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)

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)

Explanation in terms of gene regulatory networks has become standard practice in evolutionary developmental biology. In this paper, we argue that GRNs fail to provide a robust, mechanistic, and dynamic understanding of the developmental processes underlying the genotype–phenotype map. Explanations based on GRNs are limited by three main problems: the problem of genetic determinism, the problem of correspondence between network structure and function, and the problem of diachronicity, as in the unfolding of causal interactions over time. Overcoming these problems requires (...) dynamic mechanistic explanations, which rely not only on mechanistic decomposition, but also on dynamic modeling to reconstitute the causal chain of events underlying the process of development. We illustrate the power and potential of this type of explanation with a number of biological case studies that integrate empirical investigations with mathematical modeling and analysis. We conclude with general considerations on the relation between mechanism and process in evo-devo. (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)

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)

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)

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)

Both biosemiotics and evolutionary epistemology are concerned with how knowledge evolves. (Applied) Evolutionary Epistemology thereby focuses on identifying the units, levels, and mechanisms or processes that underlie the evolutionary development of knowing and knowledge, while biosemiotics places emphasis on the study of how signs underlie the development of meaning. We compare the two schools of thought and analyze how in delineating their research program, biosemiotics runs into several problems that are overcome by evolutionary epistemologists. For one, by emphasizing signs, biosemiotics (...) needs to delineate a semiotic threshold, which is a problem not encountered by evolutionary epistemologists. Instead, the latter recognizes that all organisms are knowers that evolve knowledge, which they recognize to extend toward phenomena produced by organisms such as behavior, cognition, language, culture, science, and technology. Secondly, biosemiotics attempts at continuing adaptationist notions on how organisms relate to their environment, while especially Applied Evolutionary Epistemology comes to redefine the nature of the organism–environment relationship in such a way that it recognizes the spatiotemporal boundedness of existence, which in turn makes adaptationist accounts obsolete. (shrink)

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