R. H. Lock, 1879–1915

The posthumous fourth edition of Variation, Heredity, and Evolution (1916) carries a biographical note by “B. L.,” Lock’s widow Bella, formerly Bella Sidney Woolf (1877–1960), eldest daughter of Sidney Woolf, QC. Her brother Leonard married Virginia Stephen, the second daughter of Sir Leslie Stephen and better known today under her married name of Virginia Woolf. The details of Lock’s early life, given below, are based largely on this biographical note.

Robert Heath Lock was born on January 19, 1879 at Eton College where his father, the Reverend John Bascombe Lock, was a master. His mother Emily was the daughter of Edwin Baily of Cirencester. He was educated at Charterhouse School and entered Gonville and Caius College with an entrance exhibition on October 1 1898, by which time his father, a mathematician, had moved to Cambridge and then become bursar of the college. The son prospered in the Natural Sciences Tripos, attending Bateson’s lectures “On the practical study of evolution” and being placed in the first class in both part I (1900) and part II (Botany 1902). While still an undergraduate Lock accompanied Bateson abroad, who wrote to his wife Beatrice from Tours, France, in April 1902, commenting on Lock, “My little companion ... [who] knows no French of any kind, and when I say no French I mean it. ‘What is French for ‘to eat?’ ... German, he reads with fair ease. He has a prodigious memory for verse: knows most of Hamlet, Macbeth and Gilbert and Sullivan by heart, and can recite more ephemeral verses than ever I heard of” (Cock and Fordyce 2008, p. 290).

His Tripos results gained Lock the Frank Smart studentship of the college for two years, which he spent at the Royal Botanic Gardens at Peradeniya in Ceylon (Sri Lanka), where another Caian (as members of the college are called) who had also been a Frank Smart student, J. C. Willis, was director. During this time, Lock engaged in an extensive correspondence with Bateson. Described by Cock and Forsdyke (2008, p. 290), these authors remark “In principle, the correspondence was all that a free and open scientific correspondence should be.”

Returning to England, Lock was elected a Drosier fellow of the college on January 11, 1905, finding himself next in seniority to R. C. Punnett, who had been elected in 1901 (for Punnett, see the Perspectives by Edwards 2012, GENETICS, Vol. 192, No. 9). The interval of three years reflected not only their difference in age but the timing of their graduation, Lock entering Caius in the term after Punnett graduated. Though they did not overlap then, they did in the academic year 1901–1902 when Punnett was junior fellow and Lock a fourth-year undergraduate. Very probably their friendship started at that time. On p. 43 of Variation, Heredity, and Evolution (1906) Lock refers to “my friend Mr. R. C. Punnett,” and there is much evidence of friendly banter between them later on in the Fellows’ Betting Book. Punnett had joined Bateson as a collaborator at the start of 1904, so when Lock returned at the end of the year he would soon witness their famous discovery of “partial coupling” in the sweet pea (later “linkage”; for details see Edwards 2012). Lock was also appointed curator of the University Herbarium in the Botany School.

As bachelor fellows, Punnett and Lock both lived in college, and will have done much of their writing in their college rooms. These were in the 16th century buildings of Caius Court (M2 and K3, respectively). The preface of Punnett’s book Mendelism is dated May 1905, so that for the first months of 1905 he was writing Mendelism and Lock was writing Variation, Heredity, and Evolution within a few yards of each other and in daily contact as resident college fellows. There was surely an understanding between them that they were engaged in quite different tasks, Punnett writing a short essay on just Mendelism, published in Cambridge, and Lock a survey of recent progress in genetics and evolution published in London by John Murray, Darwin’s publisher. “The idea of writing this little book,” he wrote in his preface (p. vii), “occurred to me while reading Mr. W. C. D. Whetham’s volume on The Recent Development of Physical Science.” “I found the story of the modern progress of physics so interesting as to encourage the belief that a similar account of the subjects with which I was myself more particularly familiar might prove of a like interest to other people.”

Lock held his fellowship for six years, though not resident for the last two-and-a-half. In August 1906 he attended the Third International Conference on Hybridization and Plant Breeding (later styled “Genetics”) in London with Bateson, Punnett, R. H. Biffen (another former Frank Smart student) and T. B. Wood (also a Caian). The “Newnham College Mendelians” (Richmond 2001) were also well represented, so Bateson, who was president of the congress, was supported by a substantial Cambridge contingent of his colleagues, helpers, and former students, mostly from Caius and Newnham. Further information about the “Cambridge Mendelians” is given by Richmond (2006a) and also in her account of the 1909 Darwin Celebration by the University (Richmond 2006b).

In the spring of 1908 Lock returned to Peradeniya as assistant director of the gardens under Willis. “We understand,” says the Caius College magazine, “that he is to have complete charge of the Economic Development of the Gardens.” There he met his Trinity College contemporary Leonard Woolf and, on the voyage out, Leonard’s sister Bella who was visiting him. At the end of June 1909, Punnett also traveled to Ceylon (to study mimicry) on a three-month trip, visiting Lock. No doubt their conversation naturally turned to the appointment of a first director of the newly founded John Innes Horticultural Institute then under consideration. Lock was one of three who had been short listed from among 30 applicants (Cock and Forsdyke 2008, p. 382) but the council deemed none satisfactory and ultimately, in November, Bateson, who had not applied, was offered the directorship. He accepted. There are further letters from Lock to Bateson during this period.

In 1910 Lock returned to England on short leave and married Bella on July 7. Two days previously he had dined in college and settled a bet with Mr. Duncan Jones, whom he had bet on March 11, 1906 “will be engaged to be married before him.” He had won, and Mr. Duncan Jones paid (with the customary wine after dinner). Returning to Ceylon with Bella, Lock conducted experiments on rubber tapping and rice breeding, but was also heavily engaged in administrative work, especially as acting director when Willis was on leave.

In 1911 Willis moved to Rio de Janiero as director of the botanic gardens there, but Lock, not wishing to succeed him in Ceylon, returned to England in 1912, spending the first few months after his return writing a second book Rubber and Rubber Planting (1913). In June 1912 Bateson was invited to return to Cambridge for the newly created Arthur Balfour Professorship of Genetics, but he declined it and suggested Punnett (see Edwards 2012). Had the professorship been filled by a board of electors rather than Arthur Balfour and the prime minister, Lock might have stood a chance, but Punnett was effectively the sitting tenant and the possibility might never have crossed Lock’s mind. Only with hindsight can we see what a good choice he might have been.

He then took up a post of inspector at the Board of Agriculture and Fisheries. When the First World War broke out in 1914 he sought to join the forces, but was asked instead to add the chairmanship of a Vegetable Drying and Fruit Preserving Committee to his existing duties, a government body charged with furthering work on food preservation as a wartime necessity.

Struck down by a severe bout of influenza in February 1915 from which he never fully recovered, and overworked with his wartime duties, Lock died from heart failure at Eastbourne on June 26, 1915 at the age of 36. He is buried in the Ascension Burial Ground, Cambridge, where also lie his parents John and Emily. His widow, Bella, went on, as Bella Woolf, to be a successful author.

Robert and Bella Lock had no children, but Robert’s three brothers were also educated at Caius College, and two of them started family lines that have ensured four and five generations of Locks at Caius, a tradition that had started with John Bascombe in 1867.

The influence of Variation, Heredity, and Evolution on the Columbia school

The 2000 reprint of A. H. Sturtevant’s A History of Genetics (1965) says, on the back cover:

In the small “Fly Room” at Columbia University, T. H. Morgan and his students, A. H. Sturtevant, C. B. Bridges, and H. J. Muller, carried out the work that laid the foundations of modern, chromosomal genetics. The excitement of those times, when the whole field of genetics was being created, is captured by this book, written by one of those present at the beginning. ... it is worth remembering that the world’s first genetic map was created in 1913 by A. H. Sturtevant .. .

Appendix A in the book gives a chronology, recording for 1906 “Lock: suggested the relation between linkage and the exchange of parts between homologous chromosomes” (2000, p. 137). In the text, Sturtevant wrote (2000, p. 43) “The possibility that linkage might result from genes lying on the same chromosome had been suggested by Lock in 1906, in his elaboration of de Vries’ idea that exchange of material between homologous chromosomes could account for independent segregation.” The reference is to the American edition of Variation, Heredity, and Evolution (Dutton, New York 1906; identical to the British one) and Chapter IX Recent Cytology. Lock was, of course, familiar with the recent discovery of partial coupling of characters in the sweet pea found by Bateson and Punnett in 1904 and 1905, later called linkage, and on p. 187 he briefly alludes to it. Then on p. 252, after describing de Vries’s (erroneous) hypothesis for independent segregation, Lock writes:

In cases where the phenomenon of correlation or coupling has been observed we must suppose that there is some mechanism which causes the representative particles of the respective characters concerned to remain in company during the process by which the other allelomorphs are being reassorted between the chromosomes. Of this process of coupling the cytologists have not yet been able to observe any visible indication in the behaviour of the chromosomes, any more than they can really see the redistribution of the supposed factors carried by the chromosomes. But apart from this it must be allowed that the facts of the experiment and of microscopic observation fit in with one another in a remarkable way, and that the Mendelian theory throws considerable light on the minute features of cell anatomy.

E. B. Wilson, professor of zoology at Columbia, used Lock’s book for his course “Heredity, variation and the chromosomes.” The minds of the geneticists of the Columbia school, for whom Variation, Heredity, and Evolution had been their textbook, were thus exposed to the germ of an idea about a chromosomal explanation for linkage.

H. J. Muller had taken general biology in his freshman year, much of it taught by J. H. MacGregor, and

After his first successful year, Muller spent the summer as a hotel clerk in New Jersey. He knew that in the fall he would be taking Wilson’s course on the cell, and he knew from MacGregor that the text for it would be R. H. Lock’s Recent Progress in the Study of Variation, Heredity, and Evolution (1906). Muller read the book while he hid, out of sight, under the stairwell of the hotel when the bell wasn’t ringing, and he again experienced the electrifying thrill he had when he first read Mendeleev’s periodic tables. Here was a system that brought evolution, cell biology, and plant and animal breeding into one common scheme (Carlson 1981 p. 27).

In the words of Crew (1969), it was Lock’s book “that did so much to attract Muller to genetics.” Carlson (2013) attended Muller’s course “Mutation and the gene” in 1955 and prints a page of his lecture notes recording: “1906. R. H. Lock. Recent progress in the study of variation and evolution (sic)—The completely modern viewpoint for its time.” Muller (1943) himself wrote that in 1908 he took E. B. Wilson’s “thrilling one semester course on heredity and the chromosomes, variation and evolution. In this course, the text chosen by Wilson was Lock’s extraordinary book of 1906—too far “ahead of its time” to be sufficiently remembered now—which, with less caution and fewer qualifications than employed by Wilson himself, advocated the sufficiency of Mendelism, multiple factors, the chromosome theory (including exchange of linearly arranged genes during parasynapsis, after de Vries), and the natural selection of mutations, as the basis of all heredity and evolution.”

Muller (1916) did not mention Lock in his Ph.D. thesis “The mechanism of crossing-over,” and neither had Morgan (1911) when he advanced his hypothesis that “partial coupling—linkage—is a reflection of ‘associations of factors’ that are located near together in the chromosomes.” By then Morgan was able to rely on the cytological observations of Janssens (in 1909; see the Perspectives by Koszul et al. 2012, in GENETICS, Vol. 191, Num. 2). But when all four of the Columbia school combined to publish The Mechanism of Mendelian Heredity (Morgan et al. 1915) in Chapter I they wrote “In the same year [1906] ... Lock called attention to the possible relation between the chromosome hypothesis and linkage” (p. 5) and did not there mention Janssens. Consciously or subconsciously, they had absorbed Lock’s point. But memory of Lock was fading: they gave him no initials, either there or in the index, and his book is not in the bibliography. By the time Whitehouse (1965) wrote his Toward an Understanding of the Mechanism of Heredity with its substantial historical content, Lock had disappeared almost completely, squeezed out between de Vries and Morgan rather than perceptively linking the two. Besides Sturtevant (1965) perhaps only Cock (1983), in his study of Bateson’s reaction to the chromosome theory, remembered: “Lock, in particular, gave vigorous support to the chromosome theory as early as 1906.” Indeed, in his second edition (1909, p. 201) when he added a report of the segregation ratios proposed by Bateson and Punnett on their “reduplication hypothesis” (see Edwards 2012), Lock did not mention the hypothesis by name. His account of their idea evinces skepticism and concludes “There is good reason for hoping that we shall soon be able to describe the facts of correlation [i.e. partial coupling] in terms as definite as those which can now be applied to the phenomena of segregation.” Janssens was just round the corner.

Wilson attended the 1909 Darwin Centenary Celebrations in Cambridge, staying with the Batesons (Cock and Forsdyke 2008, p. 321) and receiving an honorary degree from the university. It seems probable that Lock was in Ceylon at the time and missed the opportunity to meet the most influential user of his book.

From a historical point of view, it was unfortunate but understandable that when Doncaster inserted new material about linkage in Lock’s fourth edition (pp. 200–202, 269–271) he removed Lock’s suggestion for the phenomenon of coupling, quoted above, and replaced it with a description of Morgan’s work.

R. A. Fisher’s Sources of Information About Genetics, 1909–1911

I first came to Cambridge in 1909, the year in which the centenary of Darwin’s birth and the jubilee of the publication of The Origin of Species were being celebrated. The new school of geneticists using Mendel’s laws of inheritance was full of activity and confidence, and the shops were full of books good and bad from which one could see how completely many writers of this movement believed that Darwin’s position had been discredited (Fisher 1947).

“The fiftieth anniversary of the publication of The Origin of Species was being celebrated, apart from other things, by the publication of Bateson’s book Mendel’s Principles of Inheritance” (Fisher 1959; actually Mendel’s Principles of Heredity). Fisher bought the book. He also received as a college prize the book of essays Darwin and Modern Science (Seward 1909) published by Cambridge University Press as part of the Darwin Celebrations (“a remarkable collection of able essays,” Fisher was to say in 1959).

This is all the definite information there is about Fisher’s reading in genetics, and includes Mendel’s paper, which was given in translation in Mendel’s Principles of Heredity. But among the “books good and bad” that he encountered then, or soon after, we must surely include Punnett’s Mendelism in one of its first two editions (1905, 1907). Punnett, as we have seen, was a resident fellow of Caius, and Fisher certainly came to know him during his undergraduate days (Mazumdar 1992, p. 99). However, Fisher may not have met Lock. The only time when they might both have been in college was when Lock dined there on July 5, 1910 just before his wedding (see above). But Lock had presented his second edition (1909) to the college library in July, and even if Punnett had not drawn attention to it Fisher would surely have come across it. An undergraduate who notes “books good and bad” on a subject of consuming interest to him in the Cambridge bookshops is hardly likely to ignore his own college library. Be that as it may, Fisher never referred to Lock’s book in his writings, but then he never referred to Punnett’s either.

There is some evidence that Fisher used this particular copy of Lock’s second edition. The book has been moderately well used, and of all the potential Caius undergraduate readers of it, Fisher seems the most likely. But more direct evidence appears on p. 87. Lock there reproduces a diagram from Wallace’s Darwinism (1889, p. 58). Neither axis is labeled, and in the Caius copy someone has added “No of birds” for the x-axis and someone else (the hand is different) an indistinct “length of wing” or some such for the y-axis. As one who is familiar with Fisher’s spidery handwriting, often very small because of his extreme short sightedness, I believe “No of birds” (Figure 2) is in his hand. I can cite other instances of Fisher having annotated library books in Cambridge, including one other in the Caius library. There is also of course the possibility that Fisher remembered the diagram from Wallace, whose Darwinism he had read by 1915 (Fisher 1915).

• Download figure
• Open in new tab
• Download powerpoint

Figure 2

From the 2nd (1909) edition of Recent Progress in the Study of Variation, Heredity, and Evolution, annotated “No of birds” in a hand probably R. A. Fisher’s.

By the time Fisher gave his talk “Mendelism and Biometry” on November 10, 1911 (printed in Bennett 1983, p. 51) he was to varying degrees familiar with the work of Weismann, de Vries, and Johannsen. His first example of Mendelism is the Andalusian fowl, which he could have got from any of the three books of Bateson, Punnett, and Lock. He is familiar with the work of Pearson, and in particular with the problem of parent–offspring correlation and the differing views of Pearson and of Yule (whom he does not mention by name). The only source among the books so far mentioned seems to be Lock (1909), Chapter VIII Mendelism (continued) with a section “Mendelism and Biometry”—the very title of Fisher’s talk—“researches of Yule and Pearson” (p. 227). Fisher mentions the good fit of Mendel’s data to his hypothesis, revealing an apparent familiarity with Weldon (1902), a paper in Biometrika (see Edwards 1986, Magnello 1998). This seems to be a direct result of perusing Biometrika itself. He has read Galton’s Hereditary Genius (1869). Lock (1909) devoted a whole new Chapter X Eugenics (p. 282) to Galton and his work.

In addition to this evidence for Lock’s book in its second (1909) edition having been an important source for Fisher, it contains a number of quite suggestive remarks bearing on some of Fisher’s later work. We have already noted the opening of the topic of the correlation between relatives, which matured into Fisher (1918). On the subject of classification Lock wrote in his introduction (p. 10)

... though at first sight it may appear almost paradoxical, ... it is quite possible for groups to be perfectly distinct, although individual members of them may have deviated so far, each from its proper type, as to render impossible the task of deciding from their appearance which group any of these individuals belong to.

In Fisher’s (1925, p. 37) Statistical Methods for Research Workers we find the same thought:

When a large number of individuals are measured in respect of physical dimensions, weight, colour, density, etc., it is possible to describe with some accuracy the population of which our experience may be regarded as a sample. By this means it may be possible to distinguish it from other populations differing in their genetic origin, or in environmental circumstances. Thus local races may be very different as populations, although individuals may overlap in all characters;

Furthermore, this follows two pages after a figure (his figure 3) constructed on the same strange principle as Wallace’s diagram referred to above, which was reproduced by Lock (p. 87). As Fisher was still a fellow of Caius when writing Statistical Methods he would have had ready access to Lock in the library.

A remarkably prescient remark is made by Lock on p. 99, to the effect that the standard deviation “happens to be determinable with greater accuracy from an actual series of variates” than the mean quartile, and in a footnote he gives the root-mean-square derivation. Fisher (1920) explored the whole question in intricate detail and proved that The whole of the information respecting σ [the standard deviation of a normal distribution], which a sample provides, is summed up in the value of σ₂ [the sample standard deviation calculated from the mean-square deviation from the mean]. This is the first appearance of the notion of statistical information and led to the concept of sufficient statistics. Fisher (1950) notes that the paper arose from an examination of a statement by A. S. Eddington, but that does not preclude an initial stimulus from his reading of Lock.

Finally, when “quasi-linkage,” or association between Mendelian factors on different chromosomes, was first discussed (Michie 1953, Wallace 1953) Fisher suggested the word “affinity” to describe it (Wallace 1958). Lock (1906, p. 185, 1909 p. 197) had written of “the existence of some kind of affinity occurring in the same individual between allelomorphs which belong to distinct pairs.”

But perhaps equal to all these pieces of evidence is the unique feature of Lock’s book that it brought together (to quote from its chapter headings) evolution, the theory of natural selection, biometry, the theory of mutation, Mendelism, cytology, and eugenics, all in a single volume. Nowhere else could the young Fisher have found such a guide to the subjects that fascinated him over and above his student work for the Mathematical Tripos. He had to look no further than his college library for the book, which would lay out the ground that he was to till so fruitfully in The Genetical Theory of Natural Selection (1930). Lock’s Recent Progress in the Study of Variation, Heredity, and Evolution was, I suggest, as much an inspiration for R. A. Fisher as it had been for H. J. Muller.