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Why was Mendel's Work Ignored?

Author(s): Elizabeth B. Gasking
Source: Journal of the History of Ideas, Vol. 20, No. 1 (Jan., 1959), pp. 60-84
Published by: University of Pennsylvania Press
Stable URL: http://www.jstor.org/stable/2707967
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WHY WAS MENDEL'S WORK IGNORED?

BY ELIZABETH B. GASKING

In 1866 Gregor Mendel put forward the basic laws of heredity

and thereby provided a foundation for the modern theory of genetics.

His results were published in a journal which circulated to a hundred

and twenty universities and learned societies, but their full signifi-

cance was ignored until 1900. After this long interval their im-

portance was recognized, and Mendel was soon world-famous. Why

was he first ignored for 34 years, and then acclaimed as a great man?

To answer this question, one must recount in some detail the

development of ideas about inheritance in the intervening years, and

set them against a changing background of theoretical preoccupa-

tions.' It will then be seen that the neglect of Mendel's results was

no accident. For one cannot just say that this paper on plant breed-

ing by an unknown monk was one which all readers of the Proceed-

ings of the Scientific Society of Brunn for 1866 all happened to skip;

nor was it simply stumbled upon againi in 1900. Bateson implies

that Mendel's paper was overlooked because it had the bad luck

to appear just when everyone was distracted by the controversy

over Darwin's Origin of Species.2 But this explanation, too, is in-

sufficient: had Mendel's work appeared before Darwin's, its fate

would have been no different. Darwin in fact paved the way for

Mendel's subsequent recognition, and this came about largely as a

consequence of later development, by Galton and others, of Darwin's

ideas. One is forced to conclude that Mendel was ignored because his

whole way of looking at the phenomena of inheritance was foreign

to the scientific thought of his time. Like Darwin, Mendel was a

conceptual innovator, with a quite novel way of thinking about

species; and until 1900 there was no place in the general framework

of biological theory into which his work could have fitted.

In one important respect, Mendel's way of thinking was more

like a farmer's than a biologist's. The professional systematist divides

living things into kinds, and assigns an individual to a kind on ac-

count of a complex of likenesses and differences: whether he counts a

particular specimen as a 'rose ' or not, for instance, will depend on

whether, all things considered, it strikes him as closer to plants al-

ready so classified than to any other group. Whereas the mathe-

1 A brief outline of the facts is given in an appendix to W. Bateson's Mendel's

Principles of Heredity (Cambridge, 1909), and a good deal of material bearing on

the question is to be found scattered throughout H. Iltis's Life of Mendel (Berlin,

1924), English trans. (London, 1932). Most general histories of biology also dis-

cuss the matter (usually drawing heavily on Iltis's work), but I know of no de-

tailed account of the events which seeks systematically to relate them to the shift-

ing focus of theoretical biology.

2William Bateson, Mendel's Principles of Heredity (Cambridge, 1909), 37.

60

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No property belongs to a rose 'by definition': what is to count as a member of the class is determined in a less formal manner. however. not with the complete nature of a species. Galton. and had no general- still less a theoretical-object in view. have a different problem. and it immediately attracted a considerable number of workers. Most of these breeders. but rather with a particular property: they want cattle of larger size. or else to dismiss it as being irrelevant to their own crucial problem of the origin of species. They are concerned.33. on the other hand. and with the ways in which the forms of the hybrid reflect the parental 'essences'. Farmers and stock-breeders. Through the newly-formed local agricultural societies practical men were able. he comes to recognize the 'specific natures' or 'essences' of his species. Hence they frequently failed to record the results of their work completely or in any detail.57 on Tue. The arti- ficial pollination of plants also had become possible. aimed merely to produce a plant or animal having certain desired characteristics. Being concerned with complex overall like- nesses. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . Men have practised selective breeding of animals for many centuries. and the importance of inheritance for them lies in the results of crossing plants or animals having this particular property in different forms or degrees. or whatever it may be. and no general conclusions can be drawn from their works alone. and individual scientists were often able to profit from the experience of practical breeders: Darwin. and so differed funda- mentally from that of other biologists. the experienced systematist develops a 'feel' for the species and sub-species he studies-to use an older terminology. the systematist's class of roses is not thus limited in advance.13. either to misinterpret his work as a confused at- tempt to investigate the nature of species. They were concerned with crosses between species. WHY WAS MENDEL S WORK IGNORED? 61 matician's class of triangles is limited in advance by a formal defi- nition of the properties of a triangle. beet with a higher sugar-content. This development was of great practical importance. Mendel's contemporaries therefore tended. to pool their experience. but during the eighteenth century the 'agricultural revolution' drew new attention to the sub- ject. The fairly widespread interest in breeding was nevertheless im- portant for science in other ways. since the new types so produced could often be propagated vegetatively. Plant-Hybridization and the Problem of Species before Mendel In the hundred years before Mendel started his research there were two motives for studying inheritance in animals and plants-a practical motive and a theoretical one. for the first time. and Bateson all This content downloaded from 200. Mendel's interest in inheritance was similar. thanks to Camerarius' work on the sexual process in the higher hermaphroditic plants. It acted as a stimulus to scientific inquiry.

had been worked on by many others before Mendel used it in his investigations: one early worker in particular.3 Apple seedlings do not bear fruit for some years. " The Fecundation of Vegetables. The pea. Other workers. were also breeding peas. He also remarked. 6 John Goss.33. his results. first. 235.4 His method of cross-pollination also was similar to Mendel's. dur- ing an inquiry to see whether it was really possible to confer charac- ters on apples by artificial pollination. and that. grapes. some of the practical re- sults obtained by breeders strike one as of special interest. was popular for a long while. This content downloaded from 200.[= green-] seeded type. stud- farmers or seedsmen. whom Mendel actually mentioned. Goss6 actually noted that. Trans. Andrew Knight. this variety is often mentioned as a stable one in subsequent literature. for reasons almost exactly the same as those Mendel was to give seventy years later. (1799). plums." Phil. V (1824). Retrospectively. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . Soc. such as Goss and Seton. It was perfectly consistent with the aims of these workers that Goss. Thus Herbert. all the subsequent generations were blue-seeded: the white seeds meanwhile continued to produce both white and blue-seeded plants. Knight used this method to improve not only apples but also cherries. Having satisfied himself that characters conferred by artificial pollination could be transmitted. and there was a belief.57 on Tue. Later he returned to work on peas again. if plants grown from these seeds were planted and allowed to self-fertilize. too. GASKING acknowledge help received from friends who were fanciers. and made no attempt to follow it up." 5 produced about 1820. do to some extent foreshadow Mendel's own." Trans. I (1826). providing a useful crop. was content to record it as a fact. that amongst the next generation blue seeds appeared once more. Hort. the hybrid produced only white seeds. even used it in 1787 as an experimental plant. this time in order to produce an improved variety: " Knight's Blue Dwarf. did a great deal of work 3 The general role of the male in plant fertilization was in dispute at this time. He selected the pea as the most suitable experimental plant. shared by Linnaeus. " On Variations in Colour of Peas Occasioned by Cross-Impreg- nation. if a white-seeded variety were crossed with a blue. having made this discovery. so he decided to try the method first with an annual. 4 A. 62 ELIZABETH B. in the hope of establishing a more general law.13. that the pollen was responsible for the vegetative parts alone. developing new hybrid ornamental plants and shrubs between 1800 and 1860. strawberries and many other cultivated plants. Knight. and though the object of his inquiry was differ- ent.. as far as they go. Many other horticulturalists. 197. 5 Gardeners' Magazine. dis- played the same lack of interest in general conclusions.

soon altered his view." This content downloaded from 200. He suggested instead that in the beginning all genera were made up of one or two species alone. Linnaeus states: " We count as many species as have been created from the beginning. where he described new hybrid rhododendrons. the individ- uals are produced from eggs. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions .7 This view led them to suppose first. that the variations between these members would be minor ones. in many genera the species seemed to form an almost continu- ous chain of a kind which was difficult to square with the idea of special creation. like the well-known mule. The theoretical interests of this group of naturalists arose from the systematic studies which had begun early in the eighteenth century. More- over. The aim of their 'experiments in hybridization' was not to produce new domestic forms but rather to throw light on the origin of species. although starting out with a belief in the special creation of each separate species.13. the resulting 'un- natural' offspring would always be. that all members of any one species would be interfertile. in- fertile. Even when it was possible to apply the interfertility test- and this could not often be done-the results were surprising. the present species having sprung from hybrids between these few original species. apart from a few notable ex- ceptions. believed that each species of plant or animal was the result of a separate divine act of creation.' It soon became clear to the early systematists that in many cases these beliefs were untrue. therefore. In 1741 he began a series of experiments in hybridization to test this view. There might in fact be such great vari- ations between members of a single one of their taxonomic species that the question whether any new specimen was a variant of species A or the first example of a new species B became more difficult to decide.33. Before that time naturalists. hollyhocks.57 on Tue. and violas which he had produced. Linnaeus. Using Camerarius' 7 Thus in an early edition (1735) of Fundamenta Botanica. which happened to be fertile and to breed more or less true. and finally. WHY WAS MENDEL S WORK IGNORED? 63 on ornamental bulbs of the family Amaryllidaceae. In fact they tended to refer to the product of any species- cross as a ' mule. publishing a book on this subject (1837) and later many articles in the Journal of Horti- culture. deviating much from the type. Nevertheless some hybridists throughout this period were doing work of a more theoretical kind. Yet in spite of the range of his work on hybrids. he neither looked for nor observed the general rules governing their production. that were it proved possible for true species to interbreed. secondly. To these men we now turn. and each egg produces progeny in all respects like the parent. due to environmental differences and never. even when in- herited.

no matter how disparate they appeared to be. The hybrid off- spring (he believed) should derive the form of their flowers from the female parent and other characters from the plant providing the pollen. Knight. cit. 9 The practice of distinguishing between interspecific crosses as 'mules. makes me much disposed to believe that hybrid plants have been mistaken for mules. loc. 64 ELIZABETH B. so that. Many of Linnaeus' hybrids were fertile. but in the next generation the qualities of the separate parents tended to reappear and. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . but my total want of success in many endeavours to produce a single mule plant. 9 and to doubt (with all the deference I feel for the opinions of Linnaeus and his illustrious followers) whether nature ever did or ever will permit the production of such a monster. Thus in 1819 and 1822 the Prussian Academy set the question. since none of them continued to breed true. they could not be regarded as new species. The more con- servative school insisted that species were and had always been distinct. The other school-the "'illustrious followers" of Linnaeus-con- tinued to believe that members of two distinct species might occasion- ally mate and produce a fertile mule. they might be lucky enough to bring about an actual case. in spite of their master's in- ability to demonstrate this. "Do hybrids between species occur in the plant kingdom? ". . GASKING techniques. There were two chief schools of thought. but he noticed that the hybrids were in many respects inter- mediate between the two parents. Petersburg and gave prizes for work on this problem. Mirabilis and Trogopogon. A similar tendency appears in German. if only more investiga- tors turned their attention to the subject. and in 1828 Wiegmann received a prize from the Brunswick Scientific Society for a monograph entitled " Ueber die Bastarderzeugung im Pflanzen- reich. was later given up.13. . possibly under the influence of Darwin's views.' This content downloaded from 200.33. he cross-pollinated different species of the genera Veronica.57 on Tue. and. In the years after 1761 many learned societies followed the ex- ample of St. both culti- 8 A. Providing that such a mule were mor- phologically distinct and continued to breed true. this was conclusive evidence that they belonged to the same species. if two individuals produced a fertile hybrid. Petersburg in 1761 for a monograph on the subject of hybridization." describing crosses between plants of many kinds. Petersburg should have offered such a prize at all is an indication of the interest the new views aroused.' and reserving the word 'hybrid' for a cross between varieties. Knight apparently subscribed to this view. Nevertheless his report won the prize offered by the Academy of Sciences at St. Verbascum. it would (they held) be the progenitor of a new species. for he remarked: 8 . where-early writers speak of 'Bastard' while Mendel in his first work uses the word 'Hybriden. The fact that the Academy of St.

and Lecoq were all probably stimulated by this competition into publishing the results of their research. He remarked that a hybrid often resembled one parent more closely than the other. even in closely related varieties. Almost at once Lecoq published the results of his work on Mirabilis and Godron his on the hybrids of Datura. cross-fertilization might sometimes be possible only if one type were constantly used as the female parent-a lack of reciprocity which had been com- mented on before. and his comments on it appear at the end of his classic monograph. Gartner's work did not bring the controversy over the status of hybrids to an end.13. In it he described a vast series of experiments which involved many different species but gave inconclusive results. The prize was awarded to Gartner in 1837. Gartner pro- duced a large book'0 which was accepted as the most authoritative work on the subject. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . Twelve years later. and thus deserved to be regarded as true species. Even as late as 1861 the Paris Academy offered a prize for a solution of the problem. Naudin had previously undertaken a long series of experiments in an attempt to discover a difference in behavior between inter-varietal and inter-specific hybrids: the essay which he now wrote on this subject is in many ways the most interesting work of its kind. Mendel obtained a copy of this book soon after he had begun his experiments on peas. yet how their own assumptions about the nature of a species still prevented them from taking the final step. and what economic and ornamental plants can be produced and multiplied in this way? " The exact wording of the question is interesting. This content downloaded from 200. and that the offspring of hybrids were variable. for it acknowledges that the problem has two distinct aspects-a theoretical one and a horticultural one-and seems to be a deliberate attempt to bring them together. finally. " Do hybrids which re- produce themselves by their own fecundation sometimes preserve in- variable characters for several generations and are they able to be- come the types of constant races?" Naudin. However. the Dutch Academy at Haarlem ran a competition for the best monograph on the question. WHY WAS MENDEL S WORK IGNORED? 65 vated and wild.33. he claimed that.57 on Tue. Godron. 1849). he claimed that some of his hybrids were constant forms which bred true. In 1830 and again in 1836. 10 Versuche und Beobachtungen iuber die Bastarderzeugung im Pflanzenreich (Stuttgart. From these results he disputed both the idea that hybrids were usually intermediate in form and the idea that they were necessarily infertile. "What does experience teach regarding the production of new species and varie- ties through the artificial fertilization of the flowers of one with the pollen of the other. It shows how very close some of these men came to Mendel's dis- coveries.

' In a hybrid the essences of the two parents were at first mixed. Naudin was one of the few who wrote explicitly about specific essences. but Naudin did not follow up the suggestion further and the idea that specific essences were simple and unique was inevitably misleading. but in one form or another the idea dominated all thinking about hybridization throughout the century before Mendel.13. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . 66 ELIZABETH B. Iltis' inquiries.57 on Tue. When he entered the 11 C." Annales des Sciences Naturelles Botaniques. Naudin. 5th series (Paris. Iltis tells us. It led to extreme vagueness: experimenters would report simply that a hybrid " resembled " the maternal or paternal species more or less closely. Mendel's work was thus completely novel. while the two other possible combinations would yield further hybrid offspring. GASKING Naudin 11 suggested that there was in every plant a hereditary material which he called the 'specific essence. Just because they saw a species as something simple. hav- ing its own essence. or that it was intermediate-they did not even trouble to say in what respects (flower-color.33. although conducted 30 years after Mendel's death (1884). This hint of segregation comes near to Mendel's own theory. they were unable to direct their attentions to the individual characters possessed by its members. and so missed completely the regularity which existed at this level. he spent much of his early life helping his father in the garden and fields. etc. but Dr. if both reproductive parts were paternal in essence. Mendel was born in the small village of Heinzendorf in Moravia in 1822. retaining throughout his life his love of country life and knowledge of farming. the new individual would resemble the original male stock. His father was a smallholder and. III. This content downloaded from 200. height.) the resemblances ap- peared. and how far was he stimulated by it? There is much internal evidence about this in his monograph. Dr. "De l'hybridite consideree comme cause de variabilite dans les vegetaux. The Character of Mendel's Enquiry How much of his predecessors' work did Mendel know. the resulting plant would resemble the original female parent. 1865). it was assumed that he would eventually inherit the farm. For this reason. If a pollen grain having an exclusively-maternal essence later fused with an ovule whose essence was of the same type. have thrown much additional light on the question." The final and complete segregation of these essences (he thought) occurred as the sex cells formed. but in the course of development separated out: hence the hybrid was " a living mosaic whose various elements are so intermingled that they cannot be distinguished by eye. as Gregor was the only son. In tackling the theory of inheritance as a problem about unit-characters rather than unit-species.

57 on Tue. where he studied physics. since the monastery gardens were famous for them: there is indeed a portrait of Mendel holding a fuchsia. he chose to become a teacher of science. mathe- This content downloaded from 200. were made with ornamental plants have already afforded evidence that hybrids. in the hope that they might also confirm the laws he had discovered. at any rate it is clear than Mendel was from the start well-versed in the practical side of breeding and familiar with the breeder's craft. An aged gardener declared that Mendel had worked on many other plants also. After finishing his secondary education he entered the monastery: then. From notes he submitted to this society. in previous years. it was natural that his hobby should be the care of the gardens and orchards. WHY WAS MENDEL S WORK IGNORED? 67 monastery at K6niginkloster. In addition he grafted and perhaps cross-pollinated the trees in his orchard. when the experiments began. where he also kept bees. It is not clear exactly what plants Mendel worked with.13. instead of tak- ing charge of a parish. and when. are not ex- actly intermediate between the parent species. His practical experience is reflected in the opening words of his classic paper: Experience of artificial fertilization such as is effected with ornamental plants in order to obtain new varieties in colour has led to the experiments which will here be discussed. Additional evidence of this early interest turned up in the Brunn monastery library where many of the numerous books on horticultural subjects bore notes in Mendel's handwriting dating from before 1857. but after failing in the teachers' examination he was permitted to go to the University of Vienna for two years. he was responsible also for the conduct of the farms owned by the monastery. In order to qualify himself he studied physics and biology. he became abbot. and was so well thought of as an orchardist that he was ap- pointed examiner for the local diploma in fruit-growing. later in life. but fuchsias were probably among them. It is therefore clear that Mendel had practical experience of the uses of cross-pollination before starting his famous series of experi- ments. He was at the same time well-informed about the scientific prob- lems which were exercising his contemporaries. Later on in the same paper he comments: Experiments which. having helped to found the latter and acted for a time as its deputy president. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . though there is no indication what these were. as a rule. it is clear that he was conducting hy- bridization experiments with his bees almost until his death. Until his death he was a member of both the Moravian and the Silesian Agricultural Societies and of the local apicultural society.33. and a local nurseryman's list contains a variety named after him.

01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . because hybrids between minor variations would. His very choice of experimental plants is consistent only with this aim. One can easily underestimate the originality and importance of this step. even when describing the results of their interspecific crosses. on their view. and he may well have believed that it was " impossible to draw a sharp line between species and varieties " even before 1856. Since he attended lectures given by one of the well known systematists of the day. The current obsession with 'specific essences' had prevented other workers from paying due attention to particular characters. may well start an investigation with one aim in view and alter his objective in the light of his preliminary dis- coveries: this need not be mentioned in his final report. and Mendel's commentators often overlook this. yet he worked for many years on garden peas and seems to have been indifferent whether his crosses were between species or only between varieties: The positions [he wrote] which may be assigned to [the plants] in a clas- sificatory system are quite immaterial for the purpose of the experiments in question. for theoretical biologists re- quired plants used in experiments on the origin of species to have two qualifications which Mendel's plants did not satisfy. A scientist. because biologists regarded the effects of domes- tication with suspicion and were anxious in their experiments to re- produce natural conditions as closely as possible. although Mendel was aware of this controversy.33. GASKING matics. However. but for laws governing the inheritance of par- ticular characters. he must have heard of the cur- rent controversy about the origin of species. and no one influenced by the idea of such essences would have dreamt of looking for the sort of laws which Mendel sought. throw no light on the nature of a species: a truly new species must come from a fertile 'mule.' Further.13. they must be wild plants. It has so far been found as impossible to draw sharp lines be- tween the hybrids of species and varieties as between species and varieties themselves. To begin with they must be true species. We tend.57 on Tue. to think of Mendel's ideas as dating from their This content downloaded from 200. in the first place. Mendel could not have been ignorant of these requirements. But in Mendel's case there is evi- dence that he was from the outset looking for laws governing the inheritance of particular characters. and biology. Mendel had rejected this idea before he set out to look for his laws. On the other hand. then his choice of garden plants and his indiffer- ence to their classificatory position was fully justified. for a scientific paper is not an autobiography. if Mendel started out in 1856 to look not for the origin of species. it seems clear that his experiments were not directly connected with it. 68 ELIZABETH B. it is true. Evidently.

No. As he wrote in the final chapter of the Origin of Species: When the views entertained in this volume on the origin of species or when analogous views are generally admitted.57 on Tue. Having rejected specific essences at least three years before the ap- pearance of Darwin's book. forgetting that. and in English translation as an Appendix to Bateson's book Mendel's Principles of Heredity (Cambridge." 12 In the introduction Mendel wrote: Those who survey the work done in this department will arrive at the con- viction that amongst all the numerous experiments made. we shall at least be freed from the vain search for the undiscovered and undiscoverable essence of the term species. they must have been planned before 1856." Proceedings of the Sci- entific Society of Brunn. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . William Bateson 13 wrote: It is to the clear conception of these three primary necessities that the whole success of Mendel's work is due. Even to appre- ciate this step of Darwin's can be difficult now. Hereafter we shall be compelled to acknowledge that the only distinction between species and well marked varieties is that the latter are known or believed to be connected by intermediate grades whereas species were formerly so connected . 121. this conception was absolutely new in his day. This content downloaded from 200. 1909).. not one has been carried to such an extent and in such a way as to make it possible to deter- mine the number of different forms under which the offspring of the hybrids appear. but the importance Darwin attached to it is clear. or definitely to ascertain their statistical relations. Close observation of his early crosses suggested to Mendel the further idea that the inheritance of these features might be governed by general laws and this idea too was completely novel.13.. . WHY WAS MENDEL S WORK IGNORED? 69 publication in 1865. .33. " Versuche ueber Pflanzenhybriden. The results of Mendel's inquiries were published in a paper en- titled " Experiments on Plant Hybridization.. Mendel was able to concentrate his at- tention on the behavior of single morphological features within any cross. for the revolution he achieved has been so complete. 13 Comments appear as footnotes to the English translation supervised by Bateson. for mean- while the publication of Darwin's Origin of Species in 1859 had effectively banished 'specific essences' from biology. since the experiments continued for ten years. 12Gregor Mendel. Systematists will be able to pursue their labours as at present. 1866: now available in Ostwald's Klassiker der Exacten Wissenschaften. or to arrange these forms with certainty according to their separate generation. By 1865 Mendel's views on species were no longer so remarkable. we can dimly foresee that there will be a considerable revolution in natural history. So far as I know. Commenting on this paragraph. but they will not be incessantly haunted by the shadowy doubt whether this or that form be in essence a species.

which appeared in 1866. This is simply not true. and Wichura. GASKING It was only because Mendel understood exactly what information was needed in order to establish the laws of heredity that he was able to achieve his aim: the "three primary necessities" to which Bateson refers in fact set out these requirements. the Brunn Scientific Society had been recently formed and the volume of Proceedings containing Mendel's work was only the fourth to be published. they did not always report their results in a way that made close comparison possible. but there the resemblance ended: both in its statisti- cal analysis of the inheritance of particular characters and in its theoretical interpretation Mendel's paper was entirely unlike all that had gone before. Publication in a new and obscure This content downloaded from 200. The audience to whom Mendel first lectured comprised many intelligent men of whom some were competent scientists. in which he noted that. Copies of these Proceedings were sent to societies and universities throughout Europe and America. Neither hybridists nor breeders had been fumbling towards the goal that Mendel reached. so that. he took the trouble to include at the end a section com- paring his results with those of Gartner. and it was published in their Proceedings for 1865. as they had other aims. The novelty of his enquiry was something Mendel did not entirely overlook.70 ELIZABETH B. since these hybrids were reported to breed true. Despite this.33. As a result the paper got no advance publicity from members of the original audience. As we have already seen. and it is a failure of historical insight to sup- pose that they were. Although the whole monograph is only about twenty pages in length. further. Mendel's monograph was readily available in most of the important centers of learning. by the end of 1866. without any reversion to the forms of earlier generations. though none were theoretical biologists of international repute. Koelreuter.13. always producing offspring like themselves. owing to their inability to recognize clearly what observations were required. Nor were the experiments which Mendel went on to describe like those of his predecessors either. But in one respect both Mendel's introduction and Bateson's comments give a misleading im- pression: they imply that other workers had looked for similar laws of inheritance unsuccessfully.57 on Tue. Yet its significance was ignored. he felt that in the majority of their cases his laws would have applied: only in the case of Wichura's Salix hybrids could he see no way of applying them. Both used the same technique of cross-pollination. The Immediate Reception of Mendet's Work Mendel read his paper to the local Scientific Society at Brunn (Brno). 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . their aims were quite different.

For while Mendel was busy working. Since the opposition also con- centrated on these aspects. an obscure format does not often seriously delay recognition. and in the dazzling light cast by the new ideas the pressing problems of earlier generations sank into insignificance. On the other hand. a scientist's paper must be carefully read and judged im- portant by his fellow-specialists. We have already seen that hybridists would tend to ignore Mendel's paper. and we need look no further for an explanation of the delay before Mendel's contribution was recognized. This will happen only if other specialists are working on similar problems. because he had not aimed at the production of a new species and had used garden varieties as his experimental plants instead of distinct species of wild plants. it should be remembered that there were at this time fewer scientific publications than now. to add to the geological and geographical evidence for evolution. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . Most of those who accepted Darwin's thesis now set themselves to trace evolutionary ancestries in the plant and animal kingdoms. there was a general re-orientation of in- terest throughout biology. both on the Continent of Europe and in America. for converts to Darwinism.13. In any case. the resulting controversy was at its height. or-particularly on the Continent-to study the implica- tions of evolution for embryology. More fundamental obstacles to recognition must there- fore be sought. WHY WAS MENDEL S WORK IGNORED? 71 journal (it might accordingly be argued) does not help to make a man's work known. Whereas before 1859 there had been no great theory or question to direct attention toward the problems of heredity. the opposition of the churches had been voiced. and those were commonly of a more general nature.57 on Tue.33. or are working in fields in which the paper in question appears relevant to an important general problem. and in any case (as we shall see) Mendel sent copies of his paper to some of the leading botanists of his day. this was not generally so: horti- culturalists and theoretical biologists alike were turning their at- tention in other directions. hybridists were by 1866 a dying race: after 1861 no more prizes were offered by learned societies for work in this field. a positive swing away from them. Many of the younger biologists had already accepted both the idea of evolution and the theory of natural selec- tion. In Mendel's case. What these were. there was now. " Keeping up with the literature" was less difficult and the paper had a better chance of being widely read in 1866 than it would have today. our study of the historical back- ground of Mendel's work does something to suggest. Even This content downloaded from 200. By 1866 Darwin's work was well-known. Even today. To be of in- fluence. a revolution had occurred in biology: in 1859 Darwin had published the Origin of Species.

33.13. there might still have been individual biologists who would have appreci- ated it. writing in 1902. and in any case there was no reason why Mendel should have thought in 1866 of sending a reprint to him. I could quote stronger expressions used by agricul- turalists. 15 The Life and Letters of Charles Darwin. William Bateson. The ability of living things to vary. The Duke of Argyll14 drew attention to this fact in his criticism of Darwin. Little wonder that. Mendel's Principles of Heredity. 16 Appendix to Bateson. saying: Strictly speaking. in such an intel- lectual climate. was a presumption they ac- cepted with natural piety. 72 ELIZAETH B. III. The younger biologists did indeed agree with him and for the time being attention was firmly focussed on natural selection. F. but only a theory on the causes which led to the relative success or failure of such new forms as may be born into the world. Mendel's paper was passed over by the rank and file of biologists. Mr. 5. Nothing in Darwin's printed works at that time suggested that he would have been a particularly suitable 14The Duke of Argyll's address to the Royal Society of Edinburgh. Darwin was right. who had also voiced this opinion. II. cit. Bateson may well have been right. therefore. Proceedings. 1883). we may feel some doubt about Darwin's power drastically to have altered the course of history. 1864. 121. Darwin's theory is not a theory on the origin of species at all. Dec. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . ed. 35.57 on Tue. But no one will attack the breeder for using such expressions and the rising generation will not blame me. Yet even though the paper were not of general interest. GASKING those biologists who remained interested in the detailed manner in which new species were formed concentrated less on the origin or variations than on the effects of selection. In a letter to Lyell. for in 1866 Darwin was working on related problems and his book Variations in Animals and Plants under Domestication appeared in 1868. But when we consider the reception given to that work. for his selective power is of such importance relatively to that of the slight spontaneous variations. said: '6 Had Mendel's work come into the hands of Darwin it is not too much to say that the history of the development of evolutionary philosophy would have been very different from that which we have witnessed. and to hand on an intensified version of these variations. had they received copies of it. Man does make his artificial breeds. Darwin (London. Darwin re- plied: 1" I will cry 'peccavi' when I hear the Duke or you attacking the breeders for saying that man has made his improved shorthorn cattle or pouter pidgeons or bantams. This content downloaded from 200.

' This comes from the first letter from Nageli to Mendel. Mendel an- nounced his intention of working with other plants and perhaps with the Hieraciums. far from acclaiming the paper. Kerner. having studied them in detail some twenty years before. when Darwin's writings had made the problem of species a burning one. to start up a correspondence with the notorious Darwin. of Saxon Scientific Society." Proc. and may have sent copies to others. In a short letter accompanying the monograph. We do not know whether Kerner acknowledged it: he never mentioned Mendel in his writings. Yet. He was engaged on this work when he received Mendel's monograph. He thought. Correns. a pupil of Niageli's. One of the known recipients was A. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . As he later explained: In the year 1864. he was a knowledgeable systematist. and did not rule out the possibility that some of the intermediate species might have arisen through hybridization. As an evolutionist Naigeli was particularly interested in poly- morphic genera. This content downloaded from 200. quite rightly. The other re- cipient was Professor Niageli of Munich: he did reply. He read Mendel's monograph with care and made notes on it. a monk. I resumed my studies of the genus Hieracium. To this Niigeli replied enthusiastically: It would seem to me especially valuable if you were able to effect hybrid fertilisations in Hieracium.57 on Tue. with special reference to the phenomenon which might justify inferences regarding the origin of varieties in species. a famous hybridist and systematist. At first sight. Niigeli was just the man to under- stand and appreciate Mendel's work. and it would perhaps have been embarrassing at this period for Mendel. Much of this interest- ing correspondence survived and was published in 1905 by C.13. referred to below as 'Letters. and he seems to have conducted breeding experiments. He even 17 C. Correns. XIX (1905). on whose many forms he was already an authority.33. he replied gently urging Mendel to take up very different work on the genus Hieracium. He chose this genus as being a polymorphic one with many integrates. and in fact corresponded with Mendel for several years. WHY WAS MENDEL S WORK IGNORED? 73 recipient. Mendel certainly sent reprints of his paper to two leading bi- ologists. " Gregor Mendels Briefe an Carl Naigeli. that these genera had never been properly investigated.'7 He evidently recognized that Mendel was a patient and careful ex- perimenter. though there is some evidence that he was aware of Mendel's later interest in the Hieraciums. He had already been an evolu- tionist before 1859. for this will soon be the group about whose in- termediate forms we shall have the most precise knowledge. and was pleased to have his collaboration.

Prealtum because your H. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . he had no reason to postulate anything as implausible as the segregation of these rudiments. he insisted. Since all Naigeli's work on Hieracium was undertaken to clarify the species question. he seems to have misunderstood Mendel's aims. like the older hybridists. Either way. 'A' [the pure dominant segregated out from the selfed hybrid] has half 'a' in its body and when inbred cannot lose that element. he must have regarded them as misconceived. The notes N'ageli made on the monograph contained the remark: The constant forms require to be tested further. glaucum. GASKING offered to supply Mendel with material. in spite of the fact that Nageli did read Mendel's monograph with some care. p. The only reason for accepting such a hypothesis at that time was that it would explain Mendel's results.33. he wrote in his first letter: Your design to experiment on plants of other kinds is excellent and I am convinced that with these others you will get notably different results. Mendel could only reply that all had in fact done so 18 This letter was discovered amongst Mendel's papers too late to be included in Correns' collection of letters.18 So.. Later.57 on Tue. To the suggestion that his 'pure forms' would not continue to re- main constant. 74 ELIZABETH B. If this was the case.e. Nor did Niigeli believe that the laws Mendel put forward were really valid. prealtum (?) seems to me suspect [i. since they were conducted on cultivated plants which were not even distinct species. for instance. Nageli wrote: I have also sent you H. and to have believed that the experiments described in it had the usual aim of clarifying the traditional questions about species. when Mendel was engaged on this work. there would be no reason for taking the theory seriously either. and suggested suitable species for him to begin on. both of them from Isarkies near Munich. may not be a 'true' species]. Perhaps this was because he would discount any results obtained with cultivated varieties. 191. This content downloaded from 200. that hybridization experiments in this field should be made on wild and true species. It appears in the Life of Mendel by Iltis. or perhaps it was because his ex- perience with Hieracium would certainly have given different results. and these Niigeli could not take seriously. prealtum and H.13. If the laws did not apply generally. but since the facts of chromosome behaviour and reduction-division were unknown at that time. I expect that they would be found to vary once more. Naigeli himself held a theory of heredity of his own. which did envisage that characters were de- termined by rudiments in the sex cells.

as with beans earlier. Mendel now turned to this problem. The genus Hieracium. we can see why the work was so fruitless. Mendel continued to experiment widely. since the florets are extremely small and the arrangement of the parts made the standard methods of cross-pollination almost useless. In the original monograph he had suggested that this phenomenon might be explicable if flower- color were determined not by one but by two pairs of factors. As Niigeli had urged.33. the later results had sug- gested that the problem was not as simple as the pea experiments had led him to believe. and they became generally known only after their eventual publication in 1905. These results puzzled him. some specimens seemed to be fertilizing themselves. and by 1869 he had succeeded in producing only six hybrid specimens: one of these was infertile. No one in the 1860's seemed interested in his laws or his theory.13. Some of his results agreed with those which he had obtained with Pisum. and in particular to the case of Hieracium.57 on Tue. botanists were still hoping that hybridizing wild species might throw light on the origin of intermediate forms in polymorphic genera. which Niigeli sowed that summer. and con- vinced him that the relations he had discovered for Pisum did not hold here. further. like some other genera of the Compositae. WHY WAS MENDEL S WORK IGNORED? 75 during the five or six generations he had observed. At Brunn and else- where. but the experi- ment seems not to have continued. His eye- sight suffered. Bit by bit the opposition of expert opinion was beginning to tell on Mendel. however careful Mendel was. contenting himself with reporting the results from time to time in letters to Nageli. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . he found a great range of color amongst the hybrids. like the Salix hybrids reported by Wichura. Experiments with Hieracium required enormous patience and skill. and the other five bred true to their hybrid form in all subsequent generations. In particular. He never bothered to publish any account of his newer experiments. so perhaps he sought to turn his attention to problems his colleagues were more interested in. with different-colored stocks. The choice could hardly have proved more unfortunate. either for practical or for theoretical reasons. Others. is both This content downloaded from 200. as we now know-whether his theory was applicable in cases of this kind: We thus obtain from the different colors. In spite of Nageli's indifference. and in the next few years he worked on no less than twenty- six different genera. In retrospect. however. and neither side ever mentioned the Pisum monograph in their subsequent correspondence. He sent seeds from such plants. But after this second discovery he began to doubt-needlessly. figures from which the deduction of a developmental formula is impracticable. did not.

Soc. rather sadly: I must express some scruples in describing in this place an account of ex- periments just begun. In some species and specimens. reporting new results. and the uncertainty whether it will be granted to me to bring the same to a conclusion have de- termined me to make the present communication. Most hybrids of Hieracium do in fact reproduce apogamously. and concludes: Whether from this circumstance one may venture to draw the conclusion that the polymorphism of the genera Saltix and Hieracium is connected with the special conditions of their hybrids is still an open question which may well be raised but not as yet answered. When this happens. There follows an account of the production of the six hybrids and their behavior. in a lecture which can be found in the subsequent volume of Proceedings. a new embryo may yet be produced. and must have been very disappointed at finding this apparent refutation of his earlier theories. Nevertheless he faithfully reported the results to the Brunn Scientific Society in June 1869. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . Sci. Brunn (1869).13. He remarks how differently these new hybrids (and Wichura's Salix hybrids) had performed from his own earlier Pisum hybrids. But the conviction that the prosecution of the pro- posed experiments will demand a whole series of years. there is no reduction division in the ovules and no fertilization. Mendel could not recognize the deceptive character of his re- sults. In 1872. even if pollen is produced. and must have been much more intelligible to his contemporaries. Mendel. He had been made abbot in 1868 and had less time for his hobbies. GASKING variable and atypical in its methods of reproduction. " Experiments in Hieracium Hybridisation. 76 ELIZABETH B. however. and therefore remain constant in form. as a product of sexual fusion: such an offspring will be identical with its 'parent' cytologically. This Hieracium paper was really Mendel's botanical swan song. He continued the Hieracium experiments for a while and still wrote to Niigeli. but quite frequently. This content downloaded from 200. fertilization may be normal. developing apogamously from a diploid cell and not. in the way it appears to do.33.57 on Tue." Proc. This paper was very different from his earlier monograph. the strain on his 19 G. to which Mendel adds. He reports the aims of his enquiry as follows: 19 From the nature of the subject it is clear that without an exact knowledge of the structure and fertility of the hybrids and the condition of their off- spring through several generations no one can undertake to determine the possible influence exercised by hybridization over the multiplicity of inter- mediate forms of Hieracium.

published in 1881: this was the reference which led later workers to the original monograph. d. pro- vided very quickly. but could not. 0. and the characteristic changes in the chromosomes in this process were studied This content downloaded from 200. Mendel and his work sank into complete obscurity. towards the end of the nineteenth century. in W. it was perhaps inevitable that his discovery should not have been appreci- ated. however complete in itself. Focke recorded Mendel's claim that there was a constant numerical ratio between the various types of offspring of the hybrid pea. Focke's work Die Pflanzenmischlinge. for. though he seems. The additional pieces were.57 on Tue. Mendel now spent his limited leisure on his bees and his meteorological studies." The Rise of Cytology and Biometrics between 1865 and 1900 Mendel had produced a key piece for the jigsaw of biological theory-a much more important piece than he could have realized- but it was of no general use until the picture was sufficiently com- plete for it to be fitted in. remember how he came to read Mendel's work. to have recognized the situation and got some comfort from it. but in none of his works was Mendel ever mentioned: only in the bibliography of a work entitled Die Hieracien Mitteleuropas by Niigeli and Peter (which was published shortly after Mendel's death in 1884) is there a brief entry: Mendel: Versuche ueber Pflanzenhybriden (Verhandl. when asked. Niigeli continued to write about heredity until his death in 1891. The structure of the cell was observed in greater detail: the process of cell division. 1865. is valued until it can be fitted into the general corpus. his only re- corded comment on the fate of his first monograph was to say: " My time will come. Science is organized knowledge. 69).13. there was one other reference to the Pisum mono- graph before 1900. The most important change during this period was the advance in cytology. Given the position in biology when Mendel wrote.33. in particular reduction division. At any rate. in Brunn. This did not make it any the less of a personal tragedy for Mendel. in fact. and no piece of work. Naturhistorischer Ver. Here we can outline only the major discoveries and developments between 1865 and 1900. at least in part. Apart from this. as a result of which Mendelian genetics could take its proper place in the structure of biology. Except for these two inconspicuous refer- ences. the subject was developing at a great rate. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . WHY WAS MENDEL S WORK IGNORED? 77 eyes and the inconclusive results he had recently been obtaining in- duced him to drop the subject: he sent all his remaining specimens to Niigeli and the correspondence lapsed.

since the link between the factors and the chromosomes was demonstrated experimentally only ten years later. though long suspected. There were many other such theories. and greatly increased their understanding of the mechanism of sexual reproduction. as the controversies about evolu- tion became more intense. Darwin had surprised everyone by producing in 1868 his Variations in Plants and Animals. GASKING for the first time in the 1880's and the early 1890's. but the problem itself had changed in character. but there were many others: Spencer's theory of physiologi- cal units. 78 ELIZABETH B.57 on Tue. in this way it became clear that every sexual fusion was preceded by a reduction division. Darwin's theory of pangenesis and Galton's theory of the stipe were all produced at this time. Biological theorists therefore had to consider the possible causes of variations. yet it was becoming in- creasingly clear that the only visible difference between the zygotes of mice and men was in the number of chromosomes. Towards the end of the nineteenth century. Whereas in the eighteenth century the continuity of fixed types alone had to be explained. Into this new picture Mendel's theory fitted far more naturally. how 'form' was transmitted from one generation to the next- how it was that cats always gave birth to kittens and acorns grew into oaks. and these speculations in turn drew attention to the variations themselves. now almost completely forgotten. His hypothesis that both sexes contribute equally to the new individual gained a new meaning when the nuclei of the gametes were seen to fuse. but now it reappeared in a different guise. These cytological discoveries redirected attention to the old prob- lem. speculation about the causes of variation also revived. Towards the close of the century. In the in- This content downloaded from 200. While the majority of biologists in the 1860's were not (as we have seen) interested in heredity. Each type reproduced its kind. was now observed for the first time. the haploid and diploid phases were identified in the life-cycles of many animals and plants. the number of chromosomes in the cells of any species so remaining constant from generation to generation. This general problem had occurred to biologists at many different periods in the past. and to the facts about their inheritance.33. not only was much more known about the facts of sexual reproduction. Weismann's theory of the germ-plasm is the best-known of the theories. now every type must be thought of as gradually altering. Dis- coveries such as these altered completely the biologists' picture of the nature of living things. The actual fusion of the two sex cells. and though in 1900 this support was indirect. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . the new knowledge of chromosome segregation did at any rate make factor segregation seem a possible vera causa.13. the recognition of the haploid gametes and the diploid zygote gave an indirect plausibility to the idea of factor segregation.

this book was poorly received. especially in England.57 on Tue. one must look more closely at the situation in the 1890's. Doubt was now thrown on the old assumption that some acquired characters could be inherited: once this was questioned by Galton. had Mendel's work been lost forever. This content downloaded from 200. and he analyzed his results statistically. In order to see how it was that his monograph was actually recognized by several different workers almost simultaneously. who had always been interested in hybridization. such as height. and that the rediscovery of his monograph had at most the effect of aiding and speeding up the birth of the subject. Galton's methods resembled Mendel's: he studied particular characters. modern genetics would nevertheless be much the same today. It was a fairly complete collection of the known facts about the appearance and inheritance of variations. so now that the theory of heredity was beginning to receive attention. a new impetus was given to scientific breeding. Galton's long-sustained concern with human inheritance did much to awaken interest in the subject. nevertheless. and knowl- edge about inheritance gradually increased. written by such a well-known author. particularly amongest domestic plants and animals. in particular. instead of the overall similarity and differ- ence between individuals. than create a situation in which it was at last possible for Mendel's theories to be appreciated and understood. however.33. at the work of a group of young biologists who were leading a revolt against the orthodox Darwinian view of the origin of species. well-trained scientists started for the first time to design breeding experiments which would clarify the question of how variations arose and were passed on. did become quite influential. did something to interest biologists in problems which had previously been considered only by practical breeders. His habits of collecting data from the general public and of publishing his re- sults in popular form also attracted the attention of many laymen. and only the agricultural journals reviewed it favorably.13. The Last Phase Work on cytology and biometrics did no more. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . Paradoxically it seems that. In two crucial re- spects. creating increasing public interest in heredity. Long and complicated experiments were conducted on animals. The book. Such an as- sembly of facts in one book. As he de- veloped his novel 'biometric' methods of investigation. WHY WAS MENDEL S WORK IGNORED? 79 tellectual climate of the time. The only bi- ologist to approve of it was the systematist George Bentham. he drew into that discussion other scientists and mathematicians who would other- wise have had no more than a passing interest in biology. and later by Weismann. so one need not be surprised to find that his pupils were readier than earlier biologists to see the real significance of Mendel's work.

struck by the number of intergrades recently discovered be- tween normal familiar types. This view of the evolutionary process was based on two assumptions which were first questioned during the last decade of the nineteenth century. that large and sudden variations might occur. however. Bateson. This content downloaded from 200. Darwin allowed. This second assumption was reasonable on the common view of inheritance: according to this view. Hence. Darwin had assumed that living things formed a continuous series of minutely different forms. Each of them attacked the orthodox position from a different point of view.' Without the idea of 'segregation. the character appearing in the individual being a re- sultant of the influences exerted by these assorted particles. and sometimes as small and continuous. but he assumed that when discontinuous variations did appear they would be swamped by interbreeding. This emphasis was an understandable reaction against the common idea of species as simple natural units each with a distinct essence.33.' biologists naturally thought that the particles mixed haphazardly during re- production. 80 ELIZABETH B. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . aided perhaps by the in- heritance of acquired characters. and by the way in which the remaining gaps could be narrowed by considering also the evidence from geology. so that after a sufficient number of generations the norm for that group had significantly altered. while de Vries looked for cases in which present species could be observed giving rise to new forms. for the early Darwinians. The revolt against this orthodox view of the evolutionary process was led by three men-de Vries. but it proved to be an overemphasis and distracted attention from the possibility that new types might arise suddenly. Basically. GASKING Darwin had originally spoken about variations with two different voices. of course. species were in an important sense artificial. it was this work which dovetailed so perfectly with Mendel's to form the twin foundation-stones of modern genetics. Bateson and de Vries both pointed out that Darwin's belief in the continuity of living forms was exaggerated.57 on Tue. Johannsen set out to show that the positive part of Darwin's picture would not work. and Johannsen-all of whom attacked it well before 1900. Bateson go- ing on to show that discontinuous variations which had arisen in the past did in fact survive interbreeding.13. The research which their theoretical objections led them to undertake was still in progress in 1900. it was for him the small and continuous variations which provided the raw material for evolution: natural selection. The first assumption was contained in Darwin's idea of the nature of a species. gradually brought about a change within any group. but the effects on the adult form of the individual particles were thought to be mixed or 'blended. regarding them sometimes as being large and sudden. the material basis of inheritance might be particulate.

but which together constituted the chromosomes.33. but his work on mutation in species of Oenothera was already under way before that date and his ideas on the two subjects were interconnected. He also attacked Darwin's conclusion that ' species' is an artificial concept. (It was in this connection that Johannsen introduced his distinction between the outward appearance of that individual-the phenotype-and the sum total of the inheritable material-the genotype. De Vries. WHY WAS MENDEL S WORK IGNORED? 81 Johnannsen investigated the question of the effects of natural selection on a mixed group. he started breeding experiments. His ideas on heredity were published in 1889. but no further selection of larger or smaller beans within this line would produce offspring which differed in their average weight. According to his view. independently variable and independently inheritable. so effecting a reassortment of the unit-characters. but for one species to give rise This content downloaded from 200. by citing known examples of discontinuous variation and arguing: Let the believer in the efficacy of selection operating on continuous fluctua- tion try to breed a dwarf sweet pea from a tall race by choosing the short- est to breed from. for instance in the soil. that there are gaps in the sequence of living forms. In order to collect further evidence about discontinuous variations. the results of which supported Mendel's views. on the grounds that distinct species do exist in nature or. whose final work on mutation was published after 1900. organisms were built up of unit-characters. when the monograph was once again put into circulation. further selection had no influence: the size of the beans would fluctuate ow- ing to external differences. and they gave additional meaning both to Mendel's idea of a constant variety and to the distinction between the form of the hybrid and the constitution of the sex cells. at the very least. But if the breeder then isolated these pure lines and allowed them to self-fertilize. In this way he showed that by selection it was possible to establish pure lines whose average weight differed from that of the original group.57 on Tue. and.) Johannsen published his results in 1903.13. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . He started with a mixed population of beans and bred from the heaviest. and did much to publicize it. These unit-characters were repre- sented in potentia in the germ-cells by definite bodies-the 'pangens' -which were too minute to be independently visible. he arranged for its translation into English. In 1894 Bateson published his Material for the Study of Varia- tions. In this book he attacked the idea that small continuous varia- tions could be responsible for evolution. He regarded species as dis- tinct types whose representatives normally interbred. had actually started his work in the early 1880's.

In this paper he said: In my hybridization experiments upon races of maize and peas I have come to the same results as de Vries. but two days earlier on March 24th. de Vries added: This monograph is so rarely quoted that I did not myself become ac- quainted with it until I had conducted most of my experiments and had in- dependently deduced the above propositions. and at once sent a paper to the German Botanical Society which appeared in the April issue of their Journal. Meanwhile. which gave a fuller account of the work reported in the French paper. March 1900). des Sciences (Paris. 3 (1900). devoting many years to the most extensive experiments on peas. In a patch of Oenothera lamarckiana growing at Hilversum he thought he had found a species in such a state of change. XVIII. Subsequently.20 This was published on March 26th. GASKING to another there must be a more drastic change. While engaged in the resulting breeding experiments.13. 21 Ber. so he took specimens of the new forms back to the University where he studied their behavior and bred from them. however I found that in Brunn. during the sixties.. 1900.. but not for his mutants. Correns received de Vries' French paper at the beginning of April. 82 ELIZABETH B.57 on Tue. and the new form must possess entirely new characters. A sudden change of their sort he called a mutation: an old species (he thought) might occasionally enter into a period of change. also arrived at the same results while in- vestigating Darwin's views about vigor in peas. Abbot Gregor Mendel. after noting that his results agreed with those which an Austrian monk. during which it would give rise by mutation to one or more mutant forms. Gesellsch. in Austria. had not only obtained the same results as de Vries and myself but 20 Comptes Rendues de l'Acad.. had obtained thirty-four years before. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . he went on to test it for many different genera before publishing his results. This content downloaded from 200. I believed myself to be an innovator. Gregor Mendel. concluding that it held true for normal varieties. Deutsch.. he had read a paper Ueber das Spaltungsgesetz der Bastarde to the German Botanical Society. in Germany. Bot.2' This latter paper. de Vries sent a short note entitled Sur la loi de disjonction des hybrides to the French Academy of Sciences. he hit for himself on the law of segregation. In March 1900. contains- at long last-a reference to Mendel's monograph and does justice to the significance of his discoveries.33. At the same time Tschermak. Correns had been investigating the effect of pollen on the form of the endosperm in maize and had discovered that the endosperm originated from a second fusion within the embryo sac: in the course of this work he had conducted a series of hybridiza- tion experiments and arrived at Mendel's first law.

But this is evidently not the whole story. in which the significance of Mendel's monograph was finally recognized. reported how he had first thought he had made an entirely novel discovery. 23 Ibid. the other wrinkled ones. Deutsch. for his problems were the problems of the time. WHY WAS MENDEL S WORK IGNORED? 83 had actually given the very same explanation as far as was possible in the year 1866. in spite of the striking independent confirmation they had received. but had found that his results had been anticipated when he later read Mendel's monograph. the other remaining short. XVIII. the year 1900. immedi- ately hybridization experiments were restarted. Next he allowed these hy- brids to fertilize themselves and examined the form of the progeny: all the progeny exhibited one or other of the parental characters.22 In June 1900 Tschermak also communicated his results to the Ger- man Botanical Society: 23 he too. one plant having smooth seeds. and so on.. As soon as it was republished and translated. But though 'Mendelism' was to be the subject of heated controversy for more than a decade. He extended these experiments to subsequent genera- 22 Ber.57 on Tue. Having arrived at the laws. and so were more likely to notice numerical ratios in any result. all of them were sufficiently familiar with the controversy about the importance of variations to realize their great importance and interest. now that they were discovered. certainly marks the be- ginning of a new era in the growth of our understanding of heredity. to the fact that the later workers were more accustomed to keeping numerical records and analyzing them statistically. and the character which appeared in the hybrid he described as the 'dominant' character of each pair.. each of which resembled one of the par- ents. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . Bot. Mendel's work rap- idly became famous. This is not to say that his conclusions were immediately accepted. This was partly due. 6 (1900). In his monograph on plant hybridization Mendel first described a group of seven sets of experi- ments in which he had crossed plants varying in one clearly-marked constant and contrasting characteristic-one plant having a tall habit. close to seventy-five per cent of these progeny resembling the hybrid parent and close to twenty-five per cent resembling the other origi- nal parent. Gesellsch. 4 (1900). no doubt. again. From 1760 to 1860 many people had studied hybrids without recognizing Mendel's laws. In every one of his seven groups he obtained about 250 hybrids. What chiefly prevented the earlier workers from observing the regularity was their theoretical outlook: all the men who later rediscovered the laws had abandoned the old assumptions about 'specific essences' and so were ready to observe individual characters and their mode of transmission. XVIII.13.33. This content downloaded from 200. APPENDIX: The Argument of Mendel's Monograph Mendel's Experiments and Theory.

and any subsequent generation produced as a result of a self-fertilization would also resemble this stock. On a chance basis. In this way he came very near to formulating the later 'multifactor' hypothesis. 84 ELIZABETH B. Mendel suggested a theory to account for his laws. while the other half yield plants which remain constant and receive the dominant or the recessive characters (respectively) in equal numbers. The theory indicated that the hybrids pro- duced pollen grains and egg cells of two types. the result would be a hybrid. This content downloaded from 200. and this would continue to behave in the same way as the original hybrids. and in the light of the results enunciated his first law-now known as the Law of Segregation-as follows: The hybrids form seeds having one or other of the two differentiating char- acters and of these one half develop again the hybrid form. University of Melbourne. He discovered that some of those behaved like his peas. GASKING tions. If a pollen grain and an egg cell each bearing a factor for the recessive character fused. In addition. and all subsequent generations bred by self-fertilization would also resemble this stock.57 on Tue. and Mendel suggested that. in this case. If a pol- len grain and an egg cell both bearing the factor for the dominant character fused. It only required the ad- ditional assumption that the factors segregated independently for Mendel's second law to be explained. These results led him to the second law of heredity-the Law of Independent Assortment-which he formulated as follows: The offspring of the hybrid in which several essentially different characters are combined represent the terms of a series of combinations in which the developmental series for each pair of differentiating characters are assorted. but with beans the flower colors did not follow the pat- tern of his other results: the hybrids exhibited a range of colors. Mendel described some experiments he had under- taken with other species. So long as the different types of reproductive cells were produced in equal numbers and provided that sufficient cases were examined. If unlike egg and pollen cells fused. Half of each kind of reproductive cell carried a factor for the character of one original parent and half a factor for the character of the other parent. the color might be determined by more than one independent factor.13. there were three possible combinations which could re- sult from the fusion of the pollen grains and the egg cells. and studied their progeny. and in one case three. In the next section of the work. the resultant offspring would resemble the other original parent. the resultant offspring would share this character with one of the original parents. 01 Mar 2016 19:24:18 UTC All use subject to JSTOR Terms and Conditions . the proportions of the progeny should obey the law Mendel had discovered.33. characters. In the next set of experiments Mendel crossed plants which dif- fered with respect to two.