vestibИзвестия Национальной академии наук Беларуси. Серия биологических наукProceedings of the National Academy of Sciences of Belarus, Biological Series1029-89402524-230XThe Republican Unitary Enterprise Publishing House "Belaruskaya Navuka"vestib-281Research ArticleОБЗОРЫREVIEWSСТРАТЕГИИ УПРАВЛЕНИЯ ФОМОЗОМ РАПСА (BRASSICA NAPUS L.)BLACKLEG OF RAPE (BRASSICA NAPUS L.) MANAGEMENT STRATEGIESВолуевичЕ. А.VoluevichE. A.

д-р биол. наук, доцент, гл. науч. сотрудник

D. Sc. (Biol.), Associate Professor, Chief scientific employee

Voluevitch@yandex.ruИнститут генетики и цитологии НАН БеларусиInstitute of Genetics and Cytology of the National Academy of Sciences of Belarus20171902201701119128Copyright © Волуевич Е.А., 20172017Волуевич 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/281

Фомоз является широко распространенной болезнью рапса во всем мире, в том числе в Республике Беларусь. Для контроля этой болезни используют агротехнические приемы: севооборот, обработку почвы, оптимальные сроки посева, нормы высева семян, дозы внесения удобрений, обработки пестицидами. В зависимости от условий в зонах выращивания эту культуру рекомендуется возвращать на прежнее поле через 3–5 лет, в Беларуси – через 4–5 лет. Хорошим приемом контроля фомоза является запашка растительных остатков до осени. В некоторых странах для борьбы с фомозом проводят протравливание семян и опрыскивание почвы и растений фунгицидами. Лучший способ управления фомозом – выращивание устойчивых сортов. Поскольку Leptosphaeria maculans преодолевает главные гены устойчивости, болезнь можно контролировать путем диверсификации возделываемых сортов по генам устойчивости и их размещения. Ротацию культурных сортов, содержащих различные гены или комбинации генов устойчивости, рекомендуется проводить каждые 4 года. При этом важно осуществлять регулярный мониторинг частоты встречаемости генов вирулетности с целью определения эффективности известных главных генов устойчивости. Одной из стратегий увеличения длительности устойчивости к фомозу является применение одиночных главных генов или их различных сочетаний на генетическом фоне с неспецифической резистентностью. Длительность устойчивости можно повысить, используя смеси сортов или видов с разными генами устойчивости на одном поле и межполевую диверсификацию. Следует предусматривать и защиту рапса от L. biglobosa, который может наносить значительный ущерб, особенно сортам с эффективными главными генами устойчивости к L. maculans. Кроме того, для химической защиты рапса от L. biglobosa требуются более высокие дозы триазолов, чем для контроля L. maculans, поэтому существует большая потребность в селекции устойчивых сортов рапса как к L. maculans, так и к L. biglobosa. 

Blackleg is harmful and widespread disease of rape, including the Republic of Belarus. To control the disease are agricultural practices: crop rotation, tillage, the optimum time of sowing, seeding rate, fertilizer rates and pesticide treatments. Depending on the conditions in the areas of cultivation of this crop it is recommended to return to the previous field in 3–5 years, in Belarus – in 4–5 years. Good reception of blackleg control – plowing of crop residues to fall. Seed treatment and spraying of soil and plants with fungicides are conducted in different countries to combat blackleg disease. The best way to control of blackleg disease is cultivation of resistant varieties. Since Leptosphaeria maculans overcomes major resistance genes, the disease can be controlled through the diversification of cultivated varieties for resistance genes and their placement. Rotation of cultivars containing different genes or combinations of resistance genes is recommended to carry out every four years. Regular monitoring of virulent gene frequency is important in order to determine the effectiveness of known major resistance genes. One strategy for increasing of durable resistance to blackleg is the use of single major genes or their different combinations on genetic background with non-specific resistance. Durability of resistance can be increased using composition of species or varieties with different resistance genes on one field and interfield diversification. It is necessary and to provide of rape protect from L. biglobosa, which can cause significant damage, especially for varieties with effective major resistance genes to L. maculans. Moreover, chemical rape protection against L. biglobosa requires of higher triazole doses than to control L. maculans, so there is a great need for selection of rape cultivars with resistance to both L. maculans and L. biglobosa

рапсфомозгены устойчивостистратегии контроляrapeblacklegresistance genesstrategies of disease controlReferencesSNP markers-based map construction and genome-wide linkage analysis in Brassica napus / H. Raman [et al.] // Plant Biotechnol. J. – 2014. – Vol. 12. – P. 851–860.Raman, H., Dalton-Morgan, J., Diffey, S., Raman, R., Alamery, S., Edwards, D. and Batley, J. (2014), “SNP markers-based map construction and genome-wide linkage analysis in Brassica napus”, Plant Biotechnology Journal, vol. 12, pp. 851–860, doi: 10.1111/pbi.12186.Plant breeding: assessment of genetic diversity in crop plants and its exploitation in breeding / W. Friedt [et al.] // Progress in Botany. – 2007. – Vol. 168. – P. 151–178.Friedt, W., Snowdon, R., Ordon, F. and Ahlemeyer, J. (2007), “Plant breeding: assessment of genetic diversity in crop plants and its exploitation in breeding”, Progress in Botany, vol. 168, pp. 151–178.Conservation of the microstructure of genome segments in Brassica napus and its diploid relatives / D. Rana [et al.] // The Plant J. – 2004. – Vol. 40. – P. 725–733.Rana, D., van den Boogaart, T., O’Neill, C. M., Hynes, L., Bent, E., Macpherson, L., Park, J. Y., Lim, Y. P. and Bancroft, I. (2004), “Conservation of the microstructure of genome segments in Brassica napus and its diploid relatives”, The Plant Journal, vol. 40, pp. 725-733, doi: 10.1111/j.1365-313X.2004.02244.x.Characterization of flowering time and SSR marker analysis of spring and winter type Brassica napus L. germplasm / V. M. V. Cruz [et al.] // Euphytica. – 2007. – Vol. 153. – P. 43–57.Cruz, V. M. V., Luhman, R., Marek, L. F., Rife, C. L., Shoemaker, R. C., Brummer, E. C. and Gardner, C. A. C. (2007), “Characterization of flowering time and SSR marker analysis of spring and winter type Brassica napus L. germplasm”, Euphytica, vol. 153, pp. 43–57, doi:10.1007/s10681-006-9233-1.Bockus, W. W. The impact of reduced tillage on soilborne plant pathogens / W. W. Bockus, J. P. Shroyer // Annu. Rev. Phytopathol. – 1998. – Vol. 36. – P. 485–500.Bockus, W. W. and Shroyer, J. P. (1998), “The impact of reduced tillage on soilborne plant pathogens”, Annual Review Phytopathology, vol. 36, pp. 485–500, doi: 10.1146/annurev.phyto.36.1.485.Guo, X. W. Seasonal and diurnal patterns of spore dispersal by Leptosphaeria maculans from canola stubble in relation to environmental conditions / X. W. Guo, W. G. D., Fernando // Plant Disease. – 2005. – Vol. 89. – P. 97–104.Guo, X. W. and Fernando, W. G. D. (2005), “Seasonal and diurnal patterns of spore dispersal by Leptosphaeria maculans from canola stubble in relation to environmental conditions”, Plant Disease, vol. 89, pp. 97–104.Epidemiology and management of Leptosphaeria maculans (phoma stem canker) on oilseed rape in Australia, Canada and Europe / J. S. West [et al.] // Plant Pathol. – 2001. – Vol. 50. – P. 10–27.West, J. S., Kharband, P. D., Barbetti, M. J. and Fitt, B. D. L. (2001), “Epidemiology and management of Leptosphaeria maculans (phoma stem canker) on oilseed rape in Australia, Canada and Europe”, Plant Pathology, vol. 50, pp. 10–27, doi: 10.1046/j.1365-3059.2001.00546.x.Fernando, W. G. D. Breeding for blackleg resistance: the biology and epidemiology / W. G. D. Fernando, Y. Chen, K. Ghanbarnia // Adv. Bot. Res. – 2007. – Vol. 45. – P. 271–311.Fernando, W. G. D., Chen, Y. and Ghanbarnia, K. (2007), “Breeding for blackleg resistance: the biology and epidemiology”, Advances in Botanical Research, vol. 45, pp. 271–311.Gladders, P. Observations on the epidemiology of Leptosphaeria maculans stem canker in winter oilseed rape / P. Gladders, T. M. Musa // Plant Pathol. – 1980. – Vol. 29. – P. 28–37.Gladders, P. and Musa, T. M. (1980), “Observations on the epidemiology of Leptosphaeria maculans stem canker in winter oilseed rape”, Plant Pathology, vol. 29, pp. 28–37.McGee, D. C. Blackleg (Leptosphaeria maculans (Desm.) Ces. et de Not.) of rapeseed in Victoria: crop losses and factors which affect disease severity / D. C. McGee, R. W. Emmett // Aust. J. Agric. Res. – 1977. – Vol. 28. – P. 47–51.McGee, D. C. and Emmett, R. W. (1977), “Blackleg (Leptosphaeria maculans (Desm.) Ces. et de Not.) of rapeseed in Victoria: crop losses and factors which affect disease severity”, Australian Journal of Experimental Agriculture, vol. 28, pp. 47–51.Hammond, K. E. A systemic pathway in the infection of oilseed rape plants by Leptosphaeria maculans / K. E. Ham- mond, B. G. Lewis, T. M. Musa // Plant Pathol. – 1985. – Vol. 34, N 4. – P. 557–565.Hammond, K. E., Lewis, B. G. and Musa, T. M. (1985), “A systemic pathway in the infection of oilseed rape plants by Leptosphaeria maculans”, Plant Pathology, vol. 34, no. 4, pp. 557–565.Epidemiology of Leptosphaeria maculans in relation to forecasting stem canker severity on winter oilseed rape in the UK / J. S. West [et al.] // Ann. Appl. Biol. – 1999. – Vol. 135. – P. 535–546.West, J. S., Biddulph, J. E., Fitt, B. D. L. and Gladders, P. (1999), “Epidemiology of Leptosphaeria maculans in relation to forecasting stem canker severity on winter oilseed rape in the UK”, Annals of Applied Biology, vol. 135, pp. 535–546.Effects of timing of Leptosphaeria maculans ascospore release and fungicide regime on phoma leaf spot and phoma stem canker development on winter oilseed rape (Brassica napus) in southern England / J. S. West // Plant Pathol. – 2002b. – Vol. 51. – P. 454–463.West, J. S., Fitt, B. D. L., Leech, P. K., Biddulph, J. E., Huang, Y.-J. and Balesdent, M.-H. (2002b), “Effects of timing of Leptosphaeria maculans ascospore release and fungicide regime on phoma leaf spot and phoma stem canker development on winter oilseed rape (Brassica napus) in southern England”, Plant Pathology, vol. 51, pp. 454–463.Kutcher, H. R. Increased frequency of canola in crop rotations associated with increased blackleg disease / H. R. Kutcher, S. A. Brandt // Can. J. Plant Pathol. – 2009. – Vol. 31. – P. 123–124.Kutcher, H. R. and Brandt, S. A. (2009), “Increased frequency of canola in crop rotations associated with increased blackleg disease”, Canadian Journal of Plant Pathology, vol. 31, pp. 123–124.The Leptosphaeria maculans – Leptosphaeria biglobosa species complex in the American continent / A. Dilmaghani [et al.] // Plant Pathol. – 2009. – Vol. 58, N 6. – P. 1044–1058.Dilmaghani, A., Balesdent, M. H., Didier, J. P., Wu, C., Davey, J., Barbetti, M. J., Li, H., Moreno-Rico, O., Phillips, D., Despeghel, J. P., Vincenot, L., Gout, L. and Rouxel, T. (2009), “The Leptosphaeria maculans – Leptosphaeria biglobosa species complex in the American continent”, Plant Pathology, vol. 58, no. 6, pp. 1044–1058, doi: 10.1111/j.1365-3059.2009.02149.x.Crouch, J. H. The effect of a genome substitution on the resistance of Brassica napus to infection by Leptosphaeria maculans / J. H. Crouch, B. G. Lewis, R. F. Mithen // Plant Breeding. – 1994. – Vol. 112. – P. 265–278.Crouch, J. H., Lewis, B. G. and Mithen, R. F. (1994), “The effect of a genome substitution on the resistance of Brassica napus to infection by Leptosphaeria maculans”, Plant Breeding, vol. 112, pp. 265-278, doi: 10.1111/j.1439-0523.1994. tb00684.x.Yu, F. Identification of two novel genes for blackleg resistance in Brassica napus / F. Yu, D. J. Lydiate, S. R. Rimmer // Theor. Appl. Genet. – 2005. – Vol. 110. – P. 969–979.Yu, F., Lydiate, D. J. and Rimmer, S. R. (2005), “Identification of two novel genes for blackleg resistance in Brassica napus”, Theoretical and Applied Genetics, vol. 110, pp. 969–979, doi: 10.1007/s00122-004-1919-y.Yu, F. Identification and mapping of a third blackleg resistance locus in Brassica napus derived from B. rapa subsp. sylvestris / F. Yu, D. J. Lydiate, S. R. Rimmer // Genome. – 2008. – Vol. 51. – P. 64–72.Yu, F., Lydiate, D. J. and Rimmer, S. R. (2008), “Identification and mapping of a third blackleg resistance locus in Brassica napus derived from B. rapa subsp. sylvestris”, Genome, vol. 51, pp. 64–72, doi: 10.1139/G07-103.Screening and identification of resistance to Leptosphaeria maculans (stem canker) in Brassica napus accessions / T. Rouxel [et al.] // Euphytica. – 2003. – Vol. 133. – P. 219–231.Rouxel, T., Willner, E., Coudard, L. and Balesdent, M.-H. (2003b), “Screening and identification of resistance to Leptosphaeria maculans (stem canker) in Brassica napus accessions”, Euphytica, vol. 133, pp. 219–231.Li, C. X. Identification of a single dominant allele for resistance to blackleg in Brassica napus “Surpass 400” / C. X. Li, W. A. Cowling // Plant Breeding. – 2003a. – Vol. 122. – P. 485–488.Li, C. X. and Cowling, W. A. (2003), “Identification of a single dominant allele for resistance to blackleg in Brassica napus “Surpass 400”, Plant Breeding, vol. 122, pp. 485–488, doi: 10.1111/j.1439-0523.2003.00863.x.Quantitative resistance increases the durability of qualitative resistance to Leptosphaeria maculans in Brassica napus / H. Brun [et al.] // New Phytologist. – 2010. – Vol. 185. – P. 285–299.Brun, H., Chèvre, A. M., Fitt, B. D., Powers, S., Besnard, A.-L., Ermel, M., Huteau, V., Marquer, B., Eber, F., Renard, M. and Andrivon, D. (2010), “Quantitative resistance increases the durability of qualitative resistance to Leptosphaeria mac-ulans in Brassica napus”, New Phytologist, vol. 185, pp. 285–299, doi: 10.1111/j.1469-8137.2009.03049.x.Complexities of chromosome landing in a highly duplicated genome: toward map-based cloning of a gene controlling blackleg resistance in Brassica napus / R. Mayerhofer [et al.] // Genetics. – 2005. – Vol. 171. – P. 1977–1988.Mayerhofer, R., Wilde, K., Mayerhofer, M., Lydiate, D., Bansal, V. K., Good, A. G. and Parkin, I. A. (2005), “Complexities of chromosome landing in a highly duplicated genome: toward map-based cloning of a gene controlling blackleg resistance in Brassica napus”, Genetics, vol. 171, pp. 1977–1988, doi: 10.1534/genetics.105.049098.Effector diversification within compartments of the Leptosphaeria maculans genome affected by repeat-induced point mutations / T. Rouxel [et al.] // Nature Communications. – 2011. – Vol. 2, N 202. – P. 1–10.Rouxel, T., Grandaubert, J., Hane, J. K., Hoede, C., van de Wouw, A. P., Couloux, A., Dominguez, V., Anthouard, V., Bally, P., Bourras, S., Cozijnsen, A. J., Ciuffetti, L. M., Degrave, A., Dilmaghani, A., Duret, L., Fudal, I., Goodwin, S. B., Gout, L., Glaser, N., Linglin, J., Kema, G. H. J., Lapalu, N., Lawrence, C. B., May, K., Meyer, M., Ollivier, B., Poulain, J., Schoch, C. L., Simon, A., Spatafora, J. W., Stachowiak, A., Turgeon, B. G., Tyler, B. M., Vincent, D., Weissenbach, J., Amselem, J., Quesneville, H., Oliver, R. P., Wincker, P., Balesdent, M.-H. and Howlett, B. J. (2011), “Effector diversification within compartments of the Leptosphaeria maculans genome affected by repeat-induced point mutations”, Nature Communications, vol. 2, no. 202, pp. 1–10, doi: 10.1038/ncomms1189.A ten-year survey of populations of Leptosphaeria maculans in France indicates a rapid adaptation towards the Rlm1 resistance gene of oilseed rape / T. Rouxel [et al.] // Eur. J. Plant Pathol. – 2003. – Vol. 109. – P. 871–881.Rouxel, T., Penaud, A., Pinochet, X., Brun, H., Gout, L., Delourme, R., Schmit, J. and Balesdent, M.-H. (2003а), “A ten-year survey of populations of Leptosphaeria maculans in France indicates a rapid adaptation towards the Rlm1 resistance gene of oilseed rape”, European Journal of Plant Pathology, vol. 109, pp. 871–881.Major gene resistance to blackleg in Brassica napus overcome within three years of commercial production in southeastern Australia / S. J. Sprague [et al.] // Plant Pathol. – 2006. – Vol. 59. – P. 190–198.Sprague, S. J., Marcroft, S. J., Hayden, H. L. and Howlett, B. J. (2006), “Major gene resistance to blackleg in Brassica napus overcome within three years of commercial production in southeastern Australia”, Plant Pathology, vol. 59, pp. 190–198, doi: 10.1094/PD-90-0190.Rouxel, T. The stem canker (blackleg) fungus, Leptosphaeria maculans, enters the genomic era / T. Rouxel, M. H. Balesdent // Mol. Plant Pathol. – 2005. – Vol. 6. – P. 225–241.Rouxel, T. and Balesdent, M. H. (2005), “The stem canker (blackleg) fungus, Leptosphaeria maculans, enters the genomic era”, Mol. Plant Pathol., vol. 6, pp. 225–241, doi: 10.1111/j.1364-3703.2005.00282.x.Effect of rotation of canola (Brassica napus) cultivars with different compliments of blackleg resistance genes on disease severity / S. J. Marcroft [et al.] // Plant Pathol. – 2012. – Vol. 61. – P. 934–944.Marcroft, S. J., van de Wouw, A. P., Salsibury, P. A., Pottere, T. D. and Howlett, B. J. (2012), “Effect of rotation of canola (Brassica napus) cultivars with different compliments of blackleg resistance genes on disease severity”, Plant Pathology, vol. 61, pp. 934–944, doi: 10.1111/j.1365-3059.2011.02580.x.Breakdown of Rlm3 resistance in the Brassica napus–Leptosphaeria maculans pathosystem in western Canada / X. Zhang [et al.] // Eur. J. Plant Pathol. – 2015. – Vol. 145, N 3. – P. 1–16.Zhang, X., Peng, G., Kutcher, H. R., Balesdent, M.-H., Delourme, R. and Fernando, W. G. D. (2015), “Breakdown of Rlm3 resistance in the Brassica napus–Leptosphaeria maculans pathosystem in western Canada”, European Journal of Plant Pathology, vol. 145, no. 3, pp. 1–16, doi: 10.1007/s10658-015-0819-0.Frequency of avirulence alleles in field populations of Leptosphaeria maculans in Europe / A. Stachowiak [et al.] // Eur. J. Plant Pathol. – 2006. – Vol. 114. – P. 67–75.Stachowiak, A., Olechnowicz, J., Jedryczka, M., Rouxel, T., Balesdent, M.-H., Happstadius, I., Gladders, P., Latunde-Dada, A. and Evans, N. (2006), “Frequency of avirulence alleles in field populations of Leptosphaeria maculans in Europe”, European Journal of Plant Pathology, vol. 114, pp. 67–75, doi: 10.1007/s10658-005-2931-z.A field method for evaluating the potential durability of new resistance sources: application to the Leptosphaeria maculans/Brassica napus pathosystem / H. Brun [et al.] // Phytopathology. – 2000. – Vol. 90. – P. 961–966.Brun, H., Levivier, S., Somda, I., Ruer, D., Renard, M. and Chèvre, A. M. (2000), “A field method for evaluating the potential durability of new resistance sources: application to the Leptosphaeria maculans/Brassica napus pathosystem”, Phytopathology, vol. 90, pp. 961–966.Evolution of linked avirulence effectors in Leptosphaeria maculans is affected by genomic environment and exposure to resistance genes in host plants / A. P. Van de Wouw [et al.] // PLoS Pathol. – 2010. – Vol. 6, N 11. – P. 1–15.Van de Wouw, A. P., Cozijnsen, A. J., Hane, J. K., Brunner, P. C., McDonald, B. A., Oliver, R. P. and Howlett, B. J. (2010), “Evolution of linked avirulence effectors in Leptosphaeria maculans is affected by genomic environment and exposure to resistance genes in host plants”, PLoS Pathogens, vol. 6, no. 11, pp. 1–15.Heterochromatin-like regions as ecological niches for avirulence genes in the Leptosphaeria maculans genome: map-based cloning of AvrLm6 / I. Fudal [et al.] // MPMI. – 2007. – Vol. 20. – P. 459–470.Fudal, I., Ross, S., Gout, L., Blaise, F., Kuhn, M. L., Eckert, M. R., Cattolico, L., Bernard-Samain, S., Balesdent, M. H. and Rouxel, T. (2007), “Heterochromatin-like regions as ecological niches for avirulence genes in the Leptosphaeria maculans genome: map-based cloning of AvrLm6”, MPMI, vol. 20, pp. 459–470.Lost in the middle of nowhere: the AvrLm1 avirulence gene of the Dothideomycete Leptosphaeria maculans / L. Gout [et al.] // Mol. Microbiol. – 2006. – Vol. 60. – P. 67–80.Gout, L., Fudal, I., Kuhn, M.-L., Blaise, F., Eckert, M., Cattolico, L., Balesdent, M.-H. and Rouxel, T. (2006), “Lost in the middle of nowhere: the AvrLm1 avirulence gene of the Dothideomycete Leptosphaeria maculans”, Mol. Microbiol., vol. 60, pp. 67–80, doi:10.1111/j.1365-2958.2006.05076.x.Evolution of the frequency of the AvrLm7 allele of Leptosphaeria maculans in France under selection pressure: a 15-years survey / C. Plissonneau [et al.] // Healthy plant – healthy people: 11th conf. of the European foundation for plant pathology, Kraków, 8–13 September 2014. – Kraków, 2014. – P. 129.Plissonneau, C., Rouxel, T., Daverdin, G. Meur, L. L., Soulard, T., Cugnire, L. and Balesdent, M.-H. (2014), “Evolution of the frequency of the AvrLm7 allele of Leptosphaeria maculans in France under selection pressure: a 15-years survey”, Healthy plant – healthy people: 11th conf. of the European foundation for plant pathology, Kraków, PL, 8–13 September 2014, p. 129.Li, H. Breakdown of a Brassica rapa subsp. sylvestris single dominant blackleg resistance gene in Brassica napus rapeseed by Leptosphaeria maculans field isolates in Australia / H. Li, K. Sivasithamparam, M. J. Barbetti // Plant Dis. – 2003b. – Vol. 87. – P. 752.Li, H., Sivasithamparam, K. and Barbetti, M. J. (2003b),” Breakdown of a Brassica rapa subsp. sylvestris single dominant blackleg resistance gene in Brassica napus rapeseed by Leptosphaeria maculans field isolates in Australia”, Plant Disease, vol. 87, p. 752, doi: 10.1094/PDIS.2003.87.6.752A.Chen, C. Ability of a Leptosphaeria maculans isolate to form stem canker on Indian mustard (Brassica juncea) segregate as a single locus / C. Chen, K. Plummer, B. J. Howlett // Eur. J. Plant Pathol. – 1996. – Vol. 102. – P. 349–352.Chen, C., Plummer, K. and Howlett, B. J. (1996), “Ability of a Leptosphaeria maculans isolate to form stem canker on Indian mustard (Brassica juncea) segregate as a single locus”, European Journal of Plant Pathology, vol. 102, pp. 349–352, doi: 10.1007/BF01878129.Pathogenic variability of Leptosphaeria maculans in western Canada / H. R. Kutcher [et al.] // Can. J. Plant Pathol. – 2007. – Vol. 29, N 4. – P. 388–393.Kutcher, H. R., Keri, M., McLaren, D. L. and Rimmer, S. R. (2007), “Pathogenic variability of Leptosphaeria maculans in western Canada”, Can. J. Plant Pathol., vol. 29, no. 4, pp. 388–393, doi: 10.1080/07060660709507484.Breakdown of resistance to the fungal disease, blackleg, is averted in commercial canola (Brassica napus) crops in Australia / A. P. Van de Wouw [et al.] // Field Crops Res. – 2014b. – Vol. 166. – P. 144–151.Van de Wouw, A. P., Macroft, S. J., Ware A., Lindbeck K., Khangura R. and Howlett, B. J. (2014b), “Breakdown of resistance to the fungal disease, blackleg, is averted in commercial canola (Brassica napus) crops in Australia”, Field Crops Research, vol. 166, pp. 144–151, doi: 10.1016/j.fcr.2014.06.023.Blackleg disease of canola mitigated by resistant cultivars and four-year crop rotations in western Canada / H. R. Kut- cher [et al.] // Can. J. Plant Pathol. – 2013. – Vol. 35. – P. 209–221.Kutcher, H. R., Brandt, S. A., Smith, E. G., Ulrich, D., Malhi, S. S. and Johnston, A. M. (2013), “Blackleg disease of canola mitigated by resistant cultivars and four-year crop rotations in western Canada”, Canadian Journal of Plant Pathology, vol. 35, pp. 209–221, doi: 10.1080/07060661.2013.775600.Major gene and polygenic resistance to Leptosphaeria maculans in oilseed rape (Brassica napus) / R. Delourme [et al.] // Eur. J. Plant Pathol. – 2006. – Vol. 114. – P. 41–52.Delourme, R., Chèvre, A. M., Brun, H., Rouxel, T., Balesdent, M. H., Dias, J. S., Salisbury, P., Renard, M. and Rimmer, S. R. (2006), “Major gene and polygenic resistance to Leptosphaeria maculans in oilseed rape (Brassica napus)”, European Journal of Plant Pathology, vol. 114, pp. 41–52, doi: 10.1007/s10658-005-2108-9.Potential for using host-resistance to reduce production of pseudothecia and ascospores of Leptosphaeria maculans, the blackleg pathogen of Brassica napus / S. J. Marcroft [et al.] // Plant Pathol. – 2004. – Vol. 53. – P. 468–474.Marcroft, S. J., Sprague, S. J., Salisbury, P. A. and Howlett, B. J. (2004), “Potential for using host-resistance to reduce production of pseudothecia and ascospores of Leptosphaeria maculans, the blackleg pathogen of Brassica napus”, Plant Pathology, vol. 53, pp. 468–474, doi: 10.1111/j.1365-3059.2004.01050.x.Rimmer, S. R. Resistance genes to Leptosphaeria maculans in Brassica napus / S. R. Rimmer // Can. J. Plant Pathol. – 2006. – Vol. 28. – S. 288–297.Rimmer, S. R. (2006), “Resistance genes to Leptosphaeria maculans in Brassica napus”, Canadian Journal of Plant Pathology, vol. 28, pp. 288–297.Leach, J. E. Bacterial avirulence genes / J. E. Leach, F. F. White // Annu. Rev. Phytopathol. – 1996. – Vol. 34. – P. 153–179.Leach, J. E. and White, F. F. (1996), “Bacterial avirulence genes”, Annual Review Phytopathology, vol. 34, pp. 153–179, doi: 10.1146/annurev.phyto.34.1.153.Волуевич, Е. А. Генетическая природа полигенной устойчивости мягкой пшеницы к бурой ржавчине / Е. А. Волуевич, А. А. Булойчик, В. С. Борзяк // Докл. Нац. акад. Беларуси. – 2006. – Т. 50, № 1. – С. 59–61.Voluevich, E. A. (2006), “Genetic nature of polygenic wheat leaf rust resistance”. Doklady of the National Academy of Sciences of Belarus, vol. 50, no. 1, pp. 59–61.Волуевич, Е. А. Остаточный эффект преодоленных генов устойчивости мягкой пшеницы к биотрофным грибным патогенам / Е. А. Волуевич // Молекулярная и прикладная генетика: сб. науч. тр. / Гос. науч. учреждение “Институт генетики и цитологии Национальной академии наук Беларуси”. – Минск, 2014. – Т. 17. – С. 78–84.Voluevich, E. A. (2014), “Residual effect of defeated genes of wheat resistance to biotrophic fungal pathogens”, Molekularnaja i prikladnaya genetika: sbornik nauchnukh trydov [Molecular and Applied Genetics: digest of scientific papers], Institut genetiki i citologii Nacional’noj akademii nauk Belarusi, Minsk, BY, vol. 17, pp. 78–84.Kiyosawa, S. Genetics and epidemiological modeling of breakdown of plant disease resistance / S. Kiyosawa // Annu. Rev. Phytopathol. – 1982. – Vol. 20. – P. 93–117.Kiyosawa, S. (1982), “Genetics and epidemiological modeling of breakdown of plant disease resistance”, Annual Review Phytopathology, vol. 20, pp. 93–117, doi: 10.1146/annurev.py.20.090182.000521.Pietravalle, S. Durability of resistance and cost of virulence / S. Pietravalle, S. Lemarié, F. van den Bosch // Eur. J. Plant Pathol. – 2006. – Vol. 114, N 1. – P. 107–116.Pietravalle, S., Lemarié, S. and van den Bosch, F. (2006), “Durability of resistance and cost of virulence”, Eur. J. Plant Pathol., vol. 114, no. 1. pp. 107–116, doi:10.1007/s10658-005-3479-7.Quantitative resistance affects the speed of frequency increase but not the diversity of the virulence alleles overcoming a major resistance gene to Leptosphaeria maculans in oilseed rape / R. Delourme [et al.] // Infect. Genet. Evol. – 2014. – Vol. 27. – P. 490–499.Delourme, R., Bousset, L., Ermel, M., Duffé, P., Besnard, A. L., Marque, B., Fudal, I., Linglin, J., Chadœuf, J. and Brun, H. (2014), “Quantitative resistance affects the speed of frequency increase but not the diversity of the virulence alleles overcoming a major resistance gene to Leptosphaeria maculans in oilseed rape”, Infection, genetics and evolution, vol. 27, pp. 490–499, doi: 10.1016/j.meegid.2013.12.019.Molecular mapping of qualitative and quantitative loci for resistance to Leptosphaeria maculans causing blackleg disease in canola (Brassica napus L.) / R. Raman [et al.] // Theor. Appl. Genet. – 2012b. – Vol. 125. – P. 405–418.Raman, R., Taylor, B., Marcroft, S., Stiller, J., Eckermann, P., Coombes, N., Rehman, A., Lindbeck, K., Luckett, D., Wratten, N., Batley, J., Edwards, D., Wang, X. and Raman, H. (2012b),” Molecular mapping of qualitative and quantita- tive loci for resistance to Leptosphaeria maculans causing blackleg disease in canola (Brassica napus L.)”, Theoretical and Applied Genetics, vol. 125, pp. 405–418, doi: 10.1007/s00122-012-1842-6.World-wide importance of Phoma stem canker (Leptosphaeria maculans and L. biglobosa) on oilseed rape (Brassica napus) / B. D. L. Fitt [et al.] // Eur. J. Plant Pathol. – 2006. – Vol. 114, N 1. – P. 3–15.Fitt, B. D. L., Brun, H., Barbetti, M. J. and Rimmer, S. R. (2006), “World-wide importance of Phoma stem canker (Leptosphaeria maculans and L. biglobosa) on oilseed rape (Brassica napus)”, European Journal of Plant Pathology, vol. 114, no. 1, pp. 3–15, doi: 10.1007/s10658-005-2233-5.Strategies to prevent spread of Leptosphaeria maculans (phoma stem canker) onto oilseed rape crops in China: costs and benefits / B. D. L. Fitt [et al.] // Plant Pathol. – 2008. – Vol. 57. – P. 652–664.Fitt, B. D. L., Hu, B. C., Li, Z. Q., Liu, S. Y., Lange, R. M., Kharbanda, P. D., Butterworth, M. H. and White, R. P. (2008),” Strategies to prevent spread of Leptosphaeria maculans (phoma stem canker) onto oilseed rape crops in China: costs and benefits”, Plant Pathology, vol. 57, pp. 652–664, doi: 10.1111/j.1365-3059.2008.01841.x.Factors affecting production of inoculum of the blackleg fungus Leptosphaeria maculans in south-eastern Australia / S. J. Marcroft [et al.] // Aust. J. Exp. Agr. – 2003. – Vol. 43. – P. 1231–1236.Marcroft, S. J., Sprague, S. J., Pymer, S. J., Salisbury, P. A. and Howlett, B. J. (2003), “Factors affecting production of inoculum of the blackleg fungus Leptosphaeria maculans in south-eastern Australia”, Australian Journal of Experimental Agriculture, vol. 43, pp. 1231–1236.Blackleg resistance in rapeseed: phenotypic screen, molecular markers and genome wide linkage and association mapping / H. Raman [et al.] // 17th Australian Research Assembly on Brassicas (ARAB), Wagga Wagga, 15–17 August 2011. – Wagga Wagga, 2011. – P. 61–64.Raman, H., Raman, R., Taylor, B., Lindbeck, K., Coombes, N., Eckermann, P., Batley, J., Edwards, D., Price, A., Rehman, A., Marcroft, S., Luckett, D., Hossain, S. and Salisbury, P. (2011), “Blackleg resistance in rapeseed: phenotypic screen, molecular markers and genome wide linkage and association mapping”, 17th Australian Research Assembly on Bras- sicas (ARAB), Wagga Wagga, AU, 15–17 August 2011, pp. 61–64.Association mapping of quantitative resistance for Leptosphaeria maculans in oilseed rape (Brassica napus L.) / C. Jestin [et al.] // Mol. Breeding. – 2011. – Vol. 27. – P. 271–287.Jestin, C., Lodé, M., Vallée P., Domin, C., Falentin, C., Horvais, R., Coedel, S., Manzanares-Dauleux, M. J. and Delourme, R. (2011), “Association mapping of quantitative resistance for Leptosphaeria maculans in oilseed rape (Brassica napus L.)”, Molecular Breeding, vol. 27, pp. 271–287, doi: 10.1007/s11032-010-9429-x.Huang, Y. J. Importance of Leptosphaeria biglobosa as a cause of phoma stem canker on winter oilseed rape in the UK. Crop Production in Southern Britain: Precision Decisions for Profitable Cropping / Y. J. Huang, C. S. Karandeni-Dewage, B. D. L. Fitt // Aspects of Appl. Biol. – 2014. – Vol. 127. – P. 117–122.Huang, Y. J., Karandeni-Dewage, C. S. and Fitt, B. D. L. (2014), “Importance of Leptosphaeria biglobosa as a cause of phoma stem canker on winter oilseed rape in the UK. Crop Production in Southern Britain: Precision Decisions for Profitable Cropping”, Aspects of Applied Biology, vol. 127, pp. 117–122.Sidique, S. N. M. Effects of host resistance on colonisation of Brassica napus (oilseed rape) by Leptosphaeria maculans and L. biglobosa (phoma stem canker): Ph.D. Thesis … of Doctor of Philosophy / S. N. M. Sidique; University of Hertfordshire. – Hatfield, 2015. – 475 p.Sidique, S. N. M. (2015), “Effects of host resistance on colonisation of Brassica napus (oilseed rape) by Leptosphaeria maculans and L. biglobosa (phoma stem canker)”, Ph. D. Thesis, University of Hertfordshire, Hatfield, US.Kaczmarek, J. Characterization of two coexisting pathogen populations of Leptosphaeria spp., the cause of stem canker of Brassicas / J. Kaczmarek, M. Jedryczka // Acta Agrobotanica. – 2011. – Vol. 64. – P. 3–14.Kaczmarek, J. and Jedryczka, M. (2011), “Characterization of two coexisting pathogen populations of Leptosphaeria spp., the cause of stem canker of Brassicas”, Acta Agrobotanica, vol. 64, pp. 3–14.Improved resistance management for durable disease control: a case study of phoma stem canker of oilseed rape / J. N. Aubertot [et al.] // Eur. J. Plant Pathol. – 2006. – Vol. 114. – P. 91–106.Aubertot, J. N., West, J. S., Bousset-Vaslin, L., Salam, M. U., Barbetti, M. J. and Diggle, A. J. (2006), “Improved resistance management for durable disease control: a case study of phoma stem canker of oilseed rape”, European Journal of Plant Pathology, vol. 114, pp. 91–106.

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