The combined effects of enhanced UV-B radiation and selenium on growth, chlorophyll fluorescence and ultrastructure in strawberry (Fragaria × ananassa) and barley (Hordeum vulgare) treated in the field

The possible ameliorative effects of selenium (Se) addition to soil on the detrimental effects of enhanced UV-B radiation were tested on strawberry and barley during 4 months of field experiment in Kuopio, Central Finland. Control plants were exposed to ambient levels of UV radiation, using arrays of unenergized lamps. A control for UV-A radiation was also included in the experiment. Added Se, applied as H₂SeO₄, at the level of 0.1 mg kg⁻¹ soil (low dosage) and 1 mg kg⁻¹ soil (high dosage) increased Se concentrations in plants more than 10 and 100 times, respectively. After 4 months of exposure, strawberry and barley plants were harvested for biomass analysis. Chlorophyll fluorescence was measured using the Hansatech FMS2 fluorescence monitoring system. Leaf anatomy and ultrastructure were observed by light and transmission electron microscope. Several effects of UV and Se as well as their interaction were found, mostly for strawberry, but not for barley, indicating species-specific responses. Our results provided evidence that the high Se concentration in soil had no ameliorative effect but increased the sensitivity of strawberry to enhanced UV-B radiation in the field. Under ambient radiation, Se did not alter leaf growth of strawberry, whereas under UV-B radiation, the high Se addition significantly decreased leaf growth. Strawberry runner biomass was affected by the interaction of Se and UV. Under ambient radiation Se did not change dry weight of runners, but in combination with UV-A or UV-B radiation the high Se dosage decreased dry weight of runners by about 30%. Although the high Se concentration positively influenced on quantum efficiency of photosystem II (PSII) in strawberry leaves, it reduced runner biomass, leaf number and ratio of starch to chloroplast area. This suggests that the harmful effects of the high Se dosage on photosynthetic processes occurred as a result of changes in activity or/and biosynthesis of enzymes, rather than alteration of PSII. At the low concentration, Se effects were slight and variable.Although barley leaves accumulated higher Se concentrations than strawberry, there were no apparent changes in their growth, biomass or chlorophyll fluorescence due to Se effect either alone or in combination with UV-B. However, at the ultrastructural level, an enlargement in the peroxisome area was found due to combination of UV radiation with Se, suggesting the activation of antioxidative enzymes, possibly catalase. Decrease in mitochondrial density in barley cells in response to Se might be attributed to alteration of mitochondrial division. Increase in the proportion of cells with cytoplasmic lipid bodies due to combined effect of UV-B and Se indicated the alteration of lipid metabolism and the acceleration of cell senescence in barley. Main UV-B effects were found, mostly at the tissue and ultrastructural level in strawberry, but not in barley, indicating species-specific susceptibility to enhanced UV-B radiation. UV-B-treated strawberry plants developed marginally thinner leaves with reduced ratio of starch to chloroplast area in their cells, suggesting negative influence of UV-B on photosynthetic processes.