Genetical Theory Of Natural Selection Research Articles

R. A. Fisher on J. A. Cobb's The problem of the sex-ratio

The logic of the rarer-sex effect, concerning how natural selection acts to balance the sex ratio among newborns, was long supposed to have originated with Ronald Aylmer Fisher in his 1930 book The genetical theory of natural selection . However, the principle is now understood to have originated with John Austin Cobb in his 1914 paper ‘The problem of the sex-ratio’. Fisher did not provide a citation of Cobb's sex-ratio paper, and it has been unclear whether he was aware of its existence. Here, I show that Fisher was indeed aware of Cobb's paper in 1930, as revealed by him citing it elsewhere that same year. Fisher's willingness to highlight Cobb's sex-ratio work lends support to the view that his failure to mention it in his book reflects the less stringent citation standards of the time rather than an attempt to deceive readers as to the provenance of the rarer-sex effect.

Revisiting the number of self-incompatibility alleles in finite populations: From old models to new results.

Under gametophytic self-incompatibility (GSI), plants are heterozygous at the self-incompatibility locus (S-locus) and can only be fertilized by pollen with a different allele at that locus. The last century has seen a heated debate about the correct way of modelling the allele diversity in a GSI population that was never formally resolved. Starting from an individual-based model, we derive the deterministic dynamics as proposed by Fisher (The genetical theory of natural selection - A complete, Variorum edition, Oxford University Press, 1958) and compute the stationary S-allele frequency distribution. We find that the stationary distribution proposed by Wright (Evolution, 18, 609, 1964) is close to our theoretical prediction, in line with earlier numerical confirmation. Additionally, we approximate the invasion probability of a new S-allele, which scales inversely with the number of resident S-alleles. Lastly, we use the stationary allele frequency distribution to estimate the population size of a plant population from an empirically obtained allele frequency spectrum, which complements the existing estimator of the number of S-alleles. Our expression of the stationary distribution resolves the long-standing debate about the correct approximation of the number of S-alleles and paves the way for new statistical developments for the estimation of the plant population size based on S-allele frequencies.

R.A. Fisher, eugenics, and the campaign for family allowances in interwar Britain.

Ronald Aylmer Fisher (1890-1962) is today remembered as a giant of twentieth-century statistics, genetics and evolutionary theory. Alongside his influential scientific contributions, he was also, throughout the interwar years, a prominent figure within Britain's eugenics movement. This essay provides a close examination of his eugenical ideas and activities, focusing particularly upon his energetic advocacy of family allowances, which he hoped would boost eugenic births within the more 'desirable' middle and upper classes. Fisher's proposals, which were grounded in his distinctive explanation for the decay of civilizations throughout human history, enjoyed support from some influential figures in Britain's Eugenics Society and beyond. The ultimate failure of his campaign, though, highlights tensions both between the eugenics and family allowances movements, and within the eugenics movement itself. I show how these social and political movements represented a crucial but heretofore overlooked context for the reception of Fisher's evolutionary masterwork of 1930, The Genetical Theory of Natural Selection, with its notorious closing chapters on the causes and cures of national and racial decline.

Parental investment in the Columbian ground squirrel: empirical tests of sex allocation models

Parental allocation of resources into male or female offspring and differences in the balance of offspring sexes in natural populations are central research topics in evolutionary ecology. Fisher (Fisher, R. A. 1930. The genetical theory of natural selection, Clarendon Press, Oxford, UK) identified frequency-dependent selection as the mechanism responsible for an equal investment in the sexes of offspring at the end of parental care. Three main theories have been proposed for explaining departures from Fisherian sex ratios in light of variation in environmental (social) and individual (maternal condition) characteristics. The Trivers-Willard model (Trivers, R., and D. Willard. 1973. Natural selection of parental ability to vary the sex ratio of offspring. Science 179:90-92) of male-biased sex allocation by mothers in the best body condition is based on the competitive ability of male offspring for future access to mates and thus superior reproduction. The local resource competition model is based on competitive interactions in matrilines, as occur in many mammal species, where producing sons reduces future intrasexual competition with daughters. A final model invokes advantages of maintaining matrilines for philopatric females, despite any increased competition among females. We used 29 yr of pedigree and demographic data to evaluate these hypotheses in the Colombian ground squirrel (Urocitellus columbianus), a semisocial species characterized by strong female philopatry. Overall, male offspring were heavier than female offspring at birth and at weaning, suggesting a higher production cost. With more local kin present, mothers in the best condition biased their offspring sex ratio in favor of males, and mothers in poor condition biased offspring sex ratio in favor of females. Without co-breeding close kin, the pattern was reversed, with mothers in the best condition producing more daughters, and mothers in poor condition producing more sons. Our results do not provide strong support for any of the single-factor models of allocation to the sexes of offspring, but rather suggest combined influences of relative maternal condition and matriline dominance on offspring sex ratio.

No genetic contribution to variation in human offspring sex ratio: a total population study of 4.7 million births.

The ratio of males to females among an individual's offspring at birth (offspring sex ratio) has long been of great interest to evolutionary biologists. The human offspring sex ratio is around 1 : 1 and is understood primarily in terms of Fisher's principle (R. A. Fisher, The genetical theory of natural selection, 1930), which is based on the insight that in a population with an unequal sex ratio, each individual of the rarer sex will on average have greater reproductive value than each individual of the more common sex. Accordingly, individuals genetically predisposed to produce the rarer sex will tend to have greater fitness and thus genes predisposing to bearing that sex will increase in frequency until the population sex ratio approaches 1 : 1. An assumption of this perspective is that individuals' offspring sex ratio is heritable. However, the heritability in humans remains remarkably uncertain, with inconsistent findings and important power limitations of existing studies. To address this persistent uncertainty, we used data from the entire Swedish-born population born 1932 or later, including 3 543 243 individuals and their 4 753 269 children. To investigate whether offspring sex ratio is influenced by genetic variation, we tested the association between individuals' offspring's sex and their siblings' offspring's sex (n pairs = 14 015 421). We estimated that the heritability for offspring sex ratio was zero, with an upper 95% confidence interval of 0.002, rendering Fisher's principle and several other existing hypotheses untenable as frameworks for understanding human offspring sex ratio.

Estimation of Genetic Variance in Fitness, and Inference of Adaptation, When Fitness Follows a Log-Normal Distribution.

Additive genetic variance in relative fitness (σA2(w)) is arguably the most important evolutionary parameter in a population because, by Fisher's fundamental theorem of natural selection (FTNS; Fisher RA. 1930. The genetical theory of natural selection. 1st ed. Oxford: Clarendon Press), it represents the rate of adaptive evolution. However, to date, there are few estimates of σA2(w) in natural populations. Moreover, most of the available estimates rely on Gaussian assumptions inappropriate for fitness data, with unclear consequences. "Generalized linear animal models" (GLAMs) tend to be more appropriate for fitness data, but they estimate parameters on a transformed ("latent") scale that is not directly interpretable for inferences on the data scale. Here we exploit the latest theoretical developments to clarify how best to estimate quantitative genetic parameters for fitness. Specifically, we use computer simulations to confirm a recently developed analog of the FTNS in the case when expected fitness follows a log-normal distribution. In this situation, the additive genetic variance in absolute fitness on the latent log-scale (σA2(l)) equals (σA2(w)) on the data scale, which is the rate of adaptation within a generation. However, due to inheritance distortion, the change in mean relative fitness between generations exceeds σA2(l) and equals (exp⁡(σA2(l))-1). We illustrate why the heritability of fitness is generally low and is not a good measure of the rate of adaptation. Finally, we explore how well the relevant parameters can be estimated by animal models, comparing Gaussian models with Poisson GLAMs. Our results illustrate 1) the correspondence between quantitative genetics and population dynamics encapsulated in the FTNS and its log-normal-analog and 2) the appropriate interpretation of GLAM parameter estimates.

Some Contemporary Elaborations of Darwinian Themes

Evolutionary biology is intrinsically historical. Evolutionary theorists tend to be much more interested in both historical and conceptual/philosophical issues than their counterparts in most other disciplines. Invoking the authority of ‘‘patron saint’’ Darwin is part of their standard repertoire. Historians and philosophers, including ethical theorists, have major stakes in the ‘‘Darwin industry’’— although, somewhat disturbingly, most biomedical and environmental ethicists go about their business without much concern for recent advances in the philosophy of biology. Discussing ethics in The Descent of Man, Darwin (1871) referred to the writings of moral philosophers including Hume, Kant, and John Stuart Mill. In this issue, John Mizzoni argues that Darwin’s views did not lead him to an unfamiliar and eccentric view about the nature and content of ethics, but fitted rather well with the most representative (past and contemporary) normative ethical theories: virtue ethics, natural law ethics, social contract ethics, utilitarian ethics, deontological ethics, and care ethics. Robin Owen’s historical essay on Ronald Fisher and social insects deals with an issue that Darwin had avoided discussing in detail: the evolution of (‘‘harmonious’’) social insect colonies (in contrast to human societies that exhibit intra-communal conflict). In The Genetical Theory of Natural Selection, Fisher (1930) developed a verbal model of the evolution of eusociality by connecting selection acting on fecundity with the sterility of workers. Owen shows that Fisher’s development of the model was strongly influenced by Major Leonard Darwin, one of Charles Darwin’s sons, and argues that the Fisher-Darwin model ‘‘presages almost exactly’’ the independently derived mathematical model recently proposed by Martin Nowak, E. O. Wilson, and others. Still on the subject of eusociality, Klaus Stiefel proposes that a dishonest signaling system can be evolutionarily stable in eusocial animal societies if the amount of dishonesty is balanced by the chance of nonreproductive workers to advance to the reproductive caste in the future. He expresses this trade-off in a modified form of Hamilton’s rule, distinguishing between the real and the perceived cost of an altruistic act, and between the real and the perceived genetic relatedness between colony members. He also argues that the vertebrate neuromodulator oxytocin (likely integrated with a number of other functions related to social bonding) could serve as an internal representation of the perceived cost of an altruistic act and of perceived relatedness, and concludes with a discussion of honesty in signaling and a comparison between vertebrate and insect eusociality. Two other articles deal with different aspects of collaboration. Frans Roes discusses female inheritance and the ‘‘male retention hypothesis.’’ Permanent groups (groups with no inherent limit on group longevity) exist in several species because over generations members share important interests. Given the association between cooperation and degree of relatedness (Hamilton), he suggests that a collective interest is more likely to be achieved when members show a higher degree of relatedness. He then argues that if membership is inherited by only one sex, and this is the female sex, this results in a higher degree of relatedness between group members than when membership is inherited by both sexes, or by males only (as indeed found in the overwhelming majority of insects, fish, birds, and mammals living in permanent groups). Regarding humans, Roes also ponders whether moralizing Gods raise paternity confidence, without reaching firm conclusions in W. Callebaut (&) KLI Institute, Klosterneuburg, Austria e-mail: werner.callebaut@kli.ac.at

Autonomous-Statistical Explanations and Natural Selection

Shapiro and Sober ([2007]) claim that Walsh, Ariew, Lewens, and Matthen (henceforth WALM) give a mistaken, a priori defense of natural selection and drift as epiphenomenal. Contrary to Shapiro and Sober’s claims, we first argue that WALM’s explanatory doctrine does not require a defense of epiphenomenalism. We then defend WALM’s explanatory doctrine by arguing that the explanations provided by the modern genetical theory of natural selection are ‘autonomous-statistical explanations’ analogous to Galton’s explanation of reversion to mediocrity and an explanation of the diffusion ofgases. We then argue that whereas Sober’s theory of forces is an adequate description of Darwin’s theory, WALM’s explanatory doctrine is required to understand how themodern genetical theory of natural selection explains large-scale statistical regularities. 1 Introduction 2 Shapiro and Sober’s ‘Epiphenomenalism Do’s and Don’ts’ 3 WALM’s Explanatory Doctrine 4 Galton’s Autonomous-Statistical Explanation 5 A Second Example: The Statistical Explanation of the Diffusion of Gases 6 Distinguishing Two Theories of Evolution by Natural Selection 7 A Possible Objection: Are Statistical Laws Sufficient for Explanation? 8 Conclusion

R. A. Fisher and Social Insects: The Fisher-Darwin Model of the Evolution of Eusociality

Fisher recognized that the evolution of social insect colonies needed explaining, a point which Charles Darwin had avoided discussing in detail. Fisher, in his 1930 book The Genetical Theory of Natural Selection, outlined in detail how eusociality could evolve, and developed a verbal model by connecting selection on fecundity with the sterility of workers. Fisher saw social insect colonies as harmonious units, in contrast to human societies that exhibit intra-communal conflict. Fisher’s development of the model was strongly influenced by his correspondence with Leonard Darwin, and so I call it the Fisher-Darwin model. However, Fisher was not influenced by Lamarckian views on the evolution of social insects that were common at that time. The Fisher-Darwin model presages almost exactly the mathematical model recently proposed by Nowak and colleagues.

The origins of Austin Bradford Hill's classic textbook of medical statistics

Between January and April 1937 The Lancet published a series of seventeen articles on medical statistics by Austin Bradford Hill. The series was suggested by Dr MH (Pamela) Kettle, an assistant editor at The Lancet, and Hill was paid £3.3.0 (three pounds, three shillings, and no pence – roughly equivalent to £135 today) for each ‘installment’. The seventeen articles formed the chapters of Principles of Medical Statistics,1 which was also published in 1937. Over the next fifty-five years, the book was to run through twelve editions, and become renowned worldwide amongst physicians, epidemiologists, and medical statisticians. Hill had a very distinguished ancestry. ‘From James (b. 1724) and Sarah Hill (∼1733–1801) descended one of the most intellectual families that has ever arisen in England’2 (Gun, quoted by Hill). The family included Thomas Wright Hill (Hill's great-great grandfather, 1763–1851), an educational reformer who devised a system of shorthand, and wrote on the hexadecimal system of counting and proportional representation in parliamentary elections; Sir Rowland Hill (1795–1879), who introduced ‘the penny post’; Matthew Davenport Hill (1792–1872), a criminal-law reformer; George Birkbeck Hill (Hill's grandfather, 1835–1903), writer and critic renowned for his edition of Boswell's Life of Johnson; Rosamund Davenport Hill (1825–1902), a prominent member of the old London School Board; Sir Maurice Hill (1862–1934), a High Court judge; and Sir Leslie Scott (1869–1950), a politician and judge who was briefly the Solicitor General. (He was the eldest son of Sir John Scott (1841–1904), another distinguished judge, whose sister, Annie (1830–1902), was the wife of George Birkbeck Hill; George's sister, Laura, was the second wife of Edward Scott, Sir John's father). Hill's father, Sir Leonard Erskine Hill (1866–1952), became professor of physiology at the London Hospital (where he recruited Major Greenwood, widely regarded as the first medical statistician in Britain, in 1905), and, from 1914, head of the Department of Applied Physiology of the newly-formed Medical Research Committee (fore-runner of the Medical Research Council). Unfortunately there is no biography of Austin Bradford Hill, although brief descriptions of aspects of his life, career, and contributions to statistical and medical research are available, for example, Doll,3,4, Farewell and Johnson,5–7 Gehan and Lemak,8 Keating,9 and Statistics in Medicine.2 In 1927 Hill joined Major Greenwood's department at the London School of Hygiene and Tropical Medicine (LSHTM), having obtained an honours degree in economics at University College London in 1922, and then a grant from the Medical Research Council to examine the high mortality in young adults in rural areas of England. While carrying out this study he attended Karl Pearson's course on statistics at London University. Hill's first four papers, for all of which he was the sole author, were published in 1925. The following year he was awarded a PhD by the University of London with a thesis entitled 'A physiological and economic study of the diets of workers in rural areas as compared with those of workers resident in urban areas,’ which consisted of these four papers (we have not been able to trace the identity of Hill's examiners). In 1930 Hill obtained a DSc from the University of London, with a thesis entitled 'An investigation of sickness in various industrial occupations.’ This comprised eight (single-authored) papers published between 1925 and 1929, including the four submitted for his PhD degree. His external DSc examiner was WP Elderton, author of Frequency curves and correlation (1906) and Primer of statistics (1906). Major Greenwood was the internal examiner. By 1936 Hill had published thirty-nine book reviews, eight research reports, and sixteen papers, including nine in the British Medical Journal or The Lancet, and four in the Journal of the Royal Statistical Society. Many of the reviews were of books about population, poverty, industrial working conditions, migration, mortality, and the social conditions in London, subjects of obvious central interest at the LSHTM. However Hill also reviewed Hartshorne and May's Studies in Deceit, Burgess's Introduction to the Mathematics of Statistics, Fisher's The Genetical Theory of Natural Selection, and the second edition of Pearl's Introduction to Medical Biometry and Statistics, thus demonstrating an interest extending into statistics, genetics, and beyond. Most of Hill's papers also reflected a focus on mortality, longevity, social conditions, and industrial working conditions (including those of nurses), reflecting the rapid development of analyses of vital statistics which had occurred over the previous one hundred years. But his interests extended beyond mortality to cricket, experimental epidemiology, and medical practice.

Linking sexual selection and behavioral isolation in a colorful species of stream fish, Etheostoma barrenense

Linking sexual selection and behavioral isolation in a colorful species of stream fish, Etheostoma barrenense

Simulating the Emergence and Survival of Mutations Using a Self Regulating Multitype Branching Processes

It is difficult for an experimenter to study the emergence and survival of mutations, because mutations are rare events so that large experimental population must be maintained to ensure a reasonable chance that a mutation will be observed. In his famous book, The Genetical Theory of Natural Selection, Sir R. A. Fisher introduced branching processes into evolutionary genetics as a framework for studying the emergence and survival of mutations in an evolving population. During the lifespan of Fisher, computer technology had not advanced to a point at which it became an effective tool for simulating the phenomenon of the emergence and survival of mutations, but given the wide availability of personal desktop and laptop computers, it is now possible and financially feasible for investigators to perform Monte Carlo Simulation experiments. In this paper all computer simulation experiments were carried out within a framework of self regulating multitype branching processes, which are part of a stochastic working paradigm. Emergence and survival of mutations could also be studied within a deterministic paradigm, which raises the issue as to what sense are predictions based on the stochastic and deterministic models are consistent. To come to grips with this issue, a technique was used such that a deterministic model could be embedded in a branching process so that the predictions of both the stochastic and deterministic compared based on the same assigned values of parameters.

The change in genetic diversity down the core‐edge gradient in the eastern spadefoot toad (Pelobates syriacus)

Several hypotheses are available to predict change in genetic diversity when approaching peripheral populations. We used the eastern spadefoot toad in Israel as a model system to examine these hypotheses using population genetics analyses and network theory. Our results contradicted most of the predictions from the 'abundant centre' model, that edge populations should have lower density and lower genetic diversity than core populations. Furthermore, dispersal rate between core and peripheral populations is expected to be asymmetric, mostly directed outwards from the core population, but we did not detect such a trend. Our data did not support the hypothesis of no change or a non-linear change in genetic diversity towards the range edge. However, our results did fit the Fisher (The Genetical Theory of Natural Selection, Clarendon Press, Oxford, 1930) hypothesis, which predicts increase in genetic variability from core to edge of distribution. We attributed this finding to the much harsher climatic and abiotic conditions at the edge, which must be tolerated over generations by both tadpoles and post-metamorphic individuals in this region. Finally, our results have significant conservation implications for the survival of this species in Israel, where it is critically endangered. We identified two distinct communities, which are genetically linked through two specific rain pools in the Upper Galilee. Details on the spatial subdivision of this species are cardinal for future management and restoration of temporary wetlands in Israel.

Two Birds of Paradise in North Holland, 1592: The Gift in the Exotic

Two Birds of Paradise in North Holland, 1592: The Gift in the Exotic

R. A. Fisher. The relevance of the genetical theory of natural selection

Starting from the main statement that “. . . natural selection is not evolution. . . ”, R.A. Fisher built the foundation of the genetic theory of population in his famous work Genetical Theory of Natural Selection (1930). He rewrote the scientific paradigm proposed by Darwin in statistical terms using the calculus of probability and, most importantly, statistics. The key to his formal transposition is in the analysis of variance inwhich Fisher interpreted as phenomenical variability by means of random variability: this completely original result would become a fundamental chapter of statisticalmethod. It is not by chance that at the same time he published his statistical method for research workers in which the analysis of variance dominated his primary elements of the design of experiments.

Truthful signalling, the heritability paradox, and the Malthusian equi-marginal principle

The article shows that heritable quality differentials are consistent with the Zahavi Handicap Principle (the Truthful Signalling Hypothesis (TSH)). Earlier analyses have assumed non-heritable quality. The crucial innovation is the Malthusian equi-marginal principle: under selection pressures the relative numbers of higher- and lower-quality organisms will change until, in equilibrium, not the average but the marginal levels of quality will be equalized. Assuming kin selection, each male maximizes his own reproductive success and signals until the marginal value of more signalling is zero. We further require evolutionary stability; displacements to higher or lower population sizes must be restored to equilibrium. The article proposes an alternative to Fisher's [1958. The Genetical Theory of Natural Selection. Dover Publications, Inc., New York [Original publication 1929]] and Hamilton and Zuk's [1982. Heritable true fitness and bright birds: a role for parasites? Science 218, 384–387] suggestions. The model is solvable for ranges of parameters that constitute the stable region. We particularly consider the unit signalling costs of the high- and low-quality males, where it has been widely believed that for a TSH equilibrium the former must be lower than the latter. This article confirms our earlier result that this is not a necessary condition for a truthful signalling equilibrium, though the unit signalling costs of the high-quality males cannot be too much larger.

Preference for sons and sex ratio in two non‐western societies

Moroccan Berbers and the Bolivian Aymara are two unrelated peasant groups living in adverse environments with a still rather traditional agriculture. Precarious life conditions may be responsible for the importance given to male labor and hence for the cultural preference conferred on male descent. This preference, expressed in the social valorization of sons to the detriment of daughters, is more emphasized if the socioeconomic status of the family is lower. Interpreted according to the cost/benefit approach of Fisher (1958 Genetical Theory of Natural Selection, New York: Dover) of variations in the sex ratio and to its later developments (Trivers and Willard 1973 Science 179:90-91; Trivers 1985 Social Evolution, Menlo Park: Benjamin/Cummings; Hewlett 1991 J. Anthropol. Res. 47:1-37; Smith 1993 Ethol. Sociobiol. 14:39-44), these cultural characteristics could determine that parents bias the care allocated to their progeny in favor of sons, to the detriment of daughters. This could eventually affect their respective survival and finally modify the offspring tertiary sex ratio. This study compares differences in survival as indicators of parental care according to a child's sex and across three economic strata: poor, medium, and high. The Moroccan data express no other sex differences in survival than an advantage for daughters during the preweaning period in the medium and high strata. Within the same sex and across economic strata, a greater mortality between age 10-20 years is observed for boys of the poorest stratum. The Aymara data do not show sex differences by economic stratum, except for an advantage to daughters between birth and 5 years in the medium group, and no survival differences within the same sex across economic strata. These results suggest either that despite the social valorization of male progeny no differences in parental care according to sex occur, or that their magnitude is not great enough to contrast the survival of sons and daughters.

Impacts of flight distance on sex ratio and resource allocation to offspring in the leafcutter bee, Megachile rotundata

Fisher's theoretical prediction of equal investment in each sex for a panmictic population (The genetical theory of natural selection. Clarendon, Oxford, 1930) can be altered by a number of factors. For example, the sex ratio theory predicts variation in equal investment in each sex when the maternal fitness gains from increased investment differ between sexes. Changing sex allocation because of changing payoffs may result from different ecological situations, such as foraging conditions. We investigated the impact of foraging travel cost on relative investment in sons vs daughters. Field studies were carried out with the central-place-foraging leafcutter bee Megachile rotundata (Fabricius), which has smaller males than females. Therefore, less investment is required to produce a viable son compared with a daughter. We found that with increased flight distance to resources, females produced a greater proportion of sons. Females also invested fewer resources in individual sons and daughters and produced fewer offspring with increased flight distance.

Betrayed by Balanoglossus: William Bateson's rejection of evolutionary embryology as the basis for understanding evolution

William Bateson (1861–1926) is most remembered as the man who introduced Mendelian genetics into Britain, propounded heredity as the mechanism of evolutionary change, and coined the name ‘‘genetics’’ for this new field of scientific endeavor. Less well known is Bateson’s early professional life as an evolutionary morphologist/ evolutionary embryologist studying the wormlike marine organism, Balanoglossus. His aim was to uncover the origin of the vertebrates; was Balanoglossus ‘‘the last of the invertebrates or the first of the vertebrates’’ (Crowther, ’52: p 253). Bateson, however, switched from embryology to heredity. How Bateson came to feel betrayed by Balanoglossus and how and why he made the switch from what he termed ‘‘the embryological method’’ to genetics and the analysis of variation as the means to understand evolutionary change are the topics of this paper. It is a contribution to the history of the now flourishing field of evolutionary developmental biology (‘‘evo-devo’’), which is the current manifestation of a field with a long history seeking relationships between embryology and evolution (Bonner, ’82, 2000; Hall, ’92, ’98, 2004, 2005; Raff, ’96). The significance of Bateson’s scholarship for the relationship between embryology and evolution was captured by Peter Bowler in a Foreword to the reprinting of Bateson’s Materials for the Study of Variation: ‘‘Biologists who retain an interest in the developmental process and its relationship to evolution may find his [Bateson’s] work illuminating for its ability to identify topics and problems which have been set aside in the construction of the genetical theory of natural selection. The few who wonder if a full recognition of the role played by development will threaten the basic principles of Darwinian adaptationism can see how far one of the pioneers of genetics was willing to carry this line of argument’’ (Bowler, ’92: p xxiv–xxv).

R. A. Fisher-twice Professor of Genetics: London and Cambridge, or'A fairly well-known geneticist'

Summary. Fisher held in succession the two senior chairs of genetics in Great Britain, the Galton Professorship at University College London (1933–1943) and the Arthur Balfour Professorship at Cambridge (1943–1957), yet his fame as a statistician is such that his contributions to genetics and evolutionary biology are not well known in statistical circles. In the paper his genetical work, and that of his colleagues, is summarized, and his influence in evolutionary biology, especially through The Genetical Theory of Natural Selection (1930), is sketched.