Hatching the EGG: the first meeting of the Ecological Genetics Group

Erick Peirson | March 17, 2016, 6:57 p.m.

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David J. Harberd Anthony D. Bradshaw National Institute of Agricultural Botany Ecological Genetics Group Agrostis tenuis J.W. Gregor E. T. Jones Muriel Harberd Scottish Plant Breeding Station Welsh Plant Breeding Station Keith Jones Goginan lead mine Dennis Wilkins Tom Pritchard

The following account of the formation of the Ecological Genetics Group is taken from chapter 4 of my dissertation. It has been slightly tweaked for the web, and there may be some artifacts and typos from the conversion process. Over the next few weeks I'll revise this post with additional annotations, links, and other elaborations.

4.2 Conception

The 1956 meeting of the Fellows of the National Institute of Agricultural Botany (NIAB) was to be held in July, 1956. E. T. Jones was to receive the NIAB Cereal Award for his “Powys” variety of winter oats, and to confer with NIAB leadership about the ongoing transfer of grass and clover seed stocks from the WPBS to NIAB. For members of the SPBS, the annual NIAB meeting was seen as an obligation. “All the plant breeding stations had to get together each year,” one of the Genecology Section staff, David Harberd, recalled, “and you didn't have any choice in the matter: you went. The bosses in London just decided where you were meeting and you went” (Harberd pers. comm.).

As the NIAB meeting approached, however, circumstances were extraordinarily complicated for Harberd. Harberd’s wife, Muriel, had dropped six stone (over 80 pounds) due to complications with her pregnancy, and was still recovering as the NIAB meeting approached. So several days before the trip to Cambridge, Harberd approached Gregor and asked to be excused from the meeting. “I cannot leave Muriel just now,” Harberd explained, “She's gone through such a hell of a time, and I must be here for her.” Gregor assured Harberd that he would sort things out so that he could be excused.

Suspecting that there was more to the story, however, he probed Harberd further. “I don't think you want to go, do you?” Gregor asked. “Of course I want to go,” Harberd replied, “It's jolly good, but I just don't want to be away from home.” After all, he had lived in Cambridge for several years prior to moving to Edinburgh, and many of his friends remained in the area. Gregor pushed further: “Well now, what benefits do you think you get from these [meetings]?” Harberd replied:

Well, it's just social. There’s nobody there will be working on grasses. Last time when we were all over in Belfast, we had to go through flax breeding. Well there wasn't a single person there who knew anything about flax breeding apart from these folk. And it was fascinating to learn their problems and how they were tackling them, but we couldn't contribute. Really, it just so happened that it was fascinating, but if we're going to have a session on breeding watercress or something, I can't imagine how we're going to -- I mean it’s is a different kettle of fish altogether. I really think the money would be much better spent if, instead of the whole station decamping like that, we had an arrangement whereby all the potato breeders could gather together every so often and exchange gossips. And the barley breeders could do this, and all the grass breeders could do this. It would be cheaper, and it would be far more efficient, and we'd really make some progress. (Harberd pers. comm.)

So upon arriving at Cambridge, Gregor mentioned Harberd’s idea to Keith Jones, who had accompanied E. T. Jones from the WPBS. Jones was quite interested in a meeting specifically focused on grasses and grass breeding. Gregor was also fascinated to hear of Bradshaw’s ongoing work on lead tolerance in Agrostis, for one of his own staff members, Dennis Wilkins, had undertaken similar studies on mine tips in Lanarkshire after reading of Bradshaw’s 1952 discovery at Goginan mine. Jones and Gregor agreed to convene a meeting that would bring the staffs of their two departments together and provide an opportunity for Bradshaw and Wilkins to compare notes.

Gregor wrote to Jones around a month later to firm up their plans. Gregor wrote:

When we met at Cambridge I told you of the wish of our genecological group to pay Aberystwyth a visit some time this autumn,” , “to see the work you and your colleages are doing with the wild grasses, and you very kindly agreed to have us. […] You will remember that the suggestion was to get Bradshaw down to Aberystwyth at the same time so that we might hear about his latest work on Agrostis , and particularly to give Wilkins an opportunity to discuss with him the lead tip flora and, if possible, to visit some of the tips in his company (D1041/2/9/22/2).

Gregor proposed a meeting on a Wednesday, the 26th of September, with a field trip to the mine tips on Thursday. Jones passed the letter on to Bradshaw, writing,

I had some prolonged discussions with him at Cambridge some time ago and he is very keen to visit us. He would particularly like you to be here if you can manage it both to hear of your gene-ecological stuff and for you to take Wilkins to the lead mine areas. Watson has some interesting information on hybridisation of Agrostis under natural conditions, and I have no doubt that we should profit by discussions with them (D1041/2/9/22).

Upon hearing back from Bradshaw, Jones replied to Gregor enthusiastically, writing:

I had not forgotten about your desire to visit the W.P.B.S., and indeed we shall be looking forward to seeing you and your part. For the past month I have been trying to get in touch with the elusive Bradshaw, and at last today I received a letter from him from Fort William, where is is on holiday. As far as I can make out he will be able to come here on the 26th September. This date is also suitable for [Martin] Borrill and myself, although Mr. [Arthur R.] Beddows will probably not be here. (NLW ex 2747).

So Gregor, Harberd, Wilkins, and Patricia Watson traveled to Aberystwyth. Gregor and Wilkins passed the time by playing chess in their booth. Bradshaw came down from Bangor on his green motorbike with David Jowett, who had recently arrived from Liverpool, riding on the back. The Edinburgh party arrived at the Aberystwyth train platform in torrential rain. In 1955 the WPBS had moved its operations from its hillside campus at Penglais to a site further east at Plas Gogerddan, so none of the visitors knew which bus to catch. After asking around for directions, navigating buses, and walking in the rain, they eventually arrived at the white mansion at Plas Gogderddan that housed the new administrative offices of the WPBS. They spent the day exchanging notes, catching each other up on their latest research. That night, Harberd and Bradshaw stayed with Keith Jones at his on-campus house, named “Oregon”; when they arrived for dinner that evening, Bradshaw quipped jokingly, referring to the Oregon-R strain of Drosophila melanogaster,

Oh, Oregon. That’s a place for a wild type to live! (Harberd pers. comm.)

The visit was deemed a success, and promises were made to reconvene the following year. In April of 1957, Pat Watson wrote to Keith Jones inviting the WPBS contingent to visit Edinburgh late that summer:

We hope that you and some of your colleagues will be able to come visit us this year, and that the meeting will prove to be as interesting as we found our visit to Aberystwyth last September. As always happens, the summer months promise to be very busy but since we understand that some of you might be interested in the Edinburgh Festival also, I am writing to propose a two-day visit sometime between 15th and 20th August. (NLW ex 2747)

A third meeting was held at Bangor the following year, this time in late March or early April. Bradshaw sent the invitation for a fourth meeting to Gregor on 17 January, 1959.

We would be very pleased indeed if we could hold another of our informal discussions this Easter. We would all be very delighted to see you down here. ... Keith Jones is in Canada so we would not have him with us, but we would hope to have at least Borrill from the P.B.S (GD449/16/6).

Gregor also described the meeting as “informal” in his reply. Bradshaw’s work on lead tolerance in Agrostis also remained a dominant part of the discussion.

Many thanks for your letter inviting our herbage group to informal discussions at Bangor, an invitation we gladly accept. As to the date of the meeting, Wednesday, March 25th would suit us very well and I take it that day would also suit you. Wilkins, however, feels he would like an extra day to see your people about his lead resistance work. He therefore proposed travelling on Monday 23rd and spending the following day discussing the lead programme. (GD449/16/4)

The fifth meeting, in 1960, had a substantially different flavor from those that preceded it. Participation in the group increase substantially, and it was at this meeting that the group adopted the name Ecological Genetics Group and began to establish a somewhat more formal structure. In the next section, I briefly describe the content and focus of that meeting.

4.3. Incorporation

Thomas Owen Pirtchard (b. 13 May, 1932) was a graduate of the Agricultural Zoology Program at Bangor, and shortly after finishing his undergraduate degree went to work as Deputy Director of the Nature Conservancy in Wales. The Nature Conservancy was established in 1949 with an explicit mandate of translating ecological research into public policy for the protection of natural resources (Goodson 1993). Pritchard is credited with the origination of the phrase “environmental education” at the 1948 conference of the International Union for the Conservation of Nature and Natural Resources (IUCN) (Disinger 1984). Pritchard advocated environmental education as both vocational training for resource managers and as way to create “public awareness about environmental affairs, with the ultimate aim of realising the conservation of natural resources and stimulating enjoyment of the environment” (IUCN 1972, p. 1). Pritchard had worked with Bradshaw in the mid-1950s on the genecology of Trifolium repens in Snowdonia, and went on to receive a Ph.D. in 1958 from the University of Leeds for his work on the genecology of Euphorbia cyparissias (cypress spurge) and Hypericum perforatum (St. John’s wort). It is through this association that Pritchard became acquainted with what was becoming a regular Easter meeting of genecologists from Edinburgh, Aberystwyth, and Bangor, and during discussions with Bradshaw became excited about the prospect of hosting such a meeting at the Nature Conservancy office at Attingham Park, just east of Shrewsbury.

In his invitation to Gregor in March, 1960, Pritchard was candid about his desire to address conservation issues at the meeting. Specifically, Pritchard was interested in the conserving “the country’s gene pools.” He wrote:

A few days ago, I discussed with Dr. Bradshaw of Bangor arrangements for holding a meeting of people interested in evolution in natural populations. […] The meeting will be informal as usual and attended by about 20 people actively engaged in research on plant populations. It is expected that one or two short papers will be given to introduce topics for discussion and we would also like the group to give some time to discuss the conservation of the country’s gene pools, particularly old pastures and other areas from which breeding material has been obtained in the past. (GD449/24)

Gregor expressed some trepidation about the anticipated size of the meeting.

I hope that the meeting will be able to retain its thoroughly informal character, for it seems to us that with twenty people it shows signs of becoming just one more conference. JW Gregor (GD449/24)

In the end fifteen people, including Pritchard, arrived for the meeting on Monday, the 25th of April. Harberd was involved in an automobile accident prior to the meeting and was unable to attend. Gregor also missed the meeting, but the SPBS was represented by genecologist Frederick England.

From Bangor, Bradshaw brought his students John Aston and Roy Snaydon. The largest contingent came from the WPBS, including Les Breese, Malcolm Calder, John Cooper, Keith Jones, and Martin Borrill. Some new faces included Michael Harvey, a first-year Ph.D. student at Durham, Audrey Plack from Keele, Miss K. Luck from Cambridge, and Nature Conservancy staffer Miss E. Copeland Watts. Jones, Borrill, and Pritchard presented papers on Monday afternoon, and Goodway, Snaydon, Bradshaw and Cooper presented their recent work the following day. (GD449/25)

On Tuesday afternoon, Pritchard turned the conservation away from research updates and toward “the conservation of ecological races and subspecies.” One of the central themes of that conversation was the idea that the locally adapted plant populations, heretofore exploited by plant breeders, were a valuable natural resource and thus an important object of conservation. Pritchard drew together his notes from that conversation to write a short paper titled The conservation of ecological races, which was circulated internally at the Nature Conservancy. Pritchard wrote:

The great variety of distinct natural and manmade habitats occurring in the British Isles has been conducive to widespread differentiation of ecologic races in plant species. In this country, races differentiated in response to longestablished agricultural and forestry management systems are, to the student of evolution, at least of equal significance as those races adapted to natural habitats which are relatively extreme in terms of edaphic and climatic conditions such as in the higher mountains and along the coastline. (GD449/25)

Bradshaw’s work on Agrostis at Port Meadow in Oxford was cited in the discussion.

As an example, Port Meadow in Oxford … has been recorded as common land belonging to the Burghers of Oxford since Domesday times and, fortunately, it still exists as a habitat where grazing pressure as a selection force has been operative for centuries. The races of grasses and other species occurring there are of special interest to the research worker and plant breeder. Extreme Agrostis stolonifera x A. tenuis hybrids, for example, exist there and further study will inevitably reveal the presence of other interesting types. (GD449/25)

In his write-up, Pritchard warned about the dangers of losing habitats like Port Meadow, and the plant populations therein.

A habitat such as Port Meadow is extremely vulnerable in our time. Discounting complete destruction by ploughing or building development, it can be easily lost through simple changes in management methods. The older grazing systems are rapidly vanishing, the result being degeneration of old pasture communities and disappearance of the ecological races in them. Port Meadow still exists but there are rumours that it was recently sprayed with a chemical weedkiller to control ragwort; obviously, such a practice is bound to have devastating effects on its plant populations. (GD449/25)

The group seemed to agree that “certain ecotypes and subspecies” ought to be conserved both for their value to investigations of “experimental taxonomy and evolution” as well as to provide “useful sources of genes” for agriculturally valuable characters. Pritchard wrote:

It seems that the most urgent task at this stage is to document information on the best localities, and to take appropriate action after this has been considered. Members of the Ecological Genetics Group would be willing to co-operate in such a scheme. (GD449/25)

Finally, the group took up the issue of establishing a more formal structure. They agreed to meet annually, to limit the size of the meeting to “about 15 to 25 persons engaged in experimental studies on populations,” and that the organizers should keep minutes from each meeting. Several names were considered: “Genecology Group” was rejected, and “Evolution Group” was also deemed unfavorable “unless the word Experimental was included.” Ultimately, they adopted the name “Ecological Genetics Group.” Harberd suspected later that Keith Jones might have advocated for the name, partly due to the humor of “hatching” the “EGG” around Easter time (Harberd pers. comm.).

4.4. Diversification

Whereas the 1960 meeting established the identity and purpose of the EGG, the 1961 meeting moved the EGG into a broader institutional arena. Keith Jones, who had recently left the WPBS to work at the Jodrell Laboratory at the Royal Botanical Gardens at Kew, hosted the meeting. Bringing the EGG into southern England drew in a much broader array of participants. Harberd, who had recovered from his automobile accident, remembered the gathering as “one of our most flourishing meetings. That was when we first became not a tight little group, but much wider” (Harberd pers. comm.). In addition to new faces from the research team at Kew, many others attended from the larger Universities in southern England. The well-known American botanist George Ledyard Stebbins, from California, was visiting the UK at the time and made an appearance, along with E. J. H. Corner from Cambridge.

The earliest connections to the animal genetics community at Oxford came in 1961, albeit by way of a botanist. David A. Jones was in the second year of his Ph.D. in the Department of Zoology at Oxford. His invitation to the EGG meeting came “totally out of the blue” (Jones pers. comm.). Although Jones’ earliest graduate research was in animal genetics, even involving Lepidoptera, by 1961 his interests had shifted to cyanogenesis and lectins in plants. Cyanogeneic Trifolium was part of Bradshaw’s teaching portfolio in genetics in the 1950s, and it is possible that this shared interest, along with Jones’ general interest in polymorphism and ecological genetics, were what resulted in the invitation.

Jones’ continued involvement in the EGG opened the door to others from the zoology and animal genetics community at Oxford in subsequent years. For example, David William Snow, an ornithologist (Ph.D. Oxford, 1953), first attended the EGG in 1965. Entomologist John R. G. Turner (Ph.D. Oxford, 1969, Lepidoptera), and zoologist Bryan Clarke (Ph.D. Oxford, 1962, Cepaea) attended in 1966. Shortly after attending the EGG Clarke went on to organize the animal-oriented “Population Genetics Group,” now known as PGG or PopGroup, modeled on the EGG meetings. Meanwhile, after finishing his Ph.D. in 1961, David Jones was hired as Assistant Lecturer at Birmingham. Dennis Wilkins joined the staff at Birmingham shortly therafter. This drew in more non-plant biologists from Birmingham, including Michael J. Lawrence (Ph.D. Aberystwyth, 1958, Drosophila) in 1965, and Michael J. Kearsay (Ph.D. Birmingham, 1965, Drosophila) and Giora Simchen (Ph.D. Birmingham, 1966, yeast) in 1966.

The EGG grew and diversified over the next several decades, drawing in participants from other research groups as well as providing a “home” for the students of its central participants. Despite Gregor’s concerns about the growth of their informal annual meeting, except on rare occasions the overall attendance remained at around thirty well into the 1970s.

Interest from David Henriques Valentine (1912–1987) in 1964 brought in a new crop of genecologists from the University of Durham, where Valentine was then Professor of Botany. Valentine became involved in genecological research in the 1940s, and most of his work focused on species of Viola, and later Primula. Although Valentine’s genecology retained much of the taxonomic hue that the Gregorian school had cast aside, his interest in abrupt versus gradual speciation was a point of intellectual overlap with Bradshaw and his colleagues. Like Gregor, Valentine was also interested in the problem of professionalization and institutionalization for genecology. Several of Valentine’s students, as well as his colleague Jack Crosby, were invited to the 1964 meeting and became regular EGG attendees.

Bradshaw’s students at Bangor made up a large part of the new recruits in the 1960s, ‘70s, and ‘80s. John Aston, Janis Antonovics, Glenys Crossley, Peter Gregory, Thomas McNeilly, and John Pusey all attended in 1964. As students from Bangor, Aberystwyth, and Edinburgh took posts at other British universities they not only continued to attend EGG meetings but also brought along their students and colleagues. For example, Antonovics worked as a post-doctoral researcher at the University of Stirling between 1966 and 1970, and brought along several of his students, including Henry Ford, Heather Dickinson, and Joseph Watson. Roy Snaydon, who went on to work at the University of Reading and further developed the work on mineral nutrition, drew in a steady stream of students as well.

Bradshaw’s move to Liverpool in 1968 substantially broadened the base of attendees, as his students and colleagues joined the group. Staff and students from Aberystwyth, the Welsh Plant Breeding Station, Scottish Plant Breeding Station (later Scottish Crop Research Institute), and other agricultural research centers and provincial colleges continued to make up a large proportion of the overall attendance well into the 1990s. Over time, the EGG also attracted visitors from the European continent and North America. Bernard Dommee, who studied intraspecific variation in reproduction in Calluna vulgaris, was responsible for bringing in French students from Montpellier, starting in the late 1970s. Pieter Kakes, a genecologist at the University of Amsterdam, attended the meetings between 1975 and 1991, and brought along several Dutch botany students during that period.

Since 1988 the EGG has been recognized as a “Special Interest Group” of the British Ecological Society. The EGG has continued to hold its Easter-time meeting, the last of which was held 30 March through 1 April 2015, at Edge Hill University in Liverpool.

4.5. Conclusion

One of the most striking features of the Ecological Genetics Group over the past sixty years has been its continuity with the original community of agricultural genecologists in Scotland and Wales that began meeting informally in 1956. A preliminary analysis indicates that a very large proportion of attendees over the past several years can trace their scientific “ancestry”—in terms of supervisor-student relationships—to either the early cohorts of attendees in the 1950s and 1960s. Most recent participants have been in the third or fourth “generation” of scientists since the group began meeting in 1956. The EGG thus provides a clear continuity from the Gregorian genecology of the 1930s and ‘40s to contemporary ecological genetics in Britain.

The brief history of the Ecological Genetics Group offered in this chapter is a starting-point for further investigations into the evolution of British ecological genetics and evolutionary population ecology. While this account has highlighted key turning points and some qualitative trends, it leaves an open field for investigating the changing contexts, concepts, and content of ecological genetics in Britain in subsequent decades. The attendance records housed at the National Library of Wales are impressively comprehensive, and present a unique and attractive opportunity to reconstruct the history of ecological genetics in even greater detail. But with such an opportunity comes many challenges, the most notable of which is the problem of scale. Although attendance at the EGG meetings only rarely exceeded 50 people, the cumulative number of attendees since 1956 exceeds one thousand individual scientists. Moreover, the volume of research literature produced by the average ecologist has increased substantially since the 1960s. An analysis that pays adequate attention to the conceptual, methodological, and contextual dimensions of the investigative activities in which those scientists were engaged requires tools and methodologies that can scale with the volume of materials on which it is deployed. To that end, I have begun to draw together techniques from computational linguistics and social network analysis that can be applied to the history of the EGG, and other similar associations.


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