Conrad Hal Waddington
Conrad Hal Waddington CBE
CBE and C.B.E. are abbreviations for "Commander of the Order of the British Empire", a grade in the Order of the British Empire.Other uses include:* Chemical and Biochemical Engineering...

Royal Society of Edinburgh
The Royal Society of Edinburgh is Scotland's national academy of science and letters. It is a registered charity, operating on a wholly independent and non-party-political basis and providing public benefit throughout Scotland...

 (1905–1975) was a developmental biologist
A biologist is a scientist devoted to and producing results in biology through the study of life. Typically biologists study organisms and their relationship to their environment. Biologists involved in basic research attempt to discover underlying mechanisms that govern how organisms work...

, paleontologist
Paleontology "old, ancient", ὄν, ὀντ- "being, creature", and λόγος "speech, thought") is the study of prehistoric life. It includes the study of fossils to determine organisms' evolution and interactions with each other and their environments...

, geneticist
A geneticist is a biologist who studies genetics, the science of genes, heredity, and variation of organisms. A geneticist can be employed as a researcher or lecturer. Some geneticists perform experiments and analyze data to interpret the inheritance of skills. A geneticist is also a Consultant or...

, embryologist and philosopher who laid the foundations for systems biology
Systems biology
Systems biology is a term used to describe a number of trends in bioscience research, and a movement which draws on those trends. Proponents describe systems biology as a biology-based inter-disciplinary study field that focuses on complex interactions in biological systems, claiming that it uses...

. He had wide interests that included poetry
Poetry is a form of literary art in which language is used for its aesthetic and evocative qualities in addition to, or in lieu of, its apparent meaning...

 and painting
Painting is the practice of applying paint, pigment, color or other medium to a surface . The application of the medium is commonly applied to the base with a brush but other objects can be used. In art, the term painting describes both the act and the result of the action. However, painting is...

, as well as left-wing political leanings.


Waddington, known as "Wad" to his friends and "Con" to family, was born to Hal and Mary Ellen (Warner) Waddington, 8 November 1905. Until nearly three years of age, Waddington lived with his parents in India, where his father worked on a tea estate in the Wayanad district
Wayanad District
Wayanad District in the north-east of Kerala, India, was formed on November 1, 1980 as the 12th district by carving out areas from Kozhikode and Kannur districts. Kalpetta is the district headquarters as well as the only municipal town in the district. The region was known as Mayakshetra in the...

. 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, who 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 [chemistry] teacher," Holmyard introduced Waddington to the "Alexandrian Gnostics" and the "Arabic Alchemists." From these lessons in metaphysics, Waddington first gained an appreciation for interconnected holistic systems. Waddington reflected that this early education prepared him for Alfred North Whitehead
Alfred North Whitehead
Alfred North Whitehead, OM FRS was an English mathematician who became a philosopher. He wrote on algebra, logic, foundations of mathematics, philosophy of science, physics, metaphysics, and education...

's philosophy in the 1920s and 30s and the cybernetics
Cybernetics is the interdisciplinary study of the structure of regulatory systems. Cybernetics is closely related to information theory, control theory and systems theory, at least in its first-order form...

 of Norbert Wiener
Norbert Wiener
Norbert Wiener was an American mathematician.A famous child prodigy, Wiener later became an early researcher in stochastic and noise processes, contributing work relevant to electronic engineering, electronic communication, and control systems.Wiener is regarded as the originator of cybernetics, a...

 and others in the 1940s.

He attended Clifton College
Clifton College
Clifton College is a co-educational independent school in Clifton, Bristol, England, founded in 1862. In its early years it was notable for emphasising science in the curriculum, and for being less concerned with social elitism, e.g. by admitting day-boys on equal terms and providing a dedicated...

 and Sidney Sussex College, Cambridge
Sidney Sussex College, Cambridge
Sidney Sussex College is a constituent college of the University of Cambridge in England.The college was founded in 1596 and named after its foundress, Frances Sidney, Countess of Sussex. It was from its inception an avowedly Puritan foundation: some good and godlie moniment for the mainteynance...

. He took the Natural Sciences Tripos
The University of Cambridge, England, divides the different kinds of honours bachelor's degree by Tripos , plural Triposes. The word has an obscure etymology, but may be traced to the three-legged stool candidates once used to sit on when taking oral examinations...

, earning a First in Part II in geology in 1926. He took up a Lecturership in Zoology and was a Fellow of Christ's College
Christ's College, Cambridge
Christ's College is a constituent college of the University of Cambridge.With a reputation for high academic standards, Christ's College averaged top place in the Tompkins Table from 1980-2000 . In 2011, Christ's was placed sixth.-College history:...

 until 1942. His friends included Gregory Bateson
Gregory Bateson
Gregory Bateson was an English anthropologist, social scientist, linguist, visual anthropologist, semiotician and cyberneticist whose work intersected that of many other fields. He had a natural ability to recognize order and pattern in the universe...

, Walter Gropius
Walter Gropius
Walter Adolph Georg Gropius was a German architect and founder of the Bauhaus School who, along with Ludwig Mies van der Rohe and Le Corbusier, is widely regarded as one of the pioneering masters of modern architecture....

, C. P. Snow
C. P. Snow
Charles Percy Snow, Baron Snow of the City of Leicester CBE was an English physicist and novelist who also served in several important positions with the UK government...

, Solly Zuckerman
Solly Zuckerman
Solly Zuckerman, Baron Zuckerman, OM, KCB, FRS was a British public servant, zoologist, and scientific advisor who is best remembered as an advisor to the Allies on bombing strategy in World War II, for his work to advance the cause of nuclear non-proliferation, and for his role in bringing...

, Joseph Needham
Joseph Needham
Noel Joseph Terence Montgomery Needham, CH, FRS, FBA , also known as Li Yuese , was a British scientist, historian and sinologist known for his scientific research and writing on the history of Chinese science. He was elected a fellow of the Royal Society in 1941, and as a fellow of the British...

, and J. D. Bernal. His interests began with palaeontology but moved on to the heredity
Heredity is the passing of traits to offspring . This is the process by which an offspring cell or organism acquires or becomes predisposed to the characteristics of its parent cell or organism. Through heredity, variations exhibited by individuals can accumulate and cause some species to evolve...

 and development of living things. He also studied philosophy.

During World War II
World War II
World War II, or the Second World War , was a global conflict lasting from 1939 to 1945, involving most of the world's nations—including all of the great powers—eventually forming two opposing military alliances: the Allies and the Axis...

 he was involved in operational research with the Royal Air Force
Royal Air Force
The Royal Air Force is the aerial warfare service branch of the British Armed Forces. Formed on 1 April 1918, it is the oldest independent air force in the world...

 and became scientific advisor to the Commander in Chief of Coastal Command
RAF Coastal Command
RAF Coastal Command was a formation within the Royal Air Force . Founded in 1936, it was the RAF's premier maritime arm, after the Royal Navy's secondment of the Fleet Air Arm in 1937. Naval aviation was neglected in the inter-war period, 1919–1939, and as a consequence the service did not receive...

 from 1944 to 1945. After the war he became Professor of Animal Genetics at the University of Edinburgh
University of Edinburgh
The University of Edinburgh, founded in 1583, is a public research university located in Edinburgh, the capital of Scotland, and a UNESCO World Heritage Site. The university is deeply embedded in the fabric of the city, with many of the buildings in the historic Old Town belonging to the university...

. 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
Wesleyan University is a private liberal arts college founded in 1831 and located in Middletown, Connecticut. According to the Carnegie Foundation for the Advancement of Teaching, Wesleyan is the only Baccalaureate College in the nation that emphasizes undergraduate instruction in the arts and...

 in Middletown, Connecticut. His personal papers are largely kept at the University of Edinburgh library.

Waddington was married twice. His first marriage produced a son, C. Jake Waddington, professor of physics at the University of Minnesota
University of Minnesota
The University of Minnesota, Twin Cities is a public research university located in Minneapolis and St. Paul, Minnesota, United States. It is the oldest and largest part of the University of Minnesota system and has the fourth-largest main campus student body in the United States, with 52,557...

, but ended in 1936. He then married Justin Blanco White, daughter of the writer Amber Reeves
Amber Reeves
Amber Blanco White [née Amber Reeves] was a British feminist writer and scholar.-Early life:Reeves was born in Christchurch, New Zealand,the eldest of three children...

, with whom he had two daughters, mathematician Dusa McDuff
Dusa McDuff
Dusa McDuff is an English mathematician. She was born in London, England as the daughter of the noted biologist Conrad Hal Waddington. Her mother, Justin, born Justin Blanco White, was an architect, while her maternal grandmother was the feminist Amber Reeves, a lover of H.G. Wells and an author...

 and anthropologist Caroline Humphrey.

In the early 1930s, Waddington and many other embryologists looked for the molecules that would induce the amphibian neural tube. The search was, of course, 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
Thomas Hunt Morgan
Thomas Hunt Morgan was an American evolutionary biologist, geneticist and embryologist and science author who won the Nobel Prize in Physiology or Medicine in 1933 for discoveries relating the role the chromosome plays in heredity.Morgan received his PhD from Johns Hopkins University in zoology...

 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 30's, 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 wing (this was the essence of the approach that won the 1995 Nobel prize in medicine for Christiane Nüsslein-Volhard
Christiane Nüsslein-Volhard
Christiane Nüsslein-Volhard is a German biologist who won the Albert Lasker Award for Basic Medical Research in 1991 and the Nobel Prize in Physiology or Medicine in 1995, together with Eric Wieschaus and Edward B...

 and Eric F. Wieschaus
Eric F. Wieschaus
-External links:***, excellent profile**...

). 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
In biology, and specifically genetics, epigenetics is the study of heritable changes in gene expression or cellular phenotype caused by mechanisms other than changes in the underlying DNA sequence – hence the name epi- -genetics...

, a term that then meant the external manifestation of genetic activity.

Waddington also coined other essential concepts, such as canalisation
Canalisation (genetics)
Canalisation is a measure of the ability of a population to produce the same phenotype regardless of variability of its environment or genotype. In other words, it means robustness. The term canalisation was coined by C. H. Waddington...

, which refers to the ability of an organism to produce the same phenotype 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. He thus demonstrated that the ideas of inheritance put forward by Jean-Baptiste Lamarck
Jean-Baptiste Lamarck
Jean-Baptiste Pierre Antoine de Monet, Chevalier de la Marck , often known simply as Lamarck, was a French naturalist...

 could, in principle at least, occur.

In 1972, Waddington founded the Centre for Human Ecology.

Epigenetic landscape

Waddington's epigenetic landscape
Epigenetic landscape
Epigenetic landscape is a metaphor for biological development. Its originator, Conrad Hal Waddington, said that cell fates were established in development much like a marble rolls down to the point of lowest local elevation...

 is a metaphor for how gene regulation
Regulation of gene expression
Gene modulation redirects here. For information on therapeutic regulation of gene expression, see therapeutic gene modulation.Regulation of gene expression includes the processes that cells and viruses use to regulate the way that the information in genes is turned into gene products...

 modulates development. One is asked to imagine a number of marbles rolling down a hill towards a wall. The marbles will compete for the grooves on the slope, and come to rest at the lowest points. These points represent the eventual cell fates, that is, tissue
Biological tissue
Tissue is a cellular organizational level intermediate between cells and a complete organism. A tissue is an ensemble of cells, not necessarily identical, but from the same origin, that together carry out a specific function. These are called tissues because of their identical functioning...

 types. Waddington coined the term Chreode
Creode is a neologism coined by the biologist C.H. Waddington to represent the developmental pathway followed by a cell as it grows to form part of a specialized organ. Combining the Greek roots for "necessary" and "path," the term was inspired by the property of regulation...

 to represent this cellular developmental process. This idea was actually 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 was the first person to emphasise that evolution mainly occurred through mutations that affected developmental anatomy.

Waddington 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 1960 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, C. H. (1939). An Introduction to Modern Genetics. London : George Alien & Unwin Ltd.
  • Waddington, C. H. (1940). Organisers & genes. Cambridge: Cambridge University Press.
  • Waddington, C. H. (1941). The Scientific Attitude, Pelican Books
  • Waddington, C. H. (1946). How animals develop. London : George Allen & Unwin Ltd.
  • Waddington, C. H. (1953). The Epigenetics of birds. Cambridge : Cambridge University Press.
  • Waddington, C. H. (1956). Principles of Embryology. London : George Allen & Unwin.
  • Waddington, C. H. (1957). The Strategy of the Genes. London : George Allen & Unwin.
  • Waddington, C. H. (1959). Biological organisation cellular and subcellular : proceedings of a Symposium. London: Pergamon Press.
  • Waddington, C. H. (1960). The ethical animal. London : George Allen & Unwin.
  • Waddington, C. H. (1961). The human evolutionary system. In: Michael Banton (Ed.), Darwinism and the Study of Society. London: Tavistock.
  • Waddington, C. H. (1966). Principles of development and differentiation. New York: Macmillan Company.
  • Waddington, C. H. (1966). New patterns in genetics and development. New York: Columbia University Press.
  • Waddington, C. H., ed. (1968–72). Towards a Theoretical Biology. 4 vols. Edinburgh: Edinburgh University Press.
  • Waddington, C. H., Kenny, A.
    Anthony Kenny
    Sir Anthony John Patrick Kenny FBA is an English philosopher whose interests lie in the philosophy of mind, ancient and scholastic philosophy, the philosophy of Wittgenstein and the philosophy of religion...

    , Longuet-Higgins, H.C.
    H. Christopher Longuet-Higgins
    Hugh Christopher Longuet-Higgins FRS was both a theoretical chemist and a cognitive scientist. He was born on April 11, 1923 in Kent, England and died on March 27, 2004....

    , Lucas, J.R.
    John Lucas (philosopher)
    - Overview :John Lucas was educated at Winchester College and Balliol College, Oxford, where he studied first mathematics, then Greats , obtaining first class honors, and proceeding to an MA in Philosophy in 1954. He spent the 1957-58 academic year at Princeton University, deepening his...

     (1972). The Nature of Mind, Edinburgh: Edinburgh University Press (1971-3 Gifford Lectures
    Gifford Lectures
    The Gifford Lectures were established by the will of Adam Lord Gifford . They were established to "promote and diffuse the study of Natural Theology in the widest sense of the term — in other words, the knowledge of God." The term natural theology as used by Gifford means theology supported...

     in Edinburgh, online)
  • Waddington, C. H., 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)
  • Waddington, C. H. (1977) (published posthumously). Tools for Thought. London: Jonathan Cape Ltd.


  • Waddington C. H. 1942. Canalization of development and the inheritance of acquired characters. Nature 150:563-565.
  • Waddington, C. H. 1953. Genetic assimilation of an acquired character. Evolution 7: 118-126.
  • Waddington, C. H. 1953. Epigenetics and evolution. Symp. Soc. Exp. Biol 7:186-199.
  • Waddington, C. H. 1956. Genetic assimilation of the bithorax phenotype. Evolution 10: 1-13.
  • Waddington, C. H. 1961. Genetic assimilation. Advances Genet. 10: 257-290.
  • Waddington, C. H. 1974. A Catastrophe Theory of Evolution. Annals of the New York Academy of Sciences 231: 32-42.

External links

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