Mendel’s Paper on the
Laws of Heredity (1866):
Solving the Enigma of
the Most Famous
‘Sleeping Beauty’ in
Science
Wolf-Ekkehard Lönnig, Max Planck Institute for Plant Breeding Research
(retired), Cologne, Germany
For more than 150 years now the enigma of the
disregard of the basic laws of heredity detected
by Mendel in the 1860s for at least 34 years (from
1865 to 1900) has inspired a large number of
conjectures and speculations. The most common
of these proposals have been briefly listed and crit-
ically assessed. However, the well-argued answer
given already at the beginning of the twentieth
century by several pioneers of genetics including
Correns, de Vries, Tschermak-Seysenegg, Bate-
son, Johannsen and others, and corroborated by
further biologists and historians of biology in
the more than one hundred years that followed,
has not been adequately considered so far in the
history of science in particular and in the public
eye in general: The failure to accept the elemen-
tal laws of heredity for decades was due to the
almost unlimited predominance of Darwin’s the-
ories on heredity and evolution. Darwin and his
followers believed in the inheritance of acquired
characteristics and blending inheritance as well
as continuous evolution. Mendel rejected all three
hypotheses. On the basis of hereditary constant
elements (which he assumed to be independent
of any environmental effects), he, in contrast,
concluded ‘that species are fixed within limits
eLS subject area: Science & Society
How to cite:
Lönnig, Wolf-Ekkehard (March 2017) Mendel’s Paper on the
Laws of Heredity (1866): Solving the Enigma of the Most
Famous ‘Sleeping Beauty’ in Science. In: eLS. John Wiley &
Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0026823
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Introductory article
Article Contents
• Introduction
• The Reaction in the Scientific World from 1866
to 1900
• Conjectures and Speculations for the Oblivion
of the Genetic Laws
• The Answer by Several Pioneers of Genetics
for the Disregard of Mendel
• Mendel on Constant Heritable Elements and the
Stability of Species
• Mendel and the Darwinian Revolution
• Evolutionary Theories before 1856
• The Reaction of the Darwinian Schools after
the ‘Rediscovery’ in 1900
• The Future of the Laws of Inheritance
Discovered by Mendel
Online posting date: 6th March 2017
beyond which they cannot change’ and completely
rejected (what we today call) Lamarckism. Since
there can be no doubt concerning Darwin’s over-
whelming victory in the battle for the scientific
minds in the nineteenth century, there was no
room left for the genuine laws of heredity until
1900. Their ‘rediscovery’ strongly reinforced the
eclipse of Darwinism until the establishment of
the modern synthesis in the 1930s and 1940s.
Introduction
A Sleeping Beauty (SB) in science refers to a paper whose
importance is not recognized for several years after pub-
lication. Its citation history exhibits a long hibernation
period followed by a sudden spike of popularity (Ke et al.,
2015).
In 1866 – in 2016 its 150th anniversary – Gregor Johann Mendel
published his epoch-making paper on Experiments in Plant
Hybridization (Versuche über Pflanzen-Hybriden), which he had
read at the meetings of 8 February and 8 March 1865 to the Nat-
ural History Society of Brünn and subsequently reported in the
Proceedings of the Society about a year later. (See Figure 1 for
a portrait of Mendel and Figure 2 showing a photograph of the
original first page of the Experiments in Plant Hybridization.) See
also: Mendel, Gregor Johann
The Reaction in the Scientific
World from 1866 to 1900
For the next 34 years the reaction of the world of science was
rather meagre – to say the least. Yet, the paper was mentioned
1
 Mendel’s Paper on the Laws of Heredity (1866): Solving the Enigma of the Most Famous ‘Sleeping Beauty’ in Science
Figure 1 Portrait of Mendel about 1882. Reproduced with permission from
Mendel Museum of Masaryk University, Brno. Edited by Roland Slowik,
Dietzenbach, Germany.
in some important treatises like Focke’s Die Pflanzen-Mischlinge
(1881), the Encyclopaedia Britannica (1881), and the Cata-
logue of Scientific Papers of the Royal Society of London (1879).
Among some 6000 other papers it was listed in Jackson’s Guide
to the Literature of Botany (1881). In 1869 it had already been
quoted by H Hoffman, Professor of Botany at the University of
Giessen in a book against the ‘Darwinian Hypothesis’, and 23
years later it was referred to twice by LH Bailey (1892) who had
assumed the chair of Practical and Experimental Horticulture at
Cornell University in 1888, as well as in the following PhD the-
ses: A Blomberg 1872 (in Swedish) and I Schmalhausen 1874 (in
Russian). In 1884 the Proceedings of the Horticultural Society
even enclosed the ensuing noteworthy statement about Mendel:
‘His experiments with plant hybrids have in fact opened a new
epoch, and what he has done will never be forgotten’ (Gustafsson,
1969). However, that epoch had still to wait until 1900 (although
Olby and Gautrey (2006) were able to list five additional refer-
ences before that year).
So, taken together, these few laboriously detected references
for a period of 34 years were almost nothing in comparison
with the explosion of hundreds of citations starting in the year
1900 – for a long list of authors and their papers, see Iltis (1924)
(e-book by Springer-Verlag 2007; English edition 1932/1966).
And, in the interim, the treatise must have counted thousands of
references up to now.
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It is also worth noting that Mendel had sent his paper to
the libraries of some 120 institutions including the Royal and
Linnean Societies of Great Britain. Moreover, Mendel had 40
additional reprints at his disposal, many of which he sent to
leading biologists of Europe in 1867. In fact, his mentor professor
Gustav Niessl von Mayendorf, secretary of the Brünn Society for
the Study of Natural Sciences, asserted that Mendel’s work was
‘well known’ during his time. And in the ensuing direct quotation
he also hinted at the major reason why it was ignored, saying:
‘His work was well-known, but owing to other views prevailing
at the time it was put aside’ (Niessl von Mayendorf, 1902, 1903;
translation by Orel, 1996, p. 275; or, in the German original text:
‘Man kannte seine Arbeiten sehr wohl’). As for the ‘other views
prevailing at the time’, Niessl specified: ‘ … at a time when for the
explanation of the origin of new forms of plants the principles
of the then generally acknowledged hypothesis of Darwin were
almost exclusively decisive’ (emphasis added in both quotations).
In spite of being well known, the accomplished fact remains
that the discovery of the laws of heredity was ‘put aside’, that is,
either totally ignored or wholly rejected as irrelevant for biology
in general and heredity in particular by most scientists for the next
34 years after Mendel’s publication. And, what is more, even after
1900, this epochal identification of the basic laws of heredity was
further on viewed to be only a special case of genetics in general
and completely pointless for the theory of evolution in particular
by the ‘true Darwinians’ (Mayr, 1982) for another 37 years. That
is roughly 71 years altogether.
To convey to the reader the magnitude of this most famous
sleeping beauty in science: Imagine for a moment that the same
would have happened to the publication of the double-helix
model of the DNA structure by Watson and Crick in 1953.
Envision that the discovery would have been ignored or silently
rejected by almost all contemporary scientists as largely irrelevant
for genetics, even to the point of being ‘ridiculed’ (as for Mendel,
see below), only to be ‘rediscovered’ in 1987 and then – in
contrast to all available evidence – would still have been viewed
as totally baseless and pointless for any evolutionary questions
right until 2014. See also: DNA Structure; Watson, James
Dewey; Crick, Francis Harry Compton
Conjectures and Speculations
for the Oblivion of the Genetic
Laws
Now, up to the present, there has been an enormous amount
of conjectures and speculations regarding the reasons for this
strange neglect of Mendel’s work, for example (first the doubtful
assertion followed in brackets by a counter argument):
1. Mendel was an outsider (this could perhaps also be said
of Darwin (Wuketits, 2015) who after two years of stud-
ies dropped out of medical school in Edinburgh and then
decided to become a clergyman, enrolling at Christ’s Col-
lege, Cambridge, for the necessary BA – the first step to
prepare him for a career in the Church of England. Also,
‘Darwin was very much an amateur when he started on
 Mendel’s Paper on the Laws of Heredity (1866): Solving the Enigma of the Most Famous ‘Sleeping Beauty’ in Science

Figure 2 Photograph of the first page of the original manuscript of Versuche über Pflanzen-Hybriden (Experiments in Plant Hybridization) (1866). Reproduced
with permission from Mendel Museum of Masaryk University, Brno. Edited by Roland Slowik, Dietzenbach, Germany.

the voyage’ of the Beagle; he was not a ‘finished natural-
ist’ according to Henslow, who nevertheless recommended
him for the Voyage (cf. Himmelfarb, 1959/1996, pp. 42,
53, 82). So, why was Darwin so successful but Mendel
was not? The outsider argument appears to be doubtful for
both Darwin and Mendel (all the more so considering that
Mendel taught biology and physics at Brünn Oberrealschule
for 14 years)). See also: Mendel, Gregor Johann; Darwin,
Charles Robert
2. His seclusion in the monastery (in reality there was an
active group of different research professors affiliated with
the monastery he continually interacted with; plant hybrids
was a lively subject at Mendel’s place and time (Niessl von
Mayendorf, 1912, p. 304), also Darwin and evolution (Iltis,
1924, p. 119)).
3. He was toiling entirely unattended in a scientifically iso-
lated atmosphere (in contrast, Mendel regularly showed his
‘children’ – as he called his plants – to his many scientific
friends and likewise to his visitors from outside; moreover,

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 Mendel’s Paper on the Laws of Heredity (1866): Solving the Enigma of the Most Famous ‘Sleeping Beauty’ in Science
Figure 3 Mendel – second row from above and also second from
right-hand side – with many persons around him in Paris 1862 (although
I could not obtain a photograph with a higher pixel solution, the figure, an
excerpt from a larger picture, shows nevertheless that Mendel did not travel
alone from Paris to London). From Oswald Richter (1931). Photographer of
1862 unknown. Edited by Roland Slowik, Dietzenbach, Germany.
the notion of a solitary monk in his secluded abbey is some-
what contradicted by his many travels being usually accom-
panied by his faithful servant Josef and often also several
other travel companions: in Austria of his time (to Vienna,
Salzburg, Troppau, Ölmütz, Prerau), England (London),
France (Strasbourg, Paris), Germany (Munich, Stuttgart,
Karlsruhe, Bruchsal, Hamburg, Eystrup, Kiel, Köln), Italy
(Meran, Florenz, Rome), and so on altogether 31 voyages;
details in Richter, 1931). See Figure 3 for Mendel in Paris
1862 with many persons around him.)
4. He published his paper in an ‘obscure journal’ (however, the
Proceedings were an accepted quotable journal at his time
as shown by the references above).
5. His strange style of the paper (botany in connection with
mathematics) (yet, this had really been practised for decades
before him in plant physiology; see, for example, Stephen
Hales already in 1727).
6. Mendel’s modesty (nevertheless, he was also very patient,
persevering and even tenacious, sending reprints to more
than 160 institutions and/or leading scientists of his time
[probably even to Darwin; Galton, 2009, 2015, p. 4; and
according to Blumberg (1997/2010) of the MendelWeb ‘one
[reprint] even found its way into the library of Charles
Darwin. We know that Darwin did not read Mendel’s paper
(the pages were uncut at the time of Darwin’s death)’). In
our computer age, some people may be hardly aware of how
much time and hassle was necessary in 1867 to accomplish
such a postal task).
4 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
7. Chromosomes were not yet detected (they were, in fact,
detected but only partially understood by Nägeli in 1842,
and Hofmeister in 1848; however, the acceptance of the
fundamental laws of heredity did not depend on the iden-
tification and function of the chromosomes: intriguingly,
Boveri’s key discovery of the significance of chromosomes
for heredity in 1880 were not followed by the recognition
and acceptance of the laws detected by Mendel).
8. Mendel himself was not convinced of the universality
of his discoveries (yet, Mendel had extended his Pisum
investigations to a series of further plant species: Aquilegia,
Antirrhinum, Calceolaria, Campanula, Carex, Curcur-
bita, Dianthus, Geum, Hieracium, Ipomea, Lathyrus,
Linaria, Lychnis (Melandrium), Matthiola, Mirabilis,
Pirus, Potentilla, Prunus, Sedum (?), Tropeolum, Verbas-
cum, Veronica, Viola and Zea (Iltis, 1924, p. 103). In spite of
some troubles with Hieracium due to agamospermy (asex-
ual reproduction/clonal reproduction of seeds, discovered
only much later by Hans Winkler in 1908), it is reported
of ‘The Father of Genetics’ to have repeatedly said: ‘My
time will come’. Also, ‘I am convinced that it will not take
long before the whole world will acknowledge the results
of these studies’; Mather, 1966).
9. Mendel’s segregation ratios were too good to be true; he
invented all his data. (In contrast, the many eyewitnesses
of his experiments never doubted the reality of his investi-
gations; keen observer Niessl von Mayendorf (1903, p. 20)
spoke of the ‘precious results of protracted and extremely
careful experiments’. Klein and Klein (2013, p. 2) called
this criticism ‘nonsensical’, and Klein (2014) added on the
‘too good to be true’ objection: ‘There is no basis [for this]
anymore, absolutely no basis’, providing further arguments
against Fisher who had first raised this challenge.)
10. As shown by several recent investigations (Ke et al., 2015,
there further references), ‘sleeping beauties’ are not the
isolated exceptions in science as once thought to be but
occur regularly, for example, in physics (multidisciplinary)
in 7.6% of the publications and in plant sciences in 1.3%.
So it could perhaps be argued that Mendel‘s paper was just
one of them. (However, this answer would be totally irrele-
vant for a time in which Darwin himself as well as almost
all his scientific disciples were intently searching for the
discovery of the laws of heredity and had also produced
strongly Lamarckian-like hypotheses of their own concern-
ing that open question. So Mendel’s discoveries were not
just unknown but deliberately rejected and hence kept dead
quiet.)
Thus, all the conjectures just mentioned are either irrelevant
or definitely wrong as explanations for Mendel’s sleeping beauty
(for a detailed overview of some of the more relevant pros and
cons, see Lönnig, 2005a).
After all, the question remains to be solved what the real rea-
sons were for this strange phenomenon occurring, to repeat and
emphasise this point: at a time when almost the entire scientific
world was – in the wake of Darwin’s Origin – strongly motivated
to search for and detect the scientific explanation of heredity.
 Mendel’s Paper on the Laws of Heredity (1866): Solving the Enigma of the Most Famous ‘Sleeping Beauty’ in Science
The Answer by Several Pioneers
of Genetics for the Disregard
of Mendel
The answers given by de Vries (1901/1903); Bateson (1904,
1909, 1924/1928); Correns (1905); Baur (1911); Nilsson-Ehle
(1911); Johannsen (1909, 1926), as well as several historians
of biology and/or biologists as Niessl von Mayendorf (1903,
1906); Radl (1909); Iltis (1924); Richter (1941, 1943); von
Tschermak-Seysenegg (1951); Nilsson (1953); Stern (1962);
Stubbe (1963); Clark (1967); Krumbiegel (1967); Weiling
(1976); Callender (1988); Strickberger (1988); Bishop (1996);
Sermonti (2005); Nelson (2009); Speicher et al. (2010) and sev-
eral further authors, are in part combined and condensed in the
ensuing summary (for a full documentation of their statements,
see Lönnig, 2005a):
All the testimony and documentation of these authors leads to
the basic cause of the rejection of Mendel’s discoveries as fol-
lows: His analysis, discernment and exposition of the laws of
heredity as well as his views on evolution diametrically defied
and contradicted the ideas and convictions of Darwin and his fol-
lowers. Darwin combined his natural selection theory with his
starkly held view of the inheritance of acquired characteristics
(by blending inheritance) running through all his evolutionary
publications (and he tried to support his doubtful ideas on hered-
ity with his pangenesis hypothesis, which even the co-founder
of the synthetic theory (neo-Darwinism) in botany George Led-
yard Stebbins called (1977, p. 14) an ‘unfortunate anomaly’ and
molecular evolutionist and neo-Darwinian Jan Klein (2014) to be
‘completely wrong’). But perhaps even more important, Mendel’s
discoveries cast doubt on another definitely decisive and essential
part of Darwin’s theory: continuous evolution, for which Darwin
had postulated ‘infinitesimally small inherited variations’, ‘steps
not greater than those separating fine varieties’ and ‘insensibly
fine steps’, ‘for natural selection can act only by taking advantage
of slight successive variations; she can never take a leap, but must
advance by the shortest and slowest steps’ (emphasis added; see
for the references Darwin online). See also: Bateson, William;
Vries, Hugo de; Johannsen, Wilhelm Ludwig; History of Clas-
sical Genetics
However, modest Mendel, having demonstrated the laws of
heredity by virtually seven all-or-nothing traits (pea seeds, to
simplify somewhat: either round or wrinkled, seed colour either
yellowish or green, stem either long or short, etc.), in fact, rejected
both, the inheritance of acquired characteristics – which hypoth-
esis he had investigated for years (inter alia with Ficaria ranucu-
loides and F. calthaefolia) – as well as Darwinian evolution.
Mendel on Constant Heritable
Elements and the Stability
of Species
The laws of inheritance revealed by him were perceived to be the
laws of constant elements – stable in time, as he had emphasised
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in his correspondence with Nägeli – and relevant for both, gen-
erating a great but nevertheless finite variation in the culture
varieties of the plant breeders and equally significant for and
applicable to an abundant yet nonetheless limited divergence of
species in the wild (Mendel, 1866, pp. 36, 46, 47). In his brief
paper of 45 pp (pp. 3–47) on Experiments in Plant Hybridization,
Mendel perpetually speaks of ‘constant characters’, ‘constant
offspring’, ‘constant combinations’, ‘constant forms’, ‘constant
law’, ‘a constant species’, etc. (in such combinations, the adjec-
tive ‘constant’ occurs altogether 67 times in the German original
paper). He was convinced and also directly stated that the laws of
heredity he had discovered substantiated Carl Friedrich von Gärt-
ner’s conclusion (1848) ‘that species are fixed with limits beyond
which they cannot change’ (Mendel, 1866, pp. 46/47). And as
Theodosius Dobzhansky, perhaps the foremost founder of the
neo-Darwinian synthesis, aptly put it (1955, p. 183): ‘It is … not
a paradox to say that if someone should succeed in inventing a
universally applicable, static definition of species, he would cast
serious doubts on the validity of the theory of evolution’.
To quote Mendel on Gärtner directly (1866, p. 46):
‘Gärtner by the results of [his] transformation experi-
ments, was led to oppose the opinion of those natural-
ists who dispute the stability of plant species and believe
in a continuous evolution of vegetation. He perceives in
the complete transformation of one species into another
an indubitable proof that species are fixed within limits
beyond which they cannot change. Although this opin-
ion cannot be unconditionally accepted we find on the
other hand in Gärtner’s experiments a noteworthy confir-
mation of that supposition regarding variability of culti-
vated plants which has already been expressed.’ (Italics
added.)
Lenval A Callender of the London University Institute of Edu-
cation comments on this passage as follows (1988, p. 54):
‘Despite its clarity this paragraph has been a source of
endless confusion in the literature. If this statement is to
be taken literally, as Mendel most assuredly intended it to
be taken, then it says quite simply that he gave conditional
acceptance to the view, expressed by Gärtner, "that species
are fixed within limits beyond which they cannot change."
Nothing could be clearer. Nevertheless, interpretations of
this passage have been given which are remarkable for
their extreme departure from accepted use in both the
German and English languages’ (italics by Callender).
Bishop (1996, p. 208) concurs with and corroborates Callen-
der’s conclusion by further arguments. See also: von Gaertner,
Carl Friedrich
Also, Mendel – writing on the variability of cultivated
plants – confirmed his view on the stability of species by the
ensuing deeply thought out comment (1866, p. 36; italics added):
‘The opinion has often been expressed that the stability of the
species is greatly disturbed or entirely upset by cultivation, and
consequently there is an inclination to regard the development
of cultivated forms as a matter of chance devoid of rules; the
colouring of ornamental plants is indeed usually cited as an
5
 Mendel’s Paper on the Laws of Heredity (1866): Solving the Enigma of the Most Famous ‘Sleeping Beauty’ in Science
example of great instability. It is, however, not clear why the
simple transference into garden soil should result in such a
thorough and persistent revolution in the plant organism. No one
will seriously maintain that in the open country the development
of plants is ruled by other laws than in the garden bed. Here,
as there, typical changes must take place if the conditions of
life be altered, and the species possesses the capacity of fitting
itself to its new environment. It is willingly granted that by
cultivation the origination of new varieties is favored, and that by
man’s labour many varieties are acquired which, under natural
conditions, would be lost; but nothing justifies the assumption
that the tendency to formation of varieties is so extraordinarily
increased that the species speedily lose all stability, and their
offspring diverge into an endless series of extremely variable
forms.’ Thus, in Mendel’s view, endless evolution was neither
probable for cultivated plants nor for species in the wild (see
for further support and scientific expansion of Mendel’s views
Lönnig, 1993, 2005b, 2011, 2012, 2014).
Mendel and the Darwinian
Revolution
According to many contemporary biologists, Mendel’s seminal
genetic paper was written and published at a time of a far greater
revolution in the history of science in general and of biology in
particular: Darwin’s Origin of Species. The ensuing metamorpho-
sis of ideas in biology and of world view in general was raised by
Ernst Mayr (winner of the Swedish Crawford Prize equivalent
to the Nobel Prize) to be ‘perhaps the most fundamental of all
intellectual revolutions in the history of mankind’ (Mayr, 1972,
similarly 2000, 2009; and in agreement with him many renowned
authors, like Lewontin, Gould, and Dawkins). For most of them
the accompanying change in worldview is seen to be not only the
most fundamental but also the best that could have happened to
mankind as a whole. See also: Mayr, Ernst Walter
Thus, according to the authors mentioned before, the basic rea-
son for the neglect of the laws of heredity was essentially this: To
imply something like a static definition of the species by constant
hereditary elements right into a momentous process vigorously
favouring the Darwinian revolution (continuous evolution by nat-
ural selection without any teleology intimately combined with
the inheritance of acquired characteristics, to underscore the lat-
ter, often forgotten point once more) was met – although usually
silently – with skepticism, deliberate ignorance and strong oppo-
sition. And there is no doubt concerning Darwin’s overwhelming
victory in the battle for the scientific minds in the nineteenth cen-
tury, so much so that Mendel’s performance before the Natural
History Society of Brünn was even met with ‘scornful laughter’
(according to the information by Makowsky, one of Mendel’s
associates, to Iltis personally).
This historically most important answer can be viewed,
strangely enough, to be another sleeping beauty (sensu lato)
largely ignored by science for more than a hundred years now.
It was given by the pioneers of genetics at the beginning of the
twentieth century and has been hinted at several times indepen-
dently by some biologists and historians of science during the
6 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
following century. However, for most Darwinians it is almost
impossible to think, and much less to accept, that a man of
scientific stature like Mendel (belonging to the few names ‘that
stick out far above the Nobel Prize standard’, Klein and Klein,
2013, p. 2) could have doubted or even denied Darwinism and
evolution. Au contraire, they tried to monopolise Mendel for their
views. Referring to de Beer, Iltis and Fisher, Olby commented:
‘As Darwinians these authors were keen to exhibit Mendel as
a supporter of Darwin’, – subsequently refuting such claims
(Olby, 1997). Hence, intriguingly, the essential resolution of the
question has again not been generally accepted by the scientific
community for reasons clearly related to those of the oblivion or
rejection of Mendel’s paper at his time. (The answer concerning
Darwin was questioned recently again by Klein and Klein (2013)
and Klein (2014); see below.)
The essence of the positive conviction concerning the Dar-
winian revolution has concisely and perhaps best been summed
up in the following words of the co-founder of the modern synthe-
sis Mayr (2009) – and note, please, the deep ideological contrast
to the implications of Mendel’s findings:
‘First, Darwinism rejects all supernatural phenomena and
causations. The theory of evolution by natural selection
explains the adaptedness and diversity of the world solely
materialistically. It no longer requires God as creator
or designer [ … ]. Darwin pointed out that creation, as
described in the Bible and the origin accounts of other cul-
tures, was contradicted by almost any aspect of the natural
world. Every aspect of the “wonderful design” so admired
by the natural theologians could be explained by natu-
ral selection. [ … ] Eliminating God from science made
room for strictly scientific explanations of all natural phe-
nomena; it gave rise to positivism; it produced a powerful
intellectual and spiritual revolution, the effects of which
have lasted to this day.’ See also: Evolutionary Ideas:
The Modern Synthesis
Guiseppe Sermonti, retired professor of Genetics at the Univer-
sity of Perugia, sums up the contrast between Mendel and Darwin
in the following words (2005, p. 46):
‘What really happened was that Mendel ruled out almost
all the forces that Darwin had invoked to explain evo-
lution. For Darwin, heredity was the result of a mixing
of seminal fluids [ … ] Mendelian heredity consists in a
recombination of traits that associate with each other but
do not blend, with the result that a variant trait always has a
chance of reemerging. Darwin’s mixture theory [ … ] was
a necessary corollary to his conviction that the environ-
ment acted directly on the germinal fluids inside the body
[ … ] Mendel’s hereditary determinants were not amenable
to such influences and transmissions. They were static, per-
manent, and fully indifferent to the environment. In some
quarters their stability was considered reactionary - clerical
even, given that they came from the garden of an abbot.’
Nevertheless, it is hardly worth noting that the acknowledge-
ment, approval and acceptance of reproducible facts of science
should be entirely independent of any biases of religion and
worldview.
 Mendel’s Paper on the Laws of Heredity (1866): Solving the Enigma of the Most Famous ‘Sleeping Beauty’ in Science
Answering Klein (2014), it may be true, of course, that Mendel
may not have read Darwin before 1860 (when the first German
translation of Darwin’s Origin appeared), but in formulating
his treatise Versuche (1865 and 1866) he most probably has
also considered Darwin’s work. According to Galton (2009),
Mendel read the Origin in 1863, but as reported by Henig (2001,
p. 167) in her more popular yet quite well-researched book,
already in 1860. Moreover, at the 11 January 1865 meeting
preceding Mendel’s first presentation, the botanist and geologist
Alexander Makowsky had talked with utmost enthusiasm about
Darwin’s theory (schwungvoll und begeistert), overshadowing
Mendel’s contribution as stated by Iltis (1924, p. 119). Mendel
collected, indeed, all of Darwin’s books and Iltis commented
(p. 66): ‘Mendel bought all the works of Darwin right after
their publication and it touches us peculiarly to detect at the
monastery’s library almost the entire Darwinian literature of
the sixties and seventies’, yes, even two sets of the Origin and
Variation of Animals and Plants, additionally the Zoonomia of
Darwin’s grandfather Erasmus Darwin. Also, there were many
evolutionary books and commentaries (Büchner, Vogt and others)
in Mendel’s library being on the Index librorum prohibitorum,
which was obviously largely ignored by Mendel.
Nevertheless, as all knowledgeable authors agree, ‘the idea of
evolution had been widespread for more than one hundred years
before 1859’ (Mayr, 1972). And, looking more closely at this
question, we can extend that period before 1859 to at least 150
years further back (not to speak of Heraklit, Empedokles, Epikur
and further Greek philosophers up to ca. 2500 years ago). And
there are convincing reasons to assume that Mendel had already
had a deep knowledge of the history of evolution when he started
his experimental work with peas and further plants (see below).
Evolutionary Theories before 1856
So what was known about the topic of evolution when Mendel
started his experiments in 1856? A few examples:
Erasmus Darwin’s Zoonomia (1794; German edition
1795–1799) was in Mendel’s library (as mentioned). Inter-
estingly, George Bernard Shaw named Erasmus as one of the
founders of the theory of evolution. To convey a taste of the
evolutionary contents of this work, consider just one quotation
from Vol. 1 (I must admit that I was immediately struck by the
similarity to Charles Darwin’s style of 1859):
‘From thus meditating on the great similarity of the struc-
ture of the warm-blooded animals, and at the same time of
the great changes they undergo both before and after their
nativity; and by considering in how minute a proportion
of time many of the changes of animals above described
have been produced; would it be too bold to imagine,
that in the great length of time, since the earth began to
exist, perhaps millions of years before the commence-
ment of the history of mankind, would it be too bold to
imagine that all warm-blooded animals have arisen from
one living filament, which THE GREAT FIRST CAUSE
endued with animality, with the power of acquiring new
parts [ … ] and thus possessing the faculty of continuing
to improve by its own inherent activity, and of delivering
eLS © 2017, John Wiley & Sons, Ltd. www.els.net
down those improvements by generation to its posterity,
world without end!’ (Italics added). See also: Darwin,
Erasmus
In 1809 Lamarck published his Zoological Philosophy. An
Exposition with Regard to the Natural History of Animals (origi-
nal title: Philosophie zoologique, ou Exposition des considéra-
tions relatives à l’histoire naturelle des animaux). See also:
Jean-Baptiste Lamarck
Another climax in the history of the widespread ideas on evolu-
tion before Charles Darwin is The Cuvier–Geoffroy Debate in the
years 1820–1829 (see Appel, 1987, with ‘the most thorough anal-
ysis we have of the controversy at issue,’ according to Science).
See also: Jean-Baptiste Lamarck
Also, in this context it seems to be appropriate, if not manda-
tory, to mention the book of the botanist Carl Friedrich von Gärt-
ner of 1849 Versuche und Beobachtungen über die Bastarderzeu-
gung im Pflanzenreich. The author was quoted by Mendel 17
times directly in 1866 and before that 18 times by Darwin in
the first edition of the Origin (1859) (speaking of Gärtner as
‘that most careful observer’) and later altogether 88 times in Dar-
win’s two volumes on The Variation of Animals and Plants under
Domestication’ (1868). In his book Gärtner had given an impres-
sive in-depth discussion of the different evolutionary authors and
their hypotheses up to his time, and, what is more, presented
a range of clear counterarguments against these views. Thus, it
appears that Mendel was best informed about the pros and cons
of almost all the contemporary evolutionary theories being per-
petually discussed in science for decades before he started his
own experiments.
Although there are many further highlights and issues that
could be discussed for the long history of evolutionary theories
before Darwin, I would like to mention only a fifth strong peak
here: Robert Chamber’s Vestiges of the Natural History of Cre-
ation of 1844, translated and published in German in 1851. See
also: Evolution: History; Evolutionary Ideas: Pre-Darwinian
Thus, it would be absolutely impossible to argue that Mendel,
the indefatigable collector of evolutionary treatises, would have
known nothing of the long history of evolutionary ideas when
he started his experiments for the explicit reason, as he stated
in the introduction of his paper, that his labour appears ‘to be
the only right way by which we can finally reach the solution of
a question the importance of which cannot be overestimated in
connection with the history of the evolution of organic forms’.
One may also surmise that Mendel visiting the Great London
Exposition, the World’s Fair of 1862, could at least understand
and read some English.
The Reaction of the Darwinian
Schools after the ‘Rediscovery’
in 1900
As might have been expected by the information given above,
the ‘rediscovery’ of Mendel’s work in 1900 was accompanied
by a forceful opposition of the Darwinian schools of biology
against the validity of the newly detected paper as well as the
7
 Mendel’s Paper on the Laws of Heredity (1866): Solving the Enigma of the Most Famous ‘Sleeping Beauty’ in Science
basic laws of heredity. ‘The controversy became so bitter that
in 1903 the British periodical Nature closed its columns to
the Mendelians. The columns of Biometrica had already been
closed to them’ (Zirkle, 1964, p 68). An overview of the dra-
matic controversy between the Mendelians and Darwinians, espe-
cially between Bateson and his Darwinian opponents, has been
given by, for example, Bateson himself; see Bateson (1909,
1924/1928); Provine (1971); Bowler (1992); Lönnig (2005a) and
Richmond (2006). ‘So unexpected was the discovery that many
naturalists were convinced it was untrue, and at once proclaimed
Mendel’s conclusions as either altogether mistaken, or if true, of
very limited application’ (Bateson, 1909).
The following years were characterised by an enhanced
‘eclipse’ of Darwinism (as Julian Huxley first published the
term, perhaps following Jordan), so much so that in 1909 many
scientists celebrating Darwin’s 100th birthday and the 50th
anniversary of the publication of the Origin raised skepticism
concerning the validity of natural selection and rejected pan-
genesis in any case. Regarding Darwinism, critics correctly
noted ‘that it was non-experimental, nontestable, and ultimately,
speculative’ [Allen 2000, according to Richmond (2006, p. 451);
as for different views on the eclipse of Darwinism, compare Kel-
logg (1907) on Darwinism attacked and defended, the excellent
perhaps unsurpassed comments of the European biologist and
close eyewitness Radl (1909, pp. 539–569), on the ‘Decline of
Darwinism’, and also Largent (2009), who argues that the eclipse
never happened]. See also: Evolutionary Ideas: The Eclipse of
Darwinism
However, later in the 1930s and 1940s ‘ … Darwinism even-
tually emerged from its eclipse once it could be shown that a
more sophisticated interpretation of the new genetics would pro-
vide a firmer foundation for selection’ (Bowler, 1992, p. 14).
Or, in the words of the unequalled assessment of Gould (2002,
pp. 569/570): ‘I can imagine no contrast more stark, no reversal
so complete, as the comparison of these doubts in 1909 and the
confidence and near unanimity expressed fifty years later at the
Origin’s centennial in 1959. The success of the Modern Synthesis
established the difference. Beginning as a pluralistic marriage of
Darwin and Mendel in the 1930s the Synthesis had hardened by
1959 into a set core of commitments that, at least among epigones
and acolytes, had become formulaic and almost catechistic, if not
outright dogmatic.’ See also: Gould, Stephen Jay; Evolutionary
Ideas: The Modern Synthesis
The Future of the Laws
of Inheritance Discovered
by Mendel
In our era of molecular biology marked by polyfunctional DNA
sequences, split genes, generations of different proteins by alter-
nate mRNA splicing, prions, horizontal gene transfer, transpos-
able elements, epigenetics, and so on, the question may be raised
as to what extent the laws discovered by Mendel are still rel-
evant for biology in general and medicine in particular and
whether there may be also future prospects for them. The ques-
tions can easily be answered: Google, please, OMIM (Online
8 eLS © 2017, John Wiley & Sons, Ltd. www.els.net
Mendelian Inheritance in Man: at present more than 20 000
entries; edited at John Hopkins University School of Medicine)
or OMIA (Online Mendelian Inheritance in Animals, again with
thousands of entries; database of genes of more than 135 animal
species, authored by Frank Nicholas, Sydney, Australia). Both
sites are regularly updated. See also: Mendelian Genetic Dis-
orders; Transposons in Eukaryotes (Part B): Genomic Con-
sequences of Transposition; Epigenetic Regulation in Plants
And, last but not least, I would also like to mention a perhaps
future Mendelian project which so far has hardly been adequately
touched in genetics. The Austrian-Swedish geneticist Herbert AK
Lamprecht – working inter alia on the nature of species and
interspecific barriers at the famous Weibullsholm Plant Breeding
Institute, Landscrona, Sweden, with Pisum and several other plant
species – detected that in unique crosses between different, but
closely related species there was an unbridgeable barrier for the
segregation and recombination of special features – depending
on the cytoplasmic constitution of the mother species, which
could neither express nor recombine these unique characteristics
in a homozygous state together with fertility – producing in the
simplest case a 1 : 2 : 0 ratio instead of the usual 1 : 2 : 1
segregations as expected. For an in-depth discussion of the pros
and cons, see Lönnig (1993, pp. 210–270). If the approach is at
least basically correct, it would complete and extend Mendel’s
path of the genetic laws within species to a new non-Darwinian
testable theory on the origin of species.
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