× Triticosecale
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Triticale (; × ''Triticosecale'') is a
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 dif ...
of
wheat Wheat is a grass widely cultivated for its seed, a cereal grain that is a worldwide staple food. The many species of wheat together make up the genus ''Triticum'' ; the most widely grown is common wheat (''T. aestivum''). The archaeologi ...
(''Triticum'') and
rye Rye (''Secale cereale'') is a grass grown extensively as a grain, a cover crop and a forage crop. It is a member of the wheat tribe (Triticeae) and is closely related to both wheat (''Triticum'') and barley (genus ''Hordeum''). Rye grain is u ...
(''Secale'') first bred in laboratories during the late 19th century in
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to the ...
and
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. 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 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 legum ...
quality of wheat with the disease and environmental tolerance (including soil conditions) of rye. Only recently has it been developed into a commercially viable crop. Depending on the
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...
, 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 g ...
, 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 health foods, organic foods, local produce, and often nutritional supplements. Health food stores typically offer a wider or more specialized selection of fo ...
s and can be found in some
breakfast cereal Cereal, formally termed breakfast cereal (and further categorized as cold cereal or warm cereal), is a traditional breakfast food made from processed cereal grains. It is traditionally eaten as part of breakfast, or a snack food, primarily in ...
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 to induce polyploidy 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 is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populous ...
,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
,
Belarus Belarus,, , ; alternatively and formerly known as Byelorussia (from Russian ). officially the Republic of Belarus,; rus, Республика Беларусь, Respublika Belarus. is a landlocked country in Eastern Europe. It is bordered by R ...
,
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
and
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
. In 2014, according to the
Food and Agriculture Organization The Food and Agriculture Organization of the United Nations (FAO)french: link=no, Organisation des Nations unies pour l'alimentation et l'agriculture; it, Organizzazione delle Nazioni Unite per l'Alimentazione e l'Agricoltura is an intern ...
(FAO), 17.1 million tons were harvested in 37 countries across the world. The triticale hybrids are all
amphidiploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
, 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. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
for two
genomes In the fields of molecular biology and genetics, 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 gen ...
derived from different
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
. In other words, triticale is an allotetraploid. In earlier years, most work was done on
octoploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
triticale. Different
ploidy Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Sets of chromosomes refer to the number of mat ...
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 one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
triticale was successful enough to find commercial application. The
International Maize and Wheat Improvement Center The International Maize and Wheat Improvement Center (known - even in English - by its Spanish acronym CIMMYT for ''Centro Internacional de Mejoramiento de Maíz y Trigo'') is a non-profit research-for-development organization that develops im ...
(CIMMYT) triticale improvement program was intended to improve food production and nutrition in
developing countries A developing country is a sovereign state with a lesser developed industrial base and a lower Human Development Index (HDI) relative to other countries. However, this definition is not universally agreed upon. There is also no clear agreem ...
. Triticale was thought to have potential in the production of bread and other food products, such as
cookie A cookie is a baked or cooked snack or dessert that is typically small, flat and sweet. It usually contains flour, sugar, egg, and some type of oil, fat, or butter. It may include other ingredients such as raisins, oats, chocolate chips, n ...
s,
pasta Pasta (, ; ) is a type of food typically made from an unleavened dough of wheat flour mixed with water or eggs, and formed into sheets or other shapes, then cooked by boiling or baking. Rice flour, or legumes such as beans or lentils, are som ...
, pizza
dough Dough is a thick, malleable, sometimes elastic paste 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 yeast or other leavening ag ...
and breakfast cereals. The
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
content is higher than that of wheat, although the glutenin 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. It contains an α-amino group (which is in the protonated form under biological conditions), an α-carboxylic acid group (which is in the deprotonated −C ...
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 rota ...
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 diets ...
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 energy production by combustion (not for food). The crops are processed into solid, liquid or gaseous fuels, such as pellets, bioethanol or biogas. The fuels are burned to g ...
, and research is currently being conducted on the use of the crop's
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bi ...
in bioethanol production.


History

In the 19th century, crossing cultivars or species became better understood, allowing the controlled hybridization of more plants and animals. Semen could be taken from a male of one animal species to fertilize a female from another species that wouldn't mate with the male directly, and crops plants such as wheat or rye could more systematically be hybridized with each other. 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 million y ...
and
donkey The domestic donkey is a hoofed mammal in the family Equidae, the same family as the horse. It derives from the African wild ass, ''Equus africanus'', and may be classified either as a subspecies thereof, ''Equus africanus asinus'', or as a ...
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 pos ...
. Fifteen years later in 1888, a partially-fertile hybrid was produced by , "Tritosecale Rimpaui Wittmack". Such hybrids only germinate when the chromosomes spontaneously
double A double is a look-alike or doppelgänger; one person or being that resembles another. Double, The Double or Dubble may also refer to: Film and television * Double (filmmaking), someone who substitutes for the credited actor of a character * Th ...
. Unfortunately, "partially fertile" was all that was produced until 1937, when it was discovered that the chemical colchicine, which is used both for general plant germination and on humans with
gout Gout ( ) is a form of inflammatory arthritis characterized by recurrent attacks of a red, tender, hot and swollen joint, caused by deposition of monosodium urate monohydrate crystals. Pain typically comes on rapidly, reaching maximal intensit ...
, would force chromosome doubling by keeping them from pulling apart during cell division. Triticale then was 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 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. ...
: * × ''Triticosecale semisecale'' (Mackey) K.Hammer & Filat. – tetraploid triticale. Unstable, but used in breeding bridging. Includes the following crosses: ** ''
Triticum monococcum Einkorn wheat (from German ''Einkorn'', literally "single grain") can refer either to a wild species of wheat (''Triticum'') or to its domesticated form. The wild form is '' T. boeoticum'' (syn. ''T. m.'' ssp. ''boeoticum''), the domesticated ...
'' × ''
Secale cereale Rye (''Secale cereale'') is a grass grown extensively as a grain, a cover crop and a forage crop. It is a member of the wheat tribe (Triticeae) and is closely related to both wheat (''Triticum'') and barley (genus ''Hordeum''). Rye grain is u ...
'', genome AARR; ** Alternative crosses, genome ABRR (mixogenome A/B). * × ''Triticosecale neoblaringhemii'' A.Camus – hexaploid triticale. Stable, currently very successful in argiculture. May be produced by ''Secale cereale'' × '' Triticum turgidum'', genome AABBRR. * × ''Triticosecale rimpaui'' Wittm. – octaploid 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 b 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 an initial stage of development of 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 spe ...
and spontaneous
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
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'') studies are performed with microorganisms, cells, or biological molecules outside their normal biological context. Colloquially called "test-tube experiments", these studies in biology an ...
'' culture techniques were developed (Laibach, 1925). Colchicine 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 (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately res ...
instability, high aneuploid frequency, low
fertility Fertility is the capability to produce offspring through reproduction following the onset of sexual maturity. The fertility rate is the average number of children born by a female during her lifetime and is quantified demographically. Fertili ...
and shriveled
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
(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 (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
in the background of wheat
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
and the predominant wheat nuclear
genome In the fields of molecular biology and genetics, 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 ge ...
. This makes it difficult to realise the potential of rye in disease resistance and ecological adaptation. One of the ways to relieve this problem was to produce secalotricum, in which rye cytoplasm was used instead of that from wheat. Triticale is essentially a self-fertilizing, or naturally inbred, 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. Mo ...
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. ''Sr27'' is a stem rust 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 a member of the wheat tribe (Triticeae) and is closely related to both wheat (''Triticum'') and barley (genus ''Hordeum''). Rye grain is u ...
(
Imperial Rye Imperial is that which relates to an empire, emperor, or imperialism. Imperial or The Imperial may also refer to: Places United States * Imperial, California * Imperial, Missouri * Imperial, Nebraska * Imperial, Pennsylvania * Imperial, Texas ...
), 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
uredinia {{Short pages monitor