UV-B Response of Greening Barley Seedlings


The relationship between the greening stage of barley seedlings and their response to UV-B irradiation was studied. Etiolated barley seedlings (Hordeum vulgare L., cv. Alfa) greened 12, 24 and 48 h were exposed to UV-B irradiation (312 nm) for 5 h. As a result of UV-B treatment the rate of CO2 fixation and chlorophyll contents decreased but flavonoids, UV-B-induced compounds and carotenoids increased. The inhibition of photosynthesis in green plants was lower in comparison to greening ones. The 12 h greening plants were more sensitive to UV-B treatment than the plants greening 24 h and particularly 48 h, estimated by the quantum efficiency of PSII photochemistry and the oxygen production rate. The levels of flavonoids and UV-B induced compounds enhanced with increasing the greening time. Activity of antioxidant enzymes catalase, peroxidase and superoxide dismutase increased during the seedlings greening and as a result of UV-B irradiation, but the pattern of isoforms remained similar to those found in the controls. UV-B preferentially induced Cu,Zn-superoxide dismutase. Increase of UVB induced synthesis of antioxidant enzymes is in line with their important role in the plant response to UV-B stress. Data presented show that the response of barley seedlings to UV-B irradiation is related to the development stage of photosynthetic apparatus.


  1. 1.

    Adamse, P., Britz, S. J. (1992) Amelioration of UV-B damage under high irradiance. I. Role of photosynthesis. Photochem. Photobiol. 56, 645–650.

    CAS  Google Scholar 

  2. 2.

    Allen, D. J., McKee, I. F., Farage, P. K., Baker, N. R. (1997) Analysis of the limitation to CO2 assimilation on the exposure of leaves of two Brassica napus cultivars to UV-B. Plant Cell Environ. 20, 633–640.

    CAS  Google Scholar 

  3. 3.

    Bates, L. S., Waldren, R. O., Teare, I. D. (1973) Rapid determination of free proline for water-stress studies. Plant Soi. 39, 205–207.

    CAS  Google Scholar 

  4. 4.

    Caldwell, M. M. (1971) Solar UV irradiation and the growth and development of higher plants. In: Giese, A. C. (ed.) Photophysiology. tAcademic Press, New York, pp. 131–177.

    Google Scholar 

  5. 5.

    Caldwell, M. M., Björn, L. O., Bornman, J. F., Flint, S. D., Kulandaivelu, G., Teramura, A. H., Tevini, M. (1998) Effects of increased solar ultraviolet radiation on terrestrial ecosystems. J. Photochem. Photobiol. B Biol. 46, 40–52.

    CAS  Google Scholar 

  6. 6.

    Davis, B. J. (1964) Disc Electrophoresis-II: Method and application to human serum proteins. Ann. NY Acad. Sci. 15, 404–427.

    Google Scholar 

  7. 7.

    D’Surney, S. J., Tschaplinski, T. J., Edwards, N. T., Shugart, L. R. (1993) Biological responses of two soybean cultivars exposed to enhanced UVB radiation. Environ. Exp. Bot. 33, 347–356.

    Google Scholar 

  8. 8.

    Esterbauer, H., Cheeseman, K. (1990) Determination of aldehyde lipid peroxidation products: mal-onaldehyde and hydroxynonenal. Methods Enzymol. 186, 407–431.

    CAS  PubMed  Google Scholar 

  9. 9.

    Fedina, I., Georgieva, K., Grigorova, I. (2003) Response of barley seedlings to UV-B radiation as affected by proline and NaCl. Biol. Plant. 47, 549–554.

    CAS  Google Scholar 

  10. 10.

    Fedina, I., Velitchkova, M, Georgieva, K., Grigorova, I. (2005) UV-B induced compounds as affected by proline and NaCl in Hordeum vulgare L. cv. Alfa. Environ. Exp. Bot. 54, 182–191.

    CAS  Google Scholar 

  11. 11.

    Foyer, C. R., Lelandais, M., Kunert, K. (1994) Photooxidative stress in plants. Physiol. Plant. 92, 696–717.

    CAS  Google Scholar 

  12. 12.

    Greneche, M., Lallemand, J., Michaud, O. (1991) Comparison of different enzyme loci as a means of distinguishing ryegrass varieties by electrophoresis. Seed Sci. Technol. 19, 147–158.

    Google Scholar 

  13. 13.

    Hada, H., Hidema, J., Maekawa, M., Kumagai, T. (2003) Higher amounts of anthocyanins and UV-absorbing compounds effectively lowered CPD photorepair in purple rice (Oryza sativa L.). Plant CellEnviron. 26, 1691–1701.

    CAS  Google Scholar 

  14. 14.

    He, J., Huang, L.-K., Chow, W. S., Whitecross, M. I., Anderson, J. M. (1993) Effect of supplementary ultraviolet-B radiation on rice and pea plants. Aust. J. Plant Physiol. 20, 129–142.

    Google Scholar 

  15. 15.

    Heath, R. L., Packer, L. (1968) Photoperoxidation in isolated chloroplasts. I. Kinetics and stichiom-etry of fatty acid peroxidation. Arch. Biochem. Biophys. 125, 189–198.

    CAS  PubMed  PubMed Central  Google Scholar 

  16. 16.

    Hidema, J., Kyung Song, I. L., Sato, T., Kumagai, T. (2001) Relationship between ultraviolet-B sensitivity and cyclobutane pyrimidine dimmer photorepair in rice. J. Radiat. Res. 42, 295–303.

    CAS  PubMed  Google Scholar 

  17. 17.

    Jansen, M. A. K., Gaba, V., Greenberg, B. M. (1998) Higher plants and UV-B radiation: balancing damage, repair and acclimation. Trends Plant Sci. 3, 131–135.

    Google Scholar 

  18. 18.

    Jansen, M. A. K., Gaba, V., Greenberg, B. M., Mattoo, A. K., Edelman, M. (1996) Low threshold levels of ultraviolet-B in a background of photosynthetically active radiation trigger rapid degradation of the D2 protein of photosystem II. Plant J. 9, 693–696.

    CAS  Google Scholar 

  19. 19.

    Jordan, B. R. (1996) The effect of ultraviolet-B radiation on plants: a molecular perspective. Adv. Bot. Res. 22, 97–162.

    CAS  Google Scholar 

  20. 20.

    Jordan, B. R., James, P. E., Strid, A., Anthony, R. G. (1994) The effect of ultraviolet-B radiation on gene expression and pigment composition in etiolated and green pea leaf tissue: UV-B-induced changes are gene-specific and dependent upon the developmental stage. Plant Cell Environ. 17, 5–54.

    Google Scholar 

  21. 21.

    Kang, H. S., Hidema, J., Kumagai, T (1998) Effect of light environment during culture on UV-induced cyclobutyll pyrimidine dimmers and their photorepair in Rice (Oriza sativa L.). Photochem.

    Google Scholar 

  22. 22.

    Laemmli, U. K. (1970) Cleavage of structural proteins during the assembly of the head of bacteriophage T4. Natur. 227, 680–685.

    CAS  Google Scholar 

  23. 23.

    Li, J., Ou-Lee, T. M., Raba, R. R. G., Amundson, R. G., Last, R. L. (1993) Arabidopsis flavonoid mutants are hypersensitive to UV-B irradiation. Plant Cel. 5, 171–179.

    CAS  Google Scholar 

  24. 24.

    Lichtenthaler, H. K. (1987) Chlorophylls and carotenoids; pigments of photo synthetic biomembranes. Methods Enzymol. 148, 350–382.

    CAS  Google Scholar 

  25. 25.

    Lowry, O. H., Rosenber, N. J., Farr, A. L., Randal, R. L. (1951) Protein measurement with the Folin phenol reagent. J. Biol. Chem. 193, 265–275.

    CAS  Google Scholar 

  26. 26.

    Mackerness, S. A.-H., Butt, J. P., Jordan, B. R. (1996) Amelioration of ultraviolet-B-induced down regulation of mRNA transcript for chloroplast proteins, by irradiance, is mediated by photosynthesis. J. Plant Physiol. 148, 100–106.

    CAS  Google Scholar 

  27. 27.

    Mackerness, S. A.-H., Surplus, S. L., Jordan, B. R., Thomas, B. (1998) Effects of supplementary UV-B radiation on photo synthetic transcripts at different stages of leaf development and light levels in pea: role of active oxygen species and antioxidant enzymes. Photochem. Photobiol. 68, 88–96.

    CAS  Google Scholar 

  28. 28.

    Mirecki, R. M., Teramura, A. H. (1984) Effects of ultraviolet-B irradiance on soybean. V. The dependence of plant sensitivity on the photosynthetic photon flux density during and after leaf expansion. Plant Physiol. 74, 475–480.

    CAS  PubMed  PubMed Central  Google Scholar 

  29. 29.

    Negash, L., Björn, L. O. (1986) Stomatal closure by ultraviolet radiation. Physiol. Plant. 66, 360–364.

    Google Scholar 

  30. 30.

    Nesterenko, M. V., Tilley, M., Upton, J. (1994) A simple modification of Blum’s silver stain method allows for 30 minute detection of proteins in Polyacrylamide gels. J. Biochem. Biophys. Method. 28, 239–242.

    CAS  Google Scholar 

  31. 31.

    Ornstein, L. (1964) Enzyme Bulletin Canalco Industrial Corporation. Rockvill, Maryland.

    Google Scholar 

  32. 32.

    Radyuk, M. S., Homan, N. M. (2002) Discrete character of the development of the photosynthetic apparatus in greening barley leaves. Photosynth. Res. 72, 117–122.

    CAS  PubMed  Google Scholar 

  33. 33.

    Rajagopal, S., Sicora, C., Várkonyi, Z., Mustárdy, L., Mohanty, P. (2005) Protective effect of supplemental low intensity white light on ultraviolet-B exposure-induced impairment in cyanobacterium Spirulina platensis: Formation of air vacuoles as a possible protective measure. Photosynth. Res. 85, 181–189.

    CAS  PubMed  Google Scholar 

  34. 34.

    Rao, M. V., Paliyathy, G., Ormrod, D. P. (1996) Ultraviolet-B and ozone-induced biochemical changes in antioxidant enzymes of Arabidopsis thaliana. Plant Physiol. 110, 125–136.

    CAS  PubMed  PubMed Central  Google Scholar 

  35. 35.

    Rathore, D., Agrawa, S. B., Singh, A. (2003) Influence of supplemental UV-B radiation and mineral nutrients on biomass, pigments and yield of two cultivars of wheat (Triticum aestivum L.). Int. J. Biotronik. 32, 1–15.

    Google Scholar 

  36. 36.

    Ries, G., Heller, W., Puchta, H., Sandermann, H., Seidlitz, H. K., Hohn, B. (2000) Elevated UV-B radiation reduces genome stability in plants. Natur. 406, 98–101.

    CAS  Google Scholar 

  37. 37.

    Smirnoff, N. (1993) The role of active oxygen in the response of plants to water deficit and desiccation. New Phytol. 125, 27–58.

    CAS  Google Scholar 

  38. 38.

    Schreiber, U., Schliwa, U., Bilger, W. (1986) Continuous recording of photochemical and non-photochemical fluorescence quenching with a new type of modulation fluorometer. Photosynth. Res. 10, 51–62.

    CAS  Google Scholar 

  39. 39.

    Searles, P. S., Flint, S. D., Caldwell, M. M. (2001) A meta-analysis of plant studies simulating stratospheric ozone depletion. Oecologi. 127, 1–10.

    Google Scholar 

  40. 40.

    Sharma, P. K., Anand, P., Sankhalkar, S. (1998) Oxidative damage and changes in activities of antioxidant enzymes in wheat seedlings exposed to ultraviolet-B radiation. Curr Sci. 75, 359–366.

    CAS  Google Scholar 

  41. 41.

    Strid, Å. (1993) Increased expression of defence genes in Pisum sativum after exposure to supplementary ultraviolet-B radiation. Plant Cell Physiol. 34, 949–953.

    CAS  Google Scholar 

  42. 42.

    Strid, Å., Chow, W. S., Anderson, J. M. (1990) Effects of supplementary ultraviolet-B radiation on photosynthesis of in Pisum sativum. Biochim. Biophys. Act. 1020, 260–268.

    CAS  Google Scholar 

  43. 43.

    Takeuchi, A., Yamaguchi, T., Hidema, J., Strid, A., Kumagai, T (2002) Changes in synthesis growing under supplementary UV-B radiation. Plant Cell Environ. 25, 695–705.

    CAS  Google Scholar 

  44. 44.

    Tekchandani, K., Guruprasad, N. (1998) Modulation of guaiacol peroxidase inhibitor by UV-B in cucumber cotyledons. Plant Sci. 136, 131–137.

    CAS  Google Scholar 

  45. 45.

    Tevini, M., Iwanzik, W., Thoma, U. (1981) Some effects of enhanced UV-B radiation on the growth and composition of plants. Plant. 153, 388–394.

    CAS  Google Scholar 

  46. 46.

    Tullberg, A., Alexciev, K., Pfannschmidt, T., Allen, J. F. (2000) Photosynthetic electron flow regulates transcription of the psaB gene in pea (Pisum sativum L.) chloroplasts through the redox state of the plastoquinine pool. Plant Cell Physiol. 41, 1045–1054.

    CAS  PubMed  Google Scholar 

  47. 47.

    Tyystjarvi, E., Karunen, J. (1990) A microcomputer program and fast analog to digital converter card for the analysis of fluorescence induction transients. Photosynth. Res. 26, 127–132.

    CAS  PubMed  Google Scholar 

  48. 48.

    Willekens, H., Van Camp, W., Van Montagu, M., Inze, D., Langebartels, C., Sandermann, H. (1994) Ozone, sulfur dioxide, and ultraviolet-B have similar effects on mRNA accumulation of antioxidant genes in Nicotiana plumbaginifolia L. Plant Physiol. 106, 1007–1014.

    CAS  PubMed  PubMed Central  Google Scholar 

  49. 49.

    Woodbury, W., Spenser, A. K., Stahmann, M. A. (1971) An improved procedure using ferricyanide for detecting catalase isoenzymes. Anal. Biochem. 44, 301–305.

    CAS  PubMed  PubMed Central  Google Scholar 

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Fedina, I., Velitchkova, M., Georgieva, K. et al. UV-B Response of Greening Barley Seedlings. BIOLOGIA FUTURA 60, 195–210 (2009). https://doi.org/10.1556/ABiol.60.2009.2.7

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  • Antioxidant enzymes
  • chlorophyll fluorescence
  • flavonoids
  • oxygen evolution
  • UV-B radiation