ИЗМЕНЧИВОСТЬ РАСОВОГО СОСТАВА ЗАРАЗИХИ ПОДСОЛНЕЧНИКОВОЙ (ОROBANCHE CUMANA WALLR.) И НОВЫЕ СТРАТЕГИИ ЗАЩИТЫ ОТ ПАРАЗИТА

Заразиха подсолнечниковая – паразитическое бесхлорофилльное растение, поражающее корневую систему растения-хозяина, поглощающее из нее воду, питательные вещества и выделяющее токсичные продукты обмена. Прорастание семян заразихи происходит благодаря стриголактонам, выделяемым в почву корнями подсолнечника, которые привлекают арбускулярные  микоризные грибы (АМ-грибы). Стриголактоны являются веществами «голода» растений и относятся к новому классу фитогормонов, участвующих во многих физиологических процессах, в том числе в регулировании доступности питательных веществ корнями. У заразихи был идентифицирован специфичный рецептор KARRIKIN INSENSITIVE2 DIVERGENT (KAI2d), который участвует в дифференцированном распознавании корневых экссудатов подсолнечни - ка. В геноме паразита установлено несколько генов KAI2d, кодирующих рецепторы KAI2d. В связи с открытием перечисленных веществ обсуждаются новые стратегии защиты подсолнечника от заразихи, которые находятся на стадии разработки. На основе стриголактонов создан синтетический препарат GR24 и его аналоги с целью внесения их в почву для стимулирования прорастания семян заразихи. Аминокислота метионин сильно ингибирует раннее развитие заразихи без фитотоксичного эффекта у подсолнечника. Семена заразихи хуже распознают корни подсолнечника, которые колонизированы АМ-грибами, бактериями Rhizobium leguminosarum, Azospirillum brasilense, из-за изменения состава корневых экссудатов в колонизированных растениях.  Большой интерес среди синтезированных элиситоров представляет ацибензолар-S-метил. Он вызывает системную устойчивость к бактери альным, грибным и вирусным болезням, индуцирует синтез PR-белков, а также активизирует защитные реакции подсолнечника на внедрение заразихи и приводит к одревеснению эндодермы и торможению проникновения гаусторий семян заразихи через клеточную стенку клеток корня.

1. Albrecht H., Yoder J.I., Phillips D.A. Flavonoids promote haustoria formation in the root parasite Triphysaria versicolor. Plant Physiol. 1999;119:585-591.

2. Antonova T.S. The development of broomrape haustoria in roots of immune and susceptible sunflower varieties. Botanicheskiy zhurnal = Botanical Journal. 1978;7:1025-1029. (in Russian)

3. Antonova T.S., Araslanova N.M., Ramazanova S.A., Guchetl’ S.Z., Chelyustnikova T.A. Virulence of the broomrape affecting sunflower in the Volgograd and Rostov regions. Maslichnye kultury. Nauchnotekhnicheskiy byulleten Vserossiyskogo NII maslichnyh kultur = Oilseeds. Scientific and Technical Bulletin of the All-Russian Research Institute of Oilseeds. 2011;1:127-130. (in Russian)

4. Araslanova N.M., Antonova T.S., Ramazanova S.A., Guchetl’ S.Z., Chelyustnikova T.A. Orobanche cumana Wallr. seed germination under the influence of root excretions from nonhost species. Maslichnye kultury. Nauchno-tekhnicheskiy byulleten Vserossiyskogo NII maslichnyh kultur = Oilseeds. Scientific and Technical Bulletin of the All-Russian Research Institute of Oilseeds. 2011;1:130-134. (in Russian)

5. Bartsinsky R.M. On the physiology of the germination of seeds of the “evil” Don broomrape race. Maslichnye kultury = Oilseeds. 1932;2:42-47. (in Russian)

6. Beilin I.G. On the relationship between O. cumana and sunflower. Rastenie i sreda = Plant and Еnvironment. 1940;1:175-192. (in Russian)

7. Beilin I.G. Tsvetkovye poluparazity i parazity [Flower semiparasites and parasites]. Moscow: Nauka Publ., 1968. (in Russian)

8. Bigeard J., Colcombet J., Hirt H. Signaling mechanisms in Pattern-Triggered Immunity (PTI). Mol. Plant. 2015;8(4):521-539.

9. Bonfante P., Genre A. Plants and arbuscular mycorrhizal fungi: an evolutionary-developmental perspective. Trends Plant. 2008;13:492-498.

10. Bouwmeester H.J., Roux C., Lopez-Raez J.A., Bécard G. Rhizosphere communication of plants, parasitic plants and AM fungi. Trends Plant Sci. 2007;12:224-230.

11. Brewer P.B., Koltai H., Beveridge C.A. Diverse roles of strigolactones in plant development. Mol. Plant. 2013;6:18-28.

12. Burlov V.V., Burlov V.V. Efficiency of org genes in providing sustainability of sunflower to new broomrape races (Orobanche cumana Wallr.). Selektsіya і nasіnnitstvo = Selection and Seed Production. 2010;98:28-37. (in Ukrainian)

13. Burlov V.V., Kostyuk S.V. Inheritance of resistance to the local race of broomrape (Orobanche cumana Wallr.) in sunflower. Genetika = Genetics (Moscow). 1976;12(2):44-51. (in Russian)

14. Conn C.E., Bythell-Douglas R., Neumann D., Yoshida S., Whittington B., Westwood J.H., Shirasu K., Bond C.S., Dyer K.A., Nelson D.C. Convergent evolution of strigolactone perception enabled host detection in parasitic plants. Science. 2015;349:540-543.

15. East R. Soil science comes to life. Nature. 2013;501:8-19.

16. Evidente A., Andolfi A., Fiore M., Boari A., Vurro M. Stimulation of Orobanche ramosa seed germination by fusicoccin derivatives: a structure-activity relationship study. Phytochemistry. 2006;67:19-26.

17. Fernández-Aparicio M., Pérez-de-Luque A., Prats E., Rubiales D. Variability of interactions between barrel medic (Medicago trun catula) genotypes and Orobanche species. Ann. Appl. Biol. 2008;153: 117-126.

18. Fernández-Martínez J.M., Domínguez J., Pérez-Vich B. Update on breeding for resistance to sunflower broomrape. Helia. 2008;31: 73-84.

19. Gomez-Roldan V., Fermas S., Brewer P.B., Puech-Pagѐs V., Dun E.A., Pillot J.P., Letisse F., Matusova R., Danoun S., Portais J.-C., Bouwmeester H., Bécard G., Beveridge C.A., Rameau C., Rochange S.F. Strigolactone inhibition of shoot branching. Nature. 2008;455:189-194.

20. Goncharov S.V., Antonova T.S., Araslanova N.M., Ryzhenko Ye.N. Breeding of sunflower hybrids resistant to new broomrape races. Maslichnye kultury. Nauchno-tekhnicheskiy byulleten Vserossiyskogo NII maslichnyh kultur = Oilseeds. Scientific and Technical Bulletin of the All-Russian Research Institute of Oilseeds. 2012;1:9-12. (in Russian)

21. Gonsior G., Buschmann H., Szinicz G., Spring O., Sauerborn J. Induced resistance – an innovative approach to manage branched broomrape (Orobanche ramosa) in hemp and tobacco. Weed Sci. 2004;52:1050-1053.

22. Guchetl’ S.Z., Chelyustnikova T.A., Araslanov N.M., Antonova T.S. Tagging of the Or5 gene for resistance to the E race of broomrape Orobanche cumana Wallr. in the lines of sunflower bred at the AllRussia Research Institute of Oilseeds. Maslichnye kultury. Nauchno-tekhnicheskiy byulleten Vserossiyskogo NII maslichnyh kultur = Oilseeds. Scientific and Technical Bulletin of the All-Russian Research Institute of Oilseeds. 2012;2:157-163. (in Russian)

23. Hallmann J. Plant Interactions with Endophytic Bacteria. In: M.J. Jeger, N.J. Spence (Eds.). Biotic Interactions in Plant-Pathogen Associations. N. Y.: CABI Publ., 2001; 87-119.

24. Hemissi I., Mabrouk Y., Abdi N., Bouraoui M., Saidi M., Sifi B. Growth promotion and protection against Orobanche foetida of chickpea (Cicer aerietinum) by two Rhizobium strains under greenhouse conditions. Afr. J. Biotechnol. 2013;12:1371-1377.

25. Kaya Y., Evci Y., Pekcan V., Gucer T. Determining new broomrape infested areas, resistant lines and hybrids in Trakya region of Turke. Helia. 2004;27:211-218.

26. Khatnyansky V.I. The inheritance of sunflower resistance to infestation with a new complex of broomrape races. Byulleten nauchno-tekhnicheskoy informatsii VNIIMK = Scientific and Technical Bulletin of the All-Russia Research Institute of Oilseeds. 1982;2:3-5. (in Russian)

27. Kirichenko A. Offer: biological methods for controlling plant viruses. Bioprotection and Biopreparations are a Fair Promise. 2017;42-46. (in Ukrainian)

28. Koch L. Die Entwicklungs gechichte der Orobanchen mit besonderer Berücksichtung ihrer Beziehungen zu den Kulturpflanzen. Heidelberg, 1887.

29. Kusumoto D., Goldwasser Y., Xie X., Yoneyama K., Takeuchi Y. Resistance of red clover (Trifolium pratense) to the root parasitic plant Orobanche minor is activated by salicylate but not by jasmonate. Ann. Bot. 2007;100:537-544.

30. López-Ráez J.A., Charnikhova T., Fernandez I., Bouwmeester H., Pozo M.J. Arbuscular mycorrhizal symbiosis decreases strigolactone production in tomato. J. Plant Physiol. 2011;168:294-297.

31. Louarn J., Carbonne F., Delavault P., Bécard G., Rochange S. Reduced germination of Orobanche cumana seeds in the presence of Arbuscular Mycorrhizal fungi or their exudates. PLoS One. 2012;7(11):1-10.

32. Lugtenberg B., Kamilova F. Plant-growth-promoting rhizobacteria. Annu Rev. Microbiol. 2009;63:541-556.

33. Ma Y., Jia J.N., An Y., Wang Z., Mao J.C. Potential of some hybrid maize lines to induce germination of sunflower broomrape. Crop Sci. 2012;53:260-270.

34. Ma Y., Shui J., Inanaga S., Cheng J. Stimulatory effects of Houttuynia cordata Thunb. on seed germiation of Striga hermonthica (Del.) Benth. Allelopathy J. 2005;15:49-56.

35. Melero-Vara J.M., Dominguez J., Fernandez-Martinez J.M. Update on sunflower broomrape situation in Spain: racial status and sunflower breeding for resistance. Helia. 2000;23:45-55.

36. Metzger J. The promotion of germination of dormant weed seeds by substituted phthalimides and gibberellic acid. Weed Sci. 1983;31: 285-289.

37. Mwakaboko A.S., Zwanenburg B. Strigolactone analogs derived from ketones using a working model for germination stimulants as a blueprint. Plant Cell Physiol. 2011;52:699-715.

38. Nichiporovich A.A. On the physiology of the Don broomrape. Izvestiya po opytnomu delu Dona i Severnogo Kavkaza = Izvestiya on the Trial Case of the Don and the North Caucasus. 1929;15:237-247. (in Russian)

39. Pieterse C.M.J., Zamioudis C., Berendsen R.L., Weller D.M., Van Wees S.C.M., Bakker P. Induced systemic resistance by beneficial microbes. Ann. Rev. Phytopathol. 2014;52:347-375.

40. Plachek E.M. Sunflower immunity to broomrape infestation. Izvestiya Saratovskoy oblastnoy sel’skohozyaystvennoy opytnoy stantsii = Proceedings of the Saratov Regional Agricultural Academy. Experimental Station. 1921;3:65-82. (in Russian)

41. Pushkareva K.V. Characterization of seeds of different biological forms of broomrape. Byulleten’ Severno-Kavkazskoy kraevoy sel’skohozyaystvennoy opytnoy stantsii = Bulletin of the North Caucasus Agricultural Experiment Station. 1930;306:12-16. (in Russian)

42. Pustovoit V.S. Izbrannye trudy: Selektsiya. Semenovodstvo i nekotorye voprosy agrotekhniki podsolnechnika. [Selected Works: Breeding. Seed industry and some issues of sunflower farming]. Moscow: Kolos Publ., 1966. (in Russian)

43. Pustovoit V.S., Pustovoit G.V. Breeding of sunflower for resistance to broomrape. Zashchita rasteniy ot vrediteley i bolezney = Protection of Plants from Pests and Diseases. 1963;4:15-17. (in Russian)

44. Ranf S., Eschen-Lippold L., Pecher P., Lee J., Scheel D. Interplay between calcium signalling and early signalling elements during defence responses to microbe- or damage-associated molecular patterns. Plant J. 2011;68(1):100-113.

45. Reinhardt D. Programming good relations: development of the arbuscular mycorrhizal symbiosis. Curr. Opin. Plant Biol. 2007;10:98-105.

46. Reshetnyak N.V., Kosogova T.M., Hansiy Yu.A., Legkhoduh A.A., Fursov V.N., Malykhin I.I. Orobanche cumana Wallr. in sunflower plantings and its control. Naukoviy vіsnik Lugans’kogo natsіonalnogo agrarnogo unіversitetu = Scientific Herald of the Lugansk National Agrarian University. 2012;36:107-110. (in Ukrainian)

47. Richter A.A. On the physiology of broomrape infesting sunflower. Uchenye Zapiski Saratovskogo gos. un-ta. [Scientific Notes of the Saratov State University]. 1924;2:33-42. (in Russian)

48. Sands D.C., Pilgeram A.L. Methods for selecting hypervirulent biocontrol agents of weeds: why and how? Pest Manag. Sci. 2009;65: 581-587.

49. Shindrova P. Broomrape (Orobanche cumana Wallr.) in Bulgaria distribution and race composition. Helia. 2006;29:111-120.

50. Shreiner Ya.F. Podsolnechnaya zarazikha (Orobanche cumana Wallr.) i sposoby bor’by s neyu [Sunflower broomrape (Orobanche cumana Wallr.) аnd methods to control it]. St. Petersburg, 1904. (in Russian)

51. Thorogood C.J., Hi S.J. Compatibility interactions at the cellular level provide the basis for host specificity in the parasitic plant Orobanche. New Phytol. 2010;186:571-575.

52. Ukrainskiy V.T. Sunflower infestation with broomrape and measures to combat it. Selektsiya i semenovodstvo = Breeding and Seed Industry. 1938;1:36-43. (in Russian)

53. Velasco L., Pérez-Vich B., Jan C.C., Fernández-Martínez J.M. Inheritance of resistance to broomrape (Orobanche cumana Wallr.) race F in a sunflower line derived from wild sunflower species. Plant Breed. 2007;126:67-71.

54. Vurro M., Boari A., Evidente A., Andolfi A., Zermane N. Natural metabolites for parasitic weed management. Pest Manag. Sci. 2009;65: 566-571.

55. Waters M.T., Scaffidi A., Moulin S.L., Sun Y.K., Flematti G.R., Smith S.M. A Selaginella moellendorffii ortholog of KARRIKIN INSENSITIVE2 functions in Arabidopsis development but cannot mediate responses to karrikins or strigolactones. Plant Cell. 2015; 27:1925-1944.

56. Yoneyama K., Xie X., Kisugi T., Nomura T., Sekimoto H., Yokota T., Yoneyama K. Characterization of strigolactones exuded by Asteraceae plants. Plant Growth Regul. 2011;65:495-504.

57. Zadorozhna O.A. Molecular marker-assisted identification of the Or5 gene for broomrape resistance in sunflower. Agrarniy vіsnik Prichornomor’ya = Agrarian Bulletin of the Black Sea Region. 2012;61:23-27. (in Ukrainian)