This paper reviews the debate on the notion of biological function and on functional explanation as this takes place in philosophy. It describes the different perspectives, issues, intuitions, theories and arguments that have emerged. The author shows that the debate has been too heavily influenced by the concerns of a naturalistic philosophy of mind and argues that in order to improve our understanding of biology the attention should be shifted from the study of intuitions to the study of the actual (...) practice of biological inquiry. (shrink)

The dominant view in the cognitive science of religion (the ‘Standard Model’) is that religious belief and behaviour are not adaptive traits but rather incidental byproducts of the cognitive architecture of mind. Because evidence for the Standard Model is inconclusive, the case for it depends crucially on its alleged methodological superiority to selectionist alternatives. However, we show that the Standard Model has both methodological and evidential disadvantages when compared with selectionist alternatives. We also consider a pluralistic approach, which holds that (...) religion or various aspects of religion originated as byproducts of evolved cognitive structures but were subsequently co-opted for adaptive purposes. We argue that when properly formulated, the pluralistic approach also has certain advantages over the Standard Model. (shrink)

On dit souvent que, tandis que la biologie de l'évolution utilise un concept étiologique de fonction (la fonction d'un trait biologique n'est autre que son effet sélectionné), la physiologie prend appui sur un autre concept de fonction, celui de rôle causal. Cependant, un examen plus attentif montre que le concept non normatif de rôle causal n'est pas ce dont la physiologie générale ou la neurophysiologie ont besoin. Ces disciplines font un large usage de notions comme celles de bon fonctionnement, de (...) fonctionnement normal, de fonctionnement perturbé ou de trouble fonctionnel. Il semble ainsi qu'un concept normatif comme le concept étiologique n'est pas seulement utile dans le contexte d'explications téléologiques ou évolutionnistes, mais aussi dans celui d'explications mécanistes ou opérationnelles telles qu'elles sont couramment proposées par la physiologie. La différence entre types d'explication n'est pas une différence entre deux concepts de fonction. En particulier, l'analyse physiologique serait une tâche impossible si elle devait prendre en compte toutes les relations causales à l'intérieur de l'organisation fonctionnelle de chaque système biologique. Pour résoudre ce problème, la physiologie doit être une analyse du fonctionnement normal du système, quelle que soit la part d'idéalisation nécessaire. La notion de bon fonctionnement, conçu comme le produit de l'histoire évolutive, nous aide en particulier à comprendre pourquoi les systèmes qui fonctionnent bien se ressemblent, et pourquoi les systèmes qui dysfonctionnent peuvent le faire de tant de façons différentes. It is often claimed that although evolutionary biology uses an etiological concept of function (according to etiological theory, the function of a biological feature is its selected effect), physiology relies on a different concept of function, the causal role concept. However, upon closer scrutiny, it appears that the non-normative causal role concept is not what is needed by physiology or neurophysiology. These disciplines make a wide use of normative notions like proper functioning, normal functioning, impaired functioning, or functional disorders. In this case, it seems that a normative concept like the etiological concept is not only useful in the context of teleological or evolutionary explanations, but also in the context of mechanical or operational explanations as they are routinely offered by physiology. The difference between types of explanations is not a difference between two kinds of concepts of function. In particular, physiological analysis would be an impossible task if it had to take into account all causal links within the functional organization of each biological system. To solve this problem, physiology has to be the analysis of the proper or normal functioning of the system, even if normal functioning is usually obtained through a process of idealization. The notion of proper functioning, conceived as the product of evolutionary history, helps us understand, among other things, why properly functioning systems tend to be similar, and why malfunctioning systems may differ in so many different ways. (shrink)

Both Aristotle and Ernst Mayr present theories of dual explanation in biology, with proximal, clearly physical explanations and more distal, biology-specific explanations. Aristotle’s presentation of final cause explanations in Posterior Analytics relates final causes to the necessary material, formal, and efficient causes that mediate them. Johnson and Leunissen demonstrate the problematic nature of historical and recent interpretations and open the door for a new interpretation consistent with modern evolutionary theory. Mayr’s differentiation of proximate and ultimate/evolutionary causes provides a key to (...) appropriating Aristotle for modern use, if care is taken to differentiate Mayr’s proximate/distal and mechanical/evolutionary distinctions from his intentional/non-intentional distinction. Building on Mayr, and noting his appropriation of ancient and recent commentary, a strong case can be made for nested explanation, wherein evolutionary explanations are instantiated in systems of mechanical explanations. Biological concepts of organism, gene, function, etc., are etiologically prior to mechanical explanations of function, but temporally posterior to mechanical explanations of historical adaptation. Such an analysis sheds light on recent arguments within evolutionary biology while highlighting the importance of epistemology in contemporary science. (shrink)

A theory may be invalid, not owing to erroneous observations or the invocation of an inappropriate law, but because of the use of equivocal terms. This is demonstrated for Darwin's failed model of sympatric speciation through the principle of divergence.

In 1961, Ernst Mayr published a highly influential article on the nature of causation in biology, in which he distinguished between proximate and ultimate causes. Mayr argued that proximate causes (e.g. physiological factors) and ultimate causes (e.g. natural selection) addressed distinct ‘how’ and ‘why’ questions and were not competing alternatives. That distinction retains explanatory value today. However, the adoption of Mayr’s heuristic led to the widespread belief that ontogenetic processes are irrelevant to evolutionary questions, a belief that has (1) hindered (...) progress within evolutionary biology, (2) forged divisions between evolutionary biology and adjacent disciplines and (3) obstructed several contemporary debates in biology. Here we expand on our earlier (Laland et al. in Science 334:1512–1516, 2011) argument that Mayr’s dichotomous formulation has now run its useful course, and that evolutionary biology would be better served by a concept of reciprocal causation, in which causation is perceived to cycle through biological systems recursively. We further suggest that a newer evolutionary synthesis is unlikely to emerge without this change in thinking about causation. (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)

Within evolutionary biology, life-history theory is used to explain cross-species differences in allocation strategies regarding reproduction, maturation, and survival. Behavioral scientists have recently begun to conceptualize such strategies as a within-species individual characteristic that is predictive of behavior. Although life history theory provides an important framework for behavioral scientists, the psychometric approach to life-history strategy measurement—as operationalized by K-factors—involves conceptual entanglements. We argue that current psychometric approaches attempting to identify K-factors are based on an unwarranted conflation of functional descriptions and (...) proximate mechanisms—a conceptual mix-up that may generate unviable hypotheses and invites misinterpretation of empirical findings. The assumptions underlying generic psychometric methodology do not allow measurement of functionally defined variables; rather these methods are confined to Mayr’s proximate causal realm. We therefore conclude that K-factor scales lack validity, and that life history strategy cannot be identified with psychometrics as usual. To align theory with methodology, suggestions for alternative methods and new avenues are proposed. (shrink)

Processes produce changes: rivers erode their banks and thunderstorms cause floods. If I am right that organisms are a kind of process, then the causally efficacious behaviours of organisms are also examples of processes producing change. In this paper I shall try to articulate a view of how we should think of causation within a broadly processual ontology of the living world. Specifically, I shall argue that causation, at least in a central class of cases, is the interaction of processes, (...) that such causation is the exercise of a capacity inherent in that process and, negatively, that causation should not be understood as the instantiation of universal laws. The approach I describe has substantial similarities with the process causality articulated by Wesley Salmon and Phil Dowe for physical causation, making it plausible that the basic approach can be applied equally to the non-living world. It is an approach that builds at crucial points on the criticisms of determinism and universal causality famously articulated by Elizabeth Anscombe. (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)

Las presiones selectivas no son factores ambientales que existan independientemente de los linajes por ellas afectados. Su existencia y su configuración siempre dependen de las alternativas de evolución que el propio linaje genera y acepta; y dicha configuración, necesariamente, se va alterando conforme el propio linaje evoluciona. Aclarar eso es muy importante para desestimar algunas reticencias generadas por la dicotomía próximo-remoto. Esta polaridad continúa siendo un recurso conceptual válido e insustituible para entender la especificidad de la Biología Evolucionaria. Enmendar algunos (...) equívocos generados por Mayr en su caracterización del concepto de causa remota es una alternativa mejor que la de pensar en abandonar o en relativizar esa distinción. (shrink)

Ernst Mayr''s scientific career continues strongly 70 years after he published his first scientific paper in 1923. He is primarily a naturalist and ornithologist which has influenced his basic approach in science and later in philosophy and history of science. Mayr studied at the Natural History Museum in Berlin with Professor E. Stresemann, a leader in the most progressive school of avian systematics of the time. The contracts gained through Stresemann were central to Mayr''s participation in a three year expedition (...) to New Guinea and The Solomons, and the offer of a position in the Department of Ornithology, American Museum of Natural History, beginning in 1931. At the AMNH, Mayr was able to blend the best of the academic traditions of Europe with those of North America in developing a unified research program in biodiversity embracing systematics, biogeography and nomenclature. His tasks at the AMNH were to curate and study the huge collections amassed by the Whitney South Sea Expedition plus the just purchased Rothschild collection of birds. These studies provided Mayr with the empirical foundation essential for his 1942Systematics and the Origin of Species and his subsequent theoretical work in evolutionary biology as well as all his later work in the philosophy and history of science. Without a detailed understanding of Mayr''s empirical systematic and biogeographic work, one cannot possibly comprehend fully his immense contributions to evolutionary biology and his later analyses in the philosophy and history of science. (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)

It's been 41 years since the publication of Ernst Mayr's Cause and Effect in Biology wherein Mayr most clearly develops his version of the influential distinction between ultimate and proximate causes in biology. In critically assessing Mayr's essay I uncover false statements and red-herrings about biological explanation. Nevertheless, I argue to uphold an analogue of the ultimate/proximate distinction as it refers to two different kinds of explanations, one dynamical the other statistical.

This dissertation argues that cognition is a kind of natural agency. Natural agency is the capacity that certain systems have to act in accordance with their own norms. Natural agents are systems that bias their repertoires in response to affordances in the pursuit of their goals. Cognition is a special mode of this general phenomenon. Cognitive systems are agents that have the additional capacity to actively take their worlds to be certain ways, regardless of whether the world is really that (...) way. In this way, cognitive systems are desituated. Desituatedness is the root of specifically cognitive capacities for representation and abstraction. There are two main reasons why this view needs defending. First, natural agency is typically viewed as incompatible with natural science because it is committed to a teleological mode of explanation. Second, cognition is typically held to be categorically distinct from natural agency. This dissertation argues against both of these views. It argues against the incompatibility of agency and natural science by demonstrating that systems biology, general systems theory, and sciences that deal with complex systems have typically underappreciated conceptual and theoretical resources for grounding agency in the causal structure of the world. These conceptual resources do not, however, reduce agency to systems theory because the normativity inherent in agency demands descriptive resources beyond those of even the most sophisticated systems theory. It argues against the categorical difference between natural agency and cognition by pointing out that separating cognition from a richer web of situated, ecologically embedded relations between the agent and the world generates the frame problem, which is an insuperable obstacle to making cognition that is sufficiently responsive to the complexity of the world. Rooting cognition in natural agency is a more robust empirical bet for theorizing cognition and artificial intelligence. (shrink)

We have structured our response according to five questions arising from the commentaries: (i) What is sentience? (ii) Is sentience a necessary or sufficient condition for moral standing? (iii) What methods should guide comparative cognitive research in general, and specifically in studying invertebrates? (iv) How should we balance scientific uncertainty and moral risk? (v) What practical strategies can help reduce biases and morally dismissive attitudes toward invertebrates?

As common experience confirms, procrastination seems not only possible, but widespread. However, procrastination should not be taken for granted. Often, the procrastinator harms herself knowingly. It thus clearly seems that such a person lacks the self-concern that usually characterises us. After having spelled out what procrastination is, and having explored its main varieties, I consider the relation between procrastination and risk-taking. After this, I discuss the implications of this phenomenon for the debates about personal identity. The upshot, I argue, is (...) that reductionist accounts, according to which identity is a matter of psychological continuity, are in a better position as might have been thought. (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)