Conrad Hal Waddington (8 November 1905 – 26 September 1975) was a British
developmental biologist
Developmental biology is the study of the process by which animals and plants grow and develop. Developmental biology also encompasses the biology of regeneration, asexual reproduction, metamorphosis, and the growth and differentiation of stem c ...
,
paleontologist
Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
,
geneticist
A geneticist is a biologist or physician who studies genetics, the science of genes, heredity, and variation of organisms. A geneticist can be employed as a scientist or a lecturer. Geneticists may perform general research on genetic processes ...
,
embryologist and
philosopher who laid the foundations for
systems biology,
epigenetics
In biology, epigenetics is the study of stable phenotypic changes (known as ''marks'') that do not involve alterations in the DNA sequence. The Greek prefix '' epi-'' ( "over, outside of, around") in ''epigenetics'' implies features that are ...
, and
evolutionary developmental biology.
Although his theory of
genetic assimilation had a
Darwinian
Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations that ...
explanation, leading
evolutionary biologists
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 f ...
including
Theodosius Dobzhansky
Theodosius Grigorievich Dobzhansky (russian: Феодо́сий Григо́рьевич Добржа́нский; uk, Теодо́сій Григо́рович Добржа́нський; January 25, 1900 – December 18, 1975) was a prominent ...
and
Ernst Mayr considered that Waddington was using genetic assimilation to support so-called
Lamarckian
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 calle ...
inheritance, the acquisition of inherited characteristics through the effects of the environment during an organism's lifetime.
Waddington had wide interests that included
poetry
Poetry (derived from the Greek ''poiesis'', "making"), also called verse, is a form of literature that uses aesthetic and often rhythmic qualities of language − such as phonaesthetics, sound symbolism, and metre − to evoke meanings i ...
and
painting
Painting is the practice of applying paint, pigment, color or other medium to a solid surface (called the "matrix" or "support"). The medium is commonly applied to the base with a brush, but other implements, such as knives, sponges, and ai ...
, as well as left-wing political leanings. In his book ''The Scientific Attitude'' (1941), he touched on political topics such as
central planning
A planned economy is a type of economic system where investment, production and the allocation of capital goods takes place according to economy-wide economic plans and production plans. A planned economy may use centralized, decentralized, pa ...
, and praised
Marxism
Marxism is a left-wing to far-left method of socioeconomic analysis that uses a materialist interpretation of historical development, better known as historical materialism, to understand class relations and social conflict and a dialectical ...
as a "profound scientific philosophy".
Life
Conrad Waddington, known as "Wad" to his friends and "Con" to family, was born in
Evesham to Hal and Mary Ellen (Warner) Waddington, on 8 November 1905.
His family moved to India and until nearly three years of age, Waddington lived in India, where his father worked on a tea estate in the
Wayanad district
Wayanad () is a district in the north-east of Indian state Kerala with administrative headquarters at the municipality of Kalpetta. It is the only plateau in Kerala. The Wayanad Plateau forms a continuation of the Mysore Plateau, the southern ...
of
Kerala
Kerala ( ; ) is a state on the Malabar Coast of India. It was formed on 1 November 1956, following the passage of the States Reorganisation Act, by combining Malayalam-speaking regions of the erstwhile regions of Cochin, Malabar, South ...
. In 1910, at the age of four, he was sent to live with family in England including his aunt, uncle, and Quaker grandmother. His parents remained in India until 1928. During his childhood, he was particularly attached to a local druggist and distant relation, Dr. Doeg. Doeg, whom Waddington called "Grandpa", introduced Waddington to a wide range of sciences from chemistry to geology. During the year following the completion of his entrance exams to university, Waddington received an intense course in chemistry from
E. J. Holmyard. Aside from being "something of a genius of a
hemistryteacher," Holmyard introduced Waddington to the "Alexandrian
Gnostics" and the "Arabic
Alchemists
Alchemy (from Arabic: ''al-kīmiyā''; from Ancient Greek: χυμεία, ''khumeía'') is an ancient branch of natural philosophy, a philosophical and protoscientific tradition that was historically practiced in China, India, the Muslim world, ...
." From these lessons in metaphysics, Waddington first gained an appreciation for interconnected
holistic
Holism () is the idea that various systems (e.g. physical, biological, social) should be viewed as wholes, not merely as a collection of parts. The term "holism" was coined by Jan Smuts in his 1926 book '' Holism and Evolution''."holism, n." OED On ...
systems. Waddington reflected that this early education prepared him for
Alfred North Whitehead's philosophy in the 1920s and 30s and the
cybernetics of
Norbert Wiener and others in the 1940s.
He attended
Clifton College
''The spirit nourishes within''
, established = 160 years ago
, closed =
, type = Public schoolIndependent boarding and day school
, religion = Christian
, president =
, head_label = Head of College
, hea ...
and
Sidney Sussex College, Cambridge. He took the Natural Sciences
Tripos
At the University of Cambridge, a Tripos (, plural 'Triposes') is any of the examinations that qualify an undergraduate for a bachelor's degree or the courses taken by a student to prepare for these. For example, an undergraduate studying mathe ...
, earning a First in Part II in geology in 1926. In 1928, he was awarded an Arnold Gerstenberg Studentship in the University of Cambridge, whose purpose was to promote "the study of Moral Philosophy and Metaphysics among students of Natural Science, both men and women." He took up a Lecturership in Zoology and was a Fellow of
Christ's College until 1942. His friends included
Gregory Bateson,
Walter Gropius
Walter Adolph Georg Gropius (18 May 1883 – 5 July 1969) was a German-American architect and founder of the Bauhaus School, who, along with Alvar Aalto, Ludwig Mies van der Rohe, Le Corbusier and Frank Lloyd Wright, is widely regarded as one ...
,
C. P. Snow
Charles Percy Snow, Baron Snow, (15 October 1905 – 1 July 1980) was an English novelist and physical chemist who also served in several important positions in the British Civil Service and briefly in the UK government.''The Columbia Encyclope ...
,
Solly Zuckerman
Solomon "Solly" Zuckerman, Baron Zuckerman (30 May 1904 – 1 April 1993) was a British public servant, zoologist and operational research pioneer. He is best remembered as a scientific advisor to the Allies on bombing strategy in the Second Wo ...
,
Joseph Needham, and
John Desmond Bernal
John Desmond Bernal (; 10 May 1901 – 15 September 1971) was an Irish scientist who pioneered the use of X-ray crystallography in molecular biology. He published extensively on the history of science. In addition, Bernal wrote popular book ...
. His interests began with
palaeontology but moved on to the
heredity and development of living things. He also studied philosophy.
During
World War II
World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the vast majority of the world's countries—including all of the great powers—forming two opposing ...
he was involved in
operational research
Operations research ( en-GB, operational research) (U.S. Air Force Specialty Code: Operations Analysis), often shortened to the initialism OR, is a discipline that deals with the development and application of analytical methods to improve deci ...
with the
Royal Air Force
The Royal Air Force (RAF) is the United Kingdom's air and space force. It was formed towards the end of the First World War on 1 April 1918, becoming the first independent air force in the world, by regrouping the Royal Flying Corps (RFC) an ...
and became scientific advisor to the Commander in Chief of
Coastal Command
RAF Coastal Command was a formation within the Royal Air Force (RAF). It was founded in 1936, when the RAF was restructured into Fighter, Bomber and Coastal Commands and played an important role during the Second World War. Maritime Aviation ...
from 1944 to 1945.
After the war, in 1947, he replaced
Francis Albert Eley Crew
Francis Albert Eley Crew FRS FRSE LLD (2 March 1886 – 26 May 1973) was an English animal geneticist. He was a pioneer in his field leading to the University of Edinburgh’s place as a world leader in the science of animal genetics. He was t ...
as Professor of Animal Genetics at the
University of Edinburgh
The University of Edinburgh ( sco, University o Edinburgh, gd, Oilthigh Dhùn Èideann; abbreviated as ''Edin.'' in post-nominals) is a public research university based in Edinburgh, Scotland. Granted a royal charter by King James VI in 15 ...
. He would stay at Edinburgh for the rest of life with the exception of one year (1960–1961) when he was a Fellow on the faculty in the Center for Advanced Studies at
Wesleyan University
Wesleyan University ( ) is a private liberal arts university in Middletown, Connecticut. Founded in 1831 as a men's college under the auspices of the Methodist Episcopal Church and with the support of prominent residents of Middletown, the col ...
in Middletown, Connecticut. His personal papers are largely kept at the University of Edinburgh library.
He died in Edinburgh on 26 September 1975.
Family
Waddington was married twice. His first marriage produced a son, C. Jake Waddington, professor of physics at the
University of Minnesota
The University of Minnesota, formally the University of Minnesota, Twin Cities, (UMN Twin Cities, the U of M, or Minnesota) is a public land-grant research university in the Twin Cities of Minneapolis and Saint Paul, Minnesota, United States. ...
, but ended in 1936. He then married
Margaret Justin Blanco White
Margaret Justin Blanco White OBE ARIBA (11 December 1911 – 1 November 2001) was a Scottish architect.
Early life and education
Margaret Justin Blanco White was born at 30 Pembroke Square, Kensington, London, on 11 December 1911. Her father ...
, daughter of the writer
Amber Reeves
Amber Blanco White (' Reeves; 1 July 1887 – 26 December 1981) was a New Zealand-born British feminist writer and scholar.
Early life
Reeves was born in Christchurch, New Zealand,
the eldest of three children
of Fabian feminist Maud Pember Re ...
, with whom he had two daughters, the anthropologist
Caroline Humphrey
Caroline Humphrey, Baroness Rees of Ludlow, (''née'' Waddington; born 1 September 1943) is a British anthropologist and academic.
Biography
Humphrey's father was the biologist Conrad H. Waddington. Her mother was her father's second wife, ar ...
(1943–) and mathematician
Dusa McDuff
Dusa McDuff Fellow of the Royal Society, FRS Royal Society of Edinburgh, CorrFRSE (born 18 October 1945) is an English mathematician who works on symplectic geometry. She was the first recipient of the Ruth Lyttle Satter Prize in Mathematics, w ...
(1945–).
Evolution
In the early 1930s, Waddington and many other embryologists looked for the molecules that would induce the amphibian neural tube. The search was beyond the technology of that time, and most embryologists moved away from such deep problems. Waddington, however, came to the view that the answers to embryology lay in genetics, and in 1935 went to
Thomas Hunt Morgan's ''Drosophila'' laboratory in California, even though this was a time when most embryologists felt that genes were unimportant and just played a role in minor phenomena such as eye colour.
In the late 1930s, Waddington produced formal models about how gene regulatory products could generate developmental phenomena, showed how the mechanisms underpinning ''Drosophila'' development could be studied through a systematic analysis of mutations that affected the development of the ''
Drosophila
''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many speci ...
'' wing. In a period of great creativity at the end of the 1930s, he also discovered mutations that affected cell phenotypes and wrote his first textbook of "developmental epigenetics", a term that then meant the external manifestation of genetic activity.
Waddington introduced the concept of
canalisation, the ability of an organism to produce the same
phenotype
In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological pr ...
despite variation in genotype or environment. He also identified a mechanism called
genetic assimilation which would allow an animal's response to an environmental stress to become a fixed part of its developmental repertoire, and then went on to show that the mechanism would work.
In 1972, Waddington founded the Centre for Human Ecology in the
University of Edinburgh
The University of Edinburgh ( sco, University o Edinburgh, gd, Oilthigh Dhùn Èideann; abbreviated as ''Edin.'' in post-nominals) is a public research university based in Edinburgh, Scotland. Granted a royal charter by King James VI in 15 ...
.
Epigenetic landscape
Waddington's
epigenetic landscape is a metaphor for how
gene regulation
Regulation of gene expression, or gene regulation, includes a wide range of mechanisms that are used by cells to increase or decrease the production of specific gene products (protein or RNA). Sophisticated programs of gene expression are wi ...
modulates development. Among other metaphors, Waddington asks us to imagine a number of marbles rolling down a hill. The marbles will sample the grooves on the slope, and come to rest at the lowest points. These points represent the eventual
cell
Cell most often refers to:
* Cell (biology), the functional basic unit of life
Cell may also refer to:
Locations
* Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
fates, that is,
tissue types. Waddington coined the term
chreode to represent this cellular developmental process. The idea was based on experiment: Waddington found that one effect of mutation (which could modulate the epigenetic landscape) was to affect how cells differentiated. He also showed how mutation could affect the landscape, and used this metaphor in his discussions on evolution—he emphasised (like
Ernst Haeckel before him) that evolution mainly occurred through mutations that affected developmental anatomy.
Genetic assimilation
![Lamarck Compared to Darwin, Baldwin, Waddington](https://upload.wikimedia.org/wikipedia/commons/5/5d/Lamarck_Compared_to_Darwin%2C_Baldwin%2C_Waddington.svg)
Waddington proposed an evolutionary process, "
genetic assimilation", as a
Darwinian
Darwinism is a theory of biological evolution developed by the English naturalist Charles Darwin (1809–1882) and others, stating that all species of organisms arise and develop through the natural selection of small, inherited variations that ...
mechanism that allows certain acquired characteristic to become heritable. According to Navis, (2007) "Waddington focused his genetic assimilation work on the crossveinless trait of ''
Drosophila
''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many speci ...
''. This trait occurs with high frequency in heat-treated flies. After a few generations, the trait can be found in the population, without the application of heat, based on hidden genetic variation that Waddington asserted had been "assimilated".
Neo-Darwinism versus Lamarckism
Waddington's theory of genetic assimilation was controversial. The
evolutionary biologists
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 f ...
Theodosius Dobzhansky
Theodosius Grigorievich Dobzhansky (russian: Феодо́сий Григо́рьевич Добржа́нский; uk, Теодо́сій Григо́рович Добржа́нський; January 25, 1900 – December 18, 1975) was a prominent ...
and
Ernst Mayr both thought that Waddington was using genetic assimilation to support
Lamarckian
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 calle ...
inheritance. They denied that genetic assimilation had taken place, and asserted that Waddington had simply observed the
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. Cha ...
of genetic variants that already existed in the study population. Other biologists such as
Wallace Arthur disagree, writing that "genetic assimilation, looks, but is not Lamarckian. It is a special case of the evolution of
phenotypic plasticity
Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompa ...
".
Adam S. Wilkins wrote that "
addingtonin his lifetime... was widely perceived primarily as a critic of Neo-Darwinian evolutionary theory. His criticisms ... were focused on what he saw as unrealistic, 'atomistic' models of both gene selection and trait evolution." In particular, according to Wilkins, Waddington felt that the Neo-Darwinians badly neglected the phenomenon of extensive gene interactions and that the "randomness" of mutational effects, posited in the theory, was false.
Even though Waddington became critical of the neo-darwinian synthetic theory of evolution, he still described himself as a Darwinian, and called for an
extended evolutionary synthesis
The extended evolutionary synthesis consists of a set of theoretical concepts argued to be more comprehensive than the earlier modern synthesis of evolutionary biology that took place between 1918 and 1942. The extended evolutionary synthesis wa ...
based on his research.
Reviewing the debate in 2015, the
systems biologist Denis Noble
Denis Noble (born 16 November 1936) is a British biologist who held the Burdon Sanderson Chair of Cardiovascular Physiology at the University of Oxford from 1984 to 2004 and was appointed Professor Emeritus and co-Director of Computational Ph ...
writes however that
As an organiser
Waddington was very active in advancing biology as a discipline. He contributed to a book on the role of the sciences in times of war, and helped set up several professional bodies representing biology as a discipline.
A remarkable number of his contemporary colleagues in Edinburgh became Fellows of the Royal Society during his time there, or shortly thereafter. Waddington was an old-fashioned intellectual who lived in both the arts and science milieus of the 1950s and wrote widely. His 1969 book ''Behind Appearance; a Study of the Relations Between Painting and the Natural Sciences in This Century'' (MIT press) not only has wonderful pictures but is still worth reading.
Waddington was, without doubt, the most original and important thinker about
developmental biology of the
pre-molecular age and the medal of the British Society for Developmental Biology is named after him.
Waddington co-founded The Institute for Advanced Studies in the Humanities at the University of Edinburgh in 1969 with Professor John MacQueen, Professor of Scottish Literature and Oral Tradition.
Selected works
Books
* Waddington, C. H. (1939). ''An Introduction to Modern Genetics''. London : George Alien & Unwin.
* ––– (1940). ''Organisers & Genes''. Cambridge: Cambridge University Press.
* ––– and others (1942). ''Science and Ethics'', George Allen & Unwin.
* ––– (1946). ''How Animals Develop''. London : George Allen & Unwin.
* ––– (1948). ''The Scientific Attitude'', Pelican Books
* ––– (1953). ''The Epigenetics of birds''. Cambridge : Cambridge University Press.
* ––– (1956). ''Principles of Embryology''. London : George Allen & Unwin.
* ––– (1957). ''The Strategy of the Genes''. London : George Allen & Unwin.
* ––– (1959). ''Biological Organisation Cellular and Subcellular : Proceedings of a Symposium''. London: Pergamon Press.
* ––– (1960). ''The Ethical Animal''. London : George Allen & Unwin.
* ––– (1961). ''The Human Evolutionary System''. In: Michael Banton (Ed.), ''Darwinism and the Study of Society''. London: Tavistock.
* ––– (1961). ''The Nature of Life''. London : George, Allen, & Unwin.
* ––– (1962). ''New Patterns in Genetics and Development''. New York: Columbia University Press.
* ––– (1966). ''Principles of Development and Differentiation''. New York: Macmillan Company.
* ––– (1970). 72). ''Behind Appearance : A Study in the Relationship Between Painting and the Natural Sciences in this Century''. The MIT Press.
* –––, ed. (1968–72). ''Towards a Theoretical Biology''. 4 vols. Edinburgh: Edinburgh University Press.
* –––,
Kenny, A.,
Longuet-Higgins, H.C.,
Lucas, J.R. (1972). ''The Nature of Mind'', Edinburgh: Edinburgh University Press (1971-3
Gifford Lectures
The Gifford Lectures () are an annual series of lectures which were established in 1887 by the will of Adam Gifford, Lord Gifford. Their purpose is to "promote and diffuse the study of natural theology in the widest sense of the term – in o ...
in Edinburgh
online
* –––, Kenny, A., Longuet-Higgins, H. C., Lucas, J. R. (1973). ''The Development of Mind'', Edinburgh: Edinburgh University Press (1971-3 Gifford Lectures in Edinburgh
online
* ––– (1973) ''O.R. in World War 2: Operational Research Against the U-Boat''. London: Elek Science.
* –––, & Jantsch, E. (Eds.). (1976). (published posthumously). ''Evolution and Consciousness: Human Systems in Transition''. Addison-Wesley.
* ––– (1977) (published posthumously). ''Tools for Thought''. London: Jonathan Cape.
Papers
* Waddington, C. H. (1942). Canalization of development and the inheritance of acquired characters. ''Nature'' 150 (3811):563–565.
* --- (1946). Human Ideals and Human Progress. ''World Review'' August:29-36.
* ––– & Carter T. C. (1952). Malformations in mouse embryos induced by trypan blue. ''Nature'' 169 (4288):27-28.
* ––– (1952). Selection of the Genetic Basis for an Acquired Character. ''Nature'' 169 (4294):278.
* ––– (1953). Genetic assimilation of an acquired character. ''Evolution'' 7:118–126.
* ––– (1953). Epigenetics and evolution. ''Symposia of the Society of Experimental Biology'' 7:186–199.
* ––– (1956). Genetic assimilation of the bithorax phenotype. ''Evolution'' 10:1–13.
* ––– (1961). Genetic assimilation. ''Advances in Genetics'' 10:257–290.
* ––– (1974). A Catastrophe Theory of Evolution. ''Annals of the New York Academy of Sciences'' 231:32–42.
* ––– (1977).(published posthumously). Whitehead and Modern Science. ''Mind in Nature: The Interface of Science and Philosophy''. Ed. John B. Cobb and David R. Griffin. University Press of America.
Notes
References
External links
NAHSTE Project Record of C.H. Waddington*
ttps://web.archive.org/web/20060514181106/http://www.epigeneticsnews.com/ Epigenetics News
{{DEFAULTSORT:Waddington, Conrad Hal
1905 births
People from Evesham
1975 deaths
Academics of the University of Edinburgh
British palaeontologists
20th-century British zoologists
Extended evolutionary synthesis
Fellows of Christ's College, Cambridge
Wesleyan University faculty
Fellows of the Royal Society of Edinburgh
Fellows of the Royal Society
People educated at Clifton College
British operations researchers
Systems biologists
Theoretical biologists
Alumni of Sidney Sussex College, Cambridge
Commanders of the Order of the British Empire
Presidents of the International Union of Biological Sciences
Reeves family