Editing Triticale

{{Short description|Hybrid wheat/rye crop}}
{{Automatic taxobox
| image = Triticale.jpg
| image_caption =
| display_parents = 3
| taxon = × Triticosecale
| authority = Wittm. ex A. Camus.
| synonyms = × ''Triticale'' <small>[[Erich von Tschermak|Tscherm.-Seys.]] ex Müntzing</small>
| subdivision_ranks = Species
| subdivision = See text
}}

'''Triticale''' ({{IPAc-en|t|r|ɪ|t|ɪ|ˈ|k|eɪ|l|iː}}; × ''Triticosecale'') is a [[Hybrid (biology)|hybrid]] of [[wheat]] (''Triticum'') and [[rye]] (''Secale'') first [[plant breeding|bred]] in laboratories during the late 19th century in [[Scotland]] and [[Germany]].<ref>{{citation |author=Stace, C. A. |author-link=Stace, C. A.|year=1987 |title=Triticale: A Case of Nomenclatural Mistreatment |journal=[[Taxon (journal)|Taxon]] |volume=36 |issue=2 |pages=445–452 |jstor=1221447 |doi=10.2307/1221447}}</ref> 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]] quality of wheat with the disease and environmental tolerance (including soil conditions) of rye. Only in 1970 did the first commercial variety become available.<ref>{{cite web |url=https://www.glnc.org.au/resource/triticale/ |title= Triticale |publisher= Grains and Legumes Nutrition Council |access-date=2025-05-22}}</ref> Depending on the [[cultivar]], triticale can more or less resemble either of its parents. It is grown mostly for [[forage]] or [[fodder]], although some triticale-based foods can be purchased at [[health food store]]s and can be found in some [[breakfast cereal]]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 [[infertility|sterile]] and must be treated with [[colchicine]] to induce [[polyploidy]] and thus the ability to reproduce itself.

The primary producers of triticale are [[Poland]], [[Germany]], [[Belarus]], [[France]] and [[Russia]]. In 2014, according to the [[Food and Agriculture Organization]] (FAO), 17.1 million tons were harvested in 37 countries across the world.<ref name="FAOSTATCrops">{{cite web |url=http://faostat.fao.org/site/567/default.aspx |title=Food and Agricultural commodities production |publisher=FAO Statistics Division |access-date=2016-04-05 |archive-date=2012-04-07 |archive-url=https://web.archive.org/web/20120407180211/http://faostat.fao.org/site/567/default.aspx |url-status=dead }}</ref>

The triticale hybrids are all [[amphidiploid]], which means the plant is [[diploid]] for two [[genomes]] derived from different [[species]]. In other words, triticale is an [[allotetraploid]]. In earlier years, most work was done on [[octoploid]] triticale. Different [[ploidy]] levels have been created and evaluated over time. The tetraploids showed little promise, but [[hexaploid]] triticale was successful enough to find commercial application.<ref name="FAO_TriticaleImprovement">{{cite book |url=https://www.fao.org/4/y5553e/y5553e00.htm |title=Triticale improvement and production |first1=Mohamed |last1=Mergoum |first2=Helena |last2=Gómez-Macpherson |publisher= [[FAO]] |year=2004 |isbn=92-5-105182-8 |access-date=2024-10-03 |archive-url=https://web.archive.org/web/20241004005845/https://www.fao.org/4/y5553e/y5553e00.htm |archive-date=2024-10-04 |url-status=live }}</ref>

The [[International Maize and Wheat Improvement Center|CIMMYT]] (International Maize and Wheat Improvement Center) triticale improvement program was intended to improve food production and nutrition in [[developing countries]]. Triticale was thought to have potential in the production of bread and other food products, such as [[cookie]]s, [[pasta]], pizza [[dough]] and breakfast cereals.<ref name="FAO_TriticaleImprovement"/> The [[protein]] 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]] than wheat.<ref name="Can">{{cite web |url=https://www.thecanadianencyclopedia.ca/en/article/triticale |title=Triticale |work=Agriculture |publisher=The [[Canadian Encyclopedia]] |first=E. N. |last=Larter |access-date=2009-06-19 |archive-date=2017-09-02 |archive-url=https://web.archive.org/web/20170902095456/http://www.thecanadianencyclopedia.com/en/article/triticale/ |url-status=live }}</ref> Acceptance would require the [[Gristmill|milling]] industry to adapt to triticale, as the milling techniques employed for wheat are unsuited to triticale.<ref name="sell-et-al">Sell, J.L.; Hodgson, G.C.; Shebeski, L.H. (1962) [http://pubservices.nrc-cnrc.ca/rp-ps/absres.jsp?jcode=cjas&ftl=cjas62-026&lang=eng ''Triticale'' as a potential component of chick rations] {{Webarchive|url=https://archive.today/20130115123650/http://pubservices.nrc-cnrc.ca/rp-ps/absres.jsp?jcode=cjas&ftl=cjas62-026&lang=eng |date=2013-01-15 }} [[Canadian Journal of Animal Science]], Volume 42, Number 2</ref> Past research indicated that triticale could be used as a feed grain and, particularly, later research found that its [[starch]] is readily digested.<ref name="bird-et-al">Bird, S. H; Rowe, J. B.; Choct, M.; Stachiw, S.; Tyler, P.; Thompson, R. D. (1999) [[hdl:2123/2615|In vitro fermentation of grain and enzymatic digestion of cereal starch]] Recent Advances in Animal Nutrition, Vol 12, pp. 53–61</ref> As a feed grain, triticale is already well established and of high economic importance. It has received attention as a potential [[energy crop]], and research is currently being conducted on the use of the crop's [[biomass]] in [[bioethanol]] production. Triticale has also been used to produce [[vodka]].<ref>{{cite web |last1=France |first1=Marie |title=This distillery produces gin and vodka from locally grown bio-dynamic triticale |url=https://www.spiritshunters.com/choice/this-distillery-produces-gin-and-vodka-from-locally-grown-bio-dynamic-triticale/ |website=www.spiritshunters.com/ |date=6 May 2019 |access-date=31 January 2023}}</ref><ref>{{cite web |last1=Sharman |first1=Linda |title=WA distillery in High Spirits |url=https://www.farmweekly.com.au/story/5980720/wa-distillery-in-high-spirits/ |website=Farm Weekly Australia |date=April 2019 |access-date=31 January 2023}}</ref>

== History ==
{{More citations needed section|date=June 2021}}
[[File:Wheat, rye, triticale montage.jpg|280px|right|thumb|[[Wheat]], [[rye]], triticale|alt=The smaller grain of [[wheat]] on the left, larger kernels of [[rye]] next, and triticale on the right — triticale [[grain]] is significantly larger than wheat.]]

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<ref>{{cite web|url=http://triticale.org/triticale-history/ |title=Triticale history|publisher=International Triticale Association, Ghent University, Belgium|date=2023 }}</ref> with rye pollen (male gametes), but found that the resulting plants were sterile, much the way the offspring of a [[horse]] and [[donkey]] is an infertile [[mule]]. Fifteen years later in 1888, a partially-fertile hybrid was produced by {{ill|Wilhelm Rimpau|de|Wilhelm Rimpau (Agrarwissenschaftler)|vertical-align=sup}}, "Tritosecale Rimpaui Wittmack". Such hybrids germinate only when the chromosomes spontaneously [[doubled haploidy|double]].

Unfortunately, "partially fertile" was all that was produced until 1937. In that year, it was discovered that the chemical [[colchicine]], which is used both for general plant germination and as a treatment for [[gout]], would force chromosome doubling by keeping them from pulling apart during cell division.<ref>[https://www.carnivorousplants.org/grow/propagation/ColchicineToxicity Colchicine Treatment and Toxicity | ICPS]</ref> 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]]:<ref name=taxo-Hammer>{{cite journal |last1=Hammer |first1=Karl |last2=Filatenko |first2=Anna A. |last3=Pistrick |first3=Klaus |title=Taxonomic remarks on ''Triticum'' L. and ''×Triticosecale'' Wittm. |journal=Genetic Resources and Crop Evolution |date=January 2011 |volume=58 |issue=1 |pages=3–10 |doi=10.1007/s10722-010-9590-4|s2cid=24403249}}</ref>
* [[× Triticosecale semisecale|× ''Triticosecale semisecale'']] <small>(Mackey) K.Hammer & Filat.</small> – [[tetraploid]] triticale. Unstable, but used in [[bridging population|breeding bridging]]. Includes the following crosses:
** ''[[Triticum monococcum]]'' × ''[[Secale cereale]]'', genome AARR;
** Alternative crosses, genome ABRR ([[mixogenome]] A/B).
* [[× Triticosecale neoblaringhemii|× ''Triticosecale neoblaringhemii'']] <small>A.Camus</small> – [[hexaploid]] triticale. Stable, currently very successful in agriculture. May be produced by ''Secale cereale'' × ''[[Triticum turgidum]]'', genome AABBRR.
* [[× Triticosecale rimpaui|× ''Triticosecale rimpaui'']] <small>Wittm.</small> – [[octaploid]] triticale. Not completely stable, mainly historical importance. May be produced by ''Secale cereale'' × ''[[Triticum aestivum]]'', 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 {{section link|Taxonomy of wheat|Genome}}, with the rye genome notated as R.<ref name=taxo-Hammer/>

== Biology and genetics ==
{{More citations needed section|date=June 2021}}
Earlier work with wheat-rye crosses was difficult due to low survival of the resulting hybrid [[embryo]] 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]]'' culture techniques were developed (Laibach, 1925).{{full citation needed|date=March 2017}} [[Colchicine]] was used as a chemical agent to double the chromosomes.<ref>{{cite journal|last1=Blakeslee|first1=Albert F.|last2=Avery|first2=Amos G.|title=Methods of Inducing Doubling of Chromosomes in Plants|journal=Journal of Heredity|date=December 1937|volume=28|issue=12|pages=393–411|doi=10.1093/oxfordjournals.jhered.a104294}}</ref> After these developments, a new era of triticale [[Artificial selection|breeding]] was introduced. Earlier triticale hybrids had four reproductive disorders, namely [[meiotic]] instability, high [[aneuploid]] frequency, low [[fertility]] and shriveled [[seed]] (Muntzing 1939; Krolow 1966).{{full citation needed|date=March 2017}} Cytogenetical studies were encouraged and well funded to overcome these problems.

It is especially difficult to see the expression of rye [[genes]] in the background of wheat [[cytoplasm]] and the predominant wheat nuclear [[genome]]. This makes it difficult to realise the potential of rye in disease resistance and ecological adaptation.{{cn|date=September 2023}}

Triticale is essentially a self-fertilizing, or naturally [[inbred]] crop. This mode of reproduction results in a more [[homozygous]] 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.{{cn|date=September 2023}}

''{{ Visible anchor | Sr27 }}'' is a [[stem rust]] resistance gene which is commonly found in triticale.<ref name="Sr27-GlobalRust">{{cite web | title=''Sr27'' | website=Borlaug Global Rust Initiative | url=http://globalrust.org/gene/sr27 | access-date=2021-07-24}}</ref> Originally from [[rye]]<ref name="Singh-et-al-2011">{{cite journal | last1=Singh | first1=Ravi P. | last2=Hodson | first2=David P. | last3=Huerta-Espino | first3=Julio | last4=Jin | first4=Yue | last5=Bhavani | first5=Sridhar | last6=Njau | first6=Peter | last7=Herrera-Foessel | first7=Sybil | last8=Singh | first8=Pawan K. | last9=Singh | first9=Sukhwinder | last10=Govindan | first10=Velu|display-authors=3 | title=The Emergence of Ug99 Races of the Stem Rust Fungus is a Threat to World Wheat Production | journal=Annual Review of Phytopathology| volume=49 | issue=1 | date=2011-09-08 | issn=0066-4286 | doi=10.1146/annurev-phyto-072910-095423 | pages=465–481| pmid=21568701 }}</ref> (Imperial rye),<ref name="Park-Wellings-2012">{{cite journal | last1=Park | first1=Robert F. | last2=Wellings | first2=Colin R. | title=Somatic Hybridization in the Uredinales | journal=Annual Review of Phytopathology | volume=50 | issue=1 | date=2012-09-08 | issn=0066-4286 | doi=10.1146/annurev-phyto-072910-095405 | pages=219–239| pmid=22920559 }}</ref> now ({{as of|2021|lc=yes}}) widely found in triticale.<ref name="Upadhyaya-et-al-2021">{{cite journal|last1=Upadhyaya|first1=Narayana M.|last2=Mago|first2=Rohit|last3=Panwar|first3=Vinay|last4=Hewitt|first4=Tim|last5=Luo|first5=Ming|last6=Chen|first6=Jian|last7=Sperschneider|first7=Jana|last8=Nguyen-Phuc|first8=Hoa|last9=Wang|first9=Aihua|last10=Ortiz|first10=Diana|last11=Hac|first11=Luch|last12=Bhatt|first12=Dhara|last13=Li|first13=Feng|last14=Zhang|first14=Jianping|last15=Ayliffe|first15=Michael|last16=Figueroa|first16=Melania|last17=Kanyuka|first17=Kostya|last18=Ellis|first18=Jeffrey G.|last19=Dodds|first19=Peter N.|display-authors=3 |title=Genomics accelerated isolation of a new stem rust avirulence gene–wheat resistance gene pair|journal=Nature Plants|year=2021|volume=7|issue=9|pages=1220–1228|issn=2055-0278|doi=10.1038/s41477-021-00971-5|pmid=34294906|s2cid=236199741|url=https://repository.rothamsted.ac.uk/download/911a82b1b12af18ce55d120f48f3568494a06549fc42318ed814d03d057c0e37/4800766/10378_1_supp_129049_q96vf9_convrt.pdf}}</ref> Located on the 3A chromosome arm,<ref name="Sr27-GlobalRust" /> originally from 3R.<ref name="McIntosh-et-al-1995">{{cite book | last1=McIntosh | first1=RA | last2=Wellings | first2=CR | last3=Park | first3=RF | title=Wheat Rusts - An Atlas of Resistance Genes | publisher=[[Springer Publishing|Springer]] | isbn=9789401040419 | date=1995}}</ref> Virulence has been observed in field by [[Stem rust|''Puccinia graminis'' f. sp. ''secalis'']] (''Pgs'') and in an artificial cross ''Pgs'' {{times}} [[Stem rust|''Puccinia graminis'' f. sp. ''tritici'']] (''Pgt'').<ref name="Park-Wellings-2012" /> When successful, ''Sr27'' is among the few ''Sr''s that does not even allow the underdeveloped [[uredinia]] and slight degree of sporulation commonly allowed by most ''Sr''s.<ref name="Singh-et-al-2011" /> Instead there are [[necrotic]] or [[chlorotic]] flecks.<ref name="Roelfs-1988">{{cite journal | last=Roelfs | first=A P | title=Genetic Control of Phenotypes in Wheat Stem Rust | journal=[[Annual Review of Phytopathology]] | publisher=[[Annual Reviews (publisher)|Annual Reviews]] | volume=26 | issue=1 | year=1988 | issn=0066-4286 | doi=10.1146/annurev.py.26.090188.002031 | pages=351–367}}</ref> Deployment in triticale in [[New South Wales]] and [[Queensland]], Australia, however, rapidly showed virulence between 1982 and 1984 – the first virulence on this gene in the world.<ref name="McIntosh-Brown-1997">{{cite journal | last1=McIntosh | first1=R. A. | last2=Brown | first2=G. N. | title=Anticipatory Breeding for Resistance to Rust Diseases in Wheat | journal=[[Annual Review of Phytopathology]] | publisher=[[Annual Reviews (publisher)|Annual Reviews]] | volume=35 | issue=1 | year=1997 | issn=0066-4286 | doi=10.1146/annurev.phyto.35.1.311 | pages=311–326| pmid=15012526 }}</ref><ref name="Singh-et-al-2011" /><ref name="McIntosh-et-al-1995" /> (This was especially associated with the cultivar Coorong.)<ref name="McIntosh-Brown-1997" /><ref name="Johnson-1984">{{cite journal | last=Johnson | first=R | title=A Critical Analysis of Durable Resistance | journal=Annual Review of Phytopathology| volume=22 | issue=1 | year=1984 | issn=0066-4286 | doi=10.1146/annurev.py.22.090184.001521 | pages=309–330}}</ref> Therefore, the International Maize and Wheat Improvement Center's triticale offerings were tested and many were found to depend solely on ''Sr27''.<ref name="Johnson-1984" /><ref name="McIntosh-et-al-1995" /> Four years later, in 1988 virulence was found in [[South Africa]]. ''Sr27'' has become less common in CIMMYT triticales since the mid-'80s.<ref name="McIntosh-et-al-1995" />

== Conventional breeding approaches ==
{{Agricultural production box
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|year=2022<ref name="FAOSTAT2022">{{cite web |title=FAOSTAT |url=https://www.fao.org/faostat/en/#data/QCL/visualize |website=www.fao.org |access-date=5 March 2024}}</ref>
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The aim of a triticale breeding programme is mainly focused on the improvement of quantitative [[trait (biology)|trait]]s, such as grain yield, nutritional quality and plant height, as well as traits which are more difficult to improve, such as earlier maturity and improved test weight (a measure of bulk density). These traits are controlled by more than one [[gene]].<ref name="Manual">
{{cite web
|url=http://www1.agric.gov.ab.ca/$department/deptdocs.nsf/all/fcd10535
|title=Triticale Production Manual
|publisher= Alberta Agriculture and Food and Development, Government of Alberta, Agricultural and Rural Development
|access-date=2009-06-23}}</ref> Problems arise, however, because such [[polygenic]] traits involve the integration of several physiological processes in their expression. Thus the lack of single-gene control (or simple inheritance) results in low trait heritability (Zumelzú ''et al.'' 1998).

Since the induction of the International Maize and Wheat Improvement Center triticale breeding programme in 1964, the improvement in realised grain yield has been remarkable. In 1968, at [[Ciudad Obregón]], Sonora, in northwest Mexico, the highest yielding triticale line produced 2.4 t/ha. Today, CIMMYT has released high yielding spring triticale lines (e.g. Pollmer-2) which have surpassed the 10 t/ha yield barrier under optimum production conditions.<ref name="Hede">{{cite web |url=http://repository.cimmyt.org/xmlui/bitstream/handle/10883/632/74311.pdf?sequence=1 |publisher=Corporate Communications, International Maize and Wheat Improvement Center |title=A New Approach to Triticale Improvement |work=Research Highlights of the CIMMYT Wheat Program 1999–2000 |location=Mexico, D. F. |year=2001 |author=A. R. Hede |pages=21–26 |access-date=18 July 2013 |archive-date=4 March 2016 |archive-url=https://web.archive.org/web/20160304071638/http://repository.cimmyt.org/xmlui/bitstream/handle/10883/632/74311.pdf?sequence=1 |url-status=dead }}</ref>

Based on the commercial success of other hybrid crops, the use of hybrid triticales as a strategy for enhancing yield in favourable, as well as marginal, environments has proven successful over time. Earlier research conducted by CIMMYT made use of a chemical hybridising agent to evaluate [[heterosis]] in [[hexaploid]] triticale hybrids. To select the most promising parents for hybrid production, test crosses conducted in various environments are required, because the variance of their specific combining ability under differing environmental conditions is the most important component in evaluating their potential as parents to produce promising hybrids. The prediction of general combining ability of any triticale plant from the performance of its parents is only moderate with respect to grain yield. Commercially exploitable yield advantages of hybrid triticale [[cultivars]] is dependent on improving parent [[heterosis]] and on advances in [[inbred]]-line development.{{Citation needed|date=June 2021}}

Triticale is useful as an animal feed [[grain]]. However, it is necessary to improve its milling and bread-making quality aspects to increase its potential for human consumption. The relationship between the constituent wheat and rye [[genomes]] were noted to produce meiotic irregularities, and genome instability and incompatibility presented numerous problems when attempts were made to improve triticale. This led to two alternative methods to study and improve its reproductive performance, namely, the improvement of the number of grains per floral spikelet and its meiotic behaviour. The number of grains per spikelet has an associated low [[heritability]] value (de Zumelzú et al. 1998). In improving yield, indirect selection (the selection of correlated/related traits other than that to be improved) is not necessarily as effective as direct selection. (Gallais 1984)<ref name="Gallais">{{cite conference | first=A. | last=Gallais | title=Use of Indirect Selection in Plant Breeding |editor=Hogenboon, N. G. |display-editors=etal | book-title=Efficiency In Plant Breeding, Proc. 10th Congress Eucarpia | pages=45–60 | date=19–24 June 1983 | location=Pudoc, Wageningen, The Netherlands}}</ref>

Lodging (the toppling over of the plant stem, especially under windy conditions) resistance is a [[polygenic]]ally inherited (expression is controlled by many genes) trait, and has thus been an important breeding aim in the past.<ref name="Tikhnenko">{{cite journal | first=N. D. | last=Tikhnenko |author2=Tsvetkova, N. V. |author3=Voylokov, A. V. | title=The Effect of Parental Genotypes of Rye Lines on the Development of Quantitative Traits in Primary Octoploid Triticale: Plant Height | journal=[[Russian Journal of Genetics]] | volume=31 | issue=1 | pages=52–56 | date=23 August 2002 | doi=10.1023/A:1022070810919| s2cid=2659383 }}</ref> The use of dwarfing genes, known as ''Rht'' genes, which have been incorporated from both ''Triticum'' and ''Secale'', has resulted in a decrease of up to {{ Convert | 20 | cm }} in plant height without causing any adverse effects.{{Citation needed|date=June 2021}}

A 2013 study found that hybrids have better yield stability under [[yield stress]] than do [[inbred line]]s.<ref name = "Stability" >{{ Cite journal
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==Application of newer techniques==
Abundant information exists concerning [[R-gene]]s (for disease resistance) in wheat, and a continuously updated on-line catalogue, the Catalogue of Gene Symbols, of these genes can be found at [http://wheat.pw.usda.gov/ggpages/wgc/98/] {{Webarchive|url=https://web.archive.org/web/20060923172834/http://wheat.pw.usda.gov/ggpages/wgc/98/ |date=2006-09-23 }}. Another online database of [[cereal]] rust resistance genes is available at [https://web.archive.org/web/20061006193218/http://www.cdl.umn.edu/res_gene/res_gene.html]. Unfortunately, less is known about rye and particularly triticale R-genes. Many R-genes have been transferred to wheat from its wild relatives, and appear in such papers and catalogues, thus making them available for triticale breeding. The two mentioned databases are significant contributors to improving the genetic variability of the triticale [[gene pool]] through gene (or more specifically, allele) provision. Genetic variability is essential for progress in breeding. In addition, genetic variability can also be achieved by producing new primary triticales, which essentially means the reconstitution of triticale, and the development of various hybrids involving triticale, such as triticale-rye hybrids. In this way, some [[chromosomes]] from the R genome have been replaced by some from the D genome. The resulting so-called substitution and translocation triticale facilitates the transfer of R-genes.{{Citation needed|date=June 2021}}

===Introgression===
[[Introgression]] involves the crossing of closely related plant relatives, and results in the [[gene linkage|transfer of 'blocks' of genes]], i.e. larger segments of chromosomes compared to single genes.{{ Dubious | date=May 2023 | reason=This MAY occur during introgression.}} [[R-gene]]s are generally introduced within such blocks, which are usually incorporated/translocated/introgressed into the distal (extreme) regions of [[chromosomes]] of the crop being introgressed. Genes located in the proximal areas of chromosomes may be completely linked (very closely spaced), thus preventing or severely hampering [[genetic recombination|recombination]], which is necessary to incorporate such blocks.{{ Dubious | date=May 2023 | reason=Very strange.}}<ref name="Chelkowski">{{cite journal | url=http://jag.igr.poznan.pl/2004-Volume-45/3/abstracts/210.html | first=Jerzy | last=Chelkowski |author2=Tyrka, Miroslaw | title=Enhancing the resistance of triticale by using genes from wheat and rye | journal=Journal of Applied Genetics| volume=45 | issue=3 | pages=283–295 | date=15 October 2003 | pmid=15306719}}</ref> Molecular markers (small lengths of [[DNA]] of a characterized/known sequence) are used to 'tag' and thus track such translocations.<ref name="Lee">{{cite journal | first=T. G. | last=Lee |author2=Hong, M. J. |author3=Johnson, J. W. |author4=Bland, D. E. |author5=Kim, D. Y. |author6= Seo, Y. W. | title=Development and functional assessment of EST-derived 2RL-specific markers for 2BS.2RL translocations | journal=Theoretical and Applied Genetics| volume=119 | issue=4 | pages=663–673 | year=2009 | doi =10.1007/s00122-009-1077-3|pmid = 19543880| s2cid=22773590}}</ref> A weak [[colchicine]] solution has been employed to increase the probability of recombination in the proximal chromosome regions, and thus the introduction of the translocation to that region. The resultant translocation of smaller blocks that indeed carry the R-gene(s) of interest has decreased the probability of introducing unwanted genes.<ref name="Lukaszewski">{{cite journal | url=http://crop.scijournals.org/cgi/content/abstract/30/5/1151 | first=Adam | last=Lukaszewski | title=Frequency of 1RS.1AL and 1RS.1BL Translocations in United States Wheats | journal=[[Crop Science (journal)|Crop Science]]| volume=30 | issue=5 | pages=1151–1153 | year=1990 | doi=10.2135/cropsci1990.0011183X003000050041x | url-access=subscription }}{{Dead link|date=February 2020 |bot=InternetArchiveBot |fix-attempted=yes }}</ref>

The ''{{ Visible anchor |Sr59}}'' [[crop disease resistance gene|resistance gene]] was introgressed into wheat from the [[2R chromosome]] of [[rye]].<ref name="Sc-to-Ta" /> However this was actually done ''through'' triticale.<ref name="Sc-to-Ta" /> Triticale has been the [[amphiploid]] for several such rye{{NBSP}}⇨{{NBSP}}wheat introgressions.<ref name="Sc-to-Ta">
 {{ Cite journal
 | last3=Åhman | first3=Inger
 | last2=Gustavsson | first2=Larisa
 | last1=Herrera | first1=Leonardo
 | year=2017
 | title=A systematic review of rye (''Secale cereale'' L.) as a source of resistance to pathogens and pests in wheat (''Triticum aestivum'' L.)
 | volume=154
 | issue=1
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 | publisher=[[BioMed Central]]
 | issn=1601-5223
 | journal=[[Hereditas]]
 | doi=10.1186/s41065-017-0033-5
 | pmid=28559761
 | pmc=5445327
| doi-access=free
 }}</ref>

A 2014 study found that ''{{Vanchor|Ddw1}}'' [[dwarfing]] gene from the [[rye 5R chromosome]] also provides [[Fusarium head blight]] (FHB) resistance in this host.<ref name="Height">{{cite book|year=2019|publisher=[[Springer Nature]]|first3=Dennis|pages=363+420|first2=Shri|first1=Jameel|last1=Al-Khayri|last2=Jain|last3=Johnson|isbn=978-3-030-23108-8|title=Advances in Plant Breeding Strategies: Cereals}}</ref><ref>{{cite journal|issue=7|year=2014|first4=Thomas|first3=Bernd|first2=Hans|first1=Rasha|last4=Miedaner|last3=Hackauf|last2=Maurer|last1=Kalih|pages=1527–1536|volume=127|journal=Theoretical and Applied Genetics |issn=0040-5752|s2cid=14913718|pmid=24852306|doi=10.1007/s00122-014-2316-9|title=Effect of a rye dwarfing gene on plant height, heading stage, and Fusarium head blight in triticale (×''Triticosecale'' Wittmack)}}</ref>

===Production of doubled haploids===

[[Doubled haploidy |Doubled haploid]] (DH) plants have the potential to save much time in the development of [[inbred]] lines. This is achieved in a single generation, as opposed to many, which would otherwise occupy much physical space/facilities. DHs also express deleterious recessive [[alleles]] otherwise masked by dominance effects in a genome containing more than one copy of each chromosome (and thus more than one copy of each gene). Various techniques exist to create DHs. The ''in vitro'' culture of [[anthers]] and [[microspore]]s is most often used in [[cereals]], including triticale.<ref name="Bernard">{{cite journal |year=2009 |issue=6 |first4=Kadambot |first3=Ian |first2=Heather |first1=Maya |last4=Siddique |last3=Small |last2=Clarke |last1=Kumari |pages=393–409 |volume=28 |s2cid=85298264 |doi=10.1080/07352680903133252 |journal=Critical Reviews in Plant Sciences |title=Albinism in Plants: A Major Bottleneck in Wide Hybridization, Androgenesis and Doubled Haploid Culture |bibcode=2009CRvPS..28..393K}}</ref><ref>{{cite journal |year=1984 |first1=S. |last1=Bernard |last2=Charmet |first2=G. |title=Diallel analysis of androgenetic plant production in hexaploid Triticale (''X. triticosecale'', Wittmack) |journal=Theoretical and Applied Genetics |volume=69 |issue=1 |pages=55–61 |doi=10.1007/BF00262539 |pmid=24253624 |s2cid=37533199 |doi-access=free}}</ref><ref name="González">{{cite journal |url=http://www.fao.org/agris/search/display.do?f=./2000/v2609/HU2000000254.xml;HU2000000254 |first1=J. M. |last1=González |last2=Jouve |first2=N. |title=Improvement of Anther Culture Media for Haploid Production in Triticale |journal=Cereal Research Communications |volume=28 |issue=1–2 |pages=65–72 |year=2000 |doi=10.1007/BF03543575 |url-access=subscription }}{{Dead link |date=December 2021 |bot=InternetArchiveBot |fix-attempted=yes }}</ref><ref name="González1997">{{cite journal |first1=J. M. |last1=González |last2=Jouve |first2=N. |last3=Hernádez |first3=I. |title=Analysis of anther culture response in hexaploid triticale |journal=Plant Breeding |volume=116 |issue=3 |pages=301–304 |date=January 18, 1997 |doi=10.1111/j.1439-0523.1997.tb01003.x |doi-access=free }}</ref> These two techniques are referred to as androgenesis, which refers to the development of [[pollen]]. Many plant species and [[cultivars]] within species, including triticale, are recalcitrant in that the success rate of achieving whole newly generated (diploid) plants is very low. Genotype by culture medium interaction is responsible for varying success rates, as is a high degree of microspore abortion during culturing.<ref name= "Development-Androgenesis" >{{ Cite book |year=2009 |edition=1st |volume=3 |location=New York |publisher=Springer-Verlag |pages=|first=Marcelo |last=Carena |editor-first1=Marcelo J. |editor-last1=Carena |doi=10.1007/978-0-387-72297-9 |isbn=978-0-387-72294-8 |title=Cereals |series=Handbook of Plant Breeding |oclc=405546128 }}</ref><ref>{{ Cite journal |last1=González |first1=J. M. |last2=Jouve |first2=N. |year=2005 |title=Microspore development during in vitro androgenesis in triticale |volume=49 |issue=1 |pages=23–28 |journal=Biologia Plantarum |s2cid=35661525 |doi=10.1007/s10535-005-3028-4 |doi-access=free }}</ref><ref name="Stresses-Re-Programming">{{cite journal |last1=Shariatpanahi |first1=Mehran E. |last2=Bal |first2=Ugur |last3=Heberle-Bors |first3=Erwin |last4=Touraev |first4=Alisher |title=Stresses applied for the re-programming of plant microspores towards in vitro embryogenesis |journal=Physiologia Plantarum |publisher=John Wiley & Sons |volume=127 |issue=4 |year=2006 |doi=10.1111/j.1399-3054.2006.00675.x |pages=519–534 |s2cid=84771294 |doi-access=free}}</ref><ref>{{cite journal |last1=Tuvesson |first1=S. |last2=Ljungberg |first2=A. |last3=Johansson |first3=N. |last4=Karlsson |first4=K.-E. |last5=Suijs |first5=L. W. |last6=Josset |first6=J.-P. |title=Large-scale production of wheat and triticale double haploids through the use of a single-anther culture method |journal=Plant Breeding |publisher=Wiley-VCH GmbH |volume=119 |issue=6 |year=2000 |doi=10.1046/j.1439-0523.2000.00536.x |pages=455–459 |s2cid=84991814}}
</ref> The response of parental triticale lines to [[anther]] culture is known to be correlated to the response of their progeny.<ref name="González1997"/><ref name="Konzak">{{cite journal |url=http://rparticle.web-p.cisti.nrc.ca/rparticle/AbstractTemplateServlet?calyLang=eng&journal=gen&volume=35&year=1992&issue=6&msno=g92-146 |first1=Calvin |last1=Konzak |last2=Zhou |first2=Huaping |title=Genetic control of green plant regeneration from anther culture of wheat |journal=[[Genome (journal) |Genome]] |volume=35 |issue=6 |pages=957–961 |date=1 December 1992 |doi=10.1139/g92-146 |url-access=subscription }}{{Dead link |date=February 2020 |bot=InternetArchiveBot |fix-attempted=yes }}</ref><ref name="Anderson">{{cite journal |first1=S. B. |last1=Andersen |last2=Tuvesson |first2=I. K. D. |last3=Pedersen |first3=S. |title=Nuclear genes affecting albinism in wheat (''Triticum aestivum'' L.) anther culture |journal=Theoretical and Applied Genetics |volume=78 |issue=6 / December, 1989 |pages=879–889 |date=9 August 1989 |doi=10.1007/BF00266675 |pmid=24226023 |s2cid=9686842 }}</ref> Chromosome elimination is another method of producing DHs, and involves [[Hybrid (biology) |hybridisation]] of wheat with [[maize]] (''Zea mays'' L.), followed by [[auxin]] treatment and the artificial rescue of the resultant haploid embryos before they naturally abort. This technique is applied rather extensively to wheat.<ref name="Bennett2">{{cite conference |first1=M. D. |last1=Bennett |last2=Laurie |first2=D. A. |last3=O'Donoughue |first3=L. S. |title=Wheat x maize and other wide sexual hybrids: their potential for crop improvement and genetic manipulations |book-title=Gene Manipulation in Plant Improvement II: Proceedings of the 19th Stadler Genetics Symposium |pages=95–126 |publisher=[[Plenum Press]] |date=13–15 March 1989 |location=New York}}</ref> Its success is in large part due to the insensitivity of maize pollen to the crossability inhibitor genes known as Kr1 and Kr2 that are expressed in the floral style of many wheat cultivars.<ref name="Bennett">{{cite journal |first1=M. D. |last1=Bennett |last2=Laurie |first2=D. A. |title=The effect of the crossability loci ''Kr1'' and ''Kr2'' on fertilization frequency in hexaploid wheat x maize crosses |journal=[[Theoretical and Applied Genetics]] |volume=73 |issue=3 / January, 1987
 |pages=403–409 |date=31 August 1986 |doi=10.1007/BF00262508 |pmid=24241002 |s2cid=7530770 }}</ref> The technique is unfortunately less successful in triticale.<ref name="Marcinska">{{cite journal |url=http://www3.interscience.wiley.com/journal/119118108/abstract |archive-url=https://archive.today/20121015184323/http://www3.interscience.wiley.com/journal/119118108/abstract |url-status=dead |archive-date=15 October 2012 |first1=I. |last1=Marcińska |last2=Wodzony |first2=M. |last3=Ponitka |first3=A. |last4=Ślusarkiewicz-Jarzina |first4=A. |last5=Woźna |first5=J. |title=Production of doubled haploids in triticale (×Triticosecale Wittm.) by means of crosses with maize (Zea mays L.) using picloram and dicamba |journal=Plant Breeding |volume=117 |issue=3 |pages=211–215 |date=26 January 1998 |doi=10.1111/j.1439-0523.1998.tb01928.x|url-access=subscription }}</ref> However, ''[[Imperata cylindrica]]'' (a grass) was found to be just as effective as [[maize]] with respect to the production of DHs in both [[wheat]] and triticale.<ref name="Chaudhary">{{cite journal |url=http://www3.interscience.wiley.com/journal/118695340/abstract |archive-url=https://archive.today/20130105072243/http://www3.interscience.wiley.com/journal/118695340/abstract |url-status=dead |archive-date=January 5, 2013 |first1=H. K. |last1=Chaudhary |last2=Pratap |first2=A. |last3=Sethi |first3=G. S. |title=Relative efficiency of different Gramineae genera for haploid induction in triticale and triticale x wheat hybrids through the chromosome elimination technique |journal=Plant Breeding |volume=124 |issue=2 / April, 2005 |pages=147–153 |date=August 16, 2004 |doi=10.1111/j.1439-0523.2004.01059.x|url-access=subscription }}</ref>

===Application of molecular markers===
An important advantage of [[biotechnology]] applied to plant breeding is the speeding up of cultivar release that would otherwise take 8–12 years. It is the process of [[Selection (biology)|selection]] that is actually enhanced, i.e., retaining that which is desirable or promising and ridding that which is not. This carries with it the aim of changing the genetic structure of the plant population. The website [https://web.archive.org/web/20071018151244/http://maswheat.ucdavis.edu/protocols/protocols.htm] is a valuable resource for [[marker assisted selection]] (MAS) protocols relating to R-genes in wheat. MAS is a form of indirect [[Selection (biology)|selection]]. The Catalogue of Gene Symbols mentioned earlier is an additional source of [[molecular]] and morphological markers. Again, triticale has not been well characterised with respect to molecular markers, although an abundance of rye molecular markers makes it possible to track rye chromosomes and segments thereof within a triticale background.{{Citation needed|date=June 2021}}

Yield improvements of up to 20% have been achieved in hybrid triticale cultivars due to [[heterosis]].<!-- Góral 2002; --><ref name="Góral">{{cite journal | url=http://www.ihar.edu.pl/biblioteka/plant_breeding_and_seed_science.php | first=H. | last=Góral | title=Heterosis and Combining Ability in Spring Triticale (x ''Triticosecale'', Wittm.) | journal=Plant Breeding and Seed Science| volume=43 | pages=25–34 | year=1999 | display-authors=etal | access-date=2009-06-22 | archive-date=2009-04-23 | archive-url=https://web.archive.org/web/20090423081215/http://www.ihar.edu.pl/biblioteka/plant_breeding_and_seed_science.php | url-status=dead }}</ref><ref name="Becker">{{cite journal | url=http://www3.interscience.wiley.com/journal/118988062/abstract?CRETRY=1&SRETRY=0 | archive-url=https://archive.today/20130105075608/http://www3.interscience.wiley.com/journal/118988062/abstract?CRETRY=1&SRETRY=0 | url-status=dead | archive-date=January 5, 2013|first1=H. C.|last1=Becker|last2=Oettler|first2=G.|last3=Hoppe|first3=G.|title=Heterosis for yield and other agronomic traits of winter triticale F<sub>1</sub> and F<sub>2</sub> hybrids | journal=Plant Breeding| volume=120 | issue=4 | pages=351–353 | date= March 8, 2001 | doi=10.1046/j.1439-0523.2001.00624.x| url-access=subscription}}</ref><ref name="Burger">{{cite journal | url=http://www3.interscience.wiley.com/journal/118879174/abstract | archive-url=https://archive.today/20121015184111/http://www3.interscience.wiley.com/journal/118879174/abstract | url-status=dead | archive-date=2012-10-15 | first=H. | last=Burger |author2=Oettler, G. |author3=Melchinger, A. E. | title=Heterosis and combining ability for grain yield and other agronomic traits in winter triticale | journal=Plant Breeding| volume=122 | issue=4 | pages=318–321 | date= August 2003 | doi=10.1046/j.1439-0523.2003.00877.x| url-access=subscription }}</ref> This raises the question of what inbred lines should be crossed (to produce hybrids) with each other as parents to maximize yield in their hybrid progeny. This is termed the 'combining ability' of the parental lines. The identification of good combining ability at an early stage in the breeding programme can reduce the costs associated with 'carrying' a large number of plants (literally thousands) through it, and thus forms part of efficient selection. Combining ability is assessed by taking into consideration all available information on [[Common descent|descent]] ([[Genetics|genetic]] relatedness), [[Morphology (biology)|morphology]], qualitative (simply inherited) traits and [[biochemical]] and molecular markers. Exceptionally little information exists on the use of molecular markers to predict heterosis in triticale.<ref name="Góral2005">{{cite journal | url=http://jag.igr.poznan.pl/2005-Volume-46/2/abstracts/259.html | first=Halina | last=Góral |author2=Tyrka, Miroslaw |author3=Spiss, Ludwik | title=Assessing genetic variation to predict the breeding value of winter triticale cultivars and lines | journal=Journal of Applied Genetics | volume=46 | issue=2 |pages=125–131 | date=2 March 2005 | pmid=15876679}}</ref> Molecular markers are generally accepted as better predictors than morphological markers (of [[agronomy|agronomic]] traits) due to their insensitivity to variation in environmental conditions.{{Citation needed|date=June 2021}}

A useful molecular marker known as a [[Short tandem repeat|simple sequence repeat]] (SSR) is used in breeding with respect to selection. SSRs are segments of a genome composed of [[Variable number tandem repeat|tandem repeats]] of a short sequence of [[nucleotides]], usually two to six [[base pair]]s. They are popular tools in genetics and breeding because of their relative abundance compared to other marker types, a high degree of polymorphism (number of variants), and easy assaying by polymerase chain reaction. However, they are expensive to identify and develop. Comparative genome mapping has revealed a high degree of similarity in terms of sequence colinearity between closely related crop species. This allows the exchange of such markers within a group of related species, such as wheat, rye and triticale. One study established a 58% and 39% transferability rate to triticale from wheat and rye, respectively.<ref name="Baenziger">{{cite journal | first=P. S. | last=Baenziger |author2=Kuleung, C. |author3=Dweikat I. | title=Transferability of SSR markers among wheat, rye, and triticale | journal=Theoretical and Applied Genetics| volume=108 | issue=6 / April, 2004 | pages=1147–1150 | date=5 December 2003 | doi=10.1007/s00122-003-1532-5 | pmid=15067402| s2cid=20537650 }}</ref> Transferability refers to the phenomenon where the sequence of DNA nucleotides flanking the SSR locus (position on the [[chromosome]]) is sufficiently homologous (similar) between genomes of closely related species. Thus, DNA primers (generally, a short sequence of nucleotides used to direct the copying reaction during PCR) designed for one species can be used to detect SSRs in related species. SSR markers are available in wheat and rye, but very few, if any, are available for triticale.<ref name="Baenziger"/>

===Genetic transformation===
The [[genetic transformation]] of crops involves the incorporation of 'foreign' genes or, rather, very small DNA fragments compared to introgression discussed earlier. Amongst other uses, transformation is a useful tool to introduce new traits or characteristics into the transformed crop. Two methods are commonly employed: infectious [[bacteria]]l-mediated (usually ''[[Agrobacterium]]'') transfer and [[biolistics]], with the latter being most commonly applied to [[allopolyploid]] cereals such as triticale. ''Agrobacterium''-mediated transformation, however, holds several advantages, such as a low level of DNA rearrangement in the [[transgenic]] plant, a low number of introduced copies of the transforming DNA, stable integration of an a-priori characterized T-DNA fragment (containing the DNA expressing the trait of interest) and an expected higher level of [[transgene]] expression. Triticale has, until recently, only been transformed via biolistics, with a 3.3% success rate.<ref name="Zimny">{{cite journal | first=J. | last=Zimny |author2=Becker, D. |author3=Brettschneider, R. |author4=Lörz, H. | title=Fertile, transgenic Triticale (''×Triticosecale'' Wittmack) | journal=Molecular Breeding | volume=1 | issue=2 | pages=155–164 | date=10 October 1994 | doi=10.1007/BF01249700| s2cid=27470553 }}</ref> Little has been documented on ''Agrobacterium''-mediated transformation of wheat: while no data existed with respect to triticale until 2005, the success rate in later work was nevertheless low.<ref name="Binka">{{cite journal | first=A. | last=Binka |author2=Nadolska-Orczyk, A. |author3=Przetakiewicz, A. |author4=Kopera, K. |author5=Orczyk, W. | title=Efficient Method of Agrobacterium-mediated Transformation for Triticale (x Triticosecale Wittmack) | journal=Journal of Plant Growth Regulation| volume=24 | issue=1 / March, 2005 | pages=2–10 | date=28 July 2005 | doi=10.1007/s00344-004-0046-y| s2cid=22996555 }}</ref>

==Research==
{{More citations needed section|date=June 2021}}
Triticale holds much promise as a commercial crop, as it has the potential to address specific problems within the cereal industry. Research is currently being conducted worldwide in places like [[Stellenbosch University]] in [[South Africa]].

Conventional plant breeding has helped establish triticale as a valuable crop, especially where conditions are less favourable for wheat cultivation. Triticale being a synthesized [[grain]] notwithstanding, many initial limitations, such as an inability to reproduce due to infertility and seed shrivelling, low yield and poor nutritional value, have been largely eliminated.

[[Tissue culture]] techniques with respect to wheat and triticale have seen continuous improvements, but the isolation and culturing of individual microspores seems to hold the most promise. Many molecular markers can be applied to marker-assisted gene transfer, but the expression of R-genes in the new genetic background of triticale remains to be investigated.<ref name="Baenziger"/> More than 750 wheat microsatellite primer pairs are available in public wheat breeding programmes, and could be exploited in the development of SSRs in triticale.<ref name="Baenziger"/> Another type of molecular marker, single nucleotide polymorphism (SNP), is likely to have a significant impact on the future of triticale breeding.

==Health concerns==
{{Main|Gluten-related disorders}}
Like both its hybrid parents – wheat and rye – triticale contains [[gluten]] and is therefore unsuitable for people with gluten-related disorders, such as [[celiac disease]], [[non-celiac gluten sensitivity]] and [[wheat allergy]], among others.<ref name=TovoliMasi>{{cite journal | vauthors=Tovoli F, Masi C, Guidetti E, Negrini G, Paterini P, Bolondi L| title=Clinical and diagnostic aspects of gluten related disorders| journal=[[World Journal of Clinical Cases]]| volume=3| issue=3| pages=275–84| date=Mar 16, 2015| pmid=25789300|pmc= 4360499| doi=10.12998/wjcc.v3.i3.275| doi-access=free}}</ref>

==In fiction==
An episode of the popular TV series ''[[Star Trek]]'', "[[The Trouble with Tribbles]]", revolved around the protection of a grain developed from triticale. This grain was named "quadro-triticale" by writer [[David Gerrold]] at the suggestion of producer [[Gene Coon]], with four distinct lobes per kernel. In that episode [[Mr. Spock]] correctly attributes the ancestry of the nonfictional grain to 20th-century Canada.<ref name="Trouble-Tribbles-Vanity-Fair">{{cite magazine | last=Vinciguerra | first=Thomas | title=Star Trek: Inside "The Trouble with Tribbles," 50 Years Later | magazine=[[Vanity Fair (magazine)|Vanity Fair]] | date=2017-12-29 | url=http://www.vanityfair.com/hollywood/2017/12/star-trek-trouble-with-tribbles-50th-anniversary | access-date=2021-07-20}}</ref>

Indeed, in 1953 the [[University of Manitoba]] began the first North American triticale breeding program. Early breeding efforts concentrated on developing a high-yield, drought-tolerant human food crop species suitable for marginal wheat-producing areas.<ref>{{cite web|title=Triticale|url=http://www1.agric.gov.ab.ca/$department/deptdocs.nsf/all/agdex127|publisher=Government of Alberta, Agriculture and Rural Development|access-date=11 September 2011}}</ref> (Later in the episode, [[Pavel Chekhov|Chekov]] claims that the fictional quadro-triticale was a "Russian invention".<ref name="Gerrold-1973">{{cite book | last=Gerrold | first=David | author-link=David Gerrold | title=The Trouble With Tribbles - the birth, sale, and final production of one episode | publisher=[[Ballantine Books]] | publication-place=New York | year=1973 | isbn=978-0-345-27671-1 | oclc=9265346 | pages=1–293}} p.{{spaces}}202</ref>)

A later episode titled "[[More Tribbles, More Troubles]]", in [[Star Trek: The Animated Series|the animated series]], also written by Gerrold, dealt with "quinto-triticale", an improvement on the original, having apparently five lobes per kernel.<ref name="Gerrold-STcom">{{cite web|url=https://intl.startrek.com/article/david-gerrold-recalls-more-tribbles-and-bem|title=David Gerrold Recalls "More Tribbles" and "Bem"|date=March 10, 2011|publisher=Star Trek.com|accessdate=March 30, 2013}}</ref>

Three decades later the spinoff series ''[[Star Trek: Deep Space Nine]]'' revisited quadro-triticale and the depredations of the Tribbles in the episode "[[Trials and Tribble-ations]]".<ref name="Trials-and-Tribble-ations-STcom">{{cite web | title=20 Years Later... "Trials and Tribble-ations" | website=Star Trek.com | date=2016-11-04 | url=http://www.startrek.com/article/20-years-later-trials-and-tribble-ations | access-date=2021-07-20}}</ref>

==References==
{{Reflist}}

{{Cereals}}
{{Taxonbar|from1=Q380329|from2=Q17484014}}
{{Authority control}}

[[Category:Cereals]]
[[Category:Pooideae]]
[[Category:Energy crops]]
[[Category:Plant common names]]
<!-- [[Category:Plant nothogenera]] moved to "x Triticosecale" redirect -->