Ernst Mayr’s classical work on the nature of causation in biology has had a huge influence on biologists as well as philosophers. Although his distinction between proximate and ultimate causation recently came under criticism from those who emphasize the role of development in evolutionary processes, the formal relationship between these two notions remains elusive. Using causal graph theory, this paper offers a unified framework to systematically translate a given “proximate” causal structure into an “ultimate” evolutionary response, and illustrates evolutionary implications (...) of various kinds of causal mechanisms including epigenetic inheritance, maternal effects, and niche construction. These results not only reveal the essential interplay between proximate and ultimate causation in the study of evolution, but also provide a formal method to evaluate or discover non-standard or yet unknown evolutionary phenomena. (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)

Niche construction refers to the activities of organisms that bring about changes in their environments, many of which are evolutionarily and ecologically consequential. Advocates of niche construction theory (NCT) believe that standard evolutionary theory fails to recognize the full importance of niche construction, and consequently propose a novel view of evolution, in which niche construction and its legacy over time (ecological inheritance) are described as evolutionary processes, equivalent in importance to natural selection. Here, we subject NCT to critical evaluation, in (...) the form of a collaboration between one prominent advocate of NCT, and a team of skeptics. We discuss whether niche construction is an evolutionary process, whether NCT obscures or clarifies how natural selection leads to organismal adaptation, and whether niche construction and natural selection are of equivalent explanatory importance. We also consider whether the literature that promotes NCT overstates the significance of niche construction, whether it is internally coherent, and whether it accurately portrays standard evolutionary theory. Our disagreements reflect a wider dispute within evolutionary theory over whether the neo-Darwinian synthesis is in need of reformulation, as well as different usages of some key terms (e.g., evolutionary process). (shrink)

Concepts of cause, choice, and information are closely related. A cause is a choice that can be held responsible. It is a difference that makes a difference. Information about past causes and their effects is a valuable commodity because it can be used to guide future choices. Information about criteria of choice is generated by choosing a subset from an ensemble for ‘reasons’ and has meaning for an interpreter when it is used to achieve an end. Natural selection evolves interpreters (...) with ends. Surviving genes embody a textual record of past choices that had favorable outcomes. Consultation of these archives guides current choices. Purposive choice is well-informed difference making. (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)

Proximate and ultimate causes in evolutionary biology have come to conflate two distinctions. The first is a distinction between immediate and historical causes. The second is between explanations of mechanism and adaptive function. Mayr emphasized the first distinction but many evolutionary biologists use proximate and ultimate causes to refer to the second. I recommend that ‘ultimate cause’ be abandoned as ambiguous.

Ernst Mayr proposed a distinction between “proximate”, mechanistic, and “ultimate”, evolutionary, causes of biological phenomena. This dichotomy has influenced the thinking of many biologists, but it is increasingly perceived as impeding modern studies of evolutionary processes, including study of “niche construction” in which organisms alter their environments in ways supportive of their evolutionary success. Some still find value for this dichotomy in its separation of answers to “how?” versus “why?”questions about evolution. But “why is A?” questions about evolution necessarily take (...) the form “how does A occur?”, so this separation is illusory. Moreover, the dichotomy distorts our view of evolutionary causality, in that, contra Mayr, the action of natural selection, driven by genotype-phenotype-environment interactions which constitute adaptations, is no less “proximate” than the biological mechanisms which are altered by naturally selected genetic variants. Mayr’s dichotomy thus needs replacement by more realistic, mechanistic views of evolution. From a mechanistic viewpoint, there is a continuum of adaptations from those evolving as responses to unchanging environmental pressures to those evolving as the capacity for niche construction, and intermediate stages of this can be identified. Some biologists postulate an association of “phenotypic plasticity” (phenotype-environment covariation with genotype held constant) with capacity for niche construction. Both “plasticity” and niche construction comprise wide ranges of adaptive mechanisms, often fully heritable and resulting from case-specific evolution. Association of “plasticity” with niche construction is most likely to arise in systems wherein capacity for complex learning and behavioral flexibility have already evolved. (shrink)

Arguments for an extended evolutionary synthesis often center on the concept of “reciprocal causation.” Proponents argue that reciprocal causation is superior to standard models of evolutionary causation for at least two reasons. First, it leads to better scientific models with more predictive power. Second, it more accurately represents the causal structure of the biological world. Simply put, proponents of an extended evolutionary synthesis argue that reciprocal causation is empirically and explanatorily apt relative to competing causal frameworks. In this paper, I (...) present quantitative survey data from faculty members in biology departments at universities across the United States to evaluate this claim. The survey data indicate that a majority of the participants do not agree (i.e., most either disagree or neither agree nor disagree) that the concept of reciprocal causation confers a larger advantage on research practices. However, a majority of the participants agree that the causal framework of the extended evolutionary synthesis more accurately represents the structure of the biological world. These results demonstrate that the explanatory merits of a conceptual framework and its practical utility can come apart in interesting and informative ways. (shrink)

In this paper, we propose the expression cognitive twists for cognitive mechanisms that result from the coevolution of genes and learning. Evidence is available that at least some cultural learning mechanisms, such as imitation and language, have evolved genetically under the pressure produced by culture, even though they are mostly acquired through domain-general learning during development. Although the existence of these mechanisms is consistent with evolutionary theory, their importance has not been sufficiently emphasized by mind-centered accounts of human cognitive evolution, (...) namely evolutionary psychology and cultural evolutionary psychology. We provide concrete examples of cognitive twists, such as vocal imitation. Genetic changes in action-perception matching circuits suggest that human imitation and perhaps language are cognitive twists, namely plastic, learnable, yet genetically evolved cognitive mechanisms. We conclude that cognitive twists depict plausible evolutionary scenarios for the evolution of cognition in Homo sapiens. (shrink)

Felsefe tarihinin hiç şüphesiz en önemli filozoflarından biri olan Aristoteles (M.Ö. 384-322), aynı zamanda bilim tarihinin de en önemli kişilerindendir. Yaşadığı dönemde henüz bilim ve felsefe ayrılmamıştır; bununla birlikte Aristoteles, bilimsel ve sistemli düşünmenin, örnekleme yapmanın, bilim insanının doğaya öğrenmek için yaklaşmasının ilkelerini, ilk kez bu derece düzenli şekilde ortaya koyan kişi olmuştur. Dört neden kuramı söz konusu ilkelerin temelini oluşturmaktadır. Bu çalışmada günümüz biyolojisinin en kapsamlı tartışmalarından biri olan ‘fenotipin ne olduğu ve nasıl oluştuğu’ konusunun, Aristoteles’in dört neden kuramı (...) ile ilişkisi incelenmiştir. Genotip, fenotip ve çevrenin son derece karmaşık yolları içeren etkileşim ağı, araştırılması için birden fazla nedensel sorgulamayı gerektirir ki bu sorgulama bir anlamda Aristoteles’in dört neden kuramının, maddi, formel ve fail nedenlerinin bir arada sorgulanmasına benzerdir. (shrink)

In the 1940s, the ‘modern synthesis’ (MS) of Darwinism and genetics cast genetic mutation and recombination as the source of variability from which environmental events naturally select the fittest, such ‘natural selection’ constituting the cause of evolution. Recent biology increasingly challenges this view by casting genes as followers and awarding the leading role in the genesis of adaptations to the agency and plasticity of developing phenotypes—making natural selection a consequence of other causal processes. Both views of natural selection claim to (...) capture the core of Darwin’s arguments in On the Origin of Species. Today, historians largely concur with the MS’s reading of Origin as a book aimed to prove natural selection the cause (vera causa) of adaptive change. This paper finds the evidence for that conclusion wanting. I undertake to examine the context and meaning of all Darwin’s known uses of the phrase vera causa, documenting in particular Darwin’s resistance to the pressure to prove natural selection a vera causa in letters written early in 1860. His resistance underlines the logical dependence of natural selection, an unobservable phenomenon, on the causal processes producing the observable events captured by the laws of inheritance, variation, and the struggle for existence, established in Chapters 1–3 of Origin. (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)

This review essay reflects on recent discussions in evolutionary biology and philosophy of science on the central causes of evolution and the structure of causal explanations in evolutionary theory. In this debate, it has been argued that our view of evolutionary causation should be rethought by including more seriously developmental causes and causes of the individual acting organism. I use Tobias Uller’s and Kevin Laland’s volume Evolutionary Causation as well as recent reviews of it as a starting point to reflect (...) on the causal role of agency, individuality, and the environment in evolution. In addition, I critically discuss classical philosophical frameworks of theory change used in this debate to understand changing views of evolutionary causation. (shrink)

According to the modern synthesis, evolution is the gradual change of gene frequencies in a population. The MS is closely allied to adaptationist explanations of phenotypes, where organismic form and behavior is treated as previously selected for and owes its genesis to some remote past. However, some new theories of evolution broadly aligned with the extended evolutionary synthesis, in particular developmental plasticity theory and niche construction theory, foreground the fact that evolution is sometimes much more rapid than previously imagined, and (...) occurs through the active engagement of organisms accommodating and modifying their environments. This article describes how these contemporary theories reveal two interconnected sides of being an adaptive subject, a situated agent that modifies itself and its environment as it lives, and contributes to evolution in turn. MS and adaptationism have a generic logical structure that can be taught anywhere, but because developmental plasticity theory and niche construction theory point to an ontology that foregrounds the agency of the organism, they benefit from in situ exploration. I argue biology as a subject needs to adapt, and call for the renewed importance of field studies, outlining some elements of how such studies might be conceived. I close by considering how understanding organisms as adaptive subjects of evolution has important implications for sustainability education. (shrink)

Research on animal behavior is typically organized according to a combination of two influential frameworks: Ernst Mayr’s distinction between proximate and ultimate causes, and Niko Tinbergen’s “four questions”. My aim is to debunk two common interpretive misconceptions about Mayr’s proximate–ultimate distinction and its relationship to Tinbergen’s four questions, and to offer a new interpretation that avoids both. The first misconception is that the proximate–ultimate distinction maps cleanly onto Tinbergen’s four questions, marking a boundary between Tinbergen’s evolutionary and survival value questions (...) versus developmental and mechanistic questions. The second is that Mayr’s proximate–ultimate distinction is meant to rule out the relevance of proximate causes to evolutionary explanations. I argue that neither is plausible given the text and Mayr’s philosophical aims, namely, to argue that evolutionary biology cannot be reduced to either the physical sciences or to other areas of biology. Through a reconstruction of Mayr’s anti-reductionist argument, I develop an interpretation according to which the proximate–ultimate distinction marks two ways that teleological reasoning can be naturalistically grounded in biology, corresponding to Mayr’s distinction between teleonomic and adapted systems. Mayr distinguishes reduction, which the proximate–ultimate distinction is meant to block, from analysis, through which he allows that proximate causes, causes that are neither proximate nor ultimate, and chance can all contribute to evolutionary explanations. I conclude by suggesting some ways in which the interpretation defended here reframes our understanding of Mayr’s disagreements with some evolutionary-developmental biologists. (shrink)

Niche construction theory concerns how organisms can change selection pressures by altering the feature–factor relationship between themselves and their environment. These alterations are standardly understood to be brought about through two kinds of organism–environment interaction: perturbative and relocational niche construction. We argue that a reconceptualization is needed on the grounds that if a niche is understood as the feature–factor relationship, then there are three fundamental ways in which organisms can engage in niche construction: constitutive, relational, and external niche construction. We (...) further motivate our reconceptualization by showing some examples of organismic activities that fall outside of the current categorization of niche construction, but nonetheless should be included. We end by discussing two objections to niche construction and show how our reconceptualization helps to undercut these objections. 1Introduction2Niche Construction Theory 2.1The standard account of niche construction2.2Two problems with the standard account of niche construction3Three Kinds of Niche Construction 3.1Constitutive niche construction3.2Relational niche construction3.3External niche construction4Conceptual Improvement on Niche Construction Theory 4.1Constitutive niche construction4.2Relational niche construction4.3External niche construction5The Status of Niche Construction Theory within Evolutionary Theory 5.1Is niche construction a helpful addition to evolutionary theory?5.2Is niche construction an evolutionary process?6Conclusion. (shrink)

Ernst Mayr’s concept of dual causality in biology with the two forms of causes continues to provide an essential foundation for the philosophy of biology. They are equivalent to functional and evolutionary causes with both required for full biological explanations. The natural sciences can be classified into nomological, historical nomological and historical dual causality, the last including only biology. Because evolutionary causality is unique to biology and must be included for all complete biological explanations, biology is autonomous from the physical (...) sciences. (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)

In The Secret of Our Success, Joseph Henrich claims that human beings are unique—different from all other animals—because we engage in cumulative cultural evolution. It is the technological and social products of cumulative cultural evolution, not the intrinsic rationality or ‘smartness’ of individual humans, that enable us to live in a huge range of different habitats, and to dominate most of the creatures who share those habitats with us. We are sympathetic to this general view, the latest expression of the (...) ‘California school’s’ view of cultural evolution, and impressed by the lively and interesting way that Henrich handles evidence from anthropology, economics, and many fields of biology. However, because we think it is time for cultural evolutionists to get down to details, this essay review raises questions about Henrich’s analysis of both the cognitive processes and the selection processes that contribute to cumulative cultural evolution. In the former case, we argue that cultural evolutionists need to make more extensive use of cognitive science, and to consider the evidence that mechanisms of cultural learning are products as well as processes of cultural evolution. In the latter case, we ask whether the California school is really serious about selection, or whether it is offering a merely ‘kinetic’ view of cultural evolution, and, assuming the former, outline four potential models of cultural selection that it would be helpful to distinguish more clearly. (shrink)

Mayr’s proximate–ultimate distinction has received renewed interest in recent years. Here we discuss its role in arguments about the relevance of developmental to evolutionary biology. We show that two recent critiques of the proximate–ultimate distinction fail to explain why developmental processes in particular should be of interest to evolutionary biologists. We trace these failures to a common problem: both critiques take the proximate–ultimate distinction to neglect specific causal interactions in nature. We argue that this is implausible, and that the distinction (...) should instead be understood in the context of explanatory abstractions in complete causal models of evolutionary change. Once the debate is reframed in this way, the proximate–ultimate distinction’s role in arguments against the theoretical significance of evo-devo is seen to rely on a generally implicit premise: that the variation produced by development is abundant, small and undirected. We show that a “lean version” of the proximate–ultimate distinction can be maintained even when this isotropy assumption does not hold. Finally, we connect these considerations to biological practice. We show that the investigation of developmental constraints in evolutionary transitions has long relied on a methodology which foregrounds the explanatory role of developmental processes. It is, however, entirely compatible with the lean version of the proximate–ultimate distinction. (shrink)

The scientific investigation of music requires contributions from a diverse array of disciplines. Given the diverse methodologies, interests and research targets of the disciplines involved, we argue that there is a plurality of legitimate research questions about music, necessitating a focus on integration. In light of this we recommend a pluralistic conception of music—that there is no unitary definition divorced from some discipline, research question or context. This has important implications for how the scientific study of music ought to proceed: (...) we show that some definitions are complementary, that is, they reflect different research interests and ought to be retained and, where possible, integrated, while others are antagonistic, they represent real empirical disagreement about music’s nature and how to account for it. We illustrate this in discussion of two related issues: questions about the evolutionary function of music, and questions of the innateness of music. These debates have been, in light of pluralism, misconceived. We suggest that, in both cases, scientists ought to proceed by constructing integrated models which take into account the dynamic interaction between different aspects of music. (shrink)

Günter Wagner’s Homology, Genes, and Evolutionary Innovation is a compelling, and empirically well-supported account of the evolution of character identity and character origination which emphasizes the importance of homology and novelty as central explananda for 21st century evolutionary biology. In this essay review, I focus on the similarities and differences between the structuralist picture of evolutionary biology advocated by Wagner, and that presented by standard evolutionary theory. First, I outline the ways in which Wagner’s genetic theory of homology diverges from (...) the account of homology offered by standard evolutionary theory. Then, I consider the motivations for these divergences. Lastly, I discuss a number of concerns with Wagner’s view, and offer some concluding thoughts on the relationship between structuralism and adaptationism. (shrink)

Thought experiments de facto play many different roles in biology: economical, ethical, technical and so forth. This paper, however, is interested in whether there are any distinctive features of biological TEs as such. The question may be settled in the affirmative because TEs in biology have a function that is intimately connected with the epistemological and methodological status of biology. Peculiar to TEs in biology is the fact that the reflexive, typically human concept of finality may be profitably employed to (...) discover mechanical-experimental causal relations in all living beings—with the obvious caveat that we do not hypostatise and interpret this concept as an ontological quality, since this would land one in an implicitly animistic, pre-Galilean view of nature. From a methodical point of view, the concept of finality is an essential assumption as well as a powerful heuristic tool in the practice of biology, that is, in the investigation of living beings in an intersubjectively testable and reproducible way. (shrink)

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

Like Laland et al., I think Mayr’s distinction is problematic, but I identify a further problem with it. I argue that Mayr’s distinction is a false dichotomy, and obscures an important question about evolutionary change. I show how this question, once revealed, sheds light on some debates in evo-devo that Laland et al.’s analysis cannot, and suggest that it provides a different view about how future integration between biological disciplines might proceed.

Laland and colleagues have sought to challenge the proximate–ultimate distinction claiming that it imposes a unidirectional model of causation, is limited in its capacity to account for complex biological phenomena, and hinders progress in biology. In this article the core of their argument is critically analyzed. It is claimed that contrary to their claims Laland et al. rely upon the proximate–ultimate distinction to make their points and that their alternative conception of reciprocal causation refers to phenomena that were already accounted (...) for by standard theory. (shrink)

Natural selection is the populational process whereby, for instance, the relative number of a variant better suited to a given environment’s attributes increases over generations. In other words, a population’s makeup is altered, over generations, to suit the requirements of a particular environment. Niche construction is the process whereby an environment’s attributes can be stably modified by organisms, over generations, to suit requirements of those organisms. Should the latter process, when it occurs, be considered as significant for the complementary fit (...) between organism and environment as the former? According to mainstream evolutionary theory, random genetic mutation is the only source of novel, unlearned, trans-generationally persistent behavior. A growing number of biologists and philosophers of science, however, maintain that organisms are capable of generating such behavior, in response to, for instance, trans-generationally persistent environmental change, by way of phenotype-mediated, developmentally plastic mechanisms that do not require mutation. Should mechanisms of this sort be more generally recognized as the source of the selectable variation that selection operates on? Niche construction theory answers “yes” to both these questions, and the result is a debate over the comprehensiveness of mainstream evolutionary theory. (shrink)

The notion of biological function is fraught with difficulties—intrinsically and irremediably so, we argue. The physiological practice of functional ascription originates from a time when organisms were thought to be designed and remained largely unchanged since. In a secularized worldview, this creates a paradox which accounts of functions as selected effect attempt to resolve. This attempt, we argue, misses its target in physiology and it brings problems of its own. Instead, we propose that a better solution to the conundrum of (...) biological functions is to abandon the notion altogether, a prospect not only less daunting than it appears, but arguably the natural continuation of the naturalisation of biology. (shrink)

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

The term ‘information’ is used extensively in biology, cognitive science and the philosophy of consciousness in relation to the concepts of ‘meaning’ and ‘causation’. While ‘information’ is a term that serves a useful purpose in specific disciplines, there is much to the concept that is problematic. Part 1 is a critique of the stance that information is an independently existing entity. On this view, and in biological contexts, systems transmit, acquire, assimilate, decode and manipulate it, and in so doing, generate (...) meaning. I provide a detailed proposal in Part 2 that supports the claim that it is the dynamic form of a system that qualifies the informational nature of meaningful interactive engagement, that is, that information is dependent on dynamic form rather than that it exists independently. In Part 3, I reflect on the importance of the distinction between the independent and dependent stances by looking specifically at the implications for how we might better interpret causation and emergence. (shrink)

Research on animal behavior is typically organized according to a combination of two influential frameworks: Ernst Mayr’s distinction between proximate and ultimate causes, and Niko Tinbergen’s “four questions”. My aim is to debunk two common interpretive misconceptions about Mayr’s proximate–ultimate distinction and its relationship to Tinbergen’s four questions, and to offer a new interpretation that avoids both. The first misconception is that the proximate–ultimate distinction maps cleanly onto Tinbergen’s four questions, marking a boundary between Tinbergen’s evolutionary and survival value questions (...) versus developmental and mechanistic questions. The second is that Mayr’s proximate–ultimate distinction is meant to rule out the relevance of proximate causes to evolutionary explanations. I argue that neither is plausible given the text and Mayr’s philosophical aims, namely, to argue that evolutionary biology cannot be reduced to either the physical sciences or to other areas of biology. Through a reconstruction of Mayr’s anti-reductionist argument, I develop an interpretation according to which the proximate–ultimate distinction marks two ways that teleological reasoning can be naturalistically grounded in biology, corresponding to Mayr’s distinction between teleonomic and adapted systems. Mayr distinguishes reduction, which the proximate–ultimate distinction is meant to block, from analysis, through which he allows that proximate causes, causes that are neither proximate nor ultimate, and chance can all contribute to evolutionary explanations. I conclude by suggesting some ways in which the interpretation defended here reframes our understanding of Mayr’s disagreements with some evolutionary-developmental biologists. (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)

Despite the productivity of basic cancer research, cancer continues to be a health burden to society because this research has not yielded corresponding clinical applications. Many proposed solutions to this dilemma have revolved around implementing organizational and policy changes related to cancer research. Here I argue for a different solution: a new conceptualization of causation in cancer. Neither the standard molecular biomarker approaches nor evolutionary biology approaches to cancer fully capture its complex causal dynamics, even when considered jointly. These approaches (...) map on to Ernst Mayr’s proximate–ultimate distinction, which is an inadequate conceptualization of causation in biological systems and makes it difficult to connect developmental and evolutionary viewpoints. I propose looking to evolutionary developmental biology to overcome the distinction and integrate the proximate and ultimate causal frameworks. I use the concepts of modularity and evolvability to show how an EvoDevo perspective can be manifested in cancer translational research. This perspective on causation in cancer is better suited for integrating the complexity of current empirical results and can facilitate novel developments in the investigation and clinical treatment of cancer. (shrink)

The questions and methods of molecular biology and evolutionary biology are clearly distinct, yet a unified approach can lead to deep insights. Unfortunately, attempts to unify these approaches are fraught with pitfalls. In this informal series of questions and answers, we offer the mechanistically oriented biologist a set of steps to come up with evolutionarily reasonable and meaningful hypotheses. We emphasize the critical power and importance of carefully constructed null hypotheses, and we illustrate our ideas with examples representing a range (...) of topics, from the biology of aging, to protein structure, to speciation, and more. We also stress the importance of mathematics as the lingua franca for biologists of all stripes, and encourage mechanistic biologists to seek out quantitative collaborators to build explicit mathematical models, making their assumptions explicit, and their logic clear and testable. Biologists in all realms of inquiry stand to gain from strong bridges between our disciplines. (shrink)

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

The so-called Evolutionary Debunking Arguments are arguments that appeal to the evolutionary genealogy of our beliefs to undermine their justification. When applied to morality, such arguments are intended to undermine moral realism. In this paper I will discuss Andreas Mogensen’s recent effort to secure moral realism against EDAs. Mogensen attempts to undermine the challenge provided by EDAs in metaethics through the distinction between proximate and ultimate causes in biology. The problem with this move is that the proximate/ultimate distinction is misconceived. (...) If ultimate and proximate causes are properly understood to be complementary, such distinction cannot affect EDAs in metaethics. Therefore, I will argue, Mogensen’s argument fails and moral realism is still in danger. (shrink)

Biologists frequently draw on ideas and terminology from engineering. Evolutionary systems biology—with its circuits, switches, and signal processing—is no exception. In parallel with the frequent links drawn between biology and engineering, there is ongoing criticism against this cross-fertilization, using the argument that over-simplistic metaphors from engineering are likely to mislead us as engineering is fundamentally different from biology. In this article, we clarify and reconfigure the link between biology and engineering, presenting it in a more favorable light. We do so (...) by, first, arguing that critics operate with a narrow and incorrect notion of how engineering actually works, and of what the reliance on ideas from engineering entails. Second, we diagnose and diffuse one significant source of concern about appeals to engineering, namely that they are inherently and problematically metaphorical. We suggest that there is plenty of fertile ground left for a continued, healthy relationship between engineering and biology. (shrink)

Mayr’s distinction between proximate and ultimate explanation is justly famous, marking out a division of explanatory labor in biology. But while it is a useful heuristic in many cases, there are others in which proximate factors play an important role in shaping evolutionary trajectories, and in such cases, each project is sensitive to, and relevant to, the other. This general methodological claim is developed in the context of a discussion of human cooperation, and in particular, in a discussion on the (...) puzzling stability of the social contract over the Pleistocene–Holocene transition. For I argue that while we have a plausible account of the stability of Pleistocene cooperation, the stabilizing factors of the Pleistocene disappear in the Holocene, but cooperation does not. Holocene humans solved many collective action problems; cooperation did not collapse despite the apparent growth of free-riding elites. So the article combines a methodological claim about the interaction of proximate and evolutionary biology with a substantive one about the ecology of human cooperation. (shrink)

The validity and utility of the proximate-ultimate distinction in biology have recently been under debate. Opponents of the distinction argue that it rules out individual-level organismic processes from evolutionary explanations, thereby leading to an unfounded separation between organismic causation and evolutionary causation. Proponents of the proximate-ultimate distinction, on the other hand, argue that it serves an important epistemological role in forming different kinds of explanation-seeking questions in biology. In this paper we offer an interpretation the proximate-ultimate distinction not only as (...) a means of forming explanation-seeking questions, but also as a distinction that can help highlight the way in which individual-level organismic processes can be evolutionary causes. We do this by interpreting the distinction between proximate and ultimate causes as a distinction between structuring and triggering causes. (shrink)

Tinbergen’s classic “On Aims and Methods of Ethology” (_Zeitschrift für Tierpsychologie, 20_, 1963) proposed four levels of explanation of behavior, which he thought would soon apply to humans. This paper discusses the need for multilevel explanation; Huxley and Mayr’s prior models, and others that followed; Tinbergen’s differences with Lorenz on “the innate”; and Mayr’s ultimate/proximate distinction. It synthesizes these approaches with nine levels of explanation in three categories: phylogeny, natural selection, and genomics (ultimate causes); maturation, sensitive period effects, and routine (...) environmental effects (intermediate causes); and hormonal/metabolic processes, neural circuitry, and eliciting stimuli (proximate causes), as a respectful extension of Tinbergen’s levels. The proposed classification supports and builds on Tinbergen’s multilevel model and Mayr’s ultimate/proximate continuum, adding intermediate causes in accord with Tinbergen’s emphasis on ontogeny. It requires no modification of Standard Evolutionary Theory or The Modern Synthesis, but shows that much that critics claim was missing was in fact part of Neo-Darwinian theory (so named by J. Mark Baldwin in _The American Naturalist_ in 1896) all along, notably reciprocal causation in ontogeny, niche construction, cultural evolution, and multilevel selection. Updates of classical examples in ethology are offered at each of the nine levels, including the neuroethological and genomic findings Tinbergen foresaw. Finally, human examples are supplied at each level, fulfilling his hope of human applications as part of the biology of behavior. This broad ethological framework empowers us to explain human behavior—eventually completely—and vindicates the idea of human nature, and of humans as a part of nature. (shrink)

Pluralism is widely appealed to in many areas of philosophy of science, though what is meant by ‘pluralism’ may profoundly vary. Because explanations of behaviour have been a favoured target for pluralistic theses, the sciences of behaviour offer a rich context in which to further investigate pluralism. This is what the topical collection The Biology of Behaviour: Explanatory pluralism across the life sciences is about. In the present introduction, we briefly review major strands of pluralist theses and their motivations. We (...) highlight three distinct types of pluralisms—type pluralism, fragmentation pluralism and insular pluralism—and introduce the articles of the topical collection. (shrink)

Physical cognition involves a host of cognitive abilities that enable understanding and manipulation of the physical world. Corvids, the bird family that includes crows, ravens and jays, are renowned for their cognitive abilities, but still little is known about their folk physics. This thesis explores the origins of physical cognition in corvids by investigating its mechanisms, development,fitness value and phylogeny in a wide context that includes theoretical and empirical studies.String pulling is a valuable paradigm for addressing these questions. Many animals (...) can pull a string with food attached to its end, but uncovering the cognitive abilities involved in this behaviour requires further testing. Paper I reviews the string-pulling paradigm, which is one of the oldest tests of animal cognition.It is a highly suitable test for species comparisons, socio-ecological correlations, and phylogenetic questions. Paper II tests several corvids, apes and peafowl on a string-pulling task where the first pulls do not result in the food moving closer. Despite the absence of such visual feedback, most subjects pulled strings in completely, although corvids appeared to choose randomly.Tool use is the archetypical example of physical cognition. Many corvids are remarkably adept at using tools in experimental settings. Paper III reviews animal tool use in general. It is now clear that customary tool-using species do not necessarily outperform their non-tool-using relatives on tests of physical cognition. For corvids that frequently use tools in the wild, such as New Caledonian crows and‘Alalā, tool use appears to have a significant fitness value and may have resulted in morphological adaptations. Paper IV describes a novel tool-use mode in New Caledonian crows. They inserted sticks into objects and then moved away,thereby transporting both. One crow could not grasp the target object, which suggests that such insert-and-transport tool use facilitates control over unwieldy objects. Paper V briefly argues that some corvids have shown the ability to make novel causal interventions, although this question should be addressed in a clearer theoretical framework that makes testable predictions.Sensorimotor cognition is a set of fundamental cognitive abilities that enables the integration of sensory and motor information into practical behaviour. It underlies much of corvid physical cognition. Paper VI investigates the development of sensorimotor cognition in ravens. Their skills developed rapidly and exceeded those of some mammals. They reached the same final sensorimotor stage as great apes, albeit at a markedly accelerated rate. The propensity of corvids to cache objects was investigated in Paper VII. Ravens, which often cache food,frequently cached objects, but surprisingly at similar rates as non-food-caching jackdaws. New Caledonian crows mostly cached objects that resembled functional tools.Anthropomorphism is the attribution of human traits to animals. Fear of mistakenly anthropomorphising animals has resulted in biased principles and an uneven burden of proof, which may hinder scientific progress more than it is supposed to offer protection against making mistakes. Cognitive zoology should not be misguided by overcompensating for such potential pitfalls. (shrink)

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