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Sir Edward Bagnall Poulton, FRS HFRSE FLS (27 January 1856 – 20 November 1943) was a British
evolutionary biologist Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life fo ...
, a lifelong advocate 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 heritable traits characteristic of a population over generations. Char ...
through a period in which many scientists such as
Reginald Punnett Reginald Crundall Punnett FRS (; 20 June 1875 – 3 January 1967) was a British geneticist who co-founded, with William Bateson, the '' Journal of Genetics'' in 1910. Punnett is probably best remembered today as the creator of the Punne ...
doubted its importance. He invented the term
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species s ...
for evolution of species in the same place, and in his book ''
The Colours of Animals ''The Colours of Animals'' is a zoology book written in 1890 by Sir Edward Bagnall Poulton (1856–1943). It was the first substantial textbook to argue the case for Darwinian selection applying to all aspects of animal coloration. The book a ...
'' (1890) was the first to recognise
frequency-dependent selection 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. * In positive frequency-dependent selection, the fitness of a phenotyp ...
. Poulton is also remembered for his pioneering work on
animal coloration Animal coloration is the general appearance of an animal resulting from the reflection or emission of light from its surfaces. Some animals are brightly coloured, while others are hard to see. In some species, such as the peafowl, the male ...
. He is credited with inventing the term
aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
for warning coloration, as well as for his experiments on 'protective coloration' (
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
). Poulton became Hope Professor of Zoology at the
University of Oxford , mottoeng = The Lord is my light , established = , endowment = £6.1 billion (including colleges) (2019) , budget = £2.145 billion (2019–20) , chancellor ...
in 1893.


Life

Edward Poulton was born in
Reading, Berkshire Reading ( ) is a town and borough in Berkshire, southeast England. Located in the Thames Valley at the confluence of the rivers Thames and Kennet, the Great Western Main Line railway and the M4 motorway serve the town. Reading is east of S ...
on 27 January 1856 the son of the architect William Ford Poulton and his wife, Georgina Sabrina Bagnall. He was educated at Oakley House School in Reading. Between 1873 and 1876, Poulton studied at
Jesus College, Oxford Jesus College (in full: Jesus College in the University of Oxford of Queen Elizabeth's Foundation) is one of the constituent colleges of the University of Oxford in England. It is in the centre of the city, on a site between Turl Street, Ship S ...
under George Rolleston and the anti-Darwinian entomologist
John Obadiah Westwood John Obadiah Westwood (22 December 1805 – 2 January 1893) was an English entomologist and archaeologist also noted for his artistic talents. He published several illustrated works on insects and antiquities. He was among the first entomologist ...
, graduating with a first-class degree in natural science. He maintained an unbroken connection with the college for seventy years as scholar, lecturer and
Fellow A fellow is a concept whose exact meaning depends on context. In learned or professional societies, it refers to a privileged member who is specially elected in recognition of their work and achievements. Within the context of higher educatio ...
(appointed to a fellowship in 1898) until his death. He was a generous benefactor to Jesus College, providing silver for the high table and redecorating the Old Bursary amongst other donations. He was knighted by King
George V George V (George Frederick Ernest Albert; 3 June 1865 – 20 January 1936) was King of the United Kingdom and the British Dominions, and Emperor of India, from 6 May 1910 until his death in 1936. Born during the reign of his grandmother Quee ...
in 1935. Poulton died in
Oxford Oxford () is a city in England. It is the county town and only city of Oxfordshire. In 2020, its population was estimated at 151,584. It is north-west of London, south-east of Birmingham and north-east of Bristol. The city is home to the ...
on 20 November 1943.


Career

Poulton was throughout his career a Darwinist, believing in natural selection as the primary force in evolution. He not only admired
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended f ...
, but also defended the father of
neo-Darwinism Neo-Darwinism is generally used to describe any integration of Charles Darwin's theory of evolution by natural selection with Gregor Mendel's theory of genetics. It mostly refers to evolutionary theory from either 1895 (for the combinations of Dar ...
,
August Weismann August Friedrich Leopold Weismann FRS (For), HonFRSE, LLD (17 January 18345 November 1914) was a German evolutionary biologist. Fellow German Ernst Mayr ranked him as the second most notable evolutionary theorist of the 19th century, after C ...
. Poulton was one of the group of biologists who first translated Weismann's work into English, and he defended Weismann's idea of the continuity of the germ-plasm. In the course of these translations, he noted that recent researches had reduced or perhaps entirely removed the role of acquired characters (
Lamarckism Lamarckism, also known as Lamarckian inheritance or neo-Lamarckism, is the notion that an organism can pass on to its offspring physical characteristics that the parent organism acquired through use or disuse during its lifetime. It is also call ...
) in species formation. His 1890 book, ''
The Colours of Animals ''The Colours of Animals'' is a zoology book written in 1890 by Sir Edward Bagnall Poulton (1856–1943). It was the first substantial textbook to argue the case for Darwinian selection applying to all aspects of animal coloration. The book a ...
'', introduced the concepts of
frequency-dependent selection 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. * In positive frequency-dependent selection, the fitness of a phenotyp ...
and aposematic coloration, as well as supporting Darwin's then unpopular theories 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 heritable traits characteristic of a population over generations. Char ...
and
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (int ...
. Poulton enlarged the Hope entomological collections with his catches in the field which earned him the nickname of "Bag-all" Poulton. Many of the specimens are unmounted and held in biscuit tins (possibly acquired through Huntley & Palmer biscuits owned by his wife's family). In his 1896 book ''Charles Darwin and the Theory of Natural Selection'', Poulton described the ''
Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
'' as "incomparably the greatest work" the biological sciences had seen. Critics of natural selection, Poulton contended, had not taken the time to understand it. This is an evaluation which is much more widely held today than it was then, during the so-called
eclipse of Darwinism Julian Huxley used the phrase "the eclipse of Darwinism" to describe the state of affairs prior to what he called the "modern synthesis". During the "eclipse", evolution was widely accepted in scientific circles but relatively few biologists b ...
. The contemporary ignorance of the mechanism of
inheritance Inheritance is the practice of receiving private property, titles, debts, entitlements, privileges, rights, and obligations upon the death of an individual. The rules of inheritance differ among societies and have changed over time. Officiall ...
stood in the way of a full understanding of the mechanism of evolution. In 1897 Poulton canvassed members during meetings of the
Entomological Society of London The Royal Entomological Society is devoted to the study of insects. Its aims are to disseminate information about insects and improving communication between entomologists. The society was founded in 1833 as the Entomological Society of London ...
. He discovered that many doubted a selectionist origin for
mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. Often, mimicry f ...
. Of those he asked, only three fully supported
Batesian mimicry Batesian mimicry is a form of mimicry where a harmless species has evolved to imitate the warning signals of a harmful species directed at a predator of them both. It is named after the English naturalist Henry Walter Bates, after his work on Bu ...
and
Müllerian mimicry Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Müllerian ...
. The others doubted the inedibility/unpalatability of the models (some investigators even performed taste tests!) or were not convinced that birds were effective selective agents. External and internal forces remained popular alternatives to natural selection.


The arrival of genetics

The rediscovery of Mendel's work filled a critical gap in evolution theory, but at first this was not realised, and many thought it antithetical to selection. There was a long debate between Poulton and
Reginald Punnett Reginald Crundall Punnett FRS (; 20 June 1875 – 3 January 1967) was a British geneticist who co-founded, with William Bateson, the '' Journal of Genetics'' in 1910. Punnett is probably best remembered today as the creator of the Punne ...
, one of Bateson's disciples and the first Professor of Genetics at Oxford. Punnett's ''Mimicry in butterflies'' (1915) rejected selection as the main cause of mimicry. He noted: # The absence of transitional forms and the frequent lack of mimicry in male butterflies were unexplained by selectionist theory. # The enigma of polymorphic mimicry. Some species of butterfly mimicked not merely one, but several models. In breeding experiments these polymorphs cleanly segregated according to Mendel's law of segregation. # Evidence of birds as selective agents was slight and little was known of birds' discriminatory powers, and # The gradual accumulation of minute variations did not (in his view) fit with the facts of heredity. For Punnett, none of these observations were explained by gradual selectionism. Instead he thought mimicry had arisen from sudden mutational jumps ( saltations). Once a mimic was formed by
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...
, natural selection might play a conservative role. However, one by one, each of these objections was shown to be without substance. Evidence from field observations and experiments showed that birds were often the agents of selection in insects. Evidence that small-scale
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...
s were common arrived as soon as breeding experiments were designed to detect them: it was a consequence of experimental methods that early mutations were so noteworthy. Explanations for polymorphism were advanced by E.B. Ford,
Theodosius Dobzhansky Theodosius Grigorievich Dobzhansky (russian: Феодо́сий Григо́рьевич Добржа́нский; uk, Теодо́сій Григо́рович Добржа́нський; January 25, 1900 – December 18, 1975) was a prominent ...
and their colleagues, who developed experimental methods for populations in the wild. The gradual coming-together of field observations and experimental genetics is part of the
modern synthesis Modern synthesis or modern evolutionary synthesis refers to several perspectives on evolutionary biology, namely: * Modern synthesis (20th century), the term coined by Julian Huxley in 1942 to denote the synthesis between Mendelian genetics and s ...
which took place in the middle of the twentieth century. As has become widely accepted, mutations increase the amount of heritable variation in a population, and selection is how biologists describe the differential viability of those variants. Poulton's account is much closer to a modern view of evolution, though Punnett had framed important questions. Poulton's Presidential Address to the
British Association The British Science Association (BSA) is a charity and learned society founded in 1831 to aid in the promotion and development of science. Until 2009 it was known as the British Association for the Advancement of Science (BA). The current Chie ...
in 1937 at the age of 81 reviewed the history of evolutionary thought. He stated that the work of J.B.S. Haldane,
R.A. 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
Julian Huxley Sir Julian Sorell Huxley (22 June 1887 – 14 February 1975) was an English evolutionary biologist, eugenicist, and internationalist. He was a proponent of natural selection, and a leading figure in the mid-twentieth century modern synthesis. ...
was vitally important for showing the relationships between Mendelism and natural selection. The observations and experiments of many biologists had "immensely strengthened and confirmed" the researches on mimicry and warning colours of pioneers like Bates, Wallace, Meldola, Trimen and Müller.


Family

Poulton lived with his family at 56 Banbury Road in
North Oxford North Oxford is a suburban part of the city of Oxford in England. It was owned for many centuries largely by St John's College, Oxford and many of the area's Victorian houses were initially sold on leasehold by the College. Overview The le ...
, a large
Victorian Gothic Gothic Revival (also referred to as Victorian Gothic, neo-Gothic, or Gothick) is an architectural movement that began in the late 1740s in England. The movement gained momentum and expanded in the first half of the 19th century, as increasingly ...
house designed by John Gibbs and built in 1866. In 1881 he married Emily Palmer (d.1939), daughter of George Palmer, M.P. for Reading and head of Huntley and Palmer's biscuit company; they had five children. Three of them were dead by 1919. Their eldest son Dr. Edward Palmer Poulton of Guy's Hospital died in 1939, meaning that Sir Edward was outlived only by his daughter Margaret Lucy (1887–1965), wife of Dr Maxwell Garnett. Poulton's son, Ronald Poulton-Palmer played international rugby for England and was killed in May 1915 in World War I. His first daughter Hilda married Dr Ernest Ainsley-Walker and died in 1917. His youngest daughter, Janet Palmer, married
Charles Symonds Air Vice Marshal Sir Charles Putnam Symonds (11 April 1890 – 7 December 1978) was an English neurologist and a senior medical officer in the Royal Air Force. His initial medical training was at Guy's Hospital, followed by specialised training ...
in 1915 and died in 1919.


Legacy

Poulton is remembered as an early originator of the
biological species concept The species problem is the set of questions that arises when biologists attempt to define what a species is. Such a definition is called a species concept; there are at least 26 recognized species concepts. A species concept that works well for se ...
. According to
Ernst Mayr Ernst Walter Mayr (; 5 July 1904 – 3 February 2005) was one of the 20th century's leading evolutionary biologists. He was also a renowned taxonomist, tropical explorer, ornithologist, philosopher of biology, and historian of science. His ...
, Poulton invented the term
sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species s ...
in relation to species, and he also invented the term
aposematism Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or ...
for warning coloration. Poulton, along with
Julian Huxley Sir Julian Sorell Huxley (22 June 1887 – 14 February 1975) was an English evolutionary biologist, eugenicist, and internationalist. He was a proponent of natural selection, and a leading figure in the mid-twentieth century modern synthesis. ...
, J.B.S. Haldane,
R.A. 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 E.B. Ford, promoted the idea of natural selection through many years when it was denigrated. Poulton was succeeded by G.D. Hale Carpenter as Hope Professor of Entomology at
Oxford University Oxford () is a city in England. It is the county town and only city of Oxfordshire. In 2020, its population was estimated at 151,584. It is north-west of London, south-east of Birmingham and north-east of Bristol. The city is home to the ...
from 1933 to 1948.


Published works

Poulton had over 200 publications spanning over sixty years. * 1890. '' The Colours of Animals: Their Meaning and Use, Especially Considered in the Case of Insects''.
Kegan Paul Charles Kegan Paul (8 March 1828 – 19 July 1902) was an English clergyman, publisher and author. He began his adult life as a clergyman of the Church of England, and served the Church for more than 20 years. His religious orientation moved f ...
, London. * 1896.
Charles Darwin and the Theory of Natural Selection
'. Cassell, London. * 1904. ''What is a Species?'' (Presidential address to the Entomological Society of London, Jan 1904) ''Proc. Ent. Soc. Lond. 1903''. * 1908. ''Essays on Evolution''. London, Cassell. * 1909. ''Charles Darwin and the Origin of species; addresses, etc., in America and England in the year of the two anniversaries'' * 1915. ''Science and the Great War: The
Romanes Lecture The Romanes Lecture is a prestigious free public lecture given annually at the Sheldonian Theatre, Oxford, England. The lecture series was founded by, and named after, the biologist George Romanes, and has been running since 1892. Over the years ...
for 1915''
Clarendon Press Oxford University Press (OUP) is the university press of the University of Oxford. It is the largest university press in the world, and its printing history dates back to the 1480s. Having been officially granted the legal right to print books ...
, Oxford.


Awards and honours

*
Fellow of the Royal Society Fellowship of the Royal Society (FRS, ForMemRS and HonFRS) is an award granted by the judges of the Royal Society of London to individuals who have made a "substantial contribution to the improvement of natural knowledge, including mathematic ...
in June 1889. * President of the
Linnean Society The Linnean Society of London is a learned society dedicated to the study and dissemination of information concerning natural history, evolution, and taxonomy. It possesses several important biological specimen, manuscript and literature coll ...
(1912–1916) *
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 1914 *
Linnean Society The Linnean Society of London is a learned society dedicated to the study and dissemination of information concerning natural history, evolution, and taxonomy. It possesses several important biological specimen, manuscript and literature coll ...
's
Linnean Medal The Linnean Medal of the Linnean Society of London was established in 1888, and is awarded annually to alternately a botanist or a zoologist or (as has been common since 1958) to one of each in the same year. The medal was of gold until 1976, and ...
in 1922. *
Knighted A knight is a person granted an honorary title of knighthood by a head of state (including the Pope) or representative for service to the monarch, the church or the country, especially in a military capacity. Knighthood finds origins in the Gr ...
in 1935.
* President of the
British Association for the Advancement of Science The British Science Association (BSA) is a charity and learned society founded in 1831 to aid in the promotion and development of science. Until 2009 it was known as the British Association for the Advancement of Science (BA). The current Chie ...
for 1937.


See also

*'' Adaptive Coloration in Animals'' (book by
Hugh Cott Hugh Bamford Cott (6 July 1900 – 18 April 1987) was a British zoologist, an authority on both natural and military camouflage, and a scientific illustrator and photographer. Many of his field studies took place in Africa, where he was especia ...
)


References


External links


J. Mallet: Brief biography



Helen J. Power: Biography
(paywall) *
Natural History Museum: Poulton (correspondence with Wallace, etc)
{{DEFAULTSORT:Poulton, Edward Bagnall 1856 births 1943 deaths Evolutionary biologists British zoologists Critics of Lamarckism Alumni of Jesus College, Oxford Fellows of Jesus College, Oxford English entomologists Presidents of the Linnean Society of London Fellows of the Royal Society Presidents of the British Science Association Hope Professors of Zoology Presidents of the Oxford Union Camouflage researchers