Growth enhancement of soybean (Glycine max) upon exclusion of UV-B and UV-B/A components of solar radiation: characterization of photosynthetic parameters in leaves

Exclusion of UV (280–380 nm) radiation from the solar spectrum can be an important tool to assess the impact of ambient UV radiation on plant growth and performance of crop plants. The effect of exclusion of UV-B and UV-A from solar radiation on the growth and photosynthetic components in soybean (Glycine max) leaves were investigated. Exclusion of solar UV-B and UV-B/A radiation, enhanced the fresh weight, dry weight, leaf area as well as induced a dramatic increase in plant height, which reflected a net increase in biomass. Dry weight increase per unit leaf area was quite significant upon both UV-B and UV-B/A exclusion from the solar spectrum. However, no changes in chlorophyll a and b contents were observed by exclusion of solar UV radiation but the content of carotenoids was significantly (34–46%) lowered. Analysis of chlorophyll (Chl) fluorescence transient parameters of leaf segments suggested no change in the F _v/F _m value due to UV-B or UV-B/A exclusion. Only a small reduction in photo-oxidized signal I (P700⁺)/unit Chl was noted. Interestingly the total soluble protein content per unit leaf area increased by 18% in UV-B/A and 40% in UV-B excluded samples, suggesting a unique upregulation of biosynthesis and accumulation of biomass. Solar UV radiation thus seems to primarily affect the photomorphogenic regulatory system that leads to an enhanced growth of leaves and an enhanced rate of net photosynthesis in soybean, a crop plant of economic importance. The presence of ultra-violet components in sunlight seems to arrest carbon sequestration in plants. An erratum to this article can be found at     

Response of Barley Seedlings to UV-B Radiation as Affected by Proline and NaCl

Barley (Hordeum vulgare L. cv. Alfa) seedlings were treated for 4 d before UV-B irradiation with 0.05 mM proline or 150 mM NaCl. UV-B exposure induced synthesis of yellow coloured compounds with maximum absorbance at 438 nm. The content of these compounds was increased in proline-treated and decreased in NaCl-treated plants. UV-B radiation reduced chlorophyll/carotenoids ratio, oxygen evolution rate and photochemical efficiency of PS 2 as estimated by chlorophyll fluorescence and increased proline accumulation, H₂O₂ generation and lipid peroxidation. Exogenous proline had no effect on the parameters studied and did not change the response of plants to UV-B radiation. NaCl inhibited photochemical efficiency of PS 2, reduced oxygen evolution and increased H₂O₂ concentration and lipid peroxidation. The combination of NaCl and proline treatment led to lowering the inhibitory effect of NaCl in non UV-B irradiated seedlings. There was not relationship between the level of UV-B-induced compounds and UV-B tolerance of barley seedlings.     

Action of ultraviolet radiation (UV-B) upon cuticular waxes in some crop plants

The surface structure and composition of surface lipids were examined in leaves of barley, bean, and cucumber seedlings grown in a growth chamber under white light and low levels of ultraviolet (UV-B; 280–320 nm) radiation. The cuticular wax of cucumber cotyledons and bean leaves appeared as a thin homogeneous layer, whereas on barley leaves crystal-like structures could be observed under these irradiation conditions. Principally, the amount of cuticular wax found in barley leaves was five times greater than in bean or cucumber leaves. The prediominant wax components were primary alcohols in barley, primary alcohols and monoesters in bean, and alkanes in cucumber cotyledons. Irradiation with enhanced UV-B levels caused an increase of total wax by about 25% in all plant species investigated. Aldehydes, detected as a minor constituent of cucumber and barley wax, increased twofold. Distribution patterns of the homologs within some wax classes were different at low and enhanced UV-B levels. In general, the distribution of the homologs was shifted to shorter acyl chain lengths in wax of leaves exposed to enhanced UV-B levels. This was most apparent in cucumber wax, less in bean or barley wax. The UV-B-caused effects upon cucumber wax were mainly due to a response by the adaxial surface of the leaf.Abbreviation UV-B Ultraviolet radiation (280–320 nm)     

Ultraviolet-B radiation effects on the Mediterranean ruderal Dittrichia viscosa

Young seedlings of Dittrichia viscosa L. (syn. Inula viscosa (L.) Aiton) (Asteraceae) were extensively treated with artificial rain in order to remove the water soluble component of their epicuticular UV-B absorbing compounds. As a result, 75% of the epicuticular absorbing capacity at 300 nm was lost. The seedlings were subsequently grown in a naturaly lit glasshouse for 80 days under 0.06, 6.41 and 10.14 kJ m^-2 day ^-1 biologicaly effective UV-B radiation doses. The initial, pre-rain values of the water soluble, epicuticular UV-B absorbing potential was restored in about three weeks. During this transient period the plants were exposed to the enhanced UV-B radiation doses with part of their UV-B radiation screen removed. Although a trend for increased accumulation of epicuticular UV-B absorbing capacity was observed with increasing UV-B radiation doses, the allelopathic potential of the epicuticular material remained unchanged. Internal (cellular) UV-B absorbing compounds and chlorophylls were unaffected, but total carotenoids were increased, indicating a possible protective role against UV-B radiation damage. Leaf, stem and root dry mass were the same under all treatments but UV-B radiation caused a reduction in the dry mass invested per unit leaf area with a concomitant increase in leaf area. The importance of this UV-B radiation induced selective allocation of photosynthate to the production of assimilative surfaces is discussed.     

Responses in growth,physiology and nitrogen nutrition of dragon spruce (<Emphasis Type="Italic">Picea asperata</Emphasis>) seedlings of different ages to enhanced ultraviolet-B

In the southeast of the Qinghai–Tibetan Plateau of China, dragon spruce (Picea asperata) is a key species and widely used in reforestation processes in the area. The paper mainly studied the effects of ultraviolet-B (UV-B) on growth, physiology and nitrogen nutrition of 3- and 6-year-old dragon spruce seedlings. The experimental design included ambient UV-B (control) and enhanced UV-B (+UV-B, a 30% increase). Enhanced UV-B significantly decreased growth, needle and root nitrogen concentration, needle nitrate reductase activity and increased UV-B absorbing compounds and malondialdehyde (MDA) content of two old dragon spruce seedlings. Glutamine synthetase activity was not affected by enhanced UV-B in two old dragon spruce seedlings. On the other hand, different old seedlings also exhibited different physiological responses to enhanced UV-B radiation. Chlorophyll content, carotenoids content and soluble protein content in 3-year-old seedlings significantly reduced by enhanced UV-B, but those in 6-year-old seedlings were not affected by enhanced UV-B. Proline content of 6-year-old seedlings were increased by enhanced UV-B. Compared with the 3-year-old seedlings, the 6-year-old seedlings showed lower reduction of growth and MDA content, and accumulated more proline and UV-B absorbing compounds for protecting seedlings under enhanced UV-B. The results implicated that 3-year-old seedlings were more sensitivity to enhanced UV-B than 6-year-old seedlings.     

UV-B辐射对南亚热带森林木本植物幼苗生长的影响

研究了UV -B辐射对广东省南亚热带森林木本植物九节、鸭脚木、猴耳环、半枫荷、山乌桕和绿化树种大叶合欢幼苗生长的影响。实验结果表明 ,UV -B辐射降低叶片光合色素的含量 ;显著降低幼苗的净光合速率、蒸腾速率和气孔导度 ;降低干物质的增长 ;抑制大叶合欢幼苗根瘤的形成     

Response of barley seedlings to UV-B radiation as affected by NaCl

The response of barley seedlings, subjected to 150 mmol/L NaCl for 4 days at different light regimes (4 d in the light, 4 d in darkness and a 12 h light/dark cycle) before UV-B radiation was investigated. NaCl treatment resulted in a decrease of total chlorophyll content and an increase in H2O2, free proline and lipid peroxidation, as quantified by measurement of malondialdehyde. Significantly more proline was accumulated in the light than in darkness. The combination of UV-B and NaCl treatment produced an additive effect on most of the parameters studied. UV-B radiation reduced the chlorophyll/carotenoids ratio and photochemical efficiency of PSII as estimated by chlorophyll fluorescence. NaCl pre-exposure decreased H2O2 generation and lipid peroxidation and alleviated the inhibitory effect of UV-B on PSII activity. Proline accumulated under salt stress conditions might be one of the reasons for the observed tolerance of barley seedlings to UV-B radiation.     

NaCl induced cross-acclimation to UV-B radiation in four Barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis> L.) cultivars

The effect of pre-treatment with 200 mM NaCl on the response of four barley cultivars (Hordeum vulgare L. cv. Bülbül-89, Kalaycı-97, Tarm-92 and Tokak-157/37) to UV-B radiation was investigated. Salt stress as well as UV-B irradiation led to a decrease of the total chlorophyll (chl) content in all cultivars, except in Kalaycı-97. While carotenoids are almost not affected by NaCl treatment, UV-B irradiation caused an increase by 5–20% of carotenoid content of all cultivars. UV-B induced damages of photosynthetic apparatus were estimated by the rate of photosynthetic electron transport measured by chl fluorescence and the rate of oxygen evolution, the latter being more affected. Pre-treatment with NaCl alleviated harmful effect of UV-B irradiation on F _v/F _m and ETR, but not on oxygen evolution. UV-B-induced and UV-B-absorbing compounds with absorption at 300 and 438 nm increased as a result of UV-B treatment. The level of stress marker proline increased considerably as a result of NaCl treatment, while UV-B irradiation resulted in a pronounced increase of the level of H₂O₂. MDA enhanced in the seedlings subjected to salt and UV-B stress. Established cross-acclimation to UV-B as a result of salt treatment could be due to the increased free proline and the level of UV-B absorbing compounds in barley seedlings subjected to NaCl.     

Growth under UV-B radiation increases tolerance to high-light stress in pea and bean plants

Pea (Pisum sativum L.) and bean (Phaseolus vulgaris L.) plants were exposed to enhanced levels of UV-B radiation in a growth chamber. Leaf discs of UV-B treated and control plants were exposed to high-light (HL) stress (PAR: 1200 mol m^–2 s^–1) to study whether pre-treatment with UV-B affected the photoprotective mechanisms of the plants against photoinhibition. At regular time intervals leaf discs were taken to perform chlorophyll a fluorescence and oxygen evolution measurements to assess damage to the photosystems. Also, after 1 h of HL treatment the concentration of xanthophyll cycle pigments was determined. A significantly slower decline of maximum quantum efficiency of PSII (F _v/F _m), together with a slower decline of oxygen evolution during HL stress was observed in leaf discs of UV-B treated plants compared to controls in both plant species. This indicated an increased tolerance to HL stress in UV-B treated plants. The total pool of xanthophyll cycle pigments was increased in UV-B treated pea plants compared to controls, but in bean no significant differences were found between treatments. However, in bean plants thiol concentrations were significantly enhanced by UV-B treatment, and UV-absorbing compounds increased in both species, indicating a higher antioxidant capacity. An increased leaf thickness, together with increases in antioxidant capacity could have contributed to the higher protection against photoinhibition in UV-B treated plants.     

Effects of UV-B radiation on photosynthetic reactions in Rhizophora apiculata

Influences of UV-B radiation on Rhizophora apiculata were studied in terms of chlorophylls, their presence in protein complexes of the chloroplast, PS I and PS II photochemical activities, in vitro absorption spectrum of the chloroplast, in vivo leaf fluorescence and UV absorbing compounds. The seedlings were exposed to the various levels of UV-B radiations, equivalent to 0 (control), 10, 20, 30 and 40% stratospheric ozone depletion of the study area. The low doses of UV-B (10 and 20%) increased the reaction centre chlorophyll (10 and 8%) and activities of PS-I (98 and 39%) and PS-II (77 and 38%) respectively; whereas, 30 and 40% UV-B treatments decreased the reaction centre chlorophylls by 11 and 33% and PS II activity by 0 and 20%; while PS I activity did not show any inhibitory effect. Chloroplasts isolated from control and 10% UV-B treated plants exhibited the same level of absorption at 676 nm. In vivo leaf fluorescence was found to be diminished with UV-B radiation and at the 10% UV-B, variable fluorescence was promoted significantly by 10%. The content of UV-absorbing compounds was progressively enhanced with doses of UV-B radiation along with higher absorption at 276 and 330 nm.     

Stratospheric Ozone Depletion: High Arctic Tundra Plant Growth on Svalbard is not Affected by Enhanced UV-B after 7 years of UV-B Supplementation in the Field

The response of tundra plants to enhanced UV-B radiation simulating 15 and 30% ozone depletion was studied at two high arctic sites (Isdammen and Adventdalen, 78° N, Svalbard).The set-up of the UV-B supplementation systems is described, consisting of large and small UV lamp arrays, installed in 1996 and 2002. After 7 years of exposure to enhanced UV-B radiation, plant cover, density, morphological (leaf fresh and dry weight, leaf thickness, leaf area, reproductive and ecophysiological parameters leaf UV-B absorbance, leaf phenolic content, leaf water content) were not affected by enhanced UV-B radiation. DNA damage in the leaves was not increased with enhanced UV-B in Salix polaris and Cassiope tetragona. DNA damage in Salix polaris leaves was higher than in leaves of C. tetragona. The length of male gametophyte moss plants of Polytrichum hyperboreum was reduced with elevated UV-B as well as the number of Pedicularis hirsuta plants per plot, but the inflorescence length of Bistorta vivipara was not significantly affected. We discuss the possible causes of tolerance of tundra plants to UV-B (absence of response to enhanced UV-B) in terms of methodology (supplementation versus exclusion), ecophysiological adaptations to UV-B and the biogeographical history of polar plants     

铁核桃根与叶水浸提液对果园间作绿肥植物的化感作用及其生理机制

以西南地区具有代表性的16种绿肥植物为受体材料,采用培养皿药膜法研究了铁核桃(Juglans sigillata)根水浸提液对受体种子发芽率及幼苗鲜重、干重的化感效应;并进一步研究了铁核桃根、叶水浸提液和胡桃醌对化感效应存在明显差异的4种绿肥植物(绿豆、红三叶、白三叶、花生)种子萌发与幼苗生长以及抗氧化酶特性的影响,以筛选适宜中国西南地区核桃园种植的绿肥植物,探讨核桃根和凋落物对绿肥作物的化感作用机制。结果表明:(1)铁核桃根水浸提液对绿豆的发芽率没有影响,但对绿豆幼苗鲜重和干重有显著抑制作用,而对其他15种绿肥的发芽率和鲜重、干重均有抑制作用。(2)胡桃醌显著抑制绿豆种子萌发,而铁核桃根或叶水浸提液对绿豆种子萌发没有影响。(3)铁核桃根或叶水浸提液以及胡桃醌对绿肥植物幼苗生长的化感效应趋势一致,但核桃根或叶水浸提液的化感效应强于胡桃醌。(4)绿豆幼苗在铁核桃根或叶水浸提液以及胡桃醌处理下,超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)的活性均高于其他3种(红三叶、白三叶、花生)受体幼苗,表明绿豆清除活性氧能力高,细胞受损害程度较低,受化感作用影响最弱。研究认为,绿豆为适宜中国西南地区幼龄核桃园种植的间作绿肥植物。     

Effects of solar ultraviolet-B radiation,temperature and CO2 on growth and physiology of sunflower and maize seedlings

The effects of solar UV-B radiation, in combination with elevated temperature (4 °C ) and CO₂ (680 L L^-1 concentration, on sunflower and maize seedlings were studied from May to August in 1991 at the research station Quinta de São Pedro in Portugal (38.7°N). The ambient solar radiation of Portugal was reduced to levels of Central European latitudes by using the ozone filter technique. This radiation served as control, while the ambient solar radiation of Portugal was to simulate intense UV-B treatment (+30%). All plants were grown up to 18 days in 4 climate controlled growth chambers simulating a daily course of temperature with T_max=28 °C or 32 °C , resp., and ambient CO₂ concentrations (340 L L^-1); in one chamber the CO₂ concentration was twice as high (680 L L^-1). Under intense UV-B and at 28 °C (T_max) all growth parameters (height, leaf area, fresh and dry weight, stem elongation rate, relative growth rate) of sunflower and maize seedlings were reduced down to 35% as compared to controls. An increase in growing temperature by 4 °C , alone or in combination with doubled CO₂, compensated or even overcompensated the UV-B effect so that the treated plants were comparable to controls. Chlorophyll content, on a leaf area basis, increased under intense UV-B radiation. This increase was compensated by lower leaf areas, resulting in comparable chlorophyll contents. Similar to growth, also the net photosynthetic rates of sunflower and maize seedlings were reduced down to 29% by intense UV-B calculated on a chlorophyll basis. This reduction was compensated by an increased temperature. Doubling of CO₂ concentration had effects only on sunflower seedlings in which the photosynthetic rates were higher than in the controls. Dark respiration rates of the seedlings were not influenced by any experimental condition. Transpiration and water use efficiency (wue) were not influenced by intense UV-B. Higher temperatures led to higher transpiration rates and lower water use efficiencies, resp.. Doubling of CO₂ reduced the transpiration rate drastically while for wue maximum values were recorded.     

UV-B response of green and etiolated barley seedlings

7-d-old etiolated and green barley seedlings (Hordeum vulgare L. cv. Alfa) were irradiated with UV-B for 30 min and then kept for 24 h in light or darkness. Chlorophyll (Chl) synthesis was inhibited by about 30 % as a result of UV-B irradiation, but there were no significant changes in photochemical activity measured by variable to maximum fluorescence ratio (F_v/F_m), quantum yield (Φ_PS2) and oxygen evolution rate. Electron transport of etiolated seedlings was similar to that of green ones, nevertheless, the Chl content was more then 2-fold lower. Ribulose-1,5-bisphosphate carboxylase/oxygenase large and small subunits were diminished as a result of UV-B irradiation in etiolated and green plants, especially in those kept in the darkness. Catalase activity decreased and total superoxide dismutase activity increased in green and etiolated plants following UV-B treatment. When benzidine was used as a substrate, an isoform located between guaiacol peroxidases 2 and 3 (guaiacol peroxidase X) appeared, which was specific for UV-B treatment. As a result of irradiation, the contents of UV-B absorbing and UV-B induced compounds increased in green seedlings but not in etiolated seedlings.     

Pigment Accumulation and Photosynthesis in Developing Rye Coleoptiles

Rye seedlings (Secale cereale L.) were grown in darkness or irradiated with white light (WL) of medium intensity (100 μmol quanta . m^?2 . s^?1). In the coleoptile and primary leaf of irradiated plants synthesis of chlorophylls a and b with average ratios of 2.4 and 2.7, respectively, were measured. The chlorophyll content of the fully green primary leaf was about 10-fold larger than that of the pale green coleoptile. In darkness a large increase in carotenoid content occurred in the primary leaf, but in the coleoptile the level of carotenoids remained very low. In both organs a WL-induced increase in carotenoid synthesis was observed. The level of anthocyanin was likewise enhanced by WL. In the coleoptile and primary leaf of irradiated seedlings light-dependent oxygen evolution was measured. The specific activity of the enzyme ribulose-1,5-bisphosphate carboxylase/oxygenase was enhanced after WL treatment. The results indicate that the mature rye coleoptile is a photosynthetically active plant organ.     

Inhibition of photosynthetic activity by UV-B radiation in radish seedlings

Inhibition of primary photosynthetic reactions by UV-B radiation (280 nm-320 nm) was demonstrated in radish leaves ( Raphanus sativus cv. Saxa Treib). Detached radish cotyledons from 10-day-old seedlings were irradiated with continuous white light and increasing UV-B irradiances using cut-off filters with increasing transmission for shorter wavelengths (WG 360, WG 345, WG 320, WG 305, WG 295, WG 280). Photosynthetic activity measured in terms of chlorophyll fluorescence induction (Kautsky effect) after 2, 4, 6, 8 and 24 h irradiation decreased in a wavelength dependent way with increasing UV-B irradiance and irradiation time.
Radish seedlings grown for 10 days from the time of germination under the same UV-B irradiation conditions exhibited similar reductions of the variable fluorescence as detached cotyledons irradiated for short time periods. They additionally had lower initial fluorescence at high UV-B radiation levels, although the chlorophyll content per leaf area increased. In contrast to short term experiments, the plastoquinone and flavonoid content increased with increasing UV-B irradiance when based on leaf area.     

Impacts of ambient solar uv (280-400 nm) radiation on three tropical legumes

Tropical regions receive the highest level of global solar ultraviolet (UV) radiation especially UV-B (280-320 nm). The average daily dose of the UV-B radiation in Madurai, South India (10°N) is 10 kJ m^-2. This is approximately 50% more than the average daily UV-B radiation in many European countries. A field study was conducted using selective filters to remove either the UV-B (< 320 nm) or UV-B/A (<400 nm) of the solar spectrum, and the effects were followed inCyamopsis tetragonoloba, Vigna mungo, andVigna radiata to determine their sensitivity to UV. When compared to ambient radiation, exclusion of solar UV-B increased the seedling height, leaf area, fresh weight and dry weight and the crop yield by 50% in the case ofCyamopsis, and the extent of such increase was slightly less under UV-B/A exclusion. InV. mungo a significant reduction was seen in solar UV excluded plants whileV. radiata was found to be unaffected.     

Effects of supplementary UV-B radiation on development of damping-off in spinach caused by the soil-borne fungusFusarium oxysporum

The effects of UV-B radiation (290–320 nm) on development of damping-off of spinach (Spinacia oleracea) caused by the fungusFusarium oxysporum were examined in a growth cabinet. The incidence of disease greatly increased when experimental plants were grown in visible radiation with supplementary UV-B radiation. This increase was suppressed by increasing the irradiation of visible radiation.Fusarium oxysporum was isolated from the roots of all damping-off plants and the roots of some unwilted plants, indicating that spinach infected with the pathogen did not necessarily suffer from damping-off in 15d. Supplementary UV-B radiation suppressed the increase in growth components such as the number of leaves, the plant height and the fresh weight of aboveground plant parts, but did not affect the fresh weight of roots. The ratio of the number of plants infected with pathogen to the total number of plants was over 80% irrespective of light conditions. It was suggested that the defense response of spinach to this pathogen was greatly influenced by the physiological state of aboveground plant parts resulting from supplementary UV-B radiation.     

UV-B increases the harvest index of bean (Phaseolus vulgaris L.)

The effects of small changes in natural UV-B on the photosynthesis, pigmentation, flowering and yield of bean plants (Phaseolus vulgaris L. var. Label) were studied. To obtain a relatively natural growth environment, the plants were grown in small, half-open greenhouses of UV-transmitting Plexiglas of different thickness (3 and 5 mm), resulting in an 8% difference in the weighted UV-B reaching the plants. Although the UV-B doses used did not significantly influence photosynthesis on a leaf area basis during vegetative growth, important changes in biomass allocation were noted. A UV-B-O induced reduction in leaf area during the period of vegetative growth resulted in decreased dry weight after 57 d. During the flowering and pod-filling stages (57–79.d after planting), however, plants grown at high UV-B retained their photosynthetic capacity longer: maximal photosynthesis, chlorophyll and N content of the leaves were higher under the higher UV-B dose at a plant age of 79 d. Combined with an increased allocation under the higher UV-B dose of both N and biomass to the pods, this resulted in a small increase in yield and an important increase in harvest index with increased UV-B.     

Response of photosynthesis to high light and drought for Arabidopsis thaliana grown under a UV-B enhanced light regime

Arabidopsis thaliana grown in a light regime that included ultraviolet-B (UV-B) radiation (6 kJ m⁻² d⁻¹) had similar light-saturated photosynthetic rates but up to 50% lower stomatal conductance rates, as compared to plants grown without UV-B radiation. Growth responses of Arabidopsis to UV-B radiation included lower leaf area (25%) and biomass (10%) and higher UV-B absorbing compounds (30%) and chlorophyll content (52%). Lower stomatal conductance rates for plants grown with UV-B radiation were, in part, due to lower stomatal density on the adaxial surface. Plants grown with UV-B radiation had more capacity to down regulate photochemical efficiency of photosystem II (PSII) as shown by up to 25% lower φ_PSII and 30% higher non-photochemical quenching of chlorophyll fluorescence under saturating light. These contributed to a smaller reduction in the maximum photochemical efficiency of PSII (F _v/F _m), greater dark-recovery of F _v/F _m, and higher light-saturated carbon assimilation and stomatal conductance and transpiration rates after a four-hour high light treatment for plants grown with UV-B radiation. Plants grown with UV-B were more tolerant to a 12 day drought treatment than plants grown without UV-B as indicated by two times higher photosynthetic rates and 12% higher relative water content. UV-B-grown plants also had three times higher proline content. Higher tolerance to drought stress for Arabidopsis plants grown under UV-B radiation may be attributed to both increased proline content and decreased stomatal conductance. Growth of Arabidopsis in a UV-B-enhanced light regime increased tolerance to high light exposure and drought stress.