Both biologists and philosophers often make use of simple verbal formulations of necessary and sufficient conditions for evolution by natural selection (ENS). Such summaries go back to Darwin's Origin of Species (especially the "Recapitulation"), but recent ones are more compact.1 Perhaps the most commonly cited formulation is due to Lewontin.2 These summaries tend to have three or four conditions, where the core requirement is a combination of variation, heredity, and fitness differences. The summaries are employed in several ways. First, they (...) are often used in pedagogical contexts, and in showing the coherence of evolutionary theory in response to attacks from outside biology. Second, they are important in discussions of extensions of evolutionary principles to new domains, such as cultural change. The summaries also have intrinsic scientific and philosophical interest as attempts to capture some core principles of evolutionary theory in a highly concise way. Despite their prominence, both the proper formulation and status of these summaries are unclear. Standard formulations are subject to counterexamples, and their relations to formal models of evolutionary change are not straightforward. Here I look closely at these verbal summaries, and at how they relate to formal models. Are the summaries merely rough approximations that have no theoretical role of their own? Perhaps they could operate as theoretical statements in Darwin's time, but have now been superseded by more exact treatments. (shrink)

Debates concerning the units and levels of selection have persisted for over fifty years. One major question in this literature is whether units and levels of selection are genuine, in the sense that they are objective features of the world, or merely reflect the interests and goals of an observer. Scientists and philosophers have proposed a range of answers to this question. This Element introduces this literature and proposes a novel contribution. It defends a realist stance and offers a way (...) of delineating genuine levels of selection by invoking the notion of a functional unit. (shrink)

To naturalize the concept of teleological causality in biology it is not enough to avoid assuming backward causation or positing the existence of an inscrutable te- leological essence like the élan vital. We must also specify how the causality of or- ganisms is distinct from the causality of designed artifacts like thermostats or asym- metrically oriented processes like the ubiquitous increase of entropy. Historically, the concept of teleological causality in biology has been based on an analogy to the familiar experience (...) of purposeful action. This is experienced by us as a disposition to achieve a general type of end that is represented in advance, and which regulates the selection of efficient means to achieve it. Inspired by this analogy, to bridge the gap between biology and human agency we describe a simple molecular pro- cess called autogenesis that shows how two linked complementary self-organizing processes can give rise to higher-order relations that resemble purposeful disposi- tions, though expressed in terms of constraints on molecular processes. Because the autogenic model is described in sufficient detail to be empirically realizable, it provides a proof of principle demonstrating a simple form of teleological causality. (shrink)

We assess the arguments for recognising functionally integrated multispecies consortia as genuine biological individuals, including cases of so-called ‘holobionts’. We provide two examples in which the same core biochemical processes that sustain life are distributed across a consortium of individuals of different species. Although the same chemistry features in both examples, proponents of the holobiont as unit of evolution would recognize one of the two cases as a multispecies individual whilst they would consider the other as a compelling case of (...) ecological dependence between separate individuals. Some widely used arguments in support of the ‘holobiont’ concept apply equally to both cases, suggesting that those arguments have misidentified what is at stake when seeking to identify a new level of biological individuality. One important aspect of biological individuality is evolutionary individuality. In line with other work on the evolution of individuality, we show that our cases can be distinguished by focusing on the fitness alignment between the partners of the consortia. We conclude that much of the evidence currently presented for the ubiquity and importance of multi-species individuals is simply not to the point, at least unless the issue of biological individuality is firmly divorced from the question of evolutionary individuality. (shrink)

An idea is not a replicator because it does not consist of coded self-assembly instructions. It may retain structure as it passes from one individual to another, but does not replicate it. The cultural replicator is not an idea but an associatively-structured network of them that together form an internal model of the world, or worldview. A worldview is a primitive, uncoded replicator, like the autocatalytic sets of polymers widely believed to be the earliest form of life. Primitive replicators generate (...) self-similar structure, but because the process happens in a piecemeal manner, through bottom-up interactions rather than a top-down code, they replicate with low fidelity, and acquired characteristics are inherited. Just as polymers catalyze reactions that generate other polymers, the retrieval of an item from memory can in turn trigger other items, thus cross-linking memories, ideas, and concepts into an integrated conceptual structure. Worldviews evolve idea by idea, largely through social exchange. An idea participates in the evolution of culture by revealing certain aspects of the worldview that generated it, thereby affecting the worldviews of those exposed to it. If an idea influences seemingly unrelated fields this does not mean that separate cultural lineages are contaminating one another, because it is worldviews, not ideas, that are the basic unit of cultural evolution. (shrink)

Humans possess great capacity for behavioral and cultural change, but our ability to manage change is still limited. This article has two major objectives: first, to sketch a basic science of intentional change centered on evolution; second, to provide examples of intentional behavioral and cultural change from the applied behavioral sciences, which are largely unknown to the basic sciences community.All species have evolved mechanisms of phenotypic plasticity that enable them to respond adaptively to their environments. Some mechanisms of phenotypic plasticity (...) count as evolutionary processes in their own right. The human capacity for symbolic thought provides an inheritance system having the same kind of combinatorial diversity as does genetic recombination and antibody formation. Taking these propositions seriously allows an integration of major traditions within the basic behavioral sciences, such as behaviorism, social constructivism, social psychology, cognitive psychology, and evolutionary psychology, which are often isolated and even conceptualized as opposed to one another.The applied behavioral sciences include well-validated examples of successfully managing behavioral and cultural change at scales ranging from individuals to small groups to large populations. However, these examples are largely unknown beyond their disciplinary boundaries, for lack of a unifying theoretical framework. Viewed from an evolutionary perspective, they are examples of managing evolved mechanisms of phenotypic plasticity, including open-ended processes of variation and selection.Once the many branches of the basic and applied behavioral sciences become conceptually unified, we are closer to a science of intentional change than one might think. (shrink)

According to a once influential view of selection, it consists of repeated cycles of replication and interaction. It has been argued that this view is wrong: replication is not necessary for evolution by natural selection. I analyze the nine most influential arguments for this claim and defend the replication–interaction conception of selection against these objections. In order to do so, however, the replication–interaction conception of selection needs to be modified significantly. My proposal is that replication is not the copying of (...) an entity, the replicator, but the copying of a property. Thus, we can have a replication process without there being a replicator that is being copied. (shrink)

Recent debates between proponents of the modern evolutionary synthesis (the standard model in evolutionary biology) and those of a possible extended synthesis are a good example of the fascinating tangle among empirical, theoretical, and conceptual or philosophical matters that is the practice of evolutionary biology. In this essay, we briefly discuss two case studies from this debate, highlighting the relevance of philosophical thinking to evolutionary biologists in the hope of spurring further constructive cross-pollination between the two fields.

Darwin's theory of evolution by natural selection provided the first, and only, causal-mechanistic account of the existence of adaptations in nature. As such, it provided the first, and only, scientific alternative to the “argument from design”. That alone would account for its philosophical significance. But the theory also raises other philosophical questions not encountered in the study of the theories of physics. Unfortunately the concept of natural selection is intimately intertwined with the other basic concepts of evolutionary theory—such as the (...) concepts of fitness and adaptation —that are themselves philosophically controversial. Fortunately we can make considerable headway in getting clear on natural selection without solving all of those outstanding problems. (shrink)

The hologenome concept of evolution is a hypothesis about the evolution of animals and plants. It asserts that the evolution of animals and plants was partially triggered by their interactions with their symbiotic microbiomes. In that vein, the hologenome concept posits that the holobiont (animal host + symbionts of the microbiome) is a unit of selection. -/- The hologenome concept has been severely criticized on the basis that selection on holobionts would only be possible if there were a tight transgenerational (...) host-genotype-to-symbiont-genotype connection. As our current evidence suggests that this is not the case for most of the symbiont species that compose the microbiome of animals and plants, the opportunity for holobiont selection is very low in relation to the opportunity for selection on each of the species that compose the host microbiome. Therefore, holobiont selection will always be disrupted ‘from below’, by selection on each of the species that compose the microbiome. -/- This thesis constitutes a conceptual effort to defend philosophically the hologenome concept. I argue that the criticism according to which holobiont selection requires tight transgenerational host-genotype-to-symbiont-genotype connection is grounded on a metaphysical view of the world according to which the biological hierarchy needs to be nested, such that each new level of selection includes every entity from below. Applied to hologenomes, it entails that the hologenome is a collection of genomes, and selection of hologenomes is assumed to entail cospeciation of the host with the species that constitute its microbiome. -/- Against that interpretation, I propose the ‘stability of traits’ account, according to which hologenome evolution is the result of the action of natural selection in a non-nested hierarchical world. In that vein, hologenome evolution does not entail cospeciation, and thus it does not require tight transgenerational host-genotype-to-symbiont-genotype connection. By embracing a multilevel selection perspective, I argue that hologenome evolution results from the simultaneous action of natural selection on each of the lineages that compose the microbiome, and on the assemblage composed by the host genome plus the functional traits of its microbiome. Hologenome selection occurs when the evolution of the traits of the microbiome result from their effects on the fitness of the host, and it can take the form of multilevel selection 1, or multilevel selection 2. In both cases, hologenome selection entails the evolution of microbiome traits, as well as evolution of the host genome, rather than cospeciation of lineages. (shrink)

Reydon and Scholz raise doubts about the Darwinian status of organizational ecology by arguing that Darwinian principles are not applicable to organizational populations. Although their critique of organizational ecology’s typological essentialism is correct, they go on to reject the Darwinian status of organizational populations. This paper claims that the replicator-interactor distinction raised in modern philosophy of biology but overlooked for discussion by Reydon and Scholz provides a way forward. It is possible to conceptualize evolving Darwinian populations providing that the inheritance (...) mechanism is appropriately specified. By this approach, adaptation and selection are no longer dichotomized, and the evolutionary significance of knowledge transmission is highlighted. (shrink)

One of the most influential arguments against the claim that computers can think is that while our intentionality is intrinsic, that of computers is derived: it is parasitic on the intentionality of the programmer who designed the computer-program. Daniel Dennett chose a surprising strategy for arguing against this asymmetry: instead of denying that the intentionality of computers is derived, he endeavours to argue that human intentionality is derived too. I intend to examine that biological plausibility of Dennett’s suggestion and show (...) that Dennett’s argument for the claim that human intentionality is derived because it was designed by natural selection is based on the misunderstanding of how natural selection works. (shrink)

The improbability of a spontaneously generated self-assembling molecule has suggested that life began with a set of simpler, collectively replicating elements, such as an enclosed autocatalytic set of polymers (or autocell). Since replication occurs without a self-assembly code, acquired characteristics are inherited. Moreover, there is no strict distinction between alive and dead; one can only infer that an autocell was alive if it replicates. These features of early life render natural selection inapplicable to the description of its change-of-state because they (...) defy its underlying assumptions. Moreover, natural selection describes only randomly generated novelty; it cannot describe the emergence of form at the interface between organism and environment. Self-organization is also inadequate because it is restricted to interactions amongst parts; it too cannot account for context-driven change. A modified version of selection theory or self-organization would not work because the description of change-of-state through interaction with an incompletely specified context has a completely different mathematical structure, i.e. entails a non-Kolmogorovian probability model. It is proposed that the evolution of early life is appropriately described as lineage transformation through context-driven actualization of potential, with self-organized change-of-state being a special case of no contextual influence, and competitive exclusion of less fit individuals through a selection-like process possibly (but not necessarily) playing a secondary role. It is argued that natural selection played an important role in evolution only after genetically mediated replication was established. (shrink)

For a number of years, the debate in evolutionary biology over the ’levels of selection’ has attracted intense interest from philosophers of science. The main question concerns the level of the biological hierarchy at which natural selection occurs. Does selection act on organisms, genes, groups, colonies, demes, species, or some combination of these? According to traditional Darwinian theory the answer is the organism -- it is the differential survival and reproduction of individual organisms that drives the evolutionary process. But there (...) are alternative views too, including pluralist views which say that the choice between different levels of selection is often a matter of perspective, not empirical fact. (shrink)

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

Evolutionary, ecological and ethical studies are, at the same time, specific scientific disciplines and, from an historical point of view, structurally linked domains of research. In a context of environmental crisis, the need is increasingly emerging for a connecting epistemological framework able to express a common or convergent tendency of thought and practice aimed at building, among other things, an environmental policy management respectful of the planet’s biodiversity and its evolutionary potential.

Gers (Biol Philos, 2011) provides a positive and constructive view of the project to generalise Darwinian principles in Geoffrey Hodgson and Thorbjørn Knudsen’s Darwin’s Conjecture. We note considerable overlap with his work and ours, and also with important recent work of Godfrey-Smith ( 2009 ), which Gers cites extensively. But we also note that there are differences in research objectives between Gers and Godfrey-Smith, on the one hand, and ourselves, on the other. Gers and Godfrey-Smith focus on the elucidation of (...) the most general principles possible. Our aim is to derive principles that are sufficiently abstract to span the natural and human social worlds, and then add additional principles to help understand the Darwinian evolution of human society. Furthermore, Gers and Godfrey-Smith critique a replicator concept that is different from ours. Once these points are made apparent, the criticisms are essentially disabled, and we end up in a position with different but complementary and overlapping research projects. (shrink)

Human knowledge is a phenomenon whose roots extend from the cultural, through the neural and the biological and finally all the way down into the Precambrian “primordial soup.” The present paper reports an attempt at understanding this Greater System of Knowledge (GSK) as a hierarchical nested set of selection processes acting concurrently on several different scales of time and space. To this end, a general selection theory extending mainly from the work of Hull and Campbell is introduced. The perhaps most (...) drastic change from previous similar theories is that replication is revealed as a composite function consisting of what is referred to as memory and synthesis. This move is argued to drastically improve the fit between theory and human-related knowledge systems. The introduced theory is then used to interpret the subsystems of the GSK and their interrelations. This is done to the end of demonstrating some of the new perspectives offered by this view. (shrink)

Despite growing interest in evolutionary economics since the 1980s, a unified theoretical approach has so far been lacking. Methodological and ontological discussions within evolutionary economics have attempted to understand and help rectify this failure, but have revealed in turn further differences of perspective. One aim of this article is to show how different approaches relate to different levels of abstraction. A second purpose is to show that generalized Darwinism is some way from the most abstract level, and illustrates how it (...) may be used to move towards more specific theoretical applications. Nevertheless, there is a long way to go before these become more evident. (shrink)

Throughout his career David Hull has sought to bring the philosophy of science into closer contact with science and especially with biological science (Hull 1969, 1997b). This effort has taken many forms. Sometimes it has meant ‘either explaining basic biology to philosophers or explaining basic philosophy to biologists’ (Hull 1996, p. 77). The first of these tasks, simple as it sounds, has been responsible for revolutionary changes. It is well known that traditional philosophy of science, modeled as it was on (...) theoretical physics, proved inadequate when philosophers turned their attention to biological science. Biological examples have driven major revisions of accounts of reduction (Hull 1974; Schaffner 1993, Ch. 9), laws of nature (Beatty et al. 1997), theories (Lloyd 1988) and natural kinds (Wilson 1999, Part III). Nor is explaining basic philosophy to biologists a task to be looked down upon. It is useful, not because philosophy has all the answers, but because scientists must think about how to do science, that is doing philosophy of science and scientists frequently reinvent philosophical views with known flaws. Early in his career Hull found biological systematists in the grip of a crude operationalism about scientific concepts and said so in the pages of Systematic Zoology (Hull 1968). For the next thirty years, as biologists debated the nature of species and the correct principles of classification, Hull added a philosophical note at the same congresses and in the same journals (Hull 1970, 1976, 1980, 1997a, 1999). (shrink)

The established definition of replication in terms of the conditions of causality, similarity and information transfer is very broad. We draw inspiration from the literature on self-reproducing automata to strengthen the notion of information transfer in replication processes. To the triple conditions of causality, similarity and information transfer, we add a fourth condition that defines a “generative replicator” as a conditional generative mechanism, which can turn input signals from an environment into developmental instructions. Generative replication must have the potential to (...) enhance complexity, which in turn requires that developmental instructions are part of the information that is transmitted in replication. Demonstrating the usefulness of the generative replicator concept in the social domain, we identify social generative replicators that satisfy all of the four proposed conditions. (shrink)

Huang and Bargh’s definition of goals is ambiguous between ‘specific goals’ – the end-state of a token action I am about to perform – and ‘unspecific goals’ – the end-state of an action-type (without specifying how this would be achieved). The analogy with selfish genes pushes the authors towards the former interpretation, but the latter would provide a more robust theoretical framework.

Darwin’s Conjecture is a bold attempt to bring evolutionary explanation to the social sciences, particularly economics. The book outlines the history of Darwinian explanation in social science then puts forward a generalized replicator account of social evolution by natural selection. The authors identify habits and routines as examples of the generative replicators necessary in order that social evolution is Darwinian. This reviewer notes that the replicator approach limits the generality of this account and suggests that habits and routines might better (...) be seen as members of a Darwinian population in the tradition of Godfrey-Smith’s (2009) generalized account of natural selection instead. Furthermore, the existence of levels of social evolution will constrain what evolution can occur low in the hierarchy. (shrink)

Selection theory requires multiple, distinct, simultaneously-actualized states. In cognition, each thought or cognitive state changes the 'selection pressure' against which the next is evaluated; they are not simultaneously selected amongst. Creative thought is more a matter of honing in a vague idea through redescribing successive iterations of it from different real or imagined perspectives; in other words, actualizing potential through exposure to different contexts. It has been proven that the mathematical description of contextual change of state introduces a non-Kolmogorovian probability (...) distribution, and a classical formalism such as selection theory cannot be used. This paper argues that creative thought evolves not through a Darwinian process, but a process of context-driven actualization of potential. (shrink)

Evolutionary psychology and the selectionist theories of neural development are usually regarded as two unrelated theories addressing two logically distinct questions. The focus of evolutionary psychology is the phylogeny of the human mind, whereas the selectionist theories of neural development analyse the ontogeny of the mind. This paper will endeavour to combine these two approaches in the explanation of the human mind. Doing so might help in overcoming some of the criticisms of both theories. The first part of the paper (...) mentions three standard objections to evolutionary psychology and then outlines three philosophical problems evolutionary psychology has to offer a solution to. The second part will try to show that an approach combining evolutionary psychology and the selectionist theory of neural development might overcome some of these objections. (shrink)

In evolutionary biology and ecology, ontological and epistemological perspectives based on the replicator and the interactor have become the background that makes it possible to transcend traditional biological levels of organization and to achieve a unified view of evolution in which replication and interaction are fundamental operating processes. Using the transactional perspective proposed originally by John Dewey and Arthur Fisher Bentley, a new ontological and methodological category is proposed here: the transactor. The transactional perspective, based on the concept of the (...) transactor, bridges the dichotomy between organisms and environment that characterizes the interactional perspective on evolution and provides epistemological support for the emergentist, systemic view of evolutionary and developmental processes. (shrink)

Climate policy decisions are decisions under uncertainty and are, therefore, based on a range of future climate scenarios, describing possible consequences of alternative policies. Accordingly, the methodology for setting up such a scenario range becomes pivotal in climate policy advice. The preferred methodology of the Intergovernmental Panel on Climate Change will be characterised as ,,modal verificationism"; it suffers from severe shortcomings which disqualify it for scientific policy advice. Modal falsificationism, as a more sound alternative, would radically alter the way the (...) climate scenario range is set up. Climate science's inability to find robust upper bounds for future temperature rise in line with modal falsificationism does not disprove that methodology, rather, this very fact prescribes even more drastic efforts to curb CO2 emissions than currently proposed. (shrink)

The significant theoretical objections that have been raised against memetics have not received adequate defense, even though there is ongoing empirical research in this field. In this paper I identify the key objections to memetics as a viable explanatory tool in studies of cultural evolution. I attempt to defuse these objections by arguing that they fail to show the absence of replication, high-fidelity copying, or lineages in the cultural domain. I further respond to meme critics by arguing that, despite competing (...) explanations of cultural evolution, memetics has unique explanatory power. This is largely founded upon the increasing likelihood of formulating a workable fitness measure for memes, a memetic index. I conclude that memes must be integrated with psychological bias and population-dynamic approaches to cultural evolution. (shrink)

The dissertation criticizes two analogical applications of Darwinism to the spheres of mind and culture: the Darwinian approach to creativity and memetics. These theories rely on three basic analogies: the ontological analogy states that the basic ontological units of culture are so-called memes, which are replicators like genes; the origination analogy states that novelty in human creativity emerges in a "blind" Darwinian manner; and the explanatory units of selection analogy states that memes are "egoistic" and that they can spread independently (...) of our mind. The detailed philosophical analysis offered in the dissertation shows that these three analogies rely on either wrong or trivial statements: they provide either wrong or no new descriptions or explanations of the phenomena at hand. In Chapter One, I introduce the diverse ways, in which contemporary Darwinism is used today outside of evolutionary biology. Chapter Two explains what it means to claim that something evolves in a Darwinian manner. Chapter Three to Five address each of the three analogies separately. Darwinism applied to culture and mind, at least in the way discussed in this study, is not a dangerous idea. It leads either to wrong claims or to a re-telling in Darwinian terms of what we have already known. (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 goal of forming a science of intentional behavior requires a more richly detailed account of symbolic systems than is assumed by the authors. Cultural systems are not simply the equivalent in the ideational domain of culture of the purported Baldwin Effect in the genetic domain. © 2014 Cambridge University Press.

The aim of this paper is to outline a typologyof selection processes, and show that differentsub-categories have different explanatorypower. The basis of this typology of selectionprocesses is argued to be the difference ofreplication processes involved in them. Inorder to show this, I argue that: 1.Replication is necessary for selection and 2.Different types of replication lead todifferent types of selection. Finally, it isargued that this typology is philosophicallysignificant, since it contrasts cases ofselection (on the basis of the replicationprocesses involved in them) (...) whereby selectioncauses adaptation – and, therefore, can beused in explanations of the (real or apparent)teleology of Nature – and cases in whichselection lacks such explanatory power. (shrink)