phytosanitaryФитосанитария. Карантин растенийPlant Health and Quarantine2782-327X10.69536/FKR.2025.30.49.003phytosanitary-212Research ArticleДИАГНОСТИКАDIAGNOSTICSВироид рубцеватости плодов яблони – фитопатоген плодовых культурApple scar skin viroid – fruit crops phytopathogenhttps://orcid.org/0000-0001-9014-4179БашкироваИ. Г.BashkirovaI. G.

Башкирова Ида Геннадьевна, кандидат биологических наук, научный сотрудник научно-методического отдела вирусологии

пгт Быково, м. о. Раменский, Московская обл.

Ida Bashkirova, PhD in Biology, Researcher, Research and Methodology Department of Virology, Leading Researcher

Bykovo, Ramenskoye, Moscow Oblast

bashkirovaid@mail.ruhttps://orcid.org/0009-0004-9176-407XПриходькоЮ. Н.PrikhodkoYu. N.

Приходько Юрий Николаевич, кандидат сельскохозяйственных наук, ведущий научныйсотрудник научно-методического отдела вирусологии

пгт Быково, м. о. Раменский, Московская обл.

Yuri Prikhodko, PhD in Agriculture, Leading Researcher, Research and Methodology Department of Virology

Bykovo, Ramenskoye, Moscow Oblast

prihodko_yuri59@mail.ruhttps://orcid.org/0000-0002-7565-1241ШнейдерЮ. А.ShneyderYu A.

Шнейдер Юрий Андреевич, кандидат биологических наук, начальник научно-методического отдела вирусологии, ведущий научный сотрудник

пгт Быково, м. о. Раменский, Московская обл.

Yuri Shneyder, PhD in Biology, Head of Research and Methodology Department of Virology, Leading Researcher; Associate Professor of Plant Protection Department

Ramenskoye, Moscow Oblast

yury.shneyder@mail.ruhttps://orcid.org/0000-0001-8612-5127ЛозоваяЕ. Н.LozovayaE. N.

Лозовая Евгения Николаевна, научный сотрудник отдела аспирантуры

пгт Быково, м. о. Раменский, Московская обл.

Evgenya Lozovaya, Researcher, Postgraduate Study Department

Bykovo, Ramenskoye, Moscow Oblast

evgeniyaf@mail.ruЖиваеваТ. С.ZhivaevaT. S.

Живаева Татьяна Степановна, научный сотрудник научно-методического отдела вирусологии

пгт Быково, м. о. Раменский, Московская обл.

Tatyana Zhivaeva, Researcher, Research and Methodology Department of Virology and Bacteriology

Bykovo, Ramenskoye, Moscow Oblast

zhivaeva.vniikr@mail.ruhttps://orcid.org/0000-0001-7647-5799СелявкинС. Н.SelyavkinS. N.

Селявкин Сергей Николаевич, младший научный сотрудник Воронежского филиала ФГБУ «ВНИИКР»

г. Воронеж, Воронежская обл., 394042

Sergey Selyavkin, Junior Researcher

Voronezh

selyavkin91@mail.ruФГБУ «Всероссийский центр карантина растений» (ФГБУ «ВНИИКР»)РоссияAll-Russian Plant Quarantine Center (FGBU «VNIIKR»)Russian FederationФГБУ «Всероссийский центр карантина растений» (ФГБУ «ВНИИКР»); ФГБОУ ВО «Российский государственный аграрный университет – МСХА имени К. А. Тимирязева» (ФГБОУ ВО РГАУ-МСХА им. К. А. Тимирязева)РоссияAll-Russian Plant Quarantine Center (FGBU «VNIIKR»); Russian State Agrarian University – Moscow Timiryazev Agricultural AcademyRussian FederationВоронежский филиал ФГБУ «Всероссийский центр карантина растений» (Воронежский филиал ФГБУ «ВНИИКР»)РоссияVoronezh Branch of FGBU "VNIIKR", Voronezh, Voronezh OblastRussian Federation202524112025042337Copyright © Башкирова И.Г., Приходько Ю.Н., Шнейдер Ю.А., Лозовая Е.Н., Живаева Т.С., Селявкин С.Н., 20252025Башкирова И.Г., Приходько Ю.Н., Шнейдер Ю.А., Лозовая Е.Н., Живаева Т.С., Селявкин С.Н.Bashkirova I.G., Prikhodko Y.N., Shneyder Y.A., Lozovaya E.N., Zhivaeva T.S., Selyavkin S.N.Данная работа распространяется под лицензией Creative Commons Attribution 4.0.This work is licensed under a Creative Commons Attribution 4.0 License.https://phytosanitary.vniikr.ru/jour/article/view/212

Вироид рубцеватости плодов яблони (Apscaviroid cicatricimali, Apple scar skin viroid, ASSVd) является одним из особо опасных фитопа- тогенов, поражающих семечковые и косточковые плодовые культуры. Вироидная инфекция представляет потенциальную опасность, которая может нанести экономический ущерб производству плодовых культур. Наиболее распространенным способом передачи ASSVd является контактная передача (механически через рабочие инструменты), а также через прививку (от зараженного растения к здоровому). В статье приведена краткая обзорная информация по вироиду, а также показаны данные по апробации олигонуклеотидов для диагностики ASSVd методом ПЦР в режиме реального времени (ПЦР-РВ) с использованием отечественных реактивов. Проверку работы олигонуклеотидов осуществляли с помощью отечественных наборов реагентов компании ООО «НПФ Синтол» и ЗАО «Евроген». Для проведения экспериментов по оценке специфичности праймеров использовали различные изоляты близкородственных вироидов из семейств Avsunviroidae и Pospiviroidae. В данной работе проведен анализ олигонуклеотидов с использованием онлайн-сервиса NCBI Primer-BLAST. В исследовании проведено множественное выравнивание нук леотидных последовательностей изолятов вироида рубцева- тости плодов яблони с помощью специальных программ MAFFT и AliView. Биоинформатически проанализированы видоспецифичные олигонуклеотиды на возможность образования вторичных структур – шпилек и димеров, которые могут повлиять на диагностику фитопатогена. Установлено, что праймеры cASSVd/hASSVd и ASS-F/ASS-R способны образовывать большое количество вторичных структур, которые могут привести к ложноположительному результату. С помощью праймеров ASSVd-F91/ASSVd-R291, ASS-F/ASS-R и зонда ASS-Р возможно идентифицировать вироид рубцеватости плодов яблони методом ПЦР-РВ

Apple scar skin viroid, ASSVd (Apscaviroid cicatricimali) is a particularly dange rous phytopathogen affecting pome and stone fruit crops. Viroid infection poses a potential threat that can cause economic damage to fruit production. The most common pathways of ASSVd are contact (mechanically through tools) and grafting (from an infected plant to a healthy one). This article provides a brief overview of the viroid and presents data on the validation of oligonucleotides for ASSVd diagnostics with real-time PCR using Russian reagents. Oligonucleotide performance was tested using Russian Syntol and Evrogen reagent kit. Various isolates of closely related viroids from the Avsunviroidae and Pospiviroidae families were used to conduct experiments assessing primer specificity. In this study, oligonucleotide analysis was performed using the NCBI Primer-BLAST online service. Multiple nucleotide sequence alignments of apple fruit scarring viroid isolates were performed using specialized MAFFT and AliView software. Speciesspecific oligonucleotides were bioinformatically analyzed for the formation of secondary structures— hairpins and dimers—that may affect the diagnosis of the phytopathogen. It was found that the cASSVd/hASSVd and ASS-F/ASS-R primers are capable of forming a large number of secondary structures, which can lead to false-positive results. Using the primers ASSVd-F91/ASSVd-R291, ASSF/ ASS-R and the probe ASS-P, it is possible to identify the apple fruit scarring viroid using the real-time PCR method.

Apscaviroid cicatricimaliASSVdсемечковые культурыкосточковые культурыкарантин растенийApscaviroid cicatricimaliASSVdpome cropsstone fruit cropsplant quarantineИсследования выполнены в рамках государственного задания (рег. № ЕГИСУ НИОКТР 125031103530-8).The research was carried out within the framework of the State assignment (registration number EGISU NIOKTR 125031103530-8).ReferencesВироид латентной мозаики персика – опасный патоген плодовых культур / Приходько Ю.Н., Живаева Т.С., Лозовая Е.Н., Шнейдер Ю.А., Каримова Е.В. // Фитосанитария. Карантин растений. 2021. № 2. С. 20–26.Prikhodko Yu.N., Zhivaeva T.S., Lozovaya E.N., Shneyder Yu.A., Karimova E.V. Peach latent mosaic viroid – dangerous pathogen of fruit crops. Plant Health and Quarantine. 2021; (2): 20-26.Bae Y.S. Development of multiplex RT-PCR for apple viruses and viroid and the incidence of apple viral disease in Gyeongsangbuk-do // Master's thesis. Kyungpook National University, Daegu, Korea. 2015.Bae Y.S. Development of multiplex RT-PCR for apple viruses and viroid and the incidence of apple viral disease in Gyeongsangbuk-do // Master's thesis. Kyungpook National University, Daegu, Korea. 2015.Bak S., Seo E., Kim S.Y., Park W.H., Lee S.H. First report Apricot pseudo-chlorotic leaf spot virus infecting peach trees in South Korea // Research in Plant Disease. 2017. Т. 23. №. 1. P. 75–81.Bak S., Seo E., Kim S.Y., Park W.H., Lee S.H. First report Apricot pseudo-chlorotic leaf spot virus infecting peach trees in South Korea // Research in Plant Disease. 2017. Т. 23. №. 1. P. 75–81.Desvignes J.C., Grasseau N., Boye R., Cornaggia D., Aparicio F., Di Serio F., Flores R. Biological properties of apple scar skin viroid: isolates, host range, different sensitivity of apple cultivars, elimination, and natural transmission // Plant Disease. 1999. Т. 83. №. 8. P. 768–772.Desvignes J.C., Grasseau N., Boye R., Cornaggia D., Aparicio F., Di Serio F., Flores R. Biological properties of apple scar skin viroid: isolates, host range, different sensitivity of apple cultivars, elimination, and natural transmission // Plant Disease. 1999. Т. 83. №. 8. P. 768–772.Di Serio F., Ambrós S., Sano T., Flores R., Navarro B. Viroid diseases in pome and stone fruit trees and Koch's postulates: a critical assessment // Viruses. 2018. Т. 10. №. 11. P. 612.Di Serio F., Ambrós S., Sano T., Flores R., Navarro B. Viroid diseases in pome and stone fruit trees and Koch's postulates: a critical assessment // Viruses. 2018. Т. 10. №. 11. P. 612.Di Serio F., Malfitano M., Alioto D., Ragozzino A., Flores R. Apple dimple fruit viroid: sequence variability and its specific detection by multiplex fluorescent RT-PCR in the presence of Apple scar skin viroid // Journal of Plant Pathology. 2002. P. 27–34.Di Serio F., Malfitano M., Alioto D., Ragozzino A., Flores R. Apple dimple fruit viroid: sequence variability and its specific detection by multiplex fluorescent RT-PCR in the presence of Apple scar skin viroid // Journal of Plant Pathology. 2002. P. 27–34.Diener T.O. Potato spindle tuber “virus”: IV. A replicating, low molecular weight RNA // Virology. 1971. Т. 45. №. 2. P. 411–428.Diener T.O. Potato spindle tuber “virus”: IV. A replicating, low molecular weight RNA // Virology. 1971. Т. 45. №. 2. P. 411–428.Diener T.O. The viroid: biological oddity or evolutionary fossil? // Advances in Virus Research. 2001. Vol. 57. P. 137–184.Diener T.O. The viroid: biological oddity or evolutionary fossil? // Advances in Virus Research. 2001. Vol. 57. P. 137–184.Hadidi A., Barba M., Hong N., Hallan V. Apple scar skin viroid // Viroids and satellites. Academic press. 2017. P. 217–228.Hadidi A., Barba M., Hong N., Hallan V. Apple scar skin viroid // Viroids and satellites. Academic press. 2017. P. 217–228.Hashimoto J., Koganezawa H. Nucleotide sequence and secondary structure of apple scar skin viroid // Nucleic Acids Research. 1987. Т. 15. №. 17. P. 7045–7052.Hashimoto J., Koganezawa H. Nucleotide sequence and secondary structure of apple scar skin viroid // Nucleic Acids Research. 1987. Т. 15. №. 17. P. 7045–7052.Heo S., Kim H.R., Lee H.J. Development of a quantitative real-time nucleic acid sequence based amplification (NASBA) assay for early detection of apple scar skin viroid // The plant pathology journal. 2019. Т. 35. №. 2. P. 164.Heo S., Kim H.R., Lee H.J. Development of a quantitative real-time nucleic acid sequence based amplification (NASBA) assay for early detection of apple scar skin viroid // The plant pathology journal. 2019. Т. 35. №. 2. P. 164.Hurtt S.S., Podleckis E.V. Apple scar skin viroid is not seed transmitted or transmitted at a low rate in oriental pear // Acta Horticulturae. 1995. № 386. P. 544–550.Hurtt S.S., Podleckis E.V. Apple scar skin viroid is not seed transmitted or transmitted at a low rate in oriental pear // Acta Horticulturae. 1995. № 386. P. 544–550.Kaponi M., Kyriakopoulou P.E., Hadidi A. Viroids of the mediterranean basin // Viruses. 2024. Т. 16. №. 4. P. 612.Kaponi M., Kyriakopoulou P.E., Hadidi A. Viroids of the mediterranean basin // Viruses. 2024. Т. 16. №. 4. P. 612.Kaponi M.S., Sano T., Kyriakopoulou P.E. Natural infection of sweet cherry trees with apple scar skin viroid // Journal of Plant Pathology. 2013. P. 429–433.Kaponi M.S., Sano T., Kyriakopoulou P.E. Natural infection of sweet cherry trees with apple scar skin viroid // Journal of Plant Pathology. 2013. P. 429–433.Kim D.H., Kim H.R., Heo S., Kim S.H., Kim M.A., Shin I.S., Kim J.H., Cho K.H., Hwang J.H. Occurrence of apple scar skin viroid and relative quantity analysis using real-time RT-PCR // Research in Plant Disease. 2010. Т. 16. №. 3. P. 247–253.Kim D.H., Kim H.R., Heo S., Kim S.H., Kim M.A., Shin I.S., Kim J.H., Cho K.H., Hwang J.H. Occurrence of apple scar skin viroid and relative quantity analysis using real-time RT-PCR // Research in Plant Disease. 2010. Т. 16. №. 3. P. 247–253.Kim H.R., Lee S.H., Lee D.H., Kim J.S., Park J.W. Transmission of Apple scar skin viroid by grafting, using contaminated pruning equipment, and planting infected seeds // The Plant Pathology Journal. 2006. Т. 22. №. 1. P. 63–67.Kim H.R., Lee S.H., Lee D.H., Kim J.S., Park J.W. Transmission of Apple scar skin viroid by grafting, using contaminated pruning equipment, and planting infected seeds // The Plant Pathology Journal. 2006. Т. 22. №. 1. P. 63–67.Koganezawa H. Transmission to apple seedlings of a low molecular weight RNA extracted from apple scar skin diseased trees // Japanese Journal of Phytopathology. 1985. Т. 51. №. 2. P. 176–182.Koganezawa H. Transmission to apple seedlings of a low molecular weight RNA extracted from apple scar skin diseased trees // Japanese Journal of Phytopathology. 1985. Т. 51. №. 2. P. 176–182.Kumar S., Singh R.M., Ram R., Badyal J., Hallan V., Zaidi A.A., Varma A. Determination of major viral and sub viral pathogens incidence in apple orchards in Himachal Pradesh // Indian Journal of Virology. 2012. Т. 23. №. 1. P. 75–79.Kumar S., Singh R.M., Ram R., Badyal J., Hallan V., Zaidi A.A., Varma A. Determination of major viral and sub viral pathogens incidence in apple orchards in Himachal Pradesh // Indian Journal of Virology. 2012. Т. 23. №. 1. P. 75–79.Kwon M.J., Hwang S.L., Lee S.J., Lee D.H., Lee J.Y. Detection and distribution of apple scar skin viroid-Korean strain (ASSVd-K) from apples cultivated in Korea // The Plant Pathology Journal. 2002. Vol. 18 (6). P. 342–344.Kwon M.J., Hwang S.L., Lee S.J., Lee D.H., Lee J.Y. Detection and distribution of apple scar skin viroid-Korean strain (ASSVd-K) from apples cultivated in Korea // The Plant Pathology Journal. 2002. Vol. 18 (6). P. 342–344.Kyriakopoulou P.E., Giunchedi L., Hadidi A. Peach latent mosaic and pome fruit viroids in naturally infected cultivated pear Pyrus communis and wild pear P. amygdaliformis: implications on possible origin of these viroids in the Mediterranean region // Journal of Plant Pathology. 2001. Vol. 83 (1). P. 51–62.Kyriakopoulou P.E., Giunchedi L., Hadidi A. Peach latent mosaic and pome fruit viroids in naturally infected cultivated pear Pyrus communis and wild pear P. amygdaliformis: implications on possible origin of these viroids in the Mediterranean region // Journal of Plant Pathology. 2001. Vol. 83 (1). P. 51–62.Kyriakopoulou P.E., Hadidi A. Natural infection of wild and cultivated pears with Apple scar skin viroid in Greece // XVII International Symposium Virus and Virus-Like Diseases of Temperate Fruit Crops 472. 1997. P. 617–626.Kyriakopoulou P.E., Hadidi A. Natural infection of wild and cultivated pears with Apple scar skin viroid in Greece // XVII International Symposium Virus and Virus-Like Diseases of Temperate Fruit Crops 472. 1997. P. 617–626.Lee J.H., Park J.K., Lee D.H., Uhm J.Y., Ghim S.Y., Lee J.Y. Occurrence of apple scar skin viroid-Korean strain (ASSVd-K) in apples cultivated in Korea // The Plant Pathology Journal. 2001. Т. 17. №. 5. P. 300–304.Lee J.H., Park J.K., Lee D.H., Uhm J.Y., Ghim S.Y., Lee J.Y. Occurrence of apple scar skin viroid-Korean strain (ASSVd-K) in apples cultivated in Korea // The Plant Pathology Journal. 2001. Т. 17. №. 5. P. 300–304.Li G., Li J., Zhang H., Li J., Jia L., Zhou S., Wang Y., Sun J., Tan M., Shao J. ASSVd infection inhibits the vegetative growth of apple trees by affecting leaf metabolism // Frontiers in Plant Science. 2023. Т. 14. P. 1137630.Li G., Li J., Zhang H., Li J., Jia L., Zhou S., Wang Y., Sun J., Tan M., Shao J. ASSVd infection inhibits the vegetative growth of apple trees by affecting leaf metabolism // Frontiers in Plant Science. 2023. Т. 14. P. 1137630.Liu F.C., Chen R.F., Chen Y.X. Apple Scar Skin Disease // Academica Sinica Printery; Beijing, China. 1957. P. 43.Liu F.C., Chen R.F., Chen Y.X. Apple Scar Skin Disease // Academica Sinica Printery; Beijing, China. 1957. P. 43.Liu F.C., Wang S.Y., Tian C.F., Chen R.F. Studies on apple scar skin disease // Annual Reports of Fruit Institute, Chinese Academy of Agriculture Science. 1962. P. 71–79.Liu F.C., Wang S.Y., Tian C.F., Chen R.F. Studies on apple scar skin disease // Annual Reports of Fruit Institute, Chinese Academy of Agriculture Science. 1962. P. 71–79.Lodh S. Elucidating the fate of Apple scar skin viroid RNA in animal cells: Diss. // CSKHPKV, Palampur. 2017. P. 113.Lodh S. Elucidating the fate of Apple scar skin viroid RNA in animal cells: Diss. // CSKHPKV, Palampur. 2017. P. 113.Malandraki I., Var veri C., Olmos A ., Vassilakos N. One-step multiplex quantitative RT-PCR for the simultaneous detection of viroids and phytoplasmas of pome fruit trees // Journal of Virological Methods. 2015. Vol. 213. 12–17.Malandraki I., Var veri C., Olmos A ., Vassilakos N. One-step multiplex quantitative RT-PCR for the simultaneous detection of viroids and phytoplasmas of pome fruit trees // Journal of Virological Methods. 2015. Vol. 213. 12–17.Ohtsuka Y. A new diseased of apple, on the abnormality of fruit // Journal of Japan Society Horticultural Scienc (in Japanese). 1935. Т. 6. P. 44–53.Ohtsuka Y. A new diseased of apple, on the abnormality of fruit // Journal of Japan Society Horticultural Scienc (in Japanese). 1935. Т. 6. P. 44–53.Ohtsuka Y. On Manshu-sabika-byo of apple, graft transmission and symptom variation in cultivars // Journal of Japan Society Horticultural Science. 1938. Т. 9. P. 282–286.Ohtsuka Y. On Manshu-sabika-byo of apple, graft transmission and symptom variation in cultivars // Journal of Japan Society Horticultural Science. 1938. Т. 9. P. 282–286.Shamloul A.M., Yang X., Han L., Hadidi A. Characterization of a new variant of Apple scar skin viroid associated with pear fruit crinkle disease // Journal of Plant Pathology. 2004. P. 249–256.Shamloul A.M., Yang X., Han L., Hadidi A. Characterization of a new variant of Apple scar skin viroid associated with pear fruit crinkle disease // Journal of Plant Pathology. 2004. P. 249–256.Sipahioglu H.M., Usta M., Ocak M. Development of a rapid enzymatic cDNA amplification test for the detection of Apple Scar Skin Viroid (ASSVd) in apple trees from eastern Anatolia, Turkey // Archives of Phytopathology and Plant Protection. 2009. Т. 42. №. 4. P. 352–360.Sipahioglu H.M., Usta M., Ocak M. Development of a rapid enzymatic cDNA amplification test for the detection of Apple Scar Skin Viroid (ASSVd) in apple trees from eastern Anatolia, Turkey // Archives of Phytopathology and Plant Protection. 2009. Т. 42. №. 4. P. 352–360.Takahashi T. Plant viroid diseases occurring in Japan // Japan Agricultural Research Quarterly. 1987. Т. 21. №. 3. P. 184–191.Takahashi T. Plant viroid diseases occurring in Japan // Japan Agricultural Research Quarterly. 1987. Т. 21. №. 3. P. 184–191.Tian M., Wei S., Bian R., Luo J., Khan H.A., Tai H., Sun L. Natural cross-kingdom spread of apple scar skin viroid from apple trees to fungi // Cells. 2022. Т. 11. №. 22. P. 3686.Tian M., Wei S., Bian R., Luo J., Khan H.A., Tai H., Sun L. Natural cross-kingdom spread of apple scar skin viroid from apple trees to fungi // Cells. 2022. Т. 11. №. 22. P. 3686.Walia Y., Dhir S., Bhadoria S., Hallan V., Zaidi A.A. Molecular characterization of Apple scar skin viroid from Himalayan wild cherry // Forest Pathology. 2012. Т. 42. №. 1. P. 84–87.Walia Y., Dhir S., Bhadoria S., Hallan V., Zaidi A.A. Molecular characterization of Apple scar skin viroid from Himalayan wild cherry // Forest Pathology. 2012. Т. 42. №. 1. P. 84–87.Walia Y., Dhir S., Ram R., Zaidi A.A., Hallan V. Identification of the herbaceous host range of Apple scar skin viroid and analysis of its progeny variants // Plant pathology. 2014. Т. 63. №. 3. P. 684–690.Walia Y., Dhir S., Ram R., Zaidi A.A., Hallan V. Identification of the herbaceous host range of Apple scar skin viroid and analysis of its progeny variants // Plant pathology. 2014. Т. 63. №. 3. P. 684–690.Walia Y., Dhir S., Zaidi A.A., Hallan V. Apple scar skin viroid naked RNA is actively transmitted by the whitefly Trialeurodes vaporariorum // RNA Biology. 2015. Т. 12. №. 10. P. 1131–1138.Walia Y., Dhir S., Zaidi A.A., Hallan V. Apple scar skin viroid naked RNA is actively transmitted by the whitefly Trialeurodes vaporariorum // RNA Biology. 2015. Т. 12. №. 10. P. 1131–1138.Yazarlou A., Jafarpour B., Habili N., Randles J.W. First detection and molecular characterization of new Apple scar skin viroid variants from apple and pear in Iran //Australasian Plant Disease Notes. 2012. Т. 7. №. 1. P. 99–102.Yazarlou A., Jafarpour B., Habili N., Randles J.W. First detection and molecular characterization of new Apple scar skin viroid variants from apple and pear in Iran //Australasian Plant Disease Notes. 2012. Т. 7. №. 1. P. 99–102.CABI, 2025. Apple scar skin viroid (apple dimple) [Электронный ресурс]. URL: https://plantwiseplusknowledgebank.org/doi/10.1079/PWKB.Species.6086 (дата обращения 18.08.2025).CABI, 2025. Apple scar skin viroid (apple dimple) [Electronic resource]. URL: https://plantwiseplusknowledgebank.org/doi/10.1079/PWKB.Species.6086 (last accessed 18.08.2025).CABI/EPPO, 2025. Apple scar skin viroid [Электронныйресурс]. URL: https://www.cabidigitallibrary.org/doi/10.1079/DMP D/20123367481 (дата обращения 27.07.2025).CABI/EPPO, 2025. Apple scar skin viroid [Electronic resource]. URL: https://www.cabidigitallibrary.org/doi/10.1079/DMP D/20123367481 (last accessed 27.07.2025).EPPO Global Database, 2025. Apscaviroid cicatricimali [Электронный ресурс]. URL: https://gd.eppo.int/taxon/ASSVD0 (дата обращения: 27.07.2025).EPPO Global Database, 2025. Apscaviroid cicatricimali [Electronic resource] . URL: https://gd.eppo.int/taxon/ASSVD0 (last accessed: 27.07.2025).NCBI Primer-BLAST [Электронный ресурс]. URL: https: / / www.ncbi.nlm.nih.gov/tools/primer-blast/ (дата обращения: 08.08.2025).NCBI Primer-BLAST [Electronic resource]. URL: https://www.ncbi.nlm.nih.gov/tools/primerblast/(last accessed: 08.08.2025).Oligo Calculator, 2025. Oligonucleotide Properties Calculator Электронный ресурс]. URL: http://oligocalc.eu/ (дата обращения: 08.08.2025).Oligo Calculator, 2025. Oligonucleotide Properties Calculator [Electronic resource]. URL: http://oligocalc.eu/ (last accessed: 08.08.2025).Thermo Fisher Scientific Inc., 2025. Multiple Primer Analyzer [Электронный ресурс]. URL: https://www.thermofisher.com/ru/ru/home/brands/thermo-scientific/molecular-biology/molecularbiology-learning-center/molecular-biologyr esource-library/thermo-scientific-web-tools/multiple-primer-analyzer.html (дата обращения: 08.08.2025).Thermo Fisher Scientific Inc., 2025. Multiple Primer Analyzer [Electronic resource]. URL: https://www.thermofisher.com/ru/ru/home/brands/thermo-scientific/molecular-biology/molecularbiology-learning-center/molecular-biologyre source-library/thermo-scientific-web-tools/multiple-primer-analyzer.html (last accessed: 08.08.2025).

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