The Polemic Use of the Rediscovery by Ronald A. Fisher Lyrics
1. THE POLEMIC USE OF THE REDISCOVERY.THE tale of Mendel's discovery of the laws of inheritance, and of the sensational rediscovery of his work thirty-four years after its publication and sixteen after Mendel's death, has become traditional in the teaching of biology. A careful scrutiny can but strengthen the truth in such a tradition, and may serve to free it from such accretions as prejudice or hasty judgement may have woven into the story. Few statements are so free from these errors as that which I quote from H. F. Roberts' valuable book Plant Hybridisation before Mendel (p. 286):"The year 1900 marks the beginning of the modern period in the study of heredity. Despite the fact that there had been some development of the idea that a living organism is an aggregation of characters in the form of units of some description, there had been no attempts to ascertain by experiment, how such supposed units might behave in the offspring of a cross. In the year above mentioned the papers of Gregor Mendel came to light, being quoted almost simultaneously in the scientific contributions of three European botanists, De Vries in Holland, Correns in Germany, and Von Tschermak in Austria. Of Mendel's two papers, the important one in this connection, entitled ' Experiments in Plant Hybridization ', was read at the meetings of the Natural History Society of Brünn in Bohemia (Czecho-Slovakia) at the sessions of February 8 and March 8, 1865. This paper had passed entirely unnoticed by the scientific circles of Europe, although it appeared in 1866 in the Transactions of the Society. From its publication until 1900, Mendel's paper appears to have been completely overlooked, except for the citations in Focke's ' Pflanzenmischlinge', and the single citation of Hoffmann, elsewhere referred to."When the History of Science is taken seriously the number of enquiries which such a story suggests is somewhat formidable. We want to know first : What did Mendel discover? How did he discover it? And what did he think he had discovered? Next, what was the relevance of his discoveries to the science of his time, and what was its reaction to them? In the case of Mendel these last questions must be duplicated, for we are concerned not only with the period following the reading of his principal paper in 1865, but with that following the widespread publicity it received in 1900. This will be considered first.
Seeing how often it is taken for granted that all clouds were cleared away at the rediscovery in 1900, it is singularly difficult to ascertain exactly how Mendel's experiments were conducted and, indeed, what experiments he carried out. We have, of course, his paper, principally devoted to garden peas, entitled "Versuche über Pflanzenhybriden", printed in the transactions of the Natural History Society of Brünn, in Bohemia, in 1866, and reprinted in 1910. In 1901 it was also twice reprinted, in Flora, and in Ostwald's Klassiker der exakten Wissenschaften (No. 121). A valuable English translation, prepared for the Royal Horticultural Society, was published in 1901, and reprinted with modifications by Bateson on several occasions. I shall refer to its appearance in Bateson's book Mendel's Principles of Heredity (Cambridge, 1909).
It cannot be denied that Bateson's interest in the rediscovery was that of a zealous partisan. We must ascribe to him two elements in the legend which seem to have no other foundation : (1) The belief that Darwin's influence was responsible for the neglect of Mendel's work, and of all experimentation with similar aims ; and (2) the belief that Mendel was hostile to Darwin's theories, and fancied that his work controverted them. On the first point we may note a paragraph from Bateson's preface (p. 2) :"While the experimental study of the species problem was in full activity the Darwinian writings appeared. Evolution, from being an unsupported hypothesis, was at length shown to be so plainly deducible from ordinary experience that the reality of the process was no longer doubtful. With the triumph of the evolutionary idea curiosity as to the significance of specific differences was satisfied. The Origin was published in 1859. During the following decade, while the new views were on trial, the experimental breeders continued their work, but before 1870 the field was practically abandoned." It should be noted that Bateson here identifies experimental breeding with the hybridization of species. He ignores the fact that Mendel's advance over his predecessors was due to crossing closely allied varieties, not different species, which, as Mendel actually recognized, would differ in a large number of different factors. It is a consequence of Darwin's doctrine that the nature of the hereditary differences between species can be elucidated by studying heredity in crosses within species. So far were the new evolutionary ideas from discouraging experimental breeding that Darwin, himself, apart from other work, devoted eleven years prior to 1876 to the great series of experiments of which his book on The Effects of Cross- and Self-fertilisation in the Vegetable Kingdom is a report. Had his example been followed there would have been no such lull as succeeded his death. Like Mendel's experiments a few years earlier they seemed to lead to nothing more at the time. Today, in the light of genetic analysis, we can go further towards appreciating their significance.
Bateson's eagerness to exploit Mendel's discovery in his feud with the theory of Natural Selection shows itself again in his misrepresentation of Mendel's own views. Although he was in fact not among those responsible for the rediscovery, his advocacy created so strong an impression that he is still sometimes so described. In the biographical notice which Bateson prefixes to his reprint of Mendel's papers he writes (p. 311) :"With the views of Darwin which were at that time coming into prominence Mendel did not find himself in full agreement, and he embarked on his experiments with peas, which as we know he continued for eight years." The suggestion that Mendel was prompted by disagreement with Darwin's views to undertake his experiments is easily disproved. Mendel's experiments cannot have commenced later than 1857. Darwin's views on evolution were known only to a few friends prior to the papers which he communicated, jointly with Wallace, to the Linnean Society in 1858. That Mendel had heard of Darwin, as a geologist or an explorer, at the time his experiments with peas were commenced is, indeed, possible. More probably he knew nothing of Darwin's existence, and certainly nothing of the theory of Natural Selection, at this date. When, in 1865, Mendel reported his experiments, the situation had doubtless changed. Mendel now recognizes that the study of inheritance has a special importance in relation to evolutionary theory. He alludes to the subject, in his introductory remarks, in words which suggest not doubts, but rather a simple acceptance of the theory of evolution (p. 318) :"It requires indeed some courage to undertake a labour of such far-reaching extent; this appears, however, to be the only right way by which we can finally reach the solution of a question the importance of which cannot be overestimated in connection with the history of the evolution of organic forms," In this paper the only other mention of evolution occurs in the
concluding remarks, in which the results and opinions of Gartner are discussed. It will be seen that Mendel expressly dissociates himself from Gartner's opposition to evolution, pointing out on the other hand that Gartner's own results are easily explained by the Mendelian theory of factors (p. 361): "Gärtner by the results of these transformation experiments was led to oppose the opinion of those naturalists who dispute the stability of plant species and believe in a continuous evolution of vegetation. He perceives in the complete transformation of one species into another an indubitable proof that species are fixed within limits which they cannot change. Although this opinion cannot be unconditionally accepted we find on the other hand in Gartner's experiments a noteworthy confirmation of that supposition regarding variability of cultivated plants which has already been expressed." It is seen from these, the only two allusions to evolution in Mendel's paper, that he did not regard his work as a direct contribution to that subject. What he does claim for the laws of inheritance he established is that they make sense of many of the results of the hybridists, and that they form a necessary basis for the understanding of the evolutionary process. On this point he shows himself fully aware of the importance of what he had done. Had he considered that his results were in any degree antagonistic to the theory of selection it would have been easy for him to say this also.
Seeing how often it is taken for granted that all clouds were cleared away at the rediscovery in 1900, it is singularly difficult to ascertain exactly how Mendel's experiments were conducted and, indeed, what experiments he carried out. We have, of course, his paper, principally devoted to garden peas, entitled "Versuche über Pflanzenhybriden", printed in the transactions of the Natural History Society of Brünn, in Bohemia, in 1866, and reprinted in 1910. In 1901 it was also twice reprinted, in Flora, and in Ostwald's Klassiker der exakten Wissenschaften (No. 121). A valuable English translation, prepared for the Royal Horticultural Society, was published in 1901, and reprinted with modifications by Bateson on several occasions. I shall refer to its appearance in Bateson's book Mendel's Principles of Heredity (Cambridge, 1909).
It cannot be denied that Bateson's interest in the rediscovery was that of a zealous partisan. We must ascribe to him two elements in the legend which seem to have no other foundation : (1) The belief that Darwin's influence was responsible for the neglect of Mendel's work, and of all experimentation with similar aims ; and (2) the belief that Mendel was hostile to Darwin's theories, and fancied that his work controverted them. On the first point we may note a paragraph from Bateson's preface (p. 2) :"While the experimental study of the species problem was in full activity the Darwinian writings appeared. Evolution, from being an unsupported hypothesis, was at length shown to be so plainly deducible from ordinary experience that the reality of the process was no longer doubtful. With the triumph of the evolutionary idea curiosity as to the significance of specific differences was satisfied. The Origin was published in 1859. During the following decade, while the new views were on trial, the experimental breeders continued their work, but before 1870 the field was practically abandoned." It should be noted that Bateson here identifies experimental breeding with the hybridization of species. He ignores the fact that Mendel's advance over his predecessors was due to crossing closely allied varieties, not different species, which, as Mendel actually recognized, would differ in a large number of different factors. It is a consequence of Darwin's doctrine that the nature of the hereditary differences between species can be elucidated by studying heredity in crosses within species. So far were the new evolutionary ideas from discouraging experimental breeding that Darwin, himself, apart from other work, devoted eleven years prior to 1876 to the great series of experiments of which his book on The Effects of Cross- and Self-fertilisation in the Vegetable Kingdom is a report. Had his example been followed there would have been no such lull as succeeded his death. Like Mendel's experiments a few years earlier they seemed to lead to nothing more at the time. Today, in the light of genetic analysis, we can go further towards appreciating their significance.
Bateson's eagerness to exploit Mendel's discovery in his feud with the theory of Natural Selection shows itself again in his misrepresentation of Mendel's own views. Although he was in fact not among those responsible for the rediscovery, his advocacy created so strong an impression that he is still sometimes so described. In the biographical notice which Bateson prefixes to his reprint of Mendel's papers he writes (p. 311) :"With the views of Darwin which were at that time coming into prominence Mendel did not find himself in full agreement, and he embarked on his experiments with peas, which as we know he continued for eight years." The suggestion that Mendel was prompted by disagreement with Darwin's views to undertake his experiments is easily disproved. Mendel's experiments cannot have commenced later than 1857. Darwin's views on evolution were known only to a few friends prior to the papers which he communicated, jointly with Wallace, to the Linnean Society in 1858. That Mendel had heard of Darwin, as a geologist or an explorer, at the time his experiments with peas were commenced is, indeed, possible. More probably he knew nothing of Darwin's existence, and certainly nothing of the theory of Natural Selection, at this date. When, in 1865, Mendel reported his experiments, the situation had doubtless changed. Mendel now recognizes that the study of inheritance has a special importance in relation to evolutionary theory. He alludes to the subject, in his introductory remarks, in words which suggest not doubts, but rather a simple acceptance of the theory of evolution (p. 318) :"It requires indeed some courage to undertake a labour of such far-reaching extent; this appears, however, to be the only right way by which we can finally reach the solution of a question the importance of which cannot be overestimated in connection with the history of the evolution of organic forms," In this paper the only other mention of evolution occurs in the
concluding remarks, in which the results and opinions of Gartner are discussed. It will be seen that Mendel expressly dissociates himself from Gartner's opposition to evolution, pointing out on the other hand that Gartner's own results are easily explained by the Mendelian theory of factors (p. 361): "Gärtner by the results of these transformation experiments was led to oppose the opinion of those naturalists who dispute the stability of plant species and believe in a continuous evolution of vegetation. He perceives in the complete transformation of one species into another an indubitable proof that species are fixed within limits which they cannot change. Although this opinion cannot be unconditionally accepted we find on the other hand in Gartner's experiments a noteworthy confirmation of that supposition regarding variability of cultivated plants which has already been expressed." It is seen from these, the only two allusions to evolution in Mendel's paper, that he did not regard his work as a direct contribution to that subject. What he does claim for the laws of inheritance he established is that they make sense of many of the results of the hybridists, and that they form a necessary basis for the understanding of the evolutionary process. On this point he shows himself fully aware of the importance of what he had done. Had he considered that his results were in any degree antagonistic to the theory of selection it would have been easy for him to say this also.