60 Sahotra Sarkar
had remained unpublished.^20 However, unlike Norton, Haldane remained focused
on rates of change. The conclusion was disappointing: the temporal dynamics of
change remained similar to the situation where generations did not overlap.
At this point Haldane seems to have given up all hope of escaping from the
impasse that autosomal recessive factors cannot easily spread through a population
without strong selection on recessives. Of the different schemes he had tried, only
self-fertilization and inbreeding truly helped the recessives. Observing that most
mutations were recessive to the normal type, in Part V, he turned to mutation
as the source of recessive alleles. Fisher [1922] had analyzed the interaction of
mutation with selection earlier and Haldane [1927b] also reached the conclusion
that if the mutations are very rare, their survival depends almost entirely on
stochastic factors. Haldane managed to calculate the probability of survival of
a mutation. Assuming random mating in a diploid population, if a mutation is
dominant, and it confers a slight selective advantage,k, the probability of its
(^20) Norton’s work was eventually published — see Norton [1928]. Norton died at the age of
fifty. Years later, when R. F. Harrod [1951], who had been with Haldane at New College,
Oxford, published a biography of John Maynard Keynes, he merely mentioned] Norton’s work
as an “application of probability theorems to certain problems in genetics” that had never been
published (p. 188). Norton’s sister, Betty, complained to Haldane (B. Norton to Haldane,
January 31, 1951, Box 19, UCL) who wrote to Harrod, pointing out that Norton’s work had
eventually been published (Haldane to R. Harrod, February 2, 1951, Box 19, UCL) and:
that the paper occupies some 45 pages, and that it gives the only rigorous proof
of a great many notions which we are apt to take for granted in genetical work,
but which in fact are only true if certain conditions are fulfilled. Besides this, it
gives the methods for dealing with fairly complicated situations which sometimes
arise. I suspect that its importance will grow as the application of mathematics to
genetics advances....
May I hope that in future editions you will do justice to a man who, though
through ill health he did not achieve as much as was hoped, has certainly made a
fundamental contribution to applied mathematics.
Harrod willingly, and apologetically acquiesced, submitting an emendation for future editions
for Haldane’s approval (R. Harrod to Haldane, February 3, 1951, Box 19, UCL).
Haldane also replied to Betty Norton (Haldane to B. Norton, February 2, 1951, Box 19, UCL).
Norton’s work, he claimed,
is... beyond doubt... fundamental... and I refer to it both in lectures and in
a book which I am writing on the subject. In my own papers on the subject I had
to refer to his unpublished work, since I had independently obtained some of his
results, and of course knew his work....
I have always felt that he may have thought that I ‘jumped his claim’, on the other
hand the publication of his work was, if I remember, delayed for about seventeen
years and as we only overlapped to a relatively small extent, I felt justified in going
ahead with reference to ‘the important unpublished work of H. T. Norton’....
I very much hope that your brother did not feel that I had wronged him in any
way, but you will realise that the position was difficult. I certainly feel that full
justice should be done to his memory.
B. Norton wrote back reassuring Haldane on the last point (B. Norton to Haldane, February
3, 1951, Box 19, UCL).