Darwin without borders? Looking at ‘generalised Darwinism’ through the prism of the ‘hourglass model’

This article critically analyzes the arguments of the ‘generalized Darwinism’ recently proposed for the analysis of social-economical systems. We argue that ‘generalized Darwinism’ is both restrictive and empty. It is restrictive because it excludes alternative (non-selectionist) evolutionary mechanisms such as orthogenesis, saltationism and mutationism without any examination of their suitability for modeling socio-economic processes and ignoring their important roles in the development of contemporary evolutionary theory. It is empty, because it reduces Darwinism to an abstract triple-principle scheme (variation, selection and inheritance) thus ignoring the actual structure of Darwinism as a complex and dynamic theoretical structure inseparable from a very detailed system of theoretical constraints. Arguing against ‘generalised Darwinism’ we present our vision of the history of evolutionary biology with the help of the ‘hourglass model’ reflecting the internal dynamic of competing theories of evolution.     

Don’t Call it “Darwinism”

Evolutionary biology owes much to Charles Darwin, whose discussions of common descent and natural selection provide the foundations of the discipline. But evolutionary biology has expanded well beyond its foundations to encompass many theories and concepts unknown in the 19th century. The term “Darwinism” is, therefore, ambiguous and misleading. Compounding the problem of “Darwinism” is the hijacking of the term by creationists to portray evolution as a dangerous ideology—an “ism”—that has no place in the science classroom. When scientists and teachers use “Darwinism” as synonymous with evolutionary biology, it reinforces such a misleading portrayal and hinders efforts to present the scientific standing of evolution accurately. Accordingly, the term “Darwinism” should be abandoned as a synonym for evolutionary biology.     

Alfred Russel Wallace deserves better

During 2009, while we were celebrating Charles Darwin and his The origin of species, sadly, little was said about the critical contribution of Alfred Russel Wallace (1823–1913) to the development of the theory of evolution. Like Darwin, he was a truly remarkable nineteenth century intellect and polymath and, according to a recent book by Roy Davies (The Darwin conspiracy: origins of a scientific crime), he has a stronger claim to the Theory of Evolution by Natural Selection than has Darwin. Here we present a critical comparison between the contributions of the two scientists. Sometimes referred to as ‘The other beetle-hunter’ and largely neglected for many decades, Wallace had a far greater experience of collecting and investigating animals and plants from their native habitats than had Darwin. He was furthermore much more than a pioneer biogeographer and evolutionary theorist, and also made contributions to anthropology, ethnography, geology, land reform and social issues. However, being a more modest, self-deprecating man than Darwin, and lacking the latter’s establishment connections, Wallace’s contribution to the theory of evolution was not given the recognition it deserved and he was undoubtedly shabbily treated at the time. It is time that Wallace’s relationship with Darwin is reconsidered in preparation for 2013, the centenary of Wallace’s death, and he should be recognized as at least an equal in the Wallace-Darwin theory of evolution.     

The expanded evolutionary synthesis—a response to Godfrey-Smith,Haig, and West-Eberhard

In responding to three reviews of Evolution in Four Dimensions (Jablonka and Lamb, 2005, MIT Press), we briefly consider the historical background to the present genecentred view of evolution, especially the way in which Weismann’s theories have influenced it, and discuss the origins of the notion of epigenetic inheritance. We reaffirm our belief that all types of hereditary information—genetic, epigenetic, behavioural and cultural—have contributed to evolutionary change, and outline recent evidence, mainly from epigenetic studies, that suggests that non-DNA heritable variations are not rare and can be quite stable. We describe ways in which such variations may have influenced evolution. The approach we take leads to broader definitions of terms such as ‘units of heredity’, ‘units of evolution’, and ‘units of selection’, and we maintain that ‘information’ can be a useful concept if it is defined in terms of its effects on the receiver. Although we agree that evolutionary theory is not undergoing a Kuhnian revolution, the incorporation of new data and ideas about hereditary variation, and about the role of development in generating it, is leading to a version of Darwinism that is very different from the gene-centred one that dominated evolutionary thinking in the second half of the twentieth century.     

Sources for Imanishi Kinji’s views of sociality and evolutionary outcomes

Prior to the contribution of genetics or the modern evolutionary synthesis (MES) to natural selection theory, social ecologists searched for factors in addition to natural selection that could influence species change. The idea that sociality, not just biology, was important in determining evolutionary outcomes was prevalent in research in social ecology in the 1920s and 1930s. The influence of ‘tradition’ (or the transmission of learned behaviours between generations) and the view that animals are active in selecting their own environments, rather than passive organisms acted upon by chance, were given as much attention as natural selection theory in European ecology, while animal aggregation and cooperation studies were pursued in America. Imanishi Kinji’s personal library and his scientific notes and papers reveal that he was well aware of this literature and had been profoundly influenced by these earlier viewpoints prior to writing his view of nature in his first book, Seibutsu no Sekai (The World of Living Things, 1941). Evidence is presented to show that he developed his theories based partly on early western debates in social ecology while finding inspiration and a way to express his views in the writings of philosopher Nishida Kitarō and, perhaps, General J C Smuts. One of Imanishi’s lasting contributions is in the demonstrated results of over 40 years of subsequent ecological and ethological research by Imanishi and those trained by him that maintained the broader viewpoints on evolution that had been dropped from the western corpus of research by the 1950s. The current attempt to again get beyond natural selection theory is reflected in debates surrounding genetic and cultural evolution of cooperation, the biology of ‘traditions’ and the idea of ‘culture’ in animal societies. Imanishi Kinji is the Japanese name order, with family name first. Other Japanese names in the text are also written with family name first. A modified version of this paper appeared in Japanese in Seibutsu Kagaku, Vol. 57 No. 3, April 2006, pp 142–149.     

Tantalizing Tortoises and the Darwin-Galápagos Legend

During his historic Galápagos visit in 1835, Darwin spent nine days making scientific observations and collecting specimens on Santiago (James Island). In the course of this visit, Darwin ascended twice to the Santiago highlands. There, near springs located close to the island’s summit, he conducted his most detailed observations of Galápagos tortoises. The precise location of these springs, which has not previously been established, is here identified using Darwin’s own writings, satellite maps, and GPS technology. Photographic evidence from excursions to the areas where Darwin climbed, including repeat photography over a period of four decades, offers striking evidence of the deleterious impact of feral mammals introduced after Darwin’s visit. Exploring the impact that Darwin’s Santiago visit had on his thinking – especially focusing on his activities in the highlands – raises intriguing questions about the depth of his understanding of the evolutionary evidence he encountered while in the Galápagos. These questions and related insights provide further evidence concerning the timing of Darwin’s conversion to the theory of evolution, which, despite recent claims to the contrary, occurred only after his return to England.     

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.     

In the shadow of Darwin: Anton de Bary’s origin of myxomycetology and a molecular phylogeny of the plasmodial slime molds

In his Origin of Species (John Murray, London, 1859), Charles Darwin described the theory of descent with modification by means of natural selection and postulated that all life may have evolved from one or a few simple kinds of organisms. However, Darwin’s concept of evolutionary change is entirely based on observations of populations of animals and plants. He briefly mentioned ‘lower algae’, but ignored amoebae, bacteria and other micro-organisms. In 1859, Anton de Bary, the founder of mycology and plant pathology, published a seminal paper on the biology and taxonomy of the plasmodial slime molds (myxomycetes). These heterotrophic protists are known primarily as a large composite mass, the plasmodium, in which single nuclei are suspended in a common ‘naked’ cytoplasm that is surrounded by a plasma membrane. Here we summarize the contents of de Bary’s 1859 publication and highlight the significance of this scientific classic with respect to the establishment of the kingdom Protoctista (protists such as amoebae), the development of the protoplasmic theory of the cell, the introduction of the concept of symbiosis and the rejection of the dogma of spontaneous generation. We describe the life cycle of the myxomycetes, present new observations on the myxamoebae and propose a higher-order phylogeny based on elongation factor-1 alpha gene sequences. Our results document the congruence between the morphology-based taxonomy of the myxomycetes and molecular data. In addition, we show that free-living amoebae, common protists in the soil, are among the closest living relatives of the myxomycetes and conclude that de Bary’s ‘Amoeba-hypothesis’ on the evolutionary origin of the plasmodial slime molds may have been correct.     

Evolutionary ecologyin silico: Does mathematical modelling help in understanding ‘generic’ trends?

Motivated by the results of recent laboratory experiments, as well as many earlier field observations, that evolutionary changes can take place in ecosystems over relatively short ecological time scales, several ‘unified’ mathematical models of evolutionary ecology have been developed over the last few years with the aim of describing the statistical properties of data related to the evolution of ecosystems. Moreover, because of the availability of sufficiently fast computers, it has become possible to carry out detailed computer simulations of these models. For the sake of completeness and to put these recent developments in perspective, we begin with a brief summary of some older models of ecological phenomena and evolutionary processes. However, the main aim of this article is to review critically these ‘unified’ models, particularly those published in the physics literature, in simple language that makes the new theories accessible to a wider audience     

The molecular basis of speciation: from patterns to processes,rules to mechanisms

The empirical study of speciation has brought us closer to unlocking the origins of life’s vast diversity. By examining recently formed species, a number of general patterns, or rules, become apparent. Among fixed differences between species, sexual genes and traits are one of the most rapidly evolving and novel functional classes, and premating isolation often develops earlier than postmating isolation. Among interspecific hybrids, sterility evolves faster than inviability, the X-chromosome has a greater effect on incompatibilities than autosomes, and hybrid dysfunction affects the heterogametic sex more frequently than the homogametic sex (Haldane’s rule). Haldane’s rule, in particular, has played a major role in reviving interest in the genetics of speciation. However, the large genetic and reproductive differences between taxa and the multi-factorial nature of each rule have made it difficult to ascribe general mechanisms. Here, we review the extensive progress made since Darwin on understanding the origin of species. We revisit the rules of speciation, regarding them as landmarks as species evolve through time. We contrast these ‘rules’ of speciation to ‘mechanisms’ of speciation representing primary causal factors ranging across various levels of organization—from genic to chromosomal to organismal. To explain the rules, we propose a new ‘hierarchical faster-sex’ theory: the rapid evolution of sex and reproduction-related (SRR) genes (faster-SRR evolution), in combination with the preferential involvement of the X-chromosome (hemizygous X-effects) and sexually selected male traits (faster-male evolution). This unified theory explains a comprehensive set of speciation rules at both the prezyotic and postzygotic levels and also serves as a cohesive alternative to dominance, composite, and recent genomic conflict interpretations of Haldane’s rule.     

Thermodynamical interpretation of evolutionary dynamics on a fitness landscape in a evolution reactor,I

A theory for describing evolution as adaptive walks by a finite population with M walkers (M ≥ 1) on an anisotropic Mt. Fuji-type fitness landscape is presented, from a thermodynamical point of view. Introducing the ‘free fitness’ as the sum of a fitness term and an entropy term and ‘evolutionary force’ as the gradient of free fitness on a fitness coordinate, we demonstrate that the behavior of these theoretical walkers is almost consistent with the thermodynamical schemes. The major conclusions are as follows: (1) an adaptive walk (=evolution) is driven by an evolutionary force in the direction in which free fitness increases; (2) the expectation of the climbing rate obeys an equation analogous to the Einstein relation in Brownian motion; (3) the standard deviation of the climbing rate is a quantity analogous to the mean thermal energy of a particle, kT (×constant). In addition, on the interpretation that the walkers climb the landscape by absorbing ‘fitness information’ from the surroundings, we succeeded in quantifying the fitness information and formulating a macroscopic scheme from an informational point of view.     

Variation in performance on two grape cultivars within and among populations of grape Phylloxera from wild and cultivated habitats

When an indigenous insect becomes a pest, comparisons of performance of pest and non-pest populations on crop plants and of genetic variation in that performance may provide insight into the evolution of pest populations. To measure such genetic variation, 8–15 clones of the grape phylloxera (Daktulosphaira vitifoliae Fitch) were collected from wild grapevines in each of 3 geographically isolated sites (populations) and from commercial vineyards in northern California. A complete life table was made for clonal replicates from populations collected from wild grapevines on each of two commercial grape cultivars, the susceptibleVitis vinifera (L.) cultivar Cabernet Sauvignon, and the phylloxera-resistant rootstock ‘AxR # 1’. Variation in mean performance on these two hosts was partitioned among clones within collection sites and among sites. Performance measures included an individual analog to the intrinsic rate of increase (r), age at first oviposition, fecundity in the first ten days of reproduction, total fecundity, and longevity. The overall performance of phylloxera from the wild grapevines on the resistant cultivar AxR # 1 was greater than or equal to that on the susceptible cultivar Cabernet Sauvignon. There was significant variation among clones within populations from wild grapes in the rate of increase on ‘AxR # 1’ and marginally significant clonal variation in some of the component paramters. There was no significant variation among clones within populations on ‘Cabernet Sauvignon’ and no significant differences between populations on either crop in any trait. In a second experiment we compared the relative performance of 15–17 clones from wild grapevines and from commercial vineyards when reared on ‘Cabernet Sauvignon’ and ‘AxR # 1’. Phylloxera from commercial vineyards had much higher overall performance on ‘Cabernet Sauvignon’ than did phylloxera from the wild grapevines. Phylloxera from the commercial vineyard also had higher performance on ‘Cabernet Sauvignon’ than on ‘AxR′ 1’ but the performance of the phylloxera from wild and commercial grapes did not differ on ‘AxR # 1’. Our results show that there is genetic variation in traits related to performance on a resistant rootstock within these indigenous non-pest populations of phylloxera, but not among them. The pattern of performance of pest and non-pest populations on two commercial cultivars suggests that current levels of phylloxera performance on crop cultivars are the result of adaptation to those cultivars which has occurred while phylloxera has been associated with viticulture. Implications of these results for understanding the recent adaptation of phylloxera to ‘AxR # 1’ in California are also discussed.     

The Role of Intuitive Ontologies in Scientific Understanding – the Case of Human Evolution

Psychological evidence suggests that laypeople understand the world around them in terms of intuitive ontologies which describe broad categories of objects in the world, such as ‘person’, ‘artefact’ and ‘animal’. However, because intuitive ontologies are the result of natural selection, they only need to be adaptive; this does not guarantee that the knowledge they provide is a genuine reflection of causal mechanisms in the world. As a result, science has parted ways with intuitive ontologies. Nevertheless, since the brain is evolved to understand objects in the world according to these categories, we can expect that they continue to play a role in scientific understanding. Taking the case of human evolution, we explore relationships between intuitive ontological and scientific understanding. We show that intuitive ontologies not only shape intuitions on human evolution, but also guide the direction and topics of interest in its research programmes. Elucidating the relationships between intuitive ontologies and science may help us gain a clearer insight into scientific understanding.     

Size doesn’t matter: towards a more inclusive philosophy of biology

Philosophers of biology, along with everyone else, generally perceive life to fall into two broad categories, the microbes and macrobes, and then pay most of their attention to the latter. ‘Macrobe’ is the word we propose for larger life forms, and we use it as part of an argument for microbial equality. We suggest that taking more notice of microbes – the dominant life form on the planet, both now and throughout evolutionary history – will transform some of the philosophy of biology’s standard ideas on ontology, evolution, taxonomy and biodiversity. We set out a number of recent developments in microbiology – including biofilm formation, chemotaxis, quorum sensing and gene transfer – that highlight microbial capacities for cooperation and communication and break down conventional thinking that microbes are solely or primarily single-celled organisms. These insights also bring new perspectives to the levels of selection debate, as well as to discussions of the evolution and nature of multicellularity, and to neo-Darwinian understandings of evolutionary mechanisms. We show how these revisions lead to further complications for microbial classification and the philosophies of systematics and biodiversity. Incorporating microbial insights into the philosophy of biology will challenge many of its assumptions, but also give greater scope and depth to its investigations.     

Darwinian adaptation,population genetics and the streetcar theory of evolution

 This paper investigates the problem of how to conceive a robust theory of phenotypic adaptation in non-trivial models of evolutionary biology. A particular effort is made to develop a foundation of this theory in the context of n-locus population genetics. Therefore, the evolution of phenotypic traits is considered that are coded for by more than one gene. The potential for epistatic gene interactions is not a priori excluded. Furthermore, emphasis is laid on the intricacies of frequency-dependent selection. It is first discussed how strongly the scope for phenotypic adaptation is restricted by the complex nature of ‘reproduction mechanics’ in sexually reproducing diploid populations. This discussion shows that one can easily lose the traces of Darwinism in n-locus models of population genetics. In order to retrieve these traces, the outline of a new theory is given that I call ‘streetcar theory of evolution’. This theory is based on the same models that geneticists have used in order to demonstrate substantial problems with the ‘adaptationist programme’. However, these models are now analyzed differently by including thoughts about the evolutionary removal of genetic constraints. This requires consideration of a sufficiently wide range of potential mutant alleles and careful examination of what to consider as a stable state of the evolutionary process. A particular notion of stability is introduced in order to describe population states that are phenotypically stable against the effects of all mutant alleles that are to be expected in the long-run. Surprisingly, a long-term stable state can be characterized at the phenotypic level as a fitness maximum, a Nash equilibrium or an ESS. The paper presents these mathematical results and discusses – at unusual length for a mathematical journal – their fundamental role in our current understanding of evolution. Received 22 April 1994; received in revised form 10 July 1995     

Darwin’s contributions to genetics

Darwin’s contributions to evolutionary biology are well known, but his contributions to genetics are much less known. His main contribution was the collection of a tremendous amount of genetic data, and an attempt to provide a theoretical framework for its interpretation. Darwin clearly described almost all genetic phenomena of fundamental importance, such as prepotency (Mendelian inheritance), bud variation (mutation), heterosis, reversion (atavism), graft hybridization (Michurinian inheritance), sex-limited inheritance, the direct action of the male element on the female (xenia and telegony), the effect of use and disuse, the inheritance of acquired characters (Lamarckian inheritance), and many other observations pertaining to variation, heredity and development. To explain all these observations, Darwin formulated a developmental theory of heredity — Pangenesis — which not only greatly influenced many subsequent theories, but also is supported by recent evidence.     

Thermodynamical interpretation of evolutionary dynamics on a fitness landscape in an evolution reactor,II

In our previous report [Aita, T., Morinaga, S., Hosimi, Y., 2004. Thermodynamical interpretation of evolutionary dynamics on a fitness landscape in an evolution reactor I. Bull. Math. Biol. 66, 1371–1403], an analogy between thermodynamics and adaptive walks on a Mt. Fuji-type fitness landscape in an artificial selection system was presented. Introducing the ‘free fitness’ as the sum of a fitness term and an entropy term and ‘evolutionary force’ as the gradient of free fitness on a fitness coordinate, we demonstrated that the adaptive walk (=evolution) is driven by the evolutionary force in the direction in which free fitness increases. In this report, we examine the effect of various modifications of the original model on the properties of the adaptive walk. The modifications were as follows: first, mutation distance d was distributed obeying binomial distribution; second, the selection process obeyed the natural selection protocol; third, ruggedness was introduced to the landscape according to the NK model; fourth, a noise was included in the fitness measurement. The effect of each modification was described in the same theoretical framework as the original model by introducing ‘effective’ quantities such as the effective mutation distance or the effective screening size.     

A model for the emergence of adaptive subsystems

We investigate the interaction of learning and evolution in a changing environment. A stable learning capability is regarded as an emergent adaptive system evolved by natural selection of genetic variants. We consider the evolution of an asexual population. Each genotype can have ‘fixed’ and ‘flexible’ alleles. The former express themselves as synaptic connections that remain unchanged during ontogeny and the latter as synapses that can be adjusted through a learning algorithm. Evolution is modelled using genetic algorithms and the changing environment is represented by two optimal synaptic patterns that alternate a fixed number of times during the ‘life’ of the individuals. The amplitude of the change is related to the Hamming distance between the two optimal patterns and the rate of change to the frequency with which both exchange roles. This model is an extension of that of Hinton and Nowlan in which the fitness is given by a probabilistic measure of the Hamming distance to the optimum. We find that two types of evolutionary pathways are possible depending upon how difficult (costly) it is to cope with the changes of the environment. In one case the population loses the learning ability, and the individuals inherit fixed synapses that are optimal in only one of the environmental states. In the other case a flexible subsystem emerges that allows the individuals to adapt to the changes of the environment. The model helps us to understand how an adaptive subsystem can emerge as the result of the tradeoff between the exploitation of a congenital structure and the exploration of the adaptive capabilities practised by learning.