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Hist. Phil. Life Sci.

, 33 (2011), 3-48

What Would Have Happened if Darwin Had Known
Mendel (or Mendel’s Work)?

Pablo Lorenzano

Centro de Estudios e Investigaciones
National University of Quilmes
National Council of Scientific and Technical Investigations
Roque Sáenz Peña 352 (B1876BXD)
Bernal, Argentine

Abstract – The question posed by the title is usually answered by saying that the “syn-
thesis” between the theory of evolution by natural selection and classical genetics, which
took place in 1930s-40s, would have taken place much earlier if Darwin had been aware
of Mendel and his work. What is more, it nearly happened: it would have been enough if
Darwin had cut the pages of the offprint of Mendel’s work that was in his library and read
them! Or, if Mendel had come across Darwin in London or paid him a visit at his house in
the outskirts! (on occasion of Mendel’s trip in 1862 to that city). The aim of the present pa-
per is to provide elements for quite a different answer, based on further historical evidence,
especially on Mendel’s works, some of which mention Darwins’s studies.

Keywords – Darwin, Mendel, hybridism, speciation, evolution


The question posed in the title is usually answered, counterfactually,
by claiming some developments in evolution theory that did not occur
until the 20th century would have occurred much sooner, if Darwin
had been aware of Mendel. William Bateson (1861-1926) was an early
adherent to this notion when, in the introductory note to the first trans-
lation of Mendel’s work in English, he stated: “Had Mendel’s work
come into the hands of Darwin, it is not too much to say that the his-
tory of the development of evolutionary philosophy would have been
very different from that which we have witnessed” (Bateson 1901). This
opinion was to be repeated by Bateson (1902; 1909) as well as by Iltis
(1924) in the first comprehensive biography of Mendel. Later, in light
of developments in biology during the 20th century, the interpretation
was slightly modified; that is, it was claimed that the synthesis between
the theory of evolution by natural selection and classical genetics, the
© 2011 Stazione Zoologica Anton Dohrn

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evolutionary synthesis of the 1940s, would have occurred much before
this time.
Moreover, it is conventionally claimed that Darwin could have been
exposed to Mendel if he had only cut the pages of the offprint of Mendel’s
work that was in his library. According to this position, Darwin would
have immediately noticed that Mendel provided the mechanisms of in-
heritance that his theory of evolution required and that he had attempt-
ed to develop in his provisional hypothesis of pangenesis (Darwin 1868).
Or, the two could have met if Mendel had either met Darwin in London
or paid a visit to his house, when Mendel visited England. However,
no face-to-face encounter occurred. As Richter states (1932; 1942), al-
though Mendel traveled to Paris in August of 1862 and later went to
London to visit the World’s Fair, he neither met Darwin nor visited him
in Down. Besides, apparently Darwin had left Down with his family for
Southampton around this time, from where they went to Bournemouth,
not returning home until 30 September, by which time Mendel had al-
ready left England. Furthermore, as de Beer explains (1964, 211-212),
on 9 August Darwin wrote to Asa Gray telling him, among other things,
about his pollination experiments with Lythrium, but does not mention
any encounter with Mendel.
Things are more intriguing regarding the story about the uncut pages
of Mendel’s paper in Darwin’s library, since it has become a commonplace
in the history of biology.1 However, as Nino Strachey, curator of Down
House has unequivocally stated (responding to Robert McFetridge’s re-
quest for a copy of “Mendel’s papers” allegedly found “among Darwin’s
papers at Down House”):
The paper which is generally rumoured to have been found uncut among Dar-
win’s library is “Versuche über Pflanzenhybriden” published in Verh. naturf.
Ver. Brunn 1865. Although Darwin did receive the proceedings of some German
and Austrian natural history societies, no copy of the Brunn society proceedings
is recorded either in the 1908 catalogue of Darwin’s library or the current cata-
logue of the Darwin library at Down House. (Strachey 2000)

Strachey continues, “I have therefore been unable to trace how the ru-
mour started, or on what evidence it was based” (Strachey 2000).
A possible answer to this puzzle may be found in the debate over
whether Darwin might have known Mendel’s work through a secondary
source, likewise present in his library. In fact, two publications in the li-
brary, now at Cambridge University Library, make reference to Mendel,

1 For relatively up-to-date and allegedly authoritative reiterations of this story see Kitcher (1982, 9),

Rose (1998, 33), Henig (2000, 143-144), though none of them indicates their source.

If Darwin Had Known Mendel 5

Hoffmann (1869) and Focke (1881). The former includes Mendel’s re-
sults on Phaseolus (Hoffmann 1869, 52, n. 21) and Pisum, but nothing
exceptional was noted except that “hybrids possess a tendency to return
to their original species in the following generations.”2 The work also
casts doubt on Darwin’s reading of Gärtner’s experiments on Geum – to
which we will return later – but he did read, annotate3 and quote from
Hoffman (Darwin 1876). But there is no evidence in his texts or exten-
sive correspondence of Mendel’s name. As for the second publication,
Mendel is mentioned several times regarding his experiments on Pisum,
Phaseolus, and Hieracium. Focke notes that “Mendel’s numerous cross-
ings yielded results which were quite similar to those of Knight, however,
Mendel thought he had found constant numerical relationships” (Focke
1881, 110).4 Referring to Hieracium, there are comments about the poly-
morphic characteristic of hybrids (Focke 1881, 215) and the constancy
of individual forms (Focke 1881, 216, 217, 219, 483). Darwin acquired
the book in November 1880 (he must have obtained an advance copy,
since the book bears an 1881 publication date), just 18 months before his
death. Then, on 13 November 1880, George Romanes asked Darwin to
read a preliminary version of the entry on hybridism for the ninth edition
of the Encyclopaedia Britannica and to suggest references (Burkhardt et
al. 1994). Rather than enumerate the most influential works on hybrid-
ism, Darwin simply sent Romanes his copy of Focke’s book to “aid you
much better than I can” (Burkhardt et al. 1994). What is even more
significant is the fact that pages 108-110, where there is a brief reference
to Mendel’s experiments with peas, remain uncut. Thus, the story that
Darwin possessed an offprint of Mendel’s text with its pages uncut prob-
ably originated from this copy of Focke (1881), which was in Darwin’s
possession and did have its pages referring to Mendel’s work on Pisum
One final introductory note returns us to the paper’s title and the as-
sumption that Mendel’s work might have provided Darwin with crucial
and missing information for his evolutionary ideas. As mentioned earlier,

2 “Hybride besitzen die Neigung, in den folgenden Generationen in die Stammarten zurückzu-

schlagen.” (Hoffmann 1869, 136)
3 Punnett thinks “that Darwin must have seen that” (Punnett 1925, 606), but Olby (1985, 224)

points out that Darwin, having carefully read Hoffmann’s work (perhaps not before 1874, though) did
not mark the relevant page.
4 “Mendels zahlreiche Kreuzungen ergaben Resultate, die den Knightschen ganz ähnlich waren,

doch glaubte Mendel constante Zahlenverhältnisse zwischen den Typen der Mischlinge zu finden.”
(Focke 1881, 110)
5 In addition to the works already mentioned see Dover (2000), Gayon (1988), Sclater (2000; 2003;

2006) and (where Sclater’s and Strachey’s letters to
McFetridge are reproduced).

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Darwin’s “Provisional Hypothesis of Pangenesis,” with its mechanism
to account for the source of the variations over which natural selection
could operate, is often used to validate this assumption. However, there
is an alternative view of the relationship between Darwin and Mendel,
especially if the focus is shifted from Darwin to Mendel. In doing so
through an analysis of Mendel’s works, some of which mention Darwin’s
studies, real doubt is shed on the title’s presupposition, thus illustrating
the dangers of counterfactual history.

The Context of Mendel’s Work

Ernst Mayr distinguished two schools of breeders at the end of
the 18th and during the first half of the 19th century, “schools which
had very different interests and objectives” (Mayr 1982, 641).6 One
consisted of animal and plant breeders (Tier- und Pflanzenzüchtern),
also referred to as horticulturalists, and the second school consisted
of species hybridizers, or simply hybridists (Hybridisten). The breed-
ers were practical men who wanted to know how new and economi-
cally useful varieties could be created, thus attempting to improve the
productivity of the plants or animals they grew. Their goals included
creating better resistance to cold, brighter flower color, or better qual-
ity of wool or meat through the production of new varieties by cross-
ing with already existing varieties. Among the more prominent were
Thomas Andrew Knight (1759-1838), Alexander Seton (1824), John
Goss (1824), and Augustin Sageret (1763-1851), all of whom pro-
vided examples of the phenomena currently known as “dominance”
and “segregation” without any numerical relations (the first three
even worked on genus Pisum, made famous by Mendel; he was appar-
ently familiar with their results).7 Hybridists, in turn, possessed more
academic backgrounds. Beginning with the problem of the sexual-
ity of plants, they engaged the question raised in the 18th century of

6 We can find this distinction suggested in Roberts (1929) and, with different terminology, de-

pending on an allusion to the distinction between work performed on plants and that performed on
animals, in Bowler (1989) among others.
7 Breeders Knight and Sageret are quoted repeatedly by Gärtner (1849). Goss and Seton, on the

other hand, are only mentioned once, whence, after describing the results of his experiments with
Pisum sativum viride Gärtner claims that “These results coincide in essence with those announced
by Goss and Seton” (“Diese Resultate kommen demnach mit den von Goss und Seton bekannt
gemachten im Wesentlichen überein.” Gärtner 1849, 85). Right next to this sentence, Mendel writes
the bibliographical reference given by Gärtner, which, though it is not said there, is really a German
translation of Goss’ and Seton’s papers – with changed indications of the years of experiments – and
signed simply by “G,” which might suggest that the anonymous author describes his own experiments
(G 1837).

Regarding the changes in the concept of species in “hybridism” from Linnaeus to Mendel. see Müller-Wille & Orel (2007). his pupil Theophrastus. However. 1760. re-edited Camerarius’s work in 1749 when Kölreuter was a student in Tübingen. Rudolf Jakob Camerer or Camerarius (1665-1721) is usually considered the founder of plant sexuality. Daldberg 1755. as well as Aristotle. The first use we may label “hybridism in the broad sense. that is.” Kölreuter was a Fellow of the Academy. considered by Olby “[t]he best general account of Linnaeus’ changing views about the origin of species” (Olby 1985.9 Certain hybrids that appear in nature but that can also be produced artificially. Herodotus. with the additional purpose of finding the hidden constraints imposed by nature to the production of new species from the hybridization of preexisting species (Kölreuter 1761-1766). artificial and natural.If Darwin Had Known Mendel 7 whether new species could be created by crossing preexisting species.” 9 Regarding the changes in the concept of species and its relations to hybridization in Linnaeus. In his early writings. 1755. especially when the Assyrians and Babylonians. . where he worked as a natural historian from 1755-61. are fertile and reach the status of a new species. But it was not until much later that it was generally accepted that not only animals engaged in sex. Carl von Linné (1707-1778). see also his followers Haartman 1751. his main work being the classification of fish 8 The term ”hybridism” is used in the literature to refer both to this tradition that deals with the question of whether new species can be created by hybridization of preexisting ones. accepted this doctrine and the result- ing constancy of species (e. There are references in antiquity to the distinction between male and female in plants. Linnaeus 1736. However. and Pliny. through Kölreuter and Gärtner. he later proposed a new modified version of special creation (1744. Gråberg 1762). despite a se- ries of experiments he conducted (Camerarius 1694) that supported sex in plants. Joseph Gottlieb Kölreuter (1733-1806) accepted the challenge and performed a series of experiments with hybrids in greater number than had been done before. of a new constant species originated in the crossing of already existing species. Johann Georg Gmelin. Johann Georg Gmelin (1709- 1755) suggested that the issue should be settled experimentally (Gmelin 1749). or simply Linnaeus. Larson (1968) can also be checked. see Larson (1971). Skepticism remained even after Kölreuter’s professor. the histo- ry of which can be briefly characterized.8 The “doctrine of special creation” claimed that all existing species were originally created by an act of God. there is generation.g. This explains the decision of the Imperial Academy of Sciences of St. 1737). 270). doubts persisted. discussed dates and figs. but so did plants.” and the second “hybridism in the narrow sense. His work had as background the problem of the sexuality of plants. and to the posi- tion of those who answer affirmatively. Petersburg to offer a prize in 1759 “to strengthen or to combat the sexuality of the plants by means of new arguments and experiments.

but was sure that nature had its own means both to prevent their natural creation (Kölreuter 1761. he continued his experiments with hybrids not only with Nicotiana. where from 1763 he served as Professor of Natural History and Director of Gardens of the Margrave of Baden. 20). In 1786 Kölreuter lost his position at the garden.” In order to determine what prevented the production of hybrid plants in nature and the constant reproduction of those that were artifi- cially obtained. Verbascum. this would pro- vide powerful support for the theory of the sexuality of plants. 1766. However. but remained in Karlsruhe as Professor of Natural History. and Malvaceen. first in Leipzig. Sulz. Back in Germany. He thus supported the traditional views of special creation and rejected the “New doctrine of special creation. he created a second generation of hybrids (“back-crossed” in later terminology. . At the same time.” he tried to fertilize some of the hybrids with pollen from the father plant and others with pollen from the mother plant. 1766. Kölreuter. and then in Karlsruhe. Kölreuter had no doubts that hybrids could be produced artificially. which appeared in four parts between 1761-66. thus observing a remarkable contrast between the fertility of pure species and the sterility of hybrids. He even succeeded in creating a second gen- eration of true hybrids through the self-pollination of tobacco hybrids. In general. and “in descending degree” if fertilized with female pollen. he examined the fertility of his hybrids. according to the terminology employed by Kölreuter). and he began his experiments with hybrids that year. and Calw. con- stituted the first true hybrid produced artificially. or “in ascending degree” if fertilized with male pollen. The award was given to Linnaeus’s work Disquisitio de sexu plantarum in 1760. In his main work. Dianthus.8 pablo lorenzano and corals. he found that sterility came from “the male side” (pollen). to his opinion (Kölreuter 1761. such as Mirabilis. in others. Hybridization ex- periments on plants were considered important because if the progeny showed parental traits or if hybrid plants could be produced and analo- gies could be noticed between them and animal hybrids. 20) and to avoid the propagation as new species of those hybrids that had been obtained through experiments (Kölreuter 1763. to determine if hybrids were also sterile “from the female side. he reported the successful hybridization of a large number of plant species (Kölreuter 1761-1766). 16). In some cases he found sterility “in maximum degree”. Karl Friedrich (1749-1811). but also with other plant species. be- ing one of the judges. 1. he achieved the first success- ful hybrid crossing with two species of tobacco (Nicotiana rustica ♀ & Nicotiana paniculata ♂) which. 16. although he also studied the structure of flowers and the mechanisms of pollination (pointing to the importance of insects). had doubts about the authenticity of the hybrids described by Linnaeus.

die von mittlerer Art ist. To the production of every natural plant two similar fluid materials of different sorts are demanded. aus jenen beyden einfachen Kräften entstanden. and resembling the original male species. § 1) .” (Kölreuter 1763. This process of fecundation of offspring of hybrids with pollen from one of the original species could be continued through successive gen- erations until achieving the “return” to the original species from which the pollen came. die andere der weibliche Saame. the other female […].. the hy- brid offspring were identical. […]The one of this is male. nämlich ein Mittelsalz entsteht. entsteht eine andere. In some cases in which hybrids of the first generation were self-pollinated. sooner or later hybrid descendants returned to one or the other of the original species instead of reproducing as hybrids. even in sterile hybrids). arisen from those two simple forces. which occurs most intimately and in an orderly manner according to a definite relationship. die auf das allerinnigste und ordent- licher weise nach einem bestimmten Verhältnisse geschieht. explaining the results he achieved. […] Aus der Vereinigung und Vermischung dieser beyden Materien. independently of which of the parental species was used for fertilization. in analogy with the alchemical theory of transmutation of metals. there arises another of an intermediate sort. On the other hand. Thanks to Kölreuter’s work. Die eine davon ist der männliche. which was even greater in hybrids of other species. Hybrids of the first generation were usually all identi- cal and the majority of their traits were intermediate between the two parental species (he occasionally observed a “greater vegetative force” in the hybrid. Nonetheless. From the union and commingling of these two materials. zusammengesetzte Kraft besitz: eben so wie aus der Vereinigung eines sauren und laugenhaften ein drittes. a phenomenon known as “reversion. Hybrids from back-crossings and of the second generation were all different from each other and tended to resemble less their parental hybrid forms and more one or the other of the original species.10 10 “Zu der Erzeugung einer jeden natürlichen Pflanze werden zwey gleichförmige flüssige Materien von verschiedener Art erfordert […]. reliable and precise descrip- tions of hybrids and their descent were available for the first time in the history of biology. resembling the hybrid males of the first generation.e. und folglich auch eine mittlere.” Kölreuter expanded on his theory of fecundation (Campbell 1981). “reversed experiments” (reciprocal crosses) provided identical results. depending on which species contributed the pollen. he found that their descendants were of three types: resembling the original female species. i. and which consequently also possesses an intermediate composite force. just as through the union of an acid and an alkaline substance a third or intermediate salt originates. in a phenomenon that Kölreuter called “transmuta- tion” of one species into another.If Darwin Had Known Mendel 9 finding a slight degree of fertility.

10 pablo lorenzano Normally hybrids represented an intermediate condition of the spe- cies.”12 and in support of the requirement for two seeds (male and female) for fertilization (Olby 1985. a sense of frustration overcame him in his last years even though he continued his experiments until 1805. the combination occurred unequally. At the same time and due to what Kölreuter named the “law of the nearest affinity” (Gesetz der näheren Verwandschaft. the reversion of hybrids. die so wohl in der Chemie. § 20) 12 According to which. In exceptional cases where the hybrids varied. the external layer (or cortex) derives from the male progenitor. their intermediate character. Kölreuter’s most important results provided definitive support to the theory of the sexuality of plants and opposed the “new doctrine of special creation” proposed by Linnaeus. auf eben dasselbe eine Menge schon längst bekannter Erscheinungen. und es gründen sich. His conclusions were in complete agreement with the belief in the constancy of species and their precise differentiation of varieties and are as follows: the steril- ity of hybrids. als in der Physik. the closest affinity of pollen. . He was also never able to carry out his intention of hybridizing chickens to show that his conclu- sions applied to animals. Thus. he died one year later. whereas the central part of the flower. the identity of hybrids obtained through recipro- cal crossings. many of them ended prematurely. and the transmutation of one species into another through successive crossed pollinations. With the exception of replicating one of his experiments by the German biologist Johann Hedwig (1730-1799) in 1798. the fruit and the top of the stem come from the female progenitor. leaves and stem come from the male progenitor. thus. täglich vorkommen. The final three results were used against the spermist and ovist variations of preformation and Linnaeus’s “two- layer theory. his studies of hybrids were not reproduced until half a century after the publication of his main work and twenty years after his death. and in accordance to what was taken to be the male and female roles in the reproduction of animals. which pos- sesses “a great reach in nature and in which find its foundation a quan- tity of phenomena well-known for a long time ago that appear daily so much in chemistry as in physics”). some suffering from lack of optimal conditions and good equipment. wie es scheint. since the union blended male and female contributions equally. whereas the internal layer (or marrow) derives from the female progenitor. Augustin Sageret was 11 “[…] von einem sehr grossen Umfange in der Natur.” (Kölreuter 1766. Additionally. Kölreuter’s experiments were poorly known by his contemporaries.11 fertilization with alien pollen never takes place if the plant receives at the same time pollen from the same species and even pollen of a hybrid plant cannot fertilize if there is pol- len from one of the parental forms present. 16-18). In summary.

the Prussian Royal Academy of Sciences in Berlin offered an award to the question. Von der Sexualität der Pflanzen appeared. By the first deadline (1 January 1834) there was no an- swer. whereas Christian Konrad Sprengel (1750-1816) and Thomas A. who repeated what his teacher said (Henschel 1820). and which useful and ornamental plants can be produced and reproduced in this manner?” (Was lehrt die Erfahrung hinsichtlich der Erzeugung neuer Arten und Abarten durch die künstli- che Befruchtung von Blüthen der einen mit dem Pollen der andern. the expectation of having experiments featuring many species that would allow for a general statement. Moreover. the Dutch Academy of Sciences in Haarlem offered an award to whoever might satisfactorily answer the question. Two years later. though he did not provide any details of his experi- ments. In 1820.If Darwin Had Known Mendel 11 the first to do so. Carl Friedrich von Gärtner . “Is there hybrid fertilization in the vegetable Kingdom?” (Giebt es eine Bastardbefruchtung im Pflanzenreich?). They all gave testi- mony to the precision of Kölreuter’s work. und welche Nutz. In the absence of an answer. which surprised the critics who still denied the sexuality of plants and questioned the content and importance of his experiments. a student of Schelver. and that fertilization is not achieved by the ac- tion of pollen. Knight stood in favor of the “theory of the production by means of two types of seeds. the award was offered again in 1822. In fact. but by some exterior stimulation of the earth (Schelver 1812). This was because the specimens of the hybrid form sent in support of his work were barely recognizable. that seeds are not considered a product of fertilization. for instance.und Zierpflanzen lassen sich in dieser Weise erzeugen und vervielfältigen?). and Sergius Nawaschin (1857-1930). in 1814. additional progress in fertilization investigations was not apparent until the work of Giovanni Battista Amici (1786-1863). In 1826. “What does experience teach about the production of new species and subspecies [varieties] through the artificial fertilization of flowers of one plant with the pollen of another one. Thus. in 1812 Franz Joseph Schelver’s (1778-1832) book Kritik der Lehre von den Geschlechtern der Pflanze held that plants have no sex. was not fulfilled. In 1830. He was followed by Arend Joachim Friedrich Wiegmann (1770- 1853) and Carl Friedrich von Gärtner (1772-1850). so it was deferred until 1 January 1836. since it was considered convincing only in part. half the prize (100 ducats) was granted to Wiegmann (1828) for his work. he published Erste Fortsetzung der Kritik der Lehre von den Geschlechtern der Pflanze (Schelver 1814) and con- tinued the same notions in Zweite Fortsetzung (Schelver 1823).” Lazzaro Spallanzani (1729-1799) was against it. Eduard Strasburger (1844-1912). by August Wilhelm Eduard Theodor Henschel (1790-1856). Matthias Jakob Schleiden (1804-1881). therefore. In 1819.

J.12 pablo lorenzano did not become aware of the competition until October 1835. Gärtner devoted his life to fertilization and hybridization experiments. that hybrid fertilization occurs. cells. seeds. Gärtner showed that in some cases fer- tilization takes place by means of the use of foreign pollen (from another species). the pol- len possesses “a fertilizing force” (eine befruchtende Kraft. and that. IV).”13 Since in hybrid formation the traits of the parents considered in isolation “modify. cross. kreuzen oder vernichten. Versuche und Beobachtungen über die Bastarderzeugung im Pflanzenreich. Gärtner 1849. stimulated by the criticisms of the theory of the sexuality of plants by F. Gärtner could not meet the specified deadline (30 December 1836) because of an ill- ness.” (Gärtner 1849. III). For Gärtner the production of the plant embryo is a vital process. destroy or stand out mutually”14 giving rise to a new 13 “[…] ihre eigene Natur und Bildungskraft und ihre eigenthümliche Entwickelung und Ausbildung der Charaktere besitzt. In his book. in 1849. Schelver in 1812 and A. these factors belong to different veg- etable species and. the Dutch Academy decided to give the prize of honor to Gärtner’s work and to publish it in Dutch (1838). thus. 13). according to his opinion. After having investigated fertil- ization by means of natural and artificial pollination with pollen from the same plant in his book Beiträge zur Kenntnis der Befruchtung der vollkommenen Gewächse (1844). Gärtner did not learn of this decision until after he wrote a letter to the secretary of the Academy on 14 August 1836 who. possess “their own nature and formative force and their characteristic development and formation of the characters. On 20 May that year. mix. the sexuality of plants and of the fertil- ization process could be clearly recognized (Gärtner 1849. In hybrid fertilization. The jury then requested that Gärtner rewrite his work in a more appropriate form. 254) 14 “[…] modificiren. by return mail informed him of the demand for further elaboration. but he was finally well enough to comply in February 1837. Gärtner summarized the knowledge on hybridization and described countless experiments. or sperms – and two factors are active. one maternal and the other paternal. As this book was not widely read in Germany. revised and enlarged with the description of new experiments and discoveries. not a chemical one as Kölreuter thought (Gärtner 1849. he supported the theory of the fertilization of plants where. oder heben sich einzelne Charaktere . During fertil- ization. consequently. In this manner. Henschel in 1820 and by John Goss’ 1824 work on the change of color in peas by means of cross-fertilization. However. vermischen. He decided then to present his results to the Academy in aphoristic form. Gärtner prepared a German edition. a complete intermingling of fluids occurs – not granules.

”15 Additionally. Gärtner’s emphasis) 16 “[…] bei der Vermischung und Verbindung der Arten keine rein mathematischen Verhältnisse ihrer Faktoren darstellen. 14-15. if hybrids are spontaneously produced in nature and not only arti- ficially. Gärtner believed in the sexuality of plants. can new species be originated by means of hybridization from preexisting species. 255) 17 “Die Ergebnisse der Bastardbefruchtung und die Betrachtung über die Entstehung der Typen der Bastarde […] haben gezeigt. the origin of hybrids is casual or forced.” (Gärtner 1849. But. does this lead to the origin of new species? According to Gärtner. 15) . as much as for plants as for animals. 551-552. that the traits of the progenitors never pass pure and unmodified to the formation of the hybrid. 587-588). und noch öfter vorkom- men. and invariability of species. but dur- ing “the mixture and union of the species their factors don’t represent purely mathematical proportions (relations)”16 and “the results of the fertilization of hybrids and the consideration of the origin of the types of the hybrids [. Herbert. 152- 153. he thought hybrids were not only artificially produced. However. not only are hybrids not produced in accordance with chemical laws. Nees von Esenbeck. Lecoq. nonetheless. in accordance with the “new doctrine of special creation” or “hybridism” (Hybriditismus)? Gärtner. Puvis. but that. als bei den Thieren: dass die Charaktere der Stammeltern niemals rein und unverändert in die Bildung des Bastards übergehen. and Voigt (Gärtner 1849.. opposed this conception. Shiek.”17 Just as Kölreuter did. Kunze. 276-277) 18 “[…] dass auch in der freien Natur Pflanzenbastarde enstehen können.”18 Therefore.If Darwin Had Known Mendel 13 form. 254..” (Gärtner 1849. The reasons he gave for opposing the new doctrine of special creation are.] have shown that. it is “a general law of the production of hybrids. among others. oder nach chemischen Gesetzen erzeugt werden. no mathematical or chemi- cal proportion obtains there. als im Reiche der Thiere. but “that also in free nature hybrids of plants can be origi- nated. 4) 19 “[…] dass das Wesen der reinen Art dieser Annahme widerspricht. contrary to him. gegenseitig auf. this position was held by Knight. the following: 1. dass sich durch die hybride Zeugung verschiedene Abstufungen und Grade der Aehnlichkeit mit den Stammeltern aus den verschiedenen Arten ergeben: da aber hierin kein mathematisches oder chemisches Verhältniss stattfindet. constancy. 254) 15 “Es ist daher ein allgemeines Gesetz der Bastarderzeugung sowohl bei den Pflanzen.”19 gathering evidence in favor of the stability. in full agreement with Kölreuter.” (Gärtner 1849. “that con- tradicts the nature of pure species. dif- ferent levels and degrees of resemblance with the original parents of the different species occur.” (Gärtner 1849.” (Gärtner 1849. 12). no “essential difference” can be noticed be- tween hybrids produced artificially and hybrids as they occur in nature (Gärtner 1849. by means of hybrid production. Reichenbach. That is.” (Gärtner 1849. and that they appear even more frequently than in the kingdom of animals.

some few fertile hybrids. Labiaten. yes! of the entire plant kingdom from a few primordial types? (see before 152. but are not capable. 5. so different from that of pure species? We very much doubt that one of the researchers of nature that ascribe it that kind of genesis to plant species has ever observed and identified a true hybrid in its living and sexual behaviors. the gradual decrease of fertility and extinction of the hybrid species itself. the latter are sterile). sollten wir dann nicht noch heutzutage solche Veränderungen und Schöpfungen unter unseren Augen vor sich gehen sehen? Verträgt sich diese Hypothese mit der Wirklichkeit und der so verschiede- nen Natur der Bastarde in Vergleichung mit der der reinen Arten? Wir zweifeln sehr. 9. ja! des ganzen Gewächsreichs aus einigen wenigen Urtypen in der Phantasie herzuleiten?? (s. Labiaten. there are many absolutely sterile hybrids. Nicotiana paniculato- rustica and others. 161).” (Gärtner 1849. the great inequality and inconstancy of their fertility. 8. wie die der Umbellaten. 6. 161). the predominant influence of the fertilizing material of the original parents. they must return to one of the original par- ents.. 3. and extinguish. welche der Natur diese Art der Genesis der Pflanzenarten unterschieben. dass einer der Naturforscher. Orchideen etc. der Asclepiadeen. 588) . abandoned to the self-fecundation. (Gärtner 1849. until the “return” to the original species from which the pollen came. however. 152. However. Leguminosen. he did not be- lieve that it could be understood as the difference in fertility between them (whereas the former are fertile. of Asclepiadeen. oben S.” What could stop us from deriving in imagination the origin of whole families. Wenn dieses wirklich hätte geschehen können. contrary to Kölreuter. 4. as for instance Dianthus armeria-deltoides. eine wahre Bastardpflanze in ihren Lebens. For Gärtner. shouldn’t we see nowadays such modi- fications and creations taking place under our own eyes? Does this hypothesis correspond with the reality and nature of the hybrids. 473-475) Therefore. Leguminosen. Gärtner thought there was an essential difference between species and varieties. die Enstehung ganzer Familie.20 In agreement with Kölreuter.und Geschlechtsverhältnissen beobachtet und erkannt hat. the transformation of one species into another by means of repeated fecunda- tion of the offspring of hybrids through successive generations with pollen of one of the original species. when repeatedly inseminated artificially with their own pollen sometimes increase their fertility more and more in the successive generations. lose their reproductive capacity more and more in the successive generations (we speak here of true hybrids – not of varieties – in their natural state). even when hybrids were fertile. 7. If this could really have happened.. 20 “Was sollte nun hindern. he asserted his opposition to those who hold the “new doc- trine of special creation. 418. or self- reproduction. as well as between hybrids of varieties and hybrids of species (or “true hybrids” [wirkliche Bastarde]. as he also called them). such as Umbellaten. Orchideen etc. most fertile hybrids. as Dianthus chinensi-barbatus.14 pablo lorenzano 2.

hybrids of varieties have a much greater tendency to return to the original form by means of generation in propagation. in others to the mother) or decided bastards (decidirte Bastarde. they constitute the majority of them and are in some aspects similar to the father. mixed bastards (gemengte Bastarde. Gärtner (1849. 254. high and increased fertility. that it “is a general law of the production of hybrids. is characteristic. Gärtner attempted to locate a regularity in the manifestation of pre- dominance. hybrids of varieties differ from hybrids of pure spe- cies in the following features (Gärtner 1849. he did not believe in the sexual role of transmission (i. 5. in Linnaeus’s “two-layer theory”) because it was incompatible with the fact that reciprocal crossings are identical (Gärtner 1849. 277- 285) classified hybrids of species. special. behavior that. along with its specific form. 582). Gärtner’s emphasis). as much as for plants as for animals. and even not infrequent. since some traits prevailed more often than others.e.. they are very rare and possess an exact intermediate form). hybrids of varieties are extraordinarily more receptive to the effects of the original parental species than true hybrids. and constant in each species (Gärtner 1849. just as Kölreuter claimed. as mediated or assorted bastards (vermittelte oder gemis- chte Bastarde. 222). According to the different balance between the two intervening fac- tors and the subsequent different predominance. that the traits of progenitors never pass pure and unmodified to the formation of the hybrid” (Gärtner 1849. 3. he held not only that they were typical of a certain species. hybrids of varieties usually possess undamaged and complete male organs and from there an unshakable. 2. as already noted. in them one species has so great an influence on the form of the hybrid that the other species seems com- pletely destroyed). 163). 223). . but also. In turn. the variability of the forms in the following generations is incomparably great- er in hybrids of varieties than in hybrids of species (among many descendants of one and the same generation hardly one or two individuals are identical to another individual). 1. Hybrids of varieties are extraordinarily more receptive to the effects of the original parental species than true hybrids. 4. On the other hand. though he acknowledged there are no precise limits. But al- though he spoke of predominant individual traits. a fact consti- tuting a general rule in almost all plants and which constitutes the essen- tial difference between hybrids of plants and those of animals (Gärtner 1849. constituting its type.If Darwin Had Known Mendel 15 the nature of species is constituted by the determinate behavior of sexual forces in relation to other species.

constancy. However. Lorenzano 1995) but rather it is important to attempt to determine the questions he addressed in relation to the schools or traditions he both received and worked in. his most important conclusions and results into mediated (or assorted). 1990a. Corcos & Monaghan 1985. In his “Introductory Remarks. he intro- duced the fundamental concepts and laws later labelled “Mendel’s Laws. Herbert (1837. Callender 1988. Puvis (1837).” the “Law of Segregation” (or “Mendel’s First Law”).16 pablo lorenzano In conclusion. 1993.” Mendel clearly sets the goal of his work. Olby 1979. 1990b. suggests a different picture. and invari- ability of species. the precise difference between species and varieties and between hybrids of species and hybrids of varieties. and the “Law of Independent Assortment” (or “Mendel’s Second Law”). Thus. in so doing. Kunze and Voigt (Oken Isis 1837. read- ing his original works carefully and trying to place his research in the con- text of the 19th century. the identity of reciprocal crossings were the contribution of two “factors” (maternal and paternal) in hybrid fertilization. Nevertheless. Reichenbach. It is not important to discuss here if Mendel in fact introduced the fundamental concepts or laws of the theory later labelled “Mendelian Genetics. Linnaeus’s position of special creation has its supporters. The Main Problem Mendel Faced and Attempted to Solve According to what has become the “traditional account” in the history of genetics (Olby 1979. the stability. He held to the constancy of species and their precise differentiation into varieties. their lack of reproduction as new species and their ulterior extinction or return to one of the original species. the spontaneous production of hybrids in nature. Wiegmann (1828). but opposing the “new doctrine of special cre- ation” defended by Linnaeus. mixed and decided. .” Bowler 1989). Brannigan 1979. the in- adequacy of mathematical (or chemical) treatment or analysis of fecunda- tion and hybridization. or the “orthodox image. and the classification of hybrids of species. Bennett 1964. 1847). Johann Gregor Mendel (1822-1884) founded classical or Mendelian genetics with his “Experiments in Plant Hybridization” read in 1865 at the Natural History Society of Brünn and published the following year. Nees von Esen- beck. all of whom claimed that new species had originated from preexisting ones through hybridization. including Sageret (1826). 1969. 479).” (see Heimans 1962. 1965. Lecoq (1845). Gärtner performed his experiments with hybrids with the purpose of supporting the theory of the sexuality of plants through the production of hybrids. 1971. the impurity of traits in hybrids. Mendel was attempting to solve the problem of inheritance and.

Wer die Arbeiten auf diesem Gebiete überblickt. Wichura and others. waren die Veranlassung zu den Versuchen. wird zu der Überzeugung gelangen. has led to the experiments which will here be discussed. gab die Anregung zu weiteren Experimenten. not one has been carried out to such an extent and in such a way as to make it possible to determine the number of different forms under which the offspring of the hybrids appear. such as is effected with ornamental plants in order to obtain new variations in color. and quite recently Wichura published the results of some profound investigations into the hybrids of the Willow. mit welcher dieselben Hybridformen immer wiederkehrten. Eine endgiltige Entscheidung kann erst dann erfolgen. um neue Farben-Varianten zu erzielen. such as Kölreuter.If Darwin Had Known Mendel 17 Experience of artificial fertilization. Gärtner especially in his work “Die Bastarderzeugung im Pflanzenreiche”. 3) 22 “Dieser Aufgabe haben sorgfältige Beobachter.22 21 “Künstliche Befruchtungen. der den Umfang der Aufgabe kennt und die Schwierigkeiten zu würdigen weiss. dass man diese Formen mit Sicherheit in den einzelnen Generationen ordnen und die . Lecoq. bis Detail-Versuche aus den verschiedensten Pflanzen- Familien vorliegen. and can appreciate the difficulties with which experiments of this class have to contend. wie Kölreuter. The striking regularity with which the same hybrid forms always reappeared whenever fertilization took place between the same species induced further experiments to be undertaken. It requires indeed some courage to undertake a labor of such far-reaching extent. Die auffallende Regelmässigkeit. so kann das Niemanden Wunder nehmen. einen Theil ihres Lebens mit unermüdlicher Ausdauer geopfert. no generally applicable law governing the formation and development of hybrids has been successfully formulated can hardly be wondered at by anyone who is acquainted with the extent of the task. have devoted a part of their lives with inexhaust- ible perseverance. Gärtner. Wenn es noch nicht gelungen ist. 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. Herbert. A final decision can only be arrived at when we shall have before us the results of detailed experiments made on plants belonging to the most diverse orders. has recorded very valuable observations. a. this appears. dass es möglich wäre. or to arrange these forms with certainty according to their separate generations. That. deren Aufgabe es war. mit denen Versuche dieser Art zu kämpfen haben. Those who survey the work done in this department will arrive at the convic- tion that among all the numerous experiments made. however. ein allgemein giltiges Gesetz für die Bildung und Entwicklung der Hybriden aufzustellen. die Entwicklung der Hybriden in ihren Nachkommen zu verfolgen. Namentlich hat Gärtner in seinem Werke ‘Die Bastarderzeugung im Pflanzenreiche’ sehr schätzbare Beobachtungen niedergelegt. so oft die Befruchtung zwischen gleichen Arten geschah. Gärtner. unter welchen die Nachkommen der Hybriden auftreten. dass unter den zahlreichen Versuchen keiner in dem Umfange und in der Weise durchgeführt ist. or definitely to ascertain their statistical relations. Herbert. the object of which was to follow up the developments [Entwicklung] of the hybrids in their progeny. und in neuester Zeit wurden von Wichura gründliche Untersuchungen über die Bastarde der Weiden veröffent- licht. Lecocq. welche an Zierpflanzen deshalb vorgenommen wurden. 21 He continued: To this object numerous careful observers. die hier besprochen werden sol- len.” (Mendel 1865. so far. Wichura u. die Anzahl der verschiedenen Formen zu bestimmen.

encouraged by the experiments of plant breeders or horticulturists (crossings performed to obtain desirable modifications in individual traits). Herbert.. directed his attention to a problem related to hybrid- ists (such as Kölreuter. “evolution. Lecoq. 3-4. in which we explain the connexion between facts and laws. auf dem endlich die Lösung einer Frage erreicht werden kann. Mendel.” free of any embryological connotation. with no distinction between individual ontogeny and evolution of a lineage. 7). Thus. and contrary to Mayr.23 Additionally. Goss. This use of mathematics was also in agreement with another book read by Mendel and that will be mentioned again later. including both ontogeny and phylogeny. he established an explicit relation between this and an issue only mentioned briefly and taken for granted. he wanted to find a “generally applicable law governing the formation and development of hybrids” on the basis of a statistical analysis of his experiments. Whereas in modern German it means “development. Es gehört allerdings einiger Muth dazu.” neologisms that were in fact less common that the previous two. Gärtner. On the other hand. 23 Such an analysis. even if the latter was no longer favored in the 1840s. That is. and Wichura).” The German term “Evolution” was not introduced until the second half of the 19th century by influence of the English “evolution” and well into the 20th century the theory of evolution was still denominated “Stammesgeschichte” or “Abstammungslehre. If that were the case. expressing their proportions by combinatorial series).” as well as “Descendenztheorie” or “Evolutionstheorie.” (Mendel 1865. welche für die Entwicklungs-Geschichte der orga- nischen Formen von nicht zu unterschätzender Bedeutung ist. Another book by Ettingshausen that undoubtedly excerpted a strong influence on Mendel’s approach was Ettingshausen (1826). 37) is declared as well as that “complete theoretical knowledge. where.18 pablo lorenzano Thus. devoted to combinatorial analysis (Mendel found that traits of plants could be combined. richtige Weg zu sein. where “the importance of the mathematical viewpoint and its predomi- nance in the whole of natural knowledge” (Schleiden 1849. before going to study to the University of Vienna. Mendel should be considered as a hybrid of the two schools or traditions distinguished by Mayr. Mendel’s emphasis) The usual translation of “Entwicklung” is “development. The use of mathematics on Mendel’s part was in complete agreement with the book he studied (Baumgartner and Ettingshausen 1839) already in Olmütz. which is the problem raised by hybridists as to whether new species can be pro- duced by crossing (or hybridization of) preexisting ones. the expression “Entwicklungsgeschichte” was used with the sense of our “evolutionary history” or. and Seton and from the results they obtained on Pisum. both to embry- onic development and what later be called “evolution.” However. indessen scheint es der einzig. sich ei- ner so weit reichenden Arbeit zu unterziehen. at least for a whole year. 39).” back then – towards the middle of the 19th century – it was used to refer to any process of development. Schleiden (1849). Consequently. it is asserted that “in natural sciences there is no more science than the mathematics there con- tained” (Baumgartner and Ettingshausen 1839. a novelty Mendel was aware of and that would also distinguish him from the aforementioned Knight. though not the goal pursued. is only possible through mathematics and inasmuch it is applicable”(Schleiden 1849. this term is ambiguous. simply. It would be a mistake to suppose that Mendel always used “Entwicklung” in the sense of individual development. i. as will become clear later. Finally. during an eight week physics course based on that book. was new to the tra- ditions mentioned (although not to biological sciences in general). with whom Mendel would also study. In fact. . which where expressed qualitatively or comparatively and not quantitatively. in a Kantian spirit. Andreas von Ettingshausen was also his teacher in the University of Vienna. as Christian Doppler’s successor. gegenseitigen numerischen Verhältnisse feststellen könnte. Mendel was looking for a law that governed both development and evolution. the phrase “Bildung und Entwicklung” – here translated as “formation and development” – would be redun- dant. it was also in agree- ment with the Pythagorean perspective on life characteristic of German romantic Naturphilosophie.e.

even if Mayr acknowledged that “Mendel occasionally calls himself a hybridizer and in his paper [Mendel 1865] often refers to Kölreuter. 713). für die in Rede stehenden Versuche völlig gleichgiltig. some breeders were explicit on the role played by hybrids in the generation of new species and. hybridists also reported what happened in their crossing experiments with individual traits. the fact that “[t]he positions.” though not as a way of providing methods. rather than providing necessary and sufficient conditions for the application of the concept of species (see Carnap 1936-1937 for a “refutation” of operationalism as a way of defining. at least. thus blur- ring the original distinction (or.” because we think that what is being formulated is an operational criterion or method for distinguishing species from varieties. to an essential concept of species (Mayr 1982. as hybridists were. on the one hand. ebesowenig ist es bis jetzt gelungen. and other plant hybridizers” (Mayr 1982. in the strict logical sense those concepts later called “theoretical. as was true between hybrids of varieties and hybrids of spe- cies.24 So. nach welcher zu einer Art nur jene Individuen gehören. did not establish any distinction between varieties and species because he considered it only a matter of degree (we will return to this later when comparing Mendel’s position to Darwin’s). Independently of the fact that. 25 “Übrigens bleibt die Rangordnung. unlike his more conspicuous hybridist predeces- sors. Mendel does more than just often refer to them. In fact. on the other hand. if what Mendel regarded as “the strictest determina- tion26 of a species. Gärtner.” (“So wenig man eine scharfe Unterscheidungslinie zwischen Spezies und Varietäten zu ziehen vermag.”27 Either this was not the case. welche man denselben im Systeme gibt. One of the reasons. 713) correctly established. according to which only those individuals belong to a species which under precisely the same circumstances display pre- cisely similar characters. einen gründlichen Unterschied zwischen den Hybriden der Spezies und Varietäten aufzu- stellen”. however. ways or criteria to determine the extension of such concepts). no two of these varieties could be referred to one species. which may be assigned to them in a classificatory system are quite immaterial for the purposes of the experiments in question. Mendel. 713).If Darwin Had Known Mendel 19 Mendel should not be excluded from the hybridist tradition. such as Kölreuter and Gärtner.” (Mendel 1865. since though the plants he 24 “It has so far been found to be just as impossible to draw a sharp line between the hybrids of species and varieties as between species and varieties themselves. As Mayr (1982. Additionally. Mayr did not include Mendel in the hybridist tradition was that similar to the breeders. die unter völlig gleichen Verhältnissen auch völlig gleiche . this is a very important issue for interpreting Mendel’s work. some of the terms in which it is formulated). 24) 26 We believe it would be better here to translate “Bestimmung” as “determination” rather than “definition. 24).”25 is what allowed Mendel to try to face the problem of hybridists (raised in relation to species) with the breeders’ techniques (used on what are accepted as varieties). Mendel 1865. 27 “Wollte man die schärfste Bestimmung des Artbegriffes in Anwendung bringen. Mendel was concerned with individual traits and was not committed.

On the other hand.” (Mendel 1865. saccharatum. so könnten nicht zwei davon zu einer Art gezählt werden.” (Mendel 1865. 19). 177. nor in the Notizblatt 1 and 2) that might help us resolve the issue. as these authors correctly point out (Müller- Wille & Orel 2007. if it was applied to one or a few traits of individu- als. at the beginnings of the 20th century. bald als selbständige Arten angesehen und beschrieben wurden. the term “species” is also used by Mendel in the expression “good species” (“gute Arten”). some as sub-species of sativum. was different from it (similar to the problem of evolution. n. 56]) and from it alone it is not entirely clear (as these authors hold) that he is trying to establish a systematic distinction between “species (without qualifica- tion)” and “good species. während die übrigen bald als Unterarten von sativum. though related to heredity. in the formulation given in 1837 by Mendel’s protector Franz Cyrill Napp (1792-1867).” [“Für die Versuche dienten grösstentheils Pflanzen. “why are organisms (offspring) partly similar and partly different from their progenitors?” or. nor in the letters to Nägeli. 24) 28 “Nach der Meinung der Fachgelehrten indessen gehört die Mehrzahl der Species Pisum sativum an. and some as independent species. However. saccharatum. plants were mostly used which rank as good species and are differentiated by a large number of characters. a statement noted in numer- ous texts by the Mendel scholar and director of Mendelianum. we have not been able to find the expression “good species” in any of the other texts by Mendel relevant to this topic (neither in the paper on Hieracium. even though he only does it once (“For the experiments. Vitěslav Orel. while the rest are regarded and classed. umbellatum. cannot be identified with it). and umbellatum.”28 Or. Merkmale zeigen. it is worth noting that Mendel did not mention explicitly the problem of inheritance. see Müller-Wille & Orel (2007). the most general formu- lation of which might be “why do offspring resemble their parents?” or.20 pablo lorenzano used did not meet the stipulated requirement.” Mendel 1865. the individuals that satisfied it belonged to the same species. . though it was susceptible of being related to that of heredity. this does not affect the central point in his discussion. For Müller-Wille & Orel (2007). whom he would succeed after Napp’s death as abbot of the Augustinian monastery of Brünn. wie quadratum.” these latter being characterized as differing “in a great number of traits. “how are biological traits transmitted from parents to offspring?” or.29 On the other hand. Unfortunately. which both separates him from the tradition of “hybridists” and explains why it is appealing to those who read it later. welche als gute Arten gelten und in einer grösseren Anzahl von Merkmalen verschieden sind. The problem of hybridization.” The previous passage is also consistent with considering “species” and “good species” as identical and with these additionally differing “in a great number of traits” in the case at hand. systematic botanists (“ex- perts”) claimed that “the majority belong to the species Pisum sativum. dealing with the problem of hybridization following “die schärfste Bestimmung von Spezies” is what forces Mendel to concentrate simultaneously in the transmission of individual traits. 24) 29 For a discussion of the role of the “schärfste Bestimmung des Artbegriffes” in Mendel’s work. This latter option was what would justify Mendel’s terminology of refer- ring throughout his work to his varieties of peas as different “species” (Arten). “what is inherited and how?” Napp made this statement at the conclusion of one of the many discussions held by the Moravian sheep breeders organized in the Sheep Breeders’ Society of Brno. such as quadratum.

Mendel’s age was not particularly interested in ‘pure’ transmission genetics. 711]). 272. all his work was fundamentally about heredity (continuing the previous phrase he adds “in his work on inheritance” [Mayr 1982. in seine beiden Faktoren kein ganz vergeblicher Versuch sein. differentiation during development.] so.] so möchte die Analysierung eines Bastards mit einfacher Organisation und geometrischer Regelmässigkeit des Stengels. and so forth.. sondern folglich auch der Gewächse überhaupt auf die Spur zu kommen. w. nicht nur den Gesetzen der Formbildung der Bastarde. 66. and as a justification for his work he points out that it “should be of great interest at least to the contemporary and future natural researcher. he must be placed in a context concerning the problem of evolution. Inheritance was generally considered only in connection with other biological phenomena. astronomer. but considered that despite this. s. in the very “Preface” of his book. environmental induction (and the inheritance of acquired characters). friend of Mendel. 250) In the same line. Mendel underlined page 272 of Gärtner’s book. der Blätter u. to have . not only to trace the laws of the form-formation of the bastards but consequently also of plants in general.” Gärtner 1849. 110-111)... 713] and “as in the case of Darwin. Mendel’s marks) Additionally. Thus.If Darwin Had Known Mendel 21 Mayr admitted that this was the usual scenario in Mendel’s times (Mayr 1982. in favor of the existence of laws for the formation of the form of hybrids and plants in general. the consolidation of species characters in isolation and their breakdown (‘blending’) following the removal of the isolation barrier. see below). and despite Mayr’s latter claim to understand Mendel bet- ter and to clarify his relation to Darwin. 725)30 and placed him clearly in a context related to the problem of evolution (“As a student of Unger and of the problem of evolution” [Mayr 1982. [“[. teacher at Brno’s Technical School. where it says [.32 30 “For reasons that are not at all clear. The connection between the task sketched (finding a “generally ap- plicable law governing the formation and development of hybrids”) and the problem of the origin of new species by hybridization was familiar in such a context even for Mendel himself. such as the species problem (and that of the species hybrids). The explanation of the origin and formation of the forms of the hybrids from the elements and characters of the progenitors is just as important for plant physiol- ogy as for systematic botany.31 This is clear when Mendel underlined and marked with double line on the left margin (which is what he used to do with texts he found of utmost importance) Gärtner’s 1849 passage. Gustav Niessl von Mayendorf (1839-1919).” (Mayr 1982. see Kříženecký (1965. Secretary of the Natural History Society when Mendel gave his lectures in 1865 and editor of the Verhandlungen [Proceedings] where they were published says in his memories about Mendel that he “was much concerned with evolutionary thoughts” (“sich viel mit dem Evolutionsgedanken beschäftigte”. For an accessible reproduc- tion of such memories that would supplement those communicated to Iltis. the analysis of a bastard with simple organization and geometrical regularity of the stem. 711]). botanist. Iltis 1924. (Gärtner 1849. the leaves etc into its two factors would not be a useless attempt. 32 “Die Erklärung der Entstehung und Bildung der Formen der Bastarde aus den Elementen und Charakteren der Stammeltern ist für die Pflanzenphysiologie ebenso wichtig als für die systematische Botanik.. 725) 31 According to Iltis. it was the species ques- tion which inspired Mendel” [Mayr 1982.

250. the question is put thus “in relation with the fertility and multiplication [of hybrids] it is not explained: how. und diese Vervielfältigung der vegetabilischen Formen aufgehört. S. ix) 33 There. editor of the Catholic newspaper Wiener Kirchenzeitung.und Nachwelt von grossem Interesse sein müsse. wel- che Formen ihm besonders unterworfen seien. Wunderlich 1983. einen Status quo zu haben: um zu erfahren. Gliboff 1998. On the investigation of the origin and formation of hybrid-types from the characters of the progenitors will give even further enlightenment. 1968. and exerted a profound influence on him (Olby 1967. wenn es deren mehrere gegeben haben sollte. why and at what time would the mixing of the original types. und nach welchen Gesetzen und Normen er erfolge?”. what forms are subject to it and according to which laws does it take place?” (“dass es wenigstens dem Naturforscher der Jetzt. warum und zu welcher Zeit die Vermischung der Urtypen. Brunner accused Unger of materialism and corruption of the youth. who either dis- covered or guessed the author of the seventeen letters that appeared anonymously in the weekly supplements of the local newspaper Wiener Zeitung in 1851 (later published in book form. 153) 34 “[…] indem sich für letztere noch die Lebensfrage anknüpft: ob es stabile (abgeschlossene) Arten der vollkommenen Gewächse gibt. wie weit er sich erstrecke. 1999).34 It is likely that Franz J.” (Gärtner 1849. He was subjected to a series of attacks from 1851-56 for his public support of nascent evolutionary views. Unger did a status quo: to experiment what does the claimed change in plants consist in. und ihrer Nachkommenschaft. worin dieser behauptete Wandel der Formen bestehe. as some naturalists believe? This question has been already put above (s. and their offspring. 1971. oder ob sie im Laufe der Zeiten einer Veränderung oder Fortbildung unterworfen sind. und sie ihre Erschöpfung oder ihren Stillstand gefunden haben sollen?” Gärtner 1849. the fol- lowing: which [the explanation] is still connected for the latter [systematic botany] with the question for life: whether there are stable (closed) species of complete plants. 153):33 we have pronounced for the reasons quoted there for the stability of plant spe- cies.N. and this reproduction of vegetable forms have ended and their exhaustion and stop occurred?” (“Nun ist es aber. Mendel’s marks) . or whether they are subservient in the course of time of a change or proliferation. without the double line on the left. attacks directed by Sebastian Brunner. 153) zur Sprache gekommen: wir haben uns aus den dort angeführten Gründen für die Stabilität der Pflanzenspecies ausgesprochen. Die Untersuchung der Entstehung und Bildung der Bastardtypen aus den Charakteren der Stammeltern wird hierüber noch weitere Aufklärung geben.A. Unger was his teacher of “Anatomy and Physiology of Plants” and “Use of the Microscope” at the University of Vienna. abgesehen von der abweichenden Natur der Bastarde von der der reinen Arten. had there been many. wie einige Naturforscher glauben? Diese Frage ist schon oben (s. Gärtner 1849. what is its extension.22 pablo lorenzano He then underlined. in Hinsicht auf Fruchtbarkeit und Fortpflanzung nicht erklärlich: wie. Unger was also the first cytologist to declare that every cellular multiplication oc- curs by division (he was also a pioneer in ecology and paleobotany). Weiling 1983. Unger 1852). Unger (1800-1870) called Mendel’s at- tention in 1852 to this book.

so that the pollen tube counted as the female organ. then the result of each fertilization could be no other than that the developed . after the latter entered the interior of the embryo sac through the micropyle pushing the sac of the membrane that is before it. and. it is highly probable that the second of the only two footnotes in Mendel (1865) about the nature of fertilization in plants was meant to settle the differ- ences between two of his teachers. according to Kronfeld. yet more importantly. because even though Iltis on the basis of a letter from Kerner to Niessl. the pistil. 1871. 36 Whereas Fenzl accepted Schleiden’s theory of fertilization (according to which the embryo de- veloped in the upper part of the pollen tube. who in 1856 convinced his tutor that Amici and Hofmeister were right. Kronfeld 1908. Mendel. 387). Unger might also have motivated Mendel’s experiments of trans- plantation (see below). which stimulates the development of this germi- nal vesicle). Anton Kerner von Marilaun (1831-1898) to perform his famous experiments of transplantation. moreover. and the embryo sac were considered the male organ). How could we otherwise explain that among the offspring of the hybrids both parental forms reappear in equal numbers and with all their pe- culiarities? If the influence of the egg cell upon the pollen cell were only external. according to Tschermak (1951. following Unger. he also received support from 401 students in the Faculty of Medicine who filed a petition to the Minister of Culture and Education in support of Unger. 1889).36 with whom Mendel studied “Morphology and 35 About whom it is discussed whether he read Mendel (1865). Hofmeister (1824-1877) and. Unger first held Schleiden’s theory. 28). noting that he never addressed is- sues of religion in his scientific teachings. the aforementioned Unger and Franz Fenzl (1808-1879).If Darwin Had Known Mendel 23 not respond to the increasing attacks. But it is known that he received an offprint of the first of Mendel’s works (the note that accompanied the work sent is reproduced in Kronfeld 1908. if it fulfilled the role of a nurse only. just as he inspired one of Mendel’s classmates in Vienna. and not Fenzl who was Erich Tschermak’s grandfather and could have been his examiner the second time he failed the exam to become teacher. had its pages uncut!. n. where he proved that variations produced by plants that grow in alpine conditions were lost as soon as they were transplanted to lowland conditions (Kerner 1860.35 Finally. since the paper was within his reach in the library of Innsbruck (Barthelmess 1952. 139). 299). 47. that is already present inside the embryo sac before the arrival of the pollen tube. Eventually. opposed Schleiden’s theory and supported Amici’s. and so did W. establishes that he knew about Mendel’s experiments with Hieracium (Iltis 1924. though he filed a lawsuit against Wiener Kirchenzeitung. Schleiden’s doctoral candidate Ludwig Adolph Timotheus Radlkofer (1829-1927). a future defender of the new doctrine of special creation. Barthelmess thinks he did indeed know it. whereas the stigma. but from the mid-fifties on he subscribed Giovanni Battista Amici’s theory (according to which the embryo comes from the germinal vesicle. “Mendel’s first publication apparently also failed to impress Kerner” (“da Mendels erste Veröffentlichung an- scheinend auch auf Kerner ohne Eindruck blieb”. 77). the lawsuit against the paper was dismissed for lack of evidence and Unger announced a public clarification of his opinions. which. denying his support for pantheism or materialism and claiming that his scientific work never contradicted the Christian belief in a personal God. Thus in the footnote mentioned we read: In Pisum it is placed beyond doubt that for the formation of the new embryo a perfect union of the elements of both reproductive cells must take place.

He had written a book (Wichura 1865) that Mendel probably read before sending his lectures of 1865 to be printed. wäre demselben blos die Rolle einer Amme zugetheilt. including Gärtner and. has first expressed the opinion and more recently justified in detail that a great part of the dubi- ous forms appeared in willows were hybrids and not species.24 pablo lorenzano Taxonomy of Phanerogams” and “Research and Description of Plants” at the University of Vienna. of its placement in an evolutionary context. dass es für die Gestalt der Hybride gleichgiltig ist. neither could I fail at least to hybrid should exactly resemble the pollen parent. more interestingly.. Darwin. Ein gründlicher Beweis für die vollkommene Vereinigung des Inhaltes beider Zellen liegt wohl in der allseitig bestätigten Erfahrung. particularly the most prominent works of Koelreuter […] and Gaertner […] on fecundation of hybrids. Mendel mentioned another hybridist. this way. welche von den Stammformen die Samen. since. provide the ex- periment that confirmed the theory of which I was soon convinced [. or at any rate do so very closely. “Bei Pisum ist es wohl ausser Zweifel gestellt. as was generally accepted until then. thus. which of the parental forms is the seed parent or which the pollen parent.oder Pollenpflanze war. Das haben die bisherigen Versuche in keinerlei Weise bestätigt. Max Ernst Wichura (1817-1866). dass unter den Nachkommen der Hybriden beide Stammformen in gleicher Anzahl und mit allen ihren Eigentümlichkeiten wieder hervortreten? Wäre der Einfluss des Keimsackes auf die Pollenzelle nur ein äusserer. as well as to the experimental results obtained by others.” (Mendel 1865. Wichura also relates hybridization to the problem of the origin and mul- tiplication of species. In a series of works […] my revered friend […] Wimmer.]. that willow hybrids were not. To further the connection between Mendel’s work and the problem of the origin of species by hybridization and. Stimulated by the subject’s own interest and in order to silence once and for all the objection raised from other side. dann könnte der Erfolg einer jeden künstlichen Befruchtung kein anderer sein.. This the experi- ments so far have in no wise confirmed. 229-230) . infertile […] finally I found myself in the need of taking into account results obtained by other observers. Mendel makes his point in terms very similar to those previously used by Gärtner: Because in the crossing of species the same complete products are produced: so we find in that the strongest evidence against Schleiden’s theory of the origin of the embryo at the end of the pollen tube. as regards the form of the hybrid. An evident proof of the complete union of the contents of both cells is afforded by the experience gained on all sides that it is immaterial. oder ihr doch sehr nahe stände. Wie wollte man es sonst erklären. als dass die entwickelte Hybride ausschliesslich der Pollenpflanze gleich käme. how could the same complete embryo with identical type of development be obtained from two different species of pollen? (“Da bei der Kreuzung der Arten sich die vollkom- men gleichen Produkte in den Typen ergeben: so finden wir hierin den stärksten Beweis gegen die Schleiden’sche Entstehungstheorie des Embryo aus dem Ende des Pollenschlauchs. dass zur Bildung des neuen Embryo eine vollständige Vereinigung der Elemente beider Befruchtungszellen stattfinden müsse. I decided to produce hybrids of willows by means of artificial fertilization and. 58-59) By the way. Gärtner 1849. denn wie könnte aus zwei verschiedenen Arten von Pollen der vollkomen gleiche Embryo mit seinem ganz gleichen Entwicklungstypus hervorgehen?”. after the selec- tion of which a relatively small number of genuine clearly determinate species remained.

38 Finally. in welche die bei der Bastardbefruchtung hervortretenden Erscheinungen mit den Ansichten Darwin’s über Entstehung der Species gebracht werden können. Weidenbastarde durch künst- liche Befruchtung herzustellen und auf diese Weise der Theorie.. das bestätigende Experiment hinzuzufügen. namentlich die wichtigen Arbeiten Koelreuter […] und Gaertner […] über Bastardbefruchtung in Betracht zu ziehen. clarifying even the methodical nature of detailed experi- ments to determine the number of different forms under which the offspring of the hybrids appear. or definitely to ascertain their statistical relations. dass auch die wichtige Frage über Entstehung und Fortpflanzung der Species nur durch lange Reihen methodisch fortge- führter. after emi- grating to the USA in 1869. zur Befestigung der Ueberzeugung beizutragen. echter Arten übrig bleibe.] endlich sah ich mich genöthigt.] dass die Weidenbastarde nicht.37 Wichura’s text also contains. had adopted the idea of evolution and.If Darwin Had Known Mendel 25 note the connection that could be made between the impressions obtained in the fecundation of hybrids and Darwin’s opinions about the origin of species. 1-3) 38 “[…] die Anzahl der verschiedenen Formen zu bestimmen. die Verbindung wenigstens anzudeuten. wie man bisher angenommen hatte. as Mendel’s does. that I also pursued by the generalization of my subject-matter. Mendel’s introduction seems to be in dialogue with Wichura. Mendel’s regard for the centrality of the problem of evolution- ary history to the problem of hybridization may also be traced to (apart from hybridists and his teacher Unger39) the first of the guiding maxims 37 “In einer Reihe von Arbeiten […] hat mein verehrter Freund […] Wimmer zuerst die Ansicht aufgestellt und im Einzelnen näher begründet. auch die von an- dern Beobachtern gewonnenen Resultate. dass ein grosser Theil der unter den Weiden vorkom- menden zweifelhaften Formen Bastarde seien. Angeregt durch das Interesse des Gegenstandes und um den von anderer Seite her erhobenen Widerspruch ein für alle Mal zum Schweigen zu bringen. 4) 39 Some have noted the influence exerted in that sense by Matouš Klácel (1808-1882). unter welchen die Nachkommen der Hybriden auftreten.. My work has thereof gained scope and from the originally intended report on willow hybrids it has become a possibly abridged exposition of fecundation of hybrids in the vegetal kingdom in general explained in willow hybrids. [. a reference to the experimental approach that needed to be taken to solve the prob- lem.. nach deren Aussonderung eine verhältnissmässig gerin- ge Zahl klar bestimmter. zahlreicher Experimente ihrer endlichen Lösung entgegengeführt werden kann. Sollte es mir gelungen sein. Moreover.” (Wichura 1865.” (Mendel 1865. thus I would have achieved the main goal. unfruchtbar seien [. auch habe ich nicht umhin gekonnt. being a free thinker. Mendel’s fellow monk and teacher who. den ich bei Verallgemeinerung meines Themas verfolgte.. von der ich mich bald überzeugt hatte. or to arrange these forms with certainty according to their separate generations. dass man diese Formen mit Sicherheit in den einzelnen Generationen ordnen und die gegenseitigen numerischen Verhältnisse feststellen könnte. possibly with help from Mendel himself (an abbot by then). even wrote . If I were to succeed in contributing to enforcing the conviction that also the important issue of the origin and multiplication of species can only reach a final solution by means of long series of numerous experiments methodically performed. so würde ich den Hauptzweck erreicht haben. beschloss ich.

” (Mendel 1870a. 235) . According to Schleiden.] jede Hyp o t h e s e . Orel 1971. and so. 42 “A. […] the only chance of reaching a scientific understanding in botany. […] every hypothesis. und somit das einzige und unumgängliche methodische Hülfsmittel. 1983).40 Schleiden followed the so-called “Kantian-Friesian natural philosophy.. 1990. in Whewell’s. welches aus der Natur des Gegenstandes sich von selbst ergiebt. welche nicht d u r c h E n t w i c k l u n g s g e s c h i c h t e o r i e n t i e r t i s t . founded by Jakob Friedrich Fries (1773-1843) in Jena. for an analysis of the influence of the second see below.”43 presenting the results of an experiment to the contrary (Mendel 1870a. he discussed Charles Naudin’s (1815- 1899) and Darwin’s opinion “that a single pollen grain does not suffice for fertilization of the ovule. Mendel referred to “Darwinian theories” in his work on hybrids of Hieracium (Mendel 1869) and in letters to Carl Wilhelm von Nägeli (1817-1891). the one Bacon and Mill were concerned with. […] die e i n z i g e Möglichkeit. On 3 July 1870. ist das Studium der Entwicklungsgeschichte. to a lesser degree. is the study of the history of development/evolution [Entwicklungsgeschichte]. and empirical induction.. 146. Apelt (1854. [.42 Did Mendel know Darwin’s work? Although it does not seem likely that Darwin knew of Mendel’s work. 41-50) differentiates both conceptions of induction through his distinction between rational induction. he about Darwin’s theory (Voipio 1987. Mill or the neoinductivism yet to come. After all. Schleiden’s emphasis) 43 “dass zur genügenden Befruchtung eines Ovulum ein einziges Pollenkorn nicht ausreichend sei. based on the Kantian conception of regulative ideas and guiding maxims.” (Schleiden 1849.142. 1973. every induction in botany that is not oriented by the history of development/evolution [Entwicklungsgeschichte] must be rejected unconditionally. Schleiden’s understanding of induction is not aligned with that of Bacon. It is induction in Apelt’s sense and. and justified in terms of the developmental history of plants. recommended by Unger and ad- opted by Mendel to practice botany according to the methodological standards of his time (Orel 1979). 141. 41 As Buchdahl (1973) shows. and reformulated Fries’s general guiding principles for research. both appear in the methodological introduction to Schleiden’s 1849 text. 1972. the only and inescapable instrument self-originated in the nature of the object. Maxime der Entwicklungsgeschichte. evaluated. j e d e I n d u k t i o n i n d e r B o t a n i k i s t u n - bedingt zu verwerfen. 235). zu wissenschaftli- cher Einsicht in der Botanik zu gelangen. Maxim of the history of development/evolution [Entwicklungsgeschichte]. all inductions and hy- potheses in botany are oriented. Later. 40 In this section we’ll only consider the influence of the first of the guiding maxims.” also known as “mathematical natural philosophy” and Friesian school. on 27 September 1870.26 pablo lorenzano (“leitende Maximen”) or regulative principles (“regulative Principien”) formulated by Schleiden.41 A.

Mendel’s teaching colleague and later univer- sity professor. 240) 45 On the other hand. since “almost all the Darwinian literature of the sixties and seventies could be found in the library of the monastery of Brünn”.” 47 “[…] in der Klosterbibliothek fast die ganze darwinistische Literatur der sechziger und siebziger Jahre vorzufinden.46 Again. more appropriately. he rarely referred to Darwin nor is there a clear statement about the theory of evo- 44 “[…] hohen Grad von Selbständigkeit […]. on 25 September 1861 the Origin of Species was discussed in Brno. and also rejected the distinction between species and varieties. 46 By the way.” (Iltis 1924. Orel 1971). a meeting that was just previous to Mendel’s lecture (1865) and at which Alexander Makowsky (1833-1908). attributable to Malthus half a century before Darwin. Nevertheless. 66) . concerning hy- brids. 70). Additionally.45 Mendel did possess a copy of the second German edition (1863 trans- lation of the third English edition) of the Origin of Species. Iltis notes that Mendel purchased all of Darwin’s books as soon as they appeared in German (he could not read English). but Darwin’s book from 1868 was the one most marked and commented upon (Richter 1942. “On the Geological Succession of Organic Beings” was published in a German journal to which Mendel might have had access. 240)44 and also used the expression “struggle for existence” (“Kampf ums Dasein”) in a letter on 18 November 1873 (Mendel 1873. Unger also supported the evolutionary hypothesis. Moreover. a year after a literal translation of Darwin’s chapter. Thus. which he (with the exception of the translation of “Fertilization of British Orchids”) filled with comments. 247).If Darwin Had Known Mendel 27 expressed his agreement with Darwin’s and Rudolf Virchow’s (1821- 1902) opinions about the “high degree of independence that is typical for individual characters and whole groups of characters in animals and plants” (Mendel 1870b. Darwin’s theory of descent by modification through the means of natural selection was clearly described in the pub- lication. Mendel had access to all of Darwin’s works. The eighth chapter of the Origin. both in the German edition of Darwin (1859) and in Makowsky’s lecture the expres- sion “natural selection” is translated as “natürliche Züchtung” (“natural breeding”) and not as would be later translated. Finally. Orel 1968. it is clear he was familiar with Darwin’s ideas. it is not necessary to be a Darwinist to use this expression. was specially marked. spoke in favor of Darwin’s theories.47 Thus. many pas- sages of which he marked (Richter 1942. 1971).” (Mendel 1870b. a report on the Origin was presented at the meet- ing of January 1865 of the Natural History Society of Brno. He prob- ably also knew of the early German edition possessed by the Natural History Society of which he was a member (Orel 1971). as has been noted (Callender 1988. welche einzelnen Charakteren und ganzen Charaktergruppen an Thieren und Pflanzen zukommt. and he adds that Mendel was not restricted just to Darwin’s books. as “natürliche Auslese.

Kíenecký 1965.48 Then. This is the context to explain Mendel’s reaction to the transplantation experiments about the influence of the environment on plants. in terms of his general reaction to Darwin.] it is not clear however why the mere transplant to the garden soil can originate in the plant’s organism such a complete and persistent revolution. 66. but nothing justifies the supposition that the tendency to form new varieties has been so increased that species loose afterwards all stability. in opposition to the mechanisms advocated by Darwin.28 pablo lorenzano lution by natural selection. Darwin’s 1859 book contained few results or explanations of those phenomena Mendel had observed experimentally. type changes must appear when the living conditions change and when a species has the ca- pacity to adapt to the new environment. where he thought “half jestingly” (halb scherzhaft). though every time there was talk about Darwin’s 48 “[s]o viel sehe ich schon. dass durch die Cultur die Entstehung neuer Varietäten begünstigt und durch die Hand des Menschen manche Abänderung erhalten wird. 110). allein nichts berechtigt uns zu der Annahme. dass die Arten bald alle Selbstständigkeit verlieren und ihre Nachkommen in einer endlosen Reihe höchst veränderlicher Formen auseinander gehen. It is known that this is not the case. Hier wie dort müssen typische Abänderungen auftreten. Mendel claims: “[. wieder an Selbstständigkeit gewonnen hätten. so dürfte man er- warten. nature does not proceed that way in making species. willingly.. “as much as I now see. Granted. it would be expectable that cultivated plants. it was advisable to be careful and not get involved in a public dispute about evolution- ism in general or Darwinism in particular. Niemand wird im Ernste behaupten wollen. sich den neuen Verhältnissen anzupas- sen. even though he was known as a supporter of the Liberal Party. Es wird gerne zugegeben. If the change in living conditions were the sole cause of variation. A probable cause for Mendel’s silence was that as a priest (and later an abbot) who lived in the Austro-Hungarian Empire in counter-revolutionary or conservative times. raised through centuries under almost identical conditions. Even though Mendel sometimes dis- cussed Darwin’s notions and shared with him the basic belief in evolu- tion and genesis of new species in the course of time by natural means. als am Gartenbeete. Wäre die Aenderung in den Vegetations-Bedingungen die alleinige Ursache der Variabilität.. dass die Neigung zur Varietätenbildung so ausserordentlich gesteigert werde. 66). dass jene Culturpflanzen. dass die Entwicklung der Pflanze im freien Lande durch andere Gesetze geleitet wird. da muß noch irgend etwas anders dabei sein!” (Iltis 1924. Thus. warum das blosse Versetzen in den Gartengrund eine so durchgreifende und nachhaltige Revolution im Pflanzen-Organismus zur Folge haben müsse.” Mendel 1865. welche im freien Zustande unter- liegen müsste. 36) . regain stability. especially natu- ral selection. that cultivating favors the formation of new varieties and that by the work of man many variations that would have disappeared in the wild have been preserved. Additionally. welche Jahrhunderte hindurch unter fast gleichen Verhältnissen angebaut wurden. as much as there. wenn die Lebensbedingungen für eine Art geändert werden und diese die Fähigkeit besitzt. Here. No one would dare claim seriously that the development of the plant is directed by different laws in the country and in the garden beds. daß es die Natur auf diesem Wege im Speziesmachen nicht weiter- bringt. he still accepted and defended hybridization as the means of speciation. Das ist bekanntlich nicht der Fall.” (“Es ist jedoch nicht ein- zusehen. “Mendel […] had not been an opponent of Darwin’s theories. and that their offspring diverge in an infinite number of extremely variable forms. there must be something different!” (Iltis 1924.

who almost devoted their lives to this subject” (Darwin 1861. Darwin rea- soned that Gärtner’s rejection of them was mistaken. since it is the German translation of this one that we have record of being meticulously read by Mendel. to which he dedicated a whole chapter of the Origin of Species. but contrary to Kölreuter he does not think that this distinction can be es- tablished by the difference in fertility between these. wenn von Darwins Lehre die Rede war. Also. and that the fertility often suddenly decreases in the first few generations. yet he asserts positively that their fertility never increased. he thought that could not be all. he did not oppose Darwinism qua evolutionism but thought that it did not provide a satisfactory explanation of the ori- gin of new species. . 27). daß er sich aber immer. 66) 50 Page number corresponds to the third English edition. was to de- bate the widely held conception that these phenomena confirmed the existence of a fundamental distinction between species and varieties. Concerning the hybrids Mendel considered constant. I do not doubt that this is usually the case. daß das nicht alles sein könne.. 301). though “always with gradual decreasing fertility and general decrepi- tude of the species. Gärtner’s group of “excellently fertile hybrids” (to which Mendel also made reference).” (Iltis 1924. that there still was something missing. [. da fehle noch etwas. for six or seven. dahin äußerste. carefully guarding them from a cross with either pure parent. Darwin and His Criticism of Hybridism Darwin’s main goal in examining the phenomena of hybridism. 268). as he elsewhere claimed. Nevertheless I believe that in all these 49 “Mendel […] zwar kein Gegner der Darwinschen Theorie gewesen sei.”49 Therefore. and in one case for ten generations. as there also is between hybrids of varieties and hybrids of species. though Gärtner was enabled to rear some hybrids. This becomes much clearer after examining what Darwin and Mendel have to say about the possibility of obtaining new species by hybridization of preexisting ones (“hybridism in the narrow sense”).” Darwin observed: In regard to the sterility of hybrids in successive generations. Darwin referred to “those two conscientious and admirable observers. In the hybridism chapter in the Origin. “the whole subject of hybridism. Darwin also es- tablished that “there is no fundamental distinction between species and varieties” (Darwin 1861.. which “hybridism” (“in the narrow sense”) did seem to do. but generally greatly decreased. for Gärtner there is an essential difference between varieties and species.If Darwin Had Known Mendel 29 theory.50 As has been discussed.] is one of the greatest obstacles to the general acceptance and progress of the great principle of evolution” (Darwin 1878. Kölreuter and Gärtner.

could a pouter be produced by crossing two breeds unless one of the parent-stocks possessed the characteristic enormous crop? (Darwin 1861. referring to William Herbert’s (1778-1847) views. the doctrine is little less than absurd […]. Darwin wrote: Now let us turn to the results arrived at by the third most experienced hybridiser. they must have descended from at least seven or eight aboriginal stocks. the Hon. for instance. and have not proceeded from the rock-pigeon. namely. (Darwin 1861. Such a position inevitably led either to an infinite regress or to some version of the doctrine of special creation. but that they are all directly derived from certain aboriginal species. he noted: Great as the differences are between the breeds of pigeons. Herbert. though he accepted the existence of completely fertile and rela- tively stable hybrid forms of plants. In the case of the evolution of pigeons. The central reason was that hybridization presup- posed already existing differences. Darwin thought that without other sources of variation. for it is impossible to make the present domestic breeds by the crossing of any lesser number: how. He is as emphatic in his conclusion that some hybrids are perfectly fertile – as fertile as the pure parent-species – as are Kölreuter and Gärtner that some degree of sterility between distinct species is a universal law of nature. W. for example. If this means that new characters never spontaneously appear in our domestic races. In an early part of this chapter it was stated that Pallas33 [33 ‘Actes de l’Académie St. and Rev. including under this term several geographical races or sub-species. I will here briefly give them. As several of the reasons which have led me to this belief are in some degree applicable in other cases. which differ from each other in the most trifling respects. Petersburg. 270) On the other hand. for what made the parents or their progenitors different?” (Darwin 1868. part ii. But .30 pablo lorenzano experiments the fertility has been diminished by an independent cause. 252). namely. (Darwin 1861. vol. thus giving rise to the question of the origin of such differences. By trying to explain evolutionary change on the basis of crossings without variation “we thus only push the difficulty further back in time. that all have descended from the rock-pigeon (Columba livia). 271-272) Thus. II. If the several breeds are not varieties.’ 1780. 84. from close interbreeding. I am fully convinced that the common opinion of naturalists is correct. 23-24) This was in fact the fundamental objection to the doctrine of evo- lution solely by means of hybridization. also mentioned by Mendel. namely. hybridization by itself could not account for the evolution of species.] and a few other naturalists maintain that variability is wholly due to crossing. &c.

(Darwin 1861. eben so wenig ist es bis jetzt gelungen. 264) But Darwin was the first to admit the quandary. or in individuals of different species. is sometimes inherited and sometimes not so. a book that Mendel also read and underlined. This observation is a crucial key in interpreting his work.” presented in his book on variation in plants and animals (Darwin 1868). II. K ö l r e u t e r and G ä r t n e r.If Darwin Had Known Mendel 31 the doctrine may mean something widely different. 13-14) It was this conundrum that Darwin attempted to address with his “pro- visional hypothesis of pangenesis. in their investigations”. there is a clear distinction in Darwin between “hybridism without variation” and the theory of “descent with modifica- tion. no one can say why a peculiarity in different individuals of the same species. Nevertheless. and sometimes can hardly be 51 “So wenig man eine scharfe Unterscheidungslinie zwischen Species und Varietäten zu ziehen vermag. that the crossing of distinct species is the sole cause of the first appearance of new characters. 7) 52 “[…] die bei Pisum gemachten Beobachtungen mit den Resultaten zu vergleichen. more commonly but not exclusively to the like sex. K ö l r e u t e r und G ä r t n e r. why a peculiarity is often transmitted from one sex to both sexes.” (Mendel 1865. 38) . or to one sex alone. why the child often reverts in certain characters to its grandfather or grandmother or other more remote ancestor. Moreover. this explains why Mendel worked with hybrids of varieties and drew conclusions about an issue that was considered related to hybrids of species. bei ihren Forschungen gelangt sind. Mendel and Constant Hybrids Mendel. similar to Knight.”51 In fact. (Darwin 1868.” (Mendel 1865. and Darwin. The laws governing inheritance are quite unknown. he stated that “[i]t has so far been found to be just as impos- sible to draw a sharp line between the hybrids of species and varieties as between species and varieties themselves. In the conclusion to his 1865 paper. vol. the hybrids in outward appearance present either a form intermediate between the original species.” especially given his unequivocal support of the latter. whose purpose was “to compare the observations made regarding Pisum with the results arrived at by the two authorities in this branch of knowledge [hybridism]. did not believe in the existence of a clear-cut difference between species and varieties. or they closely resemble either the one or the other type. namely. zu welchen die beiden Autoritäten in diesem Fache.52 he noted: According to the opinion of both. einen gründlichen Unterschied zwischen den Hybriden der Species und Varietäten aufzustellen. Herbert.

Die Hybriden der Varietäten verhalten sich wie die Species- Hybriden. With regard to the form of the hybrids and their development. In der Regel behält die Mehrzahl der Individuen aus einer Befruchtung die Form der Hybride bei. nur besitzen sie eine noch grössere Veränderlichkeit der Gestalten und eine mehr ausge- sprochene Neigung. and some Dianthus hybrids. various forms which differ from the normal type [and that as] a rule. and one or other individual approaches the pollen parent. wenn die Befruchtung durch den eigenen Pollen geschah. oder sie neigen sich sämmtlich mehr nach der einen oder der anderen Seite hin. Aus den Samen derselben gehen gewöhnlich. It is otherwise with the exceptional cases cited. Sometimes the offspring have more nearly approached. according to Wichura. Mendel stated: This. In Bezug auf die Gestalt der Hybriden und ihre in der Regel erfolgende Entwicklung ist eine Übereinstimmung mit den bei Pisum gemachten Beobachtungen nicht zu verkennen. theils der anderen Stammpflanze näher gerückt. they remained identical and did not vary. Bei einzelnen sind die Nachkommen theils der einen. and. to this class belong the remarkably fertile hybrids Aquilegia atropurpureo-canadensis. 53 “Nach der übereinstimmenden Ansicht beider halten die Hybriden der äusseren Erscheinung nach entweder die Mittelform zwischen den Stammarten. Mendel continued in his characterization of the latter. Lavatera pseudolbio-thuringiaca.” (Mendel 1865. For the history of the evolution of plants this circumstance is of special importance.53 On the other hand.32 pablo lorenzano discriminated from it. The hybrids of varieties behave like hybrids of species. since constant hybrids acquire the status of new species. while in other cases they remain perfectly like the hybrid and continue constant in their offspring. the majority of individuals obtained by one fertilization maintain the hybrid form. or they all incline more to one or the other side. oder sie sind dem Typus der einen oder der anderen näher gerückt. According to Gärtner. as a rule an agreement with the observations made in Pisum is unmistakable. but they possess greater variability of form and more pronounced tendency to revert to the original types. bei einigen aber bleiben sie der Hybride vollkommen gleich und pflanzen sich unverändert fort. the variable ones that behave like those of Pisum and those he later re- ferred to as “constant”. zu den Stammformen zurückzukehren. if the fertilization was effected by their own pollen.54 Thus. 38) 54 “Das gilt jedoch nicht von allen Hybriden ohne Ausnahme. From their seeds usually arise. while some few others come more like the seed parent. 38-39. the hybrids of the Willow family. Anders verhält es sich mit den erwähnten Ausnahms-Fällen. verschiedene von dem normalen Typus abweichende Formen hervor. Geum urbano-rivale. however. We meet with an essential difference in those hybrids which remain constant in their progeny and propagate themselves as truly as the pure species. Mendel distinguished two kinds of hybrids for the first time.” (Mendel 1865. is not the case with hybrids without exception. manchmal von denselben kaum zu unterscheiden. während andere wenige der Samenpflanze ähnlicher werden und ein oder das andere Individuum der Pollenpflanze nahe kommt. Mendel’s emphasis) . some the one and some the other of the two original stocks.

only claim the value of an hypothesis for which the lack of definite data offers a wide scope. für welche bei dem Mangel an siche- ren Daten noch ein weiterer Spielraum offen stände.” The latter reproduce pure forms and achieve the status of a new species. den Dianthus Armeria-deltoides bis in die 10. die an der Pflanze vorgehenden Prozesse als Resultat der an den einzelnen Zellen vor sich gehenden Veränderungen zu erklären. Schleiden’s emphasis) 57 “[Die hier versuchte Zurückführung des wesentlichen Unterschiedes in der Entwicklung der Hybriden auf eine dauernde oder vorübergehende Verbindung der differirenden Zellelemente] kann selbstverständlich nur den Werth einer Hypothese ansprechen. Mendel’s emphasis) ..” (Mendel 1865. since it regularly propagated itself in the garden. Moreover. Gärtner hatte Gelegenheit. Mendel mentioned the “remarkably fertile hybrid” 55 “Einer wesentlichen Verschiedenheit begegnen wir bei jenen Hybriden. weil constante Hybriden die Bedeutung neuer Arten erlangen. Generation zu ver- folgen. It is interesting that.] dass im Wesentlichen das Leben der Pflanzen im Leben der Zelle enthalten seyn müsse […] jede Hypothese. however. as an example of constant hybrids.”57 What is important. welche nicht darauf abzielt. Along with a temporary union (vorübergehende Verbindung) of the dif- ferent cellular elements in variable hybrids. Mendel (1865. is that Mendel accepted that new species can originate from hybridization of preexisting ones. 40. Thus. 148. he supported the “new doctrine of special creation” advanced by Linnaeus. 146. da sich derselbe regelmässig im Garten von selbst fortpflanzte. of course. 42. 40-42) noted the adoption of the second of Schleiden’s maxims. Die Richtigkeit des Sachverhaltes ist durch vorzügliche Beobachter verbürgt und kann nicht in Zweifel gezogen wer- den.” (Mendel 1865. and cannot be doubted. Lavatera pseudolbia-thuringiaca. Maxime der Selbstständigkeit der Pflanzenzelle. nach Wichura die Hybriden der Weidenarten.. there can also be a lasting union (dauernde Verbindung) in constant hybrids.” (Schleiden 1849.If Darwin Had Known Mendel 33 The correctness of the facts is guaranteed by eminent observers. even when this expla- nation “can. jede Induktion in der Botanik ist unbedingt zu verwerfen. […] essentially the life of plants must be contained in the life of cells […] every hypothesis. Nach Gärtner ge- hören hieher die ausgezeichnet fruchtbaren Hybriden: Aquilegia atropurpurea-canadensis. Für die Entwicklungsgeschichte der Pflanzen ist dieser Umstand von be- sonderer Wichtigkeit.56 Mendel attempted to explain the differences between these two kinds of hybrids in terms of different types of unions in germ and pollen cells. Mendel’s emphasis) 56 “B. B. and contrary to Gärtner and Kölreuter.55 Mendel believed there was an essential difference between the hy- brids of Pisum and those he calls “constant. Gärtner had an opportunity of following up Dianthus Armeria-deltoides to the tenth generation. welche in ihren Nachkommen constant bleiben und sich eben so wie die reinen Arten fortpflanzen. Geum urbano-rivale und einige Dianthus-Hybriden. every induction that does not purport to explain the processes that occur in the plant as a result of the changes that take place in individual cells must be rejected unconditionally. Maxim of the autonomy of cells in plants. [.

Hoffman returns to Gärtner’s original to support his position. Ebenso Kölreuter. Erz.61 Thus. Brünn. which are the places where Gärtner refers to the mentioned hybrids. 689). From Geum urbanum + rivale Gärtner seems to have obtained remarkably fertile and constant hybrids (according to Mendel.” (Gärtner 1849. Verh. die Zeugungskraft nach und nach abnimmt und das Decrepidiren eintritt. [e]ven when very fertile hybrids reproduce regularly in a spontaneous way for 8-10 generations. After claiming that “it remained without change in type until the tenth generation and that it reproduced by itself every year in the garden until the six or even eight first generations.”62 As for Dianthus Armeria-deltoides. this was an erroneous judgment on his part. bis sie endlich steril werden und ausgehen. as “al- ways with gradually decreasing fertility and general decrepitude of the species. Ich finde dies nicht bestätigt bei der Lektüre des Originals (Bast. nat. 59 “Aus Geum urbanum + rivale soll Gärtner ausgezeichnet fruchtbare und konstante Bastarde erzogen haben (nach Mendel.34 pablo lorenzano obtained by Gärtner. Ver. as Hoffman pointed out. 40). “the most prominent ex- ample” according to Gärtner63 and that Mendel specially mentioned in the previous passage. 112) 60 “Selbst wenn sich verhältnißmäßig sehr fruchtbare Bastarde auch 8 — 10 Generationen spontan fortpflanzen. 365) 62 “[…] doch immer mit allmählich abnehmender Fruchtbarkeit und allgemeinem Decrepidiren der Art. until they finally become sterile and extinguish. 365. the reproductive force diminishes little by little and decrepitude comes in.”64 Gärtner added that “fertil- 58 In relation to that.” (Gärtner 1849.60 In fact. Mendel writes on the inner side of the cover of Gärtner (1849) “553 Über Stabilarten” (“553 On stable species” and “constt 421” (“constt” for “constant”) and on the inside of the back cover again “p. 40).” (Gärtner 1849.” (Hoffmann 1869. 422) 63 “Das auffallendste Beispiel. But. not as acquiring the status of a new species but. 40) 61 “[…] selbst bei den fruchtbarsten. Ver. the reference was likewise tendentious and incom- plete. hist. Brünn. even in the most fertile [hybrids]. when they reproduce by themselves until the eighth and tenth generation. I don’t find this established in a reading of the original (Bastarderzeugung.) Vgl. they nonetheless become decrepit and extinguish! Compare below Dianthus. 1865. IV. IV. 553) 64 “[…] welcher bis in die zehnte Generation ihne Veränderung des Typus erhalten und sich sogar in . so werden sie doch decrepid und gehen aus! (S.59 Hoffmann continued to explain why he reached this conclusion. Gärtner considered the remarkably fertile hybrids that Mendel mentioned. Unten bei Dianthus. hist. nat. Verh.” (Hoffmann 1869.58 along with the hybrids of willow obtained by Wichura. wenn sie sich auch bis in die achte bis zehente [sic] Generation selbst fortpflanzen. S. 698).” along with “constt 421” and “422!” in ink. 553.

” (Gärtner 1849. und eine stäte Fortbildung der Gewächsarten annehmen.66 However.” (Gärtner 1849. so findet sich doch anderseits in den von Gärtner angestellten Versuchen eine beachtenswerthe Bestätigung der früher über die Veränderlichkeit der Culturpflanzen ausgesprochenen Vermuthung. bis seine Zeugungskraft im zehnten Jahr völlig erloschen war” (Gärtner 1849.If Darwin Had Known Mendel 35 ity of the seeds decreased however with each generation. 553). chinensis und japonicus.67 and adds another fragment that was incomplete and modified.bis 5maligen hybriden Verbindung nichts von ihrer Selbständigkeit verlo- ren. by the results of these transformation experiments.68 The latter is important because actually Gärtner adds that the hybrid “must return to its original form or disappear. and hybrids between these species lost none of their stability after 4 or 5 generations. dass der Pflanzenspecies feste Grenzen gesetzt sind.”65 Mendel completed his famous paper with his own assessment of Gärtner. Es sieht in der vollendeten Umwandlung einer Art in die andere den unzweideutigen Beweis. über welche sie sich nicht ver- ändern kann. was led to oppose the opinion of those naturalists who dispute the stability of plant species and believe in a continuous evolution of vegetation. Dianthus Caryophyllus. He perceives in the complete transformation of one species into another an indubitable proof that species are fixed with limits beyond which they cannot change. 475) . Although this opinion cannot be unconditionally accepted we find on the other hand in Gärtner’s experiments a noteworthy confirmation of that supposition regarding variability of cultivated plants which has already been expressed. His treatment begins with a statement from Gärtner without attribution. Gärtner 1849. such as Aquilegia atropurpurea and canadensis. chinensis y japonicus. 66 “Durch den Erfolg der Umwandlungs-Versuche wurde Gärtner bewogen. Nicotiana rustica und paniculata. until its repro- ductive force was completely extinguished in the tenth year. welche die Stabilität der Pflanzenspecies bestreiten und eine stäte Fortbildung der Gewächsarten annehmen. 553) 65 “[…] dessen Fruchtbarkeit in Samen aber mit jeder Generation verminderte. 475) 68 “[…] wir finden in der wirklichen Umwandlung einer Pflanzenart in eine andere den unzwei- deutigen Beweis. über welche hinaus sie sich nicht zu ändern vermag. 46-47) 67 “who dispute the stability of plant species and believe in a continuous evolution of vegeta- tion” ([…] welche die Stabilität der Pflanzenspecies bestreiten. Gärtner.”.” (Mendel 1865. Unter den Versuchsarten kommen cultivirte Gewächse vor. dass der Species feste Grenzen gesteckt sind. Wenn auch dieser Ansicht eine bedingungslose Geltung nicht zuerkannt werden kann. wie Aquilegia atropurpurea und cana- densis. Nicotiana rustica and paniculata. The transformation of one vegetable species into another by means of sexual reproduction does seem to us to establish beyond doubt the necessity and peculiar- ity of the nature of vegetable species and their stability by means of den sechs bis acht ersten Generationen alle Jahre im Garten selber ausgesät hatte. Mendel used a liberal interpretation of Gärtner’s text. sich gegen die Meinung derjenigen Naturforscher zu kehren. Among the experimental species there were cultivated plants. und auch diese hatten nach einer 4. Dianthus Caryophyllus.

Additionally. He held this position even though he was aware of an apparent exception mentioned by Dominique Alexandre Godron (1807-1880. there is the same distinction drawn between two kinds of hybrids as there was in the first paper. Moreover. § 98) 71“[…] dass die Species in allen ihren Eigenschaften auf gewisse äussere Verhältnisse berechnet und angelegt ist. the only authors Mendel mentioned were Herbert and Henri Lecoq (1802-1871). Mendel partially reproduced Gärtner’s views. but attempted to show that the suppositions of Linnaeus that “all species owe their origin to an act of special creation. keineswegs im Widerspruche steht. 1861).36 pablo lorenzano internal forces. Godron 1855.” (Gärtner 1849. Regarding the question whether and to what extent hybridisation plays a part in the production of this wealth of forms. interpreting them as evidence of “hybridism (in the narrow sense). 475) 70 “Die Zunahme der Schwäche und Unfruchtbarkeit und das baldige Aussterben der Bastarde bei fortgesetzter Befruchtung derselben durch den eigenen Pollen. 1858.” of Darwin in his “theory of the adaptation of species. towards the end of his book.” and that of “the species is determined and organized in all its properties by certain external conditions. This was due to “the increase in weakness and infertility of the hybrids and their quick extinction by fertilization with their own pollen”70 which he considered a rule. Die Umwandelung einer Pflanzenart in eine andere durch die geschlechtliche Zeugung scheint uns daher die Nothwendigkeit und die Eigenthümlichkeit der Natur der Pflanzenspecies und ihre durch innere Kräfte gesicherte Stabilität ausser Zweifel zu setzen. § 94) .).” contrary to Wichura’s own exposition and analysis.”69 Thus.” In Mendel’s second paper on plant hybrids from 1869. Gärtner. only to question them later when he wrote that “this opinion cannot be unconditionally accepted.”71 In addition to Kölreuter. and Wichura. Herbert and Lecoq were two of several hybridists Gärtner and others referred to as in favor of the “new doctrine of special creation. where he reports in the last paragraph of the paper on Gärtner’s experiments in which the pro- cess of transformation takes place (Gärtner 1849. Mendel presented them as examples of hybrids that remained constant and reproduced pure. thus achieving the status of new species.” Therefore. where artificially produced hy- brids kept their reproductive ability. 463sqq. Wichura not only summarized his results. § 99). but the plant had not been found in nature (Wichura 1865. we find very various and conflicting views 69 “[…] sondern zur Urform zurückkehren oder untergehen muss.” (Wichura 1865. Wichura considered that hybrids did not reproduce constant indefinitely. are not in contradiction.” (Wichura 1865. when he referred to Wichura’s results with willow hy- brids (Wichura 1853/1854. 1865). Mendel presented quite a different evaluation of Gärtner’s conclusions than Gärtner made himself.

On these grounds it is regarded as inconceivable that Hieracium hybrids can constitute and maintain themselves as fully fertile and constant forms when growing near their progenitors. on the ground that they are only of short duration. dass Hieraciumbastarde sich in der Nähe ihrer Stammeltern zu vollkommen fruchtbaren und constanten Formen herausbilden und behaupten könnten. consciously because he thought that with the latter’s help he could prove the existence of constant hybrids. As he stated in one of the letters to the great Swiss botanist: The hybrid Geum urbanum + rivale deserves special attention. . […]. z. dass bei Bastarden die Selbstbefruchtung immer ausgeschlossen werde. dass dieselben aus der Transmutation untergegangener oder noch bestehender Arten herzuleiten seien. others. dass Bastarde unter den wildwachsenden Arten nicht selten gebildet werden. While some of them maintain that this phenomenon has a far-reaching influence. belongs to the few known hybrids which produce nonvariable progeny as long as they remain self-pollinated. or whether by descent with modification. 27-28) 73 “Eine grössere Aufmerksamkeit ist auch dem Bastarde Geum urbanum + rivale zugedacht. seitdem ein berühmter Hieracienkenner im Geiste der Darwin’schen Lehre die Ansicht vertritt. partly also the knowledge. wollen andere. The [suggested] causes of this are partly their restricted fertility or complete sterility. dass denselben eine wich- tigere Bedeutung aus dem Grunde nicht beizumessen sei. bei Hieracien von Bastarden überhaupt nichts wissen. Fries. obtained by experiment. Die Frage über den Ursprung der zahlreichen constanten Zwischenformen hat in neuester Zeit nicht wenig an Interesse gewonnen. theils aber in der durch Versuche erwiesenen Erfahrung. Noch andere nehmen eine vermittelnde Stellung ein und geben zu. if hybridized. or whether a hybrid that appeared naturally and exhibited some degree of fertility would return to its parental forms were the crucial issues for Mendel’s work with Hieracium. Es sei demnach undenkbar. ob und in welchem Umfange die Bastardbildung an dem Formenreichthum des genannten Geschlechtes Antheil nimmt. This plant. still maintain that no great importance is to be attached to the fact. for example.72 Whether intermediate forms of Hieracium appeared as natural constant hybrids. would behave in a fashion similar to Geum. Fries. Während einige derselben einen weit reichen- den Einfluss zugestehen.If Darwin Had Known Mendel 37 held by leading botanists. behaupten jedoch. is perhaps not without foundation. The surmise that some species of Hieracium. weil sie immer nur von kurzem Bestande sind. will have nothing to do with hybrids in Hieracia. An additional consideration was that Mendel chose Hieracium. that in hybrids self-fertilization is always prevented if pollen of one of the parent-forms reaches the stigma. begegnen wir unter den ersten Pflanzenkennern sehr ab- weichenden.” (Mendel 1869. a genus on which Nägeli was a well-known specialist. and while granting that hybrids are not rarely formed between the species in a wild state.73 72 “Bezüglich der Frage. The question of the origin of the numerous and constant intermediate forms has recently acquired no small interest since a famous Hieracium specialist has. defended the view that these forms are to be regarded as [arising] from the transmutation of lost or still existing species. Die Ursache davon liege theils in der geringen Fruchtbarkeit oder gänzlichen Sterilität derselben. sogar völlig widersprechenden Ansichten. wenn der Pollen der Stammarten auf die Narben derselben gelangt. according to Gärtner. in the spirit of the Darwinian teaching. Others take up an intermediate position. B.

Mendel not only reported on the progress of these experiments and their comparison with those obtained in Pisum in the 1869 paper. wenn man die aus der Beobachtung einiger anderer Bastarde abgeleiteten Regeln schon für Gesetze der Bastardbildung ansehen und ohe weitere Kritik auf Hieracium ausdehnen wollte. What Mendel assumed he could determine with the experiments on Hieracium was that constant hybrids could be obtained and that they could be identified with the intermediate constant forms that occur naturally. and we may be led into erroneous conclusions if we take rules deduced from observation of certain other hybrids to be Laws of hybridization. Pisum) might not be applied directly to Hieracium. dann wird mit Zuhilfenahme der Erfahrungen.38 pablo lorenzano Then. Das Verhalten der Hieracium-Bastarde in dem angedeuteten Umfange muss nothwendig durch Versuche ermittelt werden. Mendel continued his attention on Hieracium. 28) . den mögli- cherweise die Bastardbildung auf die Mannigfaltigkeit der Zwischenformen bei Hieracium ausübt. da wir eine abgeschlossenen Theorie der Bastardbildung nicht besitzen. If by the experimental method we can obtain a sufficient insight into the phenomenon of hybridization in Hieracium. 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 offspring through several generations no one can undertake to determine the possible influence exercised by hybridization over the multiplicity of intermediate forms in Hieracium. He identifies three im- Diese Pflanze gehört nach Gärtner zu den wenigen bisher bekannten Hybriden.g.” (Mendel 1866. ein competentes Urtheil in dieser Frage möglich werden. wenn man es unternehmen will. wenn die Befruchtung durch den eigenen Pollen geschieht. 196-194) 74 “Es liegt in der Sache. he also shared them in correspondence with Nägeli. und es zu irrigen Anschauungen führen könnte. a satisfactory judgment in regard to this question may become possible.” (Mendel 1869. and try to apply them to Hieracium without further consideration. dass eine genaue Kenntnis der Bastarde in Bezug auf ihre Gestalt und Fruchtbarkeit. after delineating the problem of intermediate forms and its significance in relation to evolution. then by the help of the experience which has been collected respecting the structural relations of the wild forms. den Einfluss zu beurtheilen. since he realized there was not an universal theory of hybridization so that the rules found in other hybrids (e. Gelingt es auf dem Wege des Experimentes eine genügende Einsicht in die Bastardbildung der Hieracien zu erhalten. um die es sich hier handelt. dass manche Hieracium-Arten in hybrider Verbindung ein ähnliches Verhalten wie Geum beobachten. for we do not possess a complete theory of hybridization. sowie auf das Verhalten ihrer Nachkommen durch mehrere Generationen unerlässlich ist. welche über die Vegetationsverhältnisse der verschiedenen wild wachsenden Pflanzen gesammelt wurde. welche in ihren Nachkommen unverändert bleiben.74 Mendel intended to approach the problem through an analysis both of experiments and of wild plants. The condition of the Hieracium hybrids in the range we are concerned with must necessarily be determined by experiments. […] Vielleicht ist die Vermuthung nicht ganz unbegründet.

” (Mendel 1869. allerdings noch sehr unsicheren Resultate mit jenen vergleichen. und welche ich im Jahre 1865 hier mitzutheilen die Ehre hatte. which I had the honour of communicating in the year 1865. B. obtained from the immediate crossing of two forms. in allen Fällen den gleichen Typus.”75 That is. dass es auch Bastarde gibt.If Darwin Had Known Mendel 39 portant differences between the hybrids of the two species. still very uncertain. but their posterity. ihre Nachkommen dagegen sind veränderlich und variiren nach einem bestimmten Gesetze. If finally we compare the described result. have in all cases the same type. Already in describing the Pisum experiments it was remarked that there are also hybrids whose posterity do not vary. dass z. with those obtained by crosses made between forms of Pisum. Whether from this circumstance we may venture to draw the conclusion that the polymorphism of the genera Salix and Hieracium is connected with the special condition of their hybrids is still an open question. which may well be raised but not as yet answered. are variable and follow a definite law in their variations. dass die Polymorphie der Gattungen Salix und Hieracium mit dem eigentlichen Verhalten ihrer Bastarde in Zusammenhang stehe. on the contrary. nach Wichura die Bastarde von Salix sich unverändert wie reine Arten fortpflanzen. In Pisum the hybrids. Schon bei Besprechung der Pisum-Versuche wurde darauf hingewiesen. das ist bis jetzt noch eine Frage. deren Nachkommen nicht variiren. In Hieracium we may take it we have a similar case. contrary to what happened with Pisum. welche aus Kreuzungen zwischen Pisum-Formen erhalten wurden. 31) .77 75 “Bezüglich der Gestalt der Bastarde haben wir die auffallende Erscheinung zu registriren. and that.” (Mendel 1869. Bei Hieracium scheint sich nach den bisherigen Versuchen das gerade Gegentheil davon herausstellen zu wollen. for example. dass die bis jetzt aus gleicher Befruchtung erhaltenen Formen nicht identisch sind. 29) 77 “Wenn wir schliesslich die besprochenen. so he con- cluded: The conviction is then forced on us that we have here only single terms in an unknown series which may be formed by the direct action of the pollen of one species on the egg-cells of another. so begegnen wir einer sehr wesentlichen Verschiedenheit.” (Mendel 1869. welche durch die unmittelbare Einwirkung des Pollens der einen Art auf die Keimzellen einer andren gebildet werden. we find a very real distinction. nicht aber beantworten lässt. dass wir hier nur einzelne Glieder aus noch unbekannten Reihen vor uns haben.76 The other two differences are contained in the final part of the 1869 paper. welche unmittelbar aus der Kreuzung zweier Formen gewonnen wer- den. the reciprocal crossings were not identical. we have to record the striking phenomenon that the forms hitherto obtained by similar fertilisation are not identical. according to Wichura the hybrids of Salix reproduce themselves like pure species. die sich wohl anregen. 29) 76 “Es drängt sich von selbst die Vermuthung auf. The first distinction noted that “[r]especting the structure of the hybrids. In Hieracium according to the present experiments the exactly opposite phenomenon seems to be exhibited. Ob man bei diesem Umstande die Vermuthung aussprechen dür- fe. Wir hätten demnach bei Hieracium einen analogen Fall. Bei Pisum haben die Bastarde.

the “new doctrine of special cre- ation” or “hybridism (in the narrow sense). Consequently. and to deal directly with the counterfactual in the title. On this occasion I can not resist remarking how striking it is that the hybrids of Hieracium show a behavior exactly opposite to those of Pisum.” and even thought to have found support in his experiments with Hieracium for the idea that new species can originate by hybridization from preexisting ones.” Conclusions The intent of this study was to discuss the question raised by the title “what would have happened if Darwin had known Mendel (or Mendel’s work)?” To clarify this counterfactual question and to put to rest the origin of such explanation. just as with Darwin. Mendel made the following concluding comment to Nägeli. he lists second and third differences: the first generation of hybrids of Pisum is uniform. established the existence of constant hy- brids that reach the status of new species. the latter was a great critic of the idea and devised an alternative explanation of the origin of species: Darwin’s theory of evolution by means of natural selection. established “hybridism (in the narrow sense). the central problem Mendel faced belonged to evolutionary biology. law.” (Mendel 1870a. Wir haben es hier offenbar nur mit einzelnen Erscheinungen zu thun.40 pablo lorenzano Here. thus. But while the former expressed himself in favor of a theory alternative to that of “descent with modification. Evidently we are here dealing only with individual phenomena that are the manifestation of a higher. In the following sections. what is more important is that Mendel thought he had contributed an alternative to the evolutionary explanation of the origin of constant intermediate forms in the genus Hieracium and.78 Mendel never formulated the “more fundamental” law. wie sehr es anffallen muss. 233) . he demonstrated the origin of new species by means of hybridization from preexisting ones and. In other words. it has been argued that Mendel did not meet Darwin on his trip to London in 1862 nor did Darwin possess Mendel’s best-known work on hybrids of plants as an offprint with its pages uncut. hence. whereas those of Hieracium are constant. the relationship between Mendel and Darwin was placed in a different perspective. Thus. the offspring of hybrids of Pisum are variable. if 78 “Ich kann bei dieser Gelegenheit die Bemerkung nicht unterdrücken. more fundamental. that of Hieracium is not. dass die Bastarde von Hieracium im Vergleich mit jenen von Pisum ein geradezu entgegengesetztes Verhalten beobachten.” namely. die der Ausfluss eines höheren allgemeinen Gesetzes sind. But.

but that between embryology and evolution. based on that. Sandler 2000) have pro- posed an alternative interpretation on Mendel’s experiments and. considering the good manners of an English gentleman as well as those of a farmer’s son and educated Augustinian monk. he stressed Mendel’s rejection of the importance of changes in the conditions of life for variability – “the very essence of Darwin’s conception of variation” (Olby 2009. but there would not have been the beginning of a synthesis between what was to be regarded as Darwinism and Mendelism in the 1930s and 40s. They have pointed out the nineteenth cen- tury inseparable linkage between heredity and development.If Darwin Had Known Mendel 41 Darwin had read Mendel’s text (or texts). would have had to explain in a unitary fashion not only the transmission of char- acters from parent to offspring. he would have considered him just another “hybridist. exclamation mark in the original). The discussion would have been polite and maybe a bit passionate about evolutionary subjects. . Besides he points out that however “Mendel’s results were established with strongly marked (nonblending) characters. however. 45). they would have enjoyed had a cup of tea served from the china produced by Darwin’s wife’s family or a succulent bowl of peas from Brünn’s monastery. in Darwin’s opinion.” A similar position to ours is defended by Olby (2009). 44-45). and cites Alfred Russel Wallace. But he would have also considered Mendel to be an opponent of his fundamental convictions and not a provider of that which his theory lacked. Such were not. a work that could not have contained a theory of inheritance” (Sandler and Sandler 1986. The fact that “Gregor Mendel’s 1866 paper dealt entirely with the subject of the transmission of inherited traits (that is. that Mendel’s work is better placed in an evolutionary context. is not that between heredity and development. cited by Olby 2009. 43) – as well as Darwin’s critique of hybridization as “the source of the resulting variation” (Olby 2009. 1986. We think having pro- vided enough evidence in support of the ideas that the relevant ambiguity.79� 79 Iris Sandler and Laurence Sandler (Sandler and Sandler 1985. and also tak- ing into account their shared interests. 45) and claimed that “hybridization […] had no place whatever in the natural process of species-formation” (Wallace 1908. Furthermore. in a way that “a theory of heredity. but also the embryological development of those characters” (Sandler and Sandler 1985. 755) might explain the possible lack of interest of Darwin in Mendel’s work. Based on an analysis of Darwin’s conceptions of variation and inheritance. a satisfactory explanation of the ori- gin and inheritance of the variations upon which natural selection oper- ates. 135. in the middle of the nineteenth century. on why Darwin might not have been interested in Mendel’s work in the first place. 38). that is. to be recognized as such. but to an alter- native theory of speciation – hybridism (in the narrow sense) – to his own theory – of “descent with modification. and that Darwin’s possible reaction to Mendel’s theory might have been not to an incomplete theory of heredity. but to an as yet unformulated question!” (Sandler and Sandler 1985. 11). 69. who “belittled Mendel’s achievement” (Olby 2009. for a better understanding of Mendel’s work. defined his problem in purely genetic terms. and produced a correct and amazingly complete answer. with Genetics) and therefore would have seemed to his contemporaries a partial work.” though perhaps a very good one on account of his experimental skills and for having provided a statistical treatment of his results (which is how Mendel was regarded by those contemporaries that got to know his work). if a personal encounter between Mendel and Darwin occurred. and claimed that “Mendel. relevant to evolution” (Olby 2009.

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