The extended evolutionary synthesis intends to offer a new framework for understanding evolution based mainly on empirical and theoretical findings of current studies, including heredity and evolutionary developmental biology. In this essay, we present and develop the following objections about the terminology associated with the EES literature: despite using the term "extension," EES protagonists claim new evolutionary processes, reformulate conceptual networks, and modify central assumptions of the evolutionary synthesis. Therefore, the difference between ES (...) and EES should not be described in terms of a set-subset relationship ; despite using the term "synthesis," the EES leads, at least in the short term, to a pluralism of approaches in evolutionary biology. Thus, we argue that the EES should not be described as a synthesis, but as a broad and plural interpretative framework encompassing different approaches. These objections are not directed to the proposed changes, but to the interpretation of these changes as a mere expansion of the previous evolutionary framework, as well as to the interpretation of the EES as a synthesis. Based on these objections, we seek to present some explanations for the use of the term "extended synthesis" among its protagonists. (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. (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)

To approach the issue of the recent proposal of an Extended Evolutionary Synthesis (EES) put forth by Massimo Pigliucci and Gerd Müller, I suggest to consider the EES as a metascientific view: a description of what’s new in how evolutionary biology is carried out, not only a description of recently learned aspects of evolution. Knowing ‘what is it to do research’ in evolutionary biology, today versus yesterday, can aid training, research and career choices, establishment of (...) relationships and collaborations, decision of funding and research policies, in order to make the field advance for the better. After reviewing the concepts associated to the EES proposal (categorized for convenience as mechanisms, measures, fields, perspectives and applications), I show their transience, and sketch out ongoing disagreements about the EES. Then I examine the deep difficulties, i.e., the enormity and complexity of the covered field, affecting the achievement of trusted metascientific views; the insufficiency of conceptual analysis to capture the substance of scientific research; the entanglement between empirical and metascientific concepts, between multiple chronologies, and between descriptive and normative intentions; and the ineliminable stakeholding of any reviewer involved in the reviewed field. I propose that disciplines such as scientometrics, ethnography, sociology, economics and history, combined with conceptual analysis, inspire a more rigorous approach to the evolutionary biology scientific community, more grounded and shared, confirming or transforming claims for ‘synthesis’ while preserving their maintenance goals. (shrink)

Kevin Laland and colleagues have put forward a number of arguments motivating an extended evolutionary synthesis. Here I examine Laland et al.'s central concept of reciprocal causation. Reciprocal causation features in many arguments supporting an expanded evolutionary framework, yet few of these arguments are clearly delineated. Here I clarify the concept and make explicit three arguments in which it features. I identify where skeptics can—and are—pushing back against these arguments, and highlight what I see as the (...) empirical, explanatory, and methodological issues at stake. (shrink)

Biologists and philosophers of science have recently called for an extension of evolutionary theory. This so-called ‘extended evolutionary synthesis’ seeks to integrate developmental processes, extra-genetic forms of inheritance, and niche construction into evolutionary theory in a central way. While there is often agreement in evolutionary biology over the existence of these phenomena, their explanatory relevance is questioned. Advocates of EES posit that their perspective offers better explanations than those provided by ‘standard evolutionary theory’. (...) Still, why this would be the case is unclear. Usually, such claims assume that EES’s superior explanatory status arises from the pluralist structure of EES, its different problem agenda, and a growing body of evidence for the evolutionary relevance of developmental phenomena. However, what is usually neglected in this debate is a discussion of what the explanatory standards of EES actually are, and how they differ from prevailing standards in SET. In other words, what is considered to be a good explanation in EES versus SET? To answer this question, we present a theoretical framework that evaluates the explanatory power of different evolutionary explanations of the same phenomena. This account is able to identify criteria for why and when evolutionary explanations of EES are better than those of SET. Such evaluations will enable evolutionary biology to find potential grounds for theoretical integration. (shrink)

The Extended Evolutionary Synthesis (EES) debate is gaining ground in contemporary evolutionary biology. In parallel, a number of philosophical standpoints have emerged in an attempt to clarify what exactly is represented by the EES. For Massimo Pigliucci, we are in the wake of the newest instantiation of a persisting Kuhnian paradigm; in contrast, Telmo Pievani has contended that the transition to an EES could be best represented as a progressive reformation of a prior Lakatosian scientific research (...) program, with the extension of its Neo-Darwinian core and the addition of a brand-new protective belt of assumptions and auxiliary hypotheses. Here, we argue that those philosophical vantage points are not the only ways to interpret what current proposals to ‘extend’ the Modern Synthesis-derived ‘standard evolutionary theory’ (SET) entail in terms of theoretical change in evolutionary biology. We specifically propose the image of the emergent EES as a vast network of models and interweaved representations that, instantiated in diverse practices, are connected and related in multiple ways. Under that assumption, the EES could be articulated around a paraconsistent network of evolutionary theories (including some elements of the SET), as well as models, practices and representation systems of contemporary evolutionary biology, with edges and nodes that change their position and centrality as a consequence of the co-construction and stabilization of facts and historical discussions revolving around the epistemic goals of this area of the life sciences. We then critically examine the purported structure of the EES—published by Laland and collaborators in 2015—in light of our own network-based proposal. Finally, we consider which epistemic units of Evo-Devo are present or still missing from the EES, in preparation for further analyses of the topic of explanatory integration in this conceptual framework. (shrink)

Current knowledge of the genetic, epigenetic, behavioural and symbolic systems of inheritance requires a revision and extension of the mid-twentieth-century, gene-based, 'Modern Synthesis' version of Darwinian evolutionary theory. We present the case for this by first outlining the history that led to the neo-Darwinian view of evolution. In the second section we describe and compare different types of inheritance, and in the third discuss the implications of a broad view of heredity for various aspects of evolutionary theory. (...) We end with an examination of the philosophical and conceptual ramifications of evolutionary thinking that incorporates multiple inheritance systems. (shrink)

Contemporary evolutionary biology comprises a plural landscape of multiple co-existent conceptual frameworks and strenuous voices that disagree on the nature and scope of evolutionary theory. Since the mid-eighties, some of these conceptual frameworks have denounced the ontologies of the Modern Synthesis and of the updated Standard Theory of Evolution as unfinished or even flawed. In this paper, we analyze and compare two of those conceptual frameworks, namely Niles Eldredge’s Hierarchy Theory of Evolution (with its extended ontology (...) of evolutionary entities) and the Extended Evolutionary Synthesis (with its proposal of an extended ontology of evolutionary processes), in an attempt to map some epistemic bridges (e.g. compatible views of causation; niche construction) and some conceptual rifts (e.g. extra-genetic inheritance; different perspectives on macroevolution; contrasting standpoints held in the “externalism–internalism” debate) that exist between them. This paper seeks to encourage theoretical, philosophical and historiographical discussions about pluralism or the possible unification of contemporary evolutionary biology. (shrink)

In this paper, we analyze the debate between the Modern Synthesis and the Extended Evolutionary Synthesis in light of the concept of incommensurability developed by Thomas Kuhn. In order to do so, first we briefly present both the Modern Synthesis and the Extended Evolutionary Synthesis. Then, we clarify the meaning and interpretations of incommensurability throughout Kuhn’s works, concluding that the version of this concept deployed in The Structure of Scientific Revolutions is the (...) best suited to the analysis of scientific disputes. After discussing incommensurability in Kuhn’s works, we address the question of whether the Modern Synthesis and the Extended Evolutionary Synthesis can be considered semantically, methodologically, and ontologically incommensurable, concluding that they can. Finally, we discuss three problems that arise from such a conclusion: firstly, what are the consequences of incommensurability; secondly, which mode of scientific change better explains this current dispute in evolutionary biology; and thirdly, whether rational theory comparison is possible given incommensurability. We suggest that the main consequence of incommensurability is profound disagreement, that the kind of scientific change that better explains the current dispute between the MS and the EES may be scientific specialization, and that incommensurability does not preclude rational theory comparison. (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)

Synthesising arguments motivate changes to the conceptual tools, theoretical structure, and evaluatory framework employed in a given scientific domain. Recently, a broad coalition of researchers has put forward a synthesising argument in favour of an Extended Evolutionary Synthesis (‘EES’). Often this synthesising argument is evaluated using a virtue-based approach, which construes the EES as a wholesale alternative to prevailing practice. Here I argue this virtue-based approach is not fit for purpose. Taking the central concept of niche construction (...) as a case study, I show that an agenda-based approach better captures the pragmatic and epistemological goals of the EES synthesising argument and diagnoses areas of empirical disagreement with prevailing practice. (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 Modern Synthesis (MS) gene-centered population model of evolution is currently being challenged by the Extended Evolutionary Synthesis (ESS) organism-centered developmental model of evolution. The predictions of the EES are here examined with respect to the arguments of Edward Drinker Cope (1840–1897) for an organism-centered evolutionary process in which organisms both shape and are shaped by their environments such that the activities of the organisms themselves play a role in their own evolution.

According to sources both in print and at a recent meeting, evolutionary theory is currently undergoing change which some would characterize as a New Synthesis, and others as an Extended Synthesis. This article argues that the important changes involve recognizing that there are three means by which evolutionary change can be initiated and three corresponding modes of evolutionary drift. It compares the three and goes on to discuss the scale of innovation and extended (...) or inclusive and Lamarckian inheritance. It concludes from these that “new trends in evolutionary biology” are in part a new, and in part an extended evolutionary synthesis. (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)

In this article I will analyze whether the so-called Extended Evolutionary Synthesis represents a synthesis and an extension with respect to its predecessor, Modern Synthesis. It will be argued that the MS proposes an externalist approach to evolution while the EES considers it necessary to overcome the internalism/externalism dichotomy by proposing more integrative approaches. It will be concluded that the EES cannot be considered an extension of MS and that the appeal to that extension is (...) related to sociological aspects and the epistemic value of theoretical unification that was always present in biological evolutionary thinking. (shrink)

In this article I will analyze whether the so-called Extended Evolutionary Synthesis represents a synthesis and an extension with respect to its predecessor, Modern Synthesis. It will be argued that the MS proposes an externalist approach to evolution while the EES considers it necessary to overcome the internalism/externalism dichotomy by proposing more integrative approaches. It will be concluded that the EES cannot be considered an extension of MS and that the appeal to that extension is (...) related to sociological aspects and the epistemic value of theoretical unification that was always present in biological evolutionary thinking. (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)

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)

Evolutionary theory is undergoing an intense period of discussion and reevaluation. This, contrary to the misleading claims of creationists and other pseudoscientists, is no harbinger of a crisis but rather the opposite: the field is expanding dramatically in terms of both empirical discoveries and new ideas. In this essay I briefly trace the conceptual history of evolutionary theory from Darwinism to neo-Darwinism, and from the Modern Synthesis to what I refer to as the Extended Synthesis, (...) a more inclusive conceptual framework containing among others evo–devo, an expanded theory of heredity, elements of complexity theory, ideas about evolvability, and a reevaluation of levels of selection. I argue that evolutionary biology has never seen a paradigm shift, in the philosophical sense of the term, except when it moved from natural theology to empirical science in the middle of the 19th century. The Extended Synthesis, accordingly, is an expansion of the Modern Synthesis of the 1930s and 1940s, and one that—like its predecessor—will probably take decades to complete. (shrink)

Welch :263–279, 2017) has recently proposed two possible explanations for why the field of evolutionary biology is plagued by a steady stream of claims that it needs urgent reform. It is either seriously deficient and incapable of incorporating ideas that are new, relevant and plausible or it is not seriously deficient at all but is prone to attracting discontent and to the championing of ideas that are not very relevant, plausible and/or not really new. He argues for the second (...) explanation. This paper presents a twofold critique of his analysis: firstly, the main calls for reform do not concern the field of evolutionary biology in general but rather, or more specifically, the modern evolutionary synthesis. Secondly, and most importantly, these calls are not only inspired by the factors, enumerated by Welch, but are also, and even primarily, motivated by four problematic characteristics of the modern synthesis. This point is illustrated through a short analysis of the latest reform challenge to the modern synthesis, the so-called extended evolutionary synthesis. We conclude with the suggestion that the modern synthesis should be amended, rather than replaced. (shrink)

The program of Evolutionary Ethics (EE) is based on the assumption that our moral features constitute adaptations and as such are to be explained in terms of the evolutionary process of natural selection. However, the fundamental assumption of EE was seriously put into question: the level of analysis relevant for moral features is essentially ontogeny and culture, while the explanation using natural selection applies to the level of phylogeny and genes (Sober, 1995; Ayala, 1995; Okasha, 2009). To the (...) discussion on the validity of the program of EE we propose to bring the recent program of Extended Synthesis (ES, Pigliucci & Muller, 2010), because it attempts to account for the role of the ontogeny in evolution. We conclude, nevertheless, that ES fails to properly account for the importance of ontogeny in evolutionary processes because by extending the notion of inheritance it (con-)fuses the notions of unit of inheritance and of unit of selection (against the well-known distinction made by Hull, 1980). (shrink)

In this paper, I will briefly summarize the history and current accounts of Evolutionary Medicine (EM). I will show that EM, in its current forms, is using an evolutionary understanding that carries the explanatory framework, as well as explanatory limits, of the Modern Synthesis (MS). I will then point out some essential elements that need to be seen as limiting factors within EM and analyze the limitations that are brought about by the MS understanding of it. On (...) this basis, I will argue that if the latest developments in evolutionary theory are considered – in particular, those pertaining to the inheritance mechanisms highlighted by the Extended Evolutionary Synthesis (EES), and the newly introduced evolutionary entities – EM will have a much broader explanatory scope and increased explanatory power in addition to greater relevance, which will enable its application in medical explanations. (shrink)

Two welcome extensions of evolutionary thinking have come to prominence over the last thirty years: the so-called ’extended evolutionary synthesis’ (EES) and debate about biological kinds and individuals. These two agendas have, however, remained orthogonal to one another. The EES has mostly restricted itself to widening the explanations of adaptation offered by the preceding ’modern evolutionary synthesis’ by including additional mechanisms of inheritance and variation; while discussion of biological kinds has turned toward philosophical questions (...) of essential vs. contingent properties of life forms and realist vs. epistemological approaches to categorization and classification. Here we attempt to broaden the explanatory scope of evolutionary theory by linking these two agendas. We expand on the mechanistic orientation of the EES, using new understandings of networked systems of components in order to engage the distinct intellectual challenge of the origination of “historical kinds”. With this phrase we designate a subset of natural kinds that acquires, through evolutionary processes, a quasi-independent lineage-history. Such kinds emerge in both biology and culture, and we enlarge the limited number of historical kinds that have thus far been recognized in evolutionary biology in a series of paradigmatic exemplars, from genes and cell types to rituals and music. For each exemplar we discern specific mechanisms by which it arose and persists; comparing these, we suggest a general unity in the ways in which diverse historical kinds originate. (shrink)

This chapter describes the theoretical implications of Extended Synthesis and addresses the methodological options available for determining aspects of theoretical structure. It uses a “bottom-up” approach focused on evolutionary theory in particular, as opposed to a “top-down” strategy that attempts to characterize the structure of all scientific theories. The chapter shows that there are multiple stable components contained within a broad representation of evolutionary theory. It suggests that the philosophical analysis offered in the chapter regarding the (...) structure of evolutionary theory assists attempts to recover coherence through the vehicle of an Extended Synthesis. (shrink)

In the six decades since the publication of Julian Huxley's Evolution: The Modern Synthesis, spectacular empirical advances in the biological sciences have been accompanied by equally significant developments within the core theoretical framework of the discipline. As a result, evolutionary theory today includes concepts and even entire new fields that were not part of the foundational structure of the Modern Synthesis. In this volume, sixteen leading evolutionary biologists and philosophers of science survey the conceptual changes that (...) have emerged since Huxley's landmark publication, not only in such traditional domains of evolutionary biology as quantitative genetics and paleontology but also in such new fields of research as genomics and EvoDevo. Most of the contributors to Evolution—The Extended Synthesis accept many of the tenets of the classical framework but want to relax some of its assumptions and introduce significant conceptual augmentations of the basic Modern Synthesis structure—just as the architects of the Modern Synthesis themselves expanded and modulated previous versions of Darwinism. This continuing revision of a theoretical edifice the foundations of which were laid in the middle of the nineteenth century—the reexamination of old ideas, proposals of new ones, and the synthesis of the most suitable—shows us how science works, and how scientists have painstakingly built a solid set of explanations for what Darwin called the "grandeur" of life. (shrink)

A response to Lindsay Craig's essay, The So-Called Extended Synthesis and Population Genetics.

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)

Language evolution, intended as an open problem in the evolutionary research programme, will be here analyzed from the theoretical perspective advanced by the supporters of the Extended Evolutionary Synthesis. Four factors and two associated concepts will be matched with a selection of critical examples concerning genus Homo evolution, relevant for the evolution of language, such as the evolution of hominin life-history traits, the enlargement of the social group, increased cooperation among individuals, behavioral change and innovations, heterochronic (...) modifications leading to increased synaptic plasticity. A particular form of niche construction will be considered in a multilevel framework. It will be argued that the four points mentioned above prove to be fundamental explanatory tools to understand how language might have emerged as a result of a gene-culture coevolutionary dynamics. (shrink)

Change is the fundamental idea of evolution. Explaining the extraordinary biological change we see written in the history of genomes and fossil beds is the primary occupation of the evolutionary biologist. Yet it is a surprising fact that for the majority of evolutionary research, we have rarely studied how evolution typically unfolds in nature, in changing ecological environments, over space and time. While ecology played a major role in the eventual acceptance of the population genetic viewpoint of evolution (...) in the synthetic era (circa 1918-1956), it held a lesser role in the development of evolutionary theory until the 1980s, when we began to systematically study the evolutionary dynamics of natural populations in space and time. As a result, early evolutionary theory was initially constructed in an abstract vacuum that was unrepresentative of evolution in nature. The subtle synthesis between ecology with evolutionary biology (eco-evo synthesis) over the past 40 years has progressed our knowledge of natural selection dynamics as they are found in nature, thus revealing how natural selection varies in strength, direction, form, and, more surprisingly, level of biological organization. Natural selection can no longer be reduced to lower levels of biological organization (i.e., individuals, selfish genes) over shorter timescales but should be expanded to include adaptation at higher levels and over longer timescales. Long-term and/or emergent evolutionary phenomena, such as multilevel selection or evolvability, have thus become tenable concepts within an evolutionary biology that embraces ecology and spatiotemporal change. Evolutionary biology is currently suspended at an intermediate stage of scientific progress that calls for the organization of all the recent knowledge revealed by the eco-evo synthesis into a coherent and unified theoretical framework. This is where philosophers of biology can be of particular use, acting as a bridge between the subdisciplines of biology and inventing new theoretical strategies to organize and accommodate the recent knowledge. Philosophers have recommended transitioning away from outdated philosophies that were originally derived from physics within the philosophical zeitgeist of logical positivism (i.e., monism, reductionism, and monocausation) and toward a distinct philosophy of biology that can capture the natural complexity of multifaceted biological systems within diverse ecosystems—one that embraces the emerging philosophies of pluralism, emergence, and multicausality. Therefore, I see recent advances in ecology, evolutionary biology, and the philosophy of biology as laying the groundwork for another major biological synthesis, what I refer to as the Second Synthesis because, in many respects, it is analogous to the aims and outcomes of the first major biological synthesis (but is notably distinct from the inorganic and contrived progressive movement known as the extended evolutionary synthesis). With the general development of a distinctive philosophy of science, biology has rightfully emerged as an autonomous science. Thus, while the first synthesis legitimized biology, the Second Synthesis autonomized biology and afforded biology its own philosophy, allowing biology to finally realize its full scientific potential. (shrink)

In recent years, several prominent biologists have pointed to the relatively new field of evolutionary developmental biology (EvoDevo) as evidence of an Extended Synthesis in evolutionary biology. More particularly, these biologists claim that theoretical and empirical EvoDevo research is extending the Modern Synthesis framework of evolutionary theory through investigation of evolutionarily important concepts that are not part of the framework developed during the 20th century. To describe the current changes in evolutionary biology as (...) an Extended Synthesis, however, is incorrect. Through review of Extended Synthesis arguments and analysis of the same biological concepts used to support these arguments, I argue that the foundation of the Modern Synthesis framework, theoretical population genetics, faces significant, perhaps insurmountable challenges from the concepts highlighted by EvoDevo research. As the foundation of the Modern Synthesis framework will require considerable remodeling—if possible—in light of the concepts emphasized by EvoDevo, it is incorrect to describe the ongoing changes in evolutionary biology as an Extended Synthesis. (shrink)

In recent years, several prominent biologists have pointed to the relatively new field of evolutionary developmental biology as evidence of an Extended Synthesis in evolutionary biology. More particularly, these biologists claim that theoretical and empirical EvoDevo research is extending the Modern Synthesis framework of evolutionary theory through investigation of evolutionarily important concepts that are not part of the framework developed during the 20th century. To describe the current changes in evolutionary biology as an (...) Extended Synthesis, however, is incorrect. Through review of Extended Synthesis arguments and analysis of the same biological concepts used to support these arguments, I argue that the foundation of the Modern Synthesis framework, theoretical population genetics, faces significant, perhaps insurmountable challenges from the concepts highlighted by EvoDevo research. As the foundation of the Modern Synthesis framework will require considerable remodeling—if possible—in light of the concepts emphasized by EvoDevo, it is incorrect to describe the ongoing changes in evolutionary biology as an Extended Synthesis. (shrink)

This paper argues that the Extended Synthesis, ecological information, and biosemiotics are complementary approaches whose engagement will help us explain the organism-environment interaction at the cognitive level. The Extended Synthesis, through niche construction theory, can explain the organism-environment interaction at an evolutionary level because niche construction is a process guided by information. We believe that the best account that defines information at this level is the one offered by biosemiotics and, within all kinds of biosemiotic (...) information available, we believe that ecological information is the best candidate for making sense of the organism-environment relation at the cognitive level. This entanglement of biosemiotics, ecological information and the Extended Synthesis is promising for understanding the multidimensional character of the organism-environment reciprocity as well as the relation between evolution, cognition, and meaning. (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)

Current knowledge of the genetic, epigenetic, behavioural and symbolic systems of inheritance requires a revision and extension of the mid-twentieth-century, gene-based, 'Modern Synthesis' version of Darwinian evolutionary theory. We present the case for this by first outlining the history that led to the neo-Darwinian view of evolution. In the second section we describe and compare different types of inheritance, and in the third discuss the implications of a broad view of heredity for various aspects of evolutionary theory. (...) We end with an examination of the philosophical and conceptual ramifications of evolutionary thinking that incorporates multiple inheritance systems. (shrink)

Discussions about the extended mind have ‘extended’ in various directions in the last decades. While applied to other aspects of human cognition and even consciousness, the extended-mind hypothesis has also been criticized, as it questions fundamental ideas such as the image of a dual world, divided between an external and an internal domain by the border of ‘skin and skull’, the idea of a localized and constant decision center, and the role of internal representations. We suggest that (...) the main virtue of the hypothesis is not as a theory per se, but as a vaccine against persistent metaphysical prejudices about the mind’s structure, functions and borders. Being an hypothesis about the most efficient ways to combine resources and problems, and not a theory about the mind’s a-priori constitution, the extended mind view moves the focus from ontology to pragmatics and helps purify philosophy of mind from metaphysical remainders. (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)

The developmental properties of organisms play important roles in the generation of variation necessary for evolutionary change. But how can individual development steer the course of evolution? To answer this question, we introduce developmental channeling as a disposition of individual organisms that shapes their possible developmental trajectories and evolutionary dappling as an evolutionary outcome in which the space of possible organismic forms is dappled—it is only partially filled. We then trace out the implications of the channeling-dappling framework (...) for contemporary debates in the philosophy of evolution, including evolvability, reciprocal causation, and the extended evolutionary synthesis. (shrink)

The Modern Synthesis has been receiving bad press for some time now. Back in 1983, in an article entitled “The Hardening of the Modern Synthesis” Stephen Jay Gould criticized the way the Modern Synthesis had developed since its inception in the 1930s and early 1940s (Gould 1983). Back then, those who would later become known as ‘architects’ of the synthesis were united in their call for explaining evolution at all levels in terms of causation at one (...) level: genetics. What drove changes in gene frequency remained an open question. It could be mainly selection, or drift, or some (other) form of constraint. But in the two decades that followed, the synthesis underwent a major change. By the late 1940s the synthesis had ‘hardened’ around adaptationism, according to Gould. Influential contributors like Dobzhansky, Simpson and Wright had increasingly expressed adaptationist views in the later editions of their landmark books. Not because evidence had piled up, showing that selection was in fact pervasive. Instead, Gould argued, adaptationist tendencies had been preserved by some kind of cultural inertia, and were now being revived. “Certain ‘national styles’ persisted from the eighteenth century, through Darwin’s era, and into our own time. Views on adaptation provide a good example” (Gould 1983). Gould did not just argue that some form of adaptationism had resurfaced. He became well-known for his efforts to intervene on this status quo by attempting to make evolutionary biology more ‘pluralistic’. In collaborative work with Richard Lewontin (Gould and Lewontin 1979), Elisabeth Vrba (Gould and Vrba 1982; Vrba and Gould 1986) and Niles Eldredge (Eldredge and Gould 1972; Gould and Eldredge 1977) he criticized the synthesis for its adaptationism and its lack of appreciation for hierarchical perspectives. Gould exerted his influence in a different way as well. Together with Eldredge, he had facsimiles reprinted of the first editions of two books that had shaped synthesis, but with their own critical introductions (Eldredge 1982; Gould 1982). Dobzhansky’s Genetics and the Origin of Species and Mayr’s Systematics and the Origin of Species appeared as reprints in the ‘Columbia Classics in Evolution’ series, sending an unambiguous message to the readers: these are foundational works, but they have been superseded. In the summer of 2008, some 25 years after Gould made his point about the hardening of the Modern Synthesis, a group of sixteen biologists and philosophers gathered at the Konrad Lorenz Institute for Evolution and Cognition Research (KLI) near Vienna, Austria, to discuss cutting-edge research that reaches beyond the synthesis framework. Before it even started, this workshop on the ‘Extended Synthesis’ had already attracted a fair share of attention in the blogosphere and had resulted in a news feature in Science (Pennisi 2008). After the meeting, Nature weighed in on the matter (Whitfield 2008). The results of over 3 days of presentations and extensive discussion have now been published as Evolution—The Extended Synthesis. 1 The publication of this collection of sixteen essays is accompanied by the republication of Julian Huxley’s Evolution: The Modern Synthesis; the book that introduced the term ‘Modern Synthesis’. Both books are introduced by the organizers of the KLI workshop, Massimo Pigliucci and Gerd Müller. Like Gould and Eldredge before them, Pigliucci and Müller did not reissue one of the canons of the Modern Synthesis without giving the readers some ‘guidance’. Starting with the cover, the editors proclaim boldly that this is ‘the definitive edition’ of Huxley’s book. In a new foreword, they sketch the context in which the book was written and assess some of its features. They voice some mild criticism of alleged ‘adaptationism’. But their tone is different from that of Gould and Eldredge. Pigliucci and Müller praise Huxley for his pluralistic outlook, which has again become essential in the forging of an Extended Synthesis. That makes Huxley’s book more than just an interesting but obsolete classic. Instead, it can teach valuable lessons about how to ‘soften up’ a synthesis that has become hardened over time. (shrink)

Social evolution theory conventionally takes an externalist explanatory stance, treating observed cooperation as explanandum and the positive assortment of cooperative behaviour as explanans. We ask how the circumstances bringing about this positive assortment arose in the first place. Rather than merely push the explanatory problem back a step, we move from an externalist to an interactionist explanatory stance, in the spirit of Lewontin and the Niche Construction theorists. We develop a theory of ‘social niche construction’ in which we consider biological (...) entities to be both the subject and object of their own social evolution. Some important cases of the evolution of cooperation have the side-effect of causing changes in the hierarchical level at which the evolutionary process acts. This is because the traits that act to align the fitness interests of particles in a collective can also act to diminish the extent to which those particles are bearers of heritable fitness variance, while augmenting the extent to which collectives of such particles are bearers of heritable fitness variance. In this way, we can explain upward transitions in the hierarchical level at which the Darwinian machine operates in terms of particle-level selection, even though the outcome of the process is a collective-level selection regime. Our theory avoids the logical and metaphysical paradoxes faced by other attempts to explain evolutionary transitions. (shrink)

Traditionally defined as the science of the living, or as the field that beyond anatomical structure and bodily form studies functional organization and behaviour, physiology has long been excluded from evolutionary research. The main reason for this exclusion is that physiology has a presential and futuristic outlook on life, while evolutionary theory is traditionally defined as the study of natural history. In this paper, I re-evaluate these classic science divisions and situate physiology within the history of the (...) class='Hi'>evolutionary sciences, as well as within debates on the Extended Evolutionary Synthesis and the need for a Third Way of Evolution. I then briefly point out how evolutionary physiology in particular contributes to research on function, causation, teleonomy, agency and cognition. (shrink)