C. H. Waddington
|Conrad Hal Waddington|
Conrad Hal Waddington, FRS, FRSE
|Born||8 November 1905
Evesham, Worcestershire, England
|Died||26 September 1975 (aged 69)
|Fields||Developmental biology, genetics, paleontology|
|Institutions||Cambridge University, Christ's College
University of Edinburgh
Centre for Human Ecology
|Alma mater||Cambridge University|
|Known for||Epigenetic landscape, canalisation, homeorhesis, genetic assimilation, chreod|
|Influences||Alfred North Whitehead|
|Influenced||Jean Piaget, Sanford Kwinter, Gregory Bateson, Margaret Mead|
Conrad Hal Waddington CBE FRS FRSE (8 November 1905 – 26 September 1975) was a British developmental biologist, paleontologist, geneticist, embryologist and philosopher who laid the foundations for systems biology, epigenetics, and evolutionary developmental biology.
He had wide interests that included poetry and painting, as well as left-wing political leanings. In his book The Scientific Attitude (1941), he touched on political topics such as central planning, and praised Marxism as a "profound scientific philosophy".
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 of Kerala. 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 [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's philosophy in the 1920s and 30s and the cybernetics of Norbert Wiener and others in the 1940s.
He attended Clifton College and Sidney Sussex College, Cambridge. He took the Natural Sciences Tripos, 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, C. P. Snow, Solly Zuckerman, Joseph Needham, and John Desmond Bernal. His interests began with palaeontology but moved on to the heredity and development of living things. He also studied philosophy.
After the war, in 1947, he replaced Francis Albert Eley Crew as Professor of Animal Genetics at the University of Edinburgh. 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 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, but ended in 1936. He then married Margaret Justin Blanco White, daughter of the writer Amber Reeves, with whom he had two daughters, the anthropologist Caroline Humphrey (1943-) and mathematician Dusa McDuff (1945-).
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 wing (this was the essence of the approach that won the 1995 Nobel prize in medicine for Christiane Nüsslein-Volhard and Eric F. Wieschaus). 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 also coined other essential concepts, such as canalisation, 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.
In 1972, Waddington founded the Centre for Human Ecology. 
Waddington's epigenetic landscape is a metaphor for how gene regulation modulates development. Among other metaphors, Waddington asks us to imagine a number of marbles rolling down a hill. 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 types. Waddington coined the term chreode 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 proposed an evolutionary process, "genetic assimilation", as a Darwinian 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. This is a trait that 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 has been “assimilated".
Genetic assimilation is sometimes cited as a Lamarckian mechanism, but Waddington described his mechanism as Darwinian. The biologist Wallace Arthur agreed, writing that "genetic assimilation, looks, but is not Lamarckian. It is a special case of the evolution of phenotypic plasticity".
Waddington is usually cited as a Darwinian and not a non-Darwinian. However some of the neo-Darwinian "modern synthesis" theorists described Waddington as a "non-Darwinian" as he claimed that different processes to those of microevolution drive macroevolution. According to (Wilkins, 2009) "[Waddington] in his lifetime... was widely perceived primarily as a critic of Neo-Darwinian evolutionary theory. His criticisms of Neo-Darwinian evolutionary theory were focused on what he saw as unrealistic, “atomistic” models of both gene selection and trait evolution." In particular, he 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 a false postulate. Even though Waddington became critical of the neo-Darwinian synthetic theory of evolution, he still described himself as a Darwinian and his work attempted to correct and expand neo-Darwinism, not replace it. He called for an extended evolutionary synthesis based from his research on epigenetics and genetic assimilation.
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.
- Robertson, Alan. 1977. "Conrad Hal Waddington. 8 November 1905–26 September 1975." Biographical Memoirs of Fellows of the Royal Society 23, 575-622. pp. 575-76.
- Waddington, C. H. 1975. The Evolution of an Evolutionist. Ithica, NY: Cornell University Press. Pg. 2.
- Robertson, Alan. 1977. "Conrad Hal Waddington. 8 November 1905 — 26 September 1975." Biographical Memoirs of Fellows of the Royal Society 23, 575-622. Pg 577.
- Supplement, Historical Register of the University of Cambridge, 1921-30, Cambridge: Cambridge University Press, 1932, p. 63.
- Robertson, Alan. 1977. "Conrad Hal Waddington. 8 November 1905 — 26 September 1975." Biographical Memoirs of Fellows of the Royal Society 23, 575-622. Pp. 579-580.
- Yoxen, Edward. 1986. "Form and Strategy in Biology: Reflections on the Career of C. H. Waddington." In A History of Embryology, edited by T. J Horder, J. A Witkowski, and C. C Wylie. Cambridge: Cambridge University Press. pp. 310-11.
- "Guide to the Center for Advanced Studies Records, 1958 - 1969". Wesleyan.edu. Retrieved 2010-04-04.
- Robertson, Alan. 1977. Conrad Hal Waddington. 8 November 1905 — 26 September 1975. Biographical Memoirs of Fellows of the Royal Society 23, 575-622. P. 578
- Goldberg, A. D., Allis, C. D., & Bernstein, E. (2007). Epigenetics: A landscape takes shape. Cell, 128, 635-638.
- Allen, Matthew. 2015. "Compelled by the Diagram: Thinking through C. H. Waddington’s Epigenetic Landscape." Contemporaneity 4.
- Adam R. Navis, "Conrad Hal Waddington", Embryo Project Encyclopedia (2007) ISSN 1940-5030.
- Science, Politics or Lamarckism? C. H. Waddington’s alternative approach to Darwinism by James F. Stark
- Wallace Arthur Evolution: A Developmental Approach 2011
- Elizabeth Owen, Eve Daintith The Facts on File Dictionary of Evolutionary Biology 2009 p. 235
- Wilkins, Adam S. (2008). Waddington's Unfinished Critique of Neo-Darwinian Genetics: Then and Now. Biological Theory 3: 224-232.
- Huang S. (2011). The molecular and mathematical basis of Waddington’s epigenetic landscape: A framework for post-Darwinian biology? BioEssays 34: 149-157.
-  Hall BK. 2004. In search of evolutionary developmental mechanisms: the 30-year gap between 1944 and 1974. J Exp Zool B Mol Dev Evol. 2004 Jan 15;302(1):5-18.
- Robertson, Alan. 1977. Conrad Hal Waddington. 8 November 1905 — 26 September 1975. Biographical Memoirs of Fellows of the Royal Society 23, 575-622. P. 585.
- Waddington, C. H. (1939). An Introduction to Modern Genetics. London : George Alien & Unwin.
- Waddington, C. H. (1940). Organisers & Genes. Cambridge: Cambridge University Press.
- Waddington, C. H. and others (1942). Science and Ethics, George Allen & Unwin.
- Waddington, C. H. (1946). How Animals Develop. London : George Allen & Unwin.
- Waddington, C. H. (1948). The Scientific Attitude, Pelican Books
- 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. (1961). The Nature of Life. London : George, Allen, & Unwin.
- Waddington, C. H. (1962). New Patterns in Genetics and Development. New York: Columbia University Press.
- Waddington, C. H. (1966). Principles of Development and Differentiation. New York: Macmillan Company.
- Waddington, C. H., ed. (1968–72). Towards a Theoretical Biology. 4 vols. Edinburgh: Edinburgh University Press.
- Waddington, C. H., Kenny, A., Longuet-Higgins, H.C., Lucas, J.R. (1972). The Nature of Mind, Edinburgh: Edinburgh University Press (1971-3 Gifford Lectures 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.
- Waddington, C. H. 1942. Canalization of development and the inheritance of acquired characters. Nature 150(3811):563–565.
- Waddington, C. H. & Carter T. C. 1952. Malformations in mouse embryos induced by trypan blue. Nature 169(4288):27-28.
- Waddington, C. H. 1952. Selection of the Genetic Basis for an Acquired Character. Nature 169(4294):278.
- 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.