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John Maynard Smith. 6 January 1920 − 19 April 2004:
Elected F.R.S. 1977
Brian Charlesworth and Paul Harvey
Biogr. Mems Fell. R. Soc. 2005 51, 253-265, published 1 December 2005
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JOHN MAYNARD SMITH
6 January 1920 — 19 April 2004
Biogr. Mems Fell. R. Soc. 51, 253–265 (2005)
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JOHN MAYNARD SMITH
6 January 1920 — 19 April 2004
Elected FRS 1977
BY BRIAN CHARLESWORTH1 FRS AND PAUL HARVEY2 FRS
1 Institute
of Evolutionary Biology, School of Biological Sciences,
University of Edinburgh, Edinburgh EH9 3JT, UK
2 Department of Zoology, University of Oxford, South Parks Road,
Oxford OX1 3PS, UK
FAMILY BACKGROUND AND EARLY YEARS
John Maynard Smith was born in London, the second child of Sidney Maynard Smith and
Isobel Mary (née Pitman). In 1928 the family moved to the village of Hurst in Berkshire on
the death of his father, a surgeon and freemason. John was immediately sent to boarding
school—first to a preparatory school (St Peter’s Court, Broadstairs) until 1933, then to Eton
(until 1938). His holidays were invariably spent with his sister at their maternal grandfather’s
house in Exford on Exmoor, where John became an accomplished horse-rider, stag hunter and
angler. Here, too, he developed his lifelong, self-taught interest in natural history, particularly
bird-watching. This started during his childhood in London, when he was keen on visiting the
Natural History Museum and Zoo, as well as reading any books about animals that he could
find.
He was very unhappy at school until the age of 16 years or so, and for the rest of his life
was outspoken in his criticism of Eton and similar boarding schools. In reaction to the atmosphere at Eton, and to the advance of Fascism and Nazism in the late 1930s, John became an
ardent Communist. (He had first-hand experience of Nazi Germany when he visited Berlin in
1938, where his uncle was British military attaché.) He had no formal science lessons at
school, but used Eton’s library to learn about science. It was here that he mastered the special
theory of relativity and quantum theory, and enjoyed books about biology. His particular
favourites were the essays of J. B. S. Haldane FRS, Julian Huxley (FRS 1938) and
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H. G. Wells—John was later to claim that he read Haldane because he was an author strongly
disapproved of by his schoolmasters. John read some philosophy, but never any chemistry or
geology.
He went on to university at Cambridge where, in 1941, he gained a second-class honours
degree in mechanical sciences and married Sheila Matthew (deceased, 1 August 2005), who
had taken a degree in mathematics. Sheila’s father planted tea in Ceylon, so she was sent to
boarding school (including holidays) from the age of 6 years; her family had contained no scientists but many parsons. John and Sheila had three children (Anthony, Carol and Julian) and
lived happily together until John’s death; theirs was a textbook marriage.
From 1941 to 1947 John worked as an aircraft stressman at factories in Coventry and
Reading. He had been secretary of the Communist Party at Cambridge and became local secretary of the Reading branch (he left the Party in 1956 and later a became a strong critic of
Marxism). It was in that capacity that he wrote to Haldane, himself a Communist, at
University College London, asking for advice on how to change his career from engineering
to biology. He soon started as an undergraduate, reading zoology at University College, and
graduated with a first-class honours degree in 1950.
His fellow undergraduate, Aubrey Manning, later to become Professor of Natural History
at the University of Edinburgh, has vivid memories of John’s activities as an undergraduate.
He has kindly allowed us to quote the following reminiscences.
I first met John in October 1948 when we both started to read zoology at UCL. I was 18, just out of school
and wetter behind the ears than you can imagine. He was 28, married with a family, a degree in engineering
and had been designing aircraft during the war. There were other ex-service people in our year and they all,
most especially John, helped me to grow up quickly; I was fortunate indeed. 28 seemed really old at the time,
but in fact it was John’s playfulness which first struck me. We met properly that November 5th on UCL’s ‘rag
night’. The Honours students constructed a kind of monstrous caterpillar under which we all paraded through
the West End. It was John who led the monster by a halter, wearing a baby’s cap tied under his chin and a large
nappy pinned over a scruffy pair of shorts. Later that night he and I dribbled a dustbin lid down Gower Street
until brought to a sudden halt by the shoulder of a large policeman. John never really liked the police—brushes
I imagine in his CP evangelising days—but he was civil on this occasion.
John wrote some verse about our monster and this was, I think, the first of the many verses which he set
down in the Honours Lab’s notebook. (Whatever happened to that book I wonder?) They are classics of their
type and I was glad that the journal TREE [Trends in Ecology and Evolution] recently reprinted some of them.
From memory I was able to help John reconstruct his very best composition, ‘La Femme à la Tinbergen’
(shouldn’t it have been ‘au’?), comments on fellow student Mike Ridpath’s and my obsession with the new
ethology: sign stimuli, stickleback models and all that. Referring to the elaborate courtship of ruffs towards
reeves, John wrote:
These antics formed a reflex chain
Which could be simply set in train
By patterns painted on a ball
With lines and dots and circles, all
Designed to make the ruff believe
It was the very essence, reeve.
Utterly brilliant—neither W. S. Gilbert nor G. K. Chesterton has done better!
We all devoured the splendid, if totally conventional UCL zoology course, with a year of vertebrates followed by a year of invertebrates, and so on. I vividly recall John, head with frizz of hair bent close over the
lab bench, a watchmaker’s loupe clipped onto one lens of his thick spectacles trying what he called one of his
‘blind dissections’. He was often smoking a cigarette as he did so and, peering ever closer, sometimes a smell
of burning keratin would waft up as he inadvertently singed his hair with the glowing end.
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All of us in that vintage ‘Class of 48’ came to recognise John’s brilliance. Certainly I began to learn a great
deal of biology from him right away. Whenever we have met throughout the years since those days, John
would always begin by teasing me with the idea that, as fellow students, we were rivals to get the best degree.
We did indeed both do well, but I don’t think there’s much doubt how we compared. For me, as I’m sure for
all of us, it is John’s talk one remembers. Talk indefatigable with everybody and anybody. It was not always
easy to get one’s point across in discussion. John would listen, face slightly screwed up and emitting little
‘Mm, mm’s’ signalling his intense desire to burst in again.
John went on to work for a PhD under Haldane’s supervision, but never completed it
because Peter (later Sir Peter) Medawar FRS offered him a lectureship in zoology in 1951; he
was later promoted to Reader. John was disappointed not to be offered the Weldon Chair of
Biometry in succession to Haldane, which was one reason for his eventually leaving
University College.
Several new universities were built in the UK during the 1960s, largely to accommodate
the consequences of the baby boom after the end of World War II. John turned down the
Headship of the Department of Biology at the new University of York but in 1965 started the
equivalent position—the first Dean of the School of Biological Sciences—at the University of
Sussex. He made the initial appointments there and built up the School as a leading research
institution, staying on as Dean until 1972, with a brief second term in 1982–84. He remained
a Professor of Biology at Sussex until his formal retirement in 1985, after which he stayed on
as an Emeritus Professor, going into work daily until mid-April 2004. From his time at
University College, he enjoyed teaching undergraduates. He disliked administration but had
taken on the Dean’s job at Sussex because he wanted to develop a more integrated approach
to the way in which biology was taught, as opposed to the then rather ossified standard university curriculum.
RESEARCH AND WRITINGS AT UNIVERSITY COLLEGE
John’s early work was mostly on the genetics of the common European species of fruitfly,
Drosophila subobscura, which Haldane’s laboratory was developing as a model for evolutionary studies, on the lines of Theodosius Dobzhansky’s classic use of its North American
counterpart, D. pseudoobscura. John devoted a good deal of effort to the classical genetics and
cytology of D. subobscura, observing an early example of intragenic recombination (1)*, but
failing to realize its implications for the structure of genes. His most influential work from this
period involved studies of mating behaviour and ageing in D. subobscura. He showed that
inbred males were much less successful at courting females than their outbred counterparts
(2). This caused him to become an advocate of the importance of female choice in the evolution of male-limited traits by sexual selection (4), at a time when most prominent evolutionary biologists ignored this process. The tide turned about 30 years ago, and this was to become
one of the most active areas of research in evolutionary biology (Andersson 1994). John, as
so often, was far ahead of his time.
Much the same applies to his work on ageing. His most important contribution was a series
of elegant genetic and physical manipulations of female flies, which showed that reproduction
imposes a cost in terms of survival, and significantly reduces lifespan (3). Later work has
* Numbers in this form refer to the bibliography at the end of the text.
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confirmed that ageing in female flies is accelerated by reproduction early in life, and
Drosophila is now a prime tool for investigating the evolutionary and functional biology of
ageing (Partridge & Gems 2002). John also took an interest in the problem of the genetic control of development, long before the availability of the genetic and molecular tools that have
revolutionized this field. He was strongly influenced by the ideas of Alan Turing FRS and Curt
Stern on pattern formation, and applied them in a typically creative way to the interpretation
of genetic variation in the number of ocelli in D. subobscura (6, 7).
In 1958, John published his Penguin paperback The theory of evolution (5), which ran
through several editions (now available as a Canto reprint by Cambridge University Press
(1993), with a foreword by Richard Dawkins (FRS 2001)) and which is one of the best introductions to evolution for the general reader. It displays his admirable command of a wide
range of biological knowledge and his ability to convey the gist of important ideas without
getting involved in undue detail. It must have stimulated many young people to take an interest in biology (it certainly influenced both of us as teenagers). John felt, however, that it
injured his career because senior colleagues regarded him as a lightweight popularizer rather
than a serious researcher.
RESEARCH AND WRITINGS WHILE EMPLOYED AT THE UNIVERSITY OF SUSSEX
After moving to Sussex, John soon gave up empirical research and committed himself to the
development of theory so that he could crystallize out ideas, mainly about evolution but also
on animal behaviour and, to a smaller extent, ecology. Characteristically, he identified problems during discussion with other biologists, developed a mathematical model, applied it to
case studies, and then read the relevant literature to see whether what he had done was novel.
During this period he became identified with the development of several fields of research,
notably the evolution of sex and the application of game theory to problems in evolution and
behaviour. He returned to these topics time and again, before and after writing definitive books
on them. It is therefore not possible to write a coherent temporal account of his research and
writings between 1965, when he arrived at Sussex, and 1985, when he formally retired.
Rather, it is more appropriate to classify areas of research and writing first, and then comment
on their temporal progression.
Relatively early during his time teaching at Sussex, John identified the need for an undergraduate textbook that would introduce biologists from a broad spectrum of the subject to the
use of mathematical modelling. His textbook, Mathematical ideas in biology (11), with its
worked examples and problems, was an instant success for students and teachers alike, as
broadly based biology courses developed across the country. After his training in population
genetics, John became interested in how ecologists modelled their problems. He brought his
thoughts together in a short textbook that also contained original research results, entitled
Models in ecology (16). Perhaps most notable was his treatment of cycles and time delays,
failing narrowly to discover the notion of ‘chaos in ecology’ (May 1976), and his attempts to
incorporate population genetic variation into ecological models, which laid the basis for fruitful developments by others (Roughgarden 1979).
Before arriving at Sussex, John had bought Haldane a copy of V. C. Wynne-Edwards’s
book Animal dispersion in relation to social behaviour (Wynne-Edwards 1962) to read in his
hospital bed. Wynne-Edwards (FRS 1970) was convinced that the action of selection on the
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properties of breeding groups rather than on differences between individuals was needed to
explain the phenomena he described, in particular what he thought were adaptations to avoid
overpopulation. Haldane was aware that group selection for altruistic behaviour would be a
weak force compared with individual selection for selfish behaviour, and that it could work
only under special conditions (Haldane 1932). He summarized Wynne-Edwards’s explanation
for leks as a population control measure by explaining to his younger colleague that it was as
if males put a notch on a pole in the ground every time a female was mated. When the pole
had collected enough notches, the males desisted from further mating with the expression
‘Ladies, enough is enough’.
In the 1930s, Fisher (1930) and Haldane (1932) had produced preliminary sketches of
the conditions under which individuals might be expected to be altruistic, specifically when
interacting with their close relatives. This idea was developed in detail as the theory of inclusive fitness by W. D. Hamilton (FRS 1980) and was used to explain the frequent evolution of
eusociality in haplodiploid insects (Hamilton 1964a, b). John had reviewed Hamilton’s original submission to the Journal of Theoretical Biology and saw the need to distinguish between
what he called Wynne-Edwards’s ‘group selection’ and Hamilton’s ‘inclusive fitness’ in
explaining the evolution of social behaviour (8). John subsequently used the concept of ‘kin
selection’ (as he termed selection on traits that cause individuals to benefit their relatives) to
explain previously puzzling behaviours such as the evolution of alarm calling (9) and
polyandry (14).
Group selection and selection at the species level had previously also been used to explain
the evolution and maintenance of sexual reproduction, especially by C. D. Darlington FRS in
his pioneering book on the subject (Darlington 1939). The writings of most biologists who
considered these problems invoked Darlington’s ideas until the late 1970s. However, the
weaknesses of group selection in explaining altruism also apply to the evolution of sex and
genetic systems, as was firmly pointed out by George Williams in two influential books
(Williams 1966, 1975). John was a pioneer of the development of rigorous models in this area,
based on sound population genetics principles. He was particularly concerned with the paradox of the ‘cost of sex’: the fact that an asexually reproducing female has twice the reproductive capacity of a sexual female, in terms of numbers of daughters, so that an asexual lineage
with no fitness disadvantage will rapidly take over when introduced into a sexual population.
John was not the first to recognize the twofold cost of sex, but he was the first to point out that
it raises severe problems for understanding how sexual reproduction can be maintained, at
least in species with a clear distinction between male and female germ cells.
He spent years developing his models and wrote many papers on the evolution of sex, culminating in his definitive book on the subject, The evolution of sex (20). This was a book that
laid out the problems and most of the likely solutions (some new ideas have subsequently
arrived on the scene; see, for example, Otto & Lenormand (2002)), but John was never satisfied that he fully understood why we live in a sexual rather than an asexual world. The difficulty is essentially that there are more competing theories than critical empirical evidence. It
is unusual for a scientist to become so clearly identified and respected by his contemporaries
for clarifying the nature and magnitude of a problem, without having provided a precise solution. John’s writings did, however, lead to a complete change in perspective on the evolution
of sex and genetic systems, which stimulated a wealth of both theoretical and empirical
work—Alexey Kondrashov, for example, happily acknowledges that his interest in the subject
was fired by John’s book.
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In contrast with the frustrations associated with trying to solve the problem of the evolution of sex, his application of game theory to biological questions was an extraordinary success story, marked by new understanding of a variety of biological phenomena. He had been
sent a manuscript written by George Price to referee for Nature. Again, this was essentially a
rebuttal of the group selectionist idea that, for the good of the species, animals do not escalate
aggressive behaviour in fights. Price suggested instead that animals had been selected not to
escalate so as to avoid damage to themselves through retaliation by their opponent.
John was stimulated to develop the concept of the evolutionarily stable strategy (ESS): a
population is at a stable evolutionary equilibrium when an alternative strategy, introduced at
low frequency, cannot invade. The ESS can consist of a population containing several trait values at specified frequencies, a so-called ‘mixed ESS’. The concept of frequency-dependent
fitnesses, for which ESS theory is specifically designed, was already familiar to evolutionary
biologists, who had long realized that many polymorphisms are maintained by this form of
selection. An earlier application of game theory had been described by Hamilton (1967), who
explained the evolution of unusual sex ratios by heterogeneity in the distribution of mates
(local mate competition) and used the term ‘unbeatable strategy’ for the ESS. Initially John,
together with Price (whose original manuscript remained unpublished), simply applied the
notion to animal contests in which the fitness costs of retaliation were considered substantial
(15). As different strategies were envisaged, from the original ‘hawk’ and ‘dove’ (Price, a religiously troubled soul, at one time asked that ‘doves’ be renamed ‘mice’ to avoid identification
with the Holy Spirit and consequent blasphemy), such as ‘bully’, ‘retaliator’ or ‘tit-for-tat’, so
additional or alternative ESSs emerged. Fruitful collaborations ensued, noticeably with
Geoffrey Parker (FRS 1989) on asymmetric contests (18).
This relatively simple analytical tool was quickly applied to problems outside the context
of animal battles over mates. For example, an incisive analysis of the evolution of parental
care considered cases in which the offspring were guarded by neither, one or both parents (19).
As the risks to the offspring varied, and the opportunities of producing additional offspring
differed between the sexes, so would the ESS. This extraordinarily simple but elegant analysis of parental care provided the depth and breath of insight that was so characteristic of John’s
ESS work. He summarized the approach and its biological applications in his book Evolution
and the theory of games in 1982 (21).
John made other significant contributions to evolutionary biology during this period, in
which his reputation as one of the world’s foremost evolutionary biologists was established.
In particular, he did pioneering research on the theory of molecular evolution and variation, in
response to the appearance of data on protein sequence evolution and variation in the 1960s.
His most influential contribution from this period is probably his model of ‘hitch-hiking’
(developed jointly with his mathematician colleague, John Haigh) (17). They formulated
equations describing the effect of the spread of a selectively favourable mutation on the level
of variability at nearby sites in the genome, which are not themselves under selection. With
the advent of large amounts of information on variability at the DNA sequence level, it is now
possible to screen genomes for signs of hitch-hiking events, and thus to test for the occurrence
of adaptive evolution at the molecular level (Sabeti et al. 2002). The 1974 paper provides the
conceptual foundation for such tests and is now widely cited.
An earlier paper by John on the concept of a protein space (12) has also had contemporary
influence, in the light of new ideas on how to model the process of adaptive evolution of protein and DNA sequences (Orr 2003), and the availability of data from viral systems on the
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detailed steps that actually occur during such evolution (Rokyta et al. 2005). Another paper
with John Haigh showed that data on human haemoglobin variants in European populations
indicated a recent bottleneck of small population size (13). Again, the use of data on molecular variation to infer population history is now a major research area (Marth et al. 2004).
John also wrote an influential paper on sympatric speciation, demonstrating that it is theoretically possible for a single randomly mating population to become split into two reproductively isolated populations, provided that selection acts on a polymorphic locus in such a way
that different variants are favoured in different environmental patches (10). This contradicts
the dogma established by Ernst Mayr (ForMemRS 1988) that speciation requires initial geographic isolation (Mayr 1963). As John said to one of us (B.C.) during his PhD examination,
‘I wrote my paper purely in order to annoy Ernst Mayr’, in which he was surely successful.
Sympatric speciation is now thought to be somewhat more common than previously thought,
and it is certainly not theoretically impossible (Coyne & Orr 2004); John’s paper was another
pioneering effort ahead of its time. John was also very interested in the debates on evolutionary mechanisms that flared up in response to Stephen Jay Gould and Niles Eldredge’s advocacy of evolution by punctuated equilibrium in the 1970s and 1980s. He wrote several papers
on these issues, defending neo-Darwinian ideas, while being open to ideas such as developmental constraints (22–24).
RESEARCH AND WRITINGS IN RETIREMENT
Although John formally retired in 1985, he carried on teaching at the University of Sussex for
several years and continued to produce books and original research papers at an impressive
constant rate. He went into work at the newly named John Maynard Smith biology building
on most days until he died in April 2004, after a long and debilitating illness (mesothelioma).
One of his first tasks in retirement was to write Evolutionary genetics (25), a substantial and
successful evolution textbook for undergraduates that embraced a mathematical approach to
the subject, unlike the then-standard books by Douglas Futuyma and Mark Ridley. Another
major book emerged from a collaboration with the Hungarian theoretician Eörs Szmathmáry
(28). They studied what they termed the ‘major transitions in evolution’ (events such as the
evolution of the genetic code, the origin of cells and the evolution of language). Given the subject matter, this work is speculative rather than testable, but it performs the important service
of providing mechanistically plausible solutions to some of the most challenging problems in
evolutionary biology.
In 1998 he became irritated by recent literature on animal communication: it was discursive, and confused issues that he thought had been settled. At the same time, John’s younger
colleague David Harper was equally dismayed by the lack of clear thought about empirical
data—for example, the costs of signalling were frequently not estimated in the context of the
receiver. John and David worked together on John’s last book, Animal signals, which was published shortly before his death (32).
John also became interested in understanding the extent of genetic exchange among bacteria, which could be revealed through analyses of gene sequence data. This developed into his
major research area during his retirement. His interest was aroused by data from his colleague
Brian Spratt (FRS 1993), whose laboratory is one of the foremost at generating them. John’s
approach was characteristically unusual. He would sit down with groups of sequences and
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attempt to devise his own statistical tests to determine whether recombination, mutation or
other processes had occurred (29, 31). Perhaps his major contribution, with others, was to
reveal unusually high levels of recombination in some bacterial populations, together with
patchy exchange of genomic fragments between distant relatives, causing a reassessment of
the concept of a bacterial species (26, 27). The results of these analyses have important implications for our understanding of the causes of outbreaks of bacterial diseases (31). John used
these approaches to question other established beliefs. For example, in 1999 he and his colleagues argued that there might be recombination in human mitochondrial DNA (30); this
remains a matter of debate.
PERSONALITY, THE MEDIA, AND OUTSIDE RECOGNITION
John Maynard Smith was congenial. He enjoyed talking with people—friends and colleagues—and he tended to dominate almost any conversation in which he joined. His sense of
humour was acute, and he had a fund of amusing anecdotes about Haldane, whom he intensely
admired but without any illusions about his difficult personality. Like Haldane, John claimed
to be tone deaf, and he had no interest in music, unusually for someone with mathematical
ability. He was, however, a very keen and knowledgeable gardener. He and Sheila built up a
wonderful garden containing many unusual plants around their house (the White House) on
the Sussex Downs at Kingston, near Lewes. It must have been a great wrench to them to give
it up about 18 months before John’s death, in response to his declining health.
John was a fine debater, and his powers of exposition were legendary. In 1979 he decisively
defeated Duane Gish, the creationist advocate, at a packed meeting at the University of
Sussex, to the delight of his students and colleagues. At conferences he was always to be found
in the bar, and was generally one of the last to leave. He was as happy talking with postgraduate students as with senior colleagues, and the range of topics he covered seemed to be inexhaustible.
John was extremely good-natured, rarely raising his voice or losing his temper, but always
dealing with his colleagues’ erroneous views with politeness, tact, and careful discussion. He
was also very generous in helping younger colleagues with their careers, and was extremely
good at spotting emerging talents. His ability to deal with younger scientists on an equal footing, and to treat his own work with an ironic sense of humour, was one of his outstanding characteristics, lacking in many senior figures. His lack of desire to be part of the Establishment,
although willing to do his bit for the running of science and the University of Sussex, was also
refreshing. His friends and colleagues all had great respect for John’s clarity of thought and
speech, as well as his tolerance, cheerfulness and kindness. His courage during his final illness, and tenacity at continuing his work, were inspiring.
Throughout his career, John enjoyed good relationships with the media. He was frequently
interviewed, appearing regularly on the radio and television. He also wrote many popular scientific articles and book reviews, and several collections were published in his lifetime. Again,
he was not limited to writing about evolution, behaviour and ecology—indeed, he was something of a specialist on the philosophy of science where, as ever, he enjoyed debunking pomposity and pretentiousness.
John’s work was done with pen and paper, and very basic computing facilities for running
simulations and analysing data. He rarely applied for grant support, but throughout his time at
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Sussex he attracted and worked with a steady stream of postdoctoral and sabbatical visitors
with very diverse interests, many of whom are now international leaders of their fields. John
supervised few postgraduate students of his own, mainly because he preferred people to get
on with their own ideas, but he was always a popular figure with students in the department,
frequently discussing their ideas in detail and astounding them with his ability to get at the
core of a problem.
Formal recognition for John came relatively late in his life, but was nevertheless impressive. He was awarded the Darwin, Royal and Copley medals of the Royal Society, as well as
the Balzan, Crafoord and Kyoto prizes. He turned down consideration for a knighthood on the
grounds that Sheila, his wife, would disapprove. His influence on evolutionary biology in the
second part of the last century was enormous, and is likely to endure. We end with another
reminiscence by Aubrey Manning:
His huge influence remains from this talk as much as anything. He was interested by every bright person and
every new idea. I fondly recall John, acting as an absolute magnet, sitting on the deck of a Rhine cruise boat
at one of the International Ethology Conferences. For hours he was surrounded by young people, talking, talking, exchanging good science and with great wit. There was much laughter. For me, I think my adoptive country Scotland puts it best, ‘We’ll ne’er see his like agen!’
ACKNOWLEDGEMENTS
We thank Deborah Charlesworth FRS, Aubrey Manning FRSE and Lord May FRS for their comments on the draft of
this memoir. We are very grateful to Aubrey Manning for permission to quote his reminiscences of John Maynard
Smith.
The frontispiece photograph was taken in 1997 and is reproduced by courtesy of the Prudence Cuming
Association.
REFERENCES TO OTHER AUTHORS
Andersson, M. 1994 Sexual selection. Princeton University Press.
Coyne, J. A. & Orr, H. A. 2004 Speciation. Sunderland, MA: Sinauer.
Darlington, C. D. 1939 The evolution of genetic systems. Cambridge University Press.
Fisher, R. A. 1930 The genetical theory of natural selection. Oxford: Clarendon Press.
Haldane, J. B. S. 1932 The causes of evolution. London: Longman Green.
Hamilton, W. D. 1964a The genetical evolution of social behaviour. I. J. Theor. Biol. 7, 1–16.
Hamilton, W. D. 1964b The genetical evolution of social behaviour. II. J. Theor. Biol. 7, 17–51.
Hamilton, W. D. 1967 Extraordinary sex ratios. Science 156, 477–488.
Marth, G. T., Czabarka, E., Murvai, J. & Sherry, S. T. 2004 The allele frequency spectrum in genome-wide human
variation data reveals signals of differential demographic history in three large world populations. Genetics
166, 361–372.
May, R. M. 1976 Simple mathematical models with very complicated dynamics. Nature 216, 459–467.
Mayr, E. 1963 Animal species and evolution. Harvard University Press.
Orr, H. A. 2003 The distribution of fitness effects among beneficial mutations. Genetics 163, 1519–1526.
Otto, S. P. & Lenormand, T. 2002 Resolving the paradox of sex and recombination. Nature Rev. Genet. 3, 256–261.
Partridge, L. & Gems, D. 2002 Mechanisms of ageing: public or private? Nature Rev. Genet. 3, 165–175.
Roughgarden, J. 1979 Theory of population genetics and evolutionary ecology: an introduction. New York:
Macmillan.
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Rokyta, D. R., Joyce, P., Caudle, S. B. & Wichman, H. A. 2005 An empirical test of the mutational landscape model
of adaptation using a single-stranded DNA virus. Nature Genet. 37, 441–444.
Sabeti, P. C. et al. 2002 Detecting recent positive selection in the human genome from haplotype structure. Nature
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1966
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