Strong reciprocity has recently been subject to heated debate. In this debate, the “West camp” :231–262, 2011), which is critical of the case for SR, and the “Laland camp” :1512–1516, 2011, Biol Philos 28:719–745, 2013), which is sympathetic to the case of SR, seem to take diametrically opposed positions. The West camp criticizes advocates of SR for conflating proximate and ultimate causation. SR is said to be a proximate mechanism that is put forward by its advocates as an ultimate explanation (...) of human cooperation. The West camp thus accuses advocates of SR for not heeding Mayr’s original distinction between ultimate and proximate causation. The Laland camp praises advocates of SR for revising Mayr’s distinction. Advocates of SR are said to replace Mayr’s uni-directional view on the relation between ultimate and proximate causes by the bi-directional one of reciprocal causation. The paper argues that both the West camp and the Laland camp misrepresent what advocates of SR are up to. The West camp is right that SR is a proximate cause of human cooperation. But rather than putting forward SR as an ultimate explanation, as the West camp argues, advocates of SR believe that SR itself is in need of ultimate explanation. Advocates of SR tend to take gene-culture co-evolutionary theory as the correct meta-theoretical framework for advancing ultimate explanations of SR. Appearances notwithstanding, gene-culture coevolutionary theory does not imply Laland et al.’s notion of reciprocal causation. “Reciprocal causation” suggests that proximate and ultimate causes interact simultaneously, while advocates of SR assume that they interact sequentially. I end by arguing that the best way to understand the debate is by disambiguating Mayr’s ultimate-proximate distinction. I propose to reserve “ultimate” and “proximate” for different sorts of explanations, and to use other terms for distinguishing different kinds of causes and different parts of the total causal chain producing behavior. (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)

The theoretical status of ‘niche construction’ in evolution is intensely debated. Here we substantiate the reasons for different interpretations. We consider two concepts of niche construction brought to bear on evolutionary theory; one that emphasizes how niche construction contributes to selection and another that emphasizes how it contributes to development and inheritance. We explain the rationale for claims that selective and developmental niche construction motivate conceptual change in evolutionary biology and the logic of those who reject these claims. Our analysis (...) shows how the contention arises from alternative assumptions regarding the causal independence of the processes that generate variation, differential fitness and inheritance. (shrink)

Biological evolution and cultural evolution are distinct evolutionary processes; they are apparent also in human language, where both processes contributed in shaping its evolution. However, the nature of the interaction between these two processes is still debated today. It is often claimed that the emergence of modern language was preceded by the evolution of a language-ready brain: the latter is usually intended as a product of biological evolution, while the former is believed to be the consequence of cultural processes. I (...) label this account separationist, and I argue that current evidence coming from the archaeological and fossil record suggests that cultural evolution has played a pivotal role in shaping not only modern language but also the language-ready brain. I label the latter the integrative perspective. I analyze the two accounts from two different conceptual frameworks, the Standard Evolutionary Theory and the Extended Evolutionary Synthesis, and I discuss the limits of the sharp proximate–ultimate distinction in this specific case study arguing in favor of a causal feedback model. I extend the argument to the debate on gradual and punctuational evolutionary rates, suggesting that the debate on patterns in language evolution should leave room for discussion of the nature of the processes involved in producing those patterns, providing reasons in favor of the eco-evolutionary feedback dynamics underlying the integrative perspective. (shrink)

The main purpose of this article is to consider the significance of different types of memory and non-genetic inheritance and different biosemiotic systems for the origin and evolution of language. It presents language and memory as distributed, heteronomous and system-determined processes implemented in biological and social domains. The article emphasises that language and other sign systems are both ecological and inductive systems that were caused by and always correlate with the environment and deductive systems that are inherited by and depend (...) on the internal development of organisms, individuals, and societies. The article also claims that the origin, re-occurrence and evolution of naturally-emerging sign systems presuppose their retention and accumulation in physical, biological, individual, and social types of memory and reinforcement and maintenance by conventional and deliberate social regulation and accumulation. All of this allows language and other sign systems to be situation-relevant and to be transmitted through generations without their constant reinvention. The novelty of the proposed theory of language origin and evolution is in interdisciplinary integration of biosemiotic studies, systems approaches to language and studies of inheritance systems presented by ‘Extended evolutionary synthesis’. (shrink)

The current debate over extending inheritance and its evolutionary impact has focused on adding new categories of non-genetic factors to the classical transmission of DNA, and on trying to redefine inheritance. Transmitted factors have been mainly characterized by their directions of transmission and the way they store variations. In this paper, we leave aside the issue of defining inheritance. We rather try to build an evolutionary conceptual framework that allows for tracing most, if not all forms of transmission and makes (...) sense of their different tempos and modes. We discuss three key distinctions that should in particular be the targets of theoretical and empirical investigation, and try to assess the interplay among them and evolutionary dynamics. We distinguish two channels of transmission, two measurements of the temporal dynamics of transmission, respectively across and within generations, and two types of transmitted factors according to their evolutionary relevance. By implementing these three distinctions we can then map different forms of transmission over a continuous space describing the combination of their varying dynamical features. While our aim is not to provide yet another model of inheritance, putting together these distinctions and crossing them, we manage to offer an inclusive conceptual framework of transmission, grounded in empirical observation, and coherent with evolutionary theory. This interestingly opens possibilities for qualitative and quantitative analyses, and is a necessary step, we argue, in order to question the interplay between the dynamics of evolution and the dynamics of multiple forms of transmission. (shrink)

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

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

We develop a conceptual framework that connects biological heredity and organization. We refer to heredity as the cross-generation conservation of functional elements, defined as constraints subject to organizational closure. While hereditary objects are functional constituents of biological systems, any other entity that is stable across generations—and possibly involved in the recurrence of phenotypes—belongs to their environment. The central outcome of the organizational perspective consists in extending the scope of heredity beyond the genetic domain without merging it with the broad category (...) of cross-generation stability. After discussing some implications, we conclude with a reflection on the relationship between stability and variation. 1Introduction2From Extended Heredity to Cross-generation Stability 2.1Extending the scope of heredity: A brief state of the art2.2Rethinking heredity: Conceptual challenges3Biological Heredity in Light of Organization 3.1Biological organization within organisms and beyond3.2Extending organization in time3.3What is biological heredity?4Implications and Objections 4.1Heredity as a specific kind of cross-generation stability4.2Heredity at various levels of description4.3Non-functional and dysfunctional objects5Conclusions: From Conservation to Variation. (shrink)

The rise of molecular epigenetics over the last few years promises to bring the discourse about the sociality and susceptibility to environmental influences of the brain to an entirely new level. Epigenetics deals with molecular mechanisms such as gene expression, which may embed in the organism “memories” of social experiences and environmental exposures. These changes in gene expression may be transmitted across generations without changes in the DNA sequence. Epigenetics is the most advanced example of the new postgenomic and context-dependent (...) view of the gene that is making its way into contemporary biology. In my article I will use the current emergence of epigenetics and its link with neuroscience research as an example of the new, and in a way unprecedented sociality of contemporary biology. After a review of the most important developments of epigenetic research, and some of its links with neuroscience, in the second part I reflect on the novel challenges that epigenetics presents for the social sciences for a re-conceptualization of the link between the biological and the social in a postgenomic age. Although epigenetics remains a contested, hyped, and often uncritical terrain, I claim that especially when conceptualized in broader non-genecentric frameworks,. (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 possible evolutionary significance of epigenetic memory and codes is a key problem for extended evolutionary synthesis and biosemiotics. In this paper, some less known original works are reviewed which highlight theoretical parallels between current evolutionary epigenetics, on the one hand, and its predecessors in the eco-physiology of higher nervous activity, on the other. Recently, these areas have begun to converge, with first evidence now indicating the possibility of transgenerational epigenetic inheritance of conditional associations in the mammalian nervous system, and (...) related findings in other taxa. This can serve as an interesting example of evolutionary code-making, where the molecular mechanisms underlying arbitrary associations between stimuli involve lasting changes in gene expression that may be transmitted epigenetically across generations, and which in some cases could be further assimilated into the genome over subsequent evolution. Although preliminary, such epigenetic scenarios would also offer an interesting, if so far overlooked parallel to earlier research carried out by one of I.P. Pavlov’s leading students, acad. P.K. Anokhin, and his colleagues, but also by eminent eco-physiologists of the time, several of whom offered arguments for the possibility of unconditional reflexes representing evolutionarily later, specialized, and reduced forms of associative reflexes, from which they may be derived. Although discarded under the growing dominance of modern synthesis, these early epigenetic investigations may deserve renewed attention in the modern context, and if further confirmed, could open essentially new perspectives on the morphofunctional evolution of the nervous system. (shrink)

Many biologists still believe in a sort of post-Cartesian foundation of reality wherein objects are independent of subjects which cognize them. Recent research in behaviour, cognition, and psychology, however, provides plenty of evidence to the effect that the perception of an object differs depending on the kind of animal observer, and also its personality, hormonal, and sensorial set-up etc. In the following, I argue that exposed surfaces of organisms interact with other organisms’ perception to form semiautonomous relational entities called semantic (...) organs, which participate in biological reality as discrete heritable evolutionary units. The inner dimensions and potentialities of an organism can enter the senses of another living being when effectively expressed on the outer surfaces of the former and meaningfully perceived by the latter. Semantic organs have three basic sources of variability: intrinsic, i.e., genetic, epigenetic, and developmental processes; extrinsic, meaning the biotic and abiotic environmental conditions which affect the developmental generators of intrinsic variability; and perceptual, stemming from differences in the subject-specific interpretation of a SO’s structural basis. Extrinsic and intrinsic sources of variability are, however, just precursors to semantic organs. SOs are relational entities which always come into existence through an act of perception and their actual form depends both on the physical potentialities of the bearer and the species- or group-specific interpretation of the receiver. (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)

The Central Dogma of molecular biology, which holds that DNA makes protein and not the other way around, is as influential as it is controversial. Some believe the Dogma has outlived its usefulness, either because it fails to fully capture the ins-and-outs of protein synthesis (Griffiths and Stotz, 2013; Stotz, 2006), because it turns on a confused notion of information (Sarkar, 2004), or because it problematically assumes the unidirectional flow of information from DNA to protein (Gottlieb, 2001). This paper evaluates (...) an underexplored defense of the Dogma, which relies on the assumption that the Dogma and the Inheritance of Acquired Traits, a principle which dates as far back as Jean Baptiste-Lamarck, are incompatible principles (Smith, 1993; Judson, 1979; Dawkins, 1970; Cobb, 2017; Wilkins, 2002; Graur, 2018). By appealing to empirical evidence in molecular science, I argue that this apparent incompatibility is indeed merely apparent. I conclude by briefly demonstrating how these considerations bear on the topic of conceptual pluralism in the philosophy of science (Stencel and Proszewska, 2018; Lu and Bourrat, 2018). (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)

Reading Foucault’s work on power and subjectivity alongside “developmentalist” approaches to evolutionary biology, this article endorses poststructuralist critiques of political ideals grounded in the value of subjective agency. Many political theorists embrace such critiques, of course, but those who do are often skeptical of liberal democracy, and even of normative theory itself. By contrast, those who are left to theorize liberal democracy tend to reject or ignore poststructuralist insights, and have continued to employ dubious ontological assumptions regarding human agents. Against (...) both groups, I argue that Foucault’s poststructuralism must be taken seriously, but that it is ultimately consistent with normative theory and liberal democracy. Linking poststructuralist attempts to transcend the dichotomy between agency and structure with recent efforts by evolutionary theorists to dissolve a similarly stubborn opposition between nature and nurture, I develop an anti-essentialist account of human nature and agency that vindicates poststructuralist criticism while enabling a novel defense of liberal democracy. (shrink)

In recent years, biologists and philosophers of science have argued that evolutionary theory should incorporate more seriously the idea of ‘reciprocal causation.’ This notion refers to feedback loops whereby organisms change their experiences of the environment or alter the physical properties of their surroundings. In these loops, in particular niche constructing activities are central, since they may alter selection pressures acting on organisms, and thus affect their evolutionary trajectories. This paper discusses long-standing problems that emerge when studying such reciprocal causal (...) processes between organisms and environments. By comparing past approaches to reciprocal causation from the early twentieth century with contemporary ones in niche construction theory, we identify two central reoccurring problems: All of these approaches have not been able to provide a conceptual framework that allows maintaining meaningful boundaries between organisms and environments, instead of merging the two, and integrating experiential and physical kinds of reciprocal causation. By building on case studies of niche construction research, we provide a model that is able to solve these two problems. It allows distinguishing between mutually interacting organisms and environments in complex scenarios, as well as integrating various forms of experiential and physical niche construction. (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)

In this paper, I will conduct three interrelated analyses. First, I will develop an analysis of various concepts in the history of biology that used to refer to individual-level phenomena but were then reinterpreted by the Modern Synthesis in terms of populations. Second, I argue that a similar situation can be found in contemporary biological theory. While different approaches reflect on the causal role of developing organisms in evolution, proponents of the Modern Synthesis avoid any substantial change by reinterpreting and (...) explaining individual-level phenomena from a population perspective. Finally, I will approach this debate by advocating the statistical reading of natural selection, which holds that explanations by natural selection are statistical. I will argue that the above historical conceptual reinterpretations belong to a new explanatory strategy developed by the Modern Synthesis based on population thinking. The reinterpretation of concepts at the individual level is part of the explanatory framework of the Modern Synthesis and the empty role of development within this framework. Moreover, the statistical perspective adopted here allows for the integration of two explanatory models: population-statistical and individual-causal. Finally, I will argue that this pluralistic framework can help to define the explanatory scope of the different biological approaches in order to achieve a coherent integration of development into evolution without rejecting population thinking. (shrink)

In this paper I take evolutionary biology as an example to reflect on the role of philosophy and on the transformations that philosophy is constantly stimulated to do in its own approach when dealing with science. I consider that some intellectual movements within evolutionary biology (more specifically, the various calls for 'synthesis') express metascientific views, i.e., claims about 'what it is to do research' in evolutionary biology at different times. In the construction of metascientific views I see a fundamental role (...) to be played by philosophy, and, at the same time, a need to complement the philosophical methods with many more methods coming from other sciences. What leads philosophy out of itself is its own attention to scientific practice. My humble methodological suggestions are, at this stage, only meant to help us imagine metascientific views that are built with a more scientific, interdisciplinary approach, in order to attenuate partiality, subjectivity and impressionism in describing the scientific community. And yet, we should not be naïve and imbued with the myth of 'datadriven' research, especially in this field: other complex issues about metascientific views call for a serious, constant philosophical reflection on scientific practice. (shrink)