Multiple Explanations in Darwinian Evolutionary Theory

Variational evolutionary theory as advocated by Darwin is not a single theory, but a bundle of related but independent theories, namely: (a) variational evolution; (b) gradualism rather than large leaps; (c) processes of phyletic evolution and of speciation; (d) causes for the formation of varying individuals in populations and for the action of selective agents; and (e) all organisms evolved from a common ancestor. The first four are nomological-deductive explanations and the fifth is historical-narrative. Therefore evolutionary theory must be divided into nomological and historical theories which are both testable against objective empirical observations. To be scientific, historical evolutionary theories must be based on well corroborated nomological theories, both evolutionary and functional. Nomological and general historical evolutionary theories are well tested and must be considered as strongly corroborated scientific theories. Opponents of evolutionary theory are concerned only with historical evolutionary theories, having little interest in nomological theory. Yet given a well corroborated nomological evolutionary theory, historical evolutionary theories follow automatically. If understood correctly, both forms of evolutionary theories stand on their own as corroborated scientific theories and should not be labeled as facts.     

Evolutionary epistemology as science

What credentials does evolutionary epistemology have as science? A judgement based on past performance, both in terms of advancing an empirical programme and further ng theory construction, is not much. This paper briefly outlines some of the research areas, both theoretical and empirical, that can be developed and that might secure for evolutionary epistemology a future in evolutionary biology.     

A general theory for the evolutionary dynamics of virulence

Most theory on the evolution of virulence is based on a game-theoretic approach. One potential shortcoming of this approach is that it does not allow the prediction of the evolutionary dynamics of virulence. Such dynamics are of interest for several reasons: for experimental tests of theory, for the development of useful virulence management protocols, and for understanding virulence evolution in situations where the epidemiological dynamics never reach equilibrium and/or when evolutionary change occurs on a timescale comparable to that of the epidemiological dynamics. Here we present a general theory similar to that of quantitative genetics in evolutionary biology that allows for the easy construction of models that include both within-host mutation as well as superinfection and that is capable of predicting both the short- and long-term evolution of virulence. We illustrate the generality and intuitive appeal of the theory through a series of examples showing how it can lead to transparent interpretations of the selective forces governing virulence evolution. It also leads to novel predictions that are not possible using the game-theoretic approach. The general theory can be used to model the evolution of other pathogen traits as well.     

How a Kantian Can Accept Evolutionary Metaethics

Contrary to widely held assumptions, an evolutionary metaethics need not be non-cognitivist. I define evolutionary metaethics as the claim that certain phenotypic traits expressing certain genes are both necessary and sufficient for explanation of all other phenotypic traits we consider morally significant. A review of the influential cognitivist Immanuel Kants metaethics shows that much of his ethical theory is independent of the anti-naturalist metaphysics of transcendental idealism which itself is incompatible with evolutionary metaethics. By matching those independent aspects to an evolutionary metaethics a cognitivist Kantian evolutionary metaethical theory is a possibility for researchers to consider.     

Evolution and literary theory

Presupposing that all knowledge is the study of a unitary order of nature, the author maintains that the study of literature should be included within the larger field of evolutionary theory. He outlines four elementary concepts in evolutionary theory, and he argues that these concepts should regulate our understanding of literature. On the basis of these concepts, he repudiates the antirealist and irrationalist views that, under the aegis of “poststructuralism,” have dominated academic literary studies for the past two decades. He examines the linkage between poststructuralism and standard social science, and he speculates about the ideological and disciplinary motives that have hitherto impeded evolutionary study in both the social sciences and the humanities. Finally, he distinguishes literature from science and argues that literary criticism integrates elements of both.     

Thermodynamics and the conceptual structure of evolutionary theory

Thermodynamics and evolutionary theory have spent most of their shared history in adversarial relationship to one another. The point of this paper is to consider some qualitative ways in which thermodynamics can enrich both the theory and epistemology of evolution. The "autonomy of biology" posture in evolutionary theory hangs on the supposed uniqueness of why-questions in biology. With this posture, and with the general obstruction of constructive dialogue between evolution and the physical sciences it fosters, come the perennial accusations that Darwinism deals in adaptational teleology but not mechanisms. Thermodynamics provides for a two-tiered hierarchy of causation in nature in which the why-question is rendered not only legitimate materialistically, but essential to understanding the evolutionary process in its totality--from the emergence of life to the branching of lineages in speciation.     

Reproduction and the central project of evolutionary theory

In much of the discourse of evolutionary theory, reproduction is treated as an autonomous function of the individual organism — even in discussions of sexually reproducing organisms. In this paper, I examine some of the functions and consequences of such manifestly peculiar language. In particular, I suggest that it provides crucial support for the central project of evolutionary theory — namely that of locating causal efficacy in intrinsic properties of the individual organism. Furthermore, I argue that the language of individual reproduction is maintained by certain methodological conventions that both obscure many of the problems it generates and serve to actively impede attempts to redress those difficulties that can be identified. Finally, I suggest that inclusion of the complexities introduced by sexual reproduction — in both language and methodology — may radically undermine the individualist focus of evolutionary theory.I am indepted to the Rockefeller Foundation for a Humanities Fellowship that supported this research during the spring of 1986. I am also grateful to Richard Lewontin, Diane Paul, and Lisa Lloyd for many extremely helpful conversations.     

Popper, falsifiability, and evolutionary biology

First, a brief history is provided of Popper's views on the status of evolutionary biology as a science. The views of some prominent biologists are then canvassed on the matter of falsifiability and its relation to evolutionary biology. Following that, I argue that Popper's programme of falsifiability does indeed exclude evolutionary biology from within the circumference of genuine science, that Popper's programme is fundamentally incoherent, and that the correction of this incoherence results in a greatly expanded and much more realistic concept of what is empirical, resulting in the inclusion of evolutionary biology. Finally, this expanded concept of empirical is applied to two particular problems in evolutionary biology — viz., the species problem and the debate over the theory of punctuated equilibria — and it is argued that both of them are still mainly metaphysical.     

MATHEMATICS OF KIN‐ AND GROUP‐SELECTION: FORMALLY EQUIVALENT?

Evolutionary game theory is a general mathematical framework that describes the evolution of social traits. This framework forms the basis of many multilevel selection models and is also frequently used to model evolutionary dynamics on networks. Kin selection, which was initially restricted to describe social interactions between relatives, has also led to a broader mathematical approach, inclusive fitness, that can not only describe social evolution among relatives, but also in group structured populations or on social networks. It turns out that the underlying mathematics of game theory is fundamentally different from the approach of inclusive fitness. Thus, both approaches—evolutionary game theory and inclusive fitness—can be helpful to understand the evolution of social traits in group structured or spatially extended populations.     

How evolutionary biology challenges the classical theory of rational choice

A fundamental philosophical question that arises in connection with evolutionary theory is whether the fittest patterns of behavior are always the most rational. Are fitness and rationality fully compatible? When behavioral rationality is characterized formally as in classical decision theory, the question becomes mathematically meaningful and can be explored systematically by investigating whether the optimally fit behavior predicted by evolutionary process models is decision-theoretically coherent. Upon investigation, it appears that in nontrivial evolutionary models the expected behavior is not always in accord with the norms of the standard theory of decision as ordinarily applied. Many classically irrational acts, e.g. betting on the occurrence of one event in the knowledge that the probabilities favor another, can under certain circumstances constitute adaptive behavior. One interesting interpretation of this clash is that the criterion of rationality offered by classical decision theory is simply incorrect (or at least incomplete) as it stands, and that evolutionary theory should be called upon to provide a more generally applicable theory of rationality. Such a program, should it prove feasible, would amount to the logical reduction of the theory of rational choice to evolutionary theory.     

Bioaltruism reconsidered

Altruistic behavior is often regarded as sociobiology's most central theoretical problem, but is it? Altruism in biology, bioaltruism, has many meanings, which can be grouped into two categories. The first I will callcommon bioaltruism. It is primarily of ethological relevance. The second,evolutionary bioaltruism, is a special category in evolutionary respects in that it may indeed pose a problem for evolutionary theory. These categories are logically independent. Moreover, both of them are logically different from altruism in its everyday psychological or moral sense. Sociobiological examples of bioaltruistic behavior concern bioaltruism in the first sense only, so the theoretical problem ‘altruism’ is supposed to pose, is indeed nothing but a theoretical problem and the bioaltruism that actually occurs has no evolutionary relevance. Nevertheless, evolutionary theory is relevant to our understanding of the possibility of common bioaltruism, and that possibility — even though bioaltruism is conceptually different from ethical altruism — is relevant for ethicists: it sheds light on what we can ask people to do or not to do.     

Evolutionary accounts of human behavioural diversity

Human beings persist in an extraordinary range of ecological settings, in the process exhibiting enormous behavioural diversity, both within and between populations. People vary in their social, mating and parental behaviour and have diverse and elaborate beliefs, traditions, norms and institutions. The aim of this theme issue is to ask whether, and how, evolutionary theory can help us to understand this diversity. In this introductory article, we provide a background to the debate surrounding how best to understand behavioural diversity using evolutionary models of human behaviour. In particular, we examine how diversity has been viewed by the main subdisciplines within the human evolutionary behavioural sciences, focusing in particular on the human behavioural ecology, evolutionary psychology and cultural evolution approaches. In addition to differences in focus and methodology, these subdisciplines have traditionally varied in the emphasis placed on human universals, ecological factors and socially learned behaviour, and on how they have addressed the issue of genetic variation. We reaffirm that evolutionary theory provides an essential framework for understanding behavioural diversity within and between human populations, but argue that greater integration between the subfields is critical to developing a satisfactory understanding of diversity.     

Embryology and evolution 1920-1960: worlds apart?

During the early part of the 20th century most embryologists were skeptical about the significance of Mendelian genetics to embryological development. A few embryologists began to study the developmental effects of Mendelian genes around 1940. Such work was a necessary step on the path to modern developmental biology. It occurred during the time when the Evolutionary Synthesis was integrating Mendelian and population genetics into a unified evolutionary theory. Why did the first embryological geneticists begin their study at that particular time? One possible explanation is that developmental genetics was a potential avenue of alliance between embryology and evolutionary biology, two fields that had been separated since the 1890s. To assess this possible motive it is necessary to explore the methodological contrasts that obtained between embryology and both Mendelian-chromosomal genetics and neo-Darwinian evolutionary theory. Some of these contrasts persist to the present day.     

Functional morphology and evolutionary biology

In this study the relationship between functional morpholoy and evolutionary biology is analysed by confronting the main concepts in both disciplines.Rather than only discussing this connection theoretically, the analysis is carried out by introducing important practical and experimental studies, which use aspects from both disciplines. The mentioned investigations are methodologically analysed and the consequences for extensions of the relationship are worked out. It can be shown that both disciplines have a large domain of their own and also share a large common ground. Many disagreements among evolutionary biologists can be reduced to differences in general philosophy (idealism vs. realism), selection of phenomenona (structure vs. function), definition of concepts (natural selection) and the position of the concept theory as an explaining factor (neutralists vs. selectionists, random variation, determinate selection, etc.).The significance of functional morphology for evolutionary biology, and vice versa depends on these differences. For a neo-Darwinian evolutionary theory, contributions from functional and ecological morphology are indispensable. Of ultimate importance are the notions of internal selection and constraints in the constructions determining further development. In this context the concepts of random variation and natural selection need more detailed definition.The study ends with a recommendation for future research founded in a system-theoretical or structuralistic conception.     

Evolutionary theory and systematics: relationships between process and patterns

The relevance of the Modern Evolutionary Synthesis to the foundations of taxonomy (the construction of groups, both taxa and phyla) is reexamined. The nondimensional biological species concept, and not the multidimensional, taxonomic, species notion which is based on it, represents a culmination of an evolutionary understanding. It demonstrates how established evolutionary mechanisms acting on populations of sexually reproducing organisms provide the testable ontological basis of the species category. We question the ontology and epistemology of the phylogenetic or evolutionary species concept, and find it to be a fundamentally untenable one. We argue that at best, the phylogenetic species is a taxonomic species notion which is not a theoretical concept, and therefore should not serve as foundation for taxonomic theory in general, phylogenetics, and macroevolutionary reconstruction in particular. Although both evolutionary systematists and cladists are phylogeneticists, the reconstruction of the history of life is fundamentally different in these two approaches. We maintain that all method, including taxonomic ones, must fall out of well corroborated theory. In the case of taxonomic methodology the theoretical base must be evolutionary. The axiomatic assumptions that all phena, living and fossil, must be holophyletic taxa (species, and above), resulting from splitting events, and subsequently that evaluation of evolutionary change must be based on a taxic perspective codified by the Hennig ian taxonomic species notion, are not testable premises. We discuss the relationship between some biologically, and therefore taxonomically, significant patterns in nature, and the process dependence of these patterns. Process-free establishment of deductively tested “genealogies” is a contradiction in terms; it is impossible to “recover” phylogenetic patterns without the investment of causal and processual explanations of characters to establish well tested taxonomic properties of these (such as homologies, apomorphies, synapomorphies, or transformation series). Phylogenies of either characters or of taxa are historical-narrative explanations (H-N Es), based on both inductively formulated hypotheses and tested against objective, empirical evidence. We further discuss why construction of a “genealogy”, the alleged framework for “evolutionary reconstruction”, based on a taxic, cladistic outgroup comparison and a posteriori weighting of characters is circular. We define how the procedure called null-group comparison leads to the noncircular testing of the taxonomic properties of characters against which the group phylogenies must be tested. This is the only valid rooting procedure for either character or taxon evolution. While the Hennig -principle is obviously a sound deduction from the theory of descent, cladistic reconstruction of evolutionary history itself lacks a valid methodology for testing transformation hypotheses of both characters and species. We discuss why the paleontological method is part of comparative biology with a critical time dimension ana why we believe that an “ontogenetic method” is not valid. In our view, a merger of exclusive (causal and interactive, but best described as levels of organization) and inclusive (classificatory) hierarchies has not been accomplished by a taxic scheme of evolution advocated by some. Transformational change by its very nature is not classifiable in an inclusive hierarchy, and therefore no classification can fully reflect the causal and interactive chains of events constituting phylogeny, without ignoring and contradicting large areas of corroborated evolutionary theory. Attempts to equate progressive evolutionary change with taxic schemes by Haeckel were fundamentally flawed. His ideas found 19th century expression in a taxic perception of the evolutionary process (“phylogenesis”), a merger of typology, hierarchic and taxic notions of progress, all rooted in an ontogenetic view of phylogeny. The modern schemes of genealogical hierarchies, based on punctuation and a notion of “species” individuality, have yet to demonstrate that they hold promise beyond the Haeckel ian view of progressive evolution.     

Impact of genetic architecture on the relative rates of X versus autosomal adaptive substitution

Molecular evolutionary theory predicts that the ratio of autosomal to X-linked adaptive substitution (K(A)/K(x)) is primarily determined by the average dominance coefficient of beneficial mutations. Although this theory has profoundly influenced analysis and interpretation of comparative genomic data, its predictions are based upon two unverified assumptions about the genetic basis of adaptation. The theory assumes that 1) the rate of adaptively driven molecular evolution is limited by the availability of beneficial mutations, and 2) the scaling of evolutionary parameters between the X and the autosomes (e.g., the beneficial mutation rate, and the fitness effect distribution of beneficial alleles, per X-linked versus autosomal locus) is constant across molecular evolutionary timescales. Here, we show that the genetic architecture underlying bouts of adaptive substitution can influence both assumptions, and consequently, the theoretical relationship between K(A)/K(x) and mean dominance. Quantitative predictions of prior theory apply when 1) many genomically dispersed genes potentially contribute beneficial substitutions during individual steps of adaptive walks, and 2) the population beneficial mutation rate, summed across the set of potentially contributing genes, is sufficiently small to ensure that adaptive substitutions are drawn from new mutations rather than standing genetic variation. Current research into the genetic basis of adaptation suggests that both assumptions are plausibly violated. We find that the qualitative positive relationship between mean dominance and K(A)/K(x) is relatively robust to the specific conditions underlying adaptive substitution, yet the quantitative relationship between dominance and K(A)/K(x) is quite flexible and context dependent. This flexibility may partially account for the puzzlingly variable X versus autosome substitution patterns reported in the empirical evolutionary genomics literature. The new theory unites the previously separate analysis of adaptation using new mutations versus standing genetic variation and makes several useful predictions about the interaction between genetic architecture, evolutionary genetic constraints, and effective population size in determining the ratio of adaptive substitution between autosomal and X-linked genes.     

An Evolutionary Dynamics Model Adapted to Eusocial Insects

This study aims to better understand the evolutionary processes allowing species coexistence in eusocial insect communities. We develop a mathematical model that applies adaptive dynamics theory to the evolutionary dynamics of eusocial insects, focusing on the colony as the unit of selection. The model links long-term evolutionary processes to ecological interactions among colonies and seasonal worker production within the colony. Colony population dynamics is defined by both worker production and colony reproduction. Random mutations occur in strategies, and mutant colonies enter the community. The interactions of colonies at the ecological timescale drive the evolution of strategies at the evolutionary timescale by natural selection. This model is used to study two specific traits in ants: worker body size and the degree of collective foraging. For both traits, trade-offs in competitive ability and other fitness components allows to determine conditions in which selection becomes disruptive. Our results illustrate that asymmetric competition underpins diversity in ant communities.     

Ecological aspects of the evolutionary processes

Darwin in his On the Origin of species made it clear that evolutionary change depends on the combined action of two different causes, the first being the origin of genetically based phenotypic variation in the individual organisms comprising the population and the second being the action of selective agents of the external environment placing demands on the individual organisms. For over a century following Darwin, most evolutionists focused on the origin of inherited variation and its transmission; many workers continue to regard genetics to be the core of evolutionary theory. Far less attention has been given to the exact nature of the selective agents with most evolutionists still treating this cause imprecisely to the detriment of our understanding of both nomological and historical evolutionary theory. Darwin was vague in the meaning of his new concept of "Natural Selection," using it interchangeably as one of the causes for evolutionary change and as the final outcome (= evolutionary change). In 1930, natural selection was defined clearly as "non-random, differential reproduction of genes" by R. Fisher and J.B.S. Haldane which is a statement of the outcome of evolutionary process and which omits mention of the causes bringing about this change. Evolutionists quickly accepted this outcome definition of natural selection, and have used interchangeably selection both as a cause and as the result of evolutionary change, causing great confusion. Herein, the details will be discussed of how the external environment (i.e., the environment-phenotype interaction) serves as selective agents and exerts demands on the phenotypic organisms. Included are the concepts of fitness and of the components of fitness (= adaptations) which are respectively (a) survival, (b) direct reproductive and (c) indirect reproductive features. Finally, it will be argued that historical-narrative analyses of organisms, including classification and phylogenetic history, are possible only with a full understanding of nomological evolutionary theory and with functional/adaptive studies of the employed taxonomic features in addition to the standard comparative investigations.     

Costs and constraints conspire to produce honest signaling: insights from an ant queen pheromone

Signal costs and evolutionary constraints have both been proposed as ultimate explanations for the ubiquity of honest signaling, but the interface between these two factors is unclear. Here, I propose a pluralistic interpretation, and use game theory to demonstrate that evolutionary constraints determine whether signals evolve to be costly or cheap. Specifically, when the costs or benefits of signaling are strongly influenced by the sender's quality, low-cost signals evolve. The model reaffirms that cheap and costly signals can both be honest, and predicts that expensive signals should have more positive allometric slopes than cheap ones. The new framework is applied to an experimental study of an ant queen pheromone that honestly signals fecundity. Juvenile hormone was found to have opposing, dose-dependent effects on pheromone production and fecundity and was fatal at high doses, indicating that endocrine-mediated trade-offs preclude dishonesty. Several lines of evidence suggest that the realized cost of pheromone production may be nontrivial, and the antagonistic effects of juvenile hormone indicate the presence of significant evolutionary constraints. I conclude that the honesty of queen pheromones and other signals is likely enforced by both the cost of dishonesty and a suite of evolutionary constraints.     

Morphological interpretation of darwinism

The development of evolutionary theory requires the resolution of the problem of relationships between random and regular processes in historical development of biological systems. According to the theory of natural selection, ecological factors play a leading role in evolution. Variations are nondirectional, unpredictable, and provide chaotic diversity of variants, only some of which are potentially useful. However, based on random processes, new variants that are useful for organisms and remain adaptive significance in various ecological situations are infrequent. At the same time, morphology demonstrates certain evolutionary patterns. The morphological approach takes into account the role in evolution of structural features of organism and social systems and evolutionary significance of “constructive technologies,” which distinguish morphological interpretation of evolutionary processes. The constructive and evolutionary patterns revealed in biological systems provide the basis for morphological interpretation of the principle of natural selection: both natural and artificial selection is interaction between social systems (populations, ecosystems, biogeocoenoses) and organisms composing them.