Conrad Hal Waddington (1905–1975)

1. Brief Vita

Conrad Hal Waddington was born in Evesham, England in 1905 and died in Edinburgh, Scotland in 1975.  After spending the first four years of his life in India where his father worked at a tea plantation, he returned to England and attended Clifton College School and Sydney Sussex College, University of Cambridge, where he read natural sciences and graduated in 1926 with First Class Honours in geology.  He then began postgraduate studies at Cambridge focusing on paleontology and geology but never presented his thesis on the ammonites because he changed his interest to study “’live’ biology rather than fossils” (Robertson, 1977, 578).[i] Cambridge, however, awarded Waddington an ScD in 1938 on the basis of his published work in genetics and embryology.

Waddington first became interested in evolution through his geology tutor, Gertrude Lilian Elles, and genetics through his friendship with Gregory Bateson—son of William Bateson, who introduced Mendelian genetics in Britain in 1900. His his first academic appointment was a research fellowship at the Strangeways Research Laboratory in Cambridge. In 1930 Waddington became a demonstrator in zoology at Cambridge through a part-time grant from the Medical Research Council.  In 1933 he became a lecturer of zoology at Strangeways Research Laboratory and a Fellow at Christ’s College, Cambridge.  From 1938 to 1939 Waddington traveled to the United States, where he worked at the Carnegie Institute Laboratories at Cold Spring Harbor, Columbia University, and then the California Institute of Technology.

During World War II, Waddington was assigned to the Operational Research Section Coastal Command of the Royal Air Force tasked with operational research against the German U-boats in 1942 and subsequently became officer in charge of that section in 1944. After the war, in 1945, he became chief geneticist at the National Animal Breeding and Genetics Research Organization and was then appointed to the Chair of Genetics at the University of Edinburgh.  

Waddington became a Fellow of the Royal Society of London in 1947 and Fellow of the Royal Society of Edinburgh in 1948, was named to the Finnish Academy in 1957 and named a Commander of the Order of the British Empire in 1958. In 1959 he became a foreign member of the American Academy of Arts and Sciences. He was also a Fellow of the Deutsche Akademie der Naturforscher Leopoldina in Germany. He was awarded honorary ScD degrees from the University of Montreal in 1958, Trinity College, Dublin in 1965, Prague in 1966, and Geneva in 1968, and received an LLD from Aberdeen in 1966. 

The Waddington Medal, awarded by the British Society for Developmental Biology for outstanding research performance as well as services to the subject community, is named in honor of C. H. Waddington.

2. Theoretical Biology

Waddington was a polymath of wide learning who successfully crossed the boundaries of what C. P. Snow called the “two cultures” of the arts and sciences.  It is rare to find one who made important contributions to the advance of science and writes with such a general understanding of philosophical issues.  In this regard, Waddington would most certainly qualify as a “Natural Philosopher” in the Enlightenment sense of the term.  He was the founder of epigenetics and proposed a model of genetic assimilation to compete with the Neo-Darwinian theory of evolution by natural selection.

Waddington was also one of the founders of the Theoretical Biology Club at Cambridge in the 1930s whose members advanced a philosophy of biology, “organicism,” that would offer an alternative to the reductionism of mechanistic materialism and the obscurity of vitalism in coming to terms with the dynamic, interdependent, and purposeful character of life.  This view was embraced in one form or another by E. S. Russell, John Scott Haldane, C. Lloyd Morgan, Lawrence J. Henderson, C. D. Broad, and Alfred North Whitehead.  Waddington, in particular, was devoted to the integration of genetics, development, and evolution within the context of theoretical biology. Like Whitehead, who believed that ordinary language was inadequate to express the dynamic nature of reality, Waddington was a prodigious coiner of neologisms, e.g., “epigenetics,” “epigenetic landscape,” “genetic assimilation,” “canalization,” “chreod,” and “epigenotype.” 

Whitehead’s organicism, primarily advanced in works such as Science and the Modern World (1925), Process and Reality (1929) and Nature and Life (1934), is a sustained attack on mechanistic materialism, or as he put it a “repudiation of the doctrine of ‘vacuous actuality’ (PR, xiii). Throughout Process and Reality, he referred to his position as “the philosophy of organism” and developed a metaphysics in which the basic units of existence are understood as occasions of sentient experience, i.e., actual occasions that become and perish in the temporal process (PR, xi). Reality must be conceived as experience from beginning to end; there is no radical emergence of mind or consciousness from insentient matter or living things from non-living things.  Whereas in Whitehead’s earlier works such as The Concept of Nature (1920) and The Principle of Relativity (1922), the basic units of reality are events as the foundation of his philosophy of physics, in the later metaphysical works, it looks more like the events are tiny organisms that form the basis for a biological view of the universe.  As part of his metaphysics, Whitehead formulated a mereological theory that he called “the theory of society,” according to which the extended universe is a nested continuum of organisms and environments. Up and down the continuum from subatomic particles, atoms, molecules and cells to plants and animals and to planets and galaxies and beyond, one finds smaller units of organisms nested in the larger ones, and the interdependence of wholes and parts —all part of the creative advance into novelty.  

Waddington discovered Whitehead’s work as an undergraduate at Cambridge. He said he paid more attention to Whitehead’s writings than the textbooks in the subjects he was meant to read for his exams (1975, 3).  Whitehead had such an impact on him that he abandoned geology and applied for the Arnold Gerstenberg Prize in Philosophy in 1929, which he won with an essay on the vitalist-mechanism controversy. In this essay, Waddington focused on Whitehead’s view of events and process as the fundamental constituents of the world and argued in accordance with Whitehead that biological evolution mirrors the evolving universe.

In his “Autobiographical Note” in The Evolution of an Evolutionist, entitled “The Practical Consequences of Metaphysical Beliefs on a Biologist’s Work,” Waddington acknowledged the influence of Whitehead on his scientific work (1975, 3). He also wrote an essay for a volume based on a talk he gave at Bellagio, Italy in June, 1974—“Whitehead and Modern Science”—where he explains this influence in more detail. Whitehead, he says, provided a new way of looking at the problem created by the dilemma of mechanism and objective vitalism (or reductionism and anti-reductionism), and a new vocabulary with which to express the processual and relational nature of experience (1977, 143; 1962, 19-20).[ii] As for the question―Can living things be treated as if they are nothing but mechanisms constructed of mere material components?―the mechanistic biologists and reductionists answered in the affirmative while the objective vitalists and anti-reductionists answered in the negative.  Whitehead, he thinks, resolved this dilemma by rejecting the whole conceptual framework in which the problem was framed.  Living things cannot be accounted for in concepts that were formulated for the non-living world. 

Organicists such as Waddington held that an organism does not exist at an instant, nor is it a bag of chemicals, each of which is produced by the influence of some particular gene.  It is rather a process spread out in time and the explanation of its functioning requires more than the atomistic account of genes on heredity.  Also, it is essential to an organism that its overall form and the properties of its component parts are a result of the reciprocal play of all the components on one another (1962, 53–54).

So, not only did Waddington embrace metaphysics at a time when orthodox biology rejected such speculation as unscientific, he also embraced Whitehead’s metaphysics against the status quo—a view of nature as alive and purposeful.[iii] In his view, it wasn’t a question of whether to engage in metaphysics or not; the orthodox view had accepted a metaphysical view without acknowledging it as such, a materialistic reductionism of the Democritean-Cartesian variety that Waddington called a “lousy philosophy” (Peterson, 2011, 316).  His whole point about his Whiteheadian orientation was “to illustrate the fact that metaphysical presuppositions may have a definite influence on the way in which scientific research proceeds” (1975, 10).  A metaphysical view that informs a scientist’s work is not a mere epiphenomenon.  In Waddington’s case, he said that Whitehead’s metaphysics had a definite and ascertainable influence on both his research agenda and on the experiments that he conducted in support of his theories (1975, 1).  This included the fundamental ontology of process, which proposed events or occasions of experience as basic, the concept of organism, and the importance of both atomic and continuum theories for understanding development and evolution (1975, 4-5; 1962, 53-54).[iv]

Waddington was also a proponent of panpsychism. He was more of what I would call a “closet panpsychist” by avoiding explicit mention of the term, and he often skirted around the concept of goal orientation in nature by suggesting that something more than mechanisms is needed to explain development and evolution, namely “quasi-finalistic properties” (1962, 98).  When, however, he argued that “you have either got to have consciousness or at least something of that general kind, everywhere; or nowhere” (1969, 114) it is pretty clear that he meant everywhere.  Something, he wrote,  “must go on in the simplest inanimate things which can be described in the same language as would be used to describe our self-awareness” (1962: 121). For his approach to theoretical biology, Waddington is sometimes referred to as a Whiteheadian organicist (Bard, 2017, 2).

3. Development and Evolution

Waddington’s most important scientific achievement was in advancing the concept of embryonic development as a highly integrated series of canalized pathways. His model of an “epigenetic landscape,” as a representation of developmental pathways, became fundamental to the way that most biologists viewed the organization of embryonic development.  Whitehead used the term “concrescence” to describe the process of coming together to create a novel entity which essentially invokes the concept of a subjective aim that drives the process to its endpoint. Waddington said that when he began working in embryology, he adopted a Whiteheadian perspective that development depended on activities of a very large number of genes which were brought together to result in some relatively unified type of action to form the different types of cells (1977, 143–44). He used Whitehead’s concept of concrescence, in what might be called “gene-concrescence,” to explain this biological process (1975: 9–10). 

Waddington described embryonic development in terms of attractors in an epigenetic landscape.  A ‘creode’ describes the path of change determined by initial conditions that once entered upon cannot be abandoned. The valleys in the landscape represented the usual pathways of change through which the organ developed.  As Waddington wrote in The Nature of Life: “We can say then that the heredity materials with which an organism begins life define for it a branching set of creodes. Different parts of the egg will move along one or other of these creodes, so that they will after a long process of progressive changes, finish up as one or other of a number of different end-results, as it might be a heart, muscle, nerve, kidney and so on” (1962, 64). The differentiating cells will thereby reach their endpoints unless there are powerful influences that divert the pathway and the cell or the developing organ ends up in an abnormal condition or a mutation. ‘Canalization’ is the property of developmental pathways to produce standard phenotypes despite environmental or genetic influences.  The phenotype is the actualization of the potential in the genotype given a normal developmental pathway. 

Regarding Whitehead’s influence on Waddington’s view of evolution, he wrote: “…from the Whiteheadian point of view one has to recognize that the evolving events—actual animals and plants as we meet them in real life—are influenced by environmental factors as well as genetic” (1977, 144).

Waddington stated that he put his Whiteheadian outlook to actual use in particular experimental situations. (1975, 10–11).   He wrote: “… when I started doing experiments on Drosophila evolution, in the ’40s and ’50s, I treated even that insect as a developmental system, and by manipulating the environment in which it develops was able to uncover the rather novel process of genetic assimilation. Thus my particular slant on evolution—a most unfashionable emphasis on the importance of the developing phenotype—is a fairly direct derivative from Whiteheadian-type metaphysics” (Robertson, 1977, 597).  The Neo-Darwinist view did not pay any attention to the phenotype but rather to the

genotype in explaining inheritance.  In fact, Neo-Darwinism specifically excludes the inheritance of an acquired characteristic, i.e., Lamarckian inheritance.  Against biological orthodoxy, Waddington sought to challenge this view by showing how embryos can respond to an abnormal environmental situation by producing a new character, i.e., a new phenotype, initially without genetic change.  But then after exposure of subsequent generations to the same environmental change, selection occurs, and a genetic basis for the character will be found in the population without the environmental change. A “soft” inheritance, namely, a change in phenotype in one generation, then becomes “hard” inheritance as that characteristic is programmed into the genotype of succeeding generations.[v] This is what Waddington called “genetic assimilation,” which is the origin of epigenetics, namely, the study of heritable phenotype changes that do not involve alterations in the genetic code.    

3. Conclusion

Waddington saw in Whitehead’s metaphysics the power of generalization that produced basic concepts of process, creativity and organicism which he usefully applied to heredity, development, and evolution.  Whitehead drew upon many sciences—physics, chemistry, biology, cosmology, psychology—when he formulated his metaphysical principles.  He claimed in Process and Reality that success would be measured by the extent to which the generalities of his process metaphysics pass the test of applicability beyond the restricted locus from which they originated, whether they illuminate our experience of the world, have broad explanatory power, and provide unifying concepts for the sciences (PR, 5). Waddington’s biological work might be one of the best examples of this success.  Like Whitehead, Waddington sought to explain fundamentally how novelty is possible.


[i] When Waddington began putting metaphysics to practice, he studied the ammonites as an example of Whitehead’s point that the organisms undergoing the process of evolution are themselves processes.  The ammonite’s spiral shell shows a record of its whole life history by occupying the latest-formed part of the shell, leaving behind the parts it previously inhabited.  The developmental process is preserved, unlike most types of animals in which the adult form replaces the younger stages.  (1975, 50)

[ii] In his essay, “Whitehead and Modern Science,” Waddington wrote that he wanted to write a book about Whitehead and the relevance of his thought to some of our present controversies, particularly in relation to biology, and the ‘anti-science’ movement, but he didn’t live to see it finished. (1977, 143)

[iii] See for example, Erik Peterson’s “The Excluded Philosophy of Evo-Devo? Revisiting C. H. Waddington’s Failed Attempt to Embed Alfred North Whitehead’s ‘Organicism’ in Evolutionary Biology” wherein Peterson argues that Waddington’s explicit support for Whitehead’s philosophy was one reason why Waddington’s views did not have more impact.

[iv] Understanding development required both atomic and continuum theories; genes are the basic atomic units of hereditary material further analyzed into chromosomes, proteins, amino acids, and DNA into nucleotides, and the continuum is addressed by the biological processes by which development occurs, i.e., canalized pathways (1962, 36–37).  In Whitehead’s metaphysics, actual occasions atomize the extensive continuum.  Actual occasions are analyzed into prehensions, eternal objects, subjective forms, etc. Continuity is addressed in terms of nexūs, societies, i.e., events, fields, and matter formed from the prehensive activity of actual occasions (PR, 20–30).

[v] See Noble 2015 for Waddington’s experimental success in his approach by discovering in a population under study the forms of developmental plasticity that already exist and then altering the environment in such a way to find a path that evolution could have taken.  For example, in Waddington’s 1956 paper published in Evolution, “The Genetic Assimilation of the Bithorax Phenotype,” he reported on the Bithorax mutant, a four-winged fruit fly, Drosophilia, with two thorax segments that was the product of experiments that changed the environmental temperature or used a chemical stimulus.

Works Cited and Further Readings

By C. H. Waddington

Selected Books

1940. Organisers and Genes (Cambridge: Cambridge University Press).

1946. How Animals Develop. (London: George Allen & Unwin).

1957. The Strategy of Genes: A Discussion of Some Aspects of Theoretical Biology (London: Allen & Unwin).

1961.  New Patterns in Genetics and Development (New York: Columbia University Press).

1962. The Nature of Life (New York: Atheneum),

1968–72. (ed) Towards a Theoretical Biology. 4 vols. (Edinburgh: Edinburgh University Press).

1969. Beyond Appearance A Study of the Relations between Painting and the Natural Sciences in this Century. (Edinburgh: Edinburgh University Press).

1975. The Evolution of an Evolutionist (Edinburgh: Edinburgh University Press).

Selected Articles and Chapters

1942. “Canalization of development and the inheritance of acquired characters.” Nature 150: 563.

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.

1956. “The Genetic Assimilation of the Bithorax Phenotype,” Evolution 10: 1–13.

1977. “The Process of Evolution and Notes on the Evolution of Mind,” in Mind in Nature: The Interface of Science and Philosophy. Ed. John B. Cobb and David R. Griffin. University Press of America, 27-31.

1977. “Whitehead and Modern Science,” in Mind in Nature: The Interface of Science and Philosophy. Ed. John B. Cobb and David R. Griffin. University Press of America,  143–146.


C. H. Waddington’s papers are kept in the Edinburgh University Library. <>.

Readings on Waddington

Bard, Jonathan B. L., “C. H. Waddington’s Differences with the Creators of the Modern Evolutionary Synthesis: A Tale of Two Genes,” HPLS, (2017): 1–4.

Hall, Brian K. “Waddington’s Legacy in Development and Evolution,” American Zoologist, 32: (1992): 113–122.

Needham, Joseph. “Conrad Hal Waddington, CBE, FRS.” Nature 258 (1975): 371–372.

Newth, D. R. “Conrad Hal Waddington, CBE, FRS.” Nature 258 (1975): 371.

Nicholson, Daniel J., and Dupré, John, (eds). Everything Flows: Towards a Processual Philosophy of Biology. Oxford: Oxford University Press, (2018).

Noble, Denis“Conrad Waddington and the Origin of Epigenetics”.  Journal of Experimental Biology. (2015), 218 (6): 816–818.

Peterson, Erik L.  “The Excluded Philosophy of Evo-Devo? Revisiting C. H. Waddington’s Failed Attempt to Embed Alfred North Whitehead’s ‘Organicism’ in Evolutionary Biology,” History and Philosophy of the Life Sciences, (2011), 33/3: 301–20.

Robertson, Alan. “Conrad Hal Waddington. 8 November 1905 – 26 September 1975.” Biographical Memoirs of Fellows of the Royal Academy 23 (1977): 575–622.

Slack, Jonathan M. W. “Conrad Hal Waddington: The Last Renaissance Biologist?” Nature Reviews Genetics 3 (2002): 889–895.

Author Information

Leemon B. McHenry
Department of Philosophy
California State University, Northridge
18111, Nordhoff Street, California 91330 USA

How to Cite this Article

McHenry, Leemon B., “Conrad Hal Waddington (1905–1975)”, last modified September 2023, The Whitehead Encyclopedia, Brian G. Henning and Joseph Petek (eds.), <>.