Life,the universe and everything

The Formal Darwinism project probes the connections between the dynamics of natural selection and the design of organisms. Here, I explain why this work should be of interest to philosophers, arguing that it is the natural development in a long-running scholarly enquiry into the meaning of life. I then review some of my own work which has applied the tools of Formal Darwinism to address issues concerning the units of adaptation in social evolution, leading to a deeper understanding of the adaptation of individual organisms. Finally, I sketch some directions Formal Darwinism to explore beyond the biological sciences, with a focus upon cosmology.     

Two neo-Darwinisms

There are two extant theories of evolution, each of which deserves the honourific "neo-Darwinism": Modern Synthesis Replicator theory and a theory I shall call Developmental Darwinism. The principal difference concerns the canonical unit of biological organization. Modern Synthesis replicator theory explains the process of evolution by appeal to the activities of genes or replicators. Developmental Darwinism explains the process of evolution by appeal to the capacities of organisms. In particular, it is the plasticity of organisms, manifested most distinctly during development, that causes adaptive evolution. Despite the fact that each, in its own way, traces its origin to the theory outlined by Darwin, they are radically different. The objectives of this essay are twofold: to underscore the differences between these theories, and to argue that Developmental Darwinism, though nascent, is a viable alternative to Modern Synthesis replicator theory.     

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.     

The Creativity of Natural Selection? Part I: Darwin,Darwinism, and the Mutationists

This is the first of a two-part essay on the history of debates concerning the creativity of natural selection, from Darwin through the evolutionary synthesis and up to the present. Here I focus on the mid-late nineteenth century to the early twentieth, with special emphasis on early Darwinism and its critics, the self-styled “mutationists.” The second part focuses on the evolutionary synthesis and some of its critics, especially the “neutralists” and “neo-mutationists.” Like Stephen Gould, I consider the creativity of natural selection to be a key component of what has traditionally counted as “Darwinism.” I argue that the creativity of natural selection is best understood in terms of (1) selection initiating evolutionary change, and (2) selection being responsible for the presence of the variation it acts upon, for example by directing the course of variation. I consider the respects in which both of these claims sound non-Darwinian, even though they have long been understood by supporters and critics alike to be virtually constitutive of Darwinism.     

Natural selection and the maximization of fitness

The notion that natural selection is a process of fitness maximization gets a bad press in population genetics, yet in other areas of biology the view that organisms behave as if attempting to maximize their fitness remains widespread. Here I critically appraise the prospects for reconciliation. I first distinguish four varieties of fitness maximization. I then examine two recent developments that may appear to vindicate at least one of these varieties. The first is the ‘new’ interpretation of Fisher's fundamental theorem of natural selection, on which the theorem is exactly true for any evolving population that satisfies some minimal assumptions. The second is the Formal Darwinism project, which forges links between gene frequency change and optimal strategy choice. In both cases, I argue that the results fail to establish a biologically significant maximization principle. I conclude that it may be a mistake to look for universal maximization principles justified by theory alone. A more promising approach may be to find maximization principles that apply conditionally and to show that the conditions were satisfied in the evolution of particular traits.     

Adaptation,fitness and the selection-optimality links

We critically examine a number of aspects of Grafen’s ‘formal Darwinism’ project. We argue that Grafen’s ‘selection-optimality’ links do not quite succeed in vindicating the working assumption made by behavioural ecologists and others—that selection will lead organisms to exhibit adaptive behaviour—since these links hold true even in the presence of strong genetic and developmental constraints. However we suggest that the selection-optimality links can profitably be viewed as constituting an axiomatic theory of fitness. Finally, we compare Grafen’s project with Fisher’s ‘fundamental theorem of natural selection’, and we speculate about whether Grafen’s results can be extended to a game-theoretic setting.     

Comparison and adaptation.

It has sometimes been suggested that the term adaptation should be reserved for differences with a known genetic basis. We argue that adaptation should be defined by its effects rather than by its causes as any difference between two phenotypic traits (or trait complexes) which increases the inclusive fitness of its carrier. This definition implies that some adaptations may arise by means other than natural selection. It is particularly important to bear this in mind when behavioural traits are considered. Critics of the 'adaptationist programme' have suggested that an important objection to many adaptive explanations is that they rely on ad-hoc arguments concerning the function of previously observed differences. We suggest that this is a less important problem (because evolutionary explanations generally claim some sort of generality and are therefore testable) than the difficulties arising from confounding variables. These are more widespread and more subtle than is generally appreciated. Not all differences between organisms are directly adapted to ecological variation. The form of particular traits usually constrains the form of value that other traits can take, presenting several obstacles to attempts to relate variation in morphological or behavioural characteristics directly to environmental differences. We describe some of the repercussions of differences in body size among vertebrates and ways in which these can be allowed for. In addition, a variety of evolutionary processes can produce non-adaptive differences between organisms. One way of distinguishing between these and adaptations is to investigate adaptive trends in phylogenetically different groups of species.     

Genomic imprinting and the units of adaptation

Two guiding principles identify which biological entities are able to evolve adaptations. Williams'' principle holds that, in order for an entity to evolve adaptations, there must be selection between such entities. Maynard Smith''s principle holds that, in order for an entity to evolve adaptations, selection within such entities must be absent or negligible. However, although the kinship theory of genomic imprinting suggests that parent-of-origin-specific gene expression evolves as a consequence of natural selection acting between—rather than within—individuals, it evades adaptive interpretation at the individual level and is instead viewed as an outcome of an intragenomic conflict of interest between an individual''s genes. Here, I formalize the idea that natural selection drives intragenomic conflicts of interest between genes originating from different parents. Specifically, I establish mathematical links between the dynamics of natural selection and the idea of the gene as an intentional, inclusive-fitness-maximizing agent, and I clarify the role that information about parent of origin plays in mediating conflicts of interest between genes residing in the same genome. These results highlight that the suppression of divisive information may be as important as the suppression of lower levels of selection in maintaining the integrity of units of adaptation.     

On adaptation, the assessment of adaptations, and the value of adaptive arguments in phylogenetic reconstruction

Evolutionary adaptation concerns a relative concept and the study of adaptations is directed to structures of individuals. The concept is devoid of any meaning when it is applied to species or populations. Adaptation is not synonymous with fitness or survival but does contribute to both of them. The term adaptation has a dual meaning since it refers both to the process of adaptation and to the state of being adapted. In the process of adaptation the mechanism of natural selection takes a prominent position. But the operation and effectiveness of natural selection are constrained by various limiting factors. Besides that, features may also be the result of nonadaptive evolution and only attain their present adaptive function at a later point in time. Another possibility is that features have at present a function different from the one for which they were initially designed. With respect to the state of being, the study of adaptation attempts to examine whether a particular feature indeed forms an adequate response to selection forces from the environment. Five methods or approaches generally are used to assess the adaptive significance of features, viz. the comparative, correlation, optimization, cladistic, and synthetic approach. Only the last-mentioned approach forms an adequate method since it attempts to establish, by direct analysis, which well-defined selection force exerts its influence on a certain character. The practicing taxonomist is faced with the problem that the data necessary to apply the synthetic method, generally require detailed field studies. Not all evolutionary changes are under the influence of natural selection. The presence of some features may be based on entirely different mechanisms, such as genetic drift, mutational pressure, pleiotropic gene action, allometric growth, or ecophenotypic responses. Various problems inherent to the optimization approach, and several others of practical and theoretical nature, make the morphocline method of the functional and evolutionary morphologists unsuitable as a method for phylogenetic reconstruction.     

A critique of Darwinian anthropology

Darwinian anthropology holds that human behavior is adaptive in the sense of being designed to maximize reproductive success and that measurement of reproductive differentials typically illuminates adaptation. I argue to the contrary, that adaptive design is usually manifested at the psychological rather than at the behavioral level, that measuring reproductive differentials is at best an inefficient and ambiguous way to illuminate adaptation, and that Darwin's theory of natural selection sheds light on human affairs only insofar as it promotes understanding of the psychology that underpins these affairs.     

Did Darwin really answer Paley’s question?

It is commonly thought that natural selection explains the rise of adaptive complexity. Razeto-Barry and Frick (2011) have recently argued in favour of this view, dubbing it the Creative View. I argue that the Creative View is mistaken if it claims that natural selection serves to answer Paley’s question. This is shown by a case that brings out the contrastive structure inherent in this demand for explanation. There is, however, a rather trivial sense in which specific environmental conditions are crucial for the rise of specific adaptations, but this is hardly what opponents of the Creative View are denying.     

Constraints on adaptive evolution: the functional trade-off between reproduction and fast-start swimming performance in the Trinidadian guppy (Poecilia reticulata)

The empirical study of natural selection reveals that adaptations often involve trade-offs between competing functions. Because natural selection acts on whole organisms rather than isolated traits, adaptive evolution may be constrained by the interaction between traits that are functionally integrated. Yet, few attempts have been made to characterize how and when such constraints are manifested or whether they limit the adaptive divergence of populations. Here we examine the consequences of adaptive life-history evolution on locomotor performance in the live-bearing guppy. In response to increased predation from piscivorous fish, Trinidadian guppies evolve an increased allocation of resources toward reproduction. These populations are also under strong selection for rapid fast-start swimming performance to evade predators. Because embryo development increases a female's wet mass as she approaches parturition, an increased investment in reproductive allocation should impede fast-start performance. We find evidence for adaptive but constrained evolution of fast-start swimming performance in laboratory trials conducted on second-generation lab-reared fish. Female guppies from high-predation localities attain a faster acceleration and velocity and travel a greater distance during fast-start swimming trials. However, velocity and distance traveled decline more rapidly over the course of pregnancy in these same females, thus reducing the magnitude of divergence in swimming performance between high- and low-predation populations. This functional trade-off between reproduction and swimming performance reveals how different aspects of the phenotype are integrated and highlights the complexity of adaptation at the whole-organism level.     

Neutral mutation as the source of genetic variation in life history traits

The mechanism underlying the maintenance of adaptive genetic variation is a long-standing question in evolutionary genetics. There are two concepts (mutation-selection balance and balancing selection) which are based on the phenotypic differences between alleles. Mutation - selection balance and balancing selection cannot properly explain the process of gene substitution, i.e. the molecular evolution of quantitative trait loci affecting fitness. I assume that such loci have non-essential functions (small effects on fitness), and that they have the potential to evolve into new functions and acquire new adaptations. Here I show that a high amount of neutral polymorphism at these loci can exist in real populations. Consistent with this, I propose a hypothesis for the maintenance of genetic variation in life history traits which can be efficient for the fixation of alleles with very small selective advantage. The hypothesis is based on neutral polymorphism at quantitative trait loci and both neutral and adaptive gene substitutions. The model of neutral - adaptive conversion (NAC) assumes that neutral alleles are not neutral indefinitely, and that in specific and very rare situations phenotypic (relative fitness) differences between them can appear. In this paper I focus on NAC due to phenotypic plasticity of neutral alleles. The important evolutionary consequence of NAC could be the increased adaptive potential of a population. Loci responsible for adaptation should be fast evolving genes with minimally discernible phenotypic effects, and the recent discovery of genes with such characteristics implicates them as suitable candidates for loci involved in adaptation.     

Notes from a snail island: Littorinid evolution and adaptation

The most successful study systems are built on a foundation of decades of research on the basic biology, ecology and life history of the organisms in question. Combined with new technologies, this can provide a formidable means to address important issues in evolutionary biology and molecular ecology. Littorinid marine snails are a good example of this, with a rich literature on their taxonomy, speciation, thermal tolerance and behavioural adaptations. In August 2017, an international meeting on Littorinid evolution was held at the Tjärnö Marine Research Laboratory in Western Sweden. In this meeting review, I provide a summary of some of the exciting work on parallel evolution, sexual selection and adaptation to environmental stress presented there. I argue that newly available genomic resources present an opportunity for integrating the traditionally divergent fields of speciation and environmental adaptation in Littorinid research.     

论达尔文医学(Ⅰ)

一切生物功能的设计都用查理士．达尔文的的自然选择理论来解释,是本文中贯彻始终的思想,探讨的中心自然自动所控制所挑选的适应性变化这一概念：我们与病原格斗适应性变化,病原对抗这变而产生的知识性变化。     

论达尔文医学(Ⅱ)

一切生物功能的设计都用查理士·达尔文的自然选择理论来解释,是本文中贯彻始终的思想．探讨的中心是自然选择所挑选的适应性变化这一概念：我们与病原格斗适应性变化,病原对抗这变化而产生的适应性变化,我们为了这些变化必须付出代价而出现的不适应性,以及机体设计和现在的生活环境之间的不适应性,等等．我们希望读者将在对身体的功能以及某些异常情况的进化论解释中得到教益．     

Organisms as natural purposes: the contemporary evolutionary perspective

Kant's conception of organisms as natural purposes raises a challenge to the adequacy of mechanistic explanation in biology. Certain features of organisms appear to be inexplicable by appeal to mechanical law alone. Some biological phenomena, it seems, can only be accounted for teleologically. Contemporary evolutionary biology has by and large ignored this challenge. It is widely held that Darwin's theory of natural selection gives us an adequate, wholly mechanical account of the nature of organisms. In contemporary biology, the category of the organism plays virtually no explanatory role. Contemporary evolutionary biology is a science of sub-organismal entities-replicators. I argue that recent advances in developmental biology demonstrate the inadequacy of sub-organismal mechanism. The category of the organism, construed as a 'natural purpose' should play an ineliminable role in explaining ontogenetic development and adaptive evolution. According to Kant the natural purposiveness of organisms cannot be demonstrated to be an objective principle in nature, nor can purposiveness figure in genuine explain. I attempt to argue, by appeal to recent work on self-organization, that the purposiveness of organisms is a natural phenomenon, and, by appeal to the apparatus of invariance explanation, that biological purposiveness provides genuine, ineliminable biological explanations.     

Darwinism without populations: a more inclusive understanding of the "Survival of the Fittest"

Following Wallace's suggestion, Darwin framed his theory using Spencer's expression "survival of the fittest". Since then, fitness occupies a significant place in the conventional understanding of Darwinism, even though the explicit meaning of the term 'fitness' is rarely stated. In this paper I examine some of the different roles that fitness has played in the development of the theory. Whereas the meaning of fitness was originally understood in ecological terms, it took a statistical turn in terms of reproductive success throughout the 20th Century. This has lead to the ever-increasing importance of sexually reproducing organisms and the populations they compose in evolutionary explanations. I will argue that, moving forward, evolutionary theory should look back at its ecological roots in order to be more inclusive in the type of systems it examines. Many biological systems (e.g. clonal species, colonial species, multi-species communities) can only be satisfactorily accounted for by offering a non-reproductive account of fitness. This argument will be made by examining biological systems with very small or transient population structures. I argue this has significant consequences for how we define Darwinism, increasing the significance of survival (or persistence) over that of reproduction.     

Reviving the superorganism

Individuals become functionally organized to survive and reproduce in their environments by the process of natural selection. The question of whether larger units such as groups and communities can possess similar properties of functional organization, and therefore be regarded as "superorganisms", has a long history in biological thought. Modern evolutionary biology has rejected the concept of superorganisms, explaining virtually all adaptations at the individual or gene level. We criticize the modern literature on three counts. First, individual selection in its strong form is founded on a logical contradiction, in which genes-in-individuals are treated differently than individuals-in-groups or species-in-communities. Imposing consistency clearly shows that groups and communities can be organisms in the same sense that individuals are. Furthermore, superorganisms are more than just a theoretical possibility and actually exist in nature. Second, the view that genes are the "ultimate" unit of selection is irrelevant to the question of functional organization. Third, modern evolutionary biology includes numerous conceptual frameworks for analyzing evolution in structured populations. These frameworks should be regarded as different ways of analyzing a common process which, to be correct, must converge on the same conclusions. Unfortunately, evolutionists frequently regard them as competing theories that invoke different mechanisms, such that if one is "right" the others must be "wrong". The problem of multiple frameworks is aggravated by the fact that major terms, such as "units of selection", are defined differently within each framework, yet many evolutionists who use one framework to argue against another assume shared meanings. We suggest that focusing on the concept of organism will help dispell this fog of semantic confusion, allowing all frameworks to converge on the same conclusions regarding units of functional organization.