Analysis of factors inducing the transition to the generative phase of development in the Brassicaceae family (review)

   Relevance. In the global crop production, white cabbage (Brassica oleracea L. var. capitata L.) is one of the most important vegetable crops. One of the priorities of domestic cabbage crop breeding is the creation of highly productive hybrids. The most time-consuming and lengthy stage in this process is the creation of constant parent lines.

   The purpose of this review was a systematic analysis of scientific publications devoted to the process of transition from the vegetative to the generative phase of development in representatives of the Cabbage family (Brassicaceae).

   Results. In traditional breeding, inbred lines of white cabbage are obtained through forced self-pollination for 6-12 generations. The use of the biotechnological method of culture of isolated microspores in vitro ensures the achievement of complete homozygosity of the genome during one generative cycle and contributes to the expansion of the spectrum of formation of genetic recombinant forms, including those with recessive traits. Also, in combination with the technology of obtaining doubled haploids and genomic approaches such as marker breeding, genomic selection and genome editing, "Speed Breeding" allows to increase the efficiency of breeding by reducing the breeding cycle time, providing early phenotypic assessment, efficient use of resources, increasing the accuracy of selection and genetic progress. Despite the advantages of biotechnological methods, the use of double haploid (DH) lines is associated with a number of practical difficulties, in particular, concerning the reproduction of the obtained plants. An additional problem is the lack of flowering in some DH plants after undergoing vernalization, which may be due to a number of reasons: unsuitable vernalization conditions for specific lines. There are also plants with normal vegetative development, which, nevertheless, do not set seeds.

   Conclusion. As a result of the conducted review of world and domestic practice, important stages of vernalization have been studied: temperature, duration, lighting, and stages of plant development. This knowledge will make it possible to increase the percentage of plants passing into the reproductive stage of development.

1. Soldatenko A.V., Ivanova M.I., Bondareva L.L., Tareeva M.M. Cabbage green vegetables. M., 2022. 296 p. (In Russ.) ISBN 978-5-901695-89-0. https://www.elibrary.ru/unsafi

2. Tons M.M. World Cabbage Production, 2015-2019.

3. Pivovarov V.F., Startsev V.I. Cabbage, its species and varieties (variety and methods of cultivation). M.: VNIISSOK. 2006. 192 p. https://www.elibrary.ru/qkxxur

4. Taiz L., Zeiger E. Plant Physiology. Sunderland: Sinauer Associates Inc. 2010.

5. Dunwell J.M. Haploids in flowering plants: origins and exploitation. J. Plant Biotech. 2010;8:377-424. doi: 10.1111/j.1467-7652.2009.00498.x

6. Mineykina A.I., Bondareva L.L., Shumilina D.V., Domblides E.A., Soldatenko A.V. Improvement of methods of creating hybrids of cabbage. Vegetable crops of Russia. 2019;(4):3-7. (In Russ.) doi: 10.18619/2072-9146-2019-4-3-7 https://www.elibrary.ru/faxajs6.

7. Mineykina A.I., Stebnitskaia K.S., Fomicheva M.G., Bondareva L.L., Domblides A.S., Domblides E.A. Optimization of technology steps for obtaining white cabbage dh-plants. Vavilov journal of genetics and breeding. 2025;29(4):517-529. doi: 10.18699/vjgb-25-55 https://www.elibrary.ru/gzdtco

8. Monakhos S.G. Creation of pure lines – doubled haploids of cabbage in the culture of isolated microspores and selection of F₁ hybrids based on modern biotechnology methods : methodological recommendations. М.: Publishing house RGAU - Moscow Agricultural Academy named after K.A. Timiryazev. 2014. 44 р. (In Russ.)

9. Nikitin M.A. The use of doubled haploid technology in modern cabbage crop breeding. Natural sciences. 2024;(4). (In Russ.)

10. Vishnyakova A.V., Alexandrova A. A., Monakhos S. G. Factors of direct germination of microsporogenic embryoids Brassica napus L. Izvestiya of Timiryazev Agricultural Academy. 2022;(6):43-53. (In Russ.) doi: 10.26897/0021-342Х-2022-6-43-53 https://www.elibrary.ru/bifbsl

11. Sinicyna A.A., Vishnyakova A.V., Monakhos S.G. Comparative assessment of the yield of doubled haploids of Brassica oleracea var. capitata L. and Brassica napus L. in isolated microspore culture. Potato and vegetables. 2022;(4):37-40. (In Russ.) doi: 10.25630/PAV.2022.29.31.008 https://www.elibrary.ru/hafnfc

12. Ćeran M., Miladinović D., Đorđević V., et al. Genomics-assisted speed breeding for crop improvement: present and future. Front. Sustain. Food Syst. 2024;(8):1383302. doi: 10.3389/fsufs.2024.1383302

13. Zablotskaya E.A., Mineykina A.I., Domblides E.A., Paslova T.O., Bondareva L.L. Broccoli various genotypes regenerated plants (R0) seed set after geitonogamy. Vegetable crops of Russia. 2020;(2):43-46. (In Russ.) doi: 10.18619/2072-9146-2020-2-43-46 https://www.elibrary.ru/hakswx

14. Suge H. Re- examination on the role of vernalization and photoperiod in the flowering of Brassica crops under controlled environment. Jpn. J. Breeding. 1984;(34):171-180.

15. Osadchaya T.S., Trubacheeva N.V., Kravtsova L.A., Belan I.A., Rosseeva L.P., Pershina L.A.. Study of fertility and cytogenetic variability in androgenic plants (R0 and R1) of alloplasmic introgression lines of common wheat. Vavilov Journal of Genetics and Breeding. 2016;20(3):370-377. (In Russ.) doi: 10.18699/VJ16.165 https://www.elibrary.ru/wlvkih

16. Deimling S., Flehinghaus-Roux T. Haploidy in rye. In: Mohan Jain S., Sopory S.K., Veilleux R.E. (eds) In vitro haploid production in higher plants: Volume 4: Cereals. Kluwer Academic Publ, Dordrecht, The Netherlands, 1997. P. 181–204.

17. Guo Y., Pulli S. Isolated microspore culture and plant regeneration in rye (Secale cereale L.). Plant Cell Reports. 2000;(19):875–880. doi: 10.1007/s002990000194

18. Lashermes P., Couturon E., Charrier A. Doubled haploids of Coffea canephora: development, fertility and agronomic characteristics. Euphytica. 1993;(74):149–157. doi: 10.1007/BF00033781

19. Stipic M., Campion, B. An improved protocol for androgenesis in cauliflowers (Brassica oleracea var. botrytis). J. Plant Breeding. 1997;116(2):153-157.

20. Chauvin J.E., Yang Q., LeJeune B., Herve Y. Obtention d'embryons par culture d'antheres chez le chou-fleur et le brocoli et evaluation des potentialites du materiel obtenu pour la creation varietale. Agronomie. 1993;(13):579-590.

21. Kamiński P., Dyki B., Krzyzanowska D., Gуrecka K. Diversity of diploid androgenic Brussels sprout plants of R0 and R1 generations. J. of Applied Genetics. 2005;46(1):25-33.

22. Kaminski P. Gametoclonal and somaclonal variation among head cabbage androgenic lines of R1 and R2 generations obtained from Jaguar F₁ hybrid. J. of Agricultural Science. 2011;2(2):119-128. doi: 10.5539/jas.v2n2p119

23. Zablotskaya E.A., Bondareva L.L., Shmykova N.A. Features of seed formation in double haploid lines of broccoli in different generations. Vegetable crops of Russia. 2016;(4):56-59. (In Russ.) doi: 10.18619/2072-9146-2016-4-56-59 https://www.elibrary.ru/xvrufn

24. Bartenev I.I., Gavrin D.S., Nechaeva O.M., Senyutin A.A. Heterogeneity of the seed plant population and qualitative indicators of sugar beet seeds. Sugar. 2018;(10):46-49. (In Russ.) https://www.elibrary.ru/yoxutr

25. Lizgunova T.V. Cabbage. Vegetable crops and forage roots. L: Ear. 1948. 245 p. (In Russ.)

26. Ahn J.-Y., Subburaj S., Yan F., Yao J., Chandrasekaran A., Ahn K.-G., Lee G.-J. Molecular Evaluation of the Effects of FLC Homologs and Coordinating Regulators on the Flowering Responses to Vernalization in Cabbage (Brassica oleracea var. capitata) Genotypes. Genes. 2024;(15):154. doi: 10.3390/genes15020154

27. Alessandro M.S., Galmarini C.R.. Inheritance of vernalization requirement in carrot. Journal of the American Society for Horticultural Science. 2007;132:525-529.

28. Miller J.C. Louisiana Copenhagen cabbage: methods of breeding and description. 1934.

29. Vasilevskaya V.K., Lizgunova T.V. Some features of the stage development of cabbage varieties. Proceedings on Applied Botany, Genetics and Breeding. 1951;29(1).

30. Sheen T.F. Cabbage seed production in the subtropics. 1982.

31. Lizgunova T.V. Cultivated flora of the USSR. L: Ear. 1984;9:328. (In Russ.)

32. Voronova A.F., Grinberg E.G., Domanskaya M.K. Biochemical evaluation of a collection of vegetable crops. Scientific and Technical Bulletin of the Siberian Research Institute of Plant Growing and Breeding. 1978;(4). (In Russ.)

33. Gassner G. Beitrage zur physiologischen Characteristik sommerund winterannueller Gewachse, insbesondere der Getreidepflanzen.-Bot. Ztg, 1918, 10.

34. Kuznetsov V.V., Dmitrieva G.A. Plant physiology. М.: Graduate School. 2006. 742 с.

35. Black A., Moot D., Lucas R. Development and growth characteristics of Caucasian and white clover seedlings, compared with perennial ryegrass. Grass and Forage Science. 2006;(61):442-453. doi: 10.1111/j.1365-2494.2006.00553.x

36. Monks D.P., SadatAsilan K., Moot D.J. Cardinal temperatures and thermal time requirements for germination of annual and perennial temperate pasture species. Agronomy New Zealand. 2009;(39):95-110. doi: 10.13140/2.1.1455.4242

37. EL-Eslamboly A.A.S.A., Hamed H.H. New techniques to induce flowering and produce seeds of foreign cabbage varieties as a main step for a superior hybrids production. Int. J. Environ. 2021;10(1):66-82. doi: 10.36632/ije/2021.10.1.6

38. McCormick J.I., Goodger R.A., Chynoweth R.J. Cardinal temperatures and vernalisation requirements for a selection of vegetables for seed production. 2014. P.1-83.

39. Elers B., Wiebe H.J. Flower formation of Chinese cabbage. I. Response to vernalization and photoperiods. Scientia horticulturae. 1984;22(3):219-231.

40. Kitaeva I. E. Moscow worker. 1977. Р. 57-61. (In Russ.)

41. Lizgunova T.V. Cultivated flora of the USSR.Т. VI. (In Russ.)

42. Shulyak N.V., Koroleva S.V. Response of inbred lines of white cabbage to weather conditions during vernalization. Priority areas of scientific support for the agro-industrial complex of Russia and the CIS countries. Proceedings of the International Scientific and Practical Conference with Elements of the School of Young Scientists. Krasnodar, 2018. pp. 135-141. In Russ.) https://www.elibrary.ru/bcdiok

43. Bondareva L.L., Kolesnikov I.M. Selection of early-ripening hybrids of white cabbage. Scientific works on selection and seed production. 1995;(2):150-152. (In Russ.)

44. Bondareva L.L., Razin O.A. Use of growth chambers for cabbage breeding. Vegetable crops of Russia. 2014;(4):37-39. (In Russ.) doi: 10.18619/2072-9146-2014-4-37-39 https://www.elibrary.ru/tjalbt

45. Bondareva L.L., Mineykina A.I. Accelerated reproduction of the parental lines of white cabbage using rosette plants (steckling) and artificial climate chambers. Vegetable crops of Russia. 2024;(5):26-30. (In Russ.) doi: 10.18619/2072-9146-2024-5-26-30 https://www.elibrary.ru/mpabmg

46. Lizgunova T.V. Fedulova A. P. On the vernalization of cabbage in seeds. Proceedings on Applied Botany, Genetics and Breeding. 1954;31(1). (In Russ.).

47. Michaels S.D., Amasino R.M. Memories of winter: vernalization and the competence to flower. Plant, Cell & Environment. 2000;23(11):1145-1153. doi: 10.1046/j.1365-3040.2000.00643.x

48. Petrishchev A.V. Using container growing of stecklings in hybrid seed production of white cabbage. Moscow. 2006. 156 р. https://www.elibrary.ru/nnusjv (In Russ.)

49. Zvedenyuk A.P., Kazaku V.I. White head cabbage seeds growing from rosette plants. Vegetable crops of Russia. 2013;(3):40-42. (In Russ.) doi: 10.18619/2072-9146-2013-3-40-42 https://www.elibrary.ru/rbjtoj

50. Konysbekov K., Bastaubaeva Sh., Elnazarқyzy R., Tabynbaeva L., Musagodzhaev N. Growing new sugar beet hybrid stecklings in a greenhouse complex. Izdenister Natigeler. 2021;3 (91):103–111. (In Russ.) doi: 10.37884/3-2021/12.

51. Logvinov A.V., Shevchenko A.G., Logvinov V.A., Mishchenko V.N., Koshkin S.S., Moiseev V.V., Batrakova N.V. Peculiarities of technological methods of growing crops-shteklings of mother sugar beet on irrigation. Polythematic online scientific journal of Kuban state agrarian university. 2020;(159):334-347. (In Russ.) doi: 10.21515/1990-4665-159-023. https://www.elibrary.ru/fpwgvk

52. Pivovarov V.F., Breeding and seed production of vegetable crops. М.: VNIISSOK, 2007. 816 р. (In Russ.) https://www.elibrary.ru/qkymcp

53. Bondareva L.L., Frolova S.L. Features of growth and development of seed bushes of varieties and parental lines of cabbage crops. News of FSVC. 2021;(3-4):13-19. (In Russ.) doi: 10.18619/2658-4832-2021-3-4-13-19 https://www.elibrary.ru/reneek

54. Schoen A. et al. Reducing the generation time in winter wheat cultivars using speed breeding. Crop Science. 2023;63(4):2079-2090. doi: 10.1002/csc2.20989

55. Zelenina A.S., Yanovskii A.S., Bizyakina D.O., Nagamova V.M., Rubets V.S., Blinkov A.O., Korobkova V.A., Yurkina A.I., Bespalova L.Yu., Karlov G.I., Divashuk M.G. Methodological recommendations for growing durum wheat under speed breeding conditions to solve breeding problems. Biotechnology in crop production, animal husbandry and agricultural microbiology. 2024. P. 27-28. (In Russ.) doi: 10.48397/l9911-9447-6788-e https://www.elibrary.ru/sjyaeb

56. Watson A. et al. Speed breeding is a powerful tool to accelerate crop research and breeding. 2018;4(1):23-29. doi: 10.1038/s41477-017-0083-8

57. Saifetdinov, E. AThe influence of in vitro vernalization timing on the vegetation of winter wheat and triticale in a phytotron-greenhouse complex. Proceedings of the 24^th Conference of Young Scientists with International Participation, Dedicated to Biotechnology in Plant Production, Animal Husbandry, and Agricultural Microbiology. Moscow: Federal State Budgetary Scientific Institution "All-Russian Research Institute of Agricultural Biotechnology". 2024. Р.116-117. (In Russ.) doi: 10.48397/w4265-2672-9751-v https://www.elibrary.ru/ioklty

58. Popova A.I., Trenikhina M.M. Current issues of modern breeding, biotechnology and botany. Collection of reports of the All-Russian student scientific and practical conference. М.: Russian State Agrarian University. 2024. 184 p. (In Russ.) https://www.elibrary.ru/hkacoz