Despite the burgeoning interest in new and more complex accounts of the organism-environment dyad by biologists and philosophers, little attention has been paid in the resulting discussions to the history of these ideas and to their deployment in disciplines outside biology—especially in the social sciences. Even in biology and philosophy, there is a lack of detailed conceptual models of the organism-environment relationship. This volume is designed to fill these lacunae by providing the first multidisciplinary discussion of the topic of organism-environment (...) interaction. It brings together scholars from history, philosophy, psychology, anthropology, medicine, and biology to discuss the common focus of their work: entangled life, or the complex interaction of organisms and environments. (shrink)

There is surprisingly little philosophical work on conceptually spelling out the difference between the traits on which natural selection may be said to act (e.g. “having a high running speed”) and mere circumstantial traits (e.g. “happening to be in the path of a forest fire”). I label this issue the “selectable traits problem” and, in this paper, I propose a solution for it. I first show that, contrary to our first intuition, simply equating selectable traits with heritable ones is not (...) an adequate solution. I then go on to argue that two recent philosophical solutions to this problem—due to Peter Godfrey-Smith and Pierrick Bourrat—are unconvincing because they cannot accommodate frequency-dependent selection. The way out of this difficulty is, I argue, to accept that extrinsic properties dependent on relations between intrinsic properties of the population members should also count as selectable traits. I then show that my proposal is legitimized by more than the simple accommodation of frequency-dependent selection. (shrink)

The general process view of learning, which guided research into learning for the first half of this century, has come under attack in recent years from several quarters. One form of criticism has come from proponents of the so-called biological boundaries approach to learning. These theorists have presented a variety of data showing that supposedly general laws of learning may in fact be limited in their applicability to different species and learning tasks, and they argue that the limitations are drawn (...) by the nature of each species' adaptation to the particular requirements of its natural environment. The biological boundaries approach has served an important critical function in the move away from general process learning theory, but it is limited in its ability to provide an alternative to the general process approach. In particular, the biological boundaries approach lacks generality, it is in some respects subservient to the general process tradition, and its ecological content is in too many cases limited toex post factoadaptive explanations of learning skills. A contrasting, ecological approach to learning, which can provide a true alternative to general process theory, is presented. The ecological approach begins by providing an ecological task description for naturally occurring instances of learning; this step answers the question:Whatdoes this animal learn to do? The next step is an analysis of the means by which learning occurs in the course of development, answering the question:Howdoes the animal learn to do this? On the basis of such analyses, local principles of adaptation are formulated to account for the learning abilities of individual species. More global principles are sought by generalization among these local principles and may form the basis for a general ecological theory of learning. (shrink)

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to (...) grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as “replicators” which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be “vehicles” of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units.The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to “organs” of group-level “organisms.” Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals. (shrink)

In both biology and the human sciences, social groups are sometimes treated as adaptive units whose organization cannot be reduced to individual interactions. This group-level view is opposed by a more individualistic one that treats social organization as a byproduct of self-interest. According to biologists, group-level adaptations can evolve only by a process of natural selection at the group level. Most biologists rejected group selection as an important evolutionary force during the 1960s and 1970s but a positive literature began to (...) grow during the 1970s and is rapidly expanding today. We review this recent literature and its implications for human evolutionary biology. We show that the rejection of group selection was based on a misplaced emphasis on genes as "replicators" which is in fact irrelevant to the question of whether groups can be like individuals in their functional organization. The fundamental question is whether social groups and other higher-level entities can be "vehicles" of selection. When this elementary fact is recognized, group selection emerges as an important force in nature and what seem to be competing theories, such as kin selection and reciprocity, reappear as special cases of group selection. The result is a unified theory of natural selection that operates on a nested hierarchy of units. The vehicle-based theory makes it clear that group selection is an important force to consider in human evolution. Humans can facultatively span the full range from self-interested individuals to "organs" of group-level "organisms." Human behavior not only reflects the balance between levels of selection but it can also alter the balance through the construction of social structures that have the effect of reducing fitness differences within groups, concentrating natural selection (and functional organization) at the group level. These social structures and the cognitive abilities that produce them allow group selection to be important even among large groups of unrelated individuals. (shrink)

The supervenience and multiple realizability of biological properties have been invoked to support a disunified picture of the biological sciences. I argue that supervenience does not capture the relation between fitness and an organism's physical properties. The actual relation is one of causal dependence and is, therefore, amenable to causal explanation. A case from optimality theory is presented and interpreted as a microreductive explanation of fitness difference. Such microreductions can have considerable scope. Implications are discussed for reductive physicalism in evolutionary (...) biology and for the unity of science. (shrink)

Susan Mills and John Beatty proposed a propensity interpretation of fitness (1979) to show that Darwinian explanations are not circular, but they did not address the critics' chief complaint that the principle of the survival of the fittest is either tautological or untestable. I show that the propensity interpretation cannot rescue the principle from the critics' charges. The critics, however, incorrectly assume that there is nothing more to Darwin's theory than the survival of the fittest. While Darwinians all scoff at (...) this assumption, they do not agree about what role, if any, this principle plays in Darwin's theory of natural selection. I argue that the principle has no place in Darwin's theory. His theory does include the idea that some organisms are fitter than others. But greater reproductive success is simply inferred from higher fitness. There is no reason to embody this inference in the form of a special principle of the survival of the fittest. (shrink)

Since the last decades of the twentieth and the beginning of the twenty-first century, the use of metaphysics by philosophers when approaching conceptual problems in biology has increased. Some philosophers call this tendency in philosophy of biology ‘Metaphysics of Biology’. In this paper, I aim at characterizing Metaphysics of Biology by paying attention to the diverse ways philosophers use metaphysics when addressing conceptual problems in biology. I will claim that there are two different modes of doing Metaphysics of Biology, namely (...) Metaphysics for Biology and Metaphysics in Biology. (shrink)

The notion of fitness is usually equated to reproductive success. However, this actualist approach presents some difficulties, mainly the explanatory circularity problem, which have lead philosophers of biology to offer alternative definitions in which fitness and reproductive success are distinguished. In this paper, we argue that none of these alternatives is satisfactory and, inspired by Mumford and Anjum’s dispositional theory of causation, we offer a definition of fitness as a causal dispositional property. We argue that, under this framework, the distinctiveness (...) that biologists usually attribute to fitness—namely, the fact that fitness is something different from both the physical traits of an organism and the number of offspring it leaves—can be explained, and the main problems associated with the concept of fitness can be solved. Firstly, we introduce Mumford and Anjum's dispositional theory of causation and present our definition of fitness as a causal disposition. We explain in detail each of the elements involved in our definition, namely: the relationship between fitness and the functional dispositions that compose it, the emergent character of fitness, and the context-sensitivity of fitness. Finally, we explain how fitness and realized fitness, as well as expected and realized fitness are distinguished in our approach to fitness as a causal disposition. (shrink)

Showing that the arithmetic mean number of offspring for a trait type often fails to be a predictive measure of fitness was a welcome correction to the philosophical literature on fitness. While the higher mathematical moments of a probability-weighted offspring distribution can influence fitness measurement in distinct ways, the geometric mean number of offspring is commonly singled out as the most appropriate measure. For it is well-suited to a compounding process and is sensitive to variance in offspring number. The geometric (...) mean thus proves to be a predictively efficacious measure of fitness in examples featuring discrete generations and within- or between-generation variance in offspring output. Unfortunately, this advance has subsequently led some to conclude that the arithmetic mean is never a good measure of fitness and that the geometric mean should accordingly be the default measure of fitness. We show not only that the arithmetic mean is a perfectly reasonable measure of fitness so long as one is clear about what it refers to, but also that it functions as a more general measure when properly interpreted. It must suffice as a measure of fitness in any case where the geometric mean has been effectively deployed as a measure. We conclude with a discussion about why the mathematical equivalence we highlight cannot be dismissed as merely of mathematical interest. (shrink)

The most consistent definition of fitness makes it a static property of organisms. However, this is not how fitness is used in many evolutionary models. In those models, fitness is permitted to vary with an organism’s circumstances. According to this second conception, fitness is dynamic. There is consequently tension between these two conceptions of fitness. One recently proposed solution suggests resorting to conditional properties. We argue, however, that this solution is unsatisfactory. Using a very simple model, we show that it (...) can lead to incompatible fitness values and indecision about whether selection actually occurs. (shrink)

The propensity nature of evolutionary fitness has long been appreciated and is nowadays amply discussed. The discussion has, however, on occasion followed long standing conflations in the philosophy of probability literature between propensities, probabilities, and frequencies. In this paper, I apply a more recent conception of propensities in modelling practice to some of the key issues, regarding the mathematical representation of fitness and how it may be regarded as explanatory. The ensuing complex nexus of fitness emphasises the distinction between biological (...) propensities and the probability distributions over offspring numbers that they give rise to; and how critical it is to distinguish the possession conditions of the underlying dispositional properties from those of their probabilistic manifestations. (shrink)

The propensity interpretation of fitness draws on the propensity interpretation of probability, but advocates of the former have not attended sufficiently to problems with the latter. The causal power of C to bring about E is not well-represented by the conditional probability Pr. Since the viability fitness of trait T is the conditional probability Pr, the viability fitness of the trait does not represent the degree to which having the trait causally promotes surviving. The same point holds for fertility fitness. (...) This failure of trait fitness to capture causal role can also be seen in the fact that coextensive traits must have the same fitness values even if one of them promotes survival and the other is neutral or deleterious. Although the fitness of a trait does not represent the trait’s causal power to promote survival and reproduction, variation in fitness in a population causally promotes change in trait frequencies; in this sense, fitness variation is a population-level propensity. (shrink)

Karl Popper argued in 1974 that evolutionary theory contains no testable laws and is therefore a metaphysical research program. Four years later, he said that he had changed his mind. Here we seek to understand Popper’s initial position and his subsequent retraction. We argue, contrary to Popper’s own assessment, that he did not change his mind at all about the substance of his original claim. We also explore how Popper’s views have ramifications for contemporary discussion of the nature of laws (...) and the structure of evolutionary theory. (shrink)

An increasing number of biologists are expressing discontent with the prevailing theory of neo-Darwinism. In particular, the tendency of neo-Darwinians to adopt genetic determinism and atomistic notions of both genes and organisms is seen as grossly unfair to the body of developmental theory. One faction of dissenteers, the Process Structuralists, take their inspiration from the rational morphologists who preceded Darwin. These neo-rationalists argue that a mature biology must possess universal laws and that these generative laws should be sought within organismal (...) development. Such a rational biology will only be possible once the neo-Darwinian paradigm, with its reliance on inherently stochastic processes, is overthrown.To facilitate this revolution, process structuralism launches a broad attack on the theoritical adequacy of its opponent. It is charged that neo-Darwinism is untestable and therefore its hypotheses are nothing more than adaptive stories. Further, the lamentable tendencies toward genetic determinism and atomism by modern biologists is seen as the inescapable consequences of adopting the neo-Darwinian outlook. (shrink)

The theory of natural selection is a rich systematization of biological knowledge without a first principle. When formulations of a proposed principle of natural selection are examined carefully, each is seen to be exhaustively analyzable into a proposition about sources of fitness and a proposition about consequences of fitness. But whenever the fitness of an organic variety is well defined in a given biological situation, its sources are local contingencies together with the background of laws from disciplines other than the (...) theory of natural selection; and the consequences of fitness for the long range fate of organic varieties are essentially applications of probability theory. Hence there is no role and no need for a principle of the theory of natural selection, and any generalities that may hold in that theory are derivative rather than fundamental. (shrink)

We look to mitonuclear ecology and the phenomenon of Mother’s Curse to argue that the sex of parents and offspring among populations of eukaryotic organisms, as well as the mitochondrial genome, ought to be taken into account in the conceptualization of evolutionary fitness. Subsequently, we show how characterizations of fitness considered by philosophers that do not take sex and the mitochondrial genome into account may suffer. Last, we reflect on the debate regarding the fundamentality of trait versus organism fitness and (...) gesture at the idea that the former lies at the conceptual basis of evolutionary theory. (shrink)

In several places we have argued that ‘fitness’ is a primitive term with respect to the theory of evolution properly understood. These arguments have relied heavily on the axiomatization of the theory provided by one of us. In contrast, both John Beatty and Robert Brandon have separately argued for a “propensity“ interpretation of “fitness” ; and in Brandon and Beatty they attack our view that “fitness“ is a primitive term in evolutionary theory, concluding that a definition by way of propensities (...) is possible and preferable. Here we reply to their criticisms, and argue that, at most, the view that fitness is a statistical propensity is a terminological variant on our thesis that it is a primitive in evolutionary theory. (shrink)

The propensity account of fitness intends to solve the classical tautologicity issue by identifying fitness with a disposition, the ability to survive and reproduce. As proponents recognized early on, this account requires operational independence from actual reproductive success to avoid circularity and vacuousness charges. They suggested that operational independence is achieved by measuring fitness values through optimality models. Our goal in this article is to develop this suggestion. We show that one plausible procedure by which these independent operationalizations could be (...) thought to take place, and which is in accordance with what is said in the optimality literature, is unsound. We provide two independent lines of reasoning to show this. We then provide a sketch of a more adequate account of the role of optimality models in evolutionary contexts and draw some consequences. (shrink)

Evolutionary psychology purports to explain human capacities as adaptations to an ancestral environment. A complete explanation of human language or human reasoning as adaptations depends on assessing an historical claim, that these capacities evolved under the pressure of natural selection and are prevalent because they provided systematic advantages to our ancestors. An outline of the character of the information needed in order to offer complete adaptation explanations is drawn from Robert Brandon (1990), and explanations offered for the evolution of language (...) and reasoning within evolutionary psychology are evaluated. Pinker and Bloom's (1992) defense of human language as an adaptation for verbal communication, Robert Nozick's (1993) account of the evolutionary origin of rationality, and Cosmides and Tooby's (1992) explanation of human reasoning as an adaptation for social exchange, are discussed in light of what is known, and what is not known, about the history of human evolution. In each case, though a plausible case is made that these capacities are adaptations, there is not enough known to offer even a semblance of an explanation of the origin of these capacities. These explanations of the origin of human thought and language are simply speculations lacking the kind of detailed historical information required for an evolutionary explanation of an adaptation. (shrink)

The mutual influence of science and values in biology is exhibited in several cases from the biological literature. It is argued in a number of cases, from R. A. Fisher's argument for the optimality of a 50:50 sex ratio to A. Jensen's defense of a genetic basis for intelligence, and including work on the evolution of sexual dimorphism and muted aggression, that the credence accorded the views is disproportionate with their theoretical and empirical warrant. It is, furthermore, suggested that the (...) proper explanation for the attraction and persistence of such views lies in their conformity with ideological norms. There is thus an important, if circumscribed, role for ideological critique in the evaluation of scientific theories; in particular, it lies in the explanation of the acceptance and persistence of scientific views, given independent grounds for questioning their justifiability. (shrink)

In a recent issue of Biology and Philosophy, Kenneth Waters argues that the principle of survival of the fittest should be eliminated from the theory of natural selection, because it is an untestable law of probability, and as such, has no place in evolutionary theory. His argument is impressive, but it does not do justice to the practice of biology. The principle of survival of the fittest should not be eliminated from the theory of natural selection because it is important (...) to biological practice: it plays an essential role in explanation and discovery, and in unffying the theory of natural selection. (shrink)

Elliott Sober (1987, 1993) and Orzack and Sober (forthcoming) argue that adaptationism is a very general hypothesis that can be tested by testing various particular hypotheses that invoke natural selection to explain the presence of traits in populations of organisms. In this paper, I challenge Sobers claim that adaptationism is an hypothesis and I argue that it is best viewed as a heuristic (or research strategy). Biologists would still have good reasons for employing this research strategy even if it turns (...) out that natural selection is not the most important cause of evolution. (shrink)

One contentious debate in the philosophy of biology is that between the statisticalists and causalists. The former understand core evolutionary concepts like fitness and selection to be mere statistical summaries of underlying causal processes. In this view, evolutionary changes cannot be causally explained by selection or fitness. The causalist side, on the other hand, holds that populations can change in response to selection—one can cite fitness differences or driftability in causal explanations of evolutionary change. But, on the causalist side, it (...) is often not clear how, precisely, one should understand these causes. Thus, much more could be said about what sort of causes fitness and driftability are. In this paper, I borrow Dretske's distinction between structuring and triggering causes and I suggest that fitness and driftability are structuring causes of evolution. (shrink)

A major debate in the philosophy of biology centers on the question of how we should understand the causal structure of natural selection. This debate is polarized into the causal and statistical positions. The main arguments from the statistical side are that a causal construal of the theory of natural selection's central concept, fitness, either (i) leads to inaccurate predictions about population dynamics, or (ii) leads to an incoherent set of causal commitments. In this essay, I argue that neither the (...) predictive inaccuracy nor the incoherency arguments successfully undermine the causal account of fitness. 1 Introduction2 The Importance of Trait Fitness3 Trait Fitness is not a Silver Bullet4 The Fundamental Incoherency Argument5 Car Racing and Trait Fitness Reversals6 Population Subdivisions and Evolution7 The STP and the Argument for the Incoherency of the Causal Account of Fitness8 Conclusions. (shrink)

A major debate in the philosophy of biology centers on the question of how we should understand the causal structure of natural selection. This debate is polarized into the causal and statistical positions. The main arguments from the statistical side are that a causal construal of the theory of natural selection's central concept, fitness, either (i) leads to inaccurate predictions about population dynamics, or (ii) leads to an incoherent set of causal commitments. In this essay, I argue that neither the (...) predictive inaccuracy nor the incoherency arguments successfully undermine the causal account of fitness. 1 Introduction2 The Importance of Trait Fitness3 Trait Fitness is not a Silver Bullet4 The Fundamental Incoherency Argument5 Car Racing and Trait Fitness Reversals6 Population Subdivisions and Evolution7 The STP and the Argument for the Incoherency of the Causal Account of Fitness8 Conclusions. (shrink)

In this paper, I argue (contra some recent philosophical work) that an objective distinction between natural selection and drift can be drawn. I draw this distinction by conceiving of drift, in the most fundamental sense, as an individual-level phenomenon. This goes against some other attempts to distinguish selection from drift, which have argued either that drift is a population-level process or that it is a population-level product. Instead of identifying drift with population-level features, the account introduced here can explain these (...) population-level features based on a property that I label driftability. Additionally, this account shows that biology’s “first law”—the Principle of Drift (Brandon, J Phil 102(7):319–335 2006)—is not a foundational law, but is a consequence of driftability. (shrink)

Biological fitness is a foundational concept in the theory of natural selection. Natural selection is often defined in terms of fitness differences as “any consistent difference in fitness (i.e., survival and reproduction) among phenotypically different biological entities” (Futuyma 1998, 349). And in Lewontin’s (1970) classic articulation of the theory of natural selection, he lists fitness differences as one of the necessary conditions for evolution by natural selection to occur. Despite this foundational position of fitness, there remains much debate over the (...) nature of fitness, especially whether fitness differences can truly be said to cause evolutionary change. In recent years these debates have crystalized into two camps: (1) causalists, who see fitness differences as being one of the causes of evolutionary change, and (2) statisticalists, who deny the causal efficacy of fitness and instead hold that “fitness is a mere statistical, noncausal property of trait types” (Walsh 2010, 148). (shrink)