that distinguishes the role of the genes in development (...) from that other factors. (iii) There is no simple and general causal criterion which distinguishes the role of genes in development and evolution. (iv) There is, however, an important sense in which genes but not every other developmental factor represent the phenotype. (v) The idea that genes represent features of the phenotype forces us to recognise that genes are not the only, or almost the only, replicators. Many mechanisms of replication are involved in both development and evolution. (vi) A conception of evolutionary history which recognises both genetic and non-genetic replicators, lineages of replicators and interactors has advantages over both the radical rejection of the replicator/interactor distinction and the conservative restriction of replication to genetic replication. (shrink)

I respond to Karola Stotz's criticisms of my previously published challenges to the inference from developmental systems theory to an extended view of cognition.

Developmental systems theory is an attempt to sum up the ideas of a research tradition in developmental psychobiology that goes back at least to Daniel Lehrman’s work in the 1950s. It yields a representation of evolution that is quite capable of accommodating the traditional themes of natural selection and also the new results that are emerging from evolutionary developmental biology. But it adds something else - a framework for thinking about development and evolution without the (...) distorting dichotomization of biological processes into gene and non-gene and the vestiges of the ‘black-boxing’ of developmental processes in the modern synthesis, such as the asymmetric use of the concept of information. Phenomena that are marginalized in current gene-centric conceptions, such as extra-genetic inheritance, niche construction and phenotypic plasticity are placed center stage. (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)

Many things evolve: species, languages, sports, tools, biological niches, and theories. But are these real instances of natural selection? Current assessments of the proper scope of Darwinian theory focus on the broad similarity of cultural or non-organic processes to familiar central instances of natural selection. That similarity is analysed in terms of abstract functional descriptions of evolving entities (e.g. replicators, interactors, developmental systems etc). These strategies have produced a proliferation of competing evolutionary analyses. I argue that (...) such reasoning ought not to be employed in arbitrating debates about whether particular phenomena count as instances of natural selection. My argument is based on hierarchical functional descriptions of natural selection. I suggest that natural selection ought not to be thought of as a single process but rather as a series of processes which can be analysed in terms of a hierarchy of functional descriptions (in much the same way as many people think of cognition). This, in turn, casts doubt on the idea that it is possible in principle to settle debates about whether particular phenomena count as instances of natural selection. (shrink)

Similar to parasites, malignant cells exploit the host for energy, resources and protection, thereby impairing host health and fitness. Although cancer is widespread in the animal kingdom, its impact on life history traits and strategies have rarely been documented. Devil facial tumour disease, a transmissible cancer, afflicting Tasmanian devils, provides an ideal model system to monitor the impact of cancer on host life-history, and to elucidate the evolutionary arms-race between malignant cells and their hosts. Here we provide an overview (...) of parasite-induced host life history adaptations, then both phenotypic plasticity of LH responses and changes in allele frequencies that affect LH traits of Tasmanian devils in response to DFTD are discussed. We conclude that akin to parasites, cancer can directly and indirectly affect devil LH traits and trigger host evolutionary responses. Consequently, it is important to consider oncogenic processes as a selective force in wildlife. Exposure to transmissible cancer cells, i.e., Tasmanian devil facial tumour cells, can cause changes to life history traits either as the result of phenotypic plasticity in response to exposure during the lifetime of the individual or as the result of evolutionary change in LH traits due to selection over generations.Photo credits: Frédéric Thomas, Sarah Peck, Geordie Jennings. (shrink)

In 1988, David Hull presented an evolutionary account of science. This was a direct analogy to evolutionary accounts of biological adaptation, and part of a generalized view of Darwinian selection accounts that he based upon the Universal Darwinism of Richard Dawkins. Criticisms of this view were made by, among others, Kim Sterelny, which led to it gaining only limited acceptance. Some of these criticisms are, I will argue, no longer valid in the light of developments in the (...) formal modeling of evolution, in particular that of Sergey Gavrilets’ work on adaptive landscapes. If we can usefully recast the Hullian view of science as being driven by selection in terms of Gavrilets’ and Kaufmann’s view of there being “giant components” of high-fitness networks through any realistic adaptive landscape, we may now find it useful to ask what the adaptive pressures on science are, and to extend the metaphor into a full analogy. This is in effect to reconcile the Fisherianism of the Dawkins–Hull approach to selection and replicators, with a Wrightean drift account of social constructionist views of science, preserving, it is to be hoped, the valuable aspects of both. (shrink)

Among non-monotonic systems of reasoning, non-monotonic modal logics, and autoepistemic logic in particular, have had considerable success. The presence of explicit modal operators allows flexibility in the embedding of other approaches. Also several theoretical results of interest have been established concerning these logics. In this paper we introduce non-monotonic modal logics based on many-valued logics, rather than on classical logic. This extends earlier work of ours on many-valued modal logics. Intended applications are to situations involving several reasoners, not just (...) one as in the standard development. (shrink)

Conceptions of adaptation have varied in the history of genetic Darwinism depending on whether what is taken to be focal is the process of adaptation, adapted states of populations, or discrete adaptations in individual organisms. I argue that Theodosius Dobzhansky’s view of adaptation as a dynamical process contrasts with so-called “adaptationist” views of natural selection figured as “design-without-a-designer” of relatively discrete, enumerable adaptations. Correlated with these respectively process and product oriented approaches to adaptive natural (...) class='Hi'>selection are divergent pictures of organisms themselves as developmental wholes or as “bundles” of adaptations. While even process versions of genetical Darwinism are insufficiently sensitive to the fact much of the variation on which adaptive selection works consists of changes in the timing, rate, or location of ontogenetic events, I argue that articulations of the Modern Synthesis influenced by Dobzhansky are more easily reconciled with the recent shift to evolutionary developmentalism than are versions that make discrete adaptations central. (shrink)

Classical fixpoint semantics for logic programs is based on the TP immediate consequence operator. The Kripke/Kleene, three-valued, semantics uses ΦP, which extends TP to Kleene’s strong three-valued logic. Both these approaches generalize to cover logic programming systems based on a wide class of logics, provided only that the underlying structure be that of a bilattice. This was presented in earlier papers. Recently well-founded semantics has become influential for classical logic programs. We show how the well-founded approach also extends naturally (...) to the same family of bilatticebased programming languages that the earlier fixpoint approaches extended to. Doing so provides a natural semantics for logic programming systems that have already been proposed, as well as for a large number that are of only theoretical interest. And finally, doing so simplifies the proofs of basic results about the well-founded semantics, by stripping away inessential details. (shrink)

Two things are done in this paper. First, a modal logic in which one can quantify over both objects and concepts is presented; a semantics and a tableau system are given. It is a natural modal logic, extending standard versions, and capable of addressing several well-known philosophical difficulties successfully. Second, this modal logic is used to introduce a rather different way of looking at relational databases. The idea is to treat records as possible worlds, record entries as objects, and attributes (...) as concepts, in the modal sense. This makes possible an intuitively satisfactory relational database theory. It can be extended, by the introduction of higher types, to deal with multiple-valued attributes and more complex things, though this is further than we take it here. (shrink)

Most automated theorem provers have been built around some version of resolution [4]. But resolution is an inherently Classical logic technique. Attempts to extend the method to other logics have tended to obscure its simplicity. In this paper we present a resolution style theorem prover for Intuitionistic logic that, we believe, retains many of the attractive features of Classical resolution. It is, of course, more complicated, but the complications can be given intuitive motivation. We note that a small change in (...) the system as presented here causes it to collapse back to a Classical resolution system. We present the system in some detail for the propositional case, including soundness and completeness proofs. For the first order version we are sketchier. (shrink)

Two major clarifications have greatly abetted the understanding and fruitful expansion of the theory of natural selection in recent years: the acknowledgment that interactors, not replicators, constitute the causal unit of selection; and the recognition that interactors are Darwinian individuals, and that such individuals exist with potency at several levels of organization (genes, organisms, demes, and species in particular), thus engendering a rich hierarchical theory of selection in contrast with Darwin’s own emphasis on the organismic level. But (...) a piece of the argument has been missing, and individuals at levels distinct from organisms have been denied potency (although granted existence within the undeniable logic of the theory), because they do not achieve individuality with the same devices used by organisms and therefore seem weak by comparison. We show here that different features define Darwinian individuality across scales of size and time. In particular, species-individuals may develop few emergent features as direct adaptations. The interactor approach works with emergent fitnesses, not with emergent features; and species, as a consequence of their different mechanism for achieving individuality (reproductive exclusivity among subparts, that is, among organisms), express many effects from other levels. Organisms, by contrast, suppress upwardly cascading effects, because the organismic style of individuality (by functional integration of subparts) does not permit much competition or differential reproduction of parts from within. Species do not suppress the operation of lower levels; such effects therefore become available as exaptations conferring emergent fitness—a primary source of the different strength that species achieve as effective Darwinian individuals in evolution. (shrink)

Progress has become a suspect concept in evolutionary biology, not the least because the core concepts of neo-Darwinism do not support the idea that evolution is progressive. There have been a number of attempts to account for directionality in evolution through additions to the core hypotheses of neo-Darwinism, but they do not establish progressiveness, and they are somewhat of an ad hoc collection. The standard account of fitness and adaptation can be rephrased in terms of information (...) theory. From this, an information of adaptation can be defined in terms of a fitness function. The information of adaptation is a measure of the mutual information between biota and their environment. If the actual state of adaptation lags behind the state of optimal adaptation, then it is possible for the information of adaptation to increase indefinitely. Since adaptations are functional, this suggests the possibility of progressive evolution in the sense of increasing adaptation. Keywords: evolution, information, adaptation, fitness, entropy, progress.. (shrink)

William Keith Brooks was an American zoologist at Johns Hopkins University from 1876 until his death in 1908. Over the course of his career, Brooks staunchly defended Darwinism, arguing for the centrality of natural selection in evolutionary theory at a time when alternative theories, such as neo-Lamarckism, grew prominent in American biology. In his book The Law of Heredity, Brooks addressed problems raised by Darwin’s theory of pangenesis. In modifying and developing Darwin’s pangenesis, Brooks proposed a new theory (...) of heredity that sought to avoid the pitfalls of Darwin’s hypothesis. In so doing he strengthened Darwin’s theory of natural selection by undermining arguments for the inheritance of acquired characteristics. In later attacks on neo-Lamarckism, Brooks consistently defended Darwin’s theory of natural selection on logical grounds, continued to challenge the idea of the inheritance of acquired characteristics, and argued that natural selection best explained a wide range of adaptations. Finally, he critiqued Galton’s statistical view of heredity and argued that Galton had resurrected an outmoded typological concept of species, one which Darwin and other naturalists had shown to be incorrect. Brooks’s ideas resemble the “biological species concept” of the twentieth century, as developed by evolutionary biologist Ernst Mayr and others. The late-nineteenth century was not a period of total “eclipse” of Darwinism, as biologists and historians have hitherto seen it. Although the “Modern Synthesis” refers to the reconciliation of post-Mendelian genetics with evolution by natural selection, we might adjust our understanding of how the synthesis developed by seeing it as the culmination of a longer discussion that extends back to the late-nineteenth century. (shrink)

Using the evolution of the stickleback family of subarctic fish as a touchstone, we explore the effect of new discoveries about regulatory genetics, developmental plasticity, and epigenetic inheritance on the conceptual foundations of the Modern Evolutionary Synthesis. Identifying the creativity of natural selection as the hallmark of the Modern Synthesis, we show that since its inception its adherents have pursued a variety of research projects that at first seemed to conflict with its principles, but were accommodated. We (...) situate challenges coming from developmental biology in a dialectic between innovation and tradition, suggesting on the basis of past episodes that even if developmental plasticity and epigenetic inheritance are aligned with its principles the Modern Synthesis will be significantly affected. (shrink)

Understanding good design requires addressing the question of what units undergo natural selection, thereby becoming adapted. There is, therefore, a natural connection between the formal Darwinism project (which aims to connect population genetics with the evolution of design and fitness maximization) and levels of selection issues. We argue that the formal Darwinism project offers contradictory and confusing lines of thinking concerning level(s) of selection. The project favors multicellular organisms over both the lower (cell) and (...) higher (social group) levels as the level of adaptation. Grafen offers four reasons for giving such special status to multicellular organisms: (1) they lack appreciable within-organism cell selection, (2) they have multiple features that appear contrived for the same purpose, (3) they possess a set of phenotypes, and (4) they leave offspring according to their phenotypes. We discuss why these rationales are not compelling and suggest that a more even-handed approach, in which multicellular organisms are not assumed to have special status, would be desirable for a project that aims to make progress on the foundations of evolutionary theory. (shrink)

This paper gives a detailed narrative of a controversial empirical research in postwar population genetics, the analysis of the cytological polymorphisms of an Australian grasshopper, Moraba scurra. This research intertwined key technical developments in three research areas during the 1950s and 1960s: it involved Dobzhansky’s empirical research program on cytological polymorphisms, the mathematical theory of natural selection in two-locus systems, and the building of reliable estimates of natural selection in the wild. In the mid-1950s the cytologist Michael (...) White discovered an interesting case of epistasis in populations of Moraba scurra. These observations received a wide diffusion when theoretical population geneticist Richard Lewontin represented White’s data on adaptive topographies. These topographies connected the information on the genetic structure of these grasshopper populations with the formal framework of theoretical population genetics. As such, they appeared at the time as the most successful application of two-locus models of natural selection to an empirical study system. However, this connection generated paradoxical results: in the landscapes, all grasshopper populations were located on a ridge while they were expected to reach a peak. This puzzling result fueled years of research and triggered a controversy attracting contributors from Australia, the United States and the United Kingdom. While the original problem seemed, at first, purely empirical, the subsequent controversy affected the main mathematical tools used in the study of two-gene systems under natural selection. Adaptive topographies and their underlying mathematical structure, Wright’s mean fitness equations, were submitted to close scrutiny. Suspicion eventually shifted to the statistical machinery used in data analysis, reflecting the crucial role of statistical inference in applied population genetics. In the 1950s and 1960s, population geneticists were not simply in search for new generalizations about the evolutionary process and for new evidence about genetic variation: struggling against statistical artifacts, they were searching for reliable tests and descriptors to analyze their data. (shrink)

What role does non-genetic inheritance play in evolution? In recent work we have independently and collectively argued that the existence and scope of non-genetic inheritance systems, including epigenetic inheritance, niche construction/ecological inheritance, and cultural inheritance—alongside certain other theory revisions—necessitates an extension to the neo-Darwinian Modern Synthesis (MS) in the form of an Extended Evolutionary Synthesis (EES). However, this argument has been challenged on the grounds that non-genetic inheritance systems are exclusively proximate (...) mechanisms that serve the ultimate function of calibrating organisms to stochastic environments. In this paper we defend our claims, pointing out that critics of the EES (1) conflate non-genetic inheritance with early 20th-century notions of soft inheritance; (2) misunderstand the nature of the EES in relation to the MS; (3) confuse individual phenotypic plasticity with trans-generational non-genetic inheritance; (4) fail to address the extensive theoretical and empirical literature which shows that non-genetic inheritance can generate novel targets for selection, create new genetic equilibria that would not exist in the absence of non-genetic inheritance, and generate phenotypic variation that is independent of genetic variation; (5) artificially limit ultimate explanations for traits to gene-based selection, which is unsatisfactory for phenotypic traits that originate and spread via non-genetic inheritance systems; and (6) fail to provide an explanation for biological organization. We conclude by noting ways in which we feel that an overly gene-centric theory of evolution is hindering progress in biology and other sciences. (shrink)

Darwinian evolution, as it was first conceived, has two dimensions: adaptation, that is, selection based upon “apt function”, defined as the “good fit” between an organism’s metabolic and biological demands and the environment in which it is embedded; and heredity, the transmissible memory of past apt function. Modern Darwinism has come to focus almost exclusively on hereditary memory, eclipsing the—arguably still-problematic—phenomenon of adaptation. As a result, modern Darwinism retains, at its core, certain incoherencies that, as (...) long as they remain unresolved, preclude the emergence of a fully-coherent theory of evolution. Resolving the incoherencies will involve clarifying the relationship between embodied memory and apt function. In short, adaptation is a problem of semiotics: the organism must interpret the environment to fit well into it. This is well-illustrated by the constructed environments built by colonies of social insects, such as hives or nests, and the ancillary structures that contain them, forming an organism-like system known as a superorganism. The superorganism is marked by a kind of extended physiology, in that these constructed environments often serve as adaptive interfaces between the nest and ambient environment, and are constructed to manage the matter and energy flows between environments that constitute the process of adaptation. These constructed environments are also semiotic phenomena: interpretive structures, governed by information flow between the member insects and the structures they build. I review our findings on one such example: the mounds built by the fungus-cultivating termites of the genus Macrotermes. These structures are dynamic forms that are sustained by flows of soil from deep horizons up into the mound. The form, and hence the function, of the mound is determined by several environmental cues, most notably water and wind, as well as how termites interpret these cues, and signals that flow between termites, both directly and vicariously through the structures they build. (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)

Developmental Systems Theory is a theoretical reinterpretation of biological phenomena that challenges the conventional gene-centered account of development and evolution. In this article, I focus on Griffiths and Gray’s version of DST and particularly analyze their reconceptualization of inheritance. First, I present their concept of expanded and diffused inheritance; then, I examine and criticize their rejection of the multiple inheritance system model; finally, I present and oppose Griffiths and Gray’s extension of what they call the (...) “causal parity thesis” from development to evolution. I argue that their proposal is an interesting and programmatic philosophical perspective on biological phenomena but fails to provide either additional heuristic tools for empirical investigation in biology or a more realistic representation of the biological world. (shrink)

Natural selection causes adaptation, the fit between an organism and its environment. For example, the white and grey coloration of snowy owls living and breeding around the Arctic Circle provides camouflage from both predators and prey. In this Element, we explore a variety of such outcomes of the evolutionary process, including both adaptations and alternatives to adaptations, such as nonadaptive traits inherited from ancestors. We also explore how the concept of adaptation is used in evolutionary psychology and (...) in animal behavior, and the adequacy of methods used to confirm evolutionary accounts of human traits and behaviors. (shrink)

Two questions are raised for Grafen’s formal darwinism project of aligning evolutionary dynamics under natural selection with the optimization of phenotypes for individuals of a population. The first question concerns mean fitness maximization during frequency-dependent selection; in such selection regimes, not only is mean fitness typically not maximized but it is implausible that any parameter closely related to fitness is being maximized. The second question concerns whether natural selection on inclusive fitness (...) differences can be regarded as individual selection or whether it leads to a departure from the central motivation that led to the formal darwinism project, viz., to show that “Darwinian” evolution through individual selection leads to “good design” or phenotypic adaptation through trait optimization. (shrink)

Focused on the impact of stringent intellectual property mechanisms over the uses of plant agricultural biodiversity in crop improvement, the article delves into a systematic analysis of the relationship between institutional paradigms and their technological contexts of application, identified as mass selection, controlled hybridisation, molecular breeding tools and transgenics. While the strong property paradigm has proven effective in the context of major leaps forward in genetic engineering, it faces a systematic breakdown when extended to mass selection, where (...) innovation often displays a collective nature. However, it also creates partial blockages in those innovation schemes rested between on-farm observation and genetic modification, i.e. conventional plant breeding and upstream molecular biology research tools. Neither overly strong intellectual property rights, nor the absence of well delineated protection have proven an optimal fit for these two intermediary socio-technological systems of cumulative incremental innovation. To address these challenges, the authors look at appropriate institutional alternatives which can create effective incentives for in situ agrobiodiversity conservation and the equitable distribution of technologies in plant improvement, using the flexibilities of the TRIPS Agreement, the liability rules set forth in patents or plant variety rights themselves, and other ad hoc reward regimes. (shrink)

The broader context for the formal darwinism project established by two of the commentators, in terms of reconciling the Modern Synthesis with Darwinian arguments over design and in terms of links to other types of selection and design, is discussed and welcomed. Some overselling of the project is admitted, in particular of whether it claims to consider all organic design. One important fundamental question raised in two commentaries is flagged but not answered of whether design is rightly represented (...) by an optimisation program, and another from one commentary of whether the coreplicon dissolves in the face of multi-generational imprinting. Calls for the project to be extended to design at levels above and below the individual are considered sympathetically, but judged impractical at the high level of abstraction of the project. All claims of substantive technical error are emphatically rejected. Close technical readings are welcomed that, among other things, represent the project as 'axiomatizing fitness'. The prospects for the project are set out in the light of this highly varied set of commentaries. (shrink)

The paper outlines the contours of an organizational perspective on extended inheritance. Based on theoretical studies about biological organization and extended physiology, this perspective allows for the conception of extended biological legacies while keeping a theoretically indispensable distinction between biological systems and their environment. In this context, the line of demarcation between these systems and their surroundings is modelled on an organizational criterion and on the related conceptual distinction between organizational constraints, whose specific role (...) is to harness flows of matter and energy across generations of composite biological systems, and resources exploited by those systems. Biological legacies are restricted to persisting constitutive elements responsible for the reoccurrence of organizational constraints in a given environment. The case of symbiotic transmission is presented as a paradigmatic system illustrating the main proposed conceptual clarifications. (shrink)

Phenotype, whether conventional or extended, is defined as a reflectionof an underlying genotype. Adaptation and the natural selection thatfollows from it depends upon a progressively harmonious fit betweenphenotype and environment. There is in Richard Dawkins' notion ofthe extended phenotype a paradox that seems to undercut conventionalviews of adaptation, natural selection and adaptation. In a nutshell, ifthe phenotype includes an organism's environment, how then can theorganism adapt to itself? The paradox is resolvable through aphysiological, (...) as opposed to a genetic, theory of natural selection andadaptation. (shrink)

We raise the new possibility that people diagnosed with developmental dyslexia (DD) are specialized in explorative cognitive search, and rather than having a neurocognitive disorder, play an essential role in human adaptation. Most DD research has studied educational difficulties, with theories framing differences in neurocognitive processes as deficits. However, people with DD are also often proposed to have certain strengths – particularly in realms like discovery, invention, and creativity – that deficit-centered theories cannot explain. We investigate whether these (...) strengths reflect an underlying explorative specialization. We re-examine experimental studies in psychology and neuroscience using the framework ofcognitive search, whereby many psychological processes involve a trade-off between exploration and exploitation. We report evidence of an explorative bias in DD-associated cognitive strategies. High DD prevalence and an attendant explorative bias across multiple areas of cognition suggest the existence of explorative specialization. An evolutionary perspective explains the combination of findings and challenges the view that individuals with DD have a disorder. In cooperating groups, individual specialization is favored when features that confer fitness benefits are functionally incompatible. Evidence for search specialization suggests that, as with some other social organisms, humans mediate the exploration–exploitation trade-off by specializing in complementary strategies. The existence of a system of collective cognitive search that emerges through collaboration would help to explain our species’ exceptional adaptiveness. It also aligns with evidence for substantial variability during our evolutionary history and the notion that humans are adapted not to a particular habitat but to variability itself. Specialization creates interdependence and necessitates balancing complementary strategies. Reframing DD therefore underscores the urgency of changing certain cultural practices to ensure we do not inhibit adaptation. Key improvements would remove cultural barriers to exploration and nurture explorative learning in education, academia, and the workplace, as well as emphasize collaboration over competition. Specialization in complementary search abilities represents a meta-adaptation; through collaboration, this likely enables human groups (as a species and as cultural systems) to successfully adapt. Cultural change to support this system of collaborative search may therefore be essential in confronting the challenges humanity now faces. (shrink)

I argue that Grafen’s formal darwinism project could profitably incorporate a gene’s-eye view, as informed by the major transitions framework. In this, instead of the individual being assumed to maximise its inclusive fitness, genes are assumed to maximise their inclusive fitness. Maximisation of fitness at the individual level is not a straightforward concept because the major transitions framework shows that there are several kinds of biological individual. In addition, individuals have a definable fitness, exhibit individual-level (...) adaptations and arise in a major transition, only to the extent that the inclusive-fitness interests of genes within them coincide. Therefore, as others have suggested, the fundamental level at which fitness is maximised is the gene level. Previous reconciliations of the concepts of gene-level fitness and individual-level fitness implicitly recognise this point. Adaptations always maximise the fitness of their causative genes, but may be simple or complex. Simple adaptations may be controlled by single genes and be maladaptive at higher levels, whereas complex adaptations are controlled by multiple genes and rely on those genes having coinciding fitness interests at a higher level, for a given trait. (shrink)

The article proposes to further develop the ideas of the Extended Evolutionary Synthesis by including into evolutionary research an analysis of phenomena that occur above the organismal level. We demonstrate that the current Extended Synthesis is focused more on individual traits (genetically or non-genetically inherited) and less on community system traits (synergetic/organizational traits) that characterize transgenerational biological, ecological, social, and cultural systems. In this regard, we will consider various communities that are made up of interacting populations, and (...) for which the individual members can belong to the same or to different species. Examples of communities include biofilms, ant colonies, symbiotic associations resulting in holobiont formation, and human societies. The proposed model of evolution at the level of communities revises classic theorizing on the major transitions in evolution by analyzing the interplay between community/social traits and individual traits, and how this brings forth ideas of top-down regulations of bottom-up evolutionary processes (collaboration of downward and upward causation). The work demonstrates that such interplay also includes reticulate interactions and reticulate causation. In this regard, we exemplify how community systems provide various non-genetic ‘scaffoldings’, ‘constraints’, and ‘affordances’ for individual and sociocultural evolutionary development. Such research complements prevailing models that focus on the vertical transmission of heritable information, from parent to offspring, with research that instead focusses on horizontal, oblique and even reverse information transmission, going from offspring to parent. We call this reversed information transfer the ‘offspring effect’ to contrast it from the ‘parental effect’. We argue that the proposed approach to inheritance is effective for modelling cumulative and distributed developmental process and for explaining the biological origins and evolution of language. (shrink)

In this paper, I argue that the increasing data about non-genetic inheritance requires the construction of a new conceptual framework that should complement the inclusive approaches already discussed in the literature. More precisely, I hold that this framework should be epistemologically relevant for evolutionary biologists in capturing the limits of extended inheritance and in reassessing the boundaries of biological systems that transmit traits to their offspring. I outline the first elements of an organizational account of (...) class='Hi'>extended inheritance. In this account, the category of inherited factors is neither restricted to genes nor extended to stable resources related to trans-generational similarities. Instead, it includes persisting constitutive elements appearing as difference makers for heterogeneous organizational constraints, namely for heterogeneous constitutive parts whose specific role is to harness flows of matter and energy across generations of clearly delimited extended organized systems. This both inclusive and restrictive framework opens an additional way to apprehend how extended inheritance may affect evolutionary trajectories. (shrink)

The Shape of Thought: How Mental Adaptations Evolve presents a road map for an evolutionary psychology of the twenty-first century. It brings together theory from biology and cognitive science to show how the brain can be composed of specialized adaptations, and yet also an organ of plasticity. Although mental adaptations have typically been seen as monolithic, hard-wired components frozen in the evolutionary past, The Shape of Thought presents a new view of mental adaptations as diverse and variable, with distinct functions (...) and evolutionary histories that shape how they develop, what information they use, and what they do with that information. The book describes how advances in evolutionary developmental biology can be applied to the brain by focusing on the design of the developmental systems that build it. Crucially, developmental systems can be plastic, designed by the process of natural selection to build adaptive phenotypes using the rich information available in our social and physical environments. This approach bridges the long-standing divide between "nativist" approaches to development, based on innateness, and "empiricist" approaches, based on learning. It shows how a view of humans as a flexible, culturally-dependent species is compatible with a complexly specialized brain, and how the nature of our flexibility can be better understood by confronting the evolved design of the organ on which that flexibility depends. (shrink)

The Darwinian concept of natural selection was conceived within a set of Newtonian background assumptions about systems dynamics. Mendelian genetics at first did not sit well with the gradualist assumptions of the Darwinian theory. Eventually, however, Mendelism and Darwinism were fused by reformulating natural selection in statistical terms. This reflected a shift to a more probabilistic set of background assumptions based upon Boltzmannian systems dynamics. Recent developments in molecular genetics and paleontology have put pressure on (...) Darwinism once again. Current work on self-organizing systems may provide a stimulus not only for increased problem solving within the Darwinian tradition, especially with respect to origins of life, developmental genetics, phylogenetic pattern, and energy-flow ecology, but for deeper understanding of the very phenomenon of natural selection itself. Since self-organizational phenomena depend deeply on stochastic processes, self-organizational systems dynamics advance the probability revolution. In our view, natural selection is an emergent phenomenon of physical and chemical selection. These developments suggest that natural selection may be grounded in physical law more deeply than is allowed by advocates of the autonomy of biology, while still making it possible to deny, with autonomists, that evolutionary explanations can be modeled in terms of a deductive relationship between laws and cases. We explore the relationship between, chance, self-organization, and selection as sources of order in biological systems in order to make these points. (shrink)

Evolutionary adaptation has been suggested as the hallmark of life that best accounts for life’s creativity. However, current evolutionary approaches still fail to give an adequate account of it, even if they are able to explain both the origin of novelties and the proliferation of certain traits in a population. Although modern-synthesis Darwinism is today usually appraised as too narrow a position to cope with all the complexities of developmental and structural biology—not to say biosemiotic phenomena—, (...) class='Hi'>Darwinism need not be if we separate metaphor from reality in natural selection in order to show the axiological complexity of this concept. This can shed light on the relationship between biosemiotics and biological evolution. (shrink)