What is natural selection?

‘Natural selection’ is, it seems, an ambiguous term. It is sometimes held to denote a consequence of variation, heredity, and environment, while at other times as denoting a force that creates adaptations. I argue that the latter, the force interpretation, is a redundant notion of natural selection. I will point to difficulties in making sense of this linguistic practise, and argue that it is frequently at odds with standard interpretations of evolutionary theory. I provide examples to show this; one example involving the relation between adaptations and other traits, and a second involving the relation between selection and drift.     

Explanatory unification and natural selection explanations

The debate between the dynamical and the statistical interpretations of natural selection is centred on the question of whether all explanations that employ the concepts of natural selection and drift are reducible to causal explanations. The proponents of the statistical interpretation answer negatively, but insist on the fact that selection/drift arguments are explanatory. However, they remain unclear on where the explanatory power comes from. The proponents of the dynamical interpretation answer positively and try to reduce selection/drift arguments to some of the most prominent accounts of causal explanation. In turn, they face the criticism raised by statisticalists that current accounts of causation have to be violated in some of their core conditions or otherwise used in a very loose manner in order to account for selection/drift explanations. We propose a reconciliation of both interpretations by conveying evolutionary explanations within the unificationist model of scientific explanation. Therefore, we argue that the explanatory power in natural selection arguments is a result of successful unification of individual- and population-level facts. A short case study based on research on sympatric speciation will be presented as an example of how population- and individual-level facts are unified to explain the morphological mosaic of bill shape in island scrub jays (Aphelocoma insularis).     

Human behavioral ecology and niche construction

We examine the relationship between niche construction theory (NCT) and human behavioral ecology (HBE), two branches of evolutionary science that are important sources of theory in archeology. We distinguish between formal models of niche construction as an evolutionary process, and uses of niche construction to refer to a kind of human behavior. Formal models from NCT examine how environmental modification can change the selection pressures that organisms face. In contrast, formal models from HBE predict behavior assuming people behave adaptively in their local setting, and can be used to predict when and why people engage in niche construction. We emphasize that HBE as a field is much broader than foraging theory and can incorporate social and cultural influences on decision‐making. We demonstrate how these approaches can be formally incorporated in a multi‐inheritance framework for evolutionary research, and argue that archeologists can best contribute to evolutionary theory by building and testing models that flexibly incorporate HBE and NCT elements.     

A field guide for teaching evolution in the social sciences

The theory of evolution by natural selection has begun to revolutionize our understanding of perception, cognition, language, social behavior, and cultural practices. Despite the centrality of evolutionary theory to the social sciences, many students, teachers, and even scientists struggle to understand how natural selection works. Our goal is to provide a field guide for social scientists on teaching evolution, based on research in cognitive psychology, developmental psychology, and education. We synthesize what is known about the psychological obstacles to understanding evolution, methods for assessing evolution understanding, and pedagogical strategies for improving evolution understanding. We review what is known about teaching evolution about nonhuman species and then explore implications of these findings for the teaching of evolution about humans. By leveraging our knowledge of how to teach evolution in general, we hope to motivate and equip social scientists to begin teaching evolution in the context of their own field.     

Do the evolutionary origins of our moral beliefs undermine moral knowledge?

According to some recent arguments, (Joyce in The evolution of morality, MIT Press, Cambridge, 2006; Ruse and Wilson in Conceptual issues in evolutionary biology, MIT Press, Cambridge, 1995; Street in Philos Studies 127: 109–166, 2006) if our moral beliefs are products of natural selection, then we do not have moral knowledge. In defense of this inference, its proponents argue that natural selection is a process that fails to track moral facts. In this paper, I argue that our having moral knowledge is consistent with, (a) the hypothesis that our moral beliefs are products of natural selection, and (b) the claim (or a certain interpretation of the claim) that natural selection fails to track moral facts. I also argue that natural selection is a process that could track moral facts, albeit imperfectly. I do not argue that we do have moral knowledge. I argue instead that Darwinian considerations provide us with no reason to doubt that we do, and with some reasons to suppose that we might.     

Running with the Red Queen: the role of biotic conflicts in evolution

What are the causes of natural selection? Over 40 years ago, Van Valen proposed the Red Queen hypothesis, which emphasized the primacy of biotic conflict over abiotic forces in driving selection. Species must continually evolve to survive in the face of their evolving enemies, yet on average their fitness remains unchanged. We define three modes of Red Queen coevolution to unify both fluctuating and directional selection within the Red Queen framework. Empirical evidence from natural interspecific antagonisms provides support for each of these modes of coevolution and suggests that they often operate simultaneously. We argue that understanding the evolutionary forces associated with interspecific interactions requires incorporation of a community framework, in which new interactions occur frequently. During their early phases, these newly established interactions are likely to drive fast evolution of both parties. We further argue that a more complete synthesis of Red Queen forces requires incorporation of the evolutionary conflicts within species that arise from sexual reproduction. Reciprocally, taking the Red Queen''s perspective advances our understanding of the evolution of these intraspecific conflicts.     

Constraining the adaptationism debate

This contribution to the adaptationism debate elaborates the nature of constraints and their importance in evolutionary explanation and argues that the adaptationism debate should be limited to the issue of how to privilege causes in evolutionary explanation. I argue that adaptationist explanations are deeply conceptually dependent on developmental constraints, and explanations that appeal to constraints are dependant on the results of natural selection. I suggest these explanations should be integrated into the framework of historical causal explanation. Each strategy explicitly appeals to some aspect of the evolutionary process, while implicitly appealing to others. Thus, adaptationists and anti-adaptationists can offer complementary causal explanations of the same explanandum. This eliminates much of the adaptationism debate and explains why its adversaries regularly agree with each other more than they would like. The adaptationism issue that remains is a species of the general issue of how to privilege causes in explanation. I show how a proposed solution to this general problem might be brought to bear on evolutionary explanations, and investigate some difficulties that might arise due to the nature of the evolutionary process.     

Natural selection. IV. The Price equation

The Price equation partitions total evolutionary change into two components. The first component provides an abstract expression of natural selection. The second component subsumes all other evolutionary processes, including changes during transmission. The natural selection component is often used in applications. Those applications attract widespread interest for their simplicity of expression and ease of interpretation. Those same applications attract widespread criticism by dropping the second component of evolutionary change and by leaving unspecified the detailed assumptions needed for a complete study of dynamics. Controversies over approximation and dynamics have nothing to do with the Price equation itself, which is simply a mathematical equivalence relation for total evolutionary change expressed in an alternative form. Disagreements about approach have to do with the tension between the relative valuation of abstract versus concrete analyses. The Price equation's greatest value has been on the abstract side, particularly the invariance relations that illuminate the understanding of natural selection. Those abstract insights lay the foundation for applications in terms of kin selection, information theory interpretations of natural selection and partitions of causes by path analysis. I discuss recent critiques of the Price equation by Nowak and van Veelen.     

'Molecules and monkeys': George Gaylord Simpson and the challenge of molecular evolution

In this paper, I analyze George Gaylord Simpson's response to the molecularization of evolutionary biology from his unique perspective as a paleontologist. I do so by exploring his views on early attempts to reconstruct phylogenetic relationships among primates using molecular data. Particular attention is paid to Simpson's role in the evolutionary synthesis of the 1930s and 1940s, as well as his concerns about the rise of molecular biology as a powerful discipline and world-view in the 1960s. I argue that Simpson's belief in the supremacy of natural selection as the primary driving force of evolution, as well as his view that biology was a historical science that seeks ultimate causes and highlights contingency, prevented him from acknowledging that the study of molecular evolution was an inherently valuable part of the life sciences.     

Truth, Selection and Scientific Inquiry

In this paper I examine various ways in whichphilosophers have made connections between truth andnatural selection. I introduce several versions ofthe view that mechanisms of true belief generationarise as a result of natural selection and argue thatthey fail to establish a connection between truth andnatural selection. I then turn to scientific truthsand argue that evolutionary accounts of the origin ofscientific truth generation mechanisms also fail. Iintroduce David Hull's selectionist model ofscientific development and argue that his account ofscientific success does not rely on connecting truthand natural selection. I argue that Hull's model,which severs the connection between truth andselection, can account for some aspects of scientificchange, but it still leaves us plenty of questionsabout what aspects of our individual cognitive make-upcontribute to scientific change and how they do so. I introduce an evolutionary approach to scientificcognition that shows how some of these questions canbe answered without making an explanatory appeal toselection for true belief generating mechanisms.     

"Curiously parallel": analogies of language and race in Darwin's Descent of man. A reply to Gregory Radick

In the second chapter of The descent of man (1871), Charles Darwin interrupted his discussion of the evolutionary origins of language to describe ten ways in which the formation of languages and of biological species were 'curiously' similar. I argue that these comparisons served mainly as analogies in which linguistic processes stood for aspects of biological evolution. Darwin used these analogies to recapitulate themes from On the origin of species (1859), including common descent, genealogical classification, the struggle for existence, and natural selection, among others. Skeptical of this interpretation, Gregory Radick sees the naturalistic account of language formation in the Descent comparisons as reinforcing Darwin's idea that languages and the races of mankind have both undergone progressive development. (The opposite view was that modern-day primitive peoples had degenerated from an originally civilized condition.) Yet the details of Darwin's language-species comparisons, as well as the polemical context in which they appear, show that they were not aimed at so limited a function. Rather, they addressed issues related to species transmutation in general.     

The evolution of a social construction: the case of male homosexuality

Male homosexuality has been viewed by evolutionary psychologists as a Darwinian paradox, and by other social scientists as a social construction. We argue that it is better understood as an evolutionary social construction. Male homosexuality as we now know it is an 18th-century invention, but nonexclusive same-sex sexual behavior has a long evolutionary history. According to the alliance-formation hypothesis, same-sex sexuality evolved by natural selection because it created or strengthened male-male alliances and allowed low-status males to reposition themselves in the group hierarchy and thereby increase their reproductive success. This hypothesis makes sense of some odd findings about male homosexuality and helps to explain the rise in exclusive male homosexuality in the 18th century. The sociohistorical conditions around 1700 may have resulted in an increase in same-sex sexual behavior. Cultural responses to same-sex sexuality led to the spread of exclusive homosexual behavior and to the creation of a homosexual identity. Understanding male homosexuality as an evolutionary social construction can help us move beyond the traditionally polarized debate between evolutionary psychologists and social constructionists.     

Cultural evolution and the social sciences: a case of unification?

This paper addresses the question of how to understand the relationship between Cultural Evolutionary Science (CES) and the social sciences, given that they coexist and both study cultural change. I argue that CES is best understood as having a unificatory or integrative role between evolutionary biology and the social sciences, and that it is best characterized as a bridge field; I describe the concept of a bridge field and how it relates to other non-reductionist accounts of unification or integration used in the philosophy of science literature.     

Utopianism in the British evolutionary synthesis

In this paper I propose a new interpretation of the British evolutionary synthesis. The synthetic work of J. B. S. Haldane, R. A. Fisher and J. S. Huxley was characterized by both an integration of Mendelism and Darwinism and the unification of different biological subdisciplines within a coherent framework. But it must also be seen as a bold and synthetic Darwinian program in which the biosciences served as a utopian blueprint for the progress of civilization. Describing the futuristic visions of these three scientists in their synthetic heydays, I show that, despite a number of important divergences, their biopolitical ideals could be biased toward a controlled and regimented utopian society. Their common ideals entailed a social order where liberal and democratic principles were partially or totally suspended in favor of bioscientific control and planning for the future. Finally, I will argue that the original redefinition of Darwinism that modern synthesizers proposed is a significant historical example of how Darwinism has been used and adapted in different contexts. The lesson I draw from this account is a venerable one: that, whenever we wish to define Darwinism, we need to recognize not only its scientific content and achievements but expose the other traditions and ideologies it may have supported.     

Has the combination of genetic and fossil evidence solved the riddle of modern human origins?

Debate over the origin of modern humans continues without a clear end in sight. Currently, the genetic and fossil evidence is still used to support two different interpretations of the origin of modern humans. Some researchers claim that the genetic evidence is compatible with either an Out‐of‐Africa or a Multiregional model, while other scientists argue that the evidence supports only a Multiregional model of evolution. I argue that the fossil record and archeological evidence constrain interpretation of the genetic evidence and imply that very little, if any, admixture with Eurasian archaic hominins such as the Neanderthals occurred during the spread of modern humans out of Africa.     

Selfish altruism,fierce cooperation and the predator*

This paper suggests a new way to think about a famous question: what explains cooperation in nature and in particular in humans? I argue that, for an evolutionary biologist as well as a quantitative social scientist, the triangle of two ‘teammates’ in the presence of a predator (passing and shooting in two-on-one situations) is one of the fundamental conceptual building-blocks for understanding these phenomena because in such a situation the fact that life is packaged in many distinct enclosures (and not in one big monolithic blob) can unfold its comparative advantage. I show how, in the presence of a predator, cooperative equilibria emerge among entirely selfish teammates if we infinitesimally bias the lead player in the selfish direction or assign a computational burden on the predator due to the presence of a teammate. I argue that ‘predators’ are common in the biological jungle but also in everyday human settings. Intuitively, this paper builds on the simple idea – a familiar one to a biologist observing the natural world but perhaps less so to social scientists – that everybody has enemies.     

Evolutionary medicine at twenty: rethinking adaptationism and disease

Two decades ago, the eminent evolutionary biologist George C. Williams and his physician coauthor, Randolph Nesse, formulated the evolutionary medicine research program. Williams and Nesse explicitly made adaptationism a core component of the new program, which has served to undermine the program ever since, distorting its practitioners’ perceptions of evidentiary burdens and in extreme cases has served to warp practitioner’s understandings of the relationship between evolutionary benefits/detriments and medical ones. I show that the Williams and Nesse program more particularly embraces the panselectionist variety of adaptationism (the empirical assumption that non-adaptive evolutionary processes are causally unimportant compared to natural selection), and argue that this has harmed the field. Panselectionism serves to conceal the enormous evidentiary hurdles that evolutionary medicine hypotheses face, making them appear stronger than they are. I use two examples of evolutionary medicine texts, on neonatal jaundice and on asthma, to show that some evolutionary medicine practitioners have allowed their fervent panselectionism to directly shape their recommendations for clinical practice. I argue that this escalation of panselectionism’s influence is inappropriate under Williams’ and Nesse original stated standards, despite being inspired by their program. I also show that the examples’ conflation of clinical and evolutionary considerations is inappropriate even under Christopher Boorse’s controversial evolution-rooted concepts of disease and health.     

Cultural transmission and the evolution of human behaviour: a general approach based on the Price equation

Transmitted culture can be viewed as an inheritance system somewhat independent of genes that is subject to processes of descent with modification in its own right. Although many authors have conceptualized cultural change as a Darwinian process, there is no generally agreed formal framework for defining key concepts such as natural selection, fitness, relatedness and altruism for the cultural case. Here, we present and explore such a framework using the Price equation. Assuming an isolated, independently measurable culturally transmitted trait, we show that cultural natural selection maximizes cultural fitness, a distinct quantity from genetic fitness, and also that cultural relatedness and cultural altruism are not reducible to or necessarily related to their genetic counterparts. We show that antagonistic coevolution will occur between genes and culture whenever cultural fitness is not perfectly aligned with genetic fitness, as genetic selection will shape psychological mechanisms to avoid susceptibility to cultural traits that bear a genetic fitness cost. We discuss the difficulties with conceptualizing cultural change using the framework of evolutionary theory, the degree to which cultural evolution is autonomous from genetic evolution, and the extent to which cultural change should be seen as a Darwinian process. We argue that the nonselection components of evolutionary change are much more important for culture than for genes, and that this and other important differences from the genetic case mean that different approaches and emphases are needed for cultural than genetic processes.     

Levels of selection and the formal Darwinism project

Understanding good design requires addressing the question of what units undergo natural selection, thereby becoming adapted. There is, therefore, a natural connection between the formal Darwinism project (which aims to connect population genetics with the evolution of design and fitness maximization) and levels of selection issues. We argue that the formal Darwinism project offers contradictory and confusing lines of thinking concerning level(s) of selection. The project favors multicellular organisms over both the lower (cell) and higher (social group) levels as the level of adaptation. Grafen offers four reasons for giving such special status to multicellular organisms: (1) they lack appreciable within-organism cell selection, (2) they have multiple features that appear contrived for the same purpose, (3) they possess a set of phenotypes, and (4) they leave offspring according to their phenotypes. We discuss why these rationales are not compelling and suggest that a more even-handed approach, in which multicellular organisms are not assumed to have special status, would be desirable for a project that aims to make progress on the foundations of evolutionary theory.