Fifty glorious years of PGG: the first ten percent

Welcome to No Man’s Time: too recent for history, too early to be memory. The decade or so before you were born. For many of you this lost period will be the late 1960s, when the Population Genetics Group was just beginning. There are now not many people who can remember, and archived documentation is fragmentary. I have compiled this brief account from my own memories and diaries, from my wife Sandra’s diaries, and from the recollections of Laurence Cook, John Currey, Peter O’Donald and Mark Williamson (also Charlotte Williamson’s diaries). Darren Obbard has helped me from his accumulating archive. 

The PGG originated in the burst of active research on Cepaea and related snails during the 1960s, and in a conference hosted by John Currey at the new and ambitious biology department at York. Most of the participants, including AJ Cain, BC Clarke, M Lamotte, H Wolda, LM Cook, CB Goodhart and many of Cain’s current and former graduate students, obviously were working directly with these species, but there was also a sprinkling of people with a wider interest who contributed by way of theory: J Antonovics, P O’Donald, JRG Turner and MH Williamson. Even plants were represented, by Tony Bradshaw (MW) and—as snail food—Philip Grime. 

The York meeting was a great success, and was followed by two further specialised Cepaea meetings: in Paris 1971 hosted by Maxime Lamotte, and at Groningen hosted by Henk Wolda, date so far unknown (MW, LC). Meanwhile the opportunity to build on the widened field of interest at the York meeting was seized: a more general population genetics meeting was mooted for the next year.  An extra motive was the frustration felt by many population geneticists at what they saw as the domination of Genetical Society meetings by work on intermediary fungal metabolism. My guess is that the prime mover was Bryan Clarke. The first meeting of PGG sensu stricto was therefore held in the following winter. Birmingham was selected chiefly because it was considered to approximate the least squared distance point from most of the workers in the field (tough luck Aberdeen). 

After this the meeting became annual, around Christmas, but variably before or after the change of year (hence the 1970/71 etc designations: table 1, for the first five years plus the forerunner), and peripatetic, eventually by invitation from an appropriately interested department in the host university. 

Table1. The first five PGG meetings, with the preceding Cepaea conference 
Meeting no            Season                        Date                        Place 
0                             1966/67                    3-7 Jan 67              York 
1                             1967/68                   10-11 Jan 68          Birmingham 
2                             1968/69                   16-18 Dec 68         Edinburgh 
3                             1969/70                   16-18 Dec 69         Liverpool 
4                             1970/71                   16-18 Dec 70         London [?UCL] 
5                             1971/72                   17-20 Dec 71         Bangor 

This truly was “science from the People”. There was no formal organisation, no officers, no president; but then of course, no archives. The full lists of participants may be lost for ever. I have clear recollections of some people at Birmingham, although of course there may be some false memories transferred from other meetings; Mark Williamson and Laurence Cook have garnered some more. The combined, reasonably certain roster so far is Arthur Cain, John Maynard Smith, Alan Roberston, Wilhelm Scharloo, Kennedy McWhirter, Charles Goodhart, Robert Creed, Michael Kearsey, Geoff Oxford, John Jinks, John Thoday, Terry Crawford [-Sidebotham], and probably David Jones; and of course the informants themselves! Notable absences included a number of “big names” who probably still saw their natural milieu as the Royal Society and the Genetical Society: as far as I know Henry (EB) Ford, Philip Sheppard (even when the meeting was hosted in Liverpool), and Kenneth Mather never (well, hardly ever—there is some disagreement between observers) attended PGG. Arthur Cain, who never saw himself as a geneticist, became a stalwart, as did Maynard Smith, Alan Robertson and John Thoday. 

The population size of the meetings was, by modern standards, modest. The delegates to the York Cepaea conference numbered only 24 [CW]. There was no call for parallel sessions: everybody could hear all the papers. No full programme for any of the meetings has yet been recovered, but the sessions for the Liverpool and London meetings were noted in my diary:

Table 2 The timetable of sessions for the Liverpool PGG meeting of 1969/70 
1969      am                                                                 pm 
16 Dec   Genetical architecture                      Genotype x environment 
17 Dec   Competing species                              Strong selection pressures 
18 Dec   Enzyme polymorphism                     Enzyme polymorphism 

Table 3 The timetable of sessions for the London PGG meeting of 1970/71 
1970      am                                                                 pm 
16 Dec   [No session]                                              Coadaptation 
17 Dec   Measurement of selection              Chromosomes 
18 Dec   Congenital malformations             Protein evolution 

The subjects under discussion can be surmised from the history of population genetics for the period: see B Charlesworth & D Charlesworth Heredity advance online publication, 27 July 2016; doi:10.1038/hdy.2016.55 for the broad view; as usual with history it felt somewhat different when it was happening. Memories of a few of the delivered papers have been recovered. At Birmingham [LC] the scheduled plenary lecture by Ken-Ichi Kojima was prevented by atrocious winter weather in the northern Atlantic, and Alan Robertson stepped in and gave a seminal introduction on the subject of heterozygosity and genetic load, reviewing work by Sved, Reed & Bodmer, King and Milkman. Five years later in Bangor (LC, JT) the same controversies were still current: the PGG was privileged to hear from Dick Lewontin, who was at that time preparing his “book on enzyme polymorphism” (to appear in 1974 as The genetic basis of evolutionary change) on the broad directions and challenges for population genetics as he saw them, which were vastly at variance with the overall culture of Cepaea genetics that had founded the PGG. I recall him saying to Maynard Smith over lunch, that he found it hard to credit that all selection coefficients were large. 

One further lecture at Birmingham remains firmly in my memory. Kennedy McWhirter was an unusually talented polymath: biologist, qualified barrister, linguist/translator, psychic (with the ability to communicate with ghosts); incidentally he was also the elder brother of the McWhirter (Guinness Book of Records) twins. He had worked with Ford on Maniola jurtina, and at this time held a research fellowship in Ford’s lab. His paper was on the theory of endocyclic selection, uniquely developed by Kennedy: he was suggesting that interesting results in the dynamics of polymorphism would arise if the direction of selection varied during the course of one life cycle (e.g. in favour of aa genotypes in juveniles, but against them in adults). The theory never came to adequate fruition and as far as I know was never published, but Kennedy gave a spirited presentation. For reasons that are now hard to recall, it was all extremely funny. This was unintended, but Kennedy took the laughter from the audience in somewhat puzzled good humour. It was not that we were laughing at him, or at his ideas: it was simply that the way he presented things had genuine comedy. By the end the whole lecture theatre was rocking, with the exception of Arthur Cain, who being more of a gentleman than the rest of us, was sitting with his handkerchief stuffed into his mouth, and turning bright pink. 

My thanks to the other named informants and to Brian and Deborah Charlesworth. Corrections, further memories, “I was there…”, “That was me” etc will be very much welcomed [j.r.g.turner@leeds.ac.uk; darren.obbard@ed.ac.uk]. 
© John RG Turner  August 2016