Arthur Cain
Encyclopedia
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.
) to Magdalen College, Oxford
, where he graduated with first class honors in zoology in 1941. Entering the British Army
in December 1941, Cain was commissioned Second Lieutenant
in the Royal Army Ordnance Corps
(engineering) and was later transferred to the Royal Electrical and Mechanical Engineers
(R.E.M.E.) on its formation. He was promoted to Captain
in 1942.
After leaving the military in November 1945 Cain returned to Oxford to pursue research in the Department of Zoology. He became a Departmental Demonstrator in October 1946, and received his M.A. in November 1947. From January 1949 until 1964 Cain was employed as University Demonstrator (now referred to as Lecturer) in Animal Taxonomy.
, animal taxonomy
and speciation
.
Though he conducted research with John Baker
on the histochemistry of lipids, his main work lay in the field developed by E.B. Ford
, namely, ecological genetics
. With P.M. Sheppard, Cain studied the ecological genetics of colour and banding polymorphisms
in snails. Cain and Sheppard's work on Cepaea nemoralis, one of the first studies to demonstrate natural selection by predators acting on a colour polymorphism, is now regarded as a classic. It generated a long series of further studies by Cain, including the formal genetic analysis
of the variation, the discovery of area effects and the analysis of climatic influences. With John Currey he made elegant use of sub-fossil material to follow changes in time as well as space. Later he turned to the study of variation in shell shape.
In population genetics he clarified the concept of adaptive value
. He made important contributions to the theory and practice of taxonomy
, the problems of homology
, phyletic weighting and taxonomic importance, on the status of the genus
, and on the relevance of natural selection
to our understanding of variation between taxonomic categories.
Cain was appointed Curator
of the Zoological Collections at the Oxford University Museum in 1954, a position he held for ten years in addition to his duties as University Lecturer and as Lecturer in Zoology at Saint Peter's College
(1958–1961). In 1964, he left Oxford to become Professor of Zoology at the University of Manchester
, and he later (1968) was appointed Derby Professor of Zoology at the University of Liverpool
. He received emeritus
status at Liverpool upon his retirement in 1989.
in pre-war Oxford and how it changed over the years of the modern evolutionary synthesis
. The general attitude was sceptical of natural selection. Charles Elton
, who led the emergence of ecology
as a discipline, pointed out the arctic fox
polymorphism, which can be found in all three tundra
biomes of the northern palaeoarctic. Arctic foxes (Alopex lagopus) are dimorphic: the common morph ('white') is white in winter and brownish-grey dorsally in summer; the other morph ('blue') is light brown/blue in winter and dark brown in summer. The two morphs interbreed freely. Despite the obvious advantage of white in avoiding predation, blue is actually the most frequent morph in Iceland
. Elton also gave a number of other examples which he claimed could not be explained by natural selection.
Also, Robson and Richards "showed a surprising reluctance to allow any example of natural selection; their cautious qualification that characters were non-adaptive as far as they could see became, too often, a certainty that they were non-adaptive; and their arguments were sometimes one-sided." Cain laid the blame on their "vitalistic or perhaps theistic attitudes... Robson and Richards were far from alone. Alister Hardy
... was an earnest Unitarian
and certainly a vitalist" (p7)... In Cambridge matters were even worse" (p8, giving as examples W.H. Thorpe
, Charles Raven
, Sir James Gray
and J.W.S. Pringle).
He might have added, had he known them, Ronald Fisher
and Theodosius Dobzhansky
, who were also believing Christians: Fisher from the start of his career was a leading proponent of natural selection.
In contrast to many others, E.B. Ford
appreciated that, even if a character was in itself non-adaptive, the gene or genes determining it might affect other, adaptive, characters which were always under selective influence. Ford understood the significance of pleiotropism, and knew of Fisher
's demonstration that a neutral gene derived from a single mutation could only be in about the same number of individuals as there had been generations since its inception. Also, as Cain's own research showed, much polymorphism
is maintained by differential selection in the diversity of environments within a species' range.
Cain A.J. 1968. Studies on Cepaea V. Phil. Trans. R. Soc. B 253, 499-517.
Cain A.J. 1971. Colour and banding morphs in subfossil samples of the snail Cepaea. In Creed R. (ed) Ecological genetics and evolution. Blackwell, Oxford.
Cain A.J. 1977. The efficacy of natural selection in wild populations. In The changing scene in natural sciences. Special publication #12, 111-33. Academy of Natural Sciences.
Cain A.J. 1983. Ecology and ecogenetics of terrestrial molluscan populations. In Russell-Hunter W.D. (ed) The Mollusca vol 6, p597-647. Academic Press, N.Y.
Cain A.J. and Currey J.D. 1963a. Area effects in Cepaea. Phil Trans Roy Soc B 246, 269-299.
Cain A.J. and Currey J.D. 1963b. Area effects in Cepaea on the Larkhill Artillery Ranges, Salisbury Plain. J. Linnaean Soc London (Zoology) 45, 1-15.
Cain A.J. and Currey J.D. 1968. Ecogenetics of a population of Cepaea nemoralis subject to strong area effects. Phil Trans Roy Soc B 253, 447-482.
Cain A.J., King J.M.B. and Sheppard P.M. 1960. New data on the genetics of polymorphism in the snail Cepaea nemoralis. Genetics 45, 393-411.
Cain A.J. and Provine W.B. 1991. Genes and ecology in history. In Berry R.J. et al. (eds) Genes in ecology: the 33rd Symposium of the British Ecological Society. Blackwell, Oxford.
Cain A.J. and Sheppard P.M. 1950. Selection in the polymorphic land snail Cepaea nemoralis (L.). Heredity 4, 275-94.
Cain A.J. and Sheppard P.M. 1954. Natural selection in Cepaea. Genetics 39, 89-116.
Cain A.J., Sheppard P.M. and King J.M.B. 1968. Studies on Cepaea I. The genetics of some morphs and varieties of Cepaea nemoralis (L.). Phil. Trans. R. Soc. B 253, 383-396.
Clarke B.C. 1979. The evolution of genetic diversity. Proc Roy Soc B. 205, 453-474. [a general review]
Currey J.D. and Cain A.J. 1968. Climate and selection of banding morphs in Cepaea from the climate optimum to the present day. Phil. Trans. R. Soc. B 253, 483-98.
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...
evolutionary biologist and ecologist. He was elected a Fellow of the Royal Society in 1989.
Life
Arthur James Cain was awarded an open scholarship in 1939 (DemyshipDemyship
A demyship is a form of scholarship, specifically at Magdalen College, Oxford. Oscar Wilde, Lewis Gielgud, Lord Denning andT. E. Lawrence were famous recipients. It is derived from demi-socii or half-fellows. Magdalen's founder, William of Waynflete, originally provided them for the College...
) to Magdalen College, Oxford
Magdalen College, Oxford
Magdalen College is one of the constituent colleges of the University of Oxford in England. As of 2006 the college had an estimated financial endowment of £153 million. Magdalen is currently top of the Norrington Table after over half of its 2010 finalists received first-class degrees, a record...
, where he graduated with first class honors in zoology in 1941. Entering the British Army
British Army
The British Army is the land warfare branch of Her Majesty's Armed Forces in the United Kingdom. It came into being with the unification of the Kingdom of England and Scotland into the Kingdom of Great Britain in 1707. The new British Army incorporated Regiments that had already existed in England...
in December 1941, Cain was commissioned Second Lieutenant
Second Lieutenant
Second lieutenant is a junior commissioned officer military rank in many armed forces.- United Kingdom and Commonwealth :The rank second lieutenant was introduced throughout the British Army in 1871 to replace the rank of ensign , although it had long been used in the Royal Artillery, Royal...
in the Royal Army Ordnance Corps
Royal Army Ordnance Corps
The Royal Army Ordnance Corps was a corps of the British Army. It dealt only with the supply and maintenance of weaponry, munitions and other military equipment until 1965, when it took over most other supply functions, as well as the provision of staff clerks, from the Royal Army Service...
(engineering) and was later transferred to the Royal Electrical and Mechanical Engineers
Royal Electrical and Mechanical Engineers
The Corps of Royal Electrical and Mechanical Engineers is a corps of the British Army that has responsibility for the maintenance, servicing and inspection of almost every electrical and mechanical piece of equipment within the British Army from Challenger II main battle tanks and WAH64 Apache...
(R.E.M.E.) on its formation. He was promoted to Captain
Captain (British Army and Royal Marines)
Captain is a junior officer rank of the British Army and Royal Marines. It ranks above Lieutenant and below Major and has a NATO ranking code of OF-2. The rank is equivalent to a Lieutenant in the Royal Navy and to a Flight Lieutenant in the Royal Air Force...
in 1942.
After leaving the military in November 1945 Cain returned to Oxford to pursue research in the Department of Zoology. He became a Departmental Demonstrator in October 1946, and received his M.A. in November 1947. From January 1949 until 1964 Cain was employed as University Demonstrator (now referred to as Lecturer) in Animal Taxonomy.
Career
Cain's main interests lay in evolutionary biology, ecological geneticsEcological 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....
, animal taxonomy
Systematics
Biological systematics is the study of the diversification of terrestrial life, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees...
and speciation
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...
.
Though he conducted research with John Baker
John Baker (biologist)
John Randal Baker FRS was a biologist, physical anthropologist, and professor at the University of Oxford in the mid-twentieth century. He is best remembered for his 1974 book, Race, which classifies human races in the same way in which animal subspecies are classified...
on the histochemistry of lipids, his main work lay in the field developed by E.B. Ford
E.B. Ford
Edmund Brisco "Henry" Ford FRS Hon. FRCP 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 insects which includes butterflies and moths...
, namely, 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....
. With P.M. Sheppard, Cain studied the ecological genetics of colour and banding polymorphisms
Polymorphism (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...
in snails. Cain and Sheppard's work on Cepaea nemoralis, one of the first studies to demonstrate natural selection by predators acting on a colour polymorphism, is now regarded as a classic. It generated a long series of further studies by Cain, including the formal genetic analysis
Genetic analysis
Genetic analysis can be used generally to describe methods both used in and resulting from the sciences of genetics and molecular biology, or to applications resulting from this research....
of the variation, the discovery of area effects and the analysis of climatic influences. With John Currey he made elegant use of sub-fossil material to follow changes in time as well as space. Later he turned to the study of variation in shell shape.
In population genetics he clarified the concept of adaptive value
Adaptive value
The adaptive value represents the combined influence of all characters which affect the fitness of an individual or population....
. He made important contributions to the theory and practice of taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...
, the problems of homology
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...
, phyletic weighting and taxonomic importance, on the status of the genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
, and on the relevance 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....
to our understanding of variation between taxonomic categories.
Cain was appointed Curator
Curator
A curator is a manager or overseer. Traditionally, a curator or keeper of a cultural heritage institution is a content specialist responsible for an institution's collections and involved with the interpretation of heritage material...
of the Zoological Collections at the Oxford University Museum in 1954, a position he held for ten years in addition to his duties as University Lecturer and as Lecturer in Zoology at Saint Peter's College
St Peter's College, Oxford
St Peter's College is one of the constituent colleges of the University of Oxford, Oxford, United Kingdom, located in New Inn Hall Street. It occupies the site of two of the University's oldest Inns, or medieval hostels - Bishop Trellick's, later New Inn Hall, and Rose Hall - both of which were...
(1958–1961). In 1964, he left Oxford to become Professor of Zoology at the University of Manchester
University of Manchester
The University of Manchester is a public research university located in Manchester, United Kingdom. It is a "red brick" university and a member of the Russell Group of research-intensive British universities and the N8 Group...
, and he later (1968) was appointed Derby Professor of Zoology at the University of Liverpool
University of Liverpool
The University of Liverpool is a teaching and research university in the city of Liverpool, England. It is a member of the Russell Group of large research-intensive universities and the N8 Group for research collaboration. Founded in 1881 , it is also one of the six original "red brick" civic...
. He received emeritus
Emeritus
Emeritus is a post-positive adjective that is used to designate a retired professor, bishop, or other professional or as a title. The female equivalent emerita is also sometimes used.-History:...
status at Liverpool upon his retirement in 1989.
Cain reminisces on pre-war Oxford
Towards the end of his life Cain was persuaded to reminisce about the status of natural selectionNatural 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 pre-war Oxford and how it changed over the years of the modern evolutionary synthesis
Modern evolutionary synthesis
The modern evolutionary synthesis is a union of ideas from several biological specialties which provides a widely accepted account of evolution...
. The general attitude was sceptical of natural selection. Charles Elton
Charles Elton
Charles Elton may refer to:* Charles Isaac Elton , English lawyer, politician, writer and antiquarian* Charles Sutherland Elton , English biologist* Charles Elton , Chief of Police in Los Angeles, California...
, who led the emergence of ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...
as a discipline, pointed out the arctic fox
Arctic fox
The arctic fox , also known as the white fox, polar fox or snow fox, is a small fox native to Arctic regions of the Northern Hemisphere and is common throughout the Arctic tundra biome. The Greek word alopex, means a fox and Vulpes is the Latin version...
polymorphism, which can be found in all three tundra
Tundra
In physical geography, tundra is a biome where the tree growth is hindered by low temperatures and short growing seasons. The term tundra comes through Russian тундра from the Kildin Sami word tūndâr "uplands," "treeless mountain tract." There are three types of tundra: Arctic tundra, alpine...
biomes of the northern palaeoarctic. Arctic foxes (Alopex lagopus) are dimorphic: the common morph ('white') is white in winter and brownish-grey dorsally in summer; the other morph ('blue') is light brown/blue in winter and dark brown in summer. The two morphs interbreed freely. Despite the obvious advantage of white in avoiding predation, blue is actually the most frequent morph in Iceland
Iceland
Iceland , described as the Republic of Iceland, is a Nordic and European island country in the North Atlantic Ocean, on the Mid-Atlantic Ridge. Iceland also refers to the main island of the country, which contains almost all the population and almost all the land area. The country has a population...
. Elton also gave a number of other examples which he claimed could not be explained by natural selection.
- That very phenomenon which was to be used by Fisher & Ford in studies on natural selection is here shown by cogent argument and the facts of field natural history to be [apparently] inexplicable by selection. But Elton knew that a far greater range of other characters have the same implications, namely, all, or nearly all, the differences (non-polymorphic) between closely related species. Cain.
Also, Robson and Richards "showed a surprising reluctance to allow any example of natural selection; their cautious qualification that characters were non-adaptive as far as they could see became, too often, a certainty that they were non-adaptive; and their arguments were sometimes one-sided." Cain laid the blame on their "vitalistic or perhaps theistic attitudes... Robson and Richards were far from alone. Alister Hardy
Alister Hardy
Sir Alister Clavering Hardy, FRS was an English marine biologist, expert on zooplankton and marine ecosystems...
... was an earnest Unitarian
Unitarianism
Unitarianism is a Christian theological movement, named for its understanding of God as one person, in direct contrast to Trinitarianism which defines God as three persons coexisting consubstantially as one in being....
and certainly a vitalist" (p7)... In Cambridge matters were even worse" (p8, giving as examples W.H. Thorpe
William Homan Thorpe
William Homan Thorpe FRS was Professor of Animal Ethology at the University of Cambridge, and a significant British zoologist, ethologist and ornithologist....
, Charles Raven
Charles Raven
Charles Earle Raven was an English theologian, Regius Professor of Divinity at Cambridge University, and Master of Christ's College, Cambridge...
, Sir James Gray
James Gray
James Gray may refer to:* Sir James Gray, 1st Baronet , armiger and merchant-Burgess of Edinburgh* Sir James Gray, 2nd Baronet , diplomat and antiquary* The male pseudonym of Hannah Snell...
and J.W.S. Pringle).
- What I wanted to know from all these great people was, how exactly did they know that a character was non-adaptive or neutral? They didn't know, and they couldn't know. Cain. This was the stimulus for Cain's research on evolution in natural communities.
- David Lack was the only religious man I knew at that period who did not allow his religion to dictate his view of natural selection. Cain.
He might have added, had he known them, Ronald Fisher
Ronald Fisher
Sir 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...
and 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 were also believing Christians: Fisher from the start of his career was a leading proponent of natural selection.
In contrast to many others, E.B. Ford
E.B. Ford
Edmund Brisco "Henry" Ford FRS Hon. FRCP 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 insects which includes butterflies and moths...
appreciated that, even if a character was in itself non-adaptive, the gene or genes determining it might affect other, adaptive, characters which were always under selective influence. Ford understood the significance of pleiotropism, and knew of Fisher
Ronald Fisher
Sir 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...
's demonstration that a neutral gene derived from a single mutation could only be in about the same number of individuals as there had been generations since its inception. Also, as Cain's own research showed, much polymorphism
Polymorphism (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...
is maintained by differential selection in the diversity of environments within a species' range.
Notable publications
Cain A.J. 1954. Animal species and evolution. Hutchinson, London.Cain A.J. 1968. Studies on Cepaea V. Phil. Trans. R. Soc. B 253, 499-517.
Cain A.J. 1971. Colour and banding morphs in subfossil samples of the snail Cepaea. In Creed R. (ed) Ecological genetics and evolution. Blackwell, Oxford.
Cain A.J. 1977. The efficacy of natural selection in wild populations. In The changing scene in natural sciences. Special publication #12, 111-33. Academy of Natural Sciences.
Cain A.J. 1983. Ecology and ecogenetics of terrestrial molluscan populations. In Russell-Hunter W.D. (ed) The Mollusca vol 6, p597-647. Academic Press, N.Y.
Cain A.J. and Currey J.D. 1963a. Area effects in Cepaea. Phil Trans Roy Soc B 246, 269-299.
Cain A.J. and Currey J.D. 1963b. Area effects in Cepaea on the Larkhill Artillery Ranges, Salisbury Plain. J. Linnaean Soc London (Zoology) 45, 1-15.
Cain A.J. and Currey J.D. 1968. Ecogenetics of a population of Cepaea nemoralis subject to strong area effects. Phil Trans Roy Soc B 253, 447-482.
Cain A.J., King J.M.B. and Sheppard P.M. 1960. New data on the genetics of polymorphism in the snail Cepaea nemoralis. Genetics 45, 393-411.
Cain A.J. and Provine W.B. 1991. Genes and ecology in history. In Berry R.J. et al. (eds) Genes in ecology: the 33rd Symposium of the British Ecological Society. Blackwell, Oxford.
Cain A.J. and Sheppard P.M. 1950. Selection in the polymorphic land snail Cepaea nemoralis (L.). Heredity 4, 275-94.
Cain A.J. and Sheppard P.M. 1954. Natural selection in Cepaea. Genetics 39, 89-116.
Cain A.J., Sheppard P.M. and King J.M.B. 1968. Studies on Cepaea I. The genetics of some morphs and varieties of Cepaea nemoralis (L.). Phil. Trans. R. Soc. B 253, 383-396.
Clarke B.C. 1979. The evolution of genetic diversity. Proc Roy Soc B. 205, 453-474. [a general review]
Currey J.D. and Cain A.J. 1968. Climate and selection of banding morphs in Cepaea from the climate optimum to the present day. Phil. Trans. R. Soc. B 253, 483-98.
External links
- http://www.amphilsoc.org/library/mole/c/cain.htm