hybrid
Hybrid may refer to:
Science
* Hybrid (biology), an offspring resulting from cross-breeding
** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species
** Hybridity, the property of a hybrid plant which is a union of two diff ...
of
wheat
Wheat is a group of wild and crop domestication, domesticated Poaceae, grasses of the genus ''Triticum'' (). They are Agriculture, cultivated for their cereal grains, which are staple foods around the world. Well-known Taxonomy of wheat, whe ...
(''Triticum'') and
rye
Rye (''Secale cereale'') is a grass grown extensively as a grain, a cover crop and a forage crop. It is grown principally in an area from Eastern and Northern Europe into Russia. It is much more tolerant of cold weather and poor soil than o ...
(''Secale'') first
bred
A breed is a specific group of breedable domestic animals having homogeneous appearance (phenotype), homogeneous behavior, and/or other characteristics that distinguish it from other organisms of the same species. In literature, there exist seve ...
in laboratories during the late 19th century in
Scotland
Scotland is a Countries of the United Kingdom, country that is part of the United Kingdom. It contains nearly one-third of the United Kingdom's land area, consisting of the northern part of the island of Great Britain and more than 790 adjac ...
and
Germany
Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
. Commercially available triticale is almost always a second-generation hybrid, i.e., a cross between two kinds of primary (first-cross) triticales. As a rule, triticale combines the yield potential and
grain
A grain is a small, hard, dry fruit (caryopsis) – with or without an attached husk, hull layer – harvested for human or animal consumption. A grain crop is a grain-producing plant. The two main types of commercial grain crops are cereals and ...
quality of wheat with the disease and environmental tolerance (including soil conditions) of rye. Only in 1970 did the first commercial variety become available. Depending on the
cultivar
A cultivar is a kind of Horticulture, cultivated plant that people have selected for desired phenotypic trait, traits and which retains those traits when Plant propagation, propagated. Methods used to propagate cultivars include division, root a ...
, triticale can more or less resemble either of its parents. It is grown mostly for
forage
Forage is a plant material (mainly plant leaves and stems) eaten by grazing livestock. Historically, the term ''forage'' has meant only plants eaten by the animals directly as pasture, crop residue, or immature cereal crops, but it is also used m ...
or
fodder
Fodder (), also called provender (), is any agriculture, agricultural foodstuff used specifically to feed domesticated livestock, such as cattle, domestic rabbit, rabbits, sheep, horses, chickens and pigs. "Fodder" refers particularly to food ...
, although some triticale-based foods can be purchased at
health food store
A health food store (or health food shop) is a type of grocery store that primarily sells healthful foods, organic foods, local produce, and often nutritional supplements. Health food stores typically offer a wider or more specialized selectio ...
s and can be found in some
breakfast cereal
Breakfast cereal is a category of food, including food products, made from food processing, processed cereal, cereal grains, that are eaten as part of breakfast or as a snack food, primarily in Western societies.
Although warm, cooked cereals li ...
s.
When crossing wheat and rye, wheat is used as the female parent and rye as the male parent (pollen donor). The resulting hybrid is
sterile
Sterile or sterility may refer to:
*Asepsis, a state of being free from biological contaminants
* Sterile (archaeology), a sediment deposit which contains no evidence of human activity
*Sterilization (microbiology), any process that eliminates or ...
and must be treated with
colchicine
Colchicine is a medication used to prevent and treat gout, to treat familial Mediterranean fever and Behçet's disease, and to reduce the risk of myocardial infarction. The American College of Rheumatology recommends colchicine, nonstero ...
to induce
polyploidy
Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one fro ...
and thus the ability to reproduce itself.
The primary producers of triticale are
Poland
Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...
,
Germany
Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
,
Belarus
Belarus, officially the Republic of Belarus, is a landlocked country in Eastern Europe. It is bordered by Russia to the east and northeast, Ukraine to the south, Poland to the west, and Lithuania and Latvia to the northwest. Belarus spans an a ...
,
France
France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...
and
Russia
Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
. In 2014, according to the
Food and Agriculture Organization
The Food and Agriculture Organization of the United Nations; . (FAO) is a specialized agency of the United Nations that leads international efforts to defeat hunger and improve nutrition and food security. Its Latin motto, , translates ...
(FAO), 17.1 million tons were harvested in 37 countries across the world.
The triticale hybrids are all amphidiploid, which means the plant is
diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
for two
genomes
A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
derived from different
species
A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. In other words, triticale is an
allotetraploid
Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one fro ...
. In earlier years, most work was done on
octoploid
Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one fro ...
triticale. Different
ploidy
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
levels have been created and evaluated over time. The tetraploids showed little promise, but
hexaploid
Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one fro ...
triticale was successful enough to find commercial application.
The
CIMMYT
The International Maize and Wheat Improvement Center (known – even in English – by its Spanish acronym CIMMYT for ''Centro Internacional de Mejoramiento de MaÃz y Trigo'') is a non-profit research-for-development organization that develops ...
(International Maize and Wheat Improvement Center) triticale improvement program was intended to improve food production and nutrition in
developing countries
A developing country is a sovereign state with a less-developed Secondary sector of the economy, industrial base and a lower Human Development Index (HDI) relative to developed countries. However, this definition is not universally agreed upon. ...
. Triticale was thought to have potential in the production of bread and other food products, such as
cookie
A cookie is a sweet biscuit with high sugar and fat content. Cookie dough is softer than that used for other types of biscuit, and they are cooked longer at lower temperatures. The dough typically contains flour, sugar, egg, and some type of ...
s,
pasta
Pasta (, ; ) is a type of food typically made from an Leavening agent, unleavened dough of wheat flour mixed with water or Eggs as food, eggs, and formed into sheets or other shapes, then cooked by boiling or baking. Pasta was originally on ...
, pizza
dough
Dough is a malleable, sometimes elastic paste made from flour (which itself is made from grains or from leguminous or chestnut crops). Dough is typically made by mixing flour with a small amount of water or other liquid and sometimes includes ...
and breakfast cereals. The
protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
content is higher than that of wheat, although the
glutenin
Glutenin (a type of glutelin) is a major protein within wheat flour, making up 47% of the total protein content. The glutenins are protein aggregates of high- molecular-mass (HMW) and low-molecular-mass (LMW) subunits with molar masses from ab ...
fraction is less. The grain has also been stated to have higher levels of
lysine
Lysine (symbol Lys or K) is an α-amino acid that is a precursor to many proteins. Lysine contains an α-amino group (which is in the protonated form when the lysine is dissolved in water at physiological pH), an α-carboxylic acid group ( ...
than wheat. Acceptance would require the
milling
Milling may refer to:
* Milling (minting), forming narrow ridges around the edge of a coin
* Milling (grinding), breaking solid materials into smaller pieces by grinding, crushing, or cutting in a mill
* Milling (machining), a process of using ro ...
industry to adapt to triticale, as the milling techniques employed for wheat are unsuited to triticale.Sell, J.L.; Hodgson, G.C.; Shebeski, L.H. (1962 ''Triticale'' as a potential component of chick rations Canadian Journal of Animal Science, Volume 42, Number 2 Past research indicated that triticale could be used as a feed grain and, particularly, later research found that its
starch
Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diet ...
is readily digested.Bird, S. H; Rowe, J. B.; Choct, M.; Stachiw, S.; Tyler, P.; Thompson, R. D. (1999) In vitro fermentation of grain and enzymatic digestion of cereal starch Recent Advances in Animal Nutrition, Vol 12, pp. 53–61 As a feed grain, triticale is already well established and of high economic importance. It has received attention as a potential
energy crop
Energy crops are low-cost and low-maintenance crops grown solely for renewable bioenergy production (not for food). The crops are processed into solid, liquid or gaseous fuels, such as pellets, bioethanol or biogas. The fuels are burned to ...
, and research is currently being conducted on the use of the crop's
biomass
Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
in
bioethanol
Ethanol fuel is fuel containing ethyl alcohol, the same type of alcohol as found in alcoholic beverages. It is most often used as a motor fuel, mainly as a biofuel additive for gasoline.
Several common ethanol fuel mixtures are in use a ...
production. Triticale has also been used to produce
vodka
Vodka ( ; is a clear distilled beverage, distilled alcoholic beverage. Its varieties originated in Poland and Russia. Vodka is composed mainly of water and ethanol but sometimes with traces of impurities and flavourings. Traditionally, it is ...
.
History
In the 19th century, crossing cultivars or species became better understood, allowing the controlled hybridization of more plants and animals. In 1873, Alexander Wilson first managed to manually fertilize the female organs of wheat flowers with rye pollen (male gametes), but found that the resulting plants were sterile, much the way the offspring of a
horse
The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 mi ...
and
donkey
The donkey or ass is a domesticated equine. It derives from the African wild ass, ''Equus africanus'', and may be classified either as a subspecies thereof, ''Equus africanus asinus'', or as a separate species, ''Equus asinus''. It was domes ...
is an infertile
mule
The mule is a domestic equine hybrid between a donkey, and a horse. It is the offspring of a male donkey (a jack) and a female horse (a mare). The horse and the donkey are different species, with different numbers of chromosomes; of the two ...
. Fifteen years later in 1888, a partially-fertile hybrid was produced by , "Tritosecale Rimpaui Wittmack". Such hybrids germinate only when the chromosomes spontaneously
double
Double, The Double or Dubble may refer to:
Mathematics and computing
* Multiplication by 2
* Double precision, a floating-point representation of numbers that is typically 64 bits in length
* A double number of the form x+yj, where j^2=+1
* A ...
.
Unfortunately, "partially fertile" was all that was produced until 1937. In that year, it was discovered that the chemical
colchicine
Colchicine is a medication used to prevent and treat gout, to treat familial Mediterranean fever and Behçet's disease, and to reduce the risk of myocardial infarction. The American College of Rheumatology recommends colchicine, nonstero ...
, which is used both for general plant germination and as a treatment for
gout
Gout ( ) is a form of inflammatory arthritis characterized by recurrent attacks of pain in a red, tender, hot, and Joint effusion, swollen joint, caused by the deposition of needle-like crystals of uric acid known as monosodium urate crysta ...
, would force chromosome doubling by keeping them from pulling apart during cell division. Triticale had become viable, though at that point the cost of producing the seeds was disproportionate to the yield.
By the 1960s, triticale was being produced that was far more nutritious than normal wheat. However, it was a poorly-producing crop, sometimes yielding shriveled kernels, germinating poorly or prematurely, and did not bake well.
Modern triticale has overcome most of these problems, after decades of additional breeding and gene transfer with wheat and rye. Millions of acres/hectares of the crop are grown around the world, slowly increasing toward becoming a significant source of food-calories.
Species
Triticale hybrids are currently classified by ploidy into three
nothospecies #REDIRECT Hybrid name
In botanical nomenclature, a hybrid may be given a hybrid name, which is a special kind of botanical name, but there is no requirement that a hybrid name should be created for plants that are believed to be of hybrid origin. ...
tetraploid
Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one fro ...
hexaploid
Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one fro ...
triticale. Stable, currently very successful in agriculture. May be produced by ''Secale cereale'' × ''
Triticum turgidum
''Triticum turgidum'' (with its various subspecies being known as ''pasta wheat'', ''macaroni wheat'' and ''durum wheat'') is a species of wheat. It is an annual and grows primarily in temperate areas and is native to countries around the eastern ...
octaploid
Polyploidy is a condition in which the cells of an organism have more than two paired sets of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two complete sets of chromosomes, one from ...
triticale. Not completely stable, mainly historical importance. May be produced by ''Secale cereale'' × ''
Triticum aestivum
Common wheat (''Triticum aestivum''), also known as bread wheat, is a cultivated wheat species. About 95% of wheat produced worldwide is common wheat; it is the most widely grown of all crops and the cereal with the highest monetary yield.
Ta ...
'', genome AABBDDRR.
The current treatment follows the Mac Key 2005 treatment of ''Triticum'' using a broad species concept based on genome composition. Traditional classifications used a narrow species concept based on the treatment of wheats by Dorofeev ''et al.'', 1979, and hence produced many more species names. The genome notation follows , with the rye genome notated as R.
Biology and genetics
Earlier work with wheat-rye crosses was difficult due to low survival of the resulting hybrid
embryo
An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
and spontaneous chromosome doubling. These two factors were difficult to predict and control. To improve the viability of the embryo and thus avoid its abortion, ''
in vitro
''In vitro'' (meaning ''in glass'', or ''in the glass'') Research, studies are performed with Cell (biology), cells or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in ...
'' culture techniques were developed (Laibach, 1925).
Colchicine
Colchicine is a medication used to prevent and treat gout, to treat familial Mediterranean fever and Behçet's disease, and to reduce the risk of myocardial infarction. The American College of Rheumatology recommends colchicine, nonstero ...
was used as a chemical agent to double the chromosomes. After these developments, a new era of triticale
breeding
Breeding is sexual reproduction that produces offspring, usually animals or plants. It can only occur between a male and a female animal or plant.
Breeding may refer to:
* Animal husbandry, through selected specimens such as dogs, horses, and rab ...
was introduced. Earlier triticale hybrids had four reproductive disorders, namely
meiotic
Meiosis () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one copy ...
instability, high
aneuploid
Aneuploidy is the presence of an abnormal number of chromosomes in a cell, for example a human somatic cell having 45 or 47 chromosomes instead of the usual 46. It does not include a difference of one or more complete sets of chromosomes. A cel ...
frequency, low
fertility
Fertility in colloquial terms refers the ability to have offspring. In demographic contexts, fertility refers to the actual production of offspring, rather than the physical capability to reproduce, which is termed fecundity. The fertility rate ...
and shriveled
seed
In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
(Muntzing 1939; Krolow 1966). Cytogenetical studies were encouraged and well funded to overcome these problems.
It is especially difficult to see the expression of rye
genes
In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
in the background of wheat
cytoplasm
The cytoplasm describes all the material within a eukaryotic or prokaryotic cell, enclosed by the cell membrane, including the organelles and excluding the nucleus in eukaryotic cells. The material inside the nucleus of a eukaryotic cell a ...
and the predominant wheat nuclear
genome
A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
. This makes it difficult to realise the potential of rye in disease resistance and ecological adaptation.
Triticale is essentially a self-fertilizing, or naturally
inbred
Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders an ...
crop. This mode of reproduction results in a more
homozygous
Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism.
Mos ...
genome. The crop is, however, adapted to this form of reproduction from an evolutionary point of view. Cross-fertilization is also possible, but it is not the primary form of reproduction.
' is a
stem rust
Stem rust, also known as cereal rust, black rust, red rust or red dust, is caused by the fungus ''Puccinia graminis'', which causes significant disease in cereal crops. Crop species that are affected by the disease include bread wheat, durum wh ...
resistance gene which is commonly found in triticale. Originally from
rye
Rye (''Secale cereale'') is a grass grown extensively as a grain, a cover crop and a forage crop. It is grown principally in an area from Eastern and Northern Europe into Russia. It is much more tolerant of cold weather and poor soil than o ...
(Imperial rye), now () widely found in triticale. Located on the 3A chromosome arm, originally from 3R. Virulence has been observed in field by ''Puccinia graminis'' f. sp. ''secalis'' (''Pgs'') and in an artificial cross ''Pgs'' ''Puccinia graminis'' f. sp. ''tritici'' (''Pgt''). When successful, ''Sr27'' is among the few ''Sr''s that does not even allow the underdeveloped