Молекулярные маркеры в исследованиях генетического разнообразия представителей рода Rubus L. и перспективы их применения в селекции

Род RubusL. (семейство RosaceaeJuss.), по оценкам разных систематиков, состоит из 12-16 подродов, объединяющих ~750 видов. Самые крупные по числу видов подроды - Idaeobatus (Focke) Focke, к которому относятся малины, и типовой подрод Rubus (=EubatusFocke), включающий виды ежевик. Представители рода Rubus обладают высокой пищевой и хозяйственной ценностью, а также лекарственными свойствами. Селекционные исследования направлены на расширение генетического разнообразия и создание новых сортов малин и ежевик, устойчивых к биотическим и абиотическим стрессорам и отличающихся высоким качеством плодов. Современные селекционно-генетические программы все шире включают достижения молекулярной генетики и геномики. В данной статье представлен обзор фундаментальных и прикладных исследований генетического разнообразия культивируемых и дикорастущих видов рода Rubus, выполненных на основе методов молекулярного маркирования. Рассмотрены основные типы молекулярных маркеров (RFLP, RAPD, SSR, ISSR, AFLP, SCAR, SSCP, ретротранспозонные маркеры и т. д.) и области их применения в изучении представителей рода Rubus. Приведены результаты работ по использованию методов ДНК-маркирования для решения самых разных задач, включая: исследование межвидового и внутривидового генетического разнообразия, филогенетических связей видов и надвидовых таксонов, выяснение спорных вопросов систематики, генотипирование и уточнение родословных сортов малин и ежевик, изучение сомаклональной изменчивости и др. Наиболее важным результатом в практическом плане является создание насыщенных молекулярно-генетических карт для разных видов малин и ежевик, на которых локализованы многочисленные гены и QTL, детерминирующие различные хозяйственно ценные признаки. В то же время необходимо отметить, что число маркеров, перспективных для проведения эффективного молекулярного скрининга, пока еще недостаточно.

1. Бологовская Р.П. Малины и ежевики. В: Вульф Е.В. (ред.). Культурная флора СССР. Т. 16. М.; СПб., 1936;165-226.

2. Дунаева С.Е., Кудрякова Н.В., Малышев Л.Л., Лупышева Ю.В., Гавриленко Т.А. In vitro коллекция малин и ежевик и идентификация образцов по изоферментным спектрам. Аграрная Россия. 2005;2:49-55.

3. Евдокименко С.Н., Кулагина В.Л., Якуб И.Я. Современные тенденции производства и селекции малины. Плодоводство и ягодоводство России. 2012;31(1):148-156.

4. Каган Д.И., Шестибратов К.А., Лебедев В.Г., Азарова А.Б., Филиппов М.С., Бесов С.А., Ивановская С.И., Ковалевич О.А., Барсукова М.М. Паспортизация сортов малины и ежевики и изучение их филогенетических взаимоотношений методом RAPD-анализа. В: Биотехнологические приемы в сохранении биоразнообразия и селекции растений: Сб. статей междунар. науч. конф. Минск, 2014;101-104.

5. Красовская Л.С. Рубус - Rubus L. В: Цвелёв Н.Н. (отв. ред.). Флора Восточной Европы. Т. X. СПб., 2001;362-393.

6. Лебедев В.Г., Субботина Н.М., Киркач В.В., Видягина Е.О., Поздняков И.А., Шестибратов К.А. Анализ микросателлитных локусов как первый этап на пути к маркерной селекции малины и земляники. Селекция и сорторазведение садовых культур. 2018; 5(1):65-68.

7. Овчинникова С.В. Семейство Rosaceae. В: Байков К.С. (ред.). Конспект флоры Азиатской России. Новосибирск, 2012:199-266.

8. Соболев В.В., Соболева А.Г., Андреева Г.Н., Карлов Г.И. Оценка межвидового и межсортового полиморфизма малины и маркирование признака ремонтантности с использованием ISSR-ПЦР-анализа. Изв. ТСХА. 2009;2:103-109.

9. Шнеер В.С., Родионов А.В. ДНК-штрихкоды растений. Успехи соврем. биологии. 2018;138(6):531-537.

10. Agar G., Halasz J., Ercisli S. Genetic relationships among wild and cultivated blackberries (Rubus caucasicus L.) based on amplified fragment length polymorphism markers. Plant Biosyst. 2011;145: 347-352.

11. Alice L.A., Campbell C.S. Phylogeny of Rubus (Rosaceae) based on nuclear ribosomal DNA internal transcribed spacer region sequen¬ces. Am. J. Bot. 1999;86(1):81-97.

12. Amsellem L., Noyer J., Le Bourgeois T., Hossart-McKey M. Com¬parison of genetic diversity of the invasive weed R. alceifolius Poir. (Rosaceae) in its native range in areas of introduction, using amplified fragment lenghth polymorphism markers. Mol. Ecol. 2000;9: 443-455.

13. Badjakov I., Todorovska E., Boicheva I., Atanassov I., Atanassov A. Assessment of genetic diversity in Bulgarian raspberry germplasm collection by microsatellite markers (SSR). Biotechnol. Biotechnol. Equip. 2009;19(1):43-47.

14. Bassil N.V., Nyberg A.M., Finn C.E., Clark J.R., Peace C.P., Iezzoni A. Development of a multiplexed fingerprinting set in blackberry. Acta Hortic. 2016;1133:89-96. https://doi.org/10.17660/ActaHortic.2016.1133.14.

15. Bushakra J.M., Bryant D.M., Dossett M., Vining K.J., VanBuren R., Gilmore B.S., Lee J., Mockler T.C., Finn C.E., Bassil N.V. A genetic linkage map of black raspberry (Rubus occidentalis) and the map¬ping of Ag4 conferring resistance to the aphid Amphorophora agathonica. Theor. Appl. Genet. 2015a;128:1631-1646. https://doi.org/10.1007/s00122-015-2541-x.

16. Bushakra J.M., Dossett M., Carter K.A., Vining K.J., Lee J.C., Bry¬ant D.W., VanBuren R., Lee J., Mockler T.C., Finn C.E., Bassil N.V. Characterization of aphid resistance loci in black raspberry (Rubus occidentalis L.). Mol. Breed. 2018;38:83. https://doi.org/10.1007/s11032-018-0839-5.

17. Bushakra J.M., Lewers K.S., Staton M.E., Zhebentyayeva T., Saski C.A. Developing expressed sequence tag libraries and the discovery of simple sequence repeat markers for two species of raspberry (Rubus L.) BMC Plant Biol. 2015b;15:258-269. https://doi.org/10.1007/s00122-015-2541-x.

18. Bushakra J.M., Stephens M.J., Atmadjaja A.N., Lewers K.S., Symonds V.V., Udall J.A., Chagne D., Buck E.J., Gardiner S.E. Construction of black (Rubus occidentalis) and red (R. idaeus) raspberry linkage maps and their comparison to the genomes of strawberry, apple, and peach. Theor. Appl. Genet. 2012;125(2):311-327. https://doi.org/10.1007/s00122-012-1835-5.

19. Bussemeyer D.T., Pelikan S., Kennedy R.S., Rogstad S.H. Genetic diversity of Philippine Rubus moluccanus L. (Rosaceae) populations examined with VNTR DNA probes. J. Trop. Biol. 1997;14:867-884. https://doi.org/10.1017/S0266467400011044.

20. Castillo N.R.F., Bassil N.V., Wada S., Reed B.M. Genetic stability of cryopreserved shoot tips of Rubus germplasm. In Vitro Cell. Dev. Biol. Plant. 2010b;46(3):246-256. https://doi.org/10.1007/s11627-009-9265-z.

21. Castillo N.R.F., Reed B.M., Graham J., Fernandez-Fernandez F., Bas¬sil N.V. Microsatellite markers for raspberry and blackberry. J. Am. Soc. Hortic. Sci. 2010a;135:271-278.

22. Castro P., Stafne E.T., Clark J.R., Lewers K.S. Genetic map of the pri- mocane-fruiting and thornless traits of tetraploid blackberry. Theor. Appl. Genet. 2013;126:2521-2532.

23. Cekic C., Calis O., Ozturk E.S. Genetic diversity of wild raspberry genotypes (Rubus idaeus L.) in North Anatolia based on ISSR markers. Appl. Ecol. Environ. Res. 2018;16(5):6835-6843. https://doi.org/10.15666/aeer/1605_68356843.

24. Cousineau J.C., Anderson A., Daubeny H.A., Donnelly D.J. Characterization of red raspberry cultivars and selections using isoenzyme analysis. HortScience. 1993;28(12):1185-1186.

25. Dale A., Moore P.P., McNicol R.J., Sjulin T.M., Burmistrov L.A. Genetic diversity of red raspberry cultivars throughout the world. J. Am. Soc. Hort. Sci. 1993;118:119-129.

26. Daubeny H.A., Anderson A. Germplasm enhancement in the British Columbia raspberry breeding program. Acta Hortic. 1989;262:61- 64. https://doi.org/10.17660/ActaHortic.1989.262.7.

27. Dossett M., Bassil N., Finn C. SSR fingerprinting of black raspberry cultivars shows discrepancies in identification. Acta Hortic. 2012a; 946:49-53. https://doi.org/10.17660/ActaHortic.2012.946.4.

28. Dossett M., Bassil N.V., Lewers K.S., Finn C.E. Genetic diversity in wild and cultivated black raspberry (Rubus occidentalis L.) evalu¬ated by simple sequence repeat markers. Genet. Resour. Crop Evol. 2012b;59:1849-1865.

29. Dossett M., Bushakra J.M., Gilmore B., Koch C.A., Kempler C., Finn C.E., Bassil N.V. Development and transferability of black and red raspberry microsatellite markers from shortread sequences. J. Am. Soc. Hort. Sci. 2015;140(3):243-252.

30. Ercisli S., Badjakov I., Kondakova V., Atanassov A., Todorovska E. AFLP-based genetic relationships in wild and cultivated raspberry genotypes (Rubus idaeus L.). Biotechnol. Biotechnol. Equip. 2008; 22(4):907-910.

31. Eu G., Chung B., Bandopadhyay R., Yoo N.-H., Choi D., Yun S. Phylo- genic relationships of Rubus species revealed by randomly amplified polymorphic DNA markers. J. Crop Sci. Biotech. 2008;11(1):39-44.

32. Eu G., Park M., Baek S.H., Yun S. Phylogenic relationship of Rubus cultivated in Korea revealed by chloroplast DNA spacers. Korean J. Med. Crop Sci. 2010;18(4):266-272.

33. Fazekas A.J., Burgess K.S., Kenasakurti P.R., Graham S.W., Newmas- ter S.G., Husband B.C., Percy D.M., Hajibabaei M., Barrett S.C.H. Multiple multilocus DNA barcodes from the plastid genome dis-criminate plant species equally well. PLoS One. 2008;3(7):e2802. https://doi.org/10.1371/journal.pone.0002802.

34. Fernandez G.E., Molina-Bravo R., Takeda F. What we know about heat stress in Rubus. In: Graham J., Brennan R. (Eds.). Raspberry: Breed¬ing, Challenges and Advances. Springer, 2018;29-40.

35. Fernandez-Fernandez F., Antanaviciute L., Govan C.L., Sargent D.J. Development of a multiplexed microsatellite set for fingerprinting red raspberry (Rubus idaeus) germplasm and its transferability to other Rubus species. J. Berry Res. 2011;1(4):177-187.

36. Finn C.E. Blackberries. In: Hancock J.F. (Ed.).Temperate Fruit Crop Breeding: Germplasm to Genomics. Springer, 2008;83-114.

37. Focke W.O. Species Ruborum. Monographiae generis Rubi prodromus. Biblioth. Bot. 1910;17(72):1-120.

38. Focke W.O. Species Ruborum. Monographiae generis Rubi prodromus. Biblioth. Bot. 1911;17(72):121-223.

39. Focke W.O. Species Ruborum. Monographiae generis Rubi prodromus. Biblioth. Bot. 1914;17(83):1-274.

40. Girichev G., Hanke M.V., Peil A., Flachowsky H. SSR fingerprinting of a German Rubus collection and pedigree based evaluation on trueness-to-type. Genet. Resour. Crop Evol. 2015;64:89-103. https://doi.org/10.1007/s10722-015-0345-0.

41. Gotame T.P., Cullen D.W., Graham J., Hedley P.E., Smith K., Mor¬ris J., Andersen L., Petersen K.K. Effect of short-term exposure to high-temperature on total gene expression in the leaves of four rasp¬berry (Rubus idaeus L.) cultivars. J. Hortic. Sci. Biotechnol. 2014; 89(5):532-541. https://doi.org/10.1080/14620316.2014.11513117.

42. Graham J., Gordon S.C., Williamson B. Progress towards the use of transgenic plants as an aid to control soft fruit pests and diseases. In: Proceedings of Brighton Crop Protection Conference - Pests & Diseases. Brighton, UK, 1996;3:777-781.

43. Graham J., Hackett C.A., Smith K., Woodhead M., MacKenzie K., Tier¬ney I., Cooke D., Bayer M., Jennings N. Towards an understanding of the nature of resistance to Phytophtora root rot in red raspberry. Theor. Appl. Genet. 2011;123:585-601. https://doi.org/10.1007/s00122-011-1609-5.

44. Graham J., McNicol R.J., Greig K., Van De Ven W.T.G. Identification of red raspberry cultivars and an assessment of their relatedness using fingerprints produced by random primers. J. Hortic. Sci. 1994;69:123-130.

45. Graham J., Smith K., MacKenzie K., Jorgensen L., Hackett C.A., Powell W. The construction of a genetic linkage map of red raspberry (Rubus idaeus subsp. idaeus) based on AFLPs, genomic-SSR and EST-SSR markers. Theor. Appl. Genet. 2004;109:740-749.

46. Graham J., Smith K., McCallum S., Hedley P., Cullen D., Dolan A., Milne L., McNicol J., Hackett C. Towards an understanding of crumbly fruit in red raspberry. SpringerPlus. 2015;4:223. https://doi.org/10.1186/s40064-015-1010-y.

47. Graham J., Smith K., Tierney I., MacKenzie K., Hackett C. Mapping gene H controlling cane pubescence in raspberry and its association with resistance to cane botyritis and spur blight, rust and cane spot. Theor. Appl. Genet. 2006;112:818-831.

48. Graham J., Smith K., Woodhead M., Russell J.R. Development and use of simple sequence repeat SSR markers in Rubus species. Mol. Ecol. Notes. 2002;2:250-252.

49. Graham J., Squire B., Marshall B., Harrison R.E. Spatially dependent genetic diversity within and between colonies of wild raspberry R. idaeus detected using RAPD markers. Mol. Ecol. 1997;6:1001-1008.

50. Graham J., Woodhead M., Smith K. Russell J., Marshall B., Ramsay G., Squire G. New insight into wild red raspberry populations using simple sequence repeat markers. J. Am. Soc. Hort. Sci. 2009; 134(1):109-119.

51. Gustafsson A. The Genesis of the European Blackberry Flora. Acta Univ. Lund. 1943;39(6):1-200.

52. Hackett C.A., Milne L., Smith K., Hedley P., Morris J., Simpson C.J., Preedy K., Graham J. Enhancement of Glen Moy x Latham raspberry linkage map using GbS to further understand control of development processes leading to fruit ripening. BMC Genetics. 2018;19:59. https://doi.org/10.1186/s12863-018-0666-z.

53. Hoepfner A.S., Nybom H., Carlsson U., Franzen R. DNA fingerprinting useful for monitoring cell line identity in micropropagated raspberries. Acta Agric. Scand. Sect. B. Soil Plant Sci. 1993;43:53-57.

54. Ipek A., Barut E., Gulen H., Ipek M. Genetic diversity among some blackberry cultivars and their relationship with Boysenberry assessed by AFLP markers. Afr. J. Biotechnol. 2009;8:4830-4834.

55. Jennings D.L. Raspberries and Blackberries: Their Breeding, Diseases and Growth. San Diego: Acad. Press Ltd, 1988.

56. Jibran R., Dzierzon H., Bassil N., Bushakra J.M., Edger P.P., Sullivan S., Finn C.E., Dossett M., Vining K.J., VanBuren R., Mockler T.C., Li-achko I., Davies K.M., Foster T.M., Chagne D. Chromosomescale scaffolding of the black raspberry (Rubus occidentalis L.) genome based on chromatin interaction data. Hortic. Res. 2018;5:8. https://doi.org/1038/s41438-017-0013-y.

57. Kassim A., Poette J., Paterson A., Zait D., McCallum S., Woodhead M., Smith K., Hackett C., Graham J. Environmental and seasonal influences on red raspberry anthocyanin antioxidant contents and identification of quantitative traits loci. Mol. Nutr. Food Res. 2009;53: 625-634.

58. Keane B., Smith M.K., Rogstad S.H. Genetic variation in red raspberries (Rubus idaeus L., Rosaceae) from sites differing in organic pollutants compared with synthetic repeat DNA probes. Environ. Toxicol. Chem. 1998;17:2027-2034. https://doi.org/10.1002/etc.5620171019.

59. Knight V.H. Review of Rubus species used in raspberry breeding at East Malling. Acta Hortic. 1993;352:363-372. https://doi.org/10.17660/Acta-Hortic.1993.352.52.

60. Knight V.H., Jennings D.L., McNicol R.J. Progress in the UK raspberry breeding programme. Acta Hortic. 1989;262:93-104. https://doi.org/10.17660/ActaHortic.1989.262.12.

61. Kollmann J., Steinger T., Roy B.A. Evidence of sexuality in european Rubus (Rosaceae) species based on AFLP and allozyme analysis. Am. J. Bot. 2000;87(11):1592-1598.

62. Kostamo K., Toljamo A., Antonius K., Kokko H., Karenlampi S.O. Morphological and molecular identification to secure cultivar maintenance and management of self-sterile Rubus arcticus. Ann. Bot. 2013;111(4):713-721. https://doi.org/10.1093/aob/mct029.

63. Kraft T., Nybom H. DNA fingerprinting and biometry can solve some taxonomic problems in apomictic blackberries (Rubus, subgen. Rubus). Watsonia. 1995;20:329-343.

64. Kraft T., Nybom H., Werlemark G. DNA fingerprint variation in some blackberry species (Rubus subg. Rubus, Rosaceae). Plant Syst. Evol. 1996;199:93-108.

65. Lacis G., Kota-Dombrovska I., Strautina S. Evaluation of red raspberry cultivars used for breeding and commercial growing in the Baltic region. Proc. Latvian Acad. Sci. Sect. B. 2017;71(3):203-210.

66. Lamoureux D., Sorokin A., Lefevre I., Alexanian A., Eyzaguirre P., Hausmann J.F. Investigation of genetic diversity in Russian collections of raspberry and blue honeysuckle. Plant Genet. Resour. 2011; 9(2):202-205. https://doi.org/10.1017/S1479262111000323.

67. Lee G.A., Song J.Y., Choi H.R., Chung J.W., Jeon Y.A., Lee J.R., Ma K.H., Lee M.C. Novel microsatellite markers acquired from Ru-bus coreanus Miq. and cross-amplification in other Rubus species. Molecules. 2015;20:6432-6442. https://doi.org/10.3390/molecules20046432.

68. Lee K.J., Lee G.A., Kang H.K., Lee J.R., Raveendar S., Shin M.J., Cho Y.H., Ma K.H. Genetic diversity and population structure of Rubus accessions using simple sequence repeat markers. Plant Breed. Biotech. 2016;4(3):345-351. https://doi.org/10.9787/PBB.2016.4.3.345.

69. Lewers K., Saski C., Cuthbertson B., Henry D., Staton M., Main D., Dhanaraj A., Rowland L., Tomkins J. A blackberry (Rubus L.) expressed sequence tag library for the development of simple sequence repeat markers. BMC Plant Biol. 2008;8:69. https://doi.org/10.1186/1471-2229-8-69.

70. Liang Y., Lenz R.R., Dai W. Development of retrotransposon-based molecular markers and their application in genetic mapping in chokecherry (Prunus virginiana L.). Mol. Breed. 2016;36:109. https://doi.org/10.1007/s11032-016-0535-2.

71. Lopes M., Maciel G., Mendonca D., Gil F., Da Camara A. Isolation and characterization of simple sequence repeat loci in R. hochstet-terorum and their use in other species from the Rosaceae family. Mol. Ecol. Notes. 2006;6:750-752. https://doi.org/1111/j.1471-8286.2006.01329.x.

72. López A., Barrera C., Marulanda M. Evaluation of SSR and SNP markers in R. glaucus Benth progenitors’ selection. Rev. Bras. Frutic. 2019;41(1):1-14. https://doi.org/10.1590/0100-29452019081.

73. Marieschi M., Torelli A., Poli F., Bianchi A., Bruni R. Quality control of commercial Mediterranean oregano: development of SCAR markers for the detection of the adulterants Cistus incanus L., Rubus caesius L. and Rhus coriaria L. Food Control. 2010;21:998-1003.

74. Marulanda M., Lopez A., Aguilar S. Genetic diversity of wild and cultivated Rubus species in Colombia using AFLP and SSR markers. Crop Breed. Appl. Biotechnol. 2007;7:242-252.

75. Mazzitelli L., Hancock R., Haupt S., Walker P., Pont S., McNicol J., Cardle L., Morris J., Viola R., Brennan R., Hedley P., Taylor M.A. Co-ordinated gene expression during phases of dormancy release in raspberry (R. idaeus L.) buds. J. Exp. Bot. 2006;58(5):1035-1045.

76. Meng R., Finn C. Determining ploidy level and nuclear DNA content in Rubus by flow cytometry. J. Am. Soc. Hort. Sci. 2002;127(5):767-775. https://doi.org/10.21273/JASHS.127.5.767.

77. Miyashita T., Kunitake H., Yotsukura N., Hoshino Y. Assessment of genetic relationships among cultivated and wild Rubus accessions using AFLP-markers. Sci. Hortic. 2015;193:165-173.

78. Molina-Bravo R., Fernandez G.E., Sosinski B.R. Quantitative trait locus analysis of tolerance to temperature fluctuations in winter, fruit characteristics, flower color, and prickle-free canes in raspberry. Mol. Breed. 2014;33:267-280.

79. Moore P.P. Chloroplast DNA diversity in raspberry. J. Am. Soc. Hortic. Sci. 1993;118:371-376.

80. Nybom H. Biometry and DNA fingerprinting detect limited genetic differentiation among populations of the apomictic blackberry Rubus nessensis (Rosaceae). Nordic J. Bot. 1998;18:323-333. https://doi.org/10.1111/j.1756-1051.1998.tb01884.x.

81. Nybom H., Hall H.K. Minisatellite DNA ‘fingerprints’ can distinguish Rubus cultivars and estimate their degree of relatedness. Euphytica. 1991;53:107-114.

82. Nybom H., Rogstad S.H., Schaal B.A. Genetic variation detected by use of the M13 ‘DNA fingerprint’ probe inMalus, Prunus and Rubus (Rosaceae). Theor. Appl. Genet. 1990;79:153-156.

83. Ochieng J.A., Oyoo M.E., Gesimba R.M., Korir P.C., Ojwang P.P.O., Owuoche J.O. Genetic diversity of blackberry (Rubus subgenus Ru-bus Watson) in selected counties in Kenya using simple sequence repeats (SSRs) markers. Afr. J. Biotechnol. 2018;17(39):1247-1264. https://doi.org/10.5897/AJB2018.16613.

84. Oydvin J. Important Breeding Lines and Cultivars in Raspberry Breeding (in Norwegian). St. Forsokag. Njos, 1970.

85. Parent J.G., Page D. Identification of raspberry cultivars by non-radioactive DNA fingerprinting. HortScience. 1992;27:1108-1110.

86. Parent J.G., Page D. Identification of raspberry cultivars by sequence characterized amplified region DNA analysis. HortScience. 1998; 33:140-142.

87. Raluca R.A., Pamfil D., Graham J. Mapping resistance of red raspberry (Rubus idaeus subsp. idaeus) to viral diseases - leaf spot (RLSV) and vein chlorosis (RVCV) on the genetic linkage map. USAMV-CN. 2006;63:318-319.

88. Robertson K.R. The genera Rosaceae in the southeastern United States. J. Arnold Arboretum. 1974;55:352-360.

89. Ryu J., Kim W.J., Im J., Kim S.H., Lee K.S., Jo H.J., Kim E.Y., Kang S.Y., Lee J.H., Ha B.H. Genotyping-by-sequencing based single nucleotide polymorphisms enabled Kompetitive Allele Specific PCR marker development in mutant Rubus genotypes. Electron. J. Biotechnol. 2018;35:57-62. https://doi.org/10.1016/j.ejbt.2018.08.001.

90. Saddhe A.A., Kumar K. DNA barcoding of plants: selection of core markers for taxonomic groups. Plant Sci. Today. 2018;5(1):9-13.

91. Sargent D.J., Fernandez-Fernandez F., Rys A., Knight V.H., Simpson D.W., Tobutt K.R. Mapping of A1 conferring resistance to Am-phophora idaei and dw (dwarfing habit) in red raspberry (Rubus idaeus L.) using AFLP and microsatellite markers. BMC Plant Biol. 2007;7:15.

92. Simlat M., Ptak A., Kula A., Orzel A. Assessment of genetic variability among raspberry accessions using molecular markers. Acta Sci. Pol. Hortorum Cultus. 2018;17(5):61-72. https://doi.org/10.24326/asphc.2018.5.6.

93. Simpson C.G., Cullen D.W., Hackett C.A., Smith K., Hallett P.D., McNicol J., Woodhead M. Mapping and expression of genes associated with raspberry fruit ripening and softening. Theor. Appl. Genet. 2017;130(3):557-572.

94. Stafne E., Clark J., Pelto M., Lindstrom J. Discrimination of Rubus cultivars using RAPD markers and pedigree analysis. Acta Hortic. 2003;626:119-124. https://doi.org/10.17660/ActaHortic.2003.626.16.

95. Strik B.C., Finn C.E. Blackberry production systems - a worldwide perspective. Acta Hortic. 2012;946:341-347. https://doi.org/10.17660/Acta-Hortic.2012.946.56.

96. Ukhatova Y.V., Dunaeva S.E., Antonova O.Y., Apalikova O.V., Pozd-niakova K.S., Novikova L.Y., Shuvalova L.E., Gavrilenko T.A. Cryopreservation of red raspberry cultivars from the VIR in vitro collection using a modified droplet vitrification method. In Vitro Cell. Dev. Biol. Plant. 2017;53:394-401. https://doi.org/10.1007/s11627-017-9860-3.

97. Umar G., Vasanthaiah H., Kambiranda D., Basha S., Phills B., Hunter W. Assessment of genetic diversity among selected raspberry cultivars. Proc. Fla. State Hort. Soc. 2010;123:26-28.

98. VanBuren R., Bryant D., Bushakra J.M., Vining K.J., Edger P.P., Rowley E.R., Priest H.D., Michael T.P., Lyons E., Filichkin S.A., Dossett M., Finn C.E., Bassil N.V., Mockler T.C. The genome of black raspberry (Rubus occidentalis). Plant J. 2016;87:535-547.

99. VanBuren R., Bryant D., Bushakra J.M., Vining K.J., Filichkin S., Edger P.P., Rowley E.R., Priest H.D., Michael T.P., Dossett M., Finn C.E., Bassil N.V., Mockler T.C. Sequence and analysis of the black raspberry (Rubus occidentalis) genome. In: Hytonen T., Graham J., Harrison R. (Eds.). The Genomes of Rosaceous Berries and Their Wild Relatives. Springer, 2018:185-197.

100. Wang Y., Chen Q., Chen T., Tang H., Liu L., Wang X. Phylogenetic insights into Chinese Rubus (Rosaceae) from multiple chloroplast and nuclear DNAs. Front. Plant Sci. 2016;7:968. https://doi.org/10.3389/flps.2016.00968.

101. Ward J.A., Bhangoo J., Fernandez-Fernandez F., Moore P., Swanson J.D., Viola R., Velasco R., Bassil N., Weber C.A., Sargent D.J. Saturated linkage map construction in Rubus idaeus using genotyp-ing by sequencing and genome-independent imputation. BMC Genomics. 2013;14:2.

102. Ward J.A., Boone W.E., Moore P.P., Weber C.A. Developing molecular markers for marker assisted selection for resistance for Raspberry bushy dwarf virus (RBDV) in red raspberry. Acta Hortic. 2012;946:61-66. https://doi.org/10.17660/ActaHortic.2012.946.6.

103. Waugh R., Van de Ven W.T.G., Phillips M.S., Powell W. Chloroplast diversity in the genus Rubus revealed by Southern hybridization. Plant Syst. Evol. 1990;172:65-75.

104. Weber C.A. Genetic diversity in black raspberry detected by RAPD markers. HortScience. 2003;38(2):269-272.

105. Weber C.A., Pattison J., Samuelian S. Marker assisted selection for resistance to root rot in red raspberry caused by Phytophthora fraga-riae var. rubi. Acta Hortic. 2008;777:311-316. https://doi.org/10.17660/Acta-Hortic.2008.777.46.

106. Werlemark G., Nybom H. Pollen donor impact on progenies of pseu-dogamous blackberries (Rubus subgen. Rubus). Euphytica. 2003; 133:71-80.

107. Wight H., Zhou J., Li M., Hannenhalli S., Mount S., Liu Z. Draft genome assembly and annotation of red raspberry Rubus idaeus. bioRxiv. 2019. https://doi.org/10.1101/546135.

108. Woodhead M., Smith K., McCallum S., Cardle L., Mazzitelli M., Graham J. Identification, characterisation and mapping of simple sequence repeat (SSR) markers from raspberry root and bud ESTs. Mol. Breed. 2008;22:555-563.

109. Woodhead M., Weir A., Smith K., McCallum S., MacKenzie K., Graham J. Functional markers for red raspberry. J. Am. Soc. Hort. Sci. 2010;135(5):418-427.

110. Woodhead M., Williamson S., Smith K., McCallum S., Jennings N., Hackett C., Graham J. Identification of quantitative trait loci for cane splitting in red raspberry (Rubus idaeus). Mol. Breed. 2013;31:111-122.

111. Zurn J.D., Carter K.A., Yin M.H., Worthington M., Clark J.R., Finn C.E., Bassil N. Validating blackberry seedling pedigrees and developing an improved multiplexed microsatellite fingerprinting set. J. Am. Soc. Hort. Sci. 2018;143(5):381-390.