bioselБиотехнология и селекция растенийPlant Biotechnology and Breeding2658-62662658-6258VIR10.30901/2658-6266-2025-2-o3biosel-276Research ArticleИЗУЧЕНИЕ ГЕНЕТИЧЕСКИХ РЕСУРСОВ РАСТЕНИЙ С ИСПОЛЬЗОВАНИЕМ МЕТОДОВ МОЛЕКУЛЯРНОЙ ГЕНЕТИКИSTUDY OF PLANT GENETIC RESOURCES USING MOLECULAR GENETICS METHODSМолекулярный скрининг коллекции груши, поддерживаемой на Майкопской опытной станции ВИР, на наличие маркеров генов устойчивости к паршеMolecular screening of the pear collection maintained at the Maikop Experiment Station of VIR for scab resistance gene markershttps://orcid.org/0009-0007-2474-752XГончаренкоА. О.GoncharenkoA. O.

Анастасия Олеговна Гончаренко, младший научный сотрудник, лаборатория молекулярной селекции и ДНК-паспортизации, отдел биотехнологии, ВИР

190000 Россия, Санкт-Петербург, ул. Большая Морская, 42, 44

Anastasiia O. Goncharenko, Junior Researcher, Laboratory of Molecular breeding and DNA-genotyping, Department of Biotechnology, VIR

42, 44, Bolshaya Morskaya Street, St. Petersburg, 190000 Russia

aogoncharenko97@gmail.comhttps://orcid.org/0000-0003-0768-0056БагметЛ. В.BagmetL. V.

Лариса Владимировна Багмет , кандидат биологических наук, ведущий научный сотрудник, отдел агроботаники и in situ сохранениягенетических ресурсов растений, ВИР

190000 Россия, Санкт-Петербург, ул. Большая Морская, 42, 44

Larisa V. Bagmet, Cand. Sci. (Biology), Leading Researcher, Department of Agrobotany and in situ Conservation of Plant Genetic Resources, VIR

42, 44, Bolshaya Morskaya Street, St. Petersburg, 190000 Russia

l.bagmet@vir.nw.ruhttps://orcid.org/0000-0001-5700-0384ПетроваМ. Н.PetrovaM. N.

Марина Николаевна Петрова, кандидат сельскохозяйственных наук, ведущий специалист, отдел генетических ресурсов плодовых культур, ВИР

190000 Россия, Санкт-Петербург, ул. Большая Морская, 42, 44

Marina N. Petrova, Cand. Sci. (Agriculture), Leading Specialist, Department of Genetic Resources of Fruit Crops, VIR

42, 44, Bolshaya Morskaya Street, St. Petersburg, 190000 Russia

adresspb-petrova@yandex.ruhttps://orcid.org/0000-0001-8334-8069АнтоноваО. Ю.AntonovaO. Yu.

Ольга Юрьевна Антонова, кандидат биологических наук, ведущий научный сотрудник, и.о. заведующего, лаборатория молекулярной селекции и ДНК-паспортизации, отдел биотехнологии, ВИР

190000 Россия, Санкт-Петербург, ул. Большая Морская, 42, 44

Olga Yu. Antonova, Cand. Sci. (Biology), Leading Researcher, Laboratory of Molecular Breeding and DNA-Genotyping, Department of Biotechnology, VIR

42, 44, Bolshaya Morskaya Street, St. Petersburg, 190000 Russia

olgaant326@mail.ruФедеральный исследовательский центр Всероссийский институт генетических ресурсов растений имени Н.И. ВавиловаN.I. Vavilov All-Russian Institute of Plant Genetic Resources202513112025822737Copyright © Гончаренко А.О., Багмет Л.В., Петрова М.Н., Антонова О.Ю., 20252025Гончаренко А.О., Багмет Л.В., Петрова М.Н., Антонова О.Ю.Goncharenko A.O., Bagmet L.V., Petrova M.N., Antonova O.Y.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://biosel.elpub.ru/jour/article/view/276

Актуальность. Груша (Pyrus L.) является одной из экономически важных плодовых культур, выращиваемой в 50 странах мира. Однако сорта груши поражаются многими патогенами, в частности паршой, возбудителем которой является гриб-аскомицет рода Venturia Sacc. На груше описаны два вида этого рода: Venturia nashicola S. Tanaka & S. Yamam., поражающий азиатские груши (P. pyrifolia (Burm. fil.) Nakai, P. bretschneideri Rehder, P. ussuriensis Maxim.) и Venturia pirina Aderh., специфичный для европейской груши P. communis L. Применение молекулярных маркеров для отбора устойчивых к парше сортов повысит эффективность программ селекции. Целью нашей работы была апробация маркеров генов Rvn2 и Vnk (Rvn1), контролирующих устойчивость к парше Venturia nashicola, на материале образцов коллекции Майкопской опытной станции – филиала ВИР и экспедиционных образцов. Материалы и методы: Изучена выборка из 255 образцов, включающая 246 сортов из коллекции Майкопской опытной станции – филиала ВИР, и девяти образцов, собранных экспедицией ВИР на Северном Кавказе в 2022 году. Основу выборки составили 152 сорта кавказской селекции, включая местные формы, вторую крупную подвыборку (61) образовали европейские сорта. Были использованы маркеры гена Rvn2 – PSC217/XhoI и PSC234/HaeIII, и гена Vnk (Rvn1) – STS-OPO9/SalI и STS-OPAW13, подобранные по данным литературы. Результаты: Показано широкое распространение среди образцов выборки обоих маркеров гена Rvn2 (89,4% для PSC217/XhoI и 30,9% для PSC234/HaeIII), при этом частота их встречаемости в двух основных подвыборках была примерно одинаковой. При сравнении результатов скрининга с данными по устойчивости образцов коллекции Майкопской ОС к парше груши показана низкая диагностическая ценность обоих маркеров – маркер PSC217/XhoI имел эффективность 47,2%, а маркер PSC234/HaeIII – 51,4%. Наоборот, маркеры STS-OPAW13 и STS-OPO9/SalI гена Vnk (Rvn1) присутствовали только у единичных сортов (семь) китайской и кавказской селекции. Последние, однако, согласно их родословным, были получены без использования местного оригинального материала. Заключение. В результате проведённого исследования значительной выборки образцов груши различного происхождения было показано широкое распространение в изученном материале маркеров гена Rvn2, которые, однако, продемонстрировали низкую эффективность и непригодны для молекулярного скрининга. Маркеры гена Vnk (Rvn1) были выявлены только у единичных образцов. Интерес для селекции представляет образец ‘Дан-Шансу-ли’, характеризующийся присутствием обоих маркеров гена Vnk (Rvn1) и, по предварительным данным, устойчивостью к парше груши.

Background: Pear (Pyrus sp.) is one of the economically important fruit crops grown in 50 countries worldwide. However, pear cultivars are affected by many pathogens, including scab caused by the ascomycete fungus belonging to the genus Venturia Sacc. The two pear-damaging species of this genus are Venturia nashicola S. Tanaka & S. Yamam. that affects Asian pears (P. pyrifolia (Burm.fil.) Nakai, P. bretschneideri Rehder, P. ussuriensis Maxim.), and Venturia pirina Aderh. that affects specifically the European pear P. communis L. The use of molecular markers for the selection of scab-resistant varieties will improve the efficiency of breeding programs. The aim of this work was to test the markers of the Rvn2 and Vnk (Rvn1) genes, which control scab resistance to Venturia nashicola, using the material from the collection maintained at the Maikop Experiment Station, and accessions from a collecting mission. Materials and methods: A sample of 255 accessions was studied, including 246 cultivars from the collection of the Maikop Experiment Station, and nine accessions from the VIR collecting mission to the North Caucasus in 2022. The basis of the sample were 152 cultivars of Caucasian origin, including local forms; the second large subsample (61) was made up of European cultivars. The research used the markers of the Rvn2 gene – PSC217/XhoI and PSC234/HaeIII, and of the Vnk (Rvn1) gene – STS-OPO9/SalI and STS-OPAW13, selected from the literature. Results: A wide distribution of both markers of the Rvn2 gene among the accessions was shown (89.4% for PSC217/XhoI and 30.9% for PSC234/HaeIII), while the frequency of their occurrence among the two main subsamples was approximately the same. When comparing the molecular screening results with the data on the accessions resistance to pear scab, a low diagnostic value of both markers was shown – the PSC217/XhoI marker had an efficiency of 47.2%, and that of the PSC234/HaeIII marker was 51.4%. On the contrary, the STS-OPAW13 and STS-OPO9/SalI markers of the Vnk (Rvn1) gene were present only in single cultivars (seven) bred in China and the Caucasus. However, according to their pedigrees, the latter ones were created without the use of original local material. Conclusion. The study of a large sample of pear accessions has shown a wide distribution of Rvn2 gene markers in the studied material, which, however, demonstrated low efficiency and are unsuitable for molecular screening. The Vnk (Rvn1) gene markers were detected only in few accessions. Of interest for breeding is the Chinese cultivar ‘Dan-Shansu-li’, which has both markers of the gene Vnk (Rvn1), and exhibits resistance to pear scab.

Pyrus L. sp.Venturia Saccгены устойчивостимолекулярные маркерыVnk (Rvn1)Rvn2Pyrus L. sp.Venturia Saccresistance genesmolecular markersVnk (Rvn1)Rvn2Работа выполнена в рамках Государственных заданий согласно тематическому плану ВИР по темам №1021032424343-9-4.4.4 FGEM-2022-0008 и №1021032424911-4-4.1.1; 4.4.4 FGEM-2022-0006.The research was carried out according to the State Assignment to VIR, Topics No. 1021032424343-9-4.4.4 FGEM-2022-0008 and No. 1021032424911-4-4.1.1; 4.4.4 FGEM-2022-0006.ReferencesAbe K., Kotobuki K., Sato T., Terai O. Inheritance of resistance to pear scab from European pears to Asian pears. Journal of the Japanese Society for Horticultural Science. 2000;1:1-8. [in Japanese]. DOI: 10.2503/jjshs.69.1Abe K., Kotobuki K., Sato T., Terai O. Inheritance of resistance to pear scab from European pears to Asian pears. Journal of the Japanese Society for Horticultural Science. 2000;1:1-8. [in Japanese]. DOI: 10.2503/jjshs.69.1Abe K., Saito T., Terai O., Sato Y., Kotobuki K. Genotypic difference for the susceptibility of Japanese, Chinese and European pears to Venturia nashicola, the cause of scab on Asian pears. Plant Breeding. 2008;127(4):407-412. DOI: 10.1111/j.1439-0523.2007.01482.xAbe K., Saito T., Terai O., Sato Y., Kotobuki K. Genotypic difference for the susceptibility of Japanese, Chinese and European pears to Venturia nashicola, the cause of scab on Asian pears. Plant Breeding. 2008;127(4):407-412. DOI: 10.1111/j.1439-0523.2007.01482.xАнтонова О.Ю., Клименко Н.С., Рыбаков Д.А., Фомина Н.А., Желтова В.В., Новикова Л.Ю., Гавриленко Т.А. SSR-анализ современных российских сортов картофеля с использованием ДНК номенклатурных стандартов. Биотехнология и селекция растений. 2020;3(4):77-96. DOI: 10.30901/2658-6266-2020-4-o2Antonova O.Yu., Klimenko N.S., Rybakov D.A., Fomina N.A., Zheltova V.V., Novikova L.Yu., Gavrilenko T.A. SSR analysis of modern Russian potato varieties using DNA samples of nomenclatural standards. Plant Biotechnology and Breeding. 2020;3(4):77-96. [in Russian]. DOI: 10.30901/2658-6266-2020-4-o2Bell R.L. Pears (Pyrus). Acta Horticulturae. 1991;290:657-697. DOI: 10.17660/ActaHortic.1991.290.15Bell R.L. Pears (Pyrus). Acta Horticulturae. 1991;290:657-697. DOI: 10.17660/ActaHortic.1991.290.15Bouvier L., Bourcy M., Boulay M., Tellier M., Guérif P., Denancé C., Durel C.E., Lespinasse Y. A new pear scab resistance gene Rvp1 from the European pear cultivar ‘Navara’ maps in a genomic region syntenic to an apple scab resistance gene cluster on linkage group 2. Tree Genetics & Genomes. 2012;8(1):53-60. DOI: 10.1007/s11295-011-0419-xBouvier L., Bourcy M., Boulay M., Tellier M., Guérif P., Denancé C., Durel C.E., Lespinasse Y. A new pear scab resistance gene Rvp1 from the European pear cultivar ‘Navara’ maps in a genomic region syntenic to an apple scab resistance gene cluster on linkage group 2. Tree Genetics & Genomes. 2012;8(1):53-60. DOI: 10.1007/s11295-011-0419-xBrewer L.R, Alspach P.A., Morgan C., Bus V.G.M. Resistance to scab caused by Venturia pirina in interspecific pear (Pyrus spp.) hybrids. New Zealand Journal of Crop and Horticultural Science. 2009;37(3):211-218. DOI: 10.1080/01140670909510266Brewer L.R, Alspach P.A., Morgan C., Bus V.G.M. Resistance to scab caused by Venturia pirina in interspecific pear (Pyrus spp.) hybrids. New Zealand Journal of Crop and Horticultural Science. 2009;37(3):211-218. DOI: 10.1080/01140670909510266Brown A.G. Scab resistance in progenies of varieties of the cultivated pear. Euphytica. 1960;9:247-253. DOI: 10.1007/BF00022230Brown A.G. Scab resistance in progenies of varieties of the cultivated pear. Euphytica. 1960;9:247-253. DOI: 10.1007/BF00022230Bus V.G.M., van de Weg W.E., Durel C.E., Gessler C., Calenge F., Parisi L., Rikkerink E., Gardiner S., Patocchi A., Meulenbroek E., Schouten H., Laurens F. Delineation of a scab resistance gene cluster on linkage group 2 of apple. Acta Horticulturae. 2004;663:57-62. DOI: 10.17660/ActaHortic.2004.663.3Bus V.G.M., van de Weg W.E., Durel C.E., Gessler C., Calenge F., Parisi L., Rikkerink E., Gardiner S., Patocchi A., Meulenbroek E., Schouten H., Laurens F. Delineation of a scab resistance gene cluster on linkage group 2 of apple. Acta Horticulturae. 2004;663:57-62. DOI: 10.17660/ActaHortic.2004.663.3Bus V.G.M., Laurens F.N.D., van de Weg W.E., Rusholme R.L., Rikkerink E.H.A., Gardiner S.E., Bassett H.C.M., Kodde L.P., Plummer K.M. The Vh8 locus of a new gene-for-gene interaction between Venturia inaequalis and the wild apple Malus sieversii is closely linked to the Vh2 locus in Malus pumila R12740-7A. New Phytologist. 2005a;166:1035-1049. DOI: 10.1111/j.1469-8137.2005. 01395.xBus V.G.M., Laurens F.N.D., van de Weg W.E., Rusholme R.L., Rikkerink E.H.A., Gardiner S.E., Bassett H.C.M., Kodde L.P., Plummer K.M. The Vh8 locus of a new gene-for-gene interaction between Venturia inaequalis and the wild apple Malus sieversii is closely linked to the Vh2 locus in Malus pumila R12740-7A. New Phytologist. 2005a;166:1035-1049. DOI: 10.1111/j.1469-8137.2005. 01395.xBus V.G.M., Rikkerink E.H.A., van de Weg W.E., Rusholme R.L., Gardiner S.E., Bassett H.C.M., Kodde L.P., Parisi L., Laurens F.N.D., Meulenbroek E.J., Plummer K.M. The Vh2 and Vh4 scab resistance genes in two differential hosts derived from Russian apple R12740-7A map to the same linkage group of apple. Molecular Breeding. 2005b;15:103-116. DOI: 10.1007/s11032-004-3609-5Bus V.G.M., Rikkerink E.H.A., van de Weg W.E., Rusholme R.L., Gardiner S.E., Bassett H.C.M., Kodde L.P., Parisi L., Laurens F.N.D., Meulenbroek E.J., Plummer K.M. The Vh2 and Vh4 scab resistance genes in two differential hosts derived from Russian apple R12740-7A map to the same linkage group of apple. Molecular Breeding. 2005b;15:103-116. DOI: 10.1007/s11032-004-3609-5Bus V.G.M., Rikkerink E.H.A., Caffier V., Durel C.-E., Plummer K.M. Revision of the nomenclature of the differential host-pathogen interactions of Venturia inaequalis and Malus. Annual Review of Phytopathology. 2011;49(1):391-413. DOI: 10.1146/annurev-phyto-072910-095339Bus V.G.M., Rikkerink E.H.A., Caffier V., Durel C.-E., Plummer K.M. Revision of the nomenclature of the differential host-pathogen interactions of Venturia inaequalis and Malus. Annual Review of Phytopathology. 2011;49(1):391-413. DOI: 10.1146/annurev-phyto-072910-095339Calenge F., Faure A., Goerre M., Gebhardt C., van de Weg W.E., Parisi L., Durel C.E. Quantitative trait loci (QTL) analysis reveals both broad-spectrum and isolate-specific QTL for scab resistance in an apple progeny challenged with eight isolates of Venturia inaequalis. Phytopathology. 2004;94:370-379. DOI: 10.1094/PHYTO.2004.94.4.370Calenge F., Faure A., Goerre M., Gebhardt C., van de Weg W.E., Parisi L., Durel C.E. Quantitative trait loci (QTL) analysis reveals both broad-spectrum and isolate-specific QTL for scab resistance in an apple progeny challenged with eight isolates of Venturia inaequalis. Phytopathology. 2004;94:370-379. DOI: 10.1094/PHYTO.2004.94.4.370Chevalier M., Tellier M., Lespinasse Y., Bruyninckx M. Behaviour studies of new strains of Venturia pirina isolated from ‘Conference’ cultivar on a range of pear cultivars. Acta Horticulturae. 2008;800:817-823. DOI: 10.17660/ActaHortic.2008.800.111Chevalier M., Tellier M., Lespinasse Y., Bruyninckx M. Behaviour studies of new strains of Venturia pirina isolated from ‘Conference’ cultivar on a range of pear cultivars. Acta Horticulturae. 2008;800:817-823. DOI: 10.17660/ActaHortic.2008.800.111Cho K.H., Shin I.S., Kim K.T., Suh E.J., Hong S.S., Lee H.J. Development of AFLP and CAPS markers linked to the scab resistance gene, Rvn2, in an inter-specific hybrid pear (Pyrus spp.). The Journal of Horticultural Science and Biotechnology. 2009;84(6): 619-624. DOI: 10.1080/14620316.2009.11512576Cho K.H., Shin I.S., Kim K.T., Suh E.J., Hong S.S., Lee H.J. Development of AFLP and CAPS markers linked to the scab resistance gene, Rvn2, in an inter-specific hybrid pear (Pyrus spp.). The Journal of Horticultural Science and Biotechnology. 2009;84(6): 619-624. DOI: 10.1080/14620316.2009.11512576Fisher R.A. On the interpretation of χ2 from contingency tables, and the calculation of P. Journal of the Royal Statistical Society. 1922;85(1):87-94. DOI: 10.2307/2340521Fisher R.A. On the interpretation of χ2 from contingency tables, and the calculation of P. Journal of the Royal Statistical Society. 1922;85(1):87-94. DOI: 10.2307/2340521Gygax M., Gianfranceschi L., Liebhard R., Kellerhals M., Gessler C., Patocchi A. Molecular markers linked to the apple scab resistance gene Vbj derived from Malus baccata jackii. Theoretical and Applied Genetics. 2004;109:1702-1709. DOI: 10.1007/s00122-004-1803-9Gygax M., Gianfranceschi L., Liebhard R., Kellerhals M., Gessler C., Patocchi A. Molecular markers linked to the apple scab resistance gene Vbj derived from Malus baccata jackii. Theoretical and Applied Genetics. 2004;109:1702-1709. DOI: 10.1007/s00122-004-1803-9Hemmat M, Brown S.K., Weeden N.F. Tagging and mapping scab resistance genes from R12740-7A apple. Journal of the American Society for Horticultural Science. 2002;127(3):365-370. DOI: 10.21273/JASHS.127.3.365Hemmat M, Brown S.K., Weeden N.F. Tagging and mapping scab resistance genes from R12740-7A apple. Journal of the American Society for Horticultural Science. 2002;127(3):365-370. DOI: 10.21273/JASHS.127.3.365Inglis P.W., Pappas M.D.C.R., Resende L.V., Grattapaglia D. Fast and inexpensive protocols for consistent extraction of high quality DNA and RNA from challenging plant and fungal samples for high-throughput SNP genotyping and sequencing applications. PLoS One. 2018;13(10):e0206085. DOI: 10.1371/journal.pone.0206085Inglis P.W., Pappas M.D.C.R., Resende L.V., Grattapaglia D. Fast and inexpensive protocols for consistent extraction of high quality DNA and RNA from challenging plant and fungal samples for high-throughput SNP genotyping and sequencing applications. PLoS One. 2018;13(10):e0206085. DOI: 10.1371/journal.pone.0206085Ishii H., Udagawa H., Nishimoto S., Tsuda T., Nakashima H. Scab resistance in pear species and cultivars. Acta Phytopathologica et Entomologica Hungarica. 1992;27:293-298.Ishii H., Udagawa H., Nishimoto S., Tsuda T., Nakashima H. Scab resistance in pear species and cultivars. Acta Phytopathologica et Entomologica Hungarica. 1992;27:293-298.Ishii H., Yanase H. Venturia nashicola, the scab fungus of Japanese and Chinese pears: a species distinct from V. pirina. Mycological Research. 2000;104(6):755-759. DOI: 10.1017/S0953756299001720Ishii H., Yanase H. Venturia nashicola, the scab fungus of Japanese and Chinese pears: a species distinct from V. pirina. Mycological Research. 2000;104(6):755-759. DOI: 10.1017/S0953756299001720Khajuria Y.P., Kaul S., Wani A.A., Dhar M.K. Genetics of resistance in apple against Venturia inaequalis (Wint.) Cke. Tree Genetics and Genomes. 2018;14:1-20. DOI: 10.1007/s11295-018-1226-4Khajuria Y.P., Kaul S., Wani A.A., Dhar M.K. Genetics of resistance in apple against Venturia inaequalis (Wint.) Cke. Tree Genetics and Genomes. 2018;14:1-20. DOI: 10.1007/s11295-018-1226-4[Каталог мировой коллекции ВИР]. Вып. 123. Каталог полевой устойчивости груши к основным заболеваниям / под ред. В.И. Кривченко. Ленинград; 1974.Krivchenko V.I. (ed.). [Catalogue of the VIR global collection]. Issue 123. Catalogue of field resistance of pear to major diseases (Katalog polevoy ustoychivosti grushi k osnovnym zabolevaniyam). Leningrad; 1974. [in Russian]Langford M.H., Keitt E.N. Heterothallism and variability in Venturia pirina. Phytopathology. 1942;32:357-369.Langford M.H., Keitt E.N. Heterothallism and variability in Venturia pirina. Phytopathology. 1942;32:357-369.Li J., Zhang M., Li X., Khan A., Kumar S., Allan A.C., Lin-Wang K., Espley R.V., Wang C., Wang R., Xue C., Yao G., Qin M., Sun M., Tegtmeier R., Liu H., Wei W., Ming M., Zhang S., Zhao K., Song B., Ni J., An J., Korban S.S., Wu J. Pear genetics: recent advances, new prospects, and a roadmap for the future. Horticulture Research. 2022;9:1-25. DOI: 10.1093/hr/uhab040Li J., Zhang M., Li X., Khan A., Kumar S., Allan A.C., Lin-Wang K., Espley R.V., Wang C., Wang R., Xue C., Yao G., Qin M., Sun M., Tegtmeier R., Liu H., Wei W., Ming M., Zhang S., Zhao K., Song B., Ni J., An J., Korban S.S., Wu J. Pear genetics: recent advances, new prospects, and a roadmap for the future. Horticulture Research. 2022;9:1-25. DOI: 10.1093/hr/uhab040Oh S., Han H., Kim D. A novel pear scab (Venturia nashicola) resistance gene, Rvn3, from interspecific hybrid pear (Pyrus pyrifolia × P. communis). Plants. 2021;10(12):2632. DOI: 10.3390/plants10122632Oh S., Han H., Kim D. A novel pear scab (Venturia nashicola) resistance gene, Rvn3, from interspecific hybrid pear (Pyrus pyrifolia × P. communis). Plants. 2021;10(12):2632. DOI: 10.3390/plants10122632Pierantoni L., Dondini L., Cho K.-H., Shin I.-S., Gennari F., Chiodini R., Tartarini S., Kang S.-J., Sansavini S. Pear scab resistance QTLs via a European pear (Pyrus communis) linkage map. Tree Genetics and Genomes. 2007;3:311-317. DOI: 10.1007/s11295-006-0070-0Pierantoni L., Dondini L., Cho K.-H., Shin I.-S., Gennari F., Chiodini R., Tartarini S., Kang S.-J., Sansavini S. Pear scab resistance QTLs via a European pear (Pyrus communis) linkage map. Tree Genetics and Genomes. 2007;3:311-317. DOI: 10.1007/s11295-006-0070-0Postman J.D., Spotts R.A., Calabro J. Scab resistance in Pyrus germplasm. Acta Horticulturae. 2005;671:601-608. DOI: 10.17660/ActaHortic.2005.671.84Postman J.D., Spotts R.A., Calabro J. Scab resistance in Pyrus germplasm. Acta Horticulturae. 2005;671:601-608. DOI: 10.17660/ActaHortic.2005.671.84Potter D., Eriksson T., Evans R.C., Oh S., Smedmark J.E.E., Morgan D.R., Kerr M., Robertson K.R., Arsenault M., Dickinson T.A., Campbell C.S. Phylogeny and classification of Rosaceae. Plant Systematics and Evolution. 2007;266:5-43. DOI: 10.1007/s00606-007-0539-9Potter D., Eriksson T., Evans R.C., Oh S., Smedmark J.E.E., Morgan D.R., Kerr M., Robertson K.R., Arsenault M., Dickinson T.A., Campbell C.S. Phylogeny and classification of Rosaceae. Plant Systematics and Evolution. 2007;266:5-43. DOI: 10.1007/s00606-007-0539-9Shabi E., Rotem J., Loebensten G. Physiological races of Venturia pirina on pear. Phytopathology. 1973;63:41-43. DOI: 10.1094/Phyto-63-41Shabi E., Rotem J., Loebensten G. Physiological races of Venturia pirina on pear. Phytopathology. 1973;63:41-43. DOI: 10.1094/Phyto-63-41Shabi E. Pear scab. In: A.L. Jones, H.S. Aldwinckle (eds). Compendium of Apple and Pear Diseases. St. Paul, Minnesota: APS Press; 1990. p.22-23.Shabi E. Pear scab. In: A.L. Jones, H.S. Aldwinckle (eds). Compendium of Apple and Pear Diseases. St. Paul, Minnesota: APS Press; 1990. p.22-23.Sokolova O., Moročko-Bičevska I., Bankina B. Review of pear scab caused by Venturia pyrina. Research for Rural Development. 2014;1:26-33. URL: https://lbtufb.lbtu.lv/conference/Research-for-Rural-Development/2014/LatviaResearchRuralDevel20th_volume1-26-33.pdf [дата обращения: 05.05.2025].Sokolova O., Moročko-Bičevska I., Bankina B. Review of pear scab caused by Venturia pyrina. Research for Rural Development. 2014;1:26-33. URL: https://lbtufb.lbtu.lv/conference/Research-for-Rural-Development/2014/LatviaResearchRuralDevel20th_volume1-26-33.pdf [дата обращения: 05.05.2025].Sokolova O., Moročko‐Bičevska I. Evaluation of Venturia pyrina virulence on European pear (Pyrus communis) cultivars by an in vitro methodology. Journal of Phytopathology. 2021:169(7-8):461-470. DOI: 10.1111/jph.13002Sokolova O., Moročko‐Bičevska I. Evaluation of Venturia pyrina virulence on European pear (Pyrus communis) cultivars by an in vitro methodology. Journal of Phytopathology. 2021:169(7-8):461-470. DOI: 10.1111/jph.13002Soufflet-Freslon V., Gianfranceschi L., Patocchi A., Dure C.E. Inheritance studies of apple scab resistance and identification of Rvi14, a new major gene that acts together with other broad-spectrum QTL. Genome. 2008;51:657-667. DOI: 10.1139/G08-046Soufflet-Freslon V., Gianfranceschi L., Patocchi A., Dure C.E. Inheritance studies of apple scab resistance and identification of Rvi14, a new major gene that acts together with other broad-spectrum QTL. Genome. 2008;51:657-667. DOI: 10.1139/G08-046Sugar D., Hilton R.J. Potential organic methods for management of pear scab. Acta Horticulturae. 2011;909:527-530. DOI: 10.17660/ActaHortic.2011.909.63Sugar D., Hilton R.J. Potential organic methods for management of pear scab. Acta Horticulturae. 2011;909:527-530. DOI: 10.17660/ActaHortic.2011.909.63Tanaka S., Yamamoto S. Studies in pear scab. II. Taxonomy of the causal fungus of Japanese pear scab. Annals of the Phytopathological Society of Japan. 1964;29(3):128-136. [in Japanese]. DOI: 10.3186/jjphytopath.29.128Tanaka S., Yamamoto S. Studies in pear scab. II. Taxonomy of the causal fungus of Japanese pear scab. Annals of the Phytopathological Society of Japan. 1964;29(3):128-136. [in Japanese]. DOI: 10.3186/jjphytopath.29.128Terakami S., Shoda M., Adachi Y., Gonai T., Kasumi M.,·Sawamura Y., Iketani H.,·Kotobuki K.,·Patocchi A.,·Gessler C.,·Hayashi T.,·Yamamoto T. Genetic mapping of the pear scab resistance gene Vnk of Japanese pear cultivar Kinchaku. Theoretical and Applied Genetics. 2006;113(4):743-752. DOI: 10.1007/s00122-006-0344-9Terakami S., Shoda M., Adachi Y., Gonai T., Kasumi M.,·Sawamura Y., Iketani H.,·Kotobuki K.,·Patocchi A.,·Gessler C.,·Hayashi T.,·Yamamoto T. Genetic mapping of the pear scab resistance gene Vnk of Japanese pear cultivar Kinchaku. Theoretical and Applied Genetics. 2006;113(4):743-752. DOI: 10.1007/s00122-006-0344-9Terakami S., Ogata N., Kita K., Gonai T., Saito T., Yamamoto T. Identification and genetic mapping of novel resistance gene, Rvn4, for pear scab in Chinese pear. Scientia Horticulturae. 2023;317:112032. DOI: 10.1016/j.scienta.2023.112032Terakami S., Ogata N., Kita K., Gonai T., Saito T., Yamamoto T. Identification and genetic mapping of novel resistance gene, Rvn4, for pear scab in Chinese pear. Scientia Horticulturae. 2023;317:112032. DOI: 10.1016/j.scienta.2023.112032Тхагушев Н.А. Садоводство адыгов: народные традиции, описание сортов, лесосады. Майкоп: Адыгейское республиканское книжное издательство; 2008.Thagushev N.A. Gardening of the Adyghe: folk traditions, description of varieties, forest gardens (Sadovodstvo adygov: narodnyie traditsii, opisaniye sortov, lesosady). Maykop: Adyghe Republican Book Publishing House; 2008. [in Russian]Verma L.R., Sharma R.C. Fungal and bacterial diseases of pear. In: L.R. Verma, R.C. Sharma. Diseases of horticultural crops: fruits. New Delhi: Indus Publishing company; 1999. p.140-166.Verma L.R., Sharma R.C. Fungal and bacterial diseases of pear. In: L.R. Verma, R.C. Sharma. Diseases of horticultural crops: fruits. New Delhi: Indus Publishing company; 1999. p.140-166.Won K., Bastiaanse H., Kim Y.K., Song J.H., Kang S.S., Lee H.C., Cho K.H., Brewer L., Singla G., Gardiner S.E., Chagne D., Bus V.G.M. Genetic mapping of polygenic scab (Venturia pirina) resistance in an interspecific pear family. Molecular Breeding. 2014;34(4):2179-2189. DOI: 10.1007/s11032-014-0172-6Won K., Bastiaanse H., Kim Y.K., Song J.H., Kang S.S., Lee H.C., Cho K.H., Brewer L., Singla G., Gardiner S.E., Chagne D., Bus V.G.M. Genetic mapping of polygenic scab (Venturia pirina) resistance in an interspecific pear family. Molecular Breeding. 2014;34(4):2179-2189. DOI: 10.1007/s11032-014-0172-6Won K., Choi E.D., Kim K., Shin I.S., Hohg S., Segonzac C., Sohn K.H., Deng C.H., Brewer L., Chagné D., Bus V.G.M. Genetic mapping of two quantitative resistance loci to Venturia nashicola in an interspecific pear family. Tree Genetics and Genomes. 2024;20(3):18. DOI: 10.1007/s11295-024-01650-0Won K., Choi E.D., Kim K., Shin I.S., Hohg S., Segonzac C., Sohn K.H., Deng C.H., Brewer L., Chagné D., Bus V.G.M. Genetic mapping of two quantitative resistance loci to Venturia nashicola in an interspecific pear family. Tree Genetics and Genomes. 2024;20(3):18. DOI: 10.1007/s11295-024-01650-0

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