The nature of biological autonomy and the choice of an appropriate framework for understanding it are subjects of ongoing debates in philosophy of biology and cognitive science. The enactivist view, originating with Humberto Maturana and Francisco Varela, emphasizes the concepts of organizational or operational closure and structural coupling between the organism and its environment as central in the context of autonomy. The proponents of this view contrast it with the traditional cybernetic paradigm based on inputs, outputs, feedback, and internal representations. (...) This essay takes a synoptic view of the relevant issues and situates them in the context of modern theory of systems and control, in particular the behavioral approach developed by Jan Willems. It is argued that the behavioral approach, which favors a fairly liberal notion of control as interconnection without any prefigured designation of inputs and outputs, provides a more fruitful background for understanding and modeling of biological autonomy. (shrink)

The field of the philosophy of biology is flourishing in its aim to evaluate and rethink the view inherited from the previous century ---the Modern Synthesis. Different research areas and theories have come to the fore in the last decades in order to account for different biological phenomena that, in the first instance, fall beyond the explanatory scope of the Modern Synthesis. This thesis is anchored and motivated by this revolt in the philosophy of biology. -/- The central target in (...) this context is the possibility of naturalizing teleology, a classical nightmare for the history of biology itself. This requires, principally, understanding the causes of teleological explanations without assuming an unfashioned backward causation of sorts. As the riddles of teleological explanations are about their temporal dimension, I analyze different temporal scales of biological processes: evolutionary, developmental, and physiological. -/- The first one is the one defended in the context of the Modern Synthesis. As expected, one of the aims of this thesis is to evaluate the adequacy of an evolutionary account of teleological explanation. The scrutiny is negative. Evolutionary explanations in the context of the Modern Synthesis lack the necessary causal roots to naturalize teleology. Concerning the physiological scale, a long tradition pushed up by Kant and the organicist movement in the 20th century allows us to better understand how teleological explanations can be naturalized in physiological process. The key notions in this temporal scale are self-organization and the recursive, looped character of physiological process. While the physiological scale may be suitably accounted by contemporaries views, such as Autonomous Systems Theory, different central teleological phenomena remain unexplained from a purely physiological perspective. In particular, different issues concerning the (adaptive) construction of organism ---such as plasticity, robustness, variation, novelty, inheritance--- deserve an ontogenetic analysis. -/- The principal aim of this thesis is to provide a theory of teleological development that falls beyond the Modern Synthesis' framework and is prompted by different insights from the history of biology. I call it Agential Teleosemantics. It rests on two central pillars. First, that developmental processes, beyond any gene-centered stance, can be understood in informational terms; i.e. developmental processes are about the interaction of developmental resources conveying biological information. The second ingredient is agentivity, namely the idea that development is regulated by an agentive system according to the adaptiveness of the phenotypic outcomes produced. The role of agency in Agential Teleosemantic is equivalent to the role of genes in the Modern Synthesis: it is responsible for explaining the order and the adaptive complexity in the living realm. -/- The second target of this thesis regards the possibility of naturalizing intentional explanations in cognitive science. The central project involved in such an aim is known as teleosemantics. Classical teleosemantics however is etiological: it explains the teleofunctions of representational systems in terms of evolutionary processes. The different disputes in the contemporary philosophy of biology provide two insights to analyze teleosemantics in cognitive science. First, the challenges against the Modern Synthesis must be extended to the evolutionary approach of etiological teleosemantics. Second, as Agential Teleosemantics suggests an alternative source for teleofunctions ---ontogeny, I offer an attempt to integrate Agential Teleosemantics into cognitive science in order to provide an alternative teleosemantic project to understand intentional explanations in cognitive science. (shrink)

The goal of this programmatic paper is to highlight a close connection between the core problem in the philosophy of medicine, i.e. the concept of health, and the core problem of the philosophy of mind, i.e. the concept of consciousness. I show when we look at these phenomena together, taking the evolutionary perspective of modern state-based behavioural and life-history theory used as the teleonomic tool to Darwinize the agent- and subject-side of organisms, we will be in a better position to (...) make sense of them both as natural phenomena. (shrink)

This dissertation concerns the active role of the organism in evolutionary theory. In particular, it concerns how our conception of the relationship between organism and environment, and the nature of natural selection, influences the causal and explanatory role of organismic activity and behavior in evolutionary explanations. The overarching aim is to argue that the behaviors and activities of organisms can serve both as the explananda (that which is explained) and the explanantia (that which explains) in evolutionary explanations. I attempt to (...) achieve this aim by offering three central arguments. First, that the organism-environment relation is the ontologically basic unit of biology. A common way of conceiving the relationship between organism and environment is as a duality—as two causally independent systems that overlap through interaction. I think this is a mistake. There cannot be organisms without environments and vice versa. They are codependent and codetermined. Second, that natural selection is an ecological process. By this I mean that natural selection acts primarily on organism-environment interactions. It is only in virtue of organisms interacting with their environments that there can be fitness differences amongst individual organisms in a population. The ecological approach to natural selection also entails that the nature of the system(s) of inheritance involved in the reoccurrence of favorable organism-environment interactions is, in principle, immaterial to the process of selection. It only matters for selection that organism-environment interactions actually do reoccur in subsequent generations. Third, that niche construction theory—the most well-developed theoretical and conceptual framework for studying how the activities of organisms influences evolutionary dynamics—can be seen as fully a compatible and integrated part of evolutionary theory. Together, these three arguments constitute an overarching argument. Namely that we both can, and should, take organismic activity and behavior to be crucial explanatory elements in evolutionary theory. Throughout the dissertation I also show how these three arguments have consequences for other debates in the philosophy of evolutionary theory, such as the nature of evolutionary processes, the structure of selection-based explanations, the validity and utility of the proximate-ultimate distinction, and the nature of teleology in evolutionary systems. (shrink)

Darwin’s claim about natural selection is reconstructed as an empirical claim about a causal connection leading from the match of the physiology of an individual and its environment to leaving surviving progeny. Variations in this match, Darwin claims, cause differences in the survival of the progeny. Modern concepts of fitness focus the survival side of this chain. Therefore, the assumption that evolutionary theory wants to explain reproductive success in terms of a modern concept of fitness has given rise to the (...) so-called tautology problem. It is shown that the tautology problem reappears in the treatment of fitness proxies in today’s experimental evolutionary biology when these proxies are considered to indicate fitness only. Taking Darwin’s empirical claim seriously suggests, by contrast, that fitness proxies are first of all measures of the match between organism and environment, which I call the organism’s ‘fittedness’. At the same time, they are indeed related to reproductive success. Thus looking in both directions, at fitness and at fittedness, they are janiform. Acknowledging this situation not only allows for rejection of the tautology objection, but also for integration of Darwin’s argument into current evolutionary biology. It is suggested that this helps reframe and alleviate the dispute between the Modern Synthesis and the Extended Evolutionary Synthesis. (shrink)

This paper contributes to the interdisciplinary theory of collective affective niche construction, which extends the extended mind (ExM) thesis from cognitive to affective phenomena. Although theoretically innovative, the theory lacks a detailed psychological account of how collective affectivity is scaffolded. It has also been criticized for its uncritical assumption of the subject qua the autonomous user of the affective scaffolding as disposable resources, abstracting away from embedded subjectivity in particular techno-political arrangements. We propose that the social motivation hypothesis, an account (...) grounded in recent empirical and theoretical developments in psychology as well as in the classic theory of moral sentiments, will address the former criticism by explicating the basic mechanisms of human social orientation at work in collective affective niche construction. We also begin to address the latter normative criticism in mobilizing a so-called we-mode approach to collective emotion. To make these theoretical dialectics salient, we study social media as a case of collective affective niches, focusing on the impact on subjective well-being. Finally, we briefly identify promising future directions in building a normative theory of affective niche construction on the collective level. (shrink)

Recent studies demonstrating epigenetic and developmental sensitivity to early environments, as exemplified by fields like the Developmental Origins of Health and Disease (DOHaD) and environmental epigenetics, are bringing new data and models to bear on debates about race, genetics, and society. Here, we first survey the historical prominence of models of environmental determinism in early formulations of racial thinking to illustrate how notions of direct environmental effects on bodies have been used to naturalize racial hierarchy and inequalities in the past. (...) Next, we conduct a scoping review of postgenomic work in environmental epigenetics and DOHaD that looks at the role of race/ethnicity in human health (2000–2021). Although there is substantial heterogeneity in how race is conceptualized and interpreted across studies, we observe practices that may unwittingly encourage typological thinking, including: using DNA methylation as a novel marker of racial classification; neglect of variation and reversibility within supposedly homogenous racial groups; and a tendency to label and reify whole groups as pathologized or impaired. Even in the very different politico-economic and epistemic context of contemporary postgenomic science, these trends echo deeply held beliefs in Western thinking which claimed that different environments shape different bodies and then used this logic to argue for essential differences between Europeans and non-Europeans. We conclude with a series of suggestions on interpreting and reporting findings in these fields that we feel will help researchers harness this work to benefit disadvantaged groups while avoiding the inadvertent dissemination of new and old forms of stigma or prejudice. (shrink)

According to Friedrich Schleiermacher, religiosity is rooted in feeling (Gefühl). As a result of our engagement with the world, on which we depend and which we can influence, we have both a sense of dependence and of freedom. Schleiermacher speculated that a sense of absolute dependence in reflective beings with self-consciousness (human beings) gave rise to religion. Using insights from contemporary philosophy of biology and cognitive science, I seek to naturalize Schleiermacher's ideas. I moreover show that this naturalization is in (...) line with Schleiermacher's outlook on biology, as he already had evolutionary considerations in mind when he wrote the Christian Faith (1830). While Schleiermacher rejects natural theology in a narrow sense (proofs for the existence of God), his project is natural theological in a broader sense, as it roots religion in experiences that we can examine using naturalistic theories. (shrink)

Darwin provided us with a powerful theoretical framework to explain the evolution of living systems. Natural selection alone, however, has sometimes been seen as insufficient to explain the emergence of new levels of selection. The problem is one of “circularity” for evolutionary explanations: how to explain the origins of Darwinian properties without already invoking their presence at the level they emerge. That is, how does evolution by natural selection commence in the first place? Recent results in experimental evolution suggest a (...) way forward: Paul Rainey and his collaborators have shown that Darwinian properties could be exogenously imposed via what they call “ecological scaffolding.” This could solve the “black box” dilemma faced by Darwinian explanations of new levels of organization. Yet, despite “scaffolding” recently becoming a popular term in the study of cognition, culture, and evolution, the concept has suffered from vagueness and ambiguity. This article aims to show that scaffolding can be turned into a proper scientific concept able to do explanatory work within the context of the major evolutionary transitions. Doing so will allow us to once again extend the scope of the Darwinian model of evolution by natural selection. (shrink)

This special issue for the Journal for General Philosophy of Science is devoted to exploring the impact and many ramifications of current research in evolutionary epistemology. Evolutionary epistemology is an inter- and multidisciplinary area of research that can be divided into two ever-inclusive research avenues. One research avenue expands on the EEM program and investigates the epistemology of evolution. The other research avenue builds on the EET program and researches the evolution of epistemology. Since its conception, EE has developed three (...) schools of thought: adaptationist, non-adaptationist, and applied EE. Although diverse in outlook and theoretical background, these research avenues and schools share the same agenda of understanding how knowledge evolves, and how it relates to the world. In this paper, we first explain wherefrom evolutionary epistemological schools of thought developed, and then we highlight current debates in EE by briefly reviewing the papers that form part of this special issue. (shrink)

Evolutionary biologists, evolutionary epistemologists, and biosemioticians have demonstrated that organisms not merely adapt to an external world, but that they actively construct their environmental, sociocultural, and cognitive niches. Denis Noble demonstrates that such is no different for those organisms that engage in science, and he lays bare several crucial assumptions that define the scientific dogmas and practices of evolutionary biology.

Enactivism and ecological psychology converge on the relevance of the environment in understanding perception and action. On both views, perceiving organisms are not merely passive receivers of environmental stimuli, but rather form a dynamic relationship with their environments in such a way that shapes how they interact with the world. In this paper, I suggest that while enactivism and ecological psychology enjoy a shared specification of the environment as the cognitive domain, on both accounts, the structure of the environment, itself, (...) is unspecified beyond that of contingent relations with the species-typical sensorimotor capacities of perceiving organisms. This lack of specification creates a considerable gap in theory regarding the organization of organisms as coupled with their environments. I argue that this gap can be filled by drawing from resources in developmental systems theory, namely, specifying the environmental state-space as a developmental niche that shapes and is shaped by individual organisms over developmental and, on a population scale, evolutionary time. Defining the environment as an organism’s developmental niche makes it clearer how and why certain contingencies have arisen, in turn, strengthening a joint appeal to both enactivism and ecological psychology as theories asserting complementarity between organisms and their environments. (shrink)

Explaining agency is a significant challenge for those who are interested in the sciences of the mind, and non-representationalists are no exception to this. Even though both ecological psychologists and enactivists agree that agency is to be explained by focusing on the relation between the organism and the environment, they have approached it by focusing on different aspects of the organism-environment relation. In this paper, I offer a suggestion for a radical embodied account of agency that combines ecological psychology with (...) recent trends in enactive cognitive science. According to this proposal, while enactivism focuses primarily on describing how our acquired sensorimotor schemes and habits mutually equilibrate, affecting our tendency to act upon some affordances instead of others, ecological psychology focuses on studying how perceptual information contributes to the actualization of the sensorimotor schemes and habits without mediating representations, inferences, and computations. The paper concludes by briefly exploring how this ecological-enactive theory of agency can account for how socio-cultural norms shape human agency. (shrink)

This paper aims to examine the evolutionary role of affordances, that is, the possibilities for action available in our environments. There are two allegedly competing views for explaining the evolutionary role of affordances: the first is based on natural selection; the second is based on niche construction. According to the first, affordances are resources that exert selection pressure. The second view claims that affordances are ecological inheritances in the organism’s niche that are the product of a previous alteration of the (...) environment. While there seems to be a mutually exclusive definition of affordances in each of these views, I argue in this paper that the views are not competing but, rather, complementary. In this sense, affordances play the role of either resources or ecological inheritances depending on the temporal stage of the evolutionary process. I make this argument by analyzing how natural selection and niche construction affect each other even when they function independently from each other. In this light, if these two evolutionary mechanisms exert their power in parallel but at two different stages in the evolutionary history of a given econiche, then there is room to claim that affordances can be understood as both resources and ecological inheritances. This dual aspect of affordances shows their evolutionary role. (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)

Much philosophical progress has been made in elucidating the idea of evolutionary contingency in a recent re-burgeoning of the debate. However, additional progress has been impaired on three fronts. The first relates to its characterisation: the under-specification of various contingency claims has made it difficult to conceptually pinpoint the scope to which ‘contingency’ allegedly extends, as well as which biological forms are in contention. That is—there appears to be no systematic means with which to fully specify contingency claims which has (...) led to a tendency for authors to talk past each other. Secondly, on the matter of evidence, recent research has focused on the evidential import of convergent evolution which is taken to disconfirm the evolutionary contingency thesis. However, there has been a neglect of convergent evolution’s converse: ‘evolutionary idiosyncrasies’ or the singular evolution of certain forms, which I argue is evidentially supportive of evolutionary contingency. Thirdly, evolutionary contingency has often been claimed to vary in degrees and that the debate, itself, is a matter of ‘relative significance’. However, there has been no formal method of evaluating the strength of contingency and its relative significance in a particular domain. In this paper, I address all three issues by proposing a systematic means of fully specifying contingency theses with the concept of the modal range. Secondly, I propose an account of evolutionary idiosyncrasies, investigate the explanations for their occurrences, and, subsequently, spell out their significance with respect to the evolutionary contingency thesis. Finally, having been equipped with the evidential counterpart to convergent evolution, I shall sketch a likelihood framework for evaluating, precisely on the basis of a sequence of opposing data, the strength and relative significance of evolutionary contingency in a particular domain. With this in hand, the relative observations of idiosyncrasies and convergences can be informative of the strength and relative significance of contingency in any particular domain. (shrink)

This volume provides a broad overview of issues in the philosophy of behavioral biology, covering four main themes: genetic, developmental, evolutionary, and neurobiological explanations of behavior. It is both interdisciplinary and empirically informed in its approach, addressing philosophical issues that arise from recent scientific findings in biological research on human and non-human animal behavior. Accordingly, it includes papers by professional philosophers and philosophers of science, as well as practicing scientists. Much of the work in this volume builds on presentations given (...) at the international conference, “Biological Explanations of Behavior: Philosophical Perspectives”, held in 2008 at the Leibniz Universität Hannover in Germany. The volume is intended to be of interest to a broad range of audiences, which includes philosophers (e.g., philosophers of mind, philosophers of biology, and metaethicists), as well as practicing scientists, such as biologists or psychologists whose interests relate to biological explanations of behavior. (shrink)

In this paper, I analyze the disruptive impact of Darwinian selectionism for the century-long tradition in which the environment had a direct causative role in shaping an organism’s traits. In the case of humans, the surrounding environment often determined not only the physical, but also the mental and moral features of individuals and whole populations. With its apparatus of indirect effects, random variations, and a much less harmonious view of nature and adaptation, Darwinian selectionism severed the deep imbrication of organism (...) and milieu posited by these traditional environmentalist models. This move had radical implications well beyond strictly biological debates. In my essay, I discuss the problematization of the moral idiom of environmentalism by William James and August Weismann who adopted a selectionist view of the development of mental faculties. These debates show the complex moral discourse associated with the environmentalist-selectionist dilemma. They also well illustrate how the moral reverberations of selectionism went well beyond the stereotyped associations with biological fatalism or passivity of the organism. Rereading them today may be helpful as a genealogical guide to the complex ethical quandaries unfolding in the current postgenomic scenario in which a revival of new environmentalist themes is taking place. (shrink)

Richard Lewontin is often cited as an inspiration and founder of what is now known as Niche Construction Theory. The first goal of this paper is to argue that they present distinct arguments from niche construction against Adaptationism. While Niche Construction Theory argues that natural selection is not the only adaptive evolutionary force, Lewontin rejects the externalist characterization of natural selection. The key difference lies in the types of phenomena that are allowed to count as “niche construction” and their argumentative (...) roles. The second goal is to argue that it is time to revive Lewontin’s argument. I argue that it finds renewed support in Denis Walsh’s ecological affordance framework and empirical evidence from ecological developmental biology. Reexamining the roots of Niche Construction Theory, I suggest that the rich conceptual resources therein provide a way for Niche Construction Theory to also develop a neo-Lewontinian argument against the externalism of natural selection explanations. (shrink)

Evolutionary biology is striking for its ability to explain a large and diverse range of empirical phenomena on the basis of a few general theoretical principles. This article offers a philosophical perspective on the way that evolutionary biology has come to achieve such impressive generality, by focusing on “the strategy of endogenization”. This strategy involves devising evolutionary explanations for biological features that were originally part of the background conditions, or scaffolding, against which such explanations take place. Where successful, the strategy (...) moves biology a step closer to the ideal of explaining as much as possible from evolutionary first principles. The strategy of endogenization is illustrated through a series of biological examples, historical and recent, and its philosophical implications are explored. (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)

Many have argued that there is no reason why natural selection should cause directional increases in measures such as body size or complexity across evolutionary history as a whole. In this paper I argue that this conclusion does not hold for selection for adaptations to environmental variability, and that, given the inevitability of environmental variability, trends in adaptations to variability are an expected feature of evolution by natural selection. As a concrete instance of this causal structure, I outline how this (...) may be applied to a trend in phenotypic plasticity. (shrink)

The traditional practice of establishing morphological types and investigating morphological organization has found new support from evolutionary developmental biology (evo-devo), especially with respect to the notion of body plans. Despite recurring claims that typology is at odds with evolutionary thinking, evo-devo offers mechanistic explanations of the evolutionary origin, transformation, and evolvability of morphological organization. In parallel, philosophers have developed non-essentialist conceptions of natural kinds that permit kinds to exhibit variation and undergo change. This not only facilitates a construal of species (...) and higher taxa as natural kinds, but also broadens our perspective on the diversity of kinds found in biology. There are many different natural kinds relevant to the investigative and explanatory aims of evo-devo, including homologues and developmental modules. (shrink)

David Hume's famous riddle of induction implies a second problem related to the question of whether the laws and principles of nature might change in the course of time. Claims have been made that modern developments in physics and astrophysics corroborate the translational invariance of the laws of physics in time. However, the appearance of a new general principle of nature, which might not be derivable from the known laws of physics, or that might actually be a non-physical one (this (...) means completely independent of physical science) supports the notion that the course of nature can change in time. Here it is argued that natural selection satisfies the criteria that identify a general principle of nature which so far, appears to be non-derivable from the known laws of physics and therefore, it is likely that it arose in the course of time, thus leaving open again the quest for a true solution to Hume's second problem. (shrink)

Evolutionary debunking arguments (EDAs) have attracted extensive attention in meta-ethics, as they pose an important challenge to moral realism. Mogensen (2015) suggests that EDAs contain a fallacy, by confusing two distinct forms of biological explanation – ultimate and proximate. If correct, the point is of considerable importance: evolutionary genealogies of human morality are simply irrelevant for debunking. But we argue that the actual situation is subtler: while ultimate claims do not strictly entail proximate ones, there are important evidential connections between (...) the two. Attending to these connections clears ground for a new and improved EDA. However, it also brings into view some possible problems with EDAs that have been largely neglected so far. (shrink)

The definition of biological individuality is one of the most discussed topics in philosophy of biology, but current debate has focused almost exclusively on evolution-based accounts. Moreover, several participants in this debate consider the notions of a biological individual and an organism as equivalent. In this paper, I show that the debates would be considerably enriched and clarified if philosophers took into account two elements. First, physiological fields are crucial for the understanding of biological individuality. Second, the category of biological (...) individuals should be divided into two subcategories: physiological individuals and evolutionary individuals, which suggests that the notions of organism and biological individual should not be used interchangeably. I suggest that the combination of an evolutionary and a physiological perspective will enable biologists and philosophers to supply an account of biological individuality that will be both more comprehensive and more in accordance with scientific practices. (shrink)

As a first shot, one might say that environmental ethics is concerned distinctively with the moral relations that exist between, on the one hand, human beings and, on the other, the non-human natural environment. But this really is only a first shot. For example, one might be inclined to think that at least some components of the non-human natural environment have independent moral status, that is, are morally considerable in their own right, rather than being of moral interest only to (...) the extent that they contribute to human well-being. If so, then one might be moved to claim that ethical matters involving the environment are best cashed out in terms of the dutes and responsibilities that human beings have to such components. If, however, one is inclined to deny independent moral status to the non-human natural environment or to any of its components, then one might be moved to claim that the ethical matters in question are exhaustively delineated by those moral relations existing between individual human beings, or between groups of human beings, in which the non-human natural environment figures. One key task for the environmental ethicist is to sort out which, if either, of these perspectives is the right one to adopt—as a general position or within particular contexts. I guess I don’t need to tell you that things get pretty complicated pretty quickly. (shrink)

In this paper I argue, first, that human lifeways depend on cognitive capital that has typically been built over many generations. This process of gradual accumulation produces an adaptive fit between human agents and their environments; an adaptive fit that is the result of hidden‐hand, evolutionary mechanisms. To explain distinctive features of human life, we need to understand how cultures evolve. Second, I distinguish a range of different evolutionary models of culture. Third, I argue that none of meme‐based models, dual (...) inheritance models, nor Boyd and Richerson's models fully succeed in explaining this adaptive fit between agent and the world. I then briefly develop an alternative. Finally, I explore (in a preliminary way) constraints on cultural adaptation. The processes of cultural evolution sometimes built a fit between agents and their environment, but they do not always do so. Why is folk medicine, for example, so much less reliable than folk natural history? (shrink)

In this paper, we intend to discuss if and in what sense semiosis (meaning process, cf. C. S. Peirce) can be regarded as an "emergent" process in semiotic systems. It is not our problem here to answer when or how semiosis emerged in nature. As a prerequisite for the very formulation of these problems, we are rather interested in discussing the conditions which should be fulfilled for semiosis to be characterized as an emergent process. The first step in this work (...) is to summarize a systematic analysis of the variety of emergence theories and concepts, elaborated by Achim Stephan. Along the summary of this analysis, we pose fundamental questions that have to be answered in order to ascribe a precise meaning to the term "emergence" in the context of an understanding of semiosis. After discussing a model for explaining emergence based on Salthe's hierarchical structuralism, which considers three levels at a time in a semiotic system, we present some tentative answers to those questions. (shrink)

In this article I address the issue of the ontological conditions of possibility for a naturalistic notion of emergence, trying to determine its fundamental differences from the atomist, vitalist, preformationist and potentialist alternatives. I will argue that a naturalistic notion of ontological emergence can only succeed if we explicitly refuse the atomistic fundamental ontological postulate that asserts that every entity is endowed with a set of absolutely intrinsic properties, being qualitatively immutable through its extrinsic relations. Furthermore, it will be shown (...) that, ironically enough, this metaphysical assumption is implicitly shared by all the above mentioned alternatives to Emergentism. The current article concludes that the notion of organization by itself is not enough, and that ontological emergence can only be justified by assuming a relational ontological perspective that, in opposition both to atomism and holism, defends that the existence-conditions, the identity and the causal behavior of any emergent systemic property can only be conceived, and explained, as constructed by and through specific networks of qualitatively transformative relational processes that occur between the system’s components and between the system and its environment. Additionally, I try to explain how one can make sense of the idea that an emergent phenomenon is both dependent on, and autonomous from, its emergence base. (shrink)

The writings of Joseph Henry Woodger (1894–1981) are often taken to exemplify everything that was wrongheaded, misguided, and just plain wrong with early twentieth-century philosophy of biology. Over the years, commentators have said of Woodger: (a) that he was a fervent logical empiricist who tried to impose the explanatory gold standards of physics onto biology, (b) that his philosophical work was completely disconnected from biological science, (c) that he possessed no scientific or philosophical credentials, and (d) that his work was (...) disparaged – if not altogether ignored – by the biologists and philosophers of his era. In this paper, we provide the first systematic examination of Woodger’s oeuvre, and use it to demonstrate that the four preceding claims are false. We argue that Woodger’s ideas have exerted an important influence on biology and philosophy, and submit that the current consensus on his legacy stems from a highly selective reading of his works. By rehabilitating Woodger, we hope to show that there is no good reason to continue to disregard the numerous contributions to the philosophy of biology produced in the decades prior to the professionalization of the discipline. (shrink)

A number of areas of biology raise questions about what is of value in the natural environment and how we ought to behave towards it: conservation biology, environmental science, and ecology, to name a few. Based on my experience teaching students from these and similar majors, I argue that the field of environmental ethics has much to teach these students. They come to me with pent-up questions and a feeling that more is needed to fully engage in their subjects, and (...) I believe some exposure to environmental ethics can help focus their interests and goals. I identify three primary areas in which environmental ethics can con- tribute to their education. The first is an examination of who (or what) should be considered to be part of our moral community (i.e., the community to whom we owe direct duties). Is it humans only? Or does it include all sentient life? Or all life? Or ecosystems considered holistically? Often, readings implicitly assume one or more of these answers; the goal is to make the student more sensitive to these implicit claims and to get them to think about the different reasons that support them. The second area, related to the first, is the application of the different answers concerning the extent of the ethical community to real environmental issues and problems. Students need to be aware of how the different answers concerning the moral community can imply conflicting answers for how we should act in certain cases and to think about ways to move toward conflict resolution. The third area in which environmental ethics can contribute is a more conceptual one, focusing on central concepts such as biodiversity, sustainability, species, and ecosystems. Exploring and evaluating various meanings of these terms will make students more reflective and thoughtful citizens and biologists, sensitive to the implications that different conceptual choices make. (shrink)

Holism and reductionism are usually seen as opposite and mutually exclusive approaches to nature. Recently, some have come to see them as complementary rather than mutually exclusive. In this book I have argued that, even stronger, they should be seen as mutually dependent and co-operating research programmes. I have discussed holism and reductionism in biology in general and in ecology in particular. After an introductory chapter I have provided an overview of holistic and reductionistic positions in biology, and of the (...) reduction problems to which they relate. I have argued that it is extremely important to distinguish between ontological, epistemological and methodological aspects of these problems. The overview has shown that there are actually several approaches to reduction problems in biology, which can be characterized as more or less radically or moderately holistic or reductionistic. It has shown also that there are three major ’contradistinctions’ between holism and reductionism in biology, to wit the doctrine of emergence versus the reduction thesis; functional explanations versus causal explanations; and phenomenology versus mechanicism. In later chapters I have shown that these ’contradistinctions’ are not really contradistinctions at all. In the next chapters I have discussed the terms ’reduce’ and ’reduction’ in science. In chapter 3 I have discussed the reduction of laws and theories. I have shown that there are many different types of reduction depending on the auxiliary hypotheses that are being used in addition to the reducing theory: approximation-, aggregation, correlation and/or identification-hypotheses. The distinctions between these types has later, in chapter 5, proved to be of great importance in defining the concept of ’emergence’. The major conclusion of chapter 3 was that reduction is an epistemological issue, pertaining to logical relations between statements or systems of statements. It should not be confused, therefore, with ’ontological reduction’, certainly not in one of its ordinary senses, such as diminishing, devaluating or the like. Scientific reductions are, with the exception of instrumentalistic reductions, kinds of explanation, and, moreover, non-eliminative kinds of explanation. This means that a reduced law or theory is not eliminated by the reducing theory but rather consolidated or even reinforced. The ontology of the reduced law or theory is thereby also being consolidated. In chapter 4 I have argued that the same holds for reductions of concepts. Concept reductions are supposed to be accomplished by means of so-called ontological identity relations, which are in turn supposed to connect terms in the law or theory to be reduced, which do not occur in the reducing theory, with theoretical terms of the reducing theory. Because they are called ontological identity relations, several philosophers have thought that concept reductions imply some form of ontological reduction. And because they often occur in the context of micro-reductions, some have even thought that concept reductions are themselves micro-reductions. As a result, the idea has arisen that ontological identity relations are connections between macro-concepts and micro-concepts. However, concept reductions by means of ontological identity relations cannot be micro-reductions, because micro-reduction is, by definition, reduction with an aggregation step. Moreover, ontological identity relations are relations between different representations of and the same type of thing or attribute. That is, they express different epistemological sides of the same ontological coin. Therefore, concept reductions cannot be considered ontological reductions either, certainly not in the ordinary sense of diminishing, devaluating or the like. Thus, concept reduction is, like law or theory reduction, an epistemological issue: it pertains to relations between, in this case, different concepts or representations of one and the same type of thing or attribute. ’Ontological reduction’, then, may, in the context of scientific reductions, actually be considered a contradiction of terms. In chapter 5 I have linked this conclusion to the emergence thesis and to the alleged contradistinction between this thesis and the reduction thesis. I have shown that the doctrine of emergence actually consists of two separate claims, namely an ontological thesis stating that a whole has emergent properties which the component parts do not possess, neither separately nor in other partial combinations; and an epistemological thesis stating that emergent properties of wholes are ’in principle’ irreducible. I have argued that the first claim may be regarded as a valid, universal thesis about relations between properties of wholes and properties of parts, but that the second claim is untenable. I have discussed many emergent properties of wholes, biological as well as physico-chemical, that have proved to be explainable in terms of micro-theories about the component parts and auxiliary hypotheses. Once again, however, it is important to realize that reduction is an epistemological issue, whereas emergence, in the sense of thesis, is an ontological one. It is not ’wholes’ or ’emergent properties of wholes’ that are being reduced but statements about them. Reduction is a kind of explanation, not of explaining-away. Thus, it leaves the ontology of whatever is reduced fully intact. I have developed a new definition of the term ’emergence’ which expresses emergence in terms of the auxiliary hypotheses that may be used in reductions, in particular aggregation-, correlation- and identification-hypotheses. This has even lead to two very remarkable conclusions. First, emergence may be considered the opposite of ontological identity. And second, if there were no emergence, there wouldn’t be any reductions either. to reduce.) In this respect, therefore, there is absolutely no contradistinction between holism and reductionism. On these grounds I have introduced my thesis that holism and reductionism should rather be seen as mutually dependent, and hence co-operating, research programmes than as conflicting views of nature or of relations between sciences. Holistic programmes play an important role in science as guide programmes for reductionistic programmes by discovering or developing macro-laws or -theories about phenomena at the level of wholes, which they themselves, however, cannot explain. For these explanations they depend on the fruits of reductionistic programmes. If the latter succeed in providing the explanations they act as supply programmes for the holistic guide programmes. Reductionistic programmes depend in turn on holistic programmes for providing the macro-laws or -theories that call for these explanations. In chapter 6 I have discussed an example of this mutual dependency in the form of the reduction of the Bohr-effect in animal physiology. I have shown that this law has been reduced to the theory of allostery, applied to hemoglobin molecules in red blood cells, and that this application of the theory of allostery has been reduced to the theory of chemical bonding. I have also shown that at least six research programmes were involved in these reductions, and that the relations between these programmes can be characterized very well in terms of the model of holistic guide programmes and reductionistic supply programmes. The only, but highly significant, qualification appearing from example was that the terms ’holistic’ and ’reductionistic’ are extremely relative and should always be related to a certain given level of organization. In chapter 7 I have resolved the remaining contradistinction between holism and reductionism in biology, viz. the need for functional explanations and the demand that explanations be causal. I have shown that functional explanations are perfectly legitimate explanations which in a sense can be reconstrued as causal explanations. I have argued that functional explanations are indispensable components of more comprehensive, causal-evolutionary explanations, because the latter appeal to the adaptive value, and hence the function, of the property to be explained. In that sense functions can be regarded as adaptations, and functional explanations can be regarded as a sort of ’short-hand’ for evolutionary explanations. Finally, I have shown that in the context of functional explanations, too, co-operation of holistic and reductionistic research programmes occurs. In part 2 I have applied my thesis to ecology. I have shown that in ecology too, holismreductionism disputes notwithstanding, co-operation of holistic and reductionistic research programmes occurs, or, in so far as this appears to be not the case, that it should be strived after. In chapter 8 I have given an overview of reduction problems in ecology and of the various approaches to these problems. I have noticed that in ecology, too, there are several, radical, moderate and anti-reductionistic research strategies, with respect to the levels of both ecosystems, communities and populations. I have also noticed, however, that concrete solutions to reduction problems in ecology are frustrated by what is called the intellectual immaturity or the anomalous status of ecology. This refers to the almost complete lack of general laws and theories in ecology, at least at the higher levels of communities and ecosystems. Of a number of possible causes I have lifted out two, which lend themselves to philosophical analysis and clarification. The first one is the ambiguity of a major part of the ecological vocabulary. This was the subject of chapters 9 and 10. The second is the inhibitory effect which holism-reductionism disputes have on the growth of knowledge. This was the subject of chapter 13. In chapter 9 I have discussed the concept of an community. It appeared that the term ’community’ is used for several different entities at various levels of organization. This ambiguity alone seems to be sufficient for the lack of ’general’ laws and theories about ’communities’. My purpose in chapter 9 was to try and contribute to a solution of this problem. In doing so I have argued, first, that it seems wise to use the term ’community’ only for groups of species belonging to a single taxonomic or phylogenetic group, and to use the term ’biocoenosis’ for the higher level of organization, defined as the biotic component of an ecosystem. Next I have argued that, although species within communities may of course interact with each other, interaction is itself not a necessary or sufficient condition for community membership. Finally, I have hit upon what is known as the most notorious problem in community ecology, to wit the boundary problem, as well as the problem of heterogeneity. I have found that the cause of these problems lies in the fact that communities are almost invariably being seen as groups of populations occurring together in space and time, whereas the empirical fact is rather that populations of different species mostly do not occur together in the same space, and, moreover, that the species composition of communities changes continuously. This has lead to the suggestion that a community had better be defined as the group of individuals of two or more species occurring in the area of intersection of populations of these species. It is only in such intersection areas that one can really talk of the co-occurrence of species. I have shown the empirical adequacy of this definition by discussing various salt marsh plant communities on the Dutch island of Schiermonnikoog. In chapter 10 I have clarified two other important, yet highly controversial, concepts in ecology, to wit habitat and niche. It appeared that there are at least four different habitat concepts in ecology, and as many niche concepts, the additional complication being that two of these habitat concepts correspond to two of these niche concepts, whence the distinction between habitat and niche is blurred. In addition, different habitat concepts appeared to correspond to different environment concepts and also to different biotope concepts. My purpose in chapter 10 was to disentangle all these concepts from one another and to supply each of them with a suitable term. I have done so by keeping a close eye on commonly accepted notions about habitat differentiation and niche differentiation, two ’principles’ allowing one to explain the coexistence or non-coexistence of species in communities. The results can be found in boxes 5 and 7 in chapter 10. In chapter 11 I was finally able to substantiate my claim that in ecology, too, co-operation of holistic and reductionistic research programmes occurs. I have discussed the reduction of the classical competition model of Lotka and Volterra to modern niche theory. The Lotka/Volterra model is a phenomenological model: it describes the possible effects of competition between two species. Modern niche theory, on the other hand, is a mechanistic theory, in which both the objects of competition and a mechanism are being specified. I have shown that the reduction was established with the help of a relatively simple identification hypothesis and two relatively simple aggregation hypotheses, making it yet another example of heterogeneous microreduction in biology. In the reduction, the Lotka/Volterra model acted as a holistic guide programme and modern niche theory acted as a reductionistic supply programme. In chapter 12 I have discussed another example of co-operation in the form of the approximative reduction of MacArthur and Wilson’s equilibrium theory of island biogeography. I have argued that this can be seen as a holistic model, which has been of great unifying and especially heuristic value, but which in time has been found to be a bit too simple and in some respects had to be corrected. Therefore, it can also be seen as an idealization in the sense of the model of idealization and concretization, which subsequent research programmes have concretized. Because these concretizations have a reductive character, we can see in this structure of idealization and concretization yet another example of the co-operation of holistic and reductionistic research programmes in ecology. A philosophically interesting side-conclusion was that the model of idealization and concretization thus appears to apply also to relations between research programmes, and not only to programme-internal developments. In chapter 13 I have discussed an example of the inhibitory effect that holism-reductionism disputes may have on the growth of knowledge. The example concerns a controversy in island biogeography over the role of interspecific competition in structuring communities. This controversy has lasted for about ten years, appeares to have died out rather than to have been resolved, and has produced nothing new in the field of explanations of community structure. I have shown that the major factors underlying the controversy were differing, in casu holistic and reductionistic, views of communities. Though this presented a problem for my thesis, I have argued that, ultimately, the controversy can be resolved, and in a sense has been resolved, in a way that corroborates my thesis. In the epilogue, finally, I have mentioned some remaining problems concerning the adequacy of the present reduction model in dealing with reduction in ecology. These problems pertain to the question of specificty of ecological laws and theories, the likelihood that reductions in ecology generally occur on the basis of several reducing micro-theories, and the possible relationship between the former two points. (shrink)

Developmental Systems Theory (DST) emphasises the importance of non-genetic factors in development and their relevance to evolution. A common, deflationary reaction is that it has long been appreciated that non-genetic factors are causally indispensable. This paper argues that DST can be reformulated to make a more substantive claim: that the special role played by genes is also played by some (but not all) non-genetic resources. That special role is to transmit inherited representations, in the sense of Shea (2007: Biology and (...) Philosophy, 22, 313-331). Formulating DST as the claim that there are non-genetic inherited representations turns it into a striking, empirically-testable hypothesis, driving the sort of investigations that are only now beginning to appear in the scientific literature. DST’s characteristic rejection of a gene vs. environment dichotomy is preserved, but without dissolving all potentially explanatory distinctions into an interactionist causal soup, as some have alleged. (shrink)

The category of ‘organism’ has an ambiguous status: is it scientific or is it philosophical? Or, if one looks at it from within the relatively recent field or sub-field of philosophy of biology, is it a central, or at least legitimate category therein, or should it be dispensed with? In any case, it has long served as a kind of scientific “bolstering” for a philosophical train of argument which seeks to refute the “mechanistic” or “reductionist” trend, which has been perceived (...) as dominant since the 17th century, whether in the case of Stahlian animism, Leibnizian monadology, the neo-vitalism of Hans Driesch, or, lastly, of the “phenomenology of organic life” in the 20th century, with authors such as Kurt Goldstein, Maurice Merleau-Ponty, and Georges Canguilhem. In this paper I try to reconstruct some of the main interpretive ‘stages’ or ‘layers’ of the concept of organism in order to critically evaluate it. How might ‘organism’ be a useful concept if one rules out the excesses of ‘organismic’ biology and metaphysics? Varieties of instrumentalism and what I call the ‘projective’ concept of organism are appealing, but perhaps ultimately unsatisfying. (shrink)

The question “What is an organism?”, formerly considered as essential in biology, has now been increasingly replaced by a larger question, “What is a biological individual?”. On the grounds that i) individuation is theory-dependent, and ii) physiology does not offer a theory, biologists and philosophers of biology have claimed that it is the theory of evolution by natural selection which tells us what counts as a biological individual. Here I show that one physiological field, immunology, offers a theory, which makes (...) possible a biological individuation based on physiological grounds. I give a new answer to the question of the individuation of an organism by linking together the evolutionary and the immunological approaches to biological individuation. (shrink)

Debate about adaptationism in biology continues, in part because within “the” problem of assessing adaptationism, three distinct problems are mixed together. The three problems concern the assessment of three distinct adaptationist positions, each of which asserts the central importance of adaptation and natural selection to the study of evolution, but conceives this importance in a different way. As there are three kinds of adaptationism, there are three distinct "anti-adaptationist" positions as well. Or putting it more formally, there are three different (...) dimensions here, and strongly adaptationist views, strongly anti-adaptationist views, and moderate views are possible for each dimension. (shrink)

The distinction between phenotype and genotype is fundamental to the understanding of heredity and development of organisms. The genotype of an organism is the class to which that organism belongs as determined by the description of the actual physical material made up of DNA that was passed to the organism by its parents at the organism's conception. For sexually reproducing organisms that physical material consists of the DNA contributed to the fertilized egg by the sperm and egg of its two (...) parents. For asexually reproducing organisms, for example bacteria, the inherited material is a direct copy of the DNA of its parent. The phenotype of an organism is the class to which that organism belongs as determined by the description of the physical and behavioral characteristics of the organism, for example its size and shape, its metabolic activities and its pattern of movement. (shrink)

Karl Popper has been one of the few philosophers of sciences who has influenced scientists. I evaluate Popper's influence on our understanding of evolutionary theory from his earliest publications to the present. Popper concluded that three sorts of statements in evolutionary biology are not genuine laws of nature. I take him to be right on this score. Popper's later distinction between evolutionary theory as a metaphysical research program and as a scientific theory led more than one scientist to misunderstand his (...) position on evolutionary theory as a scientific theory. In his later work Popper also introduced what he took to be improvements of evolutionary theory. Thus far these improvements have had almost no influence on evolutionary biology. I conclude by examining the influence of Popper on the reception of cladistic analysis. (shrink)

One problem faced in discussions of the evolution of intelligence is the need to get a precise fix on what is to be explained. Terms like "intelligence," "cognition" and "mind" do not have simple and agreed-upon meanings, and the differences between conceptions of intelligence have consequences for evolutionary explanation. I hope the papers in this volume will enable us to make progress on this problem. The present contribution is mostly focused on these basic and foundational issues, although the last section (...) of the paper will look at some specific models and programs of empirical work. (shrink)

The role of reciprocal causation in Extended Evolutionary Synthesis (EES) is controversial. Proponents of EES argue that reciprocal causation is a key innovation, underpinning the necessity of EES. Conversely, critics of the EES maintain that Standard Evolutionary Theory (SET) adequately encompasses the concept of reciprocal causation, challenging the need for EES. This skepticism is rooted in two primary critiques. First, the mischaracterization of causal dynamics within SET by EES advocates leads to a misrepresentation of SET. Second, the oversight of how (...) SET incorporates and acknowledges instances of reciprocal causation leads to claims about the empirical inaptness of SET. As a result, the debate has reached an impasse, with limited progress towards a constructive examination of reciprocal causation’s significance to evolutionary explanations. This paper introduces the scope argument, which examines reciprocal causation through timescales and grain of explanations. This approach revitalizes the debate in two ways. First, reframing the debate in terms of scope clarifies the role of reciprocal causation by allowing research programs to specify targets of explanation. Second, the elements of scope (timescales and grain) elucidate the epistemic advantage of reciprocal causation in the respective research programs in question. (shrink)

What explains people's propensity to ask existential questions that they have little hope of resolving, such as: Why are we here? What, if any, is our purpose? What is the structure of the universe? That humans engage in these endeavors has long puzzled evolutionary theorists, as they go beyond the immediate demands of fending for ourselves, seeking safety, finding food, and reproducing, which occupy the daily lives of other animals. In this book, philosopher Helen De Cruz draws on a wide (...) range of disciplines and thinkers, from Aristotle and 12th-century Muslim philosopher Ibn Tufayl to Jewish thinker Abraham Heschel and science writer Rachel Carson, showing how awe and wonder lie at the heart of such existential questions. She argues that there is an emotional basis to this drive for inquiry, as wonder and awe are epistemic emotions that help us think about big-picture questions and that motivate the answers we provide to those questions. (shrink)

Wright’s “adaptive landscape” has been influential in evolutionary thinking but controversial, especially because the landscape that organisms encounter is altered by the evolutionary process itsel...

Where does the human aesthetic come from? How does it develop? By introducing the notion of the ‘niche’ as a key term in an empirically and evolutionarily informed aesthetics, this paper aims to take a fresh look at these and similar questions. It also aims to shed new light on the development and functioning of the aesthetic capacity in humans and its trans-generational transmission. Drawing on recent research developments in evolutionary biology, developmental psychology, and cognitive sciences, I shall argue that (...) the human aesthetic capacity—which I understand as the capacity, involving perceptual, cognitive and emotional processes, to enter into a pleasurable/non-pleasurable and expressive relation with objects, artworks, natural phenomena, or other people—relies on constructed environmental resources for its emergence and its ontogenetic development. (shrink)

In this paper we reject the nature–culture dichotomy by means of the idea of affordance or possibility for action, which has important implications for landscape theory. Our hypothesis is that, just as the idea of affordance can serve to overcome the subjective–objective dichotomy, the ideas of landscape and ecological niche, properly defined, would allow us to also transcend the nature–culture dichotomy. First, we introduce an overview of landscape theory, emphasizing processual landscape theory as the most suitable approach for satisfying both (...) cultural and naturalist approaches. After that, we introduce the idea of affordance and we analyze a tension between sociocultural and transcultural affordances (affordances that depend on cultural conventions and affordances that depend on lawful information and bodily aspects of agents). This tension has various implications for landscape theory and ecological niches. Our proposal is that sociocultural and transcultural aspects of affordances could be systematically accommodated if we apply niche construction theory (the theory that explains the process by which organisms modify their selective environments) as a methodological framework for explaining the emergence of ecological niches. This approach will lead us to an integrative account of landscapes as the products of the interaction between human and environmental elements, making it a clear example of a concept that transcends the nature–culture dichotomy. (shrink)