Arthur Cain

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Arthur James Cain FRS (25 July 1921 – 20 August 1999) [1] was a British evolutionary biologist and ecologist. He was elected a Fellow of the Royal Society in 1989.

Contents

Career

Arthur James Cain was born and grew up in Rugby in Warwickshire, England. In 1939 he was awarded a prestigious scholarship (Demyship) 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, finishing his D. Phil. in 1948. From January 1949 until 1964 Cain was employed as University Demonstrator in Animal Taxonomy. In addition he was appointed Curator of the Zoological Collections at the Oxford University Museum in 1954, besides serving as Lecturer in Zoology at St Peter's College (1958–1961).

In 1964, Cain 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. [1] [2] [3]

Scientific activities

Cain's main interests lay in evolutionary biology, ecological genetics, animal taxonomy and speciation.

Though he initially 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. [4]

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 reminisces on pre-war Oxford

Towards the end of his life Cain was persuaded to reminisce about the status of natural selection in pre-war Oxford and how it changed over the years of the modern evolutionary synthesis. [5] [6] 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. [7]

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. [8]

Also, Robson and Richards [9] "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).

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. [10] 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. [11]

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. [12] [13] 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. [14] Also, as Cain's own research showed, much polymorphism is maintained by differential selection in the diversity of environments within a species' range. [15] [16] [17]

Notable publications

(a full bibliography listing 148 items appears as supplementary material to Bryan Clarke's obituary [1] )

Cain A.J. 1954. Animal species and their 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 B246, 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 B253, 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. Genetics45, 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.). Heredity4, 275–94.

Cain A.J. and Sheppard P.M. 1954. Natural selection in Cepaea. Genetics39, 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.

Related Research Articles

<span class="mw-page-title-main">Modern synthesis (20th century)</span> Fusion of natural selection with Mendelian inheritance

The modern synthesis was the early 20th-century synthesis of Charles Darwin's theory of evolution and Gregor Mendel's ideas on heredity into a joint mathematical framework. Julian Huxley coined the term in his 1942 book, Evolution: The Modern Synthesis. The synthesis combined the ideas of natural selection, Mendelian genetics, and population genetics. It also related the broad-scale macroevolution seen by palaeontologists to the small-scale microevolution of local populations.

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<span class="mw-page-title-main">E. B. Ford</span> British ecological geneticist (1901–1988)

Edmund Brisco "Henry" Ford 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. 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. In the wider world his best known work is Butterflies (1945).

Ecological genetics is the study of genetics in natural populations. It combines ecology, evolution, and genetics to understand the processes behind adaptation.

<span class="mw-page-title-main">Polymorphism (biology)</span> Occurrence of two or more clearly different morphs or forms in the population of a species

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Frequency-dependent selection is an evolutionary process by which the fitness of a phenotype or genotype depends on the phenotype or genotype composition of a given population.

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References

  1. 1 2 3 Clarke, B.C. (2008). "Arthur James Cain. 25 July 1921 – 20 August 1999". Biographical Memoirs of Fellows of the Royal Society . 54: 47–57. doi:10.1098/rsbm.2007.0034. S2CID   73131707.
  2. ↑ "Arthur J. Cain Papers Mss.Ms.Coll.63". American Philosophical Society. Retrieved 2 July 2024.
  3. ↑ Cook, L. (2000). "A.J. Cain F.R.S. 1921-1929". Malacological Bulletin. 34. Retrieved 2 July 2024.
  4. ↑ Cook, L.M. (2024). "Arthur Cain and ecological genetics in the Oxford Zoology Department". Archives of Natural History. 51 (1): 73–85. doi:10.3366/anh.20240897.
  5. ↑ Cain A.J. and Provine W.B. 1991. Genes and ecology in history. In Berry R.J. & others (eds) Genes in ecology: the 33rd Symposium of the British Ecological Society. Blackwell, Oxford.
  6. ↑ Huxley J. 1942. Evolution: the modern synthesis (2nd ed 1963, 3rd ed 1974)
  7. ↑ Elton C.S. 1927. Animal ecology.
  8. ↑ Cain & Provine W.B. 1991. Genes and ecology in history. In Berry R.J. Genes in ecology p5
  9. ↑ Robson G.C. and Richards O.W. 1936. The variation of animals in nature. Longmans, London.
  10. ↑ Cain & Provine W.B. 1991. Genes and ecology in history. In Berry R.J. Genes in ecology p8
  11. ↑ Cain & Provine W.B. 1991. Genes and ecology in history. In Berry R.J. Genes in ecology p9
  12. ↑ Ford E.B. 1964, 4th edn 1975. Ecological genetics. Chapman and Hall, London
  13. ↑ Ford E.B. 1965. Genetic polymorphism. All Souls Studies, Faber & Faber, London
  14. ↑ Fisher R.A. 1930. Genetical theory of natural selection p80.
  15. ↑ Cain A.J. and Currey J.D. 1963. Area effects in Cepaea. Phil Trans Roy Soc B246, 269–299.
  16. ↑ Cain A.J. and Currey J.D. 1968. Ecogenetics of a population of Cepaea nemoralis subject to strong area effects. Phil Trans Roy Soc B253, 447–482.
  17. ↑ Clarke B.C. 1979. The evolution of genetic diversity. Proc Roy Soc B. 205, 453–474.
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