E.B. Ford
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Edmund Brisco "Henry" Ford (23 April 1901 – 2 January 1988) was a British ecological geneticist. He was a leader among those British biologists who investigated the role of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
in nature. As a schoolboy Ford became interested in
lepidoptera Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...
, the group of insects which includes
butterflies Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...
and
moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as suborder Heterocera, but the group is Paraphyly, paraphyletic with respect to butterflies (s ...
s. He went on to study the genetics of natural populations, and invented the field of
ecological genetics Ecological genetics is the study of genetics in natural populations. It combines ecology, evolution, and genetics to understand the processes behind adaptation. It is virtually synonymous with the field of molecular ecology. This contrasts wit ...
. Ford was awarded the
Royal Society The Royal Society, formally The Royal Society of London for Improving Natural Knowledge, is a learned society and the United Kingdom's national academy of sciences. The society fulfils a number of roles: promoting science and its benefits, re ...
's
Darwin Medal The Darwin Medal is one of the medals awarded by the Royal Society for "distinction in evolution, biological diversity and developmental, population and organismal biology". In 1885, International Darwin Memorial Fund was transferred to the ...
in 1954. In the wider world his best known work is ''Butterflies'' (1945). Ford was a member of the UK Eugenics Society, of which he was a council member in 1933-1934, also contributing to its publications.


Education

Ford was born in
Dalton-in-Furness Dalton-in-Furness is a town in Westmorland and Furness, Cumbria, England, north east of Barrow-in-Furness. Along with the rest of the Furness peninsula, it was historically part of Lancashire. It is in the parish of Dalton Town with Newton, a ...
, near
Ulverston Ulverston is a market town and civil parish in Westmorland and Furness, Cumbria, England. Historic counties of England, Historically in Lancashire, it lies a few miles south of the Lake District Lake District National Park, National Park and j ...
, in
Lancashire Lancashire ( , ; abbreviated ''Lancs'') is a ceremonial county in North West England. It is bordered by Cumbria to the north, North Yorkshire and West Yorkshire to the east, Greater Manchester and Merseyside to the south, and the Irish Sea to ...
, England, in 1901. He was the only child of Harold Dodsworth Ford (1864–1943), a classics teacher turned Anglican clergyman, and his wife (and second cousin) Gertrude Emma Bennett. His paternal grandfather, Dr Henry Edmund Ford (1821–1909), was a professor of music at Carlisle and the organist of Carlisle Cathedral from 1842 to 1902. Ford was educated at
St Bees School St Bees School is a co-educational fee-charging school, located in the West Cumbrian village of St Bees, England. In 1583, it was founded by Edmund Grindal, the Archbishop of Canterbury, as a free grammar school for boys. The school remain ...
, Cumberland (now Cumbria), and read zoology at
Wadham College, Oxford Wadham College ( ) is a Colleges of the University of Oxford, constituent college of the University of Oxford in the United Kingdom. It is located in the centre of Oxford, at the intersection of Broad Street, Oxford, Broad Street and Parks Road ...
, (where his father had also studied), graduating B.A. in 1924, upgraded to M.A. 1927, B.Sc. (a research degree) in 1927, and taking a D.Sc in 1943.


Career and research

Ford's career was based entirely at the
University of Oxford The University of Oxford is a collegiate university, collegiate research university in Oxford, England. There is evidence of teaching as early as 1096, making it the oldest university in the English-speaking world and the List of oldest un ...
. The biologist
Arthur Cain Arthur James Cain FRS (25 July 1921 – 20 August 1999) was a British evolutionary biologist and ecologist. He was elected a Fellow of the Royal Society in 1989. Career Arthur James Cain was born and grew up in Rugby in Warwickshire, England ...
said Ford took a degree in classics before turning to zoology. Ford read zoology at Oxford, and was taught genetics by
Julian Huxley Sir Julian Sorell Huxley (22 June 1887 – 14 February 1975) was an English evolutionary biologist, eugenicist and Internationalism (politics), internationalist. He was a proponent of natural selection, and a leading figure in the mid-twentiet ...
. "The lecturer whose interests most closely reflected mine was Julian Huxley. I owe him a great debt, especially for inspiration... Even though Huxley was... only at Oxford from 1919 to 1925, he was the most powerful voice in developing the selectionist attitude there... I met Ray Lankester through E.B. Poulton. He was already an old man... but talked to me a good deal of
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
and
Pasteur Louis Pasteur (, ; 27 December 1822 – 28 September 1895) was a French chemist, pharmacist, and microbiologist renowned for his discoveries of the principles of vaccination, microbial fermentation, and pasteurization, the last of which wa ...
, both of whom he knew." Ford was appointed University Demonstrator in Zoology in 1927 and Lecturer at
University College, Oxford University College, formally The Master and Fellows of the College of the Great Hall of the University commonly called University College in the University of Oxford and colloquially referred to as "Univ", is a Colleges of the University of Oxf ...
, in 1933. Specialising in genetics, he was appointed University Reader in Genetics in 1939 and was the Director of the Genetics Laboratory, 1952–1969, and Professor of Ecological Genetics 1963–1969. Ford was one of the first scientists to be elected a
Fellow A fellow is a title and form of address for distinguished, learned, or skilled individuals in academia, medicine, research, and industry. The exact meaning of the term differs in each field. In learned society, learned or professional society, p ...
of
All Souls College All Souls College (official name: The College of All Souls of the Faithful Departed, of Oxford) is a constituent college of the University of Oxford in England. Unique to All Souls, all of its members automatically become fellows (i.e., full me ...
since the seventeenth century. He was among those opposed to the admission of women fellows beginning in 1981; once, upon encountering one such, he swung his umbrella at her and shouted "Out of my way, henbird!". Ford had a long working relationship with R.A. Fisher. By the time Ford had developed his formal definition of
genetic polymorphism A gene is said to be polymorphic if more than one allele occupies that gene's locus within a population. In addition to having more than one allele at a specific locus, each allele must also occur in the population at a rate of at least 1% to ge ...
,Ford E.B. 1940. Polymorphism and taxonomy. In Huxley J. ''The new systematics''. Oxford. Fisher had got accustomed to high selection values in nature. He was most impressed by the fact that polymorphism concealed powerful selective forces (Ford gave human blood groups as an example). Like Fisher, he continued the
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the Heredity, heritable traits characteristic of a population over generation ...
versus
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
debate with
Sewall Wright Sewall Green Wright ForMemRS HonFRSE (December 21, 1889March 3, 1988) was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. He was a founder of population genetics alongside ...
, whom Ford believed put too much emphasis on genetic drift. It was as a result of Ford's work, as well as his own, that Dobzhansky changed the emphasis in the third edition of his famous text from drift to selection. Ford was an experimental naturalist who wanted to test evolution in nature. He virtually invented the field of research known as
ecological genetics Ecological genetics is the study of genetics in natural populations. It combines ecology, evolution, and genetics to understand the processes behind adaptation. It is virtually synonymous with the field of molecular ecology. This contrasts wit ...
. His work on the wild populations of butterflies and moths was the first to show that the predictions made by R.A. Fisher were correct. He was the first to describe and define
genetic polymorphism A gene is said to be polymorphic if more than one allele occupies that gene's locus within a population. In addition to having more than one allele at a specific locus, each allele must also occur in the population at a rate of at least 1% to ge ...
, and predicted that human blood group polymorphisms might be maintained in the population by providing some protection against disease. Six years after this prediction it was found to be so, and furthermore, heterozygous advantage was decisively established by a study of AB x AB crosses. His magnum opus was ''
Ecological Genetics Ecological genetics is the study of genetics in natural populations. It combines ecology, evolution, and genetics to understand the processes behind adaptation. It is virtually synonymous with the field of molecular ecology. This contrasts wit ...
'', which ran to four editions and was widely influential.Ford E.B. 1964, 4th edn 1975. ''Ecological genetics''. Chapman and Hall, London. He laid much of the groundwork for subsequent studies in this field, and was invited as a consultant to help set up similar research groups in several other countries. Amongst Ford's many publications, perhaps the most popularly successful was the first book in the
New Naturalist The New Naturalist Library (also known as ''The New Naturalists'') is a series of books published by Collins in the United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Br ...
series, ''Butterflies''. Ford also went on in 1955 to write ''Moths'' in the same series, one of only a few to have authored more than one book in the series. Ford became Professor, and then Emeritus Professor of Ecological Genetics,
University of Oxford The University of Oxford is a collegiate university, collegiate research university in Oxford, England. There is evidence of teaching as early as 1096, making it the oldest university in the English-speaking world and the List of oldest un ...
. He was a Fellow of All Souls College, and Honorary Fellow of
Wadham College Wadham College ( ) is a constituent college of the University of Oxford in the United Kingdom. It is located in the centre of Oxford, at the intersection of Broad Street and Parks Road. Wadham College was founded in 1610 by Dorothy Wadham, a ...
. He was elected FRS in 1946, and awarded the
Darwin Medal The Darwin Medal is one of the medals awarded by the Royal Society for "distinction in evolution, biological diversity and developmental, population and organismal biology". In 1885, International Darwin Memorial Fund was transferred to the ...
in 1954.


Ecological genetics

E.B. Ford worked for many years on
genetic polymorphism A gene is said to be polymorphic if more than one allele occupies that gene's locus within a population. In addition to having more than one allele at a specific locus, each allele must also occur in the population at a rate of at least 1% to ge ...
. Polymorphism in natural populations is frequent; the key feature is the occurrence together of two or more discontinuous forms of a species in some kind of balance. So long as the proportions of each form is above mutation rate, then selection must be the cause. As early as 1930 Fisher had discussed a situation where, with alleles at a single locus, the
heterozygote Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mos ...
is more viable than either
homozygote Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mos ...
. That is a typical genetic mechanism for causing this type of polymorphism. The work involves a synthesis of field observations, taxonomy, and laboratory genetics.


Melanism in the peppered moth

Ford was the supervisor of Bernard Kettlewell during Kettlewell's famous experiments on the evolution of melanism in the peppered moth, ''
Biston betularia The peppered moth (''Biston betularia'') is a temperate species of night-flying moth. It is mostly found in the northern hemisphere in places like Asia, Europe and North America. Peppered moth evolution is an example of population genetics a ...
''. The entomologist Michael Majerus discussed criticisms that had been made of Kettlewell's experimental methods in his 1998 book '' Melanism: Evolution in Action''.Majerus M.E.N. 2004
The Peppered moth: decline of a Darwinian disciple.
(.doc download)
This book was misrepresented in reviews, and the story was picked up by creationist campaigners. In her controversial book '' Of Moths and Men'', Judith Hooper (2002) gave a critical account of Ford's supervision and relationship with Kettlewell, and implied that the work was fraudulent or at least incompetent. Careful studies of Kettlewell's surviving papers by Rudge (2005) and Young (2004) found Hooper's suggestion of fraud to be unjustified, and that "Hooper does not provide one shred of evidence to support this serious allegation".Young M. 2003
Moonshine: why the peppered moth remains an icon of evolution
Majerus himself described ''Of Moths and Men'' as "littered with errors, misrepresentations, misinterpretations and falsehoods". He concludes :"If you wade through the 200+ papers written about melanism in the peppered moth, it is difficult to come to any conclusion other than that natural selection through the agent of differential bird predation is largely responsible for the rise and fall of ''carbonaria''". Kettlewell and Helen Spurway, then the graduate student (and later the wife) of J.B.S. Haldane, were known to have shocked Ford by catching live moths as they flitted around a light, popping them in their mouths, and eating them whole. Haldane, who did not like Ford, was of the opinion that Ford and Kettlewell had attempted to capitalise on the supposed evolutionary adaptation of the main two variants of the peppered moth, for which Haldane, as early as 1924, had predicted the statistical probability of rate of change from light to melanic forms as an example of classic
Mendelian genetics Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularized ...
. In 1961, Haldane and Spurway talked to Canadian lepidopterist
Gary Botting Gary Norman Arthur Botting (born 19 July 1943) is a Canadian legal scholar and criminal defense lawyer (now retired) as well as a poet, playwright, novelist, and critic of literature and religion, in particular Jehovah's Witnesses. The author o ...
about the peppered moth and the unlikelihood of Ford and Kettlewell obtaining results that approximated Haldane's 1924 statistical calculations so closely, but the reasoning behind this view is far from clear. Botting already regarded the case of the peppered moth as tantamount to belief in Lamarckian evolution, and was of the opinion that some genetic mechanism other than bird
predation Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common List of feeding behaviours, feeding behaviours that includes parasitism and micropredation ...
was at work.


Bibliography

* * Clarke B 1995. Edmund Brisco Ford. ''Biographical Memoirs of Fellows of the Royal Society of London.'' * * * * (jointly dedicated to Ford and the
Nuffield Foundation The Nuffield Foundation is a charitable trust established in 1943 by William Morris, Lord Nuffield, the founder of Morris Motors Ltd. It aims to improve social well-being by funding research and innovation projects in education and social pol ...
) * * Teän, Isles of Scilly: the site of Ford's Common Blue (''
Polyommatus icarus The common blue butterfly or European common blue (''Polyommatus icarus'') is a butterfly in the family Lycaenidae and subfamily Polyommatinae. The butterfly is found throughout the Palearctic and has been introduced to North America. Butterfli ...
'') population study.


Works by Ford

* Ford E.B. (1931, 8th ed 1965). ''Mendelism and evolution''. Methuen, London. * Carpenter, G.D. Hale and E.B. Ford (1933) ''Mimicry''. Methuen, London. * Ford E.B. (1938, 2nd ed 1950). ''The study of heredity''. Butterworth, London. 2nd edn: Oxford University Press. * Ford E.B. (1940). Polymorphism and taxonomy. In Huxley J. ''The new systematics''. Oxford. * Ford E.B. (1942, 7th edn 1973). ''Genetics for medical students'' Chapman and Hall, London. * Ford E.B. (1945, 3rd edn 1977). ''Butterflies''.
New Naturalist The New Naturalist Library (also known as ''The New Naturalists'') is a series of books published by Collins in the United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Br ...
No. 1 Collins, London. * Ford E.B. (1951). ''British butterflies''.
Penguin Books Penguin Books Limited is a Germany, German-owned English publishing, publishing house. It was co-founded in 1935 by Allen Lane with his brothers Richard and John, as a line of the publishers the Bodley Head, only becoming a separate company the ...
, London. * Ford E.B. (1954, 3rd edn 1972). ''Moths''.
New Naturalist The New Naturalist Library (also known as ''The New Naturalists'') is a series of books published by Collins in the United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Br ...
No. 30 Collins, London. * Ford E.B. (1964, 4th edn 1975). ''Ecological genetics''. Chapman and Hall, London. * Ford E.B. (1965). ''Genetic polymorphism''. All Souls Studies, Faber & Faber, London. * Ford E.B. (1976). ''Genetics and adaptation''. Institute of Biology studies, Edward Arnold, London. * Ford E.B. (1979). ''Understanding genetics''. Faber and Faber, London. * Ford E.B. (1980). Some recollections pertaining to the evolutionary synthesis. In Mayr E. and Provine W.B. (eds) ''The evolutionary synthesis: perspectives on the unification of biology''. Harvard 1980; 1998. ffectively, this is an intellectual autobiography* Ford E.B. (1981). ''Taking genetics into the countryside''. Weidenfeld & Nicolson, London. * Ford E.B. and J.S. Haywood (1984). ''Church treasures of the Oxford district''. Alan Sutton, Gloucester.


Personal life

Ford never married, had no children, and was considered decidedly eccentric, although his eccentricity was said to be more prominent when he knew he had an audience; he was also fond of slightly surrealist practical joking. He could be markedly generous to his friends: it was "an open secret" that he made a handsome contribution to the grant of £350000 given by the Nuffield Foundation for the establishment of a Unit of Medical Genetics at the University of Liverpool; this greatly boosted the research of Cyril Clarke and Philip Sheppard. Professor Ford would come into first year biology lectures at Oxford University – which were quite large, with about 150 students, and address the mixed group “good morning gentlemen”, ignoring the ladies, who even at that time were maybe 30% of student numbers – they are now 48%. The students thought that was amusing, and decided that, for one lecture in 1965, no men would attend. So he walked in to the lecture theatre with about 50 women sitting there waiting attentively, but no men. He put his notes on the lectern and looked up. “Oh dear, nobody here today I see, might as well go home”! Picked up his notes and walked out. . Non-academic information on his life is hard to come by, mostly consisting of scattered remarks made by colleagues. He campaigned strenuously against the admission of female Fellows to
All Souls College All Souls College (official name: The College of All Souls of the Faithful Departed, of Oxford) is a constituent college of the University of Oxford in England. Unique to All Souls, all of its members automatically become fellows (i.e., full me ...
. Miriam Rothschild, an outstanding zoologist, was one of the few women with whom Ford was on good terms. Rothschild and Ford campaigned for the legalisation of male homosexuality in Britain. Ford was on good terms with
Theodosius Dobzhansky Theodosius Grigorievich Dobzhansky (; ; January 25, 1900 – December 18, 1975) was a Russian-born American geneticist and evolutionary biologist. He was a central figure in the field of evolutionary biology for his work in shaping the modern ...
, who did ground-breaking work on ecological genetics with ''
Drosophila ''Drosophila'' (), from Ancient Greek δρόσος (''drósos''), meaning "dew", and φίλος (''phílos''), meaning "loving", is a genus of fly, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or p ...
'' species: they exchanged letters and visits. Within the Department of Zoology at Oxford were a number of colleagues such as
Arthur Cain Arthur James Cain FRS (25 July 1921 – 20 August 1999) was a British evolutionary biologist and ecologist. He was elected a Fellow of the Royal Society in 1989. Career Arthur James Cain was born and grew up in Rugby in Warwickshire, England ...
with similar interests in investigating the role of natural selection; but Ford avoided the departmental teatime discussions and indeed most scientific discussion with these colleagues other than via the printed word.
Bryan Clarke Bryan Campbell Clarke (24 June 1932 – 27 February 2014) was a British Professor of genetics, latterly emeritus at the University of Nottingham. Clarke is particularly noted for his work on apostatic selection (which is a term he coine ...
wrote Ford's obituary in the ''
Biographical Memoirs of Fellows of the Royal Society The ''Biographical Memoirs of Fellows of the Royal Society'' is an academic journal on the history of science published annually by the Royal Society. It publishes obituaries of Fellows of the Royal Society. It was established in 1932 as ''Obi ...
'', but there are few other sources on his life.Marren P. 1995. ''The New Naturalists''. HarperCollins, London.


References

{{DEFAULTSORT:Ford, Edmund Brisco 1901 births 1988 deaths British evolutionary biologists English geneticists British eugenicists English science writers English lepidopterists Fellows of the Royal Society People educated at St Bees School Alumni of Wadham College, Oxford Modern synthesis (20th century) New Naturalist writers 20th-century British zoologists People from Dalton-in-Furness People associated with University College, Oxford