Why was Mendel's pioneering work on genetics initially ignored

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
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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
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, the systematist's class of roses
is not thus limited in advance. 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. Being concerned with complex overall like-
nesses, the experienced systematist develops a 'feel' for the species
and sub-species he studies-to use an older terminology, he comes to
recognize the 'specific natures' or 'essences' of his species. Farmers
and stock-breeders, on the other hand, have a different problem. They
are concerned, not with the complete nature of a species, but rather
with a particular property: they want cattle of larger size, beet with
a higher sugar-content, or whatever it may be, 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.
Mendel's interest in inheritance was similar, and so differed funda-
mentally from that of other biologists. They were concerned with
crosses between species, and with the ways in which the forms of
the hybrid reflect the parental 'essences'. Mendel's contemporaries
therefore tended, either to misinterpret his work as a confused at-
tempt to investigate the nature of species, or else to dismiss it as
being irrelevant to their own crucial problem of the origin of species.
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. Men have practised selective
breeding of animals for many centuries, but during the eighteenth
century the 'agricultural revolution' drew new attention to the sub-
ject. Through the newly-formed local agricultural societies practical
men were able, for the first time, to pool their experience. The arti-
ficial pollination of plants also had become possible, thanks to
Camerarius' work on the sexual process in the higher hermaphroditic
plants. This development was of great practical importance, since the
new types so produced could often be propagated vegetatively, and
it immediately attracted a considerable number of workers. Most
of these breeders, however, aimed merely to produce a plant or
animal having certain desired characteristics, and had no general-
still less a theoretical-object in view. Hence they frequently failed
to record the results of their work completely or in any detail, and
no general conclusions can be drawn from their works alone.
The fairly widespread interest in breeding was nevertheless im-
portant for science in other ways. It acted as a stimulus to scientific
inquiry, and individual scientists were often able to profit from the
experience of practical breeders: Darwin, Galton, and Bateson all
62 ELIZABETH B. GASKING
acknowledge help received from friends who were fanciers, stud-
farmers or seedsmen. Retrospectively, too, some of the practical re-
sults obtained by breeders strike one as of special interest. The pea,
providing a useful crop, had been worked on by many others before
Mendel used it in his investigations: one early worker in particular,
Andrew Knight, even used it in 1787 as an experimental plant, dur-
ing an inquiry to see whether it was really possible to confer charac-
ters on apples by artificial pollination.3 Apple seedlings do not bear
fruit for some years, so he decided to try the method first with an
annual. He selected the pea as the most suitable experimental plant,
for reasons almost exactly the same as those Mendel was to give
seventy years later.4 His method of cross-pollination also was
similar to Mendel's, and though the object of his inquiry was differ-
ent, his results, as far as they go, do to some extent foreshadow
Mendel's own.
Having satisfied himself that characters conferred by artificial
pollination could be transmitted, Knight used this method to improve
not only apples but also cherries, plums, grapes, strawberries and
many other cultivated plants. Later he returned to work on peas
again, this time in order to produce an improved variety: " Knight's
Blue Dwarf," 5 produced about 1820, was popular for a long while.
Other workers, such as Goss and Seton, were also breeding peas.
Goss6 actually noted that, if a white-seeded variety were crossed
with a blue- [= green-] seeded type, the hybrid produced only white
seeds. He also remarked, first, that amongst the next generation
blue seeds appeared once more, and that, if plants grown from these
seeds were planted and allowed to self-fertilize, all the subsequent
generations were blue-seeded: the white seeds meanwhile continued
to produce both white and blue-seeded plants.
It was perfectly consistent with the aims of these workers that
Goss, having made this discovery, was content to record it as a fact,
and made no attempt to follow it up, in the hope of establishing a
more general law. Many other horticulturalists, developing new
hybrid ornamental plants and shrubs between 1800 and 1860, dis-
played the same lack of interest in general conclusions. Thus
Herbert, whom Mendel actually mentioned, did a great deal of work
3 The general role of the male in plant fertilization was in dispute at this time,
and there was a belief, shared by Linnaeus, that the pollen was responsible for the
vegetative parts alone.
4 A. Knight, " The Fecundation of Vegetables," Phil. Trans. (1799), 197.
5 Gardeners' Magazine, I (1826); this variety is often mentioned as a stable
one in subsequent literature.
6 John Goss, " On Variations in Colour of Peas Occasioned by Cross-Impreg-
nation," Trans. Hort. Soc., V (1824), 235.
on ornamental bulbs of the family Amaryllidaceae, publishing a book
on this subject (1837) and later many articles in the Journal of Horti-
culture, where he described new hybrid rhododendrons, hollyhocks,
and violas which he had produced. Yet in spite of the range of his
work on hybrids, he neither looked for nor observed the general rules
governing their production.
Nevertheless some hybridists throughout this period were doing
work of a more theoretical kind. The aim of their 'experiments in
hybridization' was not to produce new domestic forms but rather to
throw light on the origin of species. To these men we now turn.
The theoretical interests of this group of naturalists arose from
the systematic studies which had begun early in the eighteenth
century. Before that time naturalists, 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.7 This view led them to suppose
first, that all members of any one species would be interfertile;
secondly, that the variations between these members would be minor
ones, due to environmental differences and never, even when in-
herited, deviating much from the type; and finally, that were it
proved possible for true species to interbreed, the resulting 'un-
natural' offspring would always be, like the well-known mule, in-
fertile. In fact they tended to refer to the product of any species-
cross as a ' mule.'
It soon became clear to the early systematists that in many cases
these beliefs were untrue. 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. Even when it was possible to apply the interfertility test-
and this could not often be done-the results were surprising. More-
over, 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.
Linnaeus, therefore, although starting out with a belief in the
special creation of each separate species, soon altered his view. He
suggested instead that in the beginning all genera were made up of
one or two species alone, the present species having sprung from
hybrids between these few original species, which happened to be
fertile and to breed more or less true. In 1741 he began a series of
experiments in hybridization to test this view. Using Camerarius'
7 Thus in an early edition (1735) of Fundamenta Botanica, Linnaeus states:
" We count as many species as have been created from the beginning; the individ-
uals are produced from eggs, and each egg produces progeny in all respects like the
parent,"
techniques, he cross-pollinated different species of the genera
Veronica, Verbascum, Mirabilis and Trogopogon. 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, but he noticed that the hybrids were in many respects inter-
mediate between the two parents. Many of Linnaeus' hybrids were
fertile, but in the next generation the qualities of the separate parents
tended to reappear and, since none of them continued to breed true,
they could not be regarded as new species. Nevertheless his report
won the prize offered by the Academy of Sciences at St. Petersburg
in 1761 for a monograph on the subject of hybridization.
The fact that the Academy of St. Petersburg should have offered
such a prize at all is an indication of the interest the new views
aroused. There were two chief schools of thought. The more con-
servative school insisted that species were and had always been
distinct; so that, if two individuals produced a fertile hybrid, this was
conclusive evidence that they belonged to the same species, no matter
how disparate they appeared to be. Knight apparently subscribed
to this view, for he remarked: 8
. . . but my total want of success in many endeavours to produce a single
mule plant, makes me much disposed to believe that hybrid plants have
been mistaken for mules; 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.
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, in spite of their master's in-
ability to demonstrate this. Providing that such a mule were mor-
phologically distinct and continued to breed true, it would (they
held) be the progenitor of a new species; and, if only more investiga-
tors turned their attention to the subject, they might be lucky
enough to bring about an actual case.
In the years after 1761 many learned societies followed the ex-
ample of St. Petersburg and gave prizes for work on this problem.
Thus in 1819 and 1822 the Prussian Academy set the question, "Do
hybrids between species occur in the plant kingdom? "; and in 1828
Wiegmann received a prize from the Brunswick Scientific Society
for a monograph entitled " Ueber die Bastarderzeugung im Pflanzen-
reich," describing crosses between plants of many kinds, both culti-
8 A. Knight, loc. cit. 9 The practice of distinguishing between interspecific
crosses as 'mules,' and reserving the word 'hybrid' for a cross between varieties,
was later given up, possibly under the influence of Darwin's views. A similar
tendency appears in German, where-early writers speak of 'Bastard' while Mendel
in his first work uses the word 'Hybriden.'
vated and wild. From these results he disputed both the idea that
hybrids were usually intermediate in form and the idea that they were
necessarily infertile.
In 1830 and again in 1836, the Dutch Academy at Haarlem ran a
competition for the best monograph on the question, "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, and what economic and ornamental plants can
be produced and multiplied in this way? " The exact wording of the
question is interesting, 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. The prize
was awarded to Gartner in 1837. Twelve years later, Gartner pro-
duced a large book'0 which was accepted as the most authoritative
work on the subject. In it he described a vast series of experiments
which involved many different species but gave inconclusive results.
He remarked that a hybrid often resembled one parent more closely
than the other, and that the offspring of hybrids were variable; he
claimed that, even in closely related varieties, 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; finally, he claimed that some of his hybrids were
constant forms which bred true, and thus deserved to be regarded as
true species. Mendel obtained a copy of this book soon after he had
begun his experiments on peas, and his comments on it appear at the
end of his classic monograph.
However, Gartner's work did not bring the controversy over the
status of hybrids to an end. Even as late as 1861 the Paris Academy
offered a prize for a solution of the problem, " 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, Godron, and Lecoq
were all probably stimulated by this competition into publishing the
results of their research. Almost at once Lecoq published the results
of his work on Mirabilis and Godron his on the hybrids of Datura.
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. It
shows how very close some of these men came to Mendel's dis-
coveries, yet how their own assumptions about the nature of a species
still prevented them from taking the final step.
10 Versuche und Beobachtungen iuber die Bastarderzeugung im Pflanzenreich
(Stuttgart, 1849).
Naudin 11 suggested that there was in every plant a hereditary
material which he called the 'specific essence.' In a hybrid the
essences of the two parents were at first mixed, 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." The final and complete segregation of these
essences (he thought) occurred as the sex cells formed. If a pollen
grain having an exclusively-maternal essence later fused with an
ovule whose essence was of the same type, the resulting plant would
resemble the original female parent; if both reproductive parts were
paternal in essence, the new individual would resemble the original
male stock; while the two other possible combinations would yield
further hybrid offspring. This hint of segregation comes near to
Mendel's own theory, but Naudin did not follow up the suggestion
further and the idea that specific essences were simple and unique
was inevitably misleading.
Naudin was one of the few who wrote explicitly about specific
essences, but in one form or another the idea dominated all thinking
about hybridization throughout the century before Mendel. It led
to extreme vagueness: experimenters would report simply that a
hybrid " resembled " the maternal or paternal species more or less
closely, or that it was intermediate-they did not even trouble to
say in what respects (flower-color, height, etc.) the resemblances ap-
peared. Just because they saw a species as something simple, hav-
ing its own essence, 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. In tackling the
theory of inheritance as a problem about unit-characters rather than
unit-species, Mendel's work was thus completely novel.
The Character of Mendel's Enquiry
How much of his predecessors' work did Mendel know, and how
far was he stimulated by it? There is much internal evidence about
this in his monograph, but Dr. Iltis' inquiries, although conducted 30
years after Mendel's death (1884), have thrown much additional light
on the question.
Mendel was born in the small village of Heinzendorf in Moravia
in 1822. His father was a smallholder and, as Gregor was the only
son, it was assumed that he would eventually inherit the farm. For
this reason, Dr. Iltis tells us, he spent much of his early life helping
his father in the garden and fields, retaining throughout his life his
love of country life and knowledge of farming. When he entered the
11 C. Naudin, "De l'hybridite consideree comme cause de variabilite dans les
vegetaux," Annales des Sciences Naturelles Botaniques, III, 5th series (Paris, 1865).
monastery at K6niginkloster, it was natural that his hobby should be
the care of the gardens and orchards, where he also kept bees; and
when, later in life, he became abbot, he was responsible also for the
conduct of the farms owned by the monastery. Until his death he
was a member of both the Moravian and the Silesian Agricultural
Societies and of the local apicultural society, having helped to found
the latter and acted for a time as its deputy president. From notes
he submitted to this society, it is clear that he was conducting hy-
bridization experiments with his bees almost until his death, in the
hope that they might also confirm the laws he had discovered. In
addition he grafted and perhaps cross-pollinated the trees in his
orchard, and was so well thought of as an orchardist that he was ap-
pointed examiner for the local diploma in fruit-growing.
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.
Later on in the same paper he comments:
Experiments which, in previous years, were made with ornamental
plants have already afforded evidence that hybrids, as a rule, are not ex-
actly intermediate between the parent species.
It is therefore clear that Mendel had practical experience of the
uses of cross-pollination before starting his famous series of experi-
ments. 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, when the experiments began. It is not clear exactly what
plants Mendel worked with, but fuchsias were probably among them,
since the monastery gardens were famous for them: there is indeed a
portrait of Mendel holding a fuchsia, and a local nurseryman's list
contains a variety named after him. An aged gardener declared that
Mendel had worked on many other plants also, though there is no
indication what these were; 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.
He was at the same time well-informed about the scientific prob-
lems which were exercising his contemporaries. After finishing his
secondary education he entered the monastery: then, instead of tak-
ing charge of a parish, he chose to become a teacher of science. In
order to qualify himself he studied physics and biology, but after
failing in the teachers' examination he was permitted to go to the
University of Vienna for two years, where he studied physics, mathe-
matics, and biology. Since he attended lectures given by one of the
well known systematists of the day, he must have heard of the cur-
rent controversy about the origin of species.
However, although Mendel was aware of this controversy, it seems
clear that his experiments were not directly connected with it. A
scientist, it is true, 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, for a scientific
paper is not an autobiography. But in Mendel's case there is evi-
dence that he was from the outset looking for laws governing the
inheritance of particular characters. His very choice of experimental
plants is consistent only with this aim, 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. To begin with
they must be true species, because hybrids between minor variations
would, on their view, throw no light on the nature of a species: a
truly new species must come from a fertile 'mule.' Further, they
must be wild plants, 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. Mendel could not
have been ignorant of these requirements, 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. 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.
On the other hand, if Mendel started out in 1856 to look not for
the origin of species, but for laws governing the inheritance of par-
ticular characters, then his choice of garden plants and his indiffer-
ence to their classificatory position was fully justified. The current
obsession with 'specific essences' had prevented other workers from
paying due attention to particular characters, even when describing
the results of their interspecific crosses, and no one influenced by the
idea of such essences would have dreamt of looking for the sort of
laws which Mendel sought. Evidently, Mendel had rejected this
idea before he set out to look for his laws, and he may well have
believed that it was " impossible to draw a sharp line between
species and varieties " even before 1856.
One can easily underestimate the originality and importance of
this step, and Mendel's commentators often overlook this. We tend,
in the first place, to think of Mendel's ideas as dating from their
publication in 1865, forgetting that, since the experiments continued
for ten years, they must have been planned before 1856. By 1865
Mendel's views on species were no longer so remarkable, for mean-
while the publication of Darwin's Origin of Species in 1859 had
effectively banished 'specific essences' from biology. Even to appre-
ciate this step of Darwin's can be difficult now, for the revolution he
achieved has been so complete; but the importance Darwin attached
to it is clear. 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, we can dimly foresee that there
will be a considerable revolution in natural history. Systematists will be
able to pursue their labours as at present; but they will not be incessantly
haunted by the shadowy doubt whether this or that form be in essence a
species.... 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 . . . we shall at least be freed from the vain
search for the undiscovered and undiscoverable essence of the term species.
Having rejected specific essences at least three years before the ap-
pearance of Darwin's book, Mendel was able to concentrate his at-
tention on the behavior of single morphological features within any
cross. 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.
The results of Mendel's inquiries were published in a paper en-
titled " Experiments on Plant Hybridization." 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, 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, or to arrange these forms with certainty according to their separate
generation, or definitely to ascertain their statistical relations.
Commenting on this paragraph, 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. So far as I know, this conception
was absolutely new in his day.
12Gregor Mendel, " Versuche ueber Pflanzenhybriden," Proceedings of the Sci-
entific Society of Brunn, 1866: now available in Ostwald's Klassiker der Exacten
Wissenschaften, No. 121, and in English translation as an Appendix to Bateson's
book Mendel's Principles of Heredity (Cambridge, 1909).
13 Comments appear as footnotes to the English translation supervised by
Bateson.
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. 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, owing to their inability to recognize
clearly what observations were required. This is simply not true.
Neither hybridists nor breeders had been fumbling towards the goal
that Mendel reached, and it is a failure of historical insight to sup-
pose that they were. As we have already seen, their aims were quite
different.
Nor were the experiments which Mendel went on to describe like
those of his predecessors either. Both used the same technique of
cross-pollination, 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.
The novelty of his enquiry was something Mendel did not entirely
overlook. Although the whole monograph is only about twenty pages
in length, he took the trouble to include at the end a section com-
paring his results with those of Gartner, Koelreuter, and Wichura, in
which he noted that, as they had other aims, they did not always
report their results in a way that made close comparison possible.
Despite this, 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, since these hybrids were reported to
breed true, always producing offspring like themselves, without any
reversion to the forms of earlier generations.
The Immediate Reception of Mendet's Work
Mendel read his paper to the local Scientific Society at Brunn
(Brno), and it was published in their Proceedings for 1865, which
appeared in 1866. Copies of these Proceedings were sent to societies
and universities throughout Europe and America, so that, by the end
of 1866, Mendel's monograph was readily available in most of the
important centers of learning. Yet its significance was ignored.
The audience to whom Mendel first lectured comprised many
intelligent men of whom some were competent scientists, though
none were theoretical biologists of international repute. As a result
the paper got no advance publicity from members of the original
audience; further, the Brunn Scientific Society had been recently
formed and the volume of Proceedings containing Mendel's work was
only the fourth to be published. Publication in a new and obscure
journal (it might accordingly be argued) does not help to make a
man's work known, and we need look no further for an explanation
of the delay before Mendel's contribution was recognized. On the
other hand, it should be remembered that there were at this time
fewer scientific publications than now, and those were commonly of
a more general nature. " 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. Even today, an obscure format does
not often seriously delay recognition, and in any case (as we shall
see) Mendel sent copies of his paper to some of the leading botanists
of his day. More fundamental obstacles to recognition must there-
fore be sought. What these were, our study of the historical back-
ground of Mendel's work does something to suggest. To be of in-
fluence, a scientist's paper must be carefully read and judged im-
portant by his fellow-specialists. This will happen only if other
specialists are working on similar problems, or are working in fields
in which the paper in question appears relevant to an important
general problem. In Mendel's case, this was not generally so: horti-
culturalists and theoretical biologists alike were turning their at-
tention in other directions.
We have already seen that hybridists would tend to ignore
Mendel's paper, 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. In any case, hybridists
were by 1866 a dying race: after 1861 no more prizes were offered by
learned societies for work in this field. For while Mendel was busy
working, a revolution had occurred in biology: in 1859 Darwin had
published the Origin of Species.
By 1866 Darwin's work was well-known, both on the Continent of
Europe and in America. Many of the younger biologists had already
accepted both the idea of evolution and the theory of natural selec-
tion; the opposition of the churches had been voiced, the resulting
controversy was at its height, and in the dazzling light cast by the
new ideas the pressing problems of earlier generations sank into
insignificance. Most of those who accepted Darwin's thesis now set
themselves to trace evolutionary ancestries in the plant and animal
kingdoms, to add to the geological and geographical evidence for
evolution, or-particularly on the Continent-to study the implica-
tions of evolution for embryology. Since the opposition also con-
centrated on these aspects, there was a general re-orientation of in-
terest throughout biology.
Whereas before 1859 there had been no great theory or question
to direct attention toward the problems of heredity, there was now,
for converts to Darwinism, a positive swing away from them. Even
72 ELIZAETH B. 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. The Duke of Argyll14
drew attention to this fact in his criticism of Darwin, saying:
Strictly speaking, therefore, Mr. Darwin's theory is not a theory on the
origin of species at all, 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.
In a letter to Lyell, who had also voiced this opinion, 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. I could quote stronger expressions used by agricul-
turalists. Man does make his artificial breeds, for his selective power is of
such importance relatively to that of the slight spontaneous variations.
But no one will attack the breeder for using such expressions and the rising
generation will not blame me.
Darwin was right. The younger biologists did indeed agree with him
and for the time being attention was firmly focussed on natural
selection. The ability of living things to vary, and to hand on an
intensified version of these variations, was a presumption they ac-
cepted with natural piety. Little wonder that, in such an intel-
lectual climate, Mendel's paper was passed over by the rank and file
of biologists.
Yet even though the paper were not of general interest, there
might still have been individual biologists who would have appreci-
ated it, had they received copies of it. William Bateson, writing in
1902, 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.
Bateson may well have been right, 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, we may feel some doubt about Darwin's
power drastically to have altered the course of history; and in any
case there was no reason why Mendel should have thought in 1866
of sending a reprint to him. 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, Dec. 5,
1864; Proceedings, III, 35.
15 The Life and Letters of Charles Darwin, F. Darwin (London, 1883), II, 121.
16 Appendix to Bateson, Mendel's Principles of Heredity, ed. cit.
recipient, and it would perhaps have been embarrassing at this period
for Mendel, a monk, to start up a correspondence with the notorious
Darwin.
Mendel certainly sent reprints of his paper to two leading bi-
ologists, and may have sent copies to others. One of the known
recipients was A. Kerner, a famous hybridist and systematist. We do
not know whether Kerner acknowledged it: he never mentioned
Mendel in his writings, though there is some evidence that he was
aware of Mendel's later interest in the Hieraciums. The other re-
cipient was Professor Niageli of Munich: he did reply, and in fact
corresponded with Mendel for several years. Much of this interest-
ing correspondence survived and was published in 1905 by C. Correns,
a pupil of Niageli's. At first sight, Niigeli was just the man to under-
stand and appreciate Mendel's work. He had already been an evolu-
tionist before 1859, he was a knowledgeable systematist, and he
seems to have conducted breeding experiments. He read Mendel's
monograph with care and made notes on it. Yet, far from acclaiming
the paper, he replied gently urging Mendel to take up very different
work on the genus Hieracium.
As an evolutionist Naigeli was particularly interested in poly-
morphic genera, and did not rule out the possibility that some of
the intermediate species might have arisen through hybridization.
He thought, quite rightly, that these genera had never been properly
investigated. As he later explained:
In the year 1864, when Darwin's writings had made the problem of species
a burning one, I resumed my studies of the genus Hieracium, with special
reference to the phenomenon which might justify inferences regarding the
origin of varieties in species.
He chose this genus as being a polymorphic one with many integrates,
on whose many forms he was already an authority, having studied
them in detail some twenty years before. He was engaged on this
work when he received Mendel's monograph.
In a short letter accompanying the monograph, Mendel an-
nounced his intention of working with other plants and perhaps with
the Hieraciums. To this Niigeli replied enthusiastically:
It would seem to me especially valuable if you were able to effect hybrid
fertilisations in Hieracium, for this will soon be the group about whose in-
termediate forms we shall have the most precise knowledge.'7
He evidently recognized that Mendel was a patient and careful ex-
perimenter, and was pleased to have his collaboration. He even
17 C. Correns, " Gregor Mendels Briefe an Carl Naigeli," Proc. of Saxon Scientific
Society, XIX (1905), referred to below as 'Letters.' This comes from the first
letter from Nageli to Mendel.
offered to supply Mendel with material, and suggested suitable
species for him to begin on. Since all Naigeli's work on Hieracium
was undertaken to clarify the species question, he insisted, like the
older hybridists, that hybridization experiments in this field should
be made on wild and true species. Later, for instance, when Mendel
was engaged on this work, Nageli wrote:
I have also sent you H. prealtum and H. glaucum, both of them from
Isarkies near Munich. Prealtum because your H. prealtum (?) seems to
me suspect [i.e., may not be a 'true' species].18
So, in spite of the fact that Nageli did read Mendel's monograph
with some care, he seems to have misunderstood Mendel's aims, and
to have believed that the experiments described in it had the usual
aim of clarifying the traditional questions about species. If this was
the case, he must have regarded them as misconceived, since they
were conducted on cultivated plants which were not even distinct
species.
Nor did Niigeli believe that the laws Mendel put forward were
really valid. Perhaps this was because he would discount any results
obtained with cultivated varieties, or perhaps it was because his ex-
perience with Hieracium would certainly have given different results.
Either way, 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.
If the laws did not apply generally, there would be no reason for
taking the theory seriously either. Naigeli himself held a theory of
heredity of his own, which did envisage that characters were de-
termined by rudiments in the sex cells; but since the facts of
chromosome behaviour and reduction-division were unknown at that
time, he had no reason to postulate anything as implausible as the
segregation of these rudiments. The only reason for accepting such a
hypothesis at that time was that it would explain Mendel's results,
and these Niigeli could not take seriously. The notes N'ageli made on
the monograph contained the remark:
The constant forms require to be tested further. I expect that they would
be found to vary once more. 'A' [the pure dominant segregated out from
the selfed hybrid] has half 'a' in its body and when inbred cannot lose
that element.
To the suggestion that his 'pure forms' would not continue to re-
main constant, 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. It appears in the Life of Mendel by Iltis, p. 191.
during the five or six generations he had observed. He sent seeds
from such plants, which Niigeli sowed that summer, but the experi-
ment seems not to have continued, and neither side ever mentioned
the Pisum monograph in their subsequent correspondence.
In spite of Nageli's indifference, Mendel continued to experiment
widely, and in the next few years he worked on no less than twenty-
six different genera. Some of his results agreed with those which he
had obtained with Pisum. Others, however, did not. In particular,
with different-colored stocks, as with beans earlier, he found a great
range of color amongst the hybrids. 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. But
after this second discovery he began to doubt-needlessly, as we now
know-whether his theory was applicable in cases of this kind:
We thus obtain from the different colors, figures from which the deduction
of a developmental formula is impracticable.
Bit by bit the opposition of expert opinion was beginning to tell
on Mendel. He never bothered to publish any account of his newer
experiments, contenting himself with reporting the results from
time to time in letters to Nageli, and they became generally known
only after their eventual publication in 1905. No one in the 1860's
seemed interested in his laws or his theory; the later results had sug-
gested that the problem was not as simple as the pea experiments
had led him to believe; so perhaps he sought to turn his attention to
problems his colleagues were more interested in. At Brunn and else-
where, botanists were still hoping that hybridizing wild species might
throw light on the origin of intermediate forms in polymorphic
genera. As Niigeli had urged, Mendel now turned to this problem,
and in particular to the case of Hieracium.
The choice could hardly have proved more unfortunate, either
for practical or for theoretical reasons. Experiments with Hieracium
required enormous patience and skill, since the florets are extremely
small and the arrangement of the parts made the standard methods
of cross-pollination almost useless; further, however careful Mendel
was, some specimens seemed to be fertilizing themselves. His eye-
sight suffered, and by 1869 he had succeeded in producing only six
hybrid specimens: one of these was infertile, and the other five bred
true to their hybrid form in all subsequent generations, like the Salix
hybrids reported by Wichura. These results puzzled him, and con-
vinced him that the relations he had discovered for Pisum did not
hold here.
In retrospect, we can see why the work was so fruitless. The
genus Hieracium, like some other genera of the Compositae, is both
variable and atypical in its methods of reproduction. In some species
and specimens, fertilization may be normal, but quite frequently,
even if pollen is produced, there is no reduction division in the ovules
and no fertilization. When this happens, a new embryo may yet be
produced, developing apogamously from a diploid cell and not, in the
way it appears to do, as a product of sexual fusion: such an offspring
will be identical with its 'parent' cytologically. Most hybrids of
Hieracium do in fact reproduce apogamously, and therefore remain
constant in form.
Mendel could not recognize the deceptive character of his re-
sults, 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, in a lecture
which can be found in the subsequent volume of Proceedings. This
paper was very different from his earlier monograph, and must have
been much more intelligible to his contemporaries. 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.
There follows an account of the production of the six hybrids and
their behavior, to which Mendel adds, rather sadly:
I must express some scruples in describing in this place an account of ex-
periments just begun. But the conviction that the prosecution of the pro-
posed experiments will demand a whole series of years, 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.
He remarks how differently these new hybrids (and Wichura's Salix
hybrids) had performed from his own earlier Pisum hybrids, 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.
This Hieracium paper was really Mendel's botanical swan song.
He had been made abbot in 1868 and had less time for his hobbies.
He continued the Hieracium experiments for a while and still wrote
to Niigeli, reporting new results. In 1872, however, the strain on his
19 G. Mendel, " Experiments in Hieracium Hybridisation," Proc. Sci. Soc. Brunn
(1869).
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. Mendel now spent his
limited leisure on his bees and his meteorological studies.
Niigeli continued to write about heredity until his death in 1891,
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. d. Naturhistorischer
Ver. in Brunn, 1865, 69).
Apart from this, there was one other reference to the Pisum mono-
graph before 1900, in W. 0. Focke's work Die Pflanzenmischlinge,
published in 1881: this was the reference which led later workers to
the original monograph. Focke recorded Mendel's claim that there
was a constant numerical ratio between the various types of offspring
of the hybrid pea, but could not, when asked, remember how he came
to read Mendel's work. Except for these two inconspicuous refer-
ences, Mendel and his work sank into complete obscurity.
Science is organized knowledge, and no piece of work, however
complete in itself, is valued until it can be fitted into the general
corpus. Given the position in biology when Mendel wrote, it was
perhaps inevitable that his discovery should not have been appreci-
ated. This did not make it any the less of a personal tragedy for
Mendel, though he seems, at least in part, to have recognized the
situation and got some comfort from it. At any rate, his only re-
corded comment on the fate of his first monograph was to say: " My
time will come."
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. The additional pieces were, in fact, pro-
vided very quickly, for, towards the end of the nineteenth century,
the subject was developing at a great rate. Here we can outline only
the major discoveries and developments between 1865 and 1900, as a
result of which Mendelian genetics could take its proper place in the
structure of biology.
The most important change during this period was the advance in
cytology. The structure of the cell was observed in greater detail:
the process of cell division, in particular reduction division, and the
characteristic changes in the chromosomes in this process were studied
for the first time in the 1880's and the early 1890's. The actual fusion
of the two sex cells, though long suspected, was now observed for the
first time. Towards the close of the century, the haploid and diploid
phases were identified in the life-cycles of many animals and plants;
in this way it became clear that every sexual fusion was preceded by
a reduction division, the number of chromosomes in the cells of any
species so remaining constant from generation to generation. Dis-
coveries such as these altered completely the biologists' picture of
the nature of living things, and greatly increased their understanding
of the mechanism of sexual reproduction. Into this new picture
Mendel's theory fitted far more naturally. 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; the recognition
of the haploid gametes and the diploid zygote gave an indirect
plausibility to the idea of factor segregation; and though in 1900
this support was indirect, since the link between the factors and the
chromosomes was demonstrated experimentally only ten years later,
the new knowledge of chromosome segregation did at any rate make
factor segregation seem a possible vera causa.
These cytological discoveries redirected attention to the old prob-
lem, 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. Each type reproduced its kind; 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. This general
problem had occurred to biologists at many different periods in the
past, but now it reappeared in a different guise; not only was much
more known about the facts of sexual reproduction, but the problem
itself had changed in character. Whereas in the eighteenth century
the continuity of fixed types alone had to be explained, now every
type must be thought of as gradually altering. Biological theorists
therefore had to consider the possible causes of variations. Towards
the end of the nineteenth century, as the controversies about evolu-
tion became more intense, speculation about the causes of variation
also revived, and these speculations in turn drew attention to the
variations themselves, and to the facts about their inheritance.
Weismann's theory of the germ-plasm is the best-known of the
theories, but there were many others: Spencer's theory of physiologi-
cal units, Darwin's theory of pangenesis and Galton's theory of the
stipe were all produced at this time. There were many other such
theories, now almost completely forgotten.
While the majority of biologists in the 1860's were not (as we
have seen) interested in heredity, Darwin had surprised everyone by
producing in 1868 his Variations in Plants and Animals. In the in-
tellectual climate of the time, this book was poorly received, and
only the agricultural journals reviewed it favorably. The only bi-
ologist to approve of it was the systematist George Bentham, who
had always been interested in hybridization. The book, nevertheless,
did become quite influential. It was a fairly complete collection of
the known facts about the appearance and inheritance of variations,
particularly amongest domestic plants and animals. Such an as-
sembly of facts in one book, written by such a well-known author,
did something to interest biologists in problems which had previously
been considered only by practical breeders; so now that the theory
of heredity was beginning to receive attention, 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. Doubt
was now thrown on the old assumption that some acquired characters
could be inherited: once this was questioned by Galton, and later by
Weismann, a new impetus was given to scientific breeding. Long
and complicated experiments were conducted on animals, and knowl-
edge about inheritance gradually increased.
Galton's long-sustained concern with human inheritance did much
to awaken interest in the subject, especially in England. As he de-
veloped his novel 'biometric' methods of investigation, he drew into
that discussion other scientists and mathematicians who would other-
wise have had no more than a passing interest in biology. 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,
creating increasing public interest in heredity. In two crucial re-
spects, Galton's methods resembled Mendel's: he studied particular
characters, such as height, instead of the overall similarity and differ-
ence between individuals, and he analyzed his results statistically; 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.
The Last Phase
Work on cytology and biometrics did no more, however, than
create a situation in which it was at last possible for Mendel's theories
to be appreciated and understood. In order to see how it was that
his monograph was actually recognized by several different workers
almost simultaneously, one must look more closely at the situation
in the 1890's, in particular, at the work of a group of young biologists
who were leading a revolt against the orthodox Darwinian view of
the origin of species. Paradoxically it seems that, had Mendel's work
been lost forever, modern genetics would nevertheless be much the
same today, and that the rediscovery of his monograph had at most
the effect of aiding and speeding up the birth of the subject.
Darwin had originally spoken about variations with two different
voices, regarding them sometimes as being large and sudden, and
sometimes as small and continuous. Basically, however, it was for
him the small and continuous variations which provided the raw
material for evolution: natural selection, aided perhaps by the in-
heritance of acquired characters, gradually brought about a change
within any group, so that after a sufficient number of generations the
norm for that group had significantly altered. This view of the
evolutionary process was based on two assumptions which were first
questioned during the last decade of the nineteenth century. The
first assumption was contained in Darwin's idea of the nature of a
species; struck by the number of intergrades recently discovered be-
tween normal familiar types, and by the way in which the remaining
gaps could be narrowed by considering also the evidence from
geology, Darwin had assumed that living things formed a continuous
series of minutely different forms. Hence, for the early Darwinians,
species were in an important sense artificial. This emphasis was an
understandable reaction against the common idea of species as simple
natural units each with a distinct essence, but it proved to be an
overemphasis and distracted attention from the possibility that new
types might arise suddenly.
Darwin allowed, of course, that large and sudden variations might
occur, but he assumed that when discontinuous variations did appear
they would be swamped by interbreeding. This second assumption
was reasonable on the common view of inheritance: according to this
view, the material basis of inheritance might be particulate, but the
effects on the adult form of the individual particles were thought to
be mixed or 'blended.' Without the idea of 'segregation,' biologists
naturally thought that the particles mixed haphazardly during re-
production, the character appearing in the individual being a re-
sultant of the influences exerted by these assorted particles.
The revolt against this orthodox view of the evolutionary process
was led by three men-de Vries, Bateson, and Johannsen-all of
whom attacked it well before 1900. The research which their
theoretical objections led them to undertake was still in progress in
1900; it was this work which dovetailed so perfectly with Mendel's
to form the twin foundation-stones of modern genetics. Each of
them attacked the orthodox position from a different point of view.
Johannsen set out to show that the positive part of Darwin's picture
would not work; Bateson and de Vries both pointed out that Darwin's
belief in the continuity of living forms was exaggerated, Bateson go-
ing on to show that discontinuous variations which had arisen in the
past did in fact survive interbreeding, while de Vries looked for cases
in which present species could be observed giving rise to new forms.
Johnannsen investigated the question of the effects of natural
selection on a mixed group. He started with a mixed population of
beans and bred from the heaviest. 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. But if the breeder then
isolated these pure lines and allowed them to self-fertilize, further
selection had no influence: the size of the beans would fluctuate ow-
ing to external differences, for instance in the soil, but no further
selection of larger or smaller beans within this line would produce
offspring which differed in their average weight. (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.) Johannsen
published his results in 1903, 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.
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, 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.
He also attacked Darwin's conclusion that ' species' is an artificial
concept, on the grounds that distinct species do exist in nature or, at
the very least, that there are gaps in the sequence of living forms.
In order to collect further evidence about discontinuous variations,
he started breeding experiments, the results of which supported
Mendel's views; and, when the monograph was once again put into
circulation, he arranged for its translation into English, and did
much to publicize it.
De Vries, whose final work on mutation was published after 1900,
had actually started his work in the early 1880's. His ideas on
heredity were published in 1889, 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. According to his view,
organisms were built up of unit-characters, independently variable
and independently inheritable. 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, but which
together constituted the chromosomes. He regarded species as dis-
tinct types whose representatives normally interbred, so effecting a
reassortment of the unit-characters; but for one species to give rise
to another there must be a more drastic change, 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, during
which it would give rise by mutation to one or more mutant forms.
In a patch of Oenothera lamarckiana growing at Hilversum he
thought he had found a species in such a state of change, so he took
specimens of the new forms back to the University where he studied
their behavior and bred from them. While engaged in the resulting
breeding experiments, he hit for himself on the law of segregation;
he went on to test it for many different genera before publishing his
results, concluding that it held true for normal varieties, but not for
his mutants.
In March 1900, de Vries sent a short note entitled Sur la loi de
disjonction des hybrides to the French Academy of Sciences.20 This
was published on March 26th, 1900, but two days earlier on March
24th, he had read a paper Ueber das Spaltungsgesetz der Bastarde to
the German Botanical Society.2' This latter paper, which gave a
fuller account of the work reported in the French paper, contains-
at long last-a reference to Mendel's monograph and does justice to
the significance of his discoveries; after noting that his results agreed
with those which an Austrian monk, Gregor Mendel, had obtained
thirty-four years before, 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.
Meanwhile, in Germany, 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. At the same time
Tschermak, in Austria, also arrived at the same results while in-
vestigating Darwin's views about vigor in peas.
Correns received de Vries' French paper at the beginning of April,
and at once sent a paper to the German Botanical Society which
appeared in the April issue of their Journal. 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.... I believed myself to be an innovator.
Subsequently, however I found that in Brunn, during the sixties, Abbot
Gregor Mendel, devoting many years to the most extensive experiments on
peas, had not only obtained the same results as de Vries and myself but
20 Comptes Rendues de l'Acad. des Sciences (Paris, March 1900).
21 Ber. Deutsch. Bot. Gesellsch., XVIII, 3 (1900).
had actually given the very same explanation as far as was possible in the
year 1866.22
In June 1900 Tschermak also communicated his results to the Ger-
man Botanical Society: 23 he too, reported how he had first thought
he had made an entirely novel discovery, but had found that his
results had been anticipated when he later read Mendel's monograph.
From 1760 to 1860 many people had studied hybrids without
recognizing Mendel's laws; now that they were discovered, immedi-
ately hybridization experiments were restarted. This was partly due,
no doubt, to the fact that the later workers were more accustomed to
keeping numerical records and analyzing them statistically, and so
were more likely to notice numerical ratios in any result. But this is
evidently not the whole story. 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. Having arrived
at the laws, again, all of them were sufficiently familiar with the
controversy about the importance of variations to realize their great
importance and interest.
As soon as it was republished and translated, Mendel's work rap-
idly became famous, for his problems were the problems of the time.
This is not to say that his conclusions were immediately accepted, in
spite of the striking independent confirmation they had received.
But though 'Mendelism' was to be the subject of heated controversy
for more than a decade, the year 1900, in which the significance of
Mendel's monograph was finally recognized, certainly marks the be-
ginning of a new era in the growth of our understanding of heredity.
APPENDIX: The Argument of Mendel's Monograph
Mendel's Experiments and Theory. 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, the other remaining short; one plant having smooth seeds, the
other wrinkled ones; and so on. In every one of his seven groups he
obtained about 250 hybrids, each of which resembled one of the par-
ents, and the character which appeared in the hybrid he described as
the 'dominant' character of each pair. 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,
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. He extended these experiments to subsequent genera-
22 Ber. Deutsch. Bot. Gesellsch., XVIII, 4 (1900). 23 Ibid., XVIII, 6 (1900).
tions, 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, while the other
half yield plants which remain constant and receive the dominant or the
recessive characters (respectively) in equal numbers.
In the next set of experiments Mendel crossed plants which dif-
fered with respect to two, and in one case three, characters, and
studied their progeny. 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.
In the next section of the work, Mendel suggested a theory to
account for his laws. The theory indicated that the hybrids pro-
duced pollen grains and egg cells of two types. 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. On a
chance basis, there were three possible combinations which could re-
sult from the fusion of the pollen grains and the egg cells. If a pol-
len grain and an egg cell both bearing the factor for the dominant
character fused, the resultant offspring would share this character
with one of the original parents, and all subsequent generations bred
by self-fertilization would also resemble this stock. If a pollen grain
and an egg cell each bearing a factor for the recessive character fused,
the resultant offspring would resemble the other original parent, and
any subsequent generation produced as a result of a self-fertilization
would also resemble this stock. If unlike egg and pollen cells fused,
the result would be a hybrid, and this would continue to behave in
the same way as the original hybrids. So long as the different types
of reproductive cells were produced in equal numbers and provided
that sufficient cases were examined, the proportions of the progeny
should obey the law Mendel had discovered. It only required the ad-
ditional assumption that the factors segregated independently for
Mendel's second law to be explained.
In addition, Mendel described some experiments he had under-
taken with other species. He discovered that some of those behaved
like his peas, but with beans the flower colors did not follow the pat-
tern of his other results: the hybrids exhibited a range of colors, and
Mendel suggested that, in this case, the color might be determined
by more than one independent factor. In this way he came very near
to formulating the later 'multifactor' hypothesis.
University of Melbourne.

Why was Mendel's pioneering work on genetics initially ignored

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Why was Mendel's pioneering work on genetics initially ignored

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Why was Mendel's Work Ignored?

Author(s): 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

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

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-

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

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

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

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

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

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,

ing focus of theoretical biology.

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

matician's class of triangles is limited in advance by a formal defi-

nition of the properties of a triangle, the systematist's class of roses

is not thus limited in advance. 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. Being concerned with complex overall like-

nesses, the experienced systematist develops a 'feel' for the species

and sub-species he studies-to use an older terminology, he comes to

recognize the 'specific natures' or 'essences' of his species. Farmers

and stock-breeders, on the other hand, have a different problem. They

a higher sugar-content, or whatever it may be, 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.

Mendel's interest in inheritance was similar, and so differed funda-

mentally from that of other biologists. They were concerned with

the hybrid reflect the parental 'essences'. Mendel's contemporaries

therefore tended, either to misinterpret his work as a confused at-

tempt to investigate the nature of species, or else to dismiss it as

being irrelevant to their own crucial problem of the origin of species.

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. Men have practised selective

breeding of animals for many centuries, but during the eighteenth

century the 'agricultural revolution' drew new attention to the sub-

ject. Through the newly-formed local agricultural societies practical

ficial pollination of plants also had become possible, thanks to

Camerarius' work on the sexual process in the higher hermaphroditic

plants. This development was of great practical importance, since the

new types so produced could often be propagated vegetatively, and

it immediately attracted a considerable number of workers. Most

of these breeders, however, aimed merely to produce a plant or