References

vestib

Известия Национальной академии наук Беларуси. Серия биологических наук

Proceedings of the National Academy of Sciences of Belarus, Biological Series

1029-89402524-230X

The Republican Unitary Enterprise Publishing House "Belaruskaya Navuka"

vestib-218

Research Article

ОБЗОРЫ

REVIEWS

ПЛЕЙОТРОПНЫЕ ЭФФЕКТЫ ГЕНОВ УСТОЙЧИВОСТИ МЯГКОЙ ПШЕНИЦЫ (TRITICUM AESTIVUM L.) К БИОТРОФНЫМ ГРИБНЫМ ПАТОГЕНАМ

PLEIOTROPIC EFFECTS OF RESISTANCE GENES OF COMMON WHEAT (TRITICUM AESTIVUM L.) TO BIOTROPHIC FUNGAL PATHOGENS

Волуевич

Е. А.

Voluevich

E. A.

Voluevitch@yandex.ru

Институт генетики и цитологии НАН БеларусиInstitute of Genetics and Cytology of the National Academy of Sciences of Belarus

2016

01082016

02115125

2016

Волуевич Е.А.

Voluevich E.A.

Данная работа распространяется под лицензией Creative Commons Attribution 4.0.

This work is licensed under a Creative Commons Attribution 4.0 License.

https://vestibio.belnauka.by/jour/article/view/218

Биотрофные грибные патогены пшеницы характеризуются высокими адаптационными способностями и вредоносностью. Для успешной селекции на устойчивость необходимы эффективные доноры. Особый интерес представляют те из них, которые обладают генами устойчивости к широкому спектру болезней. В статье рассматриваются эпидемиологические особенности биотрофных грибных патогенов, их вредоносность, гены с плейотрофным эффектом на устойчивость пшеницы к болезням, приводится информация о наличии молекулярных маркеров к этим генам.

Biotrophic fungal pathogens of wheat have high adaptive capacity and harmfulness. Successful breeding for resistance requires effective donors. Of particular interest are those genes which have resistance to a broad spectrum of diseases. The review describes epidemiological features of biotrophic fungal pathogens, their harmfulness, genes with pleiotropic effect on wheat resistance to disease, molecular markers to these genes

пшеницапатогеныустойчивостьплейотропные эффекты геновмолекулярные маркеры

wheatpathogensresistancepleiotropic effects of genesmolecular markers

References1

Dangl, J. L. Plant pathogens and integrated defence responses to infection / J. L. Dangl, J. D. Jones // Nature. – 2001. – Vol. 411. – P. 826–833.

Jones, J. D. G. The plant immune system / J. D. G. Jones, J. L. Dangl // Nature. – 2006. – Vol. 444. – P. 323–329.

Лукьяненко, П. П. Селекция устойчивых к ржавчине сортов / П. П. Лукьяненко // Селекция и семеноводство. – 1968. – № 4. – С. 10–18.

Дмитриев, А. П. Исследование внутрипопуляционных процессов у Puccinia recondita Rob. ex Desm f. sp. tritici Erikss. и генофонда устойчивости пшениц Закавказья к бурой ржавчине: автореф. дис. ... канд. биол. наук: 03.00.15 / А. П. Дмитриев; ВНИИР. – Л., 1975. – 26 с.

Джонстон, Ч. О. Листовая ржавчина / Ч. О. Джонстон // Пшеница и ее улучшение / редкол.: М. М. Якубцинер [и др.]. – М., 1970. – С. 347–355.

Селекция яровой мягкой пшеницы к местной популяции и к вирулентной расе Ug99 стеблевой ржавчины в условиях Западной Сибири / В. П. Шаманин [и др.] // Информ. вестн. ВОГиС. – 2010. – Т. 14, № 1. – С. 223–231.

Mains, E. B. Effect of leaf rust (Puccinia triticina Erikss.) on yield of wheat / E. B. Mains // J. Agr. Res. – 1930. – Vol. 40, N 5. – P. 417–446.

Caldwell, R. M. Effect of leaf rust (Puccinia triticina) on yield, physical characters, and composition of winter wheat / R. M. Caldwell // J. Agr. Res. – 1934. – Vol. 48. – P. 1049–1971. 122

Relation of leaf rust infection to yield, growth, and water economy of two varieties/ C. O. Johnston [et al.] // J. Agr. Res. – 1934. – Vol. 49, N 11. – P. 955–981.

Степанов, К. М. Ржавчина зерновых культур / К. М. Степанов. – Л.: Колос, 1975. – 75 с.

Берлянд-Кожевников, В. М. Селекция пшеницы на устойчивость к основным грибным заболеваниям / В. М. Берлянд-Кожевников, М. А. Федин. – М.: ВНИИТЭИСХ, ВАСХНИЛ, 1977. – 57 с.

Мочалова, Л. Вредоносность бурой ржавчины пшеницы / Л. Мочалова // Сб. науч. тр. Мирон. НИИСХ. – 1978. – Вып. 2. – С. 136–141.

Воронкова, А. А. Генетико-иммунологические основы селекции пшеницы на устойчивость к ржавчине / А. А. Воронкова. – М.: Колос, 1980. – 191 с.

Genetic improvement of wheat resistance to dangerous races of stem rust using conventional and molecular techniques trough international cooperation / A. Kokhmetova [et al.] // 8th international wheat conf.: abstracts of oral and poster presentations, St. Peterburg, 1–4 June 2010. – St. Peterburg, 2010. – P. 274–275.

Strategies to reduce the emerging wheat stripe rust disease // Synthesis of a dialog between policy makers and scientists from 31 countries at International Wheat Stripe Rust Symposium, ICARDA, Aleppo, Syria. – 2011. – 20 p.

Murray, G. M. The relationship between wheat yield and stripe rust as affected by length of epidemic and temperature at the grain development stage of crop growth / G. M. Murray // Plant Pathol. – 1994. – Vol. 43. – Iss. 2. – P. 397–405.

Leath, S. Effects of powdery mildew, triadimenol seed treatment, and triadimefon foliar sprays on yield of winter wheat in North Carolina / S. Leath, K. L. Bowen // Phytopathology. – 1989. – Vol. 79, N 2. – P. 152–155.

Griffey, C. A. Effectiveness of adult-plant resistance in reducing grain yield loss to powdery mildew in winter wheat / C. A. Griffey, M. K. Das, E. L. Stromberg // Plant Dis. – 1993. – Vol. 77. – P. 618–622.

Рачински, Т. Съвременната селекция и борбата с ръждите и бряшнестата мана на пшеница / Т. Рачински // Раст. защита. – 1980. – Т. 28, № 8. – С. 5–10.

Johnson, R. The concept of durable resistance / R. Johnson // Phytopathology. – 1979. – Vol. 69. – P. 198–199.

Hau, B. Wind-dispersed diseases / B. Hau, C. de Vallavieille-Pope // The Epidemiology of Plant Diseases; ed.: B. M. Cooke, Jones Gareth, B. Kaye. – 2nd ed. – Heidelberg: Springer-Verlag, 2006. – Ch. 15. – P. 387–416.

Roelfs, A. P. Wheat and rye stem rust / A. P. Roelfs // The Cereal Rusts. – Vol. 2: Diseases, Distribution, Epidemiology, and Control / ed.: A. P. Roelfs, W. R. Bushnell. – Orlando: Acad. Press, 1985a. – P. 1–37.

Roelfs, A. P. Epidemiology in North America / A. P. Roelfs // The Cereal Rusts. – Vol. 2: Diseases, Distribution, Epidemiology, and Control / ed.: A. P. Roelfs, W. R. Bushnell. – Orlando: Acad. Press, 1985b. – P. 403–434.

Limpert, E. Barley mildew in Europe: Evidence of wind-dispersal of the pathogen and its implications for improved use of host resistance and of fungicides for mildew control / E. Limpert // Integrated Control of Cereal Mildews: Monitoring the Pathogen; ed.: M. S. Wolfe, E. Limpert. – Dortrecht: Martinus Nijhof Publishers, 1987. – Р. 31–33.

Line, R. F. Stripe rust of wheat and barley in North America: a retrospective historical rev. / R. F. Line // Ann. Rev. of Phytopathol. – 2002. – Vol. 40. – P. 75–118.

Hovmøller, M. S. Clonality and long-distance migration of Puccinia striiformis f. sp. tritici in north-west Europe / M. S. Hovmøller, A. F. Justesen, J. K. M. Brown // Plant Pathol. – 2002. – Vol. 51. – P. 24–32.

Assessment of epidemiological parameters and their use in epidemiological and forecasting models of cereal airborne diseases / C. de Vallavieille-Pope [et al.] // Agronomie. – 2000. – Vol. 20, N 7. – P. 715–727.

McDonald, B. A. Pathogen population genetics, evolutionary potential, and durable resistance / B. A. McDonald, C. Linde // Ann. Rev. Phytopathol. – 2002. – Vol. 40. – P. 349–379.

Будевич, Г. В. Вредоносность стеблевой ржавчины озимой пшеницы / Г. В. Будевич, Е. А. Волуевич // Ботаника. – Минск, 1979. – Вып. XXI. – С. 171–174.

Волуевич, Е. А. Распространение бурой ржавчины в различных районах БССР / Е. А. Волуевич // Вес. АН БССР. Сер. бiял. навук. – 1982. – № 5. – С. 112–113.

Коптик, И. К. Селекция озимой пшеницы (Triticum aestivum L.) в условиях Беларуси: дис. … д-ра с.-х. наук: 06.01.05 / И. К. Коптик. – Жодино, 1996. – 304 с.

Catalogue of gene symbols for wheat. 2013 [Electronic resource] / R. A. McIntosh [et al.]. – Mode of access: http:// www.shigen.nig.ac.jp/wheat/komugi/genes/download.jsp. – Date of access: 01.02.2016.

Ohm, H. W. Three components of slow leaf-rusting at different growth stages in wheat / H. W. Ohm, G. E. Shaner // Phytopathology. – 1976. – Vol. 66. – P. 1356–1360.

Wilcoxson, R. D. Genetics of slow rusting in cereals / R. D. Wilcoxson // Phytopathology. – 1981. – Vol. 71. – P. 898–992.

Associations and genetics of three components of slow rusting in leaf rust of wheat / M. K. Das [et al.] // Euphytica. – 1993. – Vol. 68. – P. 99–109.

Dyck, P. L. The association of a gene for leaf rust resistance with the chromosome 7D suppressor of stem rust resistance in common wheat / P. L. Dyck // Genome. – 1987. – Vol. 29. – P. 467–469.

McIntosh, R. A. Close genetic linkage of genes conferring adult-plant resistance to leaf rust and stripe rust in wheat / R. A. McIntosh // Plant Pathol. – 1992. – Vol. 41. – P. 523–527.

Singh, R. P. Genetic association of leaf rust resistance gene Lr34 with adult-plant resistance to stripe rust in bread wheat / R. P. Singh // Phytopathology. – 1992a. – Vol. 82. – P. 835–838.

Powdery mildew resistance and Lr34/Yr18 genes for durable resistance to leaf and stripe rust co-segregate at a locus on the short arm of chromosome 7D of wheat / W. Spielmeyer [et al.] // Theor. Appl. Genet. – 2005. – Vol. 111. – P. 731–735. 123

The adult plant rust resistance loci Lr34/Yr18 and Lr46/Yr29 are important determinants of partial resistance to powdery mildew in bread wheat line Saar / M. Lillemo [et al.] // Theor. Appl. Genet. – 2008. – Vol. 116. – P. 1155–1166.

A putative ABC transporter confers durable resistance to multiple fungal pathogens in wheat / S. G. Krattinger [et al.] // Science. – 2009. – Vol. 323. – P. 1360–1363.

Singh, R. P. Lr46: a gene conferring slow-rusting resistance to leaf rust in wheat / R. P. Singh, A. Mujeeb-Kazi, J. Huerta-Espino // Phytopathology. – 1998. – Vol. 88. – P. 890–894.

Molecular marker mapping of leaf rust resistance gene Lr46 and its association with stripe rust resistance gene Yr29 in wheat / M. William [et al.] // Phytopathology. – 2003. – Vol. 93. – P. 153–159.

New slow-rusting leaf rust and stripe rust resistance genes Lr67 and Yr46 in wheat are pleiotropic or closely linked / S. Herrera-Foessel [et al.] // Theor. Appl. Genet. – 2011. – Vol. 122. – P. 239–249.

An introgression on wheat chromosome 4DL in RL6077 (Thatcher* 6/PI 250413) confers adult plant resistance to stripe rust and leaf rust (Lr67) / C. W. Hiebert [et al.] // Theor. Appl. Genet. – 2010a. – Vol. 121. – P. 1083–1091.

Singh, R. P. Complementary genes for reaction to Puccinia recondita tritici in Triticum aestivum. I. Genetic and linkage studies / R. P. Singh, R. A. McIntosh // Can. J. Genet. Cyt. – 1984. – Vol. 26. – P. 723–735.

A multiple resistance locus on chromosome arm 3BS in wheat confers resistance to stem rust (Sr2), leaf rust (Lr27) and powdery mildew / R. Mago [et al.] // Theor. Appl. Genet. – 2011. – Vol. 123, N 4. – P. 615–623.

Microsatellite markers for genes Lr34/Yr18 and other quantitative trait loci for leaf rust and stripe rust resistance in bread wheat / K. Suenaga [et al.] // Phytopathology. – 2003. – Vol. 93, N 7. – P. 881–890.

Bossolini, E. Development of simple sequence repeat markers specific for the Lr34 resistance region of wheat using sequence information from rice and Aegilops tauschii / E. Bossolini, S. G. Krattinger, B. Keller // Theor. Appl. Genet. – 2006. – Vol. 113, N 6. – P. 1049–1062.

Molecular-genetic characterization of the Lr34/Yr18 slow rusting resistance gene region in wheat / E. S. Lagudah [et al.] // Theor. Appl. Genet. – 2006. – Vol. 114, N 1. – P. 21–30.

Singh, D. Characterisation of wheat leaf rust resistance gene Lr34 in Australian wheats using components of resistance and the linked molecular marker csLV34 / D. Singh, R. F. Park, R. A. McIntosh // Austral. J. of Agricult. Res. – 2007. – Vol. 58. – P. 1106–1114.

Analysis of the Lr34/Yr18 rust resistance region in wheat germplasm / J. A. Kolmer [et al.] // Crop sci. – 2008. – Vol. 48, N 5. – P. 1841–1852.

Gene-specific markers for the wheat gene Lr34/Yr18/Pm38 which confers resistance to multiple fungal pathogens / E. S. Lagudah [et al.] // Theor. Appl. Genet. – 2009. – Vol. 119. – P. 889–898.

Fine-mapping of the leaf rust Lr34 locus in Triticum aestivum (L.) and characterization of large germplasm collections support the ABC transporter as essential for gene function / A. Dakouri [et al.] // Theor. Appl. Genet. – 2010. – Vol. 121. – P. 373–384.

A single-nucleotide polymorphism that accounts for allelic variation in the Lr34 gene and leaf rust reaction in hard winter wheat / S. Cao [et al.] // Theor. Appl. Genet. – 2010. – Vol. 121, N 2. – P. 385–392.

Galaev, A. V. Description of the bread wheat varieties of Ukrainian and Russian breeding by alleles of locus csLV34 closely linked with multipathogen resistance gene Lr34/Yr18/Pm38 / A. V. Galaev, Yu. M. Sivolap // Cytol. and Genet. – 2015. – Vol. 49, N 1. – P. 12–18.

Suenaga, K. Tagging of leaf rust resistance genes Lr34 and Lr46, using microsatellite markers in wheat / K. Suenaga // Aust. J. Agric. Res. – 2002. – Vol. 52. – P. 1383–1388.

Leaf tip necrosis, molecular markers and b1-proteasome subunits associated with the slow rusting resistance genes Lr46/Yr29 / G. M. Rosewarne [et al.] // Theor. Appl. Genet. – 2006. – Vol. 112. – P. 500–508.

Targered mapping of ESTs linked to the adult plant resistance gene Lr46 in wheat using systeny with rice / M. MateosHernandez [et al.] // Functional and integrative genomics. – 2006. – Vol. 6, N 2. – P. 122–131.

Quantitative trait loci for slow-rusting resistance in wheat to leaf rust and stripe rust identified with multi-environment analysis / G. M. Rosewarne [et al.] // Theor. Appl. Genet. – 2008. – Vol. 116, N 8. – P. 1027–1034.

Kolmer, J. A. Genetics of leaf rust resistance in the winter wheat line CI13227 / J. A. Kolmer, M. Lin, G. Bai // Crop Sci. – 2012. – Vol. 52. – P. 2166–2172.

Development of a SNP marker assay for the Lr67 gene of wheat using a genotyping by sequencing approach / K. Forrest [et al.] // Mol. Breeding. – 2014. – Vol. 34. – P. 2109–2118.

Spielmeyer, W. Identification and validation of markers linked to broad-spectrum stem rust resistance gene Sr2 in wheat (Triticum aestivum L.) / W. Spielmeyer, P. J. Sharp, E. S. Lagudash // Crop Sci. – 2003. – Vol. 43. – P. 333–336.

An accurate DNA marker assay for stem rust resistance gene Sr2 in wheat / R. Mago [et al.] // Theor. Appl. Genet. – 2011. – Vol. 122. – P. 735–744.

Developing and optimizing markers for stem rust resistance in wheat / Long-Xi Yu [et al.] // BGRI: proceed. of oral papers and posters of 2009 Technical Workshop, Cd. Obregon, Sonora, 17–20 March 2009 / ed. R. McIntosh. – Cd. Obregon; Sonora, 2009. – P. 117–130.

Haplotype diversity of stem rust resistance loci in uncharacterized wheat lines / Long-Xi Yu [et al.] // Mol. Breeding. – 2010. – Vol. 26. – P. 667–680.

Hayden, M. J. Sequence tagged microsatellites for the Xgwm 533 locus provide new diagnostic markers to select for the presence of stem rust resistance gene Sr2 in bread wheat (Triticum aestivum L.) / M. J. Hayden, H. Kuchel, J. Chalmers // Theor. Appl. Genet. – 2004. – Vol. 109. – P. 1641–1647. 124

BAC-derived markers for assaying the stem rust resistance gene, Sr2, in wheat breeding programs / M. D. McNeil [et al.] // Mol. Breed. – 2008. – Vol. 22. – P. 15–24.

Кохметова, А. М. Идентификация источников устойчивости к стеблевой ржавчине с использованием молекулярных маркеров / А. М. Кохметова, М. Н. Атишова // Вавилов. журн. генетики и селекции. – 2012. – Т. 16, № 1. – С. 132–141.

Dyck, P. L. Genetics of leaf rust resistance in three introductions of common wheat / P. L. Dyck // Can. J. Genet. Cyt. – 1977. – Vol. 19. – P. 711–716.

Rubiales, D. Characterization of Lr34, a major gene conferring nonhypersensitive resistance to wheat leaf rust / D. Rubiales, R. E. Niks // Plant Dis. – 1995. – Vol. 79. – P. 1208–1212.

Dyck, P. L. Inheritance of adult-plant leaf rust resistance derived from the common wheat varieties exchange and Frontana / P. L. Dyck, D. J. Samborski, R. G. Anderson // Can. J. Genet. Cyt. – 1966. – Vol. 8. – P. 665–671.

Recent emergence of the wheat Lr34 multi-pathogen resistance: insights from haplotype analysis in wheat, rice, sorghum and Aegilops tauschii / S. G. Krattinger [et al.] // Theor. Appl. Genet. – 2013. – Vol. 126. – P. 663–672.

Identification and characterization of pleiotropic and co–located resistance loci to leaf rust and stripe rust in bread wheat cultivar Sujata / C. Lan [et al.] // Theor. Appl. Genet. – 2015. – Vol. 128, N 3. – P. 549–561.

Niks, E. Haustorium formation by Puccinia hordei in leaves of hypersensitive, partially resistant, and nonhost genotypes / E. Niks // Phytopathology. – 1983. – Vol. 73. – P. 64–66.

Niks, R. E. Potentially durable resistance mechanisms in plants to specialized fungal pathogens / R. E. Niks, D. Rubiales // Euphytica. – 2002. – Vol. 124, N 2. – P. 201–216.

Leaf rust resistance gene Lr67, a third adult plant slow-rusting gene conferring resistance to multiple pathogens of wheat / C. Hiebert [et al.] // 8th international wheat conf.: abstracts of oral and poster presentations, St. Peterburg, 1–4 June 2010. – St. Peterburg, 2010. – P. 264.

Lr67 and Lr34 rust resistance genes have much in common-they confer broad spectrum resistance to multiple pathogens in wheat / W. Spielmeyer [et al.] // BMC Plant Biol. – 2013. – Vol. 13, N 96. – P. 2–9.

Kolmer, J. A. Expression of a Thatcher wheat adult plant stem rust resistance QTL on chromosome 2BL is enhanced by Lr34 / J. A. Kolmer, D. F. Garvin, Y. Jin // Crop Sci. – 2011. – Vol. 51. – P. 526–533.

Kerber, E. R. Leaf rust resistance gene Lr34 associated with nonsuppression of stem rust resistance in the wheat cultivar Canthatch / E. R. Kerber, T. Aung // Phytopathology. – 1999. – Vol. 89. – P. 518–521.

Vanegas, C. D. G. Genetics of stem rust resistance in the spring wheat cultivar Thatcher and the enhancement of stem rust resistance by Lr34 / C. D. G. Vanegas, D. F. Garvin, J. A. Kolmer // Euphytica. – 2008. – Vol. 159. – P. 391–401.

Genetics and mapping of seedling resistance to Ug99 stem rust in Canadian wheat cultivars “Peace” and “AC Cadillac” / C. W. Hiebert [et al.] // Theor. Appl. Genet. – 2010с. – Vol. 122. – P. 143–149.

Identification of Ug99 stem rust resistance loci in winter wheat germplasm using genome-wide association analysis / Long-Xi Yu [et al.] // Theor. Appl. Genet. – 2012. – Vol. 125, N 4. – P. 749–758.

Lr67/Yr46 confers adult plant resistance to stem rust and powdery mildew in wheat / S. A. Herrera-Foessel [et al.] // Theor. Appl. Genet. – 2014. – Vol. 127. – P. 781–789.

Dyck, P. L. An interchromosomal reciprocal translocation in wheat involving leaf rust resistance gene Lr34 / P. L. Dyck, E. R. Kerber, T. Aung // Genome. – 1994. – Vol. 37. – P. 556–559.

Singh, R. P. Association between gene Lr34 for leaf rust resistance and leaf tip necrosis in wheat / R. P. Singh // Crop Sci. – 1992b. – Vol. 32. – P. 874–878.

Functional variability of the Lr34 durable resistance gene in transgenic wheat / J. M. Risk [et al.] // Plant Biotechnol. J. – 2012. – Vol. 10. – P. 477–487.

Lagudah, E. S. Molecular genetics of race non-specific rust resistance in wheat // BGRI: oral presentations, full paper and abstracts of 2010 Technical Workshop, St. Petersburg, 30–31 May 2010. – St. Petersburg, 2010. – P. 183–196.

Курбанова, П. М. Генетическое разнообразие яровой мягкой пшеницы по эффективной возрастной устойчи- вости к листовой ржавчине: автореф. дис. ... канд. биол. наук: 03.02.07 / П. М. Курбанова; ВНИИР. – СПб., 2011. – 20 с.

German, S. E. Effect of gene Lr34 in the enhancement of resistance to leaf rust of wheat / S. E. German, J. A. Kolmer // Theor. Appl. Genet. – 1992. – Vol. 84. – P. 97–105.

Kloppers, F. J. Effects of combinations amongst genes Lr13, Lr34 and Lr37 on components of resistance in wheat to leaf rust / F. J. Kloppers, Z. A. Pretorius // Plant Pathol. – 1997. – Vol. 46. – P. 737–750.

Li, T. A combination of leaf rust resistance gene Lr34 and lesion mimic gene lm significantly enhances adult plant resistance to Puccinia triticina in wheat / T. Li, G. H. Bai, S. L. Gu // Chin. Sci. Bull. – 2012. – Vol. 57. – P. 2113–2119.

QTL for spot blotch resistance in bread wheat line Saar co-locate to the biotrophic disease resistance loci Lr34 and Lr46 / M. Lillemo [et al.] // Theor. Appl. Genet. – 2013. – Vol. 126. – P. 711–719.

Singh, R. P. Genetics and breeding for durable resistance to leaf and stripe rusts in wheat / R. P. Singh, J. HuertaEspino, H. M. William // Turk. J. Agric. For. – 2005. – Vol. 29. – P. 121–127.

Genetic analysis of resistance to leaf rust and stripe rust in wheat cultivar Francolin #1 / C. X. Lan [et al.] // Plant Dis. – 2014. – Vol. 98. – P. 1227–1234.

Pleiotropic gene Lr46/Yr29/Pm39/Ltn2 confers slow rusting, adult plant resistance to wheat stem rust fungus / R. P. Singh [et al.] // BGRI: proceed. of 2013 Technical Workshop, New Delhi, 19–22 August 2013. – New Delhi, 2013. – P. 17.1.

Genetic characteristics of wheat resistance gene Lr46/Yr29/Pm39, Sr2/Yr30 and Lr68 and association analysis of main agronomic traits / X. Yang [et al.] // Acta Botanica Boreali – Occidentalia Sinica. – 2014. – Vol. 34, N 3. – P. 454–462.

Dyck, P. L. The inheritance of resistance to Puccinia recondita in a group of common wheat cultivars / P. L. Dyck, D. J. Samborski // Can. J. Genet. Cyt. – 1982. – Vol. 24. – P. 273–283.

Differential expression of partial rsistance to wheat leaf rust in Mexico and The Southern Cone of America / S. German [et al.] // 8th international wheat conf.: abstracts of oral and poster presentations, St. Peterburg, 1–4 June 2010. – St. Peterburg, 2010. – P. 285–286.

100

Плотникова, Л. Я. Эффективность генов возрастной устойчивости пшеницы к бурой ржавчине Lr22b, Lr34, Lr37 в Западной Сибири и цитофизиологическая основа их действия / Л. Я. Плотникова, Т. Ю. Штубей // Вавилов. журн. генетики и селекции. – 2012. – Т. 16, № 1. – C. 123–131.

101

Singh, R. P. Expression of wheat leaf rust resistance gene Lr34 in seedlings and adult plants / R. P. Singh, A. K. Gupta // Plant Dis. – 1992. – Vol. 76. – P. 489–491.

102

Singh, R. P. Genetics of adult plant resistance to leaf rust in “Frontana” and three CIMMYT wheats / R. P. Singh, S. Rajaram // Genome. – 1992c. – Vol. 35. – P. 24–31.

103

Genetic analysis of adult-plant resistance to leaf rust in five spring wheat genotypes / A. Navabi [et al.] // Plant Dis. – 2003. – Vol. 87. – P. 1522–1529.

104

Inheritance of high levels of adult plant resistance to stripe rust in five spring wheat genotypes / A. Navabi [et al.] // Crop Sci. – 2004. – Vol. 44. – P. 1156–1162.

105

Inheritance of leaf rust resistance in CIMMYT wheat Weebill 1 / J. X. Zhang [et al.] // Crop Sci. – 2008. – Vol. 48. – P. 1037–1047.

106

Singh, R. P. Breeding spring bread wheat for irrigated and rainfed production systems of developing world / R. P. Singh, R. Trethowan // Breeding major food staples / ed.: M. Kang, P. M. Priyadarshan. – Iowa: Blackwell Publishing, 2007. – P. 109–140.

107

Singh, R. P. Genetics and breeding for durable resistance to leaf and stripe rusts of wheat increasing wheat production in Central Asia through science and international cooperation / R. P. Singh, J. Huerta-Espino, M. William // Proc. of the First Central Asian Wheat Conf., Almaty, 10–13 June 2003. – Almaty, 2003. – P. 127–132.

108

Molecular and pathological characterization of slow rusting against leaf rust in common wheat / S. Kumar [et al.] // BGRI: proceed. of oral papers and posters of 2009 Technical Workshop, Cd. Obregon, Sonora, 17–20 March 2009 / ed. R. McIntosh. – Cd. Obregon, Sonora, 2009. – P. 245.

109

McIntosh, R. A. Wheat rusts: an atlas of resistance genes / R. A. McIntosh, C. R. Park R. F. Wellings. – East Melbourne: CSIRO Publications, 1995. – 200 p.

110

Slow rusting – an effective way to achieve durable resistance against leaf rust in wheat / S. Kumar [et al.] // Wheat Information Service. – 2015. – Vol. 120. – P. 26–34.

111

McFadden, E. S. A successful transfer of emmer characters to vulgare wheat / E. S. McFadden // J. Am. Soc. Agron. – 1930. – Vol. 22. – P. 1020–1034.

112

Park, R. F. Stem rust of wheat in Australia / R. F. Park // Austral. J. of Agricult. Res. – 2007. – Vol. 58. – P. 558–566.

113

Sources of resistance to stem rust race Ug99 and its variants in Canadian wheat germplasm / R. M. DePauw [et al.] // BGRI: proceed. of oral papers and posters of 2009 Technical Workshop, Cd. Obregon, Sonora, 17–20 March 2009 / ed. R. McIntosh. – Cd. Obregon; Sonora, 2009. – P. 171–178.

114

Resistance to stem rust race Ug99 in the Canadian spring wheat cultivar “Peace” / G. Humphreys [et al.] // BGRI: proceed. of oral papers and posters of 2009 Technical Workshop, Cd. Obregon, Sonora, 17–20 March 2009 / ed. R. McIntosh. – Cd. Obregon; Sonora, 2009. – P. 253.

115

Implementaion of markers in Australian wheat breeding / H. A. Eagles [et al.] // Aust. J. Agric. Res. – 2001. – Vol. 52, N 12. – P. 1349–1356.

116

Singh, R. P. Complementary genes for resistance to Puccinia recondite tritici in Triticum aestivum. I. Genetic and linkage studies / R. P. Singh, R. A. McIntosh // Can. J. Genet. Cyt. – 1984. – Vol. 26. – P. 723–735.

117

Характеристика устойчивости к возбудителю бурой ржавчины сортов и линий мягкой пшеницы из коллекции ВИР, несущих чужеродный генетический материал / А. С. Садовая [и др.] // Вавилов. журн. генетики и селекции. – 2014. – Т. 18, № 4/1. – C. 739–750.

118

Schwessinger, B. Plant innate immunity: perception of conserved microbial signatures benjamin / B. Schwessinger, P. C. Ronald // Annu. Rev. Plant Biol. – 2012. – Vol. 63. – P. 451–482.

119

Effects and interactions of genes Lr34, Lr68 and Sr2 on wheat leaf rust adult plant resistance in Uruguay / P. Silva [et al.] // Euphytica. – 2015. – Vol. 204, N 3. – P. 599–608.

120

Application of phenotypic and molecular markers to combine genes for durable resistance against rust virulences and high yield potential in wheat / M. Hussain [et al.] // Int. J. Agric. Biol. – 2015. – Vol. 17. – P. 421−430.

121

The wheat Lr34 gene provides resistance against multiple fungal pathogens in barley / J. M. Risk [et al.] // Plant Biotechnol. J. – 2013. – Vol. 11. – P. 847–854.

The authors declare that there are no conflicts of interest present.