In The Selfish Gene, Richard Dawkins crystallized the gene's eye view of evolution developed by W.D. Hamilton and others. The book provoked widespread and heated debate. Written in part as a response, The Extended Phenotype gave a deeper clarification of the central concept of the gene as the unit of selection; but it did much more besides. In it, Dawkins extended the gene's eye view to argue that the genes that sit within an organism have an influence (...) that reaches out beyond the visible traits in that body - the phenotype - to the wider environment, which can include other individuals. So, for instance, the genes of the beaver drive it to gather twigs to produce the substantial physical structure of a dam; and the genes of the cuckoo chick produce effects that manipulate the behaviour of the host bird, making it nurture the intruder as one of its own. This notion of the extended phenotype has proved to be highly influential in the way we understand evolution and the natural world. It represents a key scientific contribution to evolutionary biology, and it continues to play an important role in research in the life sciences. The Extended Phenotype is a conceptually deep book that forms important reading for biologists and students. But Dawkins' clear exposition is accessible to all who are prepared to put in a little effort. Oxford Landmark Science books are 'must-read' classics of modern science writing which have crystallized big ideas, and shaped the way we think. (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)

While niche construction theory locates animal artefacts in their constructors’ environment, hence treating them as capable of exerting selective pressure on both the constructors and their descendants, the extended phenotype concept assimilates artefacts with their constructors’ genes. Analogous contrasts apply in the case of endoparasite and brood parasite genes influencing host behaviour. The explanatory power of these competing approaches are assessed by re-examining the core chapters of Richard Dawkins’ _The Extended Phenotype_. Because animal artefacts have multiple evolutionary (...) consequences for their constructors, the extra-body effects of a gene seemingly include feedback effects on multiple other genes, a result which is more consistent with niche construction theory than with selfish gene theory. In the case of endoparasite genes influencing host behaviour, Dawkins’ argument leaves out what appears to be the key explanatory component, namely the role of the host’s own bodily systems in making it possible for such genes to exist. For action at a distance, it is unclear whether the key genes have extended effects because they sit in the body of the manipulating organism, or alternatively do not have such effects because they sit in the body of its victim. It is argued that niche construction theory offers a superior explanation in all three cases, regardless of whether the extended phenotype concept is interpreted in selfish gene or selfish organism terms. (shrink)

I could summarise Being There unkindly, but not too inaccurately, as “Language of thought: bad. Everything else: good”. For the book is a fast and approving tour of a menagerie of alternatives to the classic computational theory of the mind. Some of the usual suspects appear— neural nets, connectionism—but they are not interrogated as deeply as the new members of the gang. For example, Clark walks us through some interesting new ideas in neuroscience. But none of the gang are pressed (...) very hard. Clark sketches a lot of interesting ideas but “sketches” is the right word. This is a book for the already sympathetic. For none of the examples are developed in the detail that would be required to convince a sceptic. A second issue concerns integration. Clark develops a number of themes as the book unwinds, but by the conclusion I was still uncertain about the extent to which the ideas in Being There are intended to be a package deal. (shrink)

The paper aims at proposing an extended notion of epigenesis acknowledging an actual causal import to the phenotypic dimension for the evolutionary diversification of life forms. “Introductory remarks” section offers introductory remarks on the issue of epigenesis contrasting it with ancient and modern preformationist views. In “Transmutation of forms: phenotypic variation, diversification, and complexification” section we propose to intend epigenesis as a process of phenotypic formation and diversification dependent on environmental influences, independent of changes in the genomic nucleotide sequence, (...) and occurring during the whole life span. Then, “Evolvability and phenotypic evolution” section focuses on phenotypic plasticity and offers an overview of basic properties that allows biological systems to be evolvable—i.e. to have the potentiality of producing phenotypic variation. Successively, the emphasis is put on environmentally-induced modifications in the regulation of gene expression giving rise to phenotypic variation and diversification. After some brief considerations on the debated issue of epigenetic inheritance, the issue of culture is considered. The key point is that, in the case of humans and of the evolutionary history of the genus Homo at least, the environment is also, importantly, the cultural environment. Thus, “Bio-cultural feedback” section argues that a bio-cultural feedback should be acknowledged in the “epigenic” processes leading to phenotypic diversification and innovation in Homo evolution. Finally, “Brain plasticity and cultural neural reuse” section introduces the notion of “cultural neural reuse”, which refers to phenotypic/neural modifications induced by specific features of the cultural environment that are effective in human cultural evolution without involving genetic changes. Therefore, cultural neural reuse may be regarded as a key instance of the bio-cultural feedback and ultimately of the extended notion of epigenesis proposed in this work. (shrink)

Phenotypic plasticity integrates the insights of ecological genetics, developmental biology, and evolutionary theory. Plasticity research asks foundational questions about how living organisms are capable of variation in their genetic makeup and in their responses to environmental factors. For instance, how do novel adaptive phenotypes originate? How do organisms detect and respond to stressful environments? What is the balance between genetic or natural constraints (such as gravity) and natural selection? The author begins by defining phenotypic plasticity and detailing its history, including (...) important experiments and methods of statistical and graphical analysis. He then provides extended examples of the molecular basis of plasticity, the plasticity of development, the ecology of plastic responses, and the role of costs and constraints in the evolution of plasticity. A brief epilogue looks at how plasticity studies shed light on the nature/nurture debate in the popular media. (shrink)

The concept of the digital phenotype has been used to refer to digital data prognostic or diagnostic of disease conditions. Medical conditions may be inferred from the time pattern in an insomniac’s tweets, the Facebook posts of a depressed individual, or the web searches of a hypochondriac. This paper conceptualizes digital data as an extended phenotype of humans, that is as digital information produced by humans and affecting human behavior and culture. It argues that there are ethical (...) obligations to persons affected by generalizable knowledge of a digital phenotype, not only those who are personally identifiable or involved in data generation. This claim is illustrated by considering the health-related digital phenotypes of precision medicine and digital epidemiology. (shrink)

The paper describes the context and the origin of a particular debate that concerns the evolution of phenotypic plasticity. In 1965, British biologist A. D. Bradshaw proposed a widely cited model intended to explain the evolution of norms of reaction, based on his studies of plant populations. Bradshaw’s model went beyond the notion of the “adaptive norm of reaction” discussed before him by Dobzhansky and Schmalhausen by suggesting that “plasticity” the ability of a phenotype to be modified by the (...) environment should be genetically determined. To prove Bradshaw’s hypothesis, it became necessary for some authors to identify the pressures exerted by natural selection on phenotypic plasticity in particular traits, and thus to model its evolution. In this paper, I contrast two different views, based on quantitative genetic models, proposed in the mid-1980s: Russell Lande and Sara Via’s conception of phenotypic plasticity, which assumes that the evolution of plasticity is linked to the evolution of the plastic trait itself, and Samuel Scheiner and Richard Lyman’s view, which assumes that the evolution of plasticity is independent from the evolution of the trait. I show how the origin of this specific debate, and different assumptions about the evolution of phenotypic plasticity, depended on Bradshaw’s definition of plasticity and the context of quantitative genetics. (shrink)

The current implementation of the Neo-Darwinian model of evolution typically assumes that the set of possible phenotypes is organized into a highly symmetric and regular space. Most conveniently, a Euclidean vector space is used, representing phenotypic properties by real-valued variables. Computational work on the biophysical genotype-phenotype model of RNA folding, however, suggests a rather different picture. If phenotypes are organized according to genetic accessibility, the resulting space lacks a metric and can be formalized only in terms of a relatively (...) unfamiliar structure. Patterns of phenotypic evolution—such as punctuation, irreversibility, and modularity—result naturally from the properties of the genotype-phenotype map, which, given the genetic accessibility structure, define accessibility in the phenotype space. The classical framework, however, addresses these patterns exclusively in terms of natural selection on suitably constructed fitness landscapes. Recent work has extended the explanatory level for phenotypic evolution from fitness considerations alone to include the topological structure of phenotype space as induced by the genotype-phenotype map. Lewontin’s notion of “quasi-independence” of characters can also be formalized in topological terms: it corresponds to the assumption that a region of the phenotype space is represented by a product space of orthogonal factors. In this picture, each character corresponds to a factor of a region of the phenotype space. We consider any region of the phenotype space that has a given factorization as a “type”, i.e., as a set of phenotypes that share the same set of phenotypic characters. Thus, a theory of character identity can be developed that is based on the correspondence of local factors in different regions of the phenotype space. (shrink)

Arguments that cognition or minds can be extended regularly invoke an analogy with Dawkins’ argument that phenotypes can be extended. I argue that there are two neglected ways in which those two boundary-breaking theses are complementary. Much of the argument of The Extended Phenotype concerns parasite phenotypes expressed in the behaviour of host organisms. But the options Dawkins considers for this extended manipulation are cognitively internalist. If we view cognition as extended, we can recognise (...) a wider range of vulnerabilities for exploitation. On the other hand, the analogies drawn with Dawkins almost always emphasise the benefit to the individual agent in being cognitively extended. Taking Dawkins’ concerns about manipulation and exploitation more seriously leads to a more contested, less optimistic picture of extended cognition and minds. This second line of thinking follows Sterelny’s lead, but I argue that hostility presents worse and more pervasive problems that he allows. (shrink)

Two critiques of simple adaptationism are distinguished: anti-adaptationism and extended adaptationism. Adaptationists and anti-adaptationists share the presumption that an evolutionary explanation should identify the dominant simple cause of the evolutionary outcome to be explained. A consideration of extended-adaptationist models such as coevolution, niche construction and extended phenotypes reveals the inappropriateness of this presumption in explaining the evolution of certain important kinds of features—those that play particular roles in the regulation of organic processes, especially behavior. These biological or (...) behavioral ‘levers’ are distinctively available for adaptation and exaptation by their possessors and for co-optation by other organisms. As a result they are likely to result from a distinctive and complex type of evolutionary process that conforms neither to simple adaptationist nor to anti-adaptationist styles of explanation. Many of the human features whose evolutionary explanation is most controversial belong to this category, including the female orgasm. (shrink)

I critically assess two widely cited evolutionary biological arguments for two versions of the ‘Extended Mind Thesis’ (EMT): namely, an argument appealing to Dawkins’s ‘Extended Phenotype Thesis’ (EPT) and an argument appealing to ‘Developmental Systems Theory’ (DST). Specifically, I argue that, firstly, appealing to the EPT is not useful for supporting the EMT (in either version), as it is structured and motivated too differently from the latter to be able to corroborate or elucidate it. Secondly, I extend (...) and defend Rupert’s argument that DST also fails to support or elucidate the EMT (in either version) by showing that the considerations in favour of the former theory have no bearing on the truth of the latter. I conclude by noting that the relevance of this discussion goes beyond the debate surrounding the EMT, as it brings out some of the difficulties of introducing evolutionary biological considerations into debates in psychology and philosophy more generally. (shrink)

The Genotype-Phenotype (G-P) distinction was proposed in the context of Mendelian genetics, in the wake of late 19th century studies about heredity. In this paper, we provide a conceptual analysis that highlights that the G-P distinction was grounded on three pillars: observability, transmissibility, and causality. Originally, the genotype is the non-observable and transmissible cause of the phenotype, which is its observable and non-transmissible effect. We argue that the current developments of biology have called the validity of such pillars (...) into question. First, molecular biology has unveiled the putative material substrate of the genotype (qua DNA), making it an observable object. Second, numerous findings on nongenetic heredity suggest that some phenotypic traits can be directly transmitted. Third, recent organicist approaches to biological phenomena have emphasized the reciprocal causality between parts of a biological system, which notably applies to the relations between genotypes and phenotypes. As a consequence, we submit that the G-P distinction has lost its general validity, although it can still apply to specific situations. This calls for forging new frameworks and concepts to better describe heredity and development. (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)

Evolution by natural selection has been extended to several supraorganismic levels, but whether it can apply to ecosystems remains controversial on two main counts. First, local ecosystems are loosely individuated, so that it is unclear how they manifest heredity and fitness. Second, even if they did, the meta-ecosystem formed by this population of local ecosystems will also suffer from a very low degree of cohesion, which will jeopardize any ENS. We suggest a way to overcome both issues, focusing on (...) ecosystem properties rather than on ecosystemic individuals. First, we follow recent theoretical developments which deny the centrality of reproduction in ENS. This leads us to merge heredity and fitness in a single process, in which a local ecosystem causally influences its neighbors, in a way responsible for their phenotypic similarity. Second, we suggest that the resulting meta-ecosystem need not meet the criteria of individuality. Instead, the evolving meta-ecosystem is best understood as a homeostatic property cluster, i.e., a set of phenotypes that together evolve by ENS at the ecosystem level. We illustrate this conceptual framework with a hypothetical example of local ecosystems that vary in terms of stability, productivity, diversity, and complementarity between species. Finally, we stress that our property-based account of ecosystemic ENS can help understand evolution at other biological levels, such as early life evolution and holobionts. (shrink)

The Modern Synthesis (MS) is the current paradigm in evolutionary biology. It was actually built by expanding on the conceptual foundations laid out by its predecessors, Darwinism and neo-Darwinism. For sometime now there has been talk of a new Extended Evolutionary Synthesis (EES), and this article begins to outline why we may need such an extension, and how it may come about. As philosopher Karl Popper has noticed, the current evolutionary theory is a theory of genes, and we still (...) lack a theory of forms. The field began, in fact, as a theory of forms in Darwin’s days, and the major goal that an EES will aim for is a unification of our theories of genes and of forms. This may be achieved through an organic grafting of novel concepts onto the foundational structure of the MS, particularly evolvability, phenotypic plasticity, epigenetic inheritance, complexity theory, and the theory of evolution in highly dimensional adaptive landscapes. (shrink)

In 1941/42 Konrad Lorenz suggested that Kant's transcendental categories ofa priori knowledge could be given an empirical interpretation in Darwinian material evolutionary terms: a priori propositional knowledge was an organ subject to natural selection for adaptation to its specific environments. D. Campbell extended the conception, and termed evolution a process of knowledge. The philosophical problem of what knowledge is became a descriptive one of how knowledge developed, the normative semantic questions have been sidestepped, as if the descriptive insights would (...) automatically resolve them. This came at a time when the traditional concept of knowledge as universally true, justified beliefs had been challenged by subjectivist, intercommunicative coherence frameworks. Much of the literature on evolutionary epistemology claimed that knowledge in general, and science as its epitome in particular, evolved along lines analogous to organic biological evolution. I refer here only to the view of knowledge as an extension of material biological evolution. These theories of evolutionary epistemology, contrary to the relativist notions of naturalized epistemology, adopted strict realist positions.Although there is no contention with the claim that biological evolution provided the raw material and the constraints for human knowledge, cognition is not knowledge and knowledge is not constrained by it beyond some trivial truisms. The view that sees evolution as a knowledge/cognition process is coercing a loosely defined term into the status of a phenotypic trait on which selection could act. This disregards the intricate many-to-many relationship between correlates of knowledge and biological capacities. But even if we grant the correlates of knowledge the status of selectable traits, the heritability of alternative phenotypes would be low and unpredictable due to the high, open-ended environmental malleability of such complex characters in the course of development. Such concepts are therefore biologically inconsequential. (shrink)

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

The question of whether the modern evolutionary synthesis requires an extension has recently become a topic of discussion, and a source of controversy. We suggest that this debate is, for the most part, not about the modern synthesis at all. Rather, it is about the extent to which genetic mechanisms can be regarded as the primary determinants of phenotypic characters. The modern synthesis has been associated with the idea that phenotypes are the result of gene products, while supporters of the (...) extended synthesis have suggested that environmental factors, along with processes such as epigenetic inheritance, and niche construction play an important role in character formation. We argue that the methodology of the modern evolutionary synthesis has been enormously successful, but does not provide an accurate characterization of the origin of phenotypes. For its part, the extended synthesis has yet to be transformed into a testable theory, and accordingly, has yielded few results. We conclude by suggesting that the origin of phenotypes can only be understood by integrating findings from all levels of the organismal hierarchy. In most cases, parts and processes from a single level fail to accurately explain the presence of a given phenotypic trait. The debate between the proponents of the modern and extended syntheses is somewhat reminiscent of the duck-rabbit illusion. The two sides are probably talking about the same thing, but from different perspectives. If not, then we argue that the challenge is to do an experiment that rules out the alternative view. (shrink)

How we ought to diagnose, categorise and respond to spectrum disabilities such as autism and Attention Deficit/Hyperactivity Disorder (ADHD) is a topic of lively debate. The heterogeneity associated with ADHD and autism is described as falling on various continua of behavioural, neural, and genetic difference. These continua are varyingly described either as extending into the general population, or as being continua within a given disorder demarcation. Moreover, the interrelationships of these continua are likewise often vague and subject to diverse interpretations. (...) -/- In this paper, I explore geneticists' and self-advocates’ perspectives concerning autism and ADHD as continua. These diagnoses are overwhelmingly analysed as falling on a continuum or continua of underlying traits, which supports the notion of “the neurodiversity spectrum”, i.e., a broader swath of human neural and behavioural diversity on which some concentrations of different functioning are diagnosed. I offer a taxonomy of conceptions of the genetic, phenotypic, and endophenotypic dimensionality within and beyond these diagnostic categories, and suggest that the spectrum of neurodiversity is characteristically endophenotypic. (shrink)

It is common in attempts to extend the theory of evolution to culture to generalize from the causal basis of biological evolution, so that evolutionary theory becomes the theory of copying processes. Generalizing from the formal dynamics of evolution allows greater leeway in what kinds of things cultural entities can be, if they are to evolve. By understanding the phenomenon of cultural transmission in terms of coordinated phenotypic variability, we can have a theory of cultural evolution which allows us to (...) avoid the various difficulties with the elaboration of informational entities such as the cultural replicator, or meme. Such an account is a boon to the project of evolutionary epistemology since it confirms the presumption in favor of the general adaptiveness of culture, illuminating rather than obscuring the inherent intimacy of our relationship to (e.g.) our ideas. (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)

Advocates of an ‘extended evolutionary synthesis’ have claimed that standard evolutionary theory fails to accommodate epigenetic inheritance. The opponents of the extended synthesis argue that the evidence for epigenetic inheritance causing adaptive evolution in nature is insufficient. We suggest that the ambiguity surrounding the conception of the gene represents a background semantic issue in the debate. Starting from Haig’s gene-selectionist framework and Griffiths and Neumann-Held’s notion of the evolutionary gene, we define senses of ‘gene’, ‘environment’, and ‘phenotype’ (...) in a way that makes them consistent with gene-centric evolutionary theory. We argue that the evolutionary gene, when being materialized, need not be restricted to nucleic acids but can encompass other heritable units such as epialleles. If the evolutionary gene is understood more broadly, and the notions of environment and phenotype are defined accordingly, current evolutionary theory does not require a major conceptual change in order to incorporate the mechanisms of epigenetic inheritance. _1_ Introduction _2_ The Gene-centric Evolutionary Theory and the ‘Evolutionary Gene’ _2.1_ The evolutionary gene _2.2_ Genes, phenotypes, and environments _3_ Epigenetic Inheritance and the Gene-Centred Framework _3.1_ Treating the gene as the sole heritable material? _3.2_ Epigenetics and phenotypic plasticity _4_ Conclusion. (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)

Adaptation by means of natural selection depends on the ability of populations to maintain variation in heritable traits. According to the Modern Synthesis this variation is sustained by mutations and genetic drift. Epigenetics, evodevo, niche construction and cultural factors have more recently been shown to contribute to heritable variation, however, leading an increasing number of biologists to call for an extended view of speciation and evolution. An additional common feature across the animal kingdom is learning, defined as the ability (...) to change behavior according to novel experiences or skills. Learning constitutes an additional source for phenotypic variation, and change in behavior may induce long lasting shifts in fitness, and hence favor evolutionary novelties. Based on published studies, I demonstrate how learning about food, mate choice and habitats has contributed substantially to speciation in the canonical story of Darwin’s finches on the Galapagos Islands. Learning cannot be reduced to genetics, because it demands decisions, which requires a subject. Evolutionary novelties may hence emerge both from shifts in allelic frequencies and from shifts in learned, subject driven behavior. The existence of two principally different sources of variation also prevents the Modern Synthesis from self-referring explanations. (shrink)

Evolutionary epistemology has experienced a continuous rise over the last decades. Important new theoretical considerations and novel empirical findings have been integrated into the existing framework. In this paper, I would like to suggest three lines of research that I believe will significantly contribute to further advance EE: ontogenetic considerations, key ideas from cognitive biology, and the framework of the Extended Evolutionary Synthesis. EE, in particular the program of the evolution of epistemological mechanisms, seeks to provide a phylogenetic account (...) of the generation of cognitive processes underlying knowledge creation. Traditionally, EE and EEM have been oriented towards an account of evolutionary theory that mainly drew from the tenets of the Modern Synthesis. The Modern Synthesis largely dismisses ontogenetic processes and considers them irrelevant for evolutionary explanations. If anything, the role of development in evolution is believed to be that of a constraint. There is, however, ample evidence for a tight intertwinement of developmental and evolutionary processes. Organisms employ their cognitive apparatus to interact with the environment in order to achieve a fully functioning perceptual and cognitive nervous system. Also, ontogeny provides generative potentials to enable variations that natural selection can act upon. EEM’s agenda may, therefore, strongly benefit from bringing together ontogenetic and phylogenetic approaches. To grapple with this challenge, an alternative vision of the evolutionary theory termed Extended Evolutionary Synthesis could be used. This extended evolutionary theory explores relationships between the processes of individual development and phenotypic change during evolution and can provide a more suitable framework for EEM to draw from. In recent years, cognitive biology has gained momentum as an independent research field. Cognitive biology builds on the concepts of EEM and understands knowledge as a biogenic phenomenon. Its main objective is also the formulation of substantiated interrelations between cognition and evolution but it focuses on cognitive functionality at all levels of biological organization. It thus employs a “vertical” approach that encompasses nested hierarchies which span from single molecules, cells, and tissues to the organismal level, communities, and societies. In contrast to cognitive biology, EEM is here understood to adopt a “horizontal” approach that focuses on phylogenetic explanations of cognition and knowledge acquisition. Linking EEM with the key ideas of cognitive biology could make EEM’s research program stronger as it can more easily accommodate phylogenetic and ontogenetic questions within a hierarchical, multilevel perspective. This is of particular importance for a more comprehensive account of cognition since living systems are subject to context-dependent causal influences from different organizational levels. In addition to EEM, there is a second program of EE. This program has been labeled evolutionary epistemology of theories and understands the increase in human knowledge, such as scientific theories, as naturalistic accounts of evolution. Both, EEM and EET initially drew from the core concepts of the Modern Synthesis. Several scholars have severely criticized the analogies made between EET and the Neo-Darwinian key processes of evolution. In particular processes of random mutation, the rate of variation, natural selection as the unique driving force, and the adaptationist agenda are believed to reveal disanalogies. In contrast to the Modern Synthesis, the Extended Evolutionary Synthesis not only recognizes developmental processes but also ecological interactions and systems dynamics as well as social and cultural evolutionary reciprocity as important evolutionary processes. Concepts of the Extended Evolutionary Synthesis are therefore expected to be more fruitful for re-conceptualizing parallels between scientific theorizing and biological evolution. (shrink)

The aim of this contribution is to investigate certain selected parts of the extended evolutionary synthesis which all have a common denominator, namely evolution by meaning attribution. We start by arguing that living organisms can manipulate and interpret their genetic script via epigenetic modifications in a semiotic manner, that is, by meaning attribution. Genes do not build living beings to be transmitted to future generations. Genes have been shaped by evolution as a memory medium that is transmitted from one (...) generation to the next, but the actual reading of such scripts is modified by momentary contexts. Secondly, we show that phenotypic evolution variously re-uses already existing homologies which in evolving systems acquire a new meaning. We also suggest that the ways in which organisms perceive their environment and other living beings is an important but still largely neglected evolutionary force. Variations in perception influence the direction and intensity of sexual selection and some behaviourally mediated regimes of natural selection. Thirdly, we point out that especially if we want to study their evolution, living beings should not be considered in isolation but in their mutual coexistence, in their historically established being together. Recent attempts to view living beings as constructors of niches and holobionts seem compatible with the classical Umwelt theory. This approach seems capable of accounting for both competitiveness and cooperation, thus making the overall image of evolution more comprehensive. And finally, we argue that if we want to expand our understanding of biological evolution, in addition to variation, selection, and inheritance we also need to take into account processes which participate in meaning attribution. (shrink)

I argue that the Conceptual Ethics and Conceptual Engineering framework, in its pragmatist version as recently defended by Thomasson, provides a means of articulating and defending the conventionalist interpretation of projects of conceptual extension (e.g. the extended mind, the extended phenotype) in biology and psychology. This promises to be illuminating in both directions: it helps to make sense of, and provides an explicit methodology for, pragmatic conceptual extension in science, while offering further evidence for the value and (...) fruitfulness of the Conceptual Ethics/Engineering framework itself, in particular with respect to conceptual change within science, which has thus-far received little attention in the literature on Conceptual Ethics/Engineering. (shrink)

Since Richard Dawkins ' The Extended Phenotype got me to thinking about copying units in the mid-1980s, I have been trying to define a cerebral code by searching for what can be successfully replicated in the brain's neural circuitry, a minimum replicable unit. I indeed found such circuitry. But to explore creativity in higher intellectual function, I wanted to see if the resulting copies could compete in a Darwinian manner, the process shaping up quality as it goes. And (...) that forced me to try and boil down a lot of evolutionary biology, attempting to abstract the features that were essential from those that merely contributed to speed or stability. This isn't the place to describe the neural outcome -- it's in The Cerebral Code and, more briefly, in the seventh chapter of my other 1996 book, How Brains Think -- but this does seem an appropriate place to review what I started calling "The Six Essentials." They seem applicable to a wide range of problems within memetics9 as the field attempts to cope with evolutionary models of information transmission. For a more. (shrink)

My aim in this paper is to quickly sketch a teleological approach to the problem of isolating the impact of genes on phenotypic characters. I begin by arguing that it is a mistake to think that there will be only one analysis of genetic input suitable for all theoretical interests. My principle focus is Richard Dawkins' argument for genic selectionism. I argue that a teleological analysis of genetic input is what Dawkins requires to establish the right kind of mapping of (...) gene onto phenotype. This comes at a certain cost, however. Accepting the analysis will threaten Dawkins' claims about the teleogogical priority of gene over phenotype. (shrink)

Most accounts of cognitive scaffolding focus on ways that external structure can support or augment an agent’s cognitive capacities. We call cases where the interests of the user are served benign scaffolding and argue for the possibility and reality of hostile scaffolding. This is scaffolding which depends on the same capacities of an agent to make cognitive use of external structure as in benign cases, but that undermines or exploits the user while serving the interests of another agent. We develop (...) criteria for scaffolding being hostile and show by reference to examples including the design features of electronic gambling machines and casino management systems that hostile scaffolding exists and can be highly effective. In cases where the scaffolding is deep and permits the offloading of significant cognitive work, hostile scaffolding exploitatively manipulates cognitive processing itself. Given the extent of human reliance on scaffolding this is an important and neglected vulnerability. [This is an updated version of the paper, added in February 2023.]. (shrink)

Extending the approach to a ‘theology of science’ developed in Faith and Wisdom in Science, I expand its theme of the tension between chaos and emergent order, within the arc of the Biblical story of creation, towards a theology of evolutionary science. In addition to the material in Job, the book of Wisdom provides a remarkable account of transmutation of species, within a recapitulation of the Exodus theme, that I juxtapose with a modern genotype-phenotype theory of evolutionary dynamics, exploiting (...) analogies with statistical mechanics. The dual and connected structures of microscopic and macroscopic provide a locus for the Joban tensions of chaos and emergent order, and provide an interpretative narrative for the emergent directionality of evolution, and a theology that situates within a creation of freedom to explore the potential of the created order. (shrink)

Have Sober and Wilson salvaged a sophisticated and sound perspective for group selection from the rhetorical overkill of the selfish-gene’s-eye gang, or have they merely reinvented Hamilton’s and Maynard Smith’s alternative to group selection models, models that can do justice to all the observed and even imagined phenomena of cooperation in the biosphere? One of the main lessons I have learned in thinking about the issues raised by Unto Others over the last two years is that they are, at least (...) for me, mind-twistingly elusive and slippery. The appeal of the competing metaphors is such that there is unwitting sleight of hand in every direction, as the perspectives shift back and forth. One stern admonition might then be: eschew the metaphors, stick to the math! The problem is that the math doesn’t distinguish between the perspectives Sober and Wilson champion and those they oppose. By their own account, the mathematical models are equivalent in what they can predict—and hence equivalent in what they can, in one important sense, explain. But that is not the end of it: “When one theory achieves an insight by virtue of its perspective, the same insight can usually be explained in retrospect by the other theories. As long as the relationships among the theories are clearly understood, this kind of pluralism is a healthy part of science.”. Good point. Patrick Suppes used to delight in challenging cognitive psychologists to point to any “cognitive” phenomenon that he couldn’t model, retrospectively, in strict behaviorist terms. The wise response was to deflect the challenge. His behaviorist models were parasitic: they would never have been devised—the phenomena in question were all but invisible to the behaviorists—without the inspiration of the cognitive model they translated into behaviorese. And what Sober and Wilson say here echoes what one of their chief opponents, Richard Dawkins, had already claimed for his perspective: “The extended phenotype may not constitute a testable hypothesis in itself, but it so far changes the way we see animals and plants that it may cause us to think of testable hypotheses that we would otherwise never have dreamed of.” So what’s all the fuss about? If each side is just saying “Here’s a good way of thinking of things, a prolific generator of hypotheses to test!” they could both be right, with their different perspectives having rather different utilities in different circumstances. (shrink)

In modern evolutionary theory, selection acts on particular genes and assemblages of genes that operate through phenotypes expressed in environments. This view, however, overlooks the fact that organisms often alter their environments in pursuit of fitness needs and thus modify some environmental selection pressures. Niche construction theory introduces a reciprocal causal process that modifies natural selection relative to three general kinds of environmental components: abiota, biota (other organisms), and artifacts. The ways in which niche-constructing organisms can construct or modify the (...) components differ. Modification of abiota, for example, may have different consequences from the construction of artifacts. Some changes in abiota may simply be caused by the by-products of metabolisms and activities of organisms. Alternatively, artifacts may be “extended phenotypes” that demonstrate obvious prior “design” and “construction” by organisms in the service of fitness needs. Nevertheless, adaptation should always account for the reciprocity between constructed niches and the living agents that construct them. Looking to well-adapted nature for inspiration for human-built artifacts must account for this reciprocity between phenotype and constructed environment as well as the novel features of human architecture, including frank intentionality of design and novel culturally acquired knowledge. (shrink)

Historically, the empirical study of phenotypic diversification has fallen into two rough camps; (1) "structuralist approaches" focusing on developmental constraint, bias, and innovation (with evo-devo at the core); and (2) "adaptationist approaches" focusing on adaptation, and natural selection. Whilst debates, such as that surrounding the proposed "Extended" Evolutionary Synthesis, often juxtapose these two positions, this review focuses on the grey space in between. Specifically, here I present a novel analysis of structuralism which enables us to take a more nuanced (...) look at the motivations behind the structuralist and adaptationist positions. This makes clear how the two approaches can conflict, and points of potential commensurability. The review clarifies (a) the value of the evo-devo approach to phenotypic diversity, but also (b) how it properly relates to other predominant approaches to the same issues in evolutionary biology more broadly. (shrink)

This note argues that the charge of reductionism levelled against Richard Dawkins is false. It does so by examining the development of his notion of the genes in his books The Selfish Gene (TSG), and The Extended Phenotype (TEP).

We have values and aspirations. What of other animals? Are their "values" different from ours? Animals manifestly prefer having plenty of food to starvation, and comfort to pain, and they will work hard to obtain a mate. But beyond these "creature comforts," they seem to be largely indifferent to the prospects and anxieties that make up human life. A suitable coverall term for human aspiration would be the pursuit of happiness, bearing in mind that happiness is many different things to (...) different people. This already sets us aside from our fellow creatures. To put it vividly, Mother Nature doesn't care whether we are happy--but we care (and Mother Nature doesn't care that we care). That is, it would be naive to suppose that the process of natural selection has somehow endorsed our pursuit of happiness as the proximal mechanism for maximizing our genetic fitness. It is consistent with what we know of evolution to suppose that the process of natural selection--Mother Nature--would design us to experience however much anxiety and torment is consistent with making more grandchildren. Our values are, like everything else in our extended phenotypes, products of evolutionary processes, but we misread them if we see them to be just like the "values" of other animals, which can indeed be viewed as the straightforward result of Mother Nature's project of installing an optimally reliable fitness-enhancing set of preferences. The difference arises, I will argue, from the fact that we have culture, and culture provides a medium in which a radically different--indeed, orthogonal--set of selection pressures can re-direct evolutionary processes into unprecedented channels. (shrink)

We consider, from a physical perspective, the case where the interface between an organism and its environment becomes large enough that it acts as a buffer regulating their matter and energy exchanges. We illustrate the physiological and evolutionary role of buffers through the example of lungfish estivation. Then we ponder the relevance of buffers of this kind to the quest for a general definition of concepts like niche construction, the extended phenotype, and related ones, whose meaning is conveyed (...) at present mostly through particular examples. Finally, we comment on the potential significance of buffers to organism—environment codetermination in the sense originally suggested by Lewontin. (shrink)

Understanding community-level selection using Lewontin’s criteria requires both community-level inheritance and community-level heritability, and in the discipline of community and ecosystem genetics, these are often conflated. While there are existing studies that show the possibility of both, these studies impose community-level inheritance as a product of the experimental design. For this reason, these experiments provide only weak support for the existence of community-level selection in nature. By contrast, treating communities as interactors (in line with Hull’s replicator-interactor framework or Dawkins’s idea (...) of the “extended phenotype”) provides a more plausible and empirically supportable model for the role of ecological communities in the evolutionary process. (shrink)

Electronics can be defined as electromagnetic technology dealing with information, and meta-electronics as a field encompassing all the synergistic technologies in which electronics plays a dominant role. Examining the broad field corresponding to this definition we realize that its history starts some seventy years earlier than the customarily accepted birth of electronics, and, what is more significant, that electronics undergoes a true evolution. This new evolution creates rich, diverse structures similar to those created by the biological evolution. Like biology, electronics (...) is non-teleological, which allows for its unlimited evolutionary development. We propose electronic analogies of all essential biological categories, at all levels: population, speciation, common ancestor, phenotype, extended phenotype, co-evolution, convergent evolution, evolutionary arms race, extinction and mass extinction, hierarchical levels, generative entrenchment, genes, alleles, genome, genetic pool, recombination, mutation, genetic drift, lateral gene transfer, etc. The evolutionary algorithm operating in electronics, like a Darwinian one, includes variation within a population of device models, heredity, natural selection, and a form of selection based on aesthetics and fashion which resembles sexual selection. Algorithm is especially similar to artificial selection, thus possessing directionality in the variational part. Electronic development is orders of magnitude faster than biological, accelerated by that directionality and by other distinct, identifiable mechanisms. Speciation in electronics, as in biology, is best represented on a phylogenetic tree, which starts from a common ancestor, but lately exhibits a unification trend. If continued, this trend may lead to the appearance of a common descendant absent in biology. Our analysis may explain emerging anti-social aspects of electronics and our conclusions add new urgency to recent concerns with unchecked development of Artificial Intelligence. (shrink)

This paper contends that law is in essence an evolutionary phenomenon that can, and indeed should, be studied in the light of biological mechanisms. Law can be seen as an extended phenotype of underlying genes. In addition, legal systems can be seen as congruous to genetic mechanisms. Properties of genes have an impact on legal systems in a fractal-like manner. Hence, it is not surprising that notions of stability, replication, and reciprocity that are important in biological systems will (...) also be important in legal systems. As a result legal systems can be constructed in a way that is congruent with the genetic advantage of group members. Law, exposure, and punishment can diminish deviant behaviour and restore balance. Law may not be particularly subject to natural selection, but it will certainly be built on the foundations of natural selection. (shrink)

We examine the self-deceptive aspects of religion and nationalism. By embracing various religious or political ideals, regardless of their truth, our ancestors could have enhanced their confidence, solidified their social ties, and manipulated their reproductive rivals. This use of culture as one's extended phenotype may increase the spread of misinformation and create global webs of deception and self-deception.

Evolutionary systems biology (ESB) is a rapidly growing integrative approach that has the core aim of generating mechanistic and evolutionary understanding of genotype‐phenotype relationships at multiple levels. ESB's more specific objectives include extending knowledge gained from model organisms to non‐model organisms, predicting the effects of mutations, and defining the core network structures and dynamics that have evolved to cause particular intracellular and intercellular responses. By combining mathematical, molecular, and cellular approaches to evolution, ESB adds new insights and methods to (...) the modern evolutionary synthesis, and offers ways in which to enhance its explanatory and predictive capacities. This combination of prediction and explanation marks ESB out as a research manifesto that goes further than its two contributing fields. Here, we summarize ESB via an analysis of characteristic research examples and exploratory questions, while also making a case for why these integrative efforts are worth pursuing. (shrink)