Editing Triticale (section)

===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>