Arthur James Cain FRS (25 July 1921 – 20 August 1999) was a British

evolutionary biologist Evolutionary biology is the subfield of biology that studies the evolutionary processes such as natural selection, common descent, and speciation that produced the diversity of life on Earth. In the 1930s, the discipline of evolutionary biol ...

and

ecologist Ecology () is the natural science of the relationships among living organisms and their environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere levels. Ecology overlaps with the closely re ...

. He was elected a

Fellow of the Royal Society Fellowship of the Royal Society (FRS, ForMemRS and HonFRS) is an award granted by the Fellows of the Royal Society of London to individuals who have made a "substantial contribution to the improvement of natural science, natural knowledge, incl ...

in 1989.

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 Magdalen College ( ) is a Colleges of the University of Oxford, constituent college of the University of Oxford. It was founded in 1458 by Bishop of Winchester William of Waynflete. It is one of the wealthiest Oxford colleges, as of 2022, and ...

, 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 The Royal Army Ordnance Corps (RAOC) was a corps of the British Army. At its renaming as a Royal Corps in 1918 it was both a supply and repair corps. In the supply area it had responsibility for weapons, armoured vehicles and other military equi ...

(engineering) and was later transferred to the

Royal Electrical and Mechanical Engineers The Corps of Royal Electrical and Mechanical Engineers (REME ) is the maintenance arm of the British Army that maintains the equipment that the Army uses. The corps is described as the "British Army's professional engineers". History Prior t ...

(R.E.M.E.) on its formation. He was promoted to

captain Captain is a title, an appellative for the commanding officer of a military unit; the supreme leader or highest rank officer of a navy ship, merchant ship, aeroplane, spacecraft, or other vessel; or the commander of a port, fire or police depa ...

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 A curator (from , meaning 'to take care') is a manager or overseer. When working with cultural organizations, a curator is typically a "collections curator" or an "exhibitions curator", and has multifaceted tasks dependent on the particular ins ...

of the Zoological Collections at the

Oxford University Museum The Oxford University Museum of Natural History (OUMNH) is a museum displaying many of the University of Oxford's natural history specimens, located on Parks Road in Oxford, England. It also contains a lecture theatre which is used by the univers ...

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 The University of Manchester is a public university, public research university in Manchester, England. The main campus is south of Manchester city centre, Manchester City Centre on Wilmslow Road, Oxford Road. The University of Manchester is c ...

, and he later (1968) was appointed Derby Professor of Zoology at the

University of Liverpool The University of Liverpool (abbreviated UOL) is a Public university, public research university in Liverpool, England. Founded in 1881 as University College Liverpool, Victoria University (United Kingdom), Victoria University, it received Ro ...

. He received

emeritus ''Emeritus/Emerita'' () is an honorary title granted to someone who retires from a position of distinction, most commonly an academic faculty position, but is allowed to continue using the previous title, as in "professor emeritus". In some c ...

status at Liverpool upon his retirement in 1989.

Scientific activities

Cain's main interests lay in

evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes such as natural selection, common descent, and speciation that produced the diversity of life on Earth. In the 1930s, the discipline of evolutionary biolo ...

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 ...

, animal taxonomy and

speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...

. Though he initially conducted research with John Baker on the

histochemistry Immunohistochemistry is a form of immunostaining. It involves the process of selectively identifying antigens in cells and tissue, by exploiting the principle of antibodies binding specifically to antigens in biological tissues. Albert Hewett ...

lipids Lipids are a broad group of organic compounds which include fats, waxes, sterols, fat-soluble vitamins (such as vitamins Vitamin A, A, Vitamin D, D, Vitamin E, E and Vitamin K, K), monoglycerides, diglycerides, phospholipids, and others. The fu ...

, his main work lay in the field developed by E.B. Ford, namely,

. With P.M. Sheppard, Cain studied the ecological genetics of colour and banding polymorphisms in

snails A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...

. 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 is the overall process of studying and researching in fields of science that involve genetics and molecular biology. There are a number of applications that are developed from this research, and these are also considered parts ...

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 image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...

, the problems of homology, phyletic weighting and taxonomic importance, on the status of the

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

, and on the relevance 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 ...

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

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 Ecology () is the natural science of the relationships among living organisms and their Natural environment, environment. Ecology considers organisms at the individual, population, community (ecology), community, ecosystem, and biosphere lev ...

as a discipline, pointed out the

Arctic fox The Arctic fox (''Vulpes lagopus''), also known as the white fox, polar fox, or snow fox, is a small species of fox native to the Arctic regions of the Northern Hemisphere and common throughout the Tundra#Arctic tundra, Arctic tundra biome. I ...

polymorphism, which can be found in all three

tundra In physical geography, a tundra () is a type of biome where tree growth is hindered by frigid temperatures and short growing seasons. There are three regions and associated types of tundra: #Arctic, Arctic, Alpine tundra, Alpine, and #Antarctic ...

biomes A biome () is a distinct geographical region with specific climate, vegetation, and animal life. It consists of a biological community (ecology), community that has formed in response to its physical environment and regional climate. In 1935, Art ...

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 is a Nordic countries, Nordic island country between the Atlantic Ocean, North Atlantic and Arctic Oceans, on the Mid-Atlantic Ridge between North America and Europe. It is culturally and politically linked with Europe and is the regi ...

. 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 pparentlyinexplicable 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 Sir Alister Clavering Hardy (10 February 1896 – 22 May 1985) was a British Marine biology, marine biologist, an expert on marine ecosystems spanning organisms from zooplankton to whales. He had the artistic skill to illustrate his books with h ...

... was an earnest Unitarian and certainly a

vitalist Vitalism is a belief that starts from the premise that "living organisms are fundamentally different from non-living entities because they contain some non-physical element or are governed by different principles than are inanimate things." Wher ...

" (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. 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 Sir Ronald Aylmer Fisher (17 February 1890 – 29 July 1962) was a British polymath who was active as a mathematician, statistician, biologist, geneticist, and academic. For his work in statistics, he has been described as "a genius who a ...

and

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 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.Clarke B.C. 1979. The evolution of genetic diversity. ''Proc Roy Soc B''. 205, 453–474.

Notable publications

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

References

External links

Collection of papers held by the American Philosophical Society
{{DEFAULTSORT:Cain, Arthur James 20th-century British biologists British evolutionary biologists British ecologists Royal Electrical and Mechanical Engineers officers Royal Army Ordnance Corps officers Fellows of the Royal Society 1921 births 1999 deaths British Army personnel of World War II Academics of the University of Oxford Alumni of Magdalen College, Oxford