African wild dogs test the 'survival of the fittest' paradigm

Charles Darwin first used the term 'survival of the fittest' in the 5th edition of The origin of species. A literal interpretation implies that predators will selectively prey upon the weakest members of a population. We demonstrate that this is true for African wild dogs hunting impala.     

The genetics and evolution of female choice

Darwin based his theory of evolution on the central theme of 'the struggle for existence', 'the preservation of favoured races', 'the survival of the fittest'. In addition to this 'natural selection', he proposed a second type of selection, sexual selection. 'This depends, not on a struggle for existence, but on a struggle between males for possession of the females; the result is not death to the unsuccessful competitor, but few or no offspring. Males would compete for females, and females would choose between males. Since Darwin's work, many examples of natural selection have been observed in nature and in laboratory experiments, and knowledge of genetics has given natural selective theory a sound basis. The theory of sexual selection through male competition has also been widely accepted. However, the theory of sexual selection through female choice has had a much more chequered passage.     

Extending Darwin’s analogy: Bridging differences in concepts of selection between farmers, biologists, and plant breeders

Darwin developed his theory of evolution based on an analogy between artificial selection by breeders of his day and “natural selection.” For Darwin, selection included what biologists came to see as being composed of (1) phenotypic selection of individuals based on phenotypic differences, and, when these are based on heritable genotypic differences, (2) genetic response between generations, which can result in (3) evolution (cumulative directional genetic response over generations). The use of the term “selection” in biology and plant breeding today reflects Darwin’s assumption—phenotypic selection is only biologically significant when it results in evolution. In contrast, research shows that small-scale, traditionally-based farmers select seed as part of an integrated production and consumption system in which selection is often not part of an evolutionary process, but is still useful to farmers. Extending Darwin’s analogy to farmers can facilitate communication between farmers, biologists, and plant breeders to improve selection and crop genetic resource conservation.     

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.     

Adaptations to sexual selection and sexual conflict: insights from experimental evolution and artificial selection

Artificial selection and experimental evolution document natural selection under controlled conditions. Collectively, these techniques are continuing to provide fresh and important insights into the genetic basis of evolutionary change, and are now being employed to investigate mating behaviour. Here, we focus on how selection techniques can reveal the genetic basis of post-mating adaptations to sexual selection and sexual conflict. Alteration of the operational sex ratio of adult Drosophila over just a few tens of generations can lead to altered ejaculate allocation patterns and the evolution of resistance in females to the costly effects of elevated mating rates. We provide new data to show how male responses to the presence of rivals can evolve. For several traits, the way in which males responded to rivals was opposite in lines selected for male-biased, as opposed to female-biased, adult sex ratio. This shows that the manipulation of the relative intensity of intra- and inter-sexual selection can lead to replicable and repeatable effects on mating systems, and reveals the potential for significant contemporary evolutionary change. Such studies, with important safeguards, have potential utility for understanding sexual selection and sexual conflict across many taxa. We discuss how artificial selection studies combined with genomics will continue to deepen our knowledge of the evolutionary principles first laid down by Darwin 150 years ago.     

Charles Robert Darwin and Alfred Russel Wallace: their dispute over the units of selection

Charles Darwin and Alfred Russel Wallace independently discovered the mechanism of natural selection for evolutionary change. However, they viewed the working of selection differently. For Darwin, selection was always focused on the benefit for the individual. For Wallace, selection was as much something of benefit for the group as for the individual. This difference is traced to their different background political–economic views, with Darwin in favor of Adam Smith's view of society and Wallace following Robert Owen in being a socialist.     

一个具选择、突变、迁移的群体的遗传差分模型

假设一个群体是由“单位点—双基因”的个体所组成的,在该群体内存在选择、突变、迁移、生死等效应的作用。本文给出了在上述假设下并满足:(1)世代重叠,选择、突变、迁移、生死等效应的作用均在世代遗传之间完成;(2)群体适当大,个体间交配随机,符合孟德尔式遗传;(3)没有任何意外的灾祸等约定的群体遗传的数学模型。通过模型分析,我们能够进一步用数学语言来解释一些生命现象。模型分析指出:虽然某些群体不满足Hardy-Weinberg定律所叙述的条件,但可能具有和Hardy-Weinberg定律的结论相似的结果。该文中还就几个主要参数的变化讨论了群体遗传和进化的某些性质,如平衡等。最后,我们给出了该模型的一个数值例子。     

Evolution by epigenesis: farewell to Darwinism, neo- and otherwise.

In the last 25 years, criticism of most theories advanced by Darwin and the neo-Darwinians has increased considerably, and so did their defense. Darwinism has become an ideology, while the most significant theories of Darwin were proven unsupportable. The critics advanced other theories instead of 'natural selection' and the survival of the fittest'. 'Saltatory ontogeny' and 'epigenesis' are such new theories proposed to explain how variations in ontogeny and novelties in evolution are created. They are reviewed again in the present essay that also tries to explain how Darwinians, artificially kept dominant in academia and in granting agencies, are preventing their acceptance. Epigenesis, the mechanism of ontogenies, creates in every generation alternative variations in a saltatory way that enable the organisms to survive in the changing environments as either altricial or precocial forms. The constant production of two such forms and their survival in different environments makes it possible, over a sequence of generations, to introduce changes and establish novelties--the true phenomena of evolution. The saltatory units of evolution remain far-from-stable structures capable of self-organization and self-maintenance (autopoiesis).     

Evolution of gametocyte sex ratios in malaria and related apicomplexan (protozoan) parasites

'Survival of the fittest' is usually interpreted to mean that natural selection favours genes that maximize their transmission to the next generation. Here, we discuss recent applications of this principle to the study of gametocyte sex ratios in malaria and other apicomplexan parasites. Sex ratios matter because they are an important determinant of fitness and transmission success -- and hence of disease epidemiology and evolution. Moreover, inbreeding rates can be estimated from gametocyte sex ratios. The sex ratio is also an excellent model trait for testing the validity of important components of what is being marketed as 'Darwinian medicine'.     

Darwin's artificial selection as an experiment

Darwin used artificial selection (ASN) extensively and variedly in his theorizing. Darwin used ASN as an analogy to natural selection; he compared artificial to natural varieties, hereditary variation in nature to that in the breeding farm; and he also compared the overall effectiveness of the two processes. Most historians and philosophers of biology have argued that ASN worked as an analogical field in Darwin's theorizing. I will argue rather that this provides a limited and somewhat muddled view of Darwinian science. I say "limited" because I will show that Darwin also used ASN as a complex experimental field. And I say "muddled" because, if we concentrate on the analogical role exclusively, we conceive Darwinian science as rather disconnected from contemporary conceptions of "good science". I will argue that ASN should be conceived as a multifaceted experiment. As a traditional experiment, ASN established the efficacy of Darwin's preferred cause: natural selection. As a non-traditional experiment, ASN disclosed the nature of a crucial element in Darwin's evolutionary mechanics: the nature of hereditary variation. Finally, I will argue that the experiment conception should help us make sense of Darwin's comments regarding the "monstrous" nature of domestic breeds traditionally considered to be problematic.     

On the problems of a closed marriage: celebrating Darwin 200

Darwin devoted much of his working life to the study of plant reproductive systems. He recognized that many of the intricacies of floral morphology had been shaped by natural selection in favour of outcrossing, and he clearly established the deleterious effects of self-fertilization on progeny. Although Darwin hypothesized the adaptive significance of self-fertilization under conditions of low mate availability, he held that a strategy of pure selfing would be strongly disadvantageous in the long term. Here, I briefly review these contributions to our understanding of plant reproduction. I then suggest that investigating two very different sexual systems, one in plants and the other in animals, would throw further light on the long-term implications of a commitment to reproduction exclusively by selfing.     

Species: kinds of individuals or individuals of a kind

The “species‐as‐individuals” thesis takes species, or taxa, to be individuals. On grounds of spatiotemporal boundedness, any biological entity at any level of complexity subject to evolutionary processes is an individual. From evolutionary theory flows an ontology that does not countenance universal properties shared by evolving entities. If austere nominalism were applied to evolving entities, however, nature would be reduced to a mere flow of passing events, each one a blob in space–time and hence of passing interest only. Yet if there is genuine biodiversity in nature, if nature is genuinely carved into species, and taxa, then these evolutionary entities will be genuinely differentiated into specific kinds, each species being one of its kind. Given the fact that evolving entities have un‐sharp boundaries, an appropriately weak, “non‐essentialist” concept of natural kind has to be invoked that does not allow for strong identity conditions. The thesis of this paper is that species are not either individuals, or natural kinds. Instead, species are complex wholes (particulars, individuals) that instantiate a specific natural kind. © The Willi Hennig Society 2007.     

Logical puzzles and scientific controversies: The nature of species,viruses and living organisms

In the past, biologists believed that species were stable and permanent entities and they viewed them as natural kinds which, like the chemical elements, exist in nature independently of any human conceptualization. After Darwin, biologists came to accept that species were the products of evolution and natural selection and were not immutable natural kinds.     

Four decades of opposing natural and human-induced artificial selection acting on Windermere pike (Esox lucius)

The ability of natural selection to drive local adaptation has been appreciated ever since Darwin. Whether human impacts can impede the adaptive process has received less attention. We tested this hypothesis by quantifying natural selection and harvest selection acting on a freshwater fish (pike) over four decades. Across the time series, directional natural selection tended to favour large individuals whereas the fishery targeted large individuals. Moreover, non-linear natural selection tended to favour intermediate sized fish whereas the fishery targeted intermediate sized fish because the smallest and largest individuals were often not captured. Thus, our results unequivocally demonstrate that natural selection and fishery selection often acted in opposite directions within this natural system. Moreover, the two selective factors combined to produce reduced fitness overall and stronger stabilizing selection relative to natural selection acting alone. The long-term ramifications of such human-induced modifications to adaptive landscapes are currently unknown and certainly warrant further investigation.     

Darwin's artificial selection analogy and the generic character of "phyletic" evolution

This paper examines the way Charles Darwin applied his domestic breeding analogy to the practical workings of species evolution: that application, it is argued, centered on Darwin's distinction between methodical and unconscious selection. Methodical selection, which entailed pairing particular individuals for mating purposes, represented conditions of strict geographic isolation, obviously useful for species multiplication (speciation). By contrast, unconscious selection represented an open landmass with a large breeding population. Yet Darwin held that this latter scenario, which often would include multiple ecological subdistricts and thus partial isolation, was better suited for speciation than were isolated conditions. At the same time, many passages in Darwin's writings that apparently portrayedphyletic evolution exclusively (these including references to unconscious selection), actually applied to speciation as well, for phyletic change in a single district could constitute a local manifestation of a larger common-descent pattern. This generic use of "phyletic" change was reflected in Darwin's deployment of the unconscious selection analogy in his published writings as well as in his dispute with Moritz Wagner over the necessity of geographic isolation for speciation. We can thus understand Darwin's otherwise puzzling declaration in The Origin of Species that unconscious selection was 'more important' than the methodical approach.     

Simulating natural selection as a culling mechanism on finite populations with the hawk-dove game

The behaviors of individuals and species are often explained in terms of evolutionary stable strategies (ESSs). The analysis of ESSs determines which, if any, combinations of behaviors cannot be invaded by alternative strategies. Two assumptions required to generate an ESS (i.e., an infinite population and payoffs described only on the average) do not hold under natural conditions. Previous experiments indicated that under more realistic conditions of finite populations and stochastic payoffs, populations may evolve in trajectories that are unrelated to an ESS, even in very simple games. The simulations offered here extend earlier research by employing truncation selection with random parental selection in a hawk-dove game. Payoffs are determined in pairwise contests using either the expected outcome, or the result of a random variable. In each case, however, the mean fraction of hawks over many generations and across many independent trials does not conform to the expected ESS. Implications of these results and philosophical underpinnings of ESS theory are offered.     

The Origin of Species by Means of Mathematical Modelling

Darwin described biological species as groups of morphologically similar individuals. These groups of individuals can split into several subgroups due to natural selection, resulting in the emergence of new species. Some species can stay stable without the appearance of a new species, some others can disappear or evolve. Some of these evolutionary patterns were described in our previous works independently of each other. In this work we have developed a single model which allows us to reproduce the principal patterns in Darwin’s diagram. Some more complex evolutionary patterns are also observed. The relation between Darwin’s definition of species, stated above, and Mayr’s definition of species (group of individuals that can reproduce) is also discussed.     

Evolutionary Optimality of Body Features. Species-Specific Lifespan as a Product of Evolution

Biophysics - According to Darwin’s theory of natural selection, the fittest population survives in the course of competition for a food resource. The values of the features of individuals of...     

Darwin's principles of divergence and natural selection: Why Fodor was almost right

Darwin maintained that the principles of natural selection and divergence were the “keystones” of his theory. He introduced the principle of divergence to explain a fundamental feature of living nature: that organisms cluster into hierarchical groups, so as to be classifiable in the Linnaean taxonomic categories of variety, species, genus, and so on. Darwin’s formulation of the principle of divergence, however, induces many perplexities. In his Autobiography, he claimed that he had neglected the problem of divergence in his Essay of 1844 and only solved it in a flash during a carriage ride in the 1850s; yet he does seem to have stated the problem in the Essay and provided the solution. This initial conundrum sets three questions I wish to pursue in this essay: (1) What is the relationship of the principle of divergence to that of natural selection? Is it independent of selection, derivative of selection, or a type of selection, perhaps comparable to sexual selection? (2) What is the advantage of divergence that the principle implies—that is, why is increased divergence beneficial in the struggle for life? And (3) What led Darwin to believe he had discovered the principle only in the 1850s? The resolution of these questions has implications for Darwin’s other principle, natural selection, and permits us to readjust the common judgment made about Jerry Fodor’s screed against that latter principle.