Межвидовая гибридизация картофеля: теоретические и прикладные аспекты

Возделываемый картофель Solanum tuberosum L. является четвертой по значимости продовольственной культурой в мире и, как многие другие культурные растения, характеризуется низким уровнем генетического разнообразия. Наиболее эффективный подход к расширению генетического разнообразия селекционных сортов основан на интрогрессии генетического материала диких и культурных видов картофеля, относящихся к секции Petota Dumort. рода Solanum L., которая, по оценкам разных систематиков, включает от 112 до 235 видов. В то же время из-за барьеров презиготической и постзиготической несовместимости в генофонд селекционных сортов был интрогрессирован генетический материал лишь около 10 % дикорастущих видов картофеля. К настоящему време- ни в области молекулярной генетики, геномики и биотехнологии накоплен большой объем информации, позволяющий решать задачи кардинального расширения генетического разнообразия культурных растений. В статье приведен краткий обзор современных данных о генетических факторах, детерминирующих презиготическую и постзиготическую несовместимость, особенностях вовлечения в гибридизационный процесс самосовместимых и самонесовместимых видов секции Petota, видов c разными значениями балансового числа эндосперма (EBN), а также о возможных механизмах взаимодействия родительских геномов при формировании эндосперма гибридного семени. Показано значение теории EBN для дальнейшего развития прикладных исследований, направленных на расширение генетического разнообразия селекционного генофонда картофеля. Рассматриваются традиционные и современные подходы и методы преодоления межвидовой несовместимости и приведены примеры успешных исследований по интрогрессивной гибридизации картофеля. Статья предваряется сведениями о генетическом разнообразии видов картофеля, относящихся к секции Petota рода Solanum, а также о концепции генных пулов этих видов.

Solanum, картофель, межвидовая несовместимость, интрогрессивная гибридизация

1. Abdalla M.M., Hermsen J.G.Th. An evaluation of Solanum verrucosum Schlechtd. for its possible use in potato breeding. Euphytica. 1973; 22:19-27.

2. Adiwilaga K.D., Brown C.R. Use of 2n pollen-producing triploid hybrids to introduce tetraploid Mexican wild species germplasm to cultivated tetraploid potato gene pool. Theor. Appl. Genet. 1991;81:645-652.

3. Anisimova I.N., Gavrilenko T.A. The prospects for utilization of cytoplasmic male sterility in breeding and genetic studies and seed production of potato. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov Journal of Genetics and Breeding. 2017;21(1):83-95. DOI 10.18699/VJ17.226. (in Russian)

4. Antonova O.Yu., Gavrilenko T.A. Yaderno-tsitoplazmaticheskie vzaimootnosheniya u somaticheskikh gibridov roda Solanum L. [Nuclear-cytoplasmic interactions in somatic hybrids of the genus Solanum L.]. Struktura i ekspressiya mitokhondrial’nogo genoma rasteniy [Structure and expression of the plant mitochondrial genome]. Irkutsk, 2006;5−10. (in Russian)

5. Austin S., Pohlman J.D., Brown C.R., Mojtahedi H., Santo G.S., Douches D.S., Helgeson J.P. Interspecific somatic hybridization between Solanum tuberosum L. and S. bulbocastanum Dun. as a means of transferring nematode resistance. Am. Potato J. 1993;70:485-495.

6. Berger F. Endosperm: the crossroad of seed development. Curr. Opin. Plant Biol. 2003;6:42-50.

7. Bradeen J.M., Haynes K.G. Introduction to Potato. Ch. 1. Genetics, Genomics and Breeding of Potato. Eds. J. Bradeen, C. Kole. 1st ed. USA: Sci. Publ., 2011;1-19.

8. Bradeen J.M., Naess S.K., Song J., Haberlach G.T., Wielgus S.M., Buell C.R., Jiang J., Helgeson J.P. Concomitant reiterative BAC walking and fine genetic mapping enable physical map development for the broad-spectrum late blight resistance region, RB. Mol. Genet. Genom. 2003;269:603-611. DOI 10.1007/s00438-003-0865-8.

9. Brown C.R., Mojtahedi H., James S., Novy R.G., Love S. Development and evaluation of potato breeding lines with introgressed resistance to Columbia root-knot nematode (Meloidogyne chitwoodi). Am. J. Potato Res. 2006;83:1-8.

10. Brown C.R., Mojtahedi H., Santo G.S. Introgression of resistance to Columbia and northern root-knot nematodes from Solanum bulbocastanum into cultivated potato. Euphytica. 1995;83:71-78.

11. Brown C.R., Yang C.-P., Mojtahedi H., Santo G.S., Masuelli R. RFLP analysis of resistance to Columbia root-knot nematode derived from Solanum bulbocastanum in a BC2 population. Theor. Appl. Genet. 1996;92:572-576.

12. Bukasov S.M. Principles of potato systematics. Trudy po prikladnoy botanike, genetike i selektsii = Proceedings on Applied Botany, Genetics and Breeding. 1978;62:3-35. (in Russian)

13. Cai D., Rodríguez F., Teng Y., Ané C., Bonierbale M., Mueller L.A., Spooner D.M. Single copy nuclear gene analysis of polyploidy in wild potatoes (Solanum section Petota). BMC Evol. Biol. 2012;12:70.

14. Camadro E.L., Espinillo J.C. Germplasm transfer from the wild tetraploid species Solanum acaule Bitt. to the cultivated potato, S. tuberosum L. using 2n eggs. Am. Potato J. 1991;67:737-749.

15. Carputo D., Barone A., Cardi T., Sebastiano A., Frusciante L., Peloquin S.J. Endosperm Balance Number manipulation for direct in vivo germplasm introgression to potato from a sexually isolated relative (Solanum commersonii Dun.). Proc. Natl. Acad. Sci. USA. 1997;94:12013-12017.

16. Carputo D., Basile B., Cardi T., Frusciante L. Erwinia resistance in backcross progenies of Solanum tuberosum × S. tarijense and S. tuberosum (+) S. commersonii hybrids. Potato Res. 2000;43:135-142. DOI 10.1007/BF02357954.

17. Castillo R., Spooner D.M. Phylogenetic relationships of wild potatoes, Solanum series Conicibaccata (sect. Petota). Syst. Botany. 1997;22: 45-83.

18. Chen Y.-K.H., Palta J.P., Bamberg J.B. Freezing tolerance and tuber production in selfed and backcross progenies derived from somatic hybrids between Solanum tuberosum L. and S. commersonii Dun. Theor. Appl. Genet. 1999;99:100-107.

19. Cipar M.S., Peloquin S.J., Hougas R.W. Variability in the expression of self-incompatibility in tuber-bearing diploid Solanum species. Am. Potato J. 1964;41:155-162.

20. Clarke A.E., Newbigin E. Molecular aspects of self-incompatibility in flowering plants. Annu. Rev. Genet. 1993;27:257-279.

21. Clulow S.A., Wilkinson M.J., Waugh R., Baird E., Demaine M.J., Powell W. Cytological and molecular observations on Solanum phurejainduced dihaploid potatoes. Theor. Appl. Genet. 1991;82:545-551. DOI 10.1007/BF00226789.

22. Corell D. The Potato and its Wild Relatives. Renner. Texas: Texas Res. Foundation, 1962.

23. Davey M.R., Anthony P., Power J.B., Lowe K.C. Plant protoplasts: status and biotechnological perspectives. Biotechnol. Advances. 2005; 23:131-171. DOI 10.1016/j.biotechadv.2004.09.008.

24. de Haan S., Rodriguez F. Potato origin and production. Advances in Potato Chemistry and Technology. Eds. J. Singh, L. Kaur. London (UK): Elsevier Inc., 2016;1-32.

25. Dinu I.I., Hayes R.J., Kynast R.G., Phillips R.L., Thill C.A. Novel inter-series hybrids in Solanum, section Petota. Theor. Appl. Genet. 2005;110:403–415. DOI: 10.1007/s00122-004-1782-x.

26. Eckardt N.A. F-box proteins take center stage. Plant Cell. 2004;16: 558-561.

27. Ehlenfeldt M.K., Hanneman R.E., Jr. Genetic control of Endosperm Balance Number (EBN): three additive loci in a threshold-like system. Theor. Appl. Genet. 1988;75:825-832.

28. Eijlander R. Mechanisms of self-incompatibility and unilateral incompatibility in diploid potato (Solanum tuberosum L.) Wageningen, Netherlands: Wageningen Univ., 1998.

29. Eijlander R., ter Laak W., Hermsen J.G.Th., Ramanna M.S., Jacobsen E. Occurrence of self-compatibility, self-incompatibility and unilateral incompatibility after crossing diploid S. tuberosum (SI) with S. verrucosum (SC): I. Expression and inheritance of self-compatibility. Euphytica. 2000;115(2):127-139.

30. Fujii S., Kubo K., Takayama S. Non-self and self recognition models in plant self-incompatibility. Nature Plants. 2016;2(9):16130. DOI 10.1038/NPLANTS.2016.130.

31. Gavrilenko T.A. Sozdanie novykh form kul’turnykh rasteniy na osnove somaticheskoy gibridizatsii [Raise of novel crop forms by somatic fusion] Rigin B.V. (Ed.). Identifitsirovannyy genofond rasteniy i selektsiya [Identified plant gene pool and breeding]. St. Petersburg, VIR Publ., 2005;628-643. (in Russian)

32. Gavrilenko T. Application of molecular cytogenetics in fundamental and applied research of potato (Review). Ch. 9. Genetics, Genomics and Breeding of Potato. Eds. J. Bradeen, C. Kole. 1st ed. USA: Sci. Publ., 2011;184-206.

33. Gavrilenko T., Antonova O., Thieme R., Szczerbakowa A., Wielgat B. Inheritance of chloroplast and mitochondrial DNAs in interspecific somatic hybrids of potato. EAPR-Papers. Bilbao, Spain. 2005;II: 652-654.

34. Gavrilenko T., Pendinen G., Rokka V.-M., Antonova O., Thieme R. Homeologous chromosome pairing in distant allohaploid hybrids of the genus Solanum. Rus. J. Genetics: Appl. Res. 2015;5(3):182-190.

35. Gavrilenko T., Thieme R., Heimback U., Thieme T. Fertile somatic hybrids of Solanum etuberosum (+) dihaploid Solanum tuberosum and their backcrossing progenies: relationships of genome dosage with tuber development and resistance to potato virus Y. Euphytica. 2003;131:323-332. DOI 10.1023/A:1024041104170.

36. Gebhardt C. Bridging the gap between genome analysis and precision breeding in potato. Trends in Genetics. 2013;29(4):248-256. DOI 10.1016/j.tig.2012.11.006.

37. Gillen A.M., Novy R.G. Molecular characterization of the progeny of Solanum etuberosum identifies a genomic region associated with resistance to potato leafroll virus. Euphytica. 2007;155:403-415. DOI 10.1007/s10681-006-9342-x.

38. Glimelius K., Fahleson G., Landgren M., Sjodin C., Sundberg E. Gene transfer via somatic hybridization in plants. Trends Biotechnol. 1991;9:24-30.

39. Grun P., Aubertin M. The inheritance and expression of unilateral incompatibility in Solanum. Heredity. 1966;212:131-138. DOI 10.1038/hdy.1966.7.

40. Hanneman R.E., Jr. The reproductive biology of the potato and its implication for breeding. Potato Res. 1999;42:283-312.

41. Harlan J.R., de Wet J.M.J. Toward a rational classification of cultivated plants. Taxon. 1971;20(4):509-517. DOI 10.2307/1218252.

42. Haverkort A.J., Struik P.C., Visser R.G.F., Jacobsen E. Applied biotechnology to combat late blight in potato caused by Phytophthora infestans. Potato Res. 2009;52:249-264.

43. Hawkes J.G. A revision of the tuber-bearing Solanums. Scottish Plant Breed. Stat. Ann. Rep. 1956;37-109.

44. Hawkes J.G. The Potato: Evolution, Biodiversity and Genetic Resources. London: Belhaven Press, 1990.

45. Hawkes J.G. Origins of cultivated potatoes and species relationships. Potato Genetics. Eds. J.E. Bradshaw, G.R. Mackay. Wallingford: CAB International, 1994;3-42.

46. Helgeson J.P., Hunt G.J., Haberlach G.T., Ehlenfeldt M.K., Hunt G., Pohlman J.D., Austin S. Sexual progeny of somatic hybrids between potato and Solanum brevidens: potential for use in breeding programs. Am. Potato J. 1993;70(6):437-452.

47. Helgeson J.P., Pohlman J.D., Austin S., Haberlach G.T., Wielgus S.M., Ronis D., Zambolim L., Tooley P., McGrath J.M., James R.V., Stevenson W.R. Somatic hybrids between Solanum bulbocastanum and potato: a new source of resistance to late blight. Theor. Appl. Genet. 1998;96:738-742. DOI 10.1007/s001220050796.

48. Hermsen J.G.Th. Mechanisms and genetic implications of 2n-gamete formation. Iowa State J. Research. 1984;58:421-434.

49. Hermsen J.G.Th. Introgression of genes from wild species, including molecular and cellular approaches. Potato Genetics. Eds. J.E. Bradshaw, G.R. Mackay. Wallingford, UK: CABI, 1994;515-539.

50. Hermsen J.G.Th., Olsder J., Jansen P., Hoving E. Acceptance of selfcompatible pollen from Solanum verrucosum in dihaploids from S. tuberosum. Fertilization in Higher Plants. Ed. H.F. Linskens. Amsterdam; New York: North Holland Am. Elsevier, 1974;37-40.

51. Hermsen J.G.Th., Ramanna M.S. Barriers to hybridization of Solanum bulbocastanum Dun. and Solanum verrucosum Schlechtd. and structural hybridity in their F1 plants. Euphytica. 1976;25(1):1-10. DOI 10.1007/BF00041523.

52. Hermsen J.G.Th., Verdenius J. Selection from Solanum tuberosum group Phureja of genotypes combining high frequency haploid induction with homozygosity for embryo-spot. Euphytica. 1973;22:244-259.

53. Hijmans R.J., Gavrilenko T., Stephenson S., Bamberg J., SalasA., Spooner D. The geographic and environmental distribution of polyploidy in wild potatoes (Solanum section Petota). Global Ecol. Biogeogr. 2007;16(4):485-495. DOI 10.1111/j.1466-8238.2007.00308.x.

54. Hijmans R.J., Spooner D.M. Geographic distribution of wild potato species. Am. J. Botany. 2001;88:2101-2112.

55. Hijmans R.J., Spooner D.M., Salas A.R., Guarino L., de la Cruz J. Atlas of Wild Potatoes. Syst. Ecogeogr. Studies on Crop Genepools. 2002. www.vcru.wisc.edu/spoonerlab/pdf/Potato%20Atlas%20Final.pdf.

56. Horsman K., Gavrilenko T., Bergervoet M., Huigen D.J., Joe A.T.W., Jacobsen E. Alteration of the genomic composition of Solanum nigrum (+) potato backcross derivatives by somatic hybridization: selection of fusion hybrids by DNA measurements and GISH. Plant Breeding. 2001;120:201-207. DOI 10.1046/j.1439-0523.2001.00591.x.

57. Hosaka K., Hanneman R.E. Genetics of self-compatibility in a self-incompatible wild diploid potato species Solanum chacoense. 1. Detection of an S locus inhibitor (Sli) gene. Euphytica. 1998a;99:191-197.

58. Hosaka K., Hanneman R.E. Genetics of self-compatibility in a selfincompatible wild diploid potato species Solanum chacoense. 2. Localization of an S locus inhibitor (Sli) gene on the potato genome using DNA markers. Euphytica. 1998b;103:65-271.

59. Irikura Y. Cytogenetic studies on the haploid plants of tuber-bearing Solanum species. II. Cytogenetic investigations on haploid plants and interspecific hybrids by utilizing haploidy. Res. Bull. Hokkaido Nat. Agric. Exp. Stat. 1976;115:1-80.

60. Jansky S.H. Overcoming hybridization barriers in potato. Plant Breeding. 2006;125:1-12.

61. Jansky S.H., Dempewolf H., Camadro E.L., Simon R., Zimnoch-Guzowska E., Bisognin D.A., Bonierbale M.A. Case for crop wild relative preservation and use in potato. Crop Science. 2013;53:746-754. DOI 10.2135/cropsci2012.11.0627.

62. Jansky S.H., Hamernik A.J. The introgression of 2x 1 EBN Solanum species into the cultivated potato using Solanum verrucosum as a bridge. Genet. Resour. Crop. Evol. 2009;56:1107-1115.

63. Jansky S.H., Peloquin S.J., Yerk G.L. Use of potato haploids to put 2x wild species germplasm in usable form. Plant Breeding. 1990; 104(4):290-294. DOI 10.1111/j.1439-0523.1990.tb00438.x.

64. Johnston S.A., den Nijs T.M., Peloquin S.J., Hanneman R.E., Jr. The significance of genetic balance to endosperm development in interspecific crosses. Theor. Appl. Genet. 1980;57:5-9.

65. Johnston S.A., Hanneman R.E., Jr. Support of the endosperm balance number hypothesis utilizing some tuber-bearing Solanum species. Am. Potato J. 1980;57:7-14.

66. Kelley K., Whitworth J.L., Novy R.G. Mapping of the potato leafroll virus resistance gene, Rlretb, from Solanum etuberosum identifies interchromosomal translocations among its E-genome chromosomes 4 and 9 relative to the A-genome of Solanum L. sect. Petota. Mol. Breeding. 2009;23:489-500.

67. Kim S.R., Ahn Y.K., Kim T.G., Kang H.S., Song S.W., Kim B.C., Kang S.G. Breeding of a new cultivar ‘Jeseo’ with resistance to common scab. Kor. J. Breed. Sci. 2013;45:468-473. DOI https://doi.org/10.9787/KJBS.2013.45.4.468.

68. Kinoshita T. Reproductive barrier and genomic imprinting in the endosperm of flowering plants. Genes Genet. Syst. 2007;82:177-186.

69. Kradolfer D., Wolff P., Jiang H., Siretskiy A., Kohler С. An imprinted gene underlies postzygotic reproductive isolation in Arabidopsis thaliana. Dev. Cell. 2013;26:525-535.

70. Kubo K., Entani T., Takara A., Wang N., Fields A.M., Hua Z. Toyoda M., Kawashima S., Ando T., Isogai А., Kao T., Takayama S. Collaborative non-self recognition system in S-RNase-based selfincompatibility. Science. 2010;330:796-799.

71. Kuhl J.C., Novy R.G., Whitworth J.L., Dibble M.S., Schneider B., Hall D. Development of molecular markers closely linked to the Potato leafroll virus resistance gene, Rlretb, for use in marker-assisted selection. Am. J. Potato Res. 2016;93(3):203-212.

72. Kuhl J.C., Zarka K., Coombs J., Kirk W., Douches D.S. Late blight resistance of RB transgenic potato lines. J. Am. Soc. Hort. Sci. 2007; 132(6):783-789.

73. Laferriere L., Helgeson J., Allen C. Fertile Solanum tuberosum+S. commersonii somatic hybrids as sources of resistance to bacterial wilt caused by Ralstonia solanacearum. Theor. Appl. Genet. 1999;98: 1272-1278. DOI 10.1007/s001220051193.

74. Lamm R. Investigation of some tuber-bearing Solanum hybrids. Hereditas. 1953;39:97-112.

75. Levyy A.V., Voronkova E.V., Polyukhovich Yu.V., Yermishin A.P. Polymorphism of microsatellite DNA fragments in interspecific hybrids contrasting for late blight resistance that were derived from from Solanum stoloniferum. Vestsi NAN Belarusi. Ser. biyal. navuk = Proc. Nat. Acad. Sci. Belarus. Ser. Biol. Sci. 2015;1:43-49. (in Russian)

76. Levyy A.V., Polyukhovich Yu.V., Voronkova E.V., GukasyanO.N., Yermishin A.P. Use of Solanum verrucosum and Sv-lines for overcoming unilateral incompatibility in involvement to breeding of wild allotetraploid potato species S. stoloniferum. Tez. dokl. mezhd. nauch. konf. Sankt-Peterburg, 3–5 avgusta 2016 g. [Abstracts from the Int. Conf. “Problems of systematics and breeding of potatoˮ. Aug. 3–5, 2016]. St.-Petersburg, 2016;69-70. (in Russian)

77. Lewis D., Crowe L.K. Unilateral incompatibility in flowering plants. Heredity. 1958;12:233-256.

78. Li W., Chetelat R.T. Unilateral incompatibility gene ui1.1 encodes an S-locus F-box protein expressed in pollen of Solanum species. Proc. Natl. Acad. Sci. USA. 2015;112(14):4417-4422. DOI 10.1073/pnas.1423301112.

79. Lin B.-Y. Ploidy barrier to endosperm development in maize. Genetics. 1984;107:103-115.

80. Makhan’ko O.V. Interspecific incompatibility in potato diploid breeding. Zemlyarobstva i akhova raslin = Agriculture and Plant Protection. 2008;1:11-14. (in Russian)

81. Marks G.E. Cytogenetic studies in tuberous Solanum species. I. Genomic differentiation in the group Demissa. J. Genetics. 1955;53: 262-269.

82. Marks G.E. Cytogenetic studies in tuberous Solanum species: III. Species relationships in some South and Central American species. New Phytologist. 1965;64:293-306.

83. Marks G.E. The origin and significance of intraspecific polyploidy: experimental evidence from Solanum chacoense. Evolution. 1966;20: 552-557.

84. Martin F.W. Staining and observing pollen tubes in the style by means of fluorescence. Stain Technol. 1958;34:125-128.

85. Matsubayashi M. Phylogenetic relationships in the potato and its related species. Chromosome Engineering in Plants: Genetics, Breeding, Evolution. Pt. B. Eds. T. Tsuchiya, P.K. Gupta. Amsterdam: Elsevier, 1991;93-118.

86. McGrath J., Williams C., Haberlach G., Wielgus S., Uchytil T., Helgeson J.P. Introgression and stabilization of Erwinia tuber soft rot resistance into potato after somatic hybridization of Solanum tuberosum and S. brevidens. Am. J. Potato Res. 2002;79:19-24.

87. Mok D.W.S., Peloquin S.J. Three mechanisms of 2n pollen formation in diploid potatoes. Can. J. Genet. Cytol. 1975;17:217-225.

88. Naess S., Bradeen J., Wielgus S., Haberlach G., McGrath J., Helgeson J.P. Resistance to late blight in Solanum bulbocastanum is mapped to chromosome 8. Theor. Appl. Gen. 2000;101:697-704.

89. Naess S., Bradeen J., Wielgus S., Haberlach G., McGrath J., Helgeson J.P. Analysis of the introgression of Solanum bulbocastanum DNA into potato breeding lines. Mol. Gen. Genomics. 2001;265(4): 694-704.

90. Nishiyama I., Inomata N. Embryological studies in cross incompatibility between 2x and 4x in Brassica. Japan. J. Genetics. 1966;41(1):27-42.

91. Novy R.G., Gillen A.M., Whitworth J.L. Characterization of the expression and inheritance of potato leafroll virus (PLRV) and potato virus Y (PVY) resistance in three generations of germplasm derived from Solanum etuberosum. Theor. Appl. Genet. 2007;114:11611172. DOI 10.1007/s00122-007-0508-2.

92. Novy R.G., Nasruddin A., Ragsdale D.W., Radcliffe E.B. Genetic resistances to potato leafroll virus, potato virus Y, and green peach aphid in progeny of Solanum etuberosum. Am. J. Potato Res. 2002;79: 9-18. DOI 10.1007/BF02883518.

93. Ortiz R. Potato breeding via ploidy manipulations. Plant Breed. Rev. 1998;16:15-86.

94. Ortiz R., Ehlenfeldt M. The importance of Endosperm Balance Number in potato breeding and the evolution of tuber-bearing Solanum species. Euphytica. 1992;60:105-113.

95. Pandey K.K. Interspecific incompatibility in Solanum species. Am. J. Bot. 1962;49:874-882.

96. Peloquin S.J., Boiteux L.S., Carputo D. Meiotic mutants in potato. Valuable variants. Genetics. 1999;153(4):1493-1499.

97. Peloquin S.J., Yerk G.L., Werner J.E., Darmo E. Potato breeding with haploids and 2n gametes. Genome. 1989;31:1000-1004.

98. Pendinen G., Gavrilenko T., Jiang J., Spooner D.M. Allopolyploid speciation of the tetraploid Mexican potato species revealed by genomic in situ hybridization. Genome. 2008;51:714-720. DOI 10.1139/G08-052.

99. Pendinen G., Spooner D.M., Jiang J., Gavrilenko T. Genomic in situ hybridization (GISH) reveals both auto- and allopolyploid origins of different North and Central American hexaploid potato (Solanum section Petota) species. Genome. 2012;55(6):407-415. DOI 10.1139/G2012-027.

100. Perl A., Aviv D., Galun E. Protoplast-fusion-derived Solanum hybrids: application and phylogenetic limitations. Theor. Appl. Genet. 1990; 79(5):632-640.

101. Pidgaetskiy A.A. Vikarystannya dikogo vidu kartopli S. demissum Lindl. pri zaluchenni do selektsionnoy roboti S. bulbocastanum Dun. [Use of wild potato species S. demissum Lindl. in the involvement of S. bulbocastanum Dun. to breeding]. Kartoplyarstvo, mezhvedomstvennyy tematicheskiy sbornik [Potato Growing, an interdepartmental subject collection]. Kyiv, 1981;12:9-12. (in Ukrainian)

102. Pignatta D., Gehring M. Imprinting meets genomics: new insights and new challenges. Curr. Opin. Plant. Biol. 2012;15(5):530-535. DOI 10.1016/j.pbi.2012.09.004.

103. Polyukhovich Yu.V., Makhan’ko O.V., Savchuk A.V., Voronkova E.V., Yermishin A.P. Development of bridge lines for overcoming interspecific incompatibility in potatoes. Vestsi NAN Belarusi. Ser. biyal. navuk = Proc. Nat. Acad. Sci. Belarus. Ser. Biol. Sci. 2010;2:51-58. (in Russian)

104. Polyukhovich Yu.V., Voronkova E.V., Savchuk A.V., Yermishin A.P. Use of Solanum verrucosum for overcoming unilateral incompatibility in crosses with wild allotetraploid potato species. Kartofelevodstvo = Potato Growing (Minsk), 2013;21(1):136-145. (in Russian)

105. Ramakrishnan A.P., Ritland C.E., Blas Sevillano R.H., Riseman A. Review of potato molecular markers to enhance trait selection. Am. J. Potato Res. 2015;92(4):455-472. DOI 10.1007/s12230-015-9455-7.

106. Ramanna M.S. A re-examination of the mechanisms of 2n-gamete formation in potato and its implications for breeding. Euphytica. 1979;28:537-561.

107. Ramanna M.S. First division restitution gametes through fertile desynaptic mutants of potato. Euphytica. 1983;32:337-350.

108. Rivard S.R., Cappadocia M., Landry B.S. A comparison of RFLP maps based on another culture derived, selfed, and hybrid progenies of Solanum chacoense. Genome. 1996;39:611-621.

109. Rodríguez F., Ghislain M., Clausen A.M., Jansky S.H., Spooner D.M. Hybrid origins of cultivated potatoes. Theor. Appl. Genet. 2010; 121:1187-1198. DOI 10.1007/s00122-010-1422-6.

110. Rodríguez F., Spooner D. Nitrate reductase phylogeny of potato (Solanum sect. Petota) genomes with emphasis on the origins of the polyploid species. Syst. Botany. 2009;34:207-219.

111. Ross H. Potato breeding. Problems and perspectives. Adv. Plant Breed. 1986;13.

112. Sanetomo R., Akino S., Suzuki N., Hosaka K. Breakdown of a hybridization barrier between Solanum pinnatisectum Dunal and potato using the S locus inhibitor gene (Sli). Euphytica. 2014;197:119-132. DOI 10.1007/s10681-013-1057-1.

113. Schatlowski N., Wolff P., Santos-González J., Schoft V., Siretskiy A., Scott R., Tamaru H., Kohlera C. Hypomethylated pollen bypasses the interploidy hybridization barrier in Arabidopsis. Plant Cell. 2014;26(9):5356-3568. DOI 10.1105/tpc.114.130120.

114. Scotti N., Monti L., Cardi T. Organelle DNA variation in parental Solanum spp. genotypes and nuclear-cytoplasmic interactions in Solanum tuberosum (+) S. commersonii somatic hybrid-backcross progeny. Theor. Appl. Genet. 2003;108(1):87-94.

115. Simko I., Jansky S., Stephenson S., Spooner D. Genetics of resistance to pests and disease. Ch. 7. Potato Biology and Biotechnology: Advances and Perspectives. Ed. D. Vreugdenhil. Elsevier, 2007.

116. Song J., Bradeen J.M., Naess S.K., Raasch J.A., Wielgus S.M., Haberlach G.T., Liu J., Kuang H., Austin-Phillips S., Buell C.R., Helgeson J.P., Jiang J. Gene RB cloned from Solanum bulbocastanum confers broad spectrum resistance to potato late blight. Proc. Natl. Acad. Sci. USA. 2003;100:9128-9133. DOI 10.1073/pnas.1533501100.

117. Spooner D.M., Alvarez N., Peralta I.E., Clausen A.M. Taxonomy of wild potatoes and their relatives in southern South America (Solanum sects. Petota and Etuberosum). Syst. Bot. Monogr. 2016;100: 1-240.

118. Spooner D.M., Castillo R.T. Reexamination of series relationships of South American wild potatoes (Solanaceae: Solanum sect. Petota): evidence from chloroplast DNA restriction site variation. Am. J. Botany. 1997;84(5):671-685.

119. Spooner D.M., Ghislain M., Simon R., Jansky S.H., Gavrilenko T. Systematics, diversity, genetics, and evolution of wild and cultivated potatoes. Bot. Review. 2014;80(4):283-383. DOI 10.1007/s12229014-9146-y.

120. Spooner D.M., Rodríguez F., Polgár Z., Ballard H.E., Jr., Jansky S.H. Genomic origins of potato polyploids: GBSSI gene sequencing data. Plant Genome: Suppl. to Crop Sci. 2008;48(S1):S27-S36.

121. Spooner D.M., Sytsma K.J. Reexamination of series relationships of Mexican and Central American wild potatoes (Solanum sect. Petota): evidence from chloroplast DNA restriction site variation. Syst. Botany. 1992;17:432-448.

122. Spooner D.M., Sytsma K.J., Conti E. Chloroplast DNA evidence for genome differentiation in wild potatoes (Solanum sect. Petota: Solanaceae). Am. J. Botany. 1991;78:1354-1366.

123. Spooner D.M., van den Berg R.G., Rodríguez A., Bamberg J., Hijmans R.J., Lara-Cabrera S.I. Wild potatoes (Solanum section Petota) of North and Central America. Syst. Bot. Monogr. 2004;68: 1-209.

124. Summers D., Grun P. Reproductive isolation barriers to gene exchange between Solanum chacoense and S. commersonii (Solanaceae). Am. J. Bot. 1981;68:1240-1248.

125. Surikov I.M. Genetics of intraspecific incompatibility of male gametophyte and pistil in flowering plants. Uspekhi sovremennoy genetiki=Progress in Modern Genetics. 1972;4:119-169. (in Russian)

126. Surikov I.M. Nesovmestimost’ i embrional’naya steril’nost’ rasteniy [Incompatibility and embryonal sterility of plants]. Moscow, Agropromisdat Publ., 1991 (in Russian)

127. Tek A.L., Stevenson W.R., Helgeson J.P., Jiang J. Transfer of tuber soft rot and early blight resistances from Solanum brevidens into cultivated potato. Theor. Appl. Genet. 2004;109:249-254.

128. Thieme R., Rakosy-Tican E., Gavrilenko T., Antonova O., Schubert J., Nachtigall M., Heimbach U., Thieme T. Novel somatic hybrids (Solanum tuberosum L. + Solanum tarnii) and their fertile BC1 progenies express extreme resistance to potato virus Y and late blight. Theor. Appl. Genet. 2008;116:691-700. DOI 10.1007/s00122-0070702-2.

129. Thieme R., Rakosy-Tican E., Nachtigall M., Schubert J., Hammann T., Antonova O., Gavrilenko T., Heimbach U., Thieme T. Characterization of the multiple resistance traits of somatic hybrids between Solanum cardiophyllum Lindl. and two commercial potato cultivars. Plant Cell Rep. 2010;29:1187-1201. DOI 10.1007/s00299-010-0905-x.

130. Thieme R., Schubert J., Nachtigall M., Hammann T., Truberg B., Heimbach U., Thieme T. Wild potato species of the series Pinnatisecta – progress in their utilisation in potato breeding. Crop Plant Resistance to Biotic and Abiotic Factors: Current Potential and Future Demands: Proc. of the 3rd Intern. Symp. on Plant Protection and Plant Health in Europe, 14–16 May 2009, Berlin, Germany. Braunschweig, 2009;428-436.

131. Thompson R.D., Uhrig H., Hermsen J.G.Th., Salamini F., Kaufmann H. Investigation of a self-compatible mutation in Solanum tuberosum clones inhibiting S-allele activity in pollen differentially. Mol. Gen. Genet. 1991;226:283-288.

132. Tonosaki K., Osabe K., Kawanabe T., Fujimoto R. The importance of reproductive barriers and the effect of allopolyploidization on crop breeding. Breed. Sci. 2016;66(3):333-349. DOI 10.1270/jsbbs.15114.

133. Trognitz B.R., Schmidiche P.E. A new look at incompatibility relationships in higher plants. Sex Plant Reprod. 1993;6:183-190.

134. Veilleux R.E., De Jong H. Potato. Genetic Resources, Chromosome Engineering, and Crop Improvement. Vol. 3. Ed. R.J. Singh. Boca Raton: CRC Press, 2007;17-58.

135. Voronkova E.V., Lisovskaya V.M., Pavlyuchuk N.V., Savchuk A.V., Yermishin A.P. Resistance to virus diseases in the progeny of diploid interspecific hybrids derived from allotetraploid species Solanum acaule and S. stoloniferum. Kartofelevodstvo = Potato Growing (Minsk). 2008;14:144-152. (in Russian)

136. Voronkova E.V., Lisovskaya V.M., Yermishin A.P. Diploid hybrids between allotetraploid wild potato species Solanum acaule Bitt., Solanumstoloniferum Schltdl. and dihaploids of Solanum tuberosum L. Genetika = Genetics (Moscow). 2007;43(8):882-889. (in Russian)

137. Voronkova E.V., Pavlyuchuk N.V., Polykhovich Yu.V., Gukasyan O.N., Zharich V.M., Luksha V.I., Yermishin A.P. Resistance to PVY transferred to diploid interspecific hybrids from Solanum stoloniferum. Kartofelevodstvo = Potato Growing (Minsk). 2015;23:27-39. (in Russian)

138. Waara S., Glimelius К. The potential of somatic hybridization in crop breeding. Euphytica. 1995;85:217-233.

139. Watanabe K., Arbizu C., Schmiediche P. Potato germplasm enhancement with disomic tetraploid Solanum acaule. I. Efficiency of introgression. Genome. 1992;35:53-57.

140. Werner J.E., Peloquin S.J. Inheritance and two mechanisms of 2n egg formation in 2x potatoes. J. Heredity. 1990;81:371-374.

141. Xiao W., Brown R.C., Lemmon B.E., Harada J.J., Goldberg R.B., Fischer R.L. Regulation of seed size by hypomethylation of maternal and paternal genomes. Plant Physiol. 2006;142:1160-1168.

142. Xu B.B., Mu J.H., Nevins D.L., Grun P., Kao T.H. Cloning and sequencing of cDNAs encoding two self-incompatibility associated proteins in Solanum chacoense. Mol. Gen. Genet. 1990;224:341-346.

143. Yermishin A.P., Polyukhovich Yu.V., Voronkova E.V., Gukasyan O.N. SvSv lines, an effective tool for involvement of valuable germplasm from 1EBN diploid potato species. Vavilovskii Zhurnal Genetiki i Selektsii = Vavilov Journal of Genetics and Breeding. 2017;21(1):4250. DOI 10.18699/VJ17.222. (in Russian)

144. Yermishin A.P., Polyukhovich Y.V., Voronkova E.V., Savchuk A.V. Production of hybrids between 2 EBN bridge species Solanum verrucosum and 1 EBN diploid potato species. Am. J. Potato Res. 2014; 91(6):610-617. DOI 10/1007/s12230-014-9385-9/.

145. Yermishin A.P., Voronkova E.V., Kozlov V.A. Kartofel [Potato]. Kilchevskiy A.V., Khotyleva L.V. (Eds.) Geneticheskie osnovy selektsii rasteniy. Tom 2. Chastnaya genetika rasteniy [Genetic basis of plant breeding. V. 2. Specific genetics]. Minsk, 2010;156-234. (in Russian)