The First Notable Family Study Involved the Jukes.

JOHN Maynard Smith was 1 of the most influential evolutionary biologists of the generation that succeeded the "founding fathers" of population genetics, as he was fond of calling Fisher, Wright, and Haldane. Maynard Smith's male parent was a London surgeon, but died when John was eight years old. His mother came from a well-to-do Edinburgh family. His babyhood holidays were spent with his grandparents in rural Somerset, where, without any encouragement from adults, he developed a strong involvement in natural history (Maynard Due southmith 1985). Stag hunting was a major occupation of the local inhabitants, and John alleged that the start of each hunting season was celebrated by the local church with the anthem "As pants the hart for cooling stream/while heated in the chase/so longs my heart for Thee, O Lord/and Thy redeeming Grace." At 13, he entered Eton College, the best-known public (i.e., private) school in England. He detested this bastion of the English ruling classes, although he admitted that the mathematics teaching was very good (Maynard Smith 1985). He then studied engineering at the University of Cambridge, where he was one of the first undergraduates to become married. His wife, Sheila, is a mathematician, who later on worked on human genetics and then on bacterial genetics until her retirement from the University of Sussex.

In 1938, John visited Berlin, where his uncle was the British military attaché. He used to merits that his uncle hatched a plot to electrocute Hitler during a parade, using a sniper posted on the roof of the French embassy. This would have had the desirable effect (from the British point of view) of both eliminating Hitler and provoking a conflict between French republic and Germany. Unfortunately, the programme was vetoed by the British government. In response to what he saw happening in Berlin, and to his experiences at Eton, John joined the British Communist party, in which he was very active until 1947. Later that, his allegiance gradually faded, and he left the political party in 1956 following the brutal Soviet suppression of the Hungarian revolution (Gaynard Due southmith 1985). In afterwards years, he became a critic of Marxism, while retaining left-of-center political views.

During the war, John worked on aircraft design in factories in Coventry and Reading, just decided to alter to biology after the end of the war, having decided that aircraft were "noisy and old-fashioned." He studied zoology at University College, London (UCL), where Haldane held the Weldon Chair of Biometry. He stayed on as a graduate student of Haldane's just never took his Ph.D., as he was given an appointment in the zoology department at UCL. (If you are as good as Haldane or Maynard Smith, a Ph.D. is an unnecessary adornment.) Haldane, e'er called "Prof" by John, was his life-long hero and his colleague until 1957, when Haldane moved to India (Maynard Smith 1985). John one time wrote:

I first read a book of essays by Haldane, it was The Inequality of Man, when I was at Eton. I was led to read them considering he was regarded by at to the lowest degree some of the masters every bit a figure of immense wickedness. Although I did not know it, this chance encounter with Haldane's writings had a big influence on my future career… When, ten years afterwards, I decided to chuck in applied science to study biology, I went to Academy College, London, considering I wanted to study under Haldane … (1000aynard Due southmith 1968a, p. vii).

In common with Haldane, John was an outstandingly clear lecturer and writer, with an immense breadth of knowledge and interests. While both were adept at spotting biologically significant theoretical problems, neither used particularly elegant mathematics: they were more interested in getting useful solutions, fifty-fifty if their methods caused professional person mathematicians to grind their teeth (John was once bellyaching by an eminent theoretician's reference to the "rough and ready methods of Maynard Smith").

They were also both outstanding communicators of scientific discipline to the general public, in John's case through television as well as his writings. John'southward Penguin paperback book The Theory of Evolution (1000aynard Smith 1958c) must have stimulated the interest of many young readers in evolutionary ideas; I certainly think reading information technology avidly equally a teenager around 1960. Just, in contrast to Haldane, who was renowned for his irascibility (and for being ane of the few participants to actively savor Earth War I), John was a kindly and gentle person. In 10 years of association with him every bit a close colleague, I cannot recall any angry words between the states, even when I made a fool of myself. John oft told how he had to beg Haldane non to have daily rows, since they ruined his ability to work afterward. Haldane seemed genuinely surprised that John did not enjoy fighting. Even so, John had a keen eye for stupidity and pomposity and could occasionally let wing. He attended the funeral of George Price, who sadly committed suicide while suffering from religious obsessions. The officiating clergyman told John that Price'south problem was that he thought that he had a hotline to God, to which John responded "Only like St. Paul."

John received many scientific honors during his career, including election as Fellow of the Royal Society and Foreign Associate of the National University of Sciences. He was awarded the Darwin, Royal, and Copley Medals of the Purple Guild, every bit well as the Balzan, Crafoord, and Kyoto Prizes. He did not receive any of the honorific titles dispensed to the groovy and the practiced by the British Regime, claiming that Sheila would divorce him if he accepted one. His FRS came at the absurdly late historic period of 57. It is rumored that this reflected the antagonism that persisted for a long time between the schools of Haldane and Fisher, with much regrettably petty behavior on the part of both great men. John recalled giving a lecture to the United kingdom Genetical Society as a struggling young scientist. Shortly after he started, Fisher got up, ostentatiously put on his coat and scarf, and and so stumbled over the feet of people sitting in his row in lodge to leave the room. Haldane used to unnerve speakers he disliked past sitting in the forepart row, placing his large, domed head in his easily, and exclaiming "Oh God, Oh God!" in a penetrating vocalisation.

Despite his great fame, John remained a humorous and unpretentious person during the whole of his life, although he certainly did not display any simulated modesty. He was unusually attainable to immature scientists and was often to be seen in the bar at meetings, exchanging ideas with a crowd of colleagues, young and quondam, until tardily in the evening. He was very open to new ideas, even if there was a strong take chances that they were wrong, and even if he did not much like the person who was proposing them. He created an uncommonly heady temper at Sussex, with numerous breather visitors from overseas (in my time in that location, these included Rolf Hoekstra, David Penny, Sue Riechert, and Monty Slatkin), besides as postdocs of various nationalities (such every bit Jim Balderdash, Peter Hammerstein, David Queller, Jon Seger, Curt Strobeck, and Wolfgang Stephan). All this was achieved with very little grant funding: John did most of his work with pencil and paper or a primitive desktop reckoner. He was the kind of thinker who needed to talk through his ideas earlier they crystallized. His marvel and intellectual strengths forged many collaborations that flowed out of his love of discussion and statement (much of information technology over morning coffee or evening beer). He was non very successful as a trainer of graduate students, at least in his afterward years. This was partly, no doubt, due to the scarcity of biology graduates interested in, or able to do, theoretical piece of work, and partly to the fact that his policy was to "allow them be effectually" rather than straight a research project. He could be quite overpowering in discussion and usually dominated any conversation in which he took part (occasionally he met his friction match with some of the larger egos in the business concern). Notwithstanding, he would mind carefully to objections to his viewpoint if you were persistent plenty and was far more than interested in getting at the truth than in winning an argument. He was e'er generous in his evaluations of the achievements of others and quick to aid the careers of young people whose talents he had noticed.

John's early on work in the 1950s was mostly on the genetics of Drosophila subobscura, which Haldane's laboratory was developing as a European rival to D. pseudoobscura. The study of the population genetics of this species has been revived in contempo years, mainly by Greek and Castilian scientists (1000rimbas 1993; Navarro-Sabaté et al. 2003). This owes much to John'southward early studies. John one time said that his biggest scientific failure was to accept overlooked the significance of intragenic recombination that Thea Koske and he detected in a mapping experiment on D. subobscura (Thousandoske and Gaynard Smith 1954). If he had interpreted this correctly, he might take shared in the epoch-making discoveries of that time on the structure of the gene. It is interesting that this work was briefly cited by Pontecorvo in his classic monograph synthesizing piece of work on intragenic recombination:

… the highest recombination (0.5 per cent) so far measured between ii non-complementary (i.due east., functionally allelic) recessives in organisms college than phage is that plant by Koske and Maynard-Due southmith [sic] (1954) between two ar alleles of Drosophila subobscura (Pontecorvo 1958, p. 34).

John was very interested in animal behavior over the whole of his career, and his last book, with his colleague David Harper, is Fauna Signals (Maynard Smith and Harper 2003). His studies of the effects of inbreeding on male mating behavior and reproductive success in D. subobscura (Kaynard Southwardmith 1956) acquired him to go an advocate of the evolutionary significance of sexual selection by female person choice of mates. As John noted over 40 years afterward (Chiliadaynard Southwardmith 2000), sexual selection involving female person mate pick was largely disregarded by nearly of the leading early 20th century evolutionary biologists, with the notable exception of Fisher (1930). There is, for example, simply a unmarried reference to it in Ernst Mayr'due south Animal Species and Evolution (Mayr 1963). In 1958, John wrote a perceptive article in a Darwin centennial volume (Thouaynard Smith 1958b), in which he anticipated the "proficient genes" theory of the development of female mate option, currently the subject area of much research in behavioral ecology. He summarized his studies of mate selection in D. subobscura as follows:

At that place was an association betwixt those characteristics of males making for mating success (probably athletic ability) and those making for fitness as a parent (the production of a large quantity of sperm). It has non been shown that a similar clan exists in natural populations, but it seems very likely that it would practice so (Kaynard Smith 1958b, p. 242).

I am not sure that this last point has withal been convincingly established.

Using his engineering science preparation, John also did theoretical work on the mechanics of bird flight, suffering difficulties with mathematically ignorant reviewers, which resulted in rejection of several of his papers (Grandaynard Smith 1985). He claimed that one of them once queried a derivation that involved a differential coefficient, wondering why the d's were non cancelled in the numerator and denominator. He became used to assuming mathematical illiteracy when explaining his work to biologists and was deeply embarrassed when an anonymous visitor brought into the lab by Haldane turned out to be Alan Turing. John became a dandy admirer of Turing and used his ideas on reaction-diffusion processes (Turing 1953) in some influential work on the genetics of pattern formation (Chiliadaynard Southwardmith 1960; Chiliadaynard Due southmith and Sondhi 1960).

During the tardily 1950s and early 1960s, John pioneered the use of Drosophila as a model organism for studying the biology of aging, providing one of the primeval demonstrations of the survival cost of reproduction (Gaynard Smith 1958a) and also prove confronting the somatic mutation theory of aging (50amb and Gaynard Smith 1964). The evolution of life-history traits in general, and crumbling in particular, has become a flourishing branch of evolutionary biological science, and Drosophila is now a major tool for analyzing the functional biological science of aging (Partridge and Geuropean monetary system 2002).

In 1965, John left UCL to become the founding Dean of the Schoolhouse of Biological Sciences at the then new Academy of Sussex, situated in an bonny park on the outskirts of Brighton, which was formerly the property of the Earl of Chichester. John very finer built up a thriving group of biologists, biochemists, and experimental psychologists. This achievement was later undermined by the set on on British universities launched by the Thatcher government during the 1980s, when several of his closest colleagues left the academy around the time of his retirement in 1985. John, nevertheless, remained at Sussex for the rest of his life and was pleased to run into a renaissance accept place over the concluding decade or so, with a very active grouping in evolutionary biology emerging. He avoided condign an administrator at a college university or national level, although he became Dean again for a couple of years before retirement, in response to the strain the school was under at the time.

After moving to Sussex, John concentrated increasingly on theoretical work and eventually abased experimental piece of work. This was partly due to the time needed for his administrative work and partly considering he no longer felt overshadowed as a theoretician by Haldane, who had died in 1964. (John often said "Annihilation I could do, Haldane could do faster.") He contributed significantly to the early on development of theoretical models of molecular variation and evolution, in response to the empirical studies of protein sequence evolution and electrophoretic variation initiated in the 1960s. Unlike many British and American evolutionists at that fourth dimension, John was not at all hostile to the neutral theory of molecular evolution and variation, introduced by Motoo Kimura (Grandimura 1968) and by Jack Lester King and Thomas Jukes (Ging and Jukes 1969). He used the neutral theory equally the basis for several of his finest articles.

In particular, he and his statistician colleague John Haigh developed and analyzed the concept of "hitchhiking" (Gaynard Southmith and Haigh 1974), in which the spread of an advantageous mutation reduces variation at linked neutral loci. This idea has go very important for interpreting data on natural variation in DNA sequences, post-obit the discovery that DNA sequence variation is frequently greatly reduced in regions of the genome with low frequencies of genetic recombination (Andolfatto 2001). At that place is also increasing evidence for signatures of hitchhiking events in regions of the genome with normal levels of recombination in a variety of species, including humans (Due southabeti et al. 2002). The two Johns likewise made a very perceptive early contribution to human molecular variation, using population data on human hemoglobin variants in Europe nerveless by Hermann Lehmann's group (Fiftyehmann and Carrell 1969) to show that the amount of variation in northern European populations is inconsistent with neutral equilibrium and that there must have been a severe population clogging (Haigh and Maynard Smith 1972). Millions of dollars that have been spent on human being SNP data sets confirm this conclusion (Marth et al. 2004).

John contributed extensively to the moving ridge of theoretical piece of work on the development of sex activity and genetic systems that was initiated in the late 1960s, which freed this field from its long domination by the species-level advantage theories of Darlington (1939) and Southtebbins (1950) and replaced these by arguments based on choice amid individuals within populations. In particular, he drew attention to the paradox of the "cost of sex": the fact that a mutant that arises in a sexual species with two sexes and causes females to produce daughters asexually volition double in frequency each generation (Maynard Southwardmith 1971). Although the thought had been suggested by others previously (eastward.g., White 1945), John was the first to perceive the profound difficulty it posed for explaining the prevalence of sexual reproduction among eukaryotes. He summed upwardly the state of the field in his 1978 book The Evolution of Sex (Gaynard Smith 1978), which is still the best overview available.

John's most influential single contribution was his development, initially in collaboration with George Price, of the concept of the evolutionarily stable strategy (ESS). This invokes the principle that, for a trait value to represent an equilibrium with respect to natural pick, a necessary condition is that all possible deviant trait values are at a selective disadvantage when introduced at a low frequency into a population whose members initially all have the trait value in question. Unless fitnesses depend on the frequencies of competing phenotypes or genotypes, the ESS corresponds to the selective optimum. But in many cases, such as sex ratios or behavioral traits governing social interactions, frequency-dependent fitnesses are inherent in the biological context.

Determination of the outcome of option past calculating trajectories of cistron frequencies or of mean trait values using quantitative genetic models would be tedious and usually intractable equally far as elementary mathematical solutions are concerned. By merely testing whether rare variants are kept out of the population, the ESS approach allows informative results to be obtained in circuitous situations, e.g., the well-known event that a 1:1 allocation of resources between male and female person offspring is favored by option on nuclear genes in a randomly mating population. While this approach had been used earlier, notably by Fisher (1930) and Hamilton (1967), John's work developed the underlying logic explicitly and showed how it could be practical to many evolutionary bug, which had previously been regarded as impossibly difficult to solve with simple theoretical models. While at that place are conspicuously limitations to the ESS method, especially in cases where the genetics of a trait constrain the outcome of choice, it has proved to be an immensely useful tool. Over the by 30 years, a big theoretical and empirical literature has appeared, applying ESS methods to a very wide range of biological phenomena. For example, the prediction by ESS methods of sexual practice ratios in haplodiploid species, where they are readily controlled past maternal decisions nearly fertilizing eggs, is one of the real success stories of evolutionary biological science, in terms of relating theory to data (West et al. 2002). John's primary contributions to ESS theory are summarized in his 1982 volume (Chiliadaynard Southmith 1982).

John was very interested in full general ideas in biology and contributed to debates on such topics as grouping selection vs. kin selection (he coined the latter term: Maynard Due southmith 1964), sympatric speciation (Thousandaynard Smith 1966), punctuated equilibrium (Maynard Smith 1983), and the evolutionary office of developmental constraints (Kaynard Southwardmith et al. 1985). With Eörs Szmathmáry, he adult a set of frankly speculative ideas nigh the major events in biological development (from the evolution of life itself and the evolution of cells to the development of language), described in their 1995 book The Major Transitions in Development (Maynard Smith and Szmathmáry 1995). He also published three excellent textbooks: Mathematical Ideas in Biology (Maynard Southwardmith 1968b), Models in Environmental (Maynard Smith 1974), and Evolutionary Genetics (Maynard Smith 1989).

After his formal retirement in 1985, John turned his attending to the analysis of data on molecular variation and development in bacteria, collaborating with Brian Spratt'southward microbial genetics group, then at Sussex. This piece of work, along with that of several other bacterial population geneticists, has led to the realization that there is much more exchange of genetic information among bacterial cells in nature than was formerly believed (Maynard Smith et al. 1993). John'southward recent work involved the evolution of methods for interpreting patterns of DNA sequence variation in populations with sporadic and patchy recombinational commutation (Maynard Smith and Smith 1998; Smith et al. 2003). This has important implications for understanding bacterial pathogenicity (Thouaynard Smith et al. 2000), too equally being of groovy intrinsic involvement. It is obviously very unusual for someone to remain at the frontline of research for nearly 20 years after retirement.

In the past two years of his life, John suffered increasingly only uncomplainingly from the effects of mesothelioma, but continued working until the very stop. Despite his physical frailty, he spoke briefly at the December 2003 meeting of the U.k. Population Genetics Meeting and gave a characteristically lucid and entertaining talk on bacterial population genetics. Many of those present felt that this was their last chance to hear him give a public lecture, which unfortunately proved to exist the case. John's breadth of interests and achievements, combined with his engaging personality, were unique and will exist sadly missed.

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John Maynard Smith (right) with Sewall Wright in 1980. Wright was visiting the University of Sussex afterwards receiving the Darwin Medal from the Regal Society. The photograph was taken by Jim Balderdash.

Acknowledgments

I thank Lindell Bromham, Deborah Charlesworth, Paul Harvey, and Aubrey Manning for their comments on the manuscript.

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Source: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1448785/

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