Geoffrey Douglas Hale Carpenter
Encyclopedia
G.D. Hale Carpenter MBE
MA
DM
FRS (Eton, Berkshire
, 26 October 1882 – Oxford
, 30 January 1953) was a British entomologist and physician
. He worked first at the London School of Hygiene and Tropical Medicine, and in Uganda
, on tse-tse flies
and sleeping sickness. His main work in zoology
was on mimicry in butterflies, an interest he developed in Uganda and Tanganyika
.
He succeeded E.B. Poulton as Hope Professor of Zoology
at Oxford University from 1933–1948.
and physiologist William Benjamin Carpenter
, and a great-grandson of Lant Carpenter
, a Unitarian
minister.
Carpenter attended St. Catherine's College, Oxford, graduating in 1904. He then studied medicine at St George's Hospital
, London, graduating MB
ChB
(the standard medical degree at the University of London) in 1908. He then joined the London School of Hygiene and Tropical Medicine, and took the DM
in 1913 with a dissertation on the tsetse fly (Glossina palpalis) and sleeping sickness.
In 1919 he married Amy Frances Thomas-Peter from Cornwall
. The marriage had no issue.
, where he worked in Uganda
on the north shore of Lake Victoria
. Upon the outbreak of World War I, Carpenter was called to service in the British Army
Medical Corps. He was stationed with the troops at the border between Uganda and German East Africa
. In December 1914 he was appointed Medical Officer at the fort in Kakindu, southern Uganda. As it turned out, he had plenty of spare time, and spent it studying the local butterflies. "The hosts of butterflies at Kakindu passed beyond anything I had ever see; some days are quite unforgettable".
From May 1916 to January 1918, he was located in Tanganyika
(former German E. Africa), 200 miles south to south-east of Lake Victoria. Here he conducted experiments on palatability
with young insectivorous monkeys. He tested the edibility of cryptic
and warning colouration in insects. This was propitious, because later, back at Oxford, both he and E.B. Poulton worked on the role of predators in shaping mimicry. The standard theory was that cryptic
forms were palatable, and aposematic forms were distasteful. Then palatable mimics of distasteful forms could gain protection from predation. The question at stake was whether the observations, which dated from work by naturalists in the 19th century, could or could not be accounted for by natural selection.
Punnett's Mimicry in butterflies (1915) rejected selection as the main cause of mimicry. He noted:
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, natural selection might play a conservative role.
However, one by one, each of these objections were 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 mutations 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 and Dobzhansky
and colleagues, who developed experimental methods for populations in the wild. The question of polymorphism is discussed further in polymorphism (biology)
.
The gradual coming-together of field observations and experimental genetics is part of the evolutionary synthesis
which took place in the middle of the twentieth century. The small book by Carpenter and E.B. Ford, was the first book on ecological genetics
, a field which produced a series of classic studies uniting fieldwork with laboratory genetics. The book is a minor masterpiece of evolutionary biology.
MBE
MBE can stand for:* Mail Boxes Etc.* Management by exception* Master of Bioethics* Master of Bioscience Enterprise* Master of Business Engineering* Master of Business Economics* Mean Biased Error...
MA
Master of Arts (postgraduate)
A Master of Arts from the Latin Magister Artium, is a type of Master's degree awarded by universities in many countries. The M.A. is usually contrasted with the M.S. or M.Sc. degrees...
DM
Doctor of Medicine
Doctor of Medicine is a doctoral degree for physicians. The degree is granted by medical schools...
FRS (Eton, Berkshire
Eton, Berkshire
Eton is a town and civil parish in Berkshire, England, lying on the opposite bank of the River Thames to Windsor and connected to it by Windsor Bridge. The parish also includes the large village of Eton Wick, 2 miles west of the town, and has a population of 4,980. Eton was in Buckinghamshire until...
, 26 October 1882 – Oxford
Oxford
The city of Oxford is the county town of Oxfordshire, England. The city, made prominent by its medieval university, has a population of just under 165,000, with 153,900 living within the district boundary. It lies about 50 miles north-west of London. The rivers Cherwell and Thames run through...
, 30 January 1953) was a British entomologist and physician
Physician
A physician is a health care provider who practices the profession of medicine, which is concerned with promoting, maintaining or restoring human health through the study, diagnosis, and treatment of disease, injury and other physical and mental impairments...
. He worked first at the London School of Hygiene and Tropical Medicine, and in Uganda
Uganda
Uganda , officially the Republic of Uganda, is a landlocked country in East Africa. Uganda is also known as the "Pearl of Africa". It is bordered on the east by Kenya, on the north by South Sudan, on the west by the Democratic Republic of the Congo, on the southwest by Rwanda, and on the south by...
, on tse-tse flies
Tsetse fly
Tsetse , sometimes spelled tzetze and also known as tik-tik flies, are large biting flies that inhabit much of mid-continental Africa between the Sahara and the Kalahari deserts. They live by feeding on the blood of vertebrate animals and are the primary biological vectors of trypanosomes, which...
and sleeping sickness. His main work in zoology
Zoology
Zoology |zoölogy]]), is the branch of biology that relates to the animal kingdom, including the structure, embryology, evolution, classification, habits, and distribution of all animals, both living and extinct...
was on mimicry in butterflies, an interest he developed in Uganda and Tanganyika
Tanganyika
Tanganyika , later formally the Republic of Tanganyika, was a sovereign state in East Africa from 1961 to 1964. It was situated between the Indian Ocean and the African Great Lakes of Lake Victoria, Lake Malawi and Lake Tanganyika...
.
He succeeded E.B. Poulton as Hope Professor of Zoology
Hope Professor of Zoology
The Hope Professor of Zoology is a professorship at Oxford University. The first Hope Professor was John Obadiah Westwood. The current holder is Charles Godfray.* John Obadiah Westwood The Hope Professor of Zoology is a professorship at Oxford University. The first Hope Professor was John...
at Oxford University from 1933–1948.
Biography
Carpenter was a son of Philip Herbert Carpenter DSc FRS, a schoolmaster at Eton College, a grandson of the naturalistNaturalist
Naturalist may refer to:* Practitioner of natural history* Conservationist* Advocate of naturalism * Naturalist , autobiography-See also:* The American Naturalist, periodical* Naturalism...
and physiologist William Benjamin Carpenter
William Benjamin Carpenter
William Benjamin Carpenter MD CB FRS was an English physician, invertebrate zoologist and physiologist. He was instrumental in the early stages of the unified University of London.-Life:...
, and a great-grandson of Lant Carpenter
Lant Carpenter
Lant Carpenter, Dr. was an English educator and Unitarian minister.Lant Carpenter was born in Kidderminster, the third son of George Carpenter and his wife Mary ....
, a 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....
minister.
Carpenter attended St. Catherine's College, Oxford, graduating in 1904. He then studied medicine at St George's Hospital
St George's Hospital
Founded in 1733, St George’s Hospital is one of the UK's largest teaching hospitals. It shares its main hospital site in Tooting, England with the St George's, University of London which trains NHS staff and carries out advanced medical research....
, London, graduating MB
MB
- Computers :* Megabyte , a measure of amount of information used, for example, to quantify computer memory or storage capacity* Megabit , a measure of amount of information* MacBook* Motherboard* Message board- File format ".MB" :...
ChB
CHB
CHB may refer to:*Centre for the history of the book University of Edinburgh*Centre half-back in Australian rules football*CHB Bank of South Korea*Champion Homes, NASDAQ symbol CHB*Chang Hwa Bank of Taiwan*Children's Hospital Boston*Chronic Hepatitis B...
(the standard medical degree at the University of London) in 1908. He then joined the London School of Hygiene and Tropical Medicine, and took the DM
Doctor of Medicine
Doctor of Medicine is a doctoral degree for physicians. The degree is granted by medical schools...
in 1913 with a dissertation on the tsetse fly (Glossina palpalis) and sleeping sickness.
In 1919 he married Amy Frances Thomas-Peter from Cornwall
Cornwall
Cornwall is a unitary authority and ceremonial county of England, within the United Kingdom. It is bordered to the north and west by the Celtic Sea, to the south by the English Channel, and to the east by the county of Devon, over the River Tamar. Cornwall has a population of , and covers an area of...
. The marriage had no issue.
Career
In 1910 he joined the Colonial Medical ServiceColonial Service
The Colonial Service was the British government service which administered Britain's colonies and protectorates, under the authority of the Secretary of State for the Colonies and the Colonial Office in London....
, where he worked in Uganda
Uganda
Uganda , officially the Republic of Uganda, is a landlocked country in East Africa. Uganda is also known as the "Pearl of Africa". It is bordered on the east by Kenya, on the north by South Sudan, on the west by the Democratic Republic of the Congo, on the southwest by Rwanda, and on the south by...
on the north shore of Lake Victoria
Lake Victoria
Lake Victoria is one of the African Great Lakes. The lake was named for Queen Victoria of the United Kingdom, by John Hanning Speke, the first European to discover this lake....
. Upon the outbreak of World War I, Carpenter was called to service in 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...
Medical Corps. He was stationed with the troops at the border between Uganda and German East Africa
German East Africa
German East Africa was a German colony in East Africa, which included what are now :Burundi, :Rwanda and Tanganyika . Its area was , nearly three times the size of Germany today....
. In December 1914 he was appointed Medical Officer at the fort in Kakindu, southern Uganda. As it turned out, he had plenty of spare time, and spent it studying the local butterflies. "The hosts of butterflies at Kakindu passed beyond anything I had ever see; some days are quite unforgettable".
From May 1916 to January 1918, he was located in Tanganyika
Tanganyika
Tanganyika , later formally the Republic of Tanganyika, was a sovereign state in East Africa from 1961 to 1964. It was situated between the Indian Ocean and the African Great Lakes of Lake Victoria, Lake Malawi and Lake Tanganyika...
(former German E. Africa), 200 miles south to south-east of Lake Victoria. Here he conducted experiments on palatability
Palatability
Palatability is the hedonic reward provided by foods or fluids that are agreeable to the "palate" in regard to the homeostatic satisfaction of nutritional, water, or energy needs. The palatability of a food or fluid, unlike its flavor or taste, varies with the state of an individual: it is lower...
with young insectivorous monkeys. He tested the edibility of cryptic
Crypsis
In ecology, crypsis is the ability of an organism to avoid observation or detection by other organisms. It may be either a predation strategy or an antipredator adaptation, and methods include camouflage, nocturnality, subterranean lifestyle, transparency, and mimicry...
and warning colouration in insects. This was propitious, because later, back at Oxford, both he and E.B. Poulton worked on the role of predators in shaping mimicry. The standard theory was that cryptic
Crypsis
In ecology, crypsis is the ability of an organism to avoid observation or detection by other organisms. It may be either a predation strategy or an antipredator adaptation, and methods include camouflage, nocturnality, subterranean lifestyle, transparency, and mimicry...
forms were palatable, and aposematic forms were distasteful. Then palatable mimics of distasteful forms could gain protection from predation. The question at stake was whether the observations, which dated from work by naturalists in the 19th century, could or could not be accounted for by natural selection.
Mimicry
In England, the geneticist R.C. Punnett, and in America the ornithologist W.L. McAtee, doubted both that birds could distinguish distasteful forms, and that their predation was heavy enough to bring about the colour forms found in butterflies.Punnett's Mimicry in butterflies (1915) rejected selection as the main cause of mimicry. He noted:
- 1. The absence of transitional forms and the frequent lack of mimicry in male butterflies were unexplained by selectionist theory.
- 2. 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.
- 3. Evidence of birds as selective agents was slight and little was known of birds' discriminatory powers, and
- 4. The gradual accumulation of minute variations did not 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, natural selection might play a conservative role.
However, one by one, each of these objections were 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 mutations 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 and 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...
and colleagues, who developed experimental methods for populations in the wild. The question of polymorphism is discussed further in polymorphism (biology)
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...
.
The gradual coming-together of field observations and experimental genetics is part of the 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...
which took place in the middle of the twentieth century. The small book by Carpenter and E.B. Ford, was the first book on 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....
, a field which produced a series of classic studies uniting fieldwork with laboratory genetics. The book is a minor masterpiece of evolutionary biology.
See also
- Hobby, B.M. 1953. Geoffrey Douglas Hale Carpenter. British Medical Journal 1, p406
- Remington, Charles L. 1954. Geoffrey Douglas Hale Carpenter. Lepidopterist's News, 8, p31-43. http://research.yale.edu/peabody/jls/pdfs/1950s/1954/1954-8(1-2)31-Remington.pdf
Some publications
- Carpenter G.D.H. 1920. A Naturalist on Lake Victoria, with an account of Sleeping Sickness and the Tse-tse Fly. Unwin, London.
- Carpenter G.D.H. 1921. Experiments on the relative edibility of insects, with special reference to their coloration. Transactions of the Royal Entomological Society, London 54:1-105.
- Carpenter G.D.H. and E.B. FordE.B. FordEdmund 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...
. 1933. Mimicry. Methuen, London. - Carpenter G.D.H. 1933. Gregarious roosting habits of aposematic butterflies. Proceedings of the Entomological Society of London 8:110-111.
- Carpenter G.D.H. 1935. The Rhopalocera of Abyssinia a faunistic study. Trans. R. Entomol. Soc. Lond. 83 : 313-447
- Carpenter G.D.H. 1936. Charles Darwin and entomology. Transactions of the South-eastern Union of Scientific Societies. Papers Contributed to Congress 1936:1-23.
- Carpenter G.D.H. 1938. Audible emission of defensive froth by insects with an appendix on the anatomical structures concerned in a moth by H. Eltringham. Proceedings of the Zoological Society of London 108:243-252.
- Carpenter G.D.H. 1939. Birds as enemies of butterflies, with special reference to mimicry. Proceedings, VII Internationaler Kongress für Entomologie, Berlin 1938:1061-1074.
- Carpenter G.D.H. 1941. The relative frequency of beakmarks on butterflies of different edibility to birds. Proceedings of the Zoological Society of London (Series A) 3:223-231.
- Carpenter G.D.H. 1947. The writings of I. Portschinsky on warning colours and eyespots. Proceedings of the Entomological Society of London. Series A. General Entomology 22:103-113.
- Carpenter G.D.H. 1949. Pseudacraea eurytus (L.) (Lep. Nymphalidae): a study of a polymorphic mimic in various degrees of speciation. Transactions of the Royal Entomological Society, London 100:71-133.
- Carpenter G.D.H. 1953. The genus Euploea (Lep. Danaidae) in Microneia, Melanesia, Polynesia and Australia. A zoo-geographical study. Transactions of the Zoological Society of London 28:1-184, plates 1-9.