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Our History - Timeline


Key events include:

  • Cyril Dean Darlington joins JIHI
  • William Bateson dies and is replaced as director by Sir Alfred Daniel Hall.
  • John Burdon Sanderson Haldane joined staff as part-time head of genetical research and developed the application of mathematical theory to genetics.
  • Dorothy Cayley, JIHI mycologist, showed ‘breaking’ in tulips caused by a transmissible virus.

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Nucleic acid is found to be a major component of chromosomes but its molecular structure is thought to be too simple to carry genetic information.

William Bateson investigates T. H. Morgan’s chromosome theory by experiments on peas. He tests Morgan’s assumption that the number of independent elements cannot exceed the number of chromosomes. For the next few years Bateson also continues to work on physiological aspects of plants and forms of reproduction where segregation of chromosomes plays no part. He studies root cuttings, chimaeras (plants composed of tissues of two or more genetically distinct types), variegated plants, and rogue peas (peas that do not come true from seed), all in the hope of finding an alternative to Morgan’s chromosome theory. Bateson’s objections to the chromosome theory stemmed from his requirement that it should not only explain heredity but also differentiation and development in plants. Other genetic research at JIHI focuses on studies of linkage.

The building of the new Laboratory and Library at Merton is completed. The laboratory provides seven new bench-places, prep rooms, a dark room, and office space. The library is a handsome room, fitted with a lantern for slides. It serves as a lecture room and enables JIHI to accommodate large scientific meetings.

The first plant of economic value resulting from the application of the theory of linkage is produced at JIHI. Dorothea de Winton’s new combination of red stigma and red foliage in Primula sinensis is given to the seed firm of Suttons in Reading, Berkshire, who after purifying the petal-colour and raising a stock, exhibit the new variety as ‘Etna’, winning an RHS Award of Merit. This work laid the foundation of many other striking colour-combinations.

Morley Benjamin Crane begins a research programme on apple root-stocks with East Malling Research Station. This collaboration later results in the development of the MM and MI series of rootstocks for woolly aphid resistance and to control tree vigour. Crane’s main research is concentrated on elucidating the fertility, sterility, and cross-incompatibilities of fruit trees.

‘Evolutionary Faith and Modern Doubts’
In December 1921 William Bateson gives a plenary address to the American Association for the Advancement of Science annual meeting at Toronto. Press reports misrepresent Bateson’s comments on evolution as lending support to the campaign against teaching evolution in state schools. Bateson views the campaign as ‘a terrible example of the way in which truth can be perverted by the ignorant’. In a second lecture to the Zoology section of the meeting titled ‘The outlook of genetics’, Bateson pays homage to T. H. Morgan’s work and announces his partial conversion to chromosome theory but withholds his assent to what he calls the ‘many extensions’ of chromosome theory, including linkage theory. He is en route to visit Morgan’s lab in person and other plant and animal breeding centres in the USA including Cold Spring Harbor, Bussey Institution (Boston), Wistar Institute, and the universities of Pennsylvania (Philadelphia), Cornell (Ithaca), Michigan (Ann Arbor) and Yale.

See also A. G. Cock on Bateson’s two Toronto addresses:


WCF NewtonWilliam Bateson visits T. H. Morgan at Columbia University in New York in December. He spends a week there in laboratory sessions and at Morgan’s home and accepts the principal points of the chromosome theory of heredity.

Bateson was glad that he had visited Morgan and his team to see the cytological work for himself: ‘I was drifting into an untenable position which would soon have become ridiculous’. On his return to JIHI Bateson promotes the study of cytology by appointing W C F Newton

Bateson’s former collaborator R. C. Punnett publishes a comprehensive genetic analysis of the sweet pea in the Journal of Genetics. The result of his important investigation is consistent with the chromosome theory of linkage.

T. H. Morgan and his Drosophila team come to London. William Bateson arranges for the London branch of Spencer microscopes to lend them as many modern microscopes as they need for their demonstrations at the Royal Society and at the annual meeting of the Genetical Society which is held at JIHI in June. Here Morgan and A. H. Sturtevant demonstrate the brilliant work of the Drosophila school aided by hat-boxes filled with Drosophila melanogaster and D. simulans. This was one of the most important and successful early meetings of the Genetical Society: ‘The enthralled audience [of 120 +] had no difficulty in recognising how greatly Drosophila had contributed to the advancement of genetics… to us it seemed that the vinegar fly was to be regarded as one of the most important immigrants to enter the United States- probably with the bananas from South-East Asia’ (Crew, 1969). At the meeting Bateson’s long-time collaborator R. C. Punnett made a ‘handsome’ retraction of the doubts he had entertained concerning the identification of the chromosome with the physical mechanism of Mendelian heredity but Bateson continued to doubt aspects of the theory until his death.

See also:

F.A. E. Crew, ‘Recollections of the early days of the genetical society’, in The Genetical Society- the first fifty years edited by John Jinks. Edinburgh: Oliver and Boyd, 1969, pp. 9-15

A. G. Cock, ‘William Bateson’s rejection and eventual acceptance of chromosome theory’, Annals of Science, 40 (1983): 19-59

Donald Forsdyke, ‘Bateson’s enemies after 1926: William Coleman’

CD DarlingtonAfter a probationary period in the ‘Ladies lab’ with Caroline Pellew and Dorothea de Winton, Cyril Dean Darlington is given an ‘out-of-date second-hand microscope’ and begins cytological research with W C F Newton.   From Newton he learns extreme scepticism of all past cytological work, and particularly of the work of British cytologists, John Bretland Farmer, Lettice Digby and Reginald Ruggles Gates. Bateson, Newton and Darlington are united by a common bond of dislike for Farmer, Gates and E. W. MacBride, the first two they dislike for their (as they see it) unreliable cytological observations, the last for his anti-genetical Lamarckian views. Challenging these biologists gives Darlington’s work direction and momentum until 1930.

Darlington is encouraged to work on three sets of problems. Firstly, studies on the structure, mechanics and division of the chromosomes particularly at meiosis using Liliaceous plants with large chromosomes, this was first done in close collaboration with W C F Newton; Secondly, studies on plants with a known genetic background (and readily available at JIHI) such as Primula sinensis and Prunus species, but with chromosomes that were small and less favourable for study; and Thirdly, the study of variegation by breeding not by cytology, particularly in Vicia faba (broad beans). Darlington’s first choice is the study of meiosis in large chromosomes, the study of Prunus and Primula is very much a second choice, and the study of variegation is to him a ‘chore’ and an example of assisting the regular staff (in fact, Bateson). Darlington brings all three of these diverse studies to fruition during the course of his career, but it is the first area and the series of papers he produces on meiosis, chiasmata and crossing over and chromosome pairing in diploids and polyploids that, culminating in 1932, makes an immediate worldwide impact.


Dan Lewis, ‘Cyril Dean Darlington 1903-81’, Heredity, 48, 2 (1982): 161-67.


Audio ClipListen to Darlington describing his arrival at JIHI, taken from an interview with BJ Harrison; 1979

The seed firm Messrs Suttons of Reading, Berkshire, to whom a set of JIHI’s ‘non-bolting’ strain of Golden Tankard mangolds was sent some years ago, sell the seed commercially for the first time.

William Bateson is invited to give an address in March 1924 to mark the centenary of Birkbeck College, London. In his lecture he pays tribute to T. H. Morgan and his team of cytologists who had proved ‘some, probably all, of this group of [transferable] characters are transmitted by elements in or attached to the chromosomes. Possibly, . . . the visible distinctions were produced not by the presence or absence of a piece of chromosome material, but by an interaction between the chromosomes as a whole. . .’

Chromosomes begin to be a feature of Genetical Society meetings.

In August 1924 Bateson is invited by the Swedish Mendelian Society to tour the Scandinavian biological research stations. He sees experiments directed by Herman Nilsson-Ehle and others and also visits geneticists at Copenhagen and at Stockholm. Bateson comments that ‘collectively, the Scandinavian institutions both in purpose and method have much in common with our own’.

Bateson gives an address on his work. He discovers that the Swedes seem rather ‘half-hearted’ about ‘the chromosome cult’ and have misgivings about Morgan’s theory. Bateson ‘hammered a few wedges into the cracks’ and was pleased to find support for his distrust of American findings on chromosomes.

Bateson advises the seed firm Messrs Vilmorin of Paris on experiments to eliminate bolting from sugar beet, paralleling methods used at JIHI for mangolds.

The John Innes Trustees buy 5 acres of land for a new fruit sub-station and an 8 ft. concrete boundary wall is constructed to prevent pilfering of fruit (the JIHI is surrounded by people who do not have enough to eat). In 1925 the Trustees agree to build a Fruit Room, workshops, a carpenter’s shop and laboratories.

Plants arrive from Wisley (R.H.S.) for trials of commercial varieties of fruit under a joint committee of the Ministry of Agriculture and the Royal Horticultural Society.

William Bateson visits the new USSR to participate in the celebrations to mark the 200th Anniversary of the Russian Academy of Science. He is motivated by curiosity to see the Russian experiment and desires to help restore cultural and intellectual life in that country by reviving scientific interchange. On his return Bateson publishes an article in Nature giving his impressions of scientific work in Russia. Soviet plant geneticist Nikolai Vavilov commented that Bateson’s remarks were quite true but ‘didn’t please us very much’. However, Bateson was impressed by Vavilov’s Institute of Applied Botany and Plant Breeding near Leningrad (St. Petersburg), housed in one of the former Tsar’s palaces requisitioned by the Communists. Here the main object was to provide varieties of cereals and other economic plants suited to grow in the various climatic regions of Russia

See also:

W. Bateson and H. A. Miers, ‘Science in Russia’, Nature, 116 (7 November 1925): 681-3.

William Bateson dies after a brief and unexpected illness on 8 February 1926 at the age of 64. His last paper, ‘Segregation’ is published in the Journal of Genetics.  Here he lists his continuing difficulties with chromosome theory, argues that it leaves too many of the problems of heredity and variation in plants unexplained, and concludes that the acceptance of chromosome theory as a general theory of heredity should be postponed. Readers of the Journal, have come to identify Bateson’s ‘Merton years’ with his series of investigations on variegation, bud-sports, root cuttings, and ‘rogues’ in peas.

C D Darlington later wrote: ‘The disappearance of his powerful personality left English genetics looking very empty indeed… the development of genetics was widely resented as a threat to estab[lished] teaching of botany & zoology in the English universities. The conflict was keenest in the University of London & on the governing body of John Innes itself’. Consequently the John Innes Horticultural Institution faced an uncertain future, it was far from clear who Bateson’s scientific successor would be, or in what direction the Institution would be taken.


Sir A Daniel HalSir Alfred Daniel Hall, who is well known as a former Director of Rothamsted, is appointed Director.  Inexperienced in genetics himself, he appoints J B S Haldane to the staff as part-time head of genetical research. Hall’s actions and the arrival of Haldane come as a relief to the researchers at JIHI. C D Darlington notes that Hall might well have dismantled the genetics research, or might have followed the promptings of the cytologist R Ruggles Gates, or of E W MacBride or J B Farmer on JIHI’s Governing Council. By ‘miraculous good fortune’ Hall did none of these things but took Julian Huxley’s advice and recruited Haldane. Haldane went on to develop the application of mathematical theory to genetics.

Under Hall and Haldane biochemical research is introduced at JIHI. A fruit store is equipped as a chemical laboratory with special rooms ‘for polariscope and combustions’ and Haldane announces his intention to start ‘a systematic biochemical study of flower colour variation in the extensive genetical material available’ at JIHI.

Dorothy CayleyDorothy Cayley, JIHI’s mycologist, shows that ‘breaking’ in tulips is caused by a transmissible virus

The phenomenon of ‘breaking’ in garden tulips had been known for several centuries but the cause was not understood. ‘Breaking’ describes colour variegation of the petals into bi-colours; the original colour is broken into splashes, stripes or lines distributed on a white or yellow background. This pretty effect was nevertheless a commercial disadvantage, many tulip growers wanted to ‘rectify’ their tulips to get ‘true’ colours. Dorothy Cayley’s work showed that colour ‘breaking’ was due to ‘virus or enzyme infection’, probably spread by aphid attack, and that the effect could be artificially produced by bringing the internal tissue of a normal bulb into contact with tissue from a ‘broken’ bulb during the resting stage. Her experiments suggested the degree of breaking was proportional to the amount of infected tissue introduced. Cayley published her results in the Annals of Applied Biology in November 1928.

Hermann J. Muller, at the University of Texas at Austin, USA, reports that x-rays cause artificial gene mutations in Drosophila in a dose-dependent fashion. During the 1920s Muller established most of the principles of spontaneous gene mutation. In C D Darlington’s office at JIHI Muller’s reprints are kept in a box file annotated with ‘H. G. M.’ and a crown.


Biographical details and significance of Muller’s mutation research

W C F Newton, JIHI Cytologist, dies at the age of 32. He is succeeded by C D Darlington, and Charles Leonard Huskins is appointed to work on ‘cytology and plant breeding’. A deep antagonism develops between Huskins and Darlington sparked by Darlington’s freedom to pursue his own research and their rival opinions in cytology. Huskins severely criticises Darlington’s book after he leaves the Institution in 1930.

C D Darlington shares a course of 10 lectures with Charles Leonard Huskins comprising the first two-week summer course in cytology at the John Innes Horticultural Institution. These summer courses, which included lectures on genetics by Haldane, were held biennially until World War II and were initiated to fill a gap in higher education in cytology and genetics training. They were usually attended by 30 to 45 undergraduate and postgraduate students from Britain and overseas. The students of London University’s colleges were already offered the opportunity to learn something of the subject from annual visits to JIHI to see chromosomes and the segregating families of Primula sinensis, but in the late 1920s there was a feeling that more was needed. There were then only three departments of genetics in British Universities: the Galton Laboratory at University College, London (1904), the Department of Genetics at Cambridge (1909) and the Department of Animal Genetics at Edinburgh (1921). In addition there were a few individuals in departments of Botany and Zoology who were active practitioners of cytogenetics. Consequently opportunities for specialist training in the field were few and far between. Later Darlington commented wryly: ‘The fortnight’s course was regarded as a sufficient and permanent qualification for a teacher in genetics and cytology at most universities. A conclusion flattering to the John Innes Institution but disastrous to the progress of genetics’.


Course of instruction in genetics’ 1928

C D Darlington completes two important papers on polyploidy and Oenothera; these are published in Nature and the Journal of Genetics respectively. On polyploidy Darlington records that his very first idea came in December 1926 when he was travelling to work on the London underground with Alice Gairdner, one of Bateson’s assistants and with whom he collaborated on Campanula. They were discussing the effect of polyploidy on fertility, particularly on the tetraploid cultivar of Campanula percisifolia ‘Telham Beauty’ and the hybrid Primula kewensis. He formulated what became known later as ‘Darlington’s rule’: that there is a negative correlation between the fertility of the polyploid and that of the diploid from which it arose. This rule became the basis for the study of fertility in species, hybrids and diploids and polyploidsuntil it was added to by Ralph Riley whose research on wheat identified specific genes affecting chromosome pairing. Darlington’s paper on the evening primrose Oenothera lamarckiana united this seemingly anomalous plant with Mendelism, the chromosome theory, and his own theories of meiosis.

On Darlington’s research see also:

C D Darlington, ‘Polyploids and polyploidy’, Nature, 124 (1929): 62-64; 98-100

C D Darlington, ‘Ring formation in Oenothera and other genera’, Journal of Genetics, 20 (1929): 345-63.

Dan Lewis, ‘Cyril Dean Darlington 1903-1981’, Biographical Memoirs of Fellows of the Royal Sociey, 29 (1983): 113-157, esp. pp. 137-40.

British medical officer and geneticist Frederick Griffith, working in the Ministry of Health’s labs on a vaccine against pneumonia infections following ‘Spanish flu’,  shows that some component of heat-killed virulent bacteria can ‘transform’ a non-virulent strain to become virulent. The unknown ‘component’ involved is later (1944) identified as DNA


Biography and significance of Griffith’s work

In November 1928 C D Darlington is called to the Director’s office and told that he is not co-operating satisfactorily with other members of staff (probably C L Huskins). For this reason he is to be sent to Persia to collect tulips (and anything else of interest) at the joint expense of the Empire Marketing Board and the John Innes, in the company (or custody) of an experienced botanist.

In February 1929 Darlington embarks at Tilbury en route to Iraq and Persia to collect Prunus and Tulipa species with John MacQueen Cowan, a retired Indian Forest Officer working for the Royal Botanic Gardens at Kew. Cowan’s goal was to collect 5000 specimens of plants in flower, to be dried and sent back to Kew labelled ‘Coll. Cowan and Darlington’. Though Darlington deprecated the ‘intellectual vacuum’ at the heart of this expedition, the collection of both these genera came to be essential for Darlington’s chromosome studies. These travels also stimulated his later interest in human society. Darlington and Cowan separated at the Russian frontier in June 1929; Darlington went on to make an unescorted visit to Russian geneticist Nikolai Vavilov before returning to England, bearing amoebic dysentery and malaria. Darlington revered Vavilov; his picture hung among the few portrait photographs in Darlington’s lab.  Darlington became an activist for Vavilov’s release after his disappearance at the hands of the Russian authorities in 1940.

Example of Osterstock's illustrations of TulipsSir Alfred Daniel Hall's The Book of the Tulip is published in May, beautifully illustrated by Herbert Osterstock’s flower paintings. This and Hall’s sequel The Genus Tulipa (1940) were the last occasions that a flower painter was employed at JIHI to illustrate scientific data.

Photography became the preferred medium for recording scientific observations.

Rose Scott-MoncrieffRose Scott-Moncrieff begins work with J B S Haldane on the chemistry of flower colour with a small grant from the Department of Scientific and Industrial Research. In the early years of their collaboration Scott-Moncrieff is based in the biochemical laboratory of Professor Gowland Hopkins at the University of Cambridge where Haldane holds a Readership. Their experimental material is mainly at Merton where work on the chemistry of anthocyanins can begin on plants of ‘known genetical composition, in order to determine the precise nature of the change produced by a factor’. Haldane puts her in touch with JIHI’s geneticists to begin a biochemical survey of related genotypes. Scott-Moncrieff has already isolated the anthocyanin from purple Antirrhinum majus and begins work on the red variety and on flower colour in different races of Primula sinensis. Scott-Moncrieff is also mentored by Muriel Wheldale Onslow who in the 1910s worked on the chemical genetics of flower colour variation, first at JIHI and later in Gowland Hopkins’ lab at Cambridge.