Studying grain color diversity in the barley collection of VIR

Background. Dark color of barley grain (Hordeum vulgare L.) can be caused by the synthesis and accumulation of two types of polyphenolic pigments – anthocyanins and melanins, which perform important functions in plant life, participating in the regulation of growth and development, and protecting plants from adverse environmental conditions. The aim of this study was to investigate the diversity of barley in the VIR collection in the context of grain color.

Materials and methods. To analyze the pigment composition of the grain, 150 barley accessions with colored grains were selected from the VIR collection. Anthocyanins and melanins in grain husk were identified using qualitative reactions.

Results and discussion. It was shown that in 60% of the accessions the dark color of their grain was induced by independent accumulation of melanin, while the accessions characterized by accumulation of only anthocyanins, and those with combined accumulation of anthocyanins and melanins, were 14.6% and 14%, respectively. For 11.3% of the accessions the presence of anthocyanins and melanins in grain husk was not found; their pigmentation could presumably be associated with an increased content of other polyphenolic pigments – proanthocyanidins. Accessions with melanin in grain predominated in all identified geographic groups, while other types of pigmentation were most evenly represented in the regions with the widest genetic diversity of barleys – Africa, East Asia, and the Middle East.

Conclusion. Dark pigmentation of barley grain was shown to be mainly associated with the accumulation of melanin, and this type of pigmentation prevails in all geographical regions identified. The results obtained made it possible to describe the barley collection more fully and expand the possibilities of its utilization.

1. Aastrup S., Outtrup H., Erdal K. Location of the proanthocyanidins in the barley grain. Carlsberg Research Communications. 1984;49:105-109. DOI: 10.1007/BF02913969

2. Abdel-Aal E.S.M., Hucl P. A rapid method for quantifying total anthocyanins in blue aleurone and purple pericarp wheats. Cereal Chemistry. 1999;76(3):350-354. DOI:10.1094/CCHEM.1999.76.3.350

3. Badr A., Sch K.M.R., El Rabey H., Effgen S., Ibrahim H.H., Pozzi C. et al. On the origin and domestication history of barley (Hordeum vulgare). Molecular Biology and Evolution. 2000;17(4):499-510. DOI: 10.1093/oxfordjournals.molbev.a026330

4. Bishaw Z., Struik P.C., van Gastel A.J.G. Wheat and barley seed system in Syria: farmers, varietal perceptions, seed sources and seed management. International Journal of Plant Production. 2011;5(4):323–348. DOI: 10.22069/IJPP.2012.744

5. Choo A., Vigier B., Ho K.M., Ceccarelli S., Grando S., Franckowiak J.D. Comparison of black, purple, and yellow barleys. Genetic Resources and Crop Evolution. 2005;52(2):121-126. DOI: 10.1007/s10722-003-3086-4

6. Choo T.M. Genetic resources of Tibetan barley in China. Crop Science. 2002;42(5):1759-1760. DOI: 10.2135/cropsci2002.1759

7. Dai F., Chen Z.H., Wang X., Li Z., Jin G., Wu D. et al. Transcriptome profiling reveals mosaic genomic origins of modern cultivated barley. Proceedings of the National Academy of Sciences of the United States of America. 2014;111(37):13403-13408. DOI: 10.1073/pnas.1414335111

8. Debeaujon I., Léon-Kloosterziel K.M., Koornneef M. Influence of the testa on seed dormancy, germination, and longevity in Arabidopsis. Plant Physiology. 2000;122(2):403-414. DOI: 10.1104/pp.122.2.403

9. Di Ferdinando M., Brunetti C., Fini A., Tattini M. Flavonoids as antioxidants in plants under abiotic stresses. In: P. Ahmad, M. Prasad (eds). Abiotic Stress Responses in Plants. New York, NY: Springer; 2012. p.159-179. DOI: 10.1007/978-1-4614-0634-1_9

10. Downie A.B., Zhang D., Dirk L.M.A., Thacker R.R., Pfeiffer J.A., Drake J.L. et al. Communication between the maternal testa and the embryo and/or endosperm affect testa attributes in tomato. Plant Physiology. 2003;133(1):145-160. DOI: 10.1104/pp.103.022632

11. Glagoleva A., Kukoeva T., Mursalimov S., Khlestkina E., Shoeva O. Effects of combining the genes controlling anthocyanin and melanin synthesis in the barley grain on pigment accumulation and plant development. Agronomy. 2022;12(1):112. DOI: 10.3390/agronomy12010112

12. Glagoleva A.Y., Shoeva O.Y., Khlestkina E.K. Melanin pigment in plants: current knowledge and future perspectives. Frontiers in Plant Science. 2020;11:770. DOI: 10.3389/fpls.2020.00770

13. Hosoda K., Sasahara H., Matsushita K., Tamura Y., Miyaji M., Matsuyama H. Anthocyanin and proanthocyanidin contents, antioxidant activity, and in situ degradability of black and red rice grains. Asian­Australasian Journal of Animal Sciences. 2018;31(8):1213-1220. DOI: 10.5713/AJAS.17.0655

14. Landi M., Tattini M., Gould K.S. Multiple functional roles of anthocyanins in plant–environment interactions. Environmental and Experimental Botany. 2015;119:4-17. DOI: 10.1016/J.ENVEXPBOT.2015.05.012

15. Loskutov I.G., Blinova E.V., Gavrilova O.P., Gagkaeva T.Yu. The valuable characteristics and resistance to Fusarium disease of oat genotypes. Russian Journal of Genetics: Applied Research. 2017;7(20/3):290-298. DOI: 10.1134/S2079059717030108

16. Lukyanova M.V., Trofimovskaya A.Y., Gudkova G.N., Terentieva I.A., Jarosh N.P. Cultivated Flora of the USSR. Vol. 2, Pt 2. Barley (Yachmen). Leningrad: Agropromizdat; 1990. [in Russian]

17. Mascher M., Schuenemann V.J., Davidovich U., Marom N., Himmelbach A., Hübner S. et al. Genomic analysis of 6,000-year-old cultivated grain illuminates the domestication history of barley. Nature Genetics. 2016;48(9):1089-1093. DOI: 10.1038/ng.3611

18. Nicolas J.J., Richard-Forget F.C., Goupy P.M., Amiot M., Aubert S.Y. Enzymatic browning reactions in apple and apple products. Critical Reviews in Food Science and Nutri­ tion. 1994;34(2):109-157. DOI: 10.1080/10408399409527653

19. Shoeva O.Yu., Mock H.P., Kukoeva T.V., Börner A., Khlestkina E.K. Regulation of the flavonoid biosynthesis path-way genes in purple and black grains of Hordeum vulgare. PloS One. 2016;11(10):e0163782. DOI: 10.1371/journal.pone.0163782

20. Shoeva O.Yu., Mursalimov S.R., Gracheva N.V., Glagoleva A.Y., Börner A., Khlestkina E.K. Melanin formation in barley grain occurs within plastids of pericarp and husk cells. Scientific Reports. 2020;10(1):179. DOI: 10.1038/s41598-019-56982-y

21. Shoeva O.Yu., Strygina K.V., Khlestkina E.K. Genes determining the synthesis of flavonoid and melanin pigments in barley. Vavilov Journal of Genetics and Breeding. 2018;22(3):333-342. [in Russian] DOI: 10.18699/VJ18.369

22. Solano F. Melanin and melanin-related polymers as materials with biomedical and biotechnological applications – Cuttlefish ink and mussel foot proteins as inspired biomolecules. International Journal of Molecular Sciences. 2017;18(7):1561. DOI: 10.3390/ijms18071561

23. Vavilov N.I. Selected works in two volumes. Vol. I (Izbrannye proizvedeniya v dvukh tomakh. T. I). St. Petersburg: Nauka; 1967. [in Russian]

24. Wang Y., Ren X., Sun D., Sun G. Origin of worldwide cultivated barley revealed by NAM-1 gene and grain protein content. Frontiers in Plant Science. 2015;6:803. DOI: 10.3389/fpls.2015.00803

25. Weltzien E. Evaluation of barley (Hordeum vulgare L.) landrace populations originating from different growing regions in the Near East. Plant Breeding. 1998;101(2):95-106. DOI: 10.1111/j.1439-0523.1988.tb00273.x

26. Yudina R.S., Gordeeva E.I., Shoeva O.Yu., Tikhonova M.A., Khlestkina E.K. Anthocyanins as functional food components. Vavilov Journal of Genetics and Breeding. 2021;25(2):178-189. [in Russian] DOI: 10.18699/VJ21.022