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)

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)

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

A notion of delegated causality is introduced here. This subtle kind of causality is dual to interventional causality. Delegated causality elucidates the causal role of dynamical systems at the “edge of chaos”, explicates evident cases of downward causation, and relates emergent phenomena to Gödel’s incompleteness theorem. Apparently rich implications are noticed in biology and Chinese philosophy. The perspective of delegated causality supports cognitive interpretations of self-organization and evolution.

Many people who claim that evolution and theism are in tension assume that God, being omnipotent, could create life in different ways. For instance, Paul Draper has argued that the fact that life evolved on earth supports naturalism over theism. However, for there to be a probabilistic tension between naturalism and theism, because of the fact of evolution, a certain background assumption must be true, namely, that God could have made biological organisms and species through an act of Genesis-style special (...) creation, or some combination of evolution and special creation. But that background assumption is potentially faulty. Why? Because special creation presupposes that biological organisms and species are spatiotemporally unrestricted entities, and if the dominant paradigm in biology, and philosophy of biology, on the ontological status of organisms and species is correct, then they are spatiotemporally restricted entities. (shrink)

The causal structure of Darwinian evolution by natural selection is investigated. Its basic scheme is reproduction resulting from a feedback loop driven by internal and external causes. Causation internal to the loop connects genotype, development, phenotype, and fitness, with environmental constraints on the latter preventing runaway reproduction. External causes driving the core loop are environmental change and genetic change. This basic causal structure is complicated by modern additions such as control of mutation rate, niche construction, interactions between evolution and development, (...) and epigenetic effects. In this article it is argued that at least one of these additions, control of mutation rate, produces an additional feedback loop incorporating a continual, targeted modulation of stochasticity, and thereby produces an unusual form of causation. This form of causation entangles deterministic and stochastic causation in such a way that it imposes, in effect, an active, goal-driven role on the organisms in a line of descent. The goal is defined by the factor driving the stochasticity, which must be regarded as an intrinsic estimate made implicitly by the organism of its own fitness. The estimator itself is an evolved property of the organism, and is aligned with the apparent goal-directedness of natural selection. (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)

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)

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 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)

In this paper we argue that ecological evolutionary developmental biology accounts of cognitive modernity are compatible with cultural evolution theories of language built upon iterated learning models. Cultural evolution models show that the emergence of near universal properties of language do not require the preexistence of strong specific constraints. Instead, the development of general abilities, unrelated to informational specificity, like the copying of complex signals and sharing of communicative intentions is required for cultural evolution to yield specific properties, such as (...) language structure. We argue that eco-evo-devo provides the appropriate conceptual background to ground an account for the many interconnected genetic, environmental and developmental factors that facilitated the emergence of an organic system able to develop language through the iterated transmission of information. We use the concept of niche construction to connect evolutionary developmental accounts for sensory guided motor capacities and cultural evolution guided by iterated learning models. This integrated theoretical model aims to build bridges between biological and cultural approaches. (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)

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)

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 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)

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)

Evolutionary systems biology is an emerging hybrid approach that integrates methods, models, and data from evolutionary and systems biology. Drawing on themes that arose at a cross-disciplinary meeting on ESB in 2013, we discuss in detail some of the explanatory friction that arises in the interaction between evolutionary and systems biology. These tensions appear because of different modeling approaches, diverse explanatory aims and strategies, and divergent views about the scope of the evolutionary synthesis. We locate these discussions in the context (...) of long-running philosophical deliberations on explanation, modeling, and theoretical synthesis. We show how many of the issues central to ESB’s progress can be understood as general philosophical problems. The benefits of addressing these philosophical issues feed back into philosophy too, because ESB provides excellent examples of scientific practice for the development of philosophy of science and philosophy of biology. (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)

Climate change has historically been an evolutionary determinant for our species, affecting both hominin evolutionary innovations and extinction rates, and the early waves of migration and expansion outside Africa. Today Homo sapiens has turned itself into a major geological force, able to cause a biodiversity crisis comparable to previous mass extinction events, shaping the Earth surface and impacting biogeochemical cycles and the climate at a global level. We argue that anthropogenically-driven climate change must be understood in terms of a monumental (...) niche construction process, generating long-term ecological inheritance and eco-evolutionary feedbacks that are putting our health and well-being and those of future generations at risk. We then list five major sources of climate change counter-intuitiveness, highlighting how evolved cognitive biases and heuristics may stand in the way of providing effective responses within tight deadlines. Drawing on our framing of the climate breakdown, we finally call for an evolutionary perspective in approaching the adaptive challenge posed by climate change: we argue that putting the brakes on a genuine self-endangering evolutionary trap ultimately depends on our counteractive niche constructing abilities, played at the level of our institutional and innovation capacity. (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)

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)

We are grateful to the commentators for taking the time to respond to our article. Too many interesting and important points have been raised for us to tackle them all in this response, and so in the below we have sought to draw out the major themes. These include problems with both the term ‘ultimate causation’ and the proximate-ultimate causation dichotomy more generally, clarification of the meaning of reciprocal causation, discussion of issues related to the nature of development and phenotypic (...) plasticity and their roles in evolution, and consideration of the need for an extended evolutionary synthesis. (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)

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)

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.

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)

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)

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)

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)

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)

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)

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)

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)

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.

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)

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)

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)

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)

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)

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)

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)

Ecological developmental biology considers the phenotype as actively produced through an environmentally informed process of individual development, rather than predetermined by the genotype. Accordingly, the genotype is viewed as one among many interactants that contribute formative elements; it is understood to do so no differently from the way other organism-internal and environmental resources do. Although the EcoDevo approach is evidently particularly apt to inform approaches to human development, which mostly takes shape in rich cultural environments, it is remarkable that, at (...) least within some highly influential circles of the linguistic sciences, the study of many distinctive human cognitive traits including language remains strongly anchored to a preformationist stance. This article brings an EcoDevo approach to the biologically focused study of language to argue that an aspect of language design commonly assumed to be “blueprinted,” namely, the lexicon-syntax interface, may instead be explained as a plastic effect of ongoing developmental processes in the mind of the child, in which designated aspects of the environment fulfill a key constructive role. (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)

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)