E.B. Ford
Encyclopedia
Edmund Brisco "Henry" Ford FRS Hon. FRCP (23 April 1901 – 21 January 1988) was a British
ecological geneticist
. He was a leader among those British biologists who investigated the role of natural selection
in nature. As a schoolboy Ford became interested in lepidoptera
, the group of insect
s which includes butterflies
and moth
s. He went on to study the genetics of natural populations, and invented the field of ecological genetics
. Ford was awarded the Royal Society
's Darwin Medal
in 1954. Later, in 1968, he was awarded UNESCO
's Kalinga Prize
for the popularisation of science.
Ford was born in Papcastle
, near Cockermouth
, Cumberland
, England
in 1901. He was educated at Wadham College, Oxford University, graduating in zoology in 1924.
, 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
, who did ground-breaking work on ecological genetics with Drosophila
species: they exchanged letters and visits.
Ford has a Royal Society
biographical memoir, but there are few other sources on his life.
. "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
and Pasteur
, both of whom he knew."
Ford was appointed University Demonstrator in Zoology in 1927 and Lecturer at University College, Oxford
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
of All Souls College since the seventeenth century.
Ford had a long working relationship with R.A. Fisher. By the time Ford had developed his formal definition of genetic polymorphism, 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
versus genetic drift
debate with Sewall Wright
, 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
. 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, 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
, which ran to four editions and was widely influential. 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
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
. He was a Fellow of All Souls College, and Honorary Fellow of Wadham College. He was elected FRS in 1946, and awarded the Darwin Medal
in 1954.
. 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 is more viable than either homozygote. That is a typical genetic mechanism for causing this type of polymorphism. The work involves a synthesis of field observations, taxonomy, and laboratory genetics.
during Kettlewell's famous experiments on the evolution of melanism in the peppered moth
, Biston betularia.
The entomologist Michael Majerus
discussed criticisms that had been made of Kettlewell's experimental methods in his 1998 book Melanism: Evolution in Action
. 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”. 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".
United Kingdom
The United Kingdom of Great Britain and Northern IrelandIn the United Kingdom and Dependencies, other languages have been officially recognised as legitimate autochthonous languages under the European Charter for Regional or Minority Languages...
ecological geneticist
Ecological genetics
Ecological genetics is the study of genetics in natural populations.This contrasts with classical genetics, which works mostly on crosses between laboratory strains, and DNA sequence analysis, which studies genes at the molecular level....
. He was a leader among those British biologists who investigated the role of natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
in nature. As a schoolboy Ford became interested in lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...
, the group of insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...
s which includes butterflies
Butterfly
A butterfly is a mainly day-flying insect of the order Lepidoptera, which includes the butterflies and moths. Like other holometabolous insects, the butterfly's life cycle consists of four parts: egg, larva, pupa and adult. Most species are diurnal. Butterflies have large, often brightly coloured...
and moth
Moth
A moth is an insect closely related to the butterfly, both being of the order Lepidoptera. Moths form the majority of this order; there are thought to be 150,000 to 250,000 different species of moth , with thousands of species yet to be described...
s. He went on to study the genetics of natural populations, and invented the field of ecological genetics
Ecological genetics
Ecological genetics is the study of genetics in natural populations.This contrasts with classical genetics, which works mostly on crosses between laboratory strains, and DNA sequence analysis, which studies genes at the molecular level....
. Ford was awarded the Royal Society
Royal Society
The Royal Society of London for Improving Natural Knowledge, known simply as the Royal Society, is a learned society for science, and is possibly the oldest such society in existence. Founded in November 1660, it was granted a Royal Charter by King Charles II as the "Royal Society of London"...
's Darwin Medal
Darwin Medal
The Darwin Medal is awarded by the Royal Society every alternate year for "work of acknowledged distinction in the broad area of biology in which Charles Darwin worked, notably in evolution, population biology, organismal biology and biological diversity". First awarded in 1890, it was created in...
in 1954. Later, in 1968, he was awarded UNESCO
UNESCO
The United Nations Educational, Scientific and Cultural Organization is a specialized agency of the United Nations...
's Kalinga Prize
Kalinga Prize
The Kalinga Prize for the Popularization of Science is an award given by UNESCO for exceptional skill in presenting scientific ideas to lay people...
for the popularisation of science.
Ford was born in Papcastle
Papcastle
Papcastle is a village and civil parish in the Borough of Allerdale in the English county of Cumbria. The village is now effectively a northern extension of Cockermouth, which lies to the south of the River Derwent. It has its own parish council and lies within Bridekirk Parish for Church of...
, near Cockermouth
Cockermouth
-History:The Romans created a fort at Derventio, now the adjoining village of Papcastle, to protect the river crossing, which had become located on a major route for troops heading towards Hadrian's Wall....
, Cumberland
Cumberland
Cumberland is a historic county of North West England, on the border with Scotland, from the 12th century until 1974. It formed an administrative county from 1889 to 1974 and now forms part of Cumbria....
, England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...
in 1901. He was educated at Wadham College, Oxford University, graduating in zoology in 1924.
Life
Ford never married, had no children, and was considered decidedly eccentric. 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. Miriam RothschildMiriam Rothschild
Dame Miriam Louisa Rothschild DBE, FRS was a British natural scientist and author with contributions to zoology, entomology, and botany.-Early life:...
, 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 Dobzhansky
Theodosius Grygorovych Dobzhansky ForMemRS was a prominent geneticist and evolutionary biologist, and a central figure in the field of evolutionary biology for his work in shaping the unifying modern evolutionary synthesis...
, who did ground-breaking work on ecological genetics with Drosophila
Drosophila
Drosophila is a genus of small flies, belonging to the family Drosophilidae, whose members are often called "fruit flies" or more appropriately pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species to linger around overripe or rotting fruit...
species: they exchanged letters and visits.
Ford has a Royal Society
Royal Society
The Royal Society of London for Improving Natural Knowledge, known simply as the Royal Society, is a learned society for science, and is possibly the oldest such society in existence. Founded in November 1660, it was granted a Royal Charter by King Charles II as the "Royal Society of London"...
biographical memoir, but there are few other sources on his life.
Career
Ford's career was based entirely at Oxford University. A.J. Cain said he took a degree in classics before turning to zoology. Ford read zoology at Oxford, and was taught genetics by Julian HuxleyJulian Huxley
Sir Julian Sorell Huxley FRS was an English evolutionary biologist, humanist and internationalist. He was a proponent of natural selection, and a leading figure in the mid-twentieth century evolutionary synthesis...
. "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
Ray Lankester
Sir E. Ray Lankester KCB, FRS was a British zoologist, born in London.An invertebrate zoologist and evolutionary biologist, he held chairs at University College London and Oxford University. He was the third Director of the Natural History Museum, and was awarded the Copley Medal of the Royal...
through E.B. Poulton. He was already an old man... but talked to me a good deal of Charles Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...
and Pasteur
Pasteur
Pasteur could refer to* Louis Pasteur , French chemist and microbiologist who invented:**Pasteurization**The pasteur pipette, both named after him-Things and places named after Louis Pasteur:* Pasteur Institute* Pasteur point, level of oxygen...
, both of whom he knew."
Ford was appointed University Demonstrator in Zoology in 1927 and Lecturer at University College, Oxford
University College, Oxford
.University College , is a constituent college of the University of Oxford in England. As of 2009 the college had an estimated financial endowment of £110m...
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
Fellow
A fellow in the broadest sense is someone who is an equal or a comrade. The term fellow is also used to describe a person, particularly by those in the upper social classes. It is most often used in an academic context: a fellow is often part of an elite group of learned people who are awarded...
of All Souls College since the seventeenth century.
Ford had a long working relationship with R.A. Fisher. By the time Ford had developed his formal definition of genetic polymorphism, 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
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....
versus genetic drift
Genetic drift
Genetic drift or allelic drift is the change in the frequency of a gene variant in a population due to random sampling.The alleles in the offspring are a sample of those in the parents, and chance has a role in determining whether a given individual survives and reproduces...
debate with Sewall Wright
Sewall Wright
Sewall Green Wright was an American geneticist known for his influential work on evolutionary theory and also for his work on path analysis. With R. A. Fisher and J.B.S. Haldane, he was a founder of theoretical population genetics. He is the discoverer of the inbreeding coefficient and of...
, 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
Theodosius Dobzhansky
Theodosius Grygorovych Dobzhansky ForMemRS was a prominent geneticist and evolutionary biologist, and a central figure in the field of evolutionary biology for his work in shaping the unifying modern evolutionary synthesis...
changed the emphasis in the third edition of his famous text from drift to selection.
Ecological genetics
Ecological genetics is the study of genetics in natural populations.This contrasts with classical genetics, which works mostly on crosses between laboratory strains, and DNA sequence analysis, which studies genes at the molecular level....
. 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, and predicted that human blood group polymorphisms
Human blood group systems
The International Society of Blood Transfusion currently recognises 30 major blood group systems . Thus, in addition to the ABO antigens and Rhesus antigens, many other antigens are expressed on the red blood cell surface membrane...
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 (book)
Ecological Genetics is a 1964 book by the British biologist E.B. Ford on ecological genetics. Ford founded the field and it is considered his magnum opus. The fourth and final edition was published in 1975....
, which ran to four editions and was widely influential. 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
New Naturalist
The New Naturalist Library books are a series published by Collins in the United Kingdom, on a variety of natural history topics relevant to the British Isles...
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
University of Oxford
The University of Oxford is a university located in Oxford, United Kingdom. It is the second-oldest surviving university in the world and the oldest in the English-speaking world. Although its exact date of foundation is unclear, there is evidence of teaching as far back as 1096...
. He was a Fellow of All Souls College, and Honorary Fellow of Wadham College. He was elected FRS in 1946, and awarded the Darwin Medal
Darwin Medal
The Darwin Medal is awarded by the Royal Society every alternate year for "work of acknowledged distinction in the broad area of biology in which Charles Darwin worked, notably in evolution, population biology, organismal biology and biological diversity". First awarded in 1890, it was created in...
in 1954.
Ecological genetics
E.B. Ford worked for many years on genetic polymorphismPolymorphism (biology)
Polymorphism in biology occurs when two or more clearly different phenotypes exist in the same population of a species — in other words, the occurrence of more than one form or morph...
. 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 is more viable than either homozygote. That is a typical genetic mechanism for causing this type of polymorphism. The work involves a synthesis of field observations, taxonomy, and laboratory genetics.
Peppered moth
Ford was the supervisor of Bernard KettlewellBernard Kettlewell
Henry Bernard Davis Kettlewell was a British geneticist, lepidopterist and medical doctor, who carried out research into the influence of industrial melanism on natural selection in moths, showing why moths are darker in polluted areas.-Early life:Kettlewell was born in Howden, Yorkshire, and...
during Kettlewell's famous experiments on the evolution of melanism in the peppered moth
Peppered moth evolution
The evolution of the peppered moth over the last two hundred years has been studied in detail. Originally, the vast majority of peppered moths had light colouration, which effectively camouflaged them against the light-coloured trees and lichens which they rested upon...
, Biston betularia.
The entomologist Michael Majerus
Michael Majerus
Michael Eugene Nicolas Majerus was a geneticist and Professor of Ecology at Clare College, Cambridge, an enthusiast who became a world authority in his field of evolutionary biology. He was widely noted for his work on moths and ladybirds and as an advocate of the science of evolution...
discussed criticisms that had been made of Kettlewell's experimental methods in his 1998 book Melanism: Evolution in Action
Melanism: Evolution in Action
thumb|220px|Melanism in ActionMelanism: Evolution in Action is a book by Dr Mike Majerus, published in 1998. It is an update of Bernard Kettlewell's book The Evolution of Melanism....
. This book was misrepresented in reviews, and the story was picked up by creationist
Creationism
Creationism is the religious beliefthat humanity, life, the Earth, and the universe are the creation of a supernatural being, most often referring to the Abrahamic god. As science developed from the 18th century onwards, various views developed which aimed to reconcile science with the Genesis...
campaigners. In her controversial book Of Moths and Men
Of Moths and Men
Of Moths and Men is a controversial book by the journalist Judith Hooper about the Oxford University ecological genetics school led by E.B. Ford. The book specifically concerns Bernard Kettlewell's experiments on the peppered moth which were intended as experimental validation of evolution...
, Judith Hooper
Judith Hooper
Judith Hooper is an American journalist.Hooper has worked as an editor and writer for the magazine Omni. With her husband, Dick Teresi, she co-wrote the books The Three-Pound Universe and Would the Buddha Wear a Walkman? A Catalogue of Revolutionary Tools for Higher Consciousness...
(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”. 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".
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.Julian HuxleySir Julian Sorell Huxley FRS was an English evolutionary biologist, humanist and internationalist. He was a proponent of natural selection, and a leading figure in the mid-twentieth century evolutionary synthesis...
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 NaturalistNew NaturalistThe New Naturalist Library books are a series published by Collins in the United Kingdom, on a variety of natural history topics relevant to the British Isles...
#1 Collins, London. - Ford E.B. (1951). British butterflies. Penguin BooksPenguin BooksPenguin Books is a publisher founded in 1935 by Sir Allen Lane and V.K. Krishna Menon. Penguin revolutionised publishing in the 1930s through its high quality, inexpensive paperbacks, sold through Woolworths and other high street stores for sixpence. Penguin's success demonstrated that large...
, London. - Ford E.B. (1954, 3rd edn 1972). Moths. New NaturalistNew NaturalistThe New Naturalist Library books are a series published by Collins in the United Kingdom, on a variety of natural history topics relevant to the British Isles...
#30 HarperCollins, 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.Ernst MayrErnst Walter Mayr was one of the 20th century's leading evolutionary biologists. He was also a renowned taxonomist, tropical explorer, ornithologist, historian of science, and naturalist...
and Provine W.B. (eds) The evolutionary synthesis: perspectives on the unification of biology. Harvard 1980; 1998. [effectively, 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.
See also
- TeänTeänTeän is one of the uninhabited islands to the north of the Isles of Scilly archipelago between Tresco, 1.5 km to the west and St Martin's 300 m to the east....
, Isles of Scilly: the site of Ford's Common Blue (Polyommatus icarus) population study.
External links
- Papers co-written with R.A. FisherRonald FisherSir Ronald Aylmer Fisher FRS was an English statistician, evolutionary biologist, eugenicist and geneticist. Among other things, Fisher is well known for his contributions to statistics by creating Fisher's exact test and Fisher's equation...
are available on the University of AdelaideUniversity of AdelaideThe University of Adelaide is a public university located in Adelaide, South Australia. Established in 1874, it is the third oldest university in Australia...
's website at