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Czech Journal of Genetics and Plant Breeding, 58, 2022 (3): 113–126 Review
https://doi.org/10.17221/19/2022-CJGPB
Interspecific hybridization and plant breeding:
From historical retrospective through work of Mendel
to current crops
1 2 3
David Kopecký , Antonio Martín , Petr Smýkal *
1
Institute of Experimental Botany, Czech Academy of Sciences, Olomouc, Czech Republic
2
Instituto de Agricultura Sostenible IAS-CSIC, Cordoba, Spain
3
Department of Botany, Faculty of Sciences, Palacky University, Olomouc, Czech Republic
*Corresponding author: petr.smykal@upol.cz
Citation: Kopecký D., Martín A., Smýkal P. (2022): Interspecific hybridization and plant breeding: From historical retrospecti-
ve through work of Mendel to current crops. Czech J. Genet. Plant Breed., 58: 113–126.
Abstract: There is a relatively long history of plant hybridization traced back to ancient time, both from theoretical
as well as practical perspectives. At first considered as an evolutionary dead-end, it was soon recognized to have im-
portant role in plant speciation. Beside his work on pea, G.J. Mendel also conducted interspecific hybridization using
several species including Hieracium. Current knowledge shows that the frequent occurrence of wide hybridization
in nature is often connected with polyploidy. Interspecific hybridization has played a role in plant domestication and
numerous crops are allopolyploids, sometimes of complex hybrid origin. This has been also used in practical breeding,
extending even to intergeneric crosses which benefit from heterosis, transgressive segregation and introgression phe-
nomenon. This review aims to provide a a historical retrospective and summarize both current knowledge and the
usage of interspecific hybridization in crop breeding.
Keywords: allopolyploid; breeding; crop; domestication; heterosis; hybridization; interspecific hybrids
Historical retrospective on plant hybridization fruits is given by Theophrastus (c. 371–287 B.C.)
in Historia plantarum and De causis plantarum. He
The history of plant hybridization is related to the described the practice of artificial pollination of the
identification of male and female components of the female date tree by the inflorescence of the male,
flower. The earliest historical evidence regarding arti- and compared it to the sexual process among fish.
ficial pollination is found in Akkadian Code of Ham- In his work, Theophrastus used the terms ‘male’ and
murabi (18 century B.C.) where specific terms for ‘female’ in the modem sense (Negbi 1995).
the inflorescences of the female and male date trees During the Middle Ages, there was no novelty
and for fertilization were used. The artistic depiction in view of plant hybridization until Alonso de Her-
th rera published the first modern treatise on agri-
of artificial pollination appears first in the 9 century
B.C. in Aramaic relief from Tell Halaf in northeastern culture in his Obra de Agricultura (1513; ‘Treatise
Syria, depicting a man climbing a date palm, appar- on Agriculture’). This thesis showed that many cur-
ently to fertilize the female tree by applying the male rent breeding methods had already been developed
inflorescences to the flowers. at that time. He refered to the Roman agronomists,
The first written records on the structure and Virgilius, Varro, Plinius and Columella in particu-
function of plant flowers and their relation to the lar. Moreover, Herrera provided recommendations
Supported by the Czech Science Foundation (Grant award 20-10019S), and by the Grant Agency of the Czech Republic
and Palacký University Grant Agency (IGA 2022-002).
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Review Czech Journal of Genetics and Plant Breeding, 58, 2022 (3): 113–126
https://doi.org/10.17221/19/2022-CJGPB
about the seeds to be used for sowing cereals. For and provided examples of 17 bigeneric (intergeneric)
example, he recommended taking the grains from crosses and 17 congeneric (interspecific) crosses.
the bottom of the spike, which is similar to current Antoine Nicolas Duchesne (1766), while work-
bulk selection methods. He assumed that they will ing with a strawberry collection, documented the
perform better due to their heavier weight. He also separation of sexes in wild strawberry and identified
recognized the influence of environment in stating the garden strawberry (Fragaria × ananassa) (Dar-
that ‘no good crops are to be expected from poor seed row 1966), demonstrating that species can change.
unless favoured by good growing conditions;’ and the The 1849 work of Carl Friedrich Gärtner, Ver-
importance of specific adaptation when he wrote that suche und Beobachtungen über die Bastarderzeugung
the seed has to be harvested from similar conditions im Pflanzenreich (The Production of Hybrids in the
to those where seed was to be grown ‘from hot to hot, Plant Kingdom), contained a summary and discussion
from cold to cold, from dry to dry, from mild to mild, on the knowledge about plant hybridization (Gärtner
from humid to humid, …’. 1849). Gärtner himself noted both the uniformity
The advent of the modern history of plant hybrid- of the first hybrid generation and the diversity of the
ization can be dated back to the work of Rudolph forms in the second and successive generations. He
Jacob Camerarius. He contributed particularly stated that both parental types and entirely new
to the investigation of sexual differentiation in plants ones reappeared in these later generations and that
by identifying and defining the male (anther) and the variability was found in all of the characteristics
female (pistil) reproductive parts of the plant and of the progeny. A few years later in 1863, Charles
also by describing their function in fertilization, Naudin contrasted the uniformity of the first hy-
showingthat pollen is required for this process brid generation with ‘the extreme medley of forms’
(De sexu plantarum, 1694; ‘On the sex of plants’), in the second generation, ‘with some approaching the
and in Opuscula botanica (1697; ‘Botanical Works’). specific type of the father, others that of the mother’
Although it is Camerarius, who is most commonly (Harvey 2003). The first scientific reports targeting
considered the first author to describe plant sexuality, exclusively plant hybridization were the publications
it was actually Adam Zalužanský ze Zalužan who of Joseph Gottlieb Kölreuter about hybridization
defined male, female and bisexual plant individuals experiments (Vorlaufige Nachricht von einigen das
(dioecious and monoecious plants) for the first time Geschlecht der Pflanzen betreffenden Versuchen und
in the pioneering chapter ‘De Sexu Plantarum’ of his Beobachtungen (1761), Fortsetzung (1763), Zweyte
book Methodus Herbaria, Libri Tres from 1592. Fortsetzung (1764), and Dritte Fortsetzung (1766).
He also stressed that the individuals of both sexes Kölreuter demonstrated that hybrids from inter-
formed the same species and should not be classified specific crosses are often sterile. As a result, he
as different taxa or varieties (Funk 2013). concluded that hybrid plants are produced only
The first written reference on spontaneous plant with difficulty and are unlikely to occur in nature
hybridization was by Cotton Mather in 1716, de- without human intervention or disturbance of the
scribing crosses between Indian and yellow corn, and habitat. He also showed that F hybrids are usu-
1
between gourds and squash (Zirkle 1934). In the same ally morphologically intermediate relative to their
year, Thomas Fairchild reported the first artificial parents and that successive generations tend to re-
hybrid from a cross between carnation (Dianthus vert back to the parental forms (Mayr 1986). This
caryophyllus) and sweet William (Dianthus barbatus) discovery refuted an earlier suggestion by Linnaeus
(reviewed in Roberts 1929). Contrary to common that hybrids were constant or true-breeding and
belief, Fairchild was distraught by his success because represented new species. The character of hybrids
he regarded all plant species as created by God at the had been explained from the viewpoint of taxonomy
time of Creation. He thus feared the consequences and fertility, a practice common among the botanists
of disturbing this natural order, as many others also of that time (in Roberts 1929). Kölreuter made over
thought in his time. Therefore, his work is largely 500 crosses involving 138 species such as Nicotiana,
unknown and many authors ascribe the first artificial Dianthus, Matthiola, Hyoscyamus and Verbascum.
hybrid to C. Linnaeus experiments on Tragopogon Although he often found sterility, there were cases
in 1759 (reviewed in Roberts 1929). It was in his Plan- of fertile hybrids displaying heterosis (hybrid vigour)
tae Hybridae (1753), where he initiated the discussion not only in vegetative growth, but also in number
about the role of hybridization in plant speciation and size of flowers. By making reciprocal crosses
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Czech Journal of Genetics and Plant Breeding, 58, 2022 (3): 113–126 Review
https://doi.org/10.17221/19/2022-CJGPB
Kölreuter disproved the ancient theory of the dif- ments, often aiming for practical outcomes. For
ferent contributions of male and female parents. example, R. Geshwind applied hybridization to for-
Due to his interest in flower structure and pollina- est trees as well as to breeding roses. Actually, in his
tion, he examined the shape, size and color of the first experiments that date to 1845, prior to work
pollen grains from over 1 000 species. Kölreuter’s of Mendel on pea, he obtained hybrids of Pyrus and
attention to detail was remarkable. For example, he Sorbus trees (in Roberts 1929). He noticed that not
counted 3 154 flowers in Verbascum hybrids, and only new varieties, but also new species can occur
4 863 pollen grains in a single Hibiscus flower and among offspring upon self-pollination.
subsequently tested how many are needed for fer-
tilization. Possibly this inspired Mendel in his work. Mendel’s interest on species evolution
In contrast to his progressiveness in experimentation, (speciation)
Kölreuter was rather conservative in his conceptual
framework. He complied to common beliefs in a cre- Gärtner’s and Kölreuter’s work very likely influ-
ated and well-designed world. As a consequence, he enced Mendel’s experimental set up as well as his
admired sterility of hybrids, which supported view scientific work. Fortunately, he decided not to reca-
on sharply separated species in the world. In his view, pitulate the experiments, but applied the knowledge
if the hybrids were fertile and frequent, then there to intraspecific crosses with pea, likely inspired
would be confusion and disorder in nature. A second by previous results of Thomas A. Knight (1799).
major concept which shaped Kölreuter’s thinking was We know that after Pisum (Mendel 1866), Mendel
the assumption that laws in biology are the same as in investigated many other species from the genera
chemistry and physics. He viewed fertilization, the mix- Aquilegia, Antirrhinum, Calceolaria, Campanula,
ing of male and female gametes, as chemical process Cheiranthus, Cirsium, Dianthus, Geum, Hieracium,
resulting in traits blending. Together with essentialism Ipomoea, Linaria, Lychnis, Matthiola, Mirabilis,
this influenced his theory of inheritance. Kölreuter Phaseolus, Tropaeoleum, Verbascum and Zea (Cetl
insisted that essences are uniform, and the material 1973). By far, the largest number of experiments was
of the two parents is blended into intermediate state conducted in Hieracium (Mendel 1870), which have
after hybridization. Hybrids should be therefore exactly been often interpreted as failure. Recently, van Dijk
intermediate between parents. Thus, the observed and Ellis (2016) have shown that the assumption
exceptions puzzled him. He explained them as a result of Mendel’s studies on Hieracium arised from a mis-
of irregular or incomplete blending. However, he was understanding that could be explained by a missing
fair in admitting that the results of his experiments page in Mendel’s first letter to Carl Nägeli. Mendel’s
conflicted his fundamental beliefs (Mayr 1986). The writings clearly indicated his interest in ‘constant
dominating opinion during his time was that an off- hybrids’, hybrids which do not segregate and which
spring was already pre-formed in the female or the were ‘essentially different’ from ‘variable hybrids’ such
male and that the embryo was developed after sex and as those in Pisum. Thus, it is argued that Mendel’s
the origin predetermines the offspring’s characteristics main motivation for the Hieracium experiments was
or similarities to the parent (Bentley 1960). his interest in hybridization and speciation, rather
Kölreuter found that in general, only closely related than the inheritance of traits. Notably, even before
plants, and not always even these, can be crossed. the publication of Darwin’s ‘Origin of Species’ in 1859,
He concluded that the continued self-pollination C. Nägeli accepted that species were not constant,
of hybrids finally results in the re-appearance of the but could evolve (cited in Junker 2011). Nägeli was
original parental forms. Similarly to Mendel, Köl- an expert in the genus Hieracium, which seemed
reuter’s work has been neglected but not forgot- to be particularly suitable for studies on speciation.
ten. One reason for the relative neglect of his work It is a highly polymorphic genus consisting of many
is the unfortunate manner in which he published his distinct and easily recognizable species connected
results, as his publications consist of preliminary by a continuum of intermediate forms. In contrast
progress reports and different aspects of the same to others, Nägeli did not deny evolutionary role
cross (F , reciprocal F , F , backcrosses) were often of hybridization, especially in the early steps of spe-
1 1 2
reported in entirely different sections. ciation (van Dijk & Ellis 2016).
There were several scientists, who were Mendel’s Therefore, Mendel used different titles for his work
contemporaries, conducting hybridization experi- with pea (hybriden) and Hieracium (bastarden).
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Review Czech Journal of Genetics and Plant Breeding, 58, 2022 (3): 113–126
https://doi.org/10.17221/19/2022-CJGPB
Similarly, Bronn’s translation of Darwin’s Origin plantarum (1760), and rejected the previous view
(in Junker 1991), used the word ‘Bastard’ for hybrids of immutability. He stated that although hybridiza-
formed from different species (‘hybrid’ in Darwin) tion is widespread, the generation of hybrid lineage
and the word ‘Blendlinge’ for hybrids from different and speciation via hybridization is rare. Nowadays,
varieties of the same species (‘mongrel’ in Darwin), we know that new hybrid lineages must establish
which is identical to ‘Mischlinge’ by Focke (1881). reproductive isolation in order to overcome genetic
On the other hand, word ‘hybriden’ originates from assimilation and competition with parental species
Latin ‘hybrida’, used for ‘offspring of a tame sow and (Winge 1917; Müntzing 1930). As mentioned above,
a wild boar’, or more generally for ‘anything a product hybrid lineage may be formed through allopoly-
of two heterogeneous things’. ploid or homoploid speciation. Allopolyploid hybrid
speciation is more common (Soltis & Soltis 2009).
Occurrence of interspecific hybrids in nature A recent review found that 11% of species across
47 plant genera are of allopolyploid origin (Barker
Modern evolutionary studies of hybridization were et al. 2016). It has long been observed that crossing
laid down by three key discoveries. The first discovery two plant species or genotypes can create a hybrid
by Winge (1917, 1932) showed that genetically stable with faster growth, more biomass or greater reproduc-
and fertile hybrid species could be derived instanta- tive output than its parents (Jones 1917; East 1936).
neously by the duplication of a hybrid’s chromosome This phenomenon is called hybrid vigor or heterosis.
complement (i.e. allopolyploidy). This hypothesis Both Kölreuter (1766) and Darwin (1876) described
was soon confirmed experimentally (Dorsey 1936; the phenomenon of heterosis in their experimental
Sax 1936), and allopolyploidy is now recognized crosses of plants. It was Shull’s (1908, 1911) work
as a prominent mechanism of speciation in flowering on maize, that determined the genetic mechanism
plants and ferns (Stebbins 1959; Soltis & Soltis 2009). causing heterosis. Similar to heterosis, transgressive
A second important discovery resulted from the work segregation occurs when phenotypic trait values
of Müntzing (1930) on homoploid hybridization. He in hybrid populations fall outside the range of parental
postulated that the sorting of chromosomal rearrange- variation. Transgressive segregation is distinct from
ments in successive generations of hybrids could, heterosis because it is manifested predominantly
by chance, lead to the formation of new population in the F and following generations and may persist
2
systems that are homozygous for a unique combina- indefinitely once established (Rieseberg & Carney
tion of chromosomal sterility factors. Thus, the new 1998). Transgressive segregation demonstrates how
hybrid population would be fertile, stable, and at the hybridization can produce novel phenotypes, thus
same ploidy level as its parents, yet at least partially enable adaptation to new ecological niches and play
reproductively isolated from both parental species. a significant creative role in evolution. On the other
Müntzing (1936) also showed that polyploids were hand, where there is no fertility barrier between
generally more vigorous and hardier. The importance hybrids and parents, there is potential for gene flow.
of hybridization in plant speciation and evolution Hybrids may backcross with either or both of the
received opposing views of being either a creative parents, resulting in alien introgression forms.
evolutionary force or an evolutionary dead-end.
Natural hybridization can be defined as an inter- Interspecific hybridization and plant
breeding of individuals from two distinct populations, domestication
and individuals in those populations must be dis-
tinguishable on the basis of one or more heritable During the domestication process, several im-
characters. Both homoploid and allopolyploid hybrid portant crops have originated through interspecific
speciation involve the formation of novel genetic hybridization accompanied by the whole genome
combination, which may lead to the novel adapta- duplication (Smýkal et al. 2018; Purruganan 2019).
tions that allow persistence of the hybrid lineage, Molecular studies revealed the hybrid origin of a large
often in an environment distinct from that of either number of domesticated crops, including wheat
parent and is recognized as a common phenomenon (Marcussen et al. 2014), tall fescue (Humphreys
in plants (Soltis & Soltis 2009; Abbott et al. 2013). For et al. 1995), cotton (Wendel & Crown 2003), banana
the first time, Linnaeus admitted that new species (Heslop-Harrison & Schwarzacher 2007), oilseed
might arise by hybridization in Disquisitio de sexu rape (Lu et al. 2019), apple (Cornille et al. 2014), date
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