CO(2) Enhancement of Growth and Photosynthesis in Rice (Oryza sativa) : Modification by Increased Ultraviolet-B Radiation

Two cultivars of rice (Oryza sativa L.) IR-36 and Fujiyama-5 were grown at ambient (360 microbars) and elevated CO₂ (660 microbars) from germination through reproduction in unshaded greenhouses at the Duke University Phytotron. Growth at elevated CO₂ resulted in significant decreases in nighttime respiration and increases in photosynthesis, total biomass, and yield for both cultivars. However, in plants exposed to simultaneous increases in CO₂ and ultraviolet-B (UV-B) radiation, CO₂ enhancement effects on respiration, photosynthesis, and biomass were eliminated in IR-36 and significantly reduced in Fujiyama-5. UV-B radiation simulated a 25% depletion in stratospheric ozone at Durham, North Carolina. Analysis of the response of CO₂ uptake to internal CO₂ concentration at light saturation suggested that, for IR-36, the predominant limitation to photosynthesis with increased UV-B radiation was the capacity for regeneration of ribulose bisphosphate (RuBP), whereas for Fujiyama-5 the primary photosynthetic decrease appeared to be related to a decline in apparent carboxylation efficiency. Changes in the RuBP regeneration limitation in IR-36 were consistent with damage to the photochemical efficiency of photosystem II as estimated from the ratio of variable to maximum chlorophyll fluorescence. Little change in RuBP regeneration and photochemistry was evident in cultivar Fujiyama-5, however. The degree of sensitivity of photochemical reactions with increased UV-B radiation appeared to be related to leaf production of UV-B-absorbing compounds. Fujiyama-5 had a higher concentration of these compounds than IR-36 in all environments, and the production of these compounds in Fujiyama-5 was stimulated by UV-B fluence. Results from this study suggest that in rice alterations in growth or photosynthesis as a result of enhanced CO₂ may be eliminated or reduced if UV-B radiation continues to increase.     

Intraspecific responses of <Emphasis Type="Italic">Fagopyrum esculentum</Emphasis> to enhanced ultraviolet B radiation

The effects of supplemental UV-B radiation on crop growth, morphology, reproduction and physiology were studied in three cultivars of Fagopyrum esculentum Moench (buckwheat) originating from high elevation (Qinghai-Tibet plateau) and lower altitudes (The Sichuan Basin). Our results showed that common buckwheat was sensitive to UV-B stress. Plant growth, development, and reproduction were inhibited by elevated UV-B radiation. Plant lipid oxidation and polyphenol oxidase (PPO) activity increased with increasing UV-B radiation, along with the concentration of phenylpropanoid compounds, superoxide dismutase (SOD) activity and ascorbic acid (Asa) concentration were also enhanced at the lowest level of supplemental UV-B radiation but decreased at the higher level of enhanced UV-B. While, a cultivar originating from elevated locations had lower dry matter accumulation and was more tolerant to UV-B radiation than cultivars originating from lower elevations. The effects on leaf thickness and increased antioxidant capacity could be linked with the improved performance of cultivar originating from high elevation when exposed to enhanced UV-B radiation. We conclude that UV-B tolerance should be considered prior to introducing or breeding common buckwheat cultivars from lowland cultivation to regions at high elevation such as the Qinghai-Tibet plateau.     

Effects of UV-B radiation on soybean yield and seed quality: a 6-year field study

Two soybean [ Glycine max (L.) Merr.] cultivars, Essex and Williams, were grown in the field for 6 consecutive seasons under ambient and supplemental levels of ultravio-Set-B radiation to determine the potential for alterations in yield or seed quality with a reduction in the stratospheric ozone column. The supplemental UV-B fluences simulated a 16 or 25% ozone depletion. The data presented here represent the first field experiment conducted over multiple seasons which assesses the effects of increased UV-B radiation on seed yield. Overall, the cultivar Essex was found to be sensitive to UV-B radiation (yield reductions of 20%) while the cultivar Williams was tolerant. However, the effectiveness of UV-B radiation in altering yield was strongly influenced by the seasonal microclimate, and the 2 cultivars responded differently to these changing factors. Yield was reduced most in Essex during seasons in which water availability was high and was reduced in Williams only when water was severely limiting. The results of these experiments demonstrate the necessity for multiple-year experiments and the need to increase our understanding of the interaction between UV-B radiation and other environmental stresses in order to assess the potential consequences of stratospheric ozone depletion.     

Intraspecific differences in physiological response of 20 wheat cultivars to enhanced ultraviolet-B radiation under field conditions

Field studies were conducted to determine the potential for alterations in physiology and the intraspecific variation in sensitivity of 20 wheat (Triticum aestivum) cultivars to enhanced ultraviolet-B (UV-B, 280-315 nm) radiation. The supplemental UV-B radiation was 5 kJ m(-2), simulating a depletion of 20% stratospheric ozone. Out of 20 wheat cultivars (from South China, North China and Mexico) tested, 13 showed significant changes in total chlorophyll content. In most of these sensitive species, chlorophyll a content was strongly reduced, and chlorophyll b content decreased in a lesser extent, leading to a decrease in chlorophyll a/b ratio. However, some species had an increased chlorophyll a/b ratio under enhanced UV-B. The effect of UV-B on flavonoid content also showed intraspecific differences, a significant increase for one cultivar, decreases in 12 cultivars and no effect on the other seven cultivars. Superoxide dismutase (SOD) activity of five cultivars was significantly increased, and that of six cultivars significantly decreased. Membrane permeability of 12 cultivars significantly increased, while only that of Dali 905 was significantly decreased. Malonaldehyde (MDA) contents of eight cultivars were increased significantly, while that of three cultivars was significantly decreased. Although large intraspecific differences were found for the different parameters measured, there was no clear correlation between them under UV-B radiation.     

紫外光B辐射增强对水稻叶片内IAA和ABA含量的影响

 两个水稻品种CK46和Dular生长在人工气候箱的条件下,在0.0和13.0kJm-2 day-1(模拟臭氧浓度下降20％的UVB强度)的紫外光B(UV-B 280 nm-320 nm)下进行4周的照射处理,研究UV-B对水稻体内内源IAA和ABA含量的影响。结果表明：随着UV—B处理时间的延长,CK46和Dular叶片内的IAA含量下降。相反,UV-B辐射增强使两个品种叶片内的ABA含量上升。     

Intraspecific variation in sensitivity to ultraviolet-B radiation in endogenous hormones and photosynthetic characteristics of 10 wheat cultivars grown under field conditions

Field studies were conducted to determine the potential of altering endogenous hormones and photosynthetic characteristics and intraspecific variation in sensitivity of 10 wheat (Triticum aestivum) cultivars (four tolerant, two middle sensitive and four sensitive) to enhanced ultraviolet-B (UV-B, 280–315 nm) radiation under field conditions. The supplemental UV-B radiation was 5.00 kJ m⁻², simulating a depletion of 20% stratospheric ozone. Responses were cultivar-specific. Out of the 10 tested wheat cultivars, six showed significant decrease in IAA content. UV-B radiation significantly increased ZR content in two wheat cultivars and significantly decreased in five cultivars. ABA content of three wheat cultivars was increased significantly, while that of five cultivars was decreased significantly. UV-B radiation significantly increased the stomatal conductance of three cultivars, and significantly decreased that of four cultivars. Intercellular CO₂ concentrations were significantly increased in five cultivars and significantly decreased in one cultivar (Mianyang 20). Transpiration rate of three cultivars significantly increased, while that of three cultivars significantly decreased. UV-B radiation significantly decreased the net photosynthetic rate of six cultivars. Intraspecific differences were found for the different measured parameters. For seven measured parameters, UV-B radiation had significant effects on five wheat cultivars, while no effect on the others. Significant correlations were observed between net photosynthetic rate and stomatal conductance, intercellular CO₂ concentrations and transpiration rate in eight cultivars. UV-B radiation might change stomatal conductance, intercellular CO₂ concentrations and transpiration rate, thus resulting in changes in net photosynthetic rate.     

The effect of enhanced ultraviolet-B radiation on growth,photosynthesis and stable carbon isotope composition (δ13C) of two soybean cultivars (Glycine max) under field conditions

Two Chinese cultivars of Glycine max, namely Heidou and Jindou, were exposed to ambient and supplemental levels of ultraviolet-B (UV-B) radiation simulating a 24% depletion in stratospheric ozone over a 9-week growing period at an outdoor experimental site. Enhanced UV-B irradiation significantly reduced leaf, stem and root biomass, and plant height in the Heidou cultivar. These changes were associated with a diminished photosynthetic (net CO₂) rate, stomatal conductance, transpiration rate and water use efficiency, and accompanied by decreased foliar chlorophyll a and b, and total carotenoid concentrations and elevated foliar flavonoid levels. In contrast, the Jindou cultivar displayed only a significantly reduced stem mass and stomatal conductance, but no changes in pigment composition under elevated UV-B. The greater tolerance of elevated UV-B exposures by the Jindou cultivar was attributed partly to its higher foliar flavonoid content, smaller leaf size, thicker leaf cuticle and scabrous (hairy) lamina. Nevertheless both the Heidou cultivar and the less UV-B sensitive Jindou cultivar displayed an altered carbon isotope composition (δ¹³C) in their tissues following exposure to elevated UV-B. Such carbon isotope composition changes in plant tissues suggested a means of early detection of photosynthetic disruption in plants with anticipated increase in UV-B due to stratospheric ozone depletion.     

Influence of supplemental UV-B radiation on photosynthetic characteristics of rice plants

In a field experiment with rice (Oryza sativa L. cv. Saket 4) grown under ambient and supplemental ultraviolet-B (UV-B) radiation at 20 % ozone depletion, differences in gas exchange, concentrations of photosynthetic pigments, anthocyanins and flavonoids, biomass accumulation, catalase and peroxidase activities, and contents of ascorbic acid and phenol were determined. Decline in photosynthesis was associated with reductions in stomatal conductance and concentrations of photosynthetic pigments. Enhanced UV-B radiation (eUV-B) increased the contents of flavonoid and phenolic compounds in leaves. Peroxidase activity increased and catalase activity was always lower at eUV-B. The total plant biomass decreased at eUV-B.     

Influence of supplemental UV-B radiation on photosynthetic characteristics of rice plants

In a field experiment with rice (Oryza sativa L. cv. Saket 4) grown under ambient and supplemental ultraviolet-B (UV-B) radiation at 20 % ozone depletion, differences in gas exchange, concentrations of photosynthetic pigments, anthocyanins and flavonoids, biomass accumulation, catalase and peroxidase activities, and contents of ascorbic acid and phenol were determined. Decline in photosynthesis was associated with reductions in stomatal conductance and concentrations of photosynthetic pigments. Enhanced UV-B radiation (eUV-B) increased the contents of flavonoid and phenolic compounds in leaves. Peroxidase activity increased and catalase activity was always lower at eUV-B. The total plant biomass decreased at eUV-B.     

Influence of ultraviolet-B (UV-B) radiation on photosynthetic and growth characteristics in field-grown cassava (Manihot esculentum Crantz)

The effects of ultraviolet-B (UV-B between 290 and 320 nm) on photosynthesis and growth characteristics were investigated in field grown cassava (Manihot esculentum Crantz). Plants were grown at ambient and ambient plus a 5.5kJ m^?2 d^?1 supplementation of UV-B radiation for 95 d. The supplemental UV-B fluence used in this experiment simulated a 15% depletion in stratospheric ozone at the equator (0°N). Carbon dioxide exchange, oxygen evolution, and the ratio of variable to maximum fluorescence (F_v/F_m) were determined for fully expanded leaves after 64–76 d of UV-B exposure. AH plants were harvested after 95 d of UV-B exposure, assayed for chlorophyll and UV-B absorbing compounds, and separated into leaves, petioles, stems and roots. Exposure to UV-B radiation had no effect on in situ rates of photosynthesis or dark respiration. No difference in the concentration of UV-B absorbing compounds was observed between treatments. A 2-d daytime diurnal comparison of F_v to F_m ratios indicated a significant decline in F_v/F_m ratios and a subsequent increase in photoinhibition under enhanced UV-B radiation if temperature or PPF exceeded 35°C or 1800μmol m^?2 s^?1, respectively. However, UV-B effects on fluorescence kinetics appeared to be temporal since maximal photosynthetic rates as determined by oxygen evolution at saturated CO₂ and PPF remained unchanged. Although total biomass was unaltered with UV-B exposure, alterations in the growth characteristics of cassava grown with supplemental UV-B radiation are consistent with auxin destruction and reduced apical dominance. Changes in growth included an alteration of biomass partitioning with a significant increase in shoot/root ratio noted for plants receiving supplemental UV-B radiation. The increase in shoot/root ratio was due primarily to a significant decrease in root weight (–32%) with UV-B exposure. Because root production determines the harvest-able portion of cassava, UV-B radiation may still influence the yield of an important tropical agronomic species, even though photosynthesis and total dry biomass may not be directly affected.     

Interaction of Elevated Ultraviolet-B Radiation and CO(2) on Productivity and Photosynthetic Characteristics in Wheat, Rice, and Soybean

Wheat (Triticum aestivum L. cv Bannock), rice (Oryza sativa L. cv IR-36), and soybean (Glycine max [L.] Merr cv Essex) were grown in a factorial greenhouse experiment to determine if CO₂-induced increases in photosynthesis, biomass, and yield are modified by increases in ultraviolet (UV)-B radiation corresponding to stratospheric ozone depletion. The experimental conditions simulated were: (a) an increase in CO₂ concentration from 350 to 650 microliters per liter; (b) an increase in UV-B radiation corresponding to a 10% ozone depletion at the equator; and (c) a and b in combination. Seed yield and total biomass increased significantly with elevated CO₂ in all three species when compared to the control. However, with concurrent increases in UV-B and CO₂, no increase in either seed yield (wheat and rice) or total biomass (rice) was observed with respect to the control. In contrast, CO₂-induced increases in seed yield and total plant biomass were maintained or increased in soybean within the elevated CO₂, UV-B environment. Whole leaf gas exchange indicated a significant increase in photosynthesis, apparent quantum efficiency (AQE) and water-use-efficiency (WUE) with elevated CO₂ in all 3 species. Including elevated UV-B radiation with high CO₂ eliminated the effect of high CO₂ on photosynthesis and WUE in rice and the increase in AQE associated with high CO₂ in all species. Elevated CO₂ did not change the apparent carboxylation efficiency (ACE) in the three species although the combination of elevated CO₂ and UV-B reduced ACE in wheat and rice. The results of this experiment illustrate that increased UV-B radiation may modify CO₂-induced increases in biomass, seed yield and photosynthetic parameters and suggest that available data may not adequately characterize the potential effect of future, simultaneous changes in CO₂ concentration and UV-B radiation.     

水稻对UV-B辐射增强的抗性遗传及其生理生化特性研究

本研究结果表明紫外线B(UVB,280～330nm)能严重影响水稻秧苗生长发育,从而引起植株矮化,叶片干枯,叶绿素含量降低,单株干物质减少甚至死亡。不同水稻基因型对紫外线的反应存在明显差异。在3个供试品种(HR、M63、Hr)中,HR最抗UVB危害。遗传分析表明水稻抗UVB的性状为数量遗传性状且受多对隐性基因控制,生化分析表明水稻秧苗经UVB处理后,在0～3d内,随着时间推移,秧苗保护酶活性明显增强,之后,保护酶活性逐惭下降,MDA含量相应增加。抗UVB品种HR比感UVB品种Hr保护酶活性一直较高,膜质过氧化程度较低,反映了保护酶对抵抗UVB危害的重要作用。     

Effects of enhanced UV-B radiation on the hormonal content of vegetative and reproductive tissues of two tomato cultivars and their relationships with reproductive characteristics

The effects of enhanced UV-B radiation on hormone changes in vegetative and reproductive tissues of tomato (Lycopersicon esculentum Mill.) and their relationships with reproductive characteristics were studied. Two cultivars, TongHui (TH) and XiaGuang (XG), were grown in the field for one growing season under ambient (Control), ambient plus 2.54 kJ m^–2 d^–1 (T1) or ambient plus 4.25 kJ m^–2 d^–1 (T2) of supplemental ultraviolet-B (280–320 nm). The number of open flowers increased significantly in the TH cultivar under T2 while it declined in the XG cultivar under T1. Although pollen germination from both cultivars was inhibited by UV-B treatment, fruit number was enhanced in the TH cultivar at both UV-B doses and in the XG cultivar at the low dose. On the other hand, seed size (dry weight) was reduced in the XG cultivar by both UV-B doses and in the TH cultivar at the low UV-B dose. The final germination rates of seeds from control and UV-B treated plants of both cultivars showed no significant differences (p > 0.05), while germination was delayed in the TH cultivar at both doses of UV-B and in the XG cultivar only for T2. To determine the mechanism of UV-B's effects on developmental processes, hormone concentrations in leaves, pistils and seeds were analyzed using ELISA on partially purified extracts. The results suggested that enhanced UV-B radiation induced hormone changes in both vegetative and reproductive tissues. The alteration of flower number may be associated with the changes of ZR in leaves under enhanced UV-B radiation and the delayed germination may due to the changes in seed ABA and GAs.     

Influence of supplemental ultraviolet-B on indoleacetic acid and calmodulin in the leaves of rice (Oryza sativa L.)

IR68 and Dular rice cultivars were grown under ambient, 13.0 (simulating 20% ozone depletion) and 19.1 (simulating 40% ozone depletion) kJ m^-2 day^-1 of biologically effective ultraviolet-B (UV-B_BE) for 4 weeks. Plant height and leaf area were significantly reduced by supplemental UV-B_BE radiation. Greater reduction in leaf area than of plant height was observed. A decrease in indole-3-acetic acid (IAA) content and increase in peroxidase and IAA oxidase activities of UV-B treated plants in both cultivars were observed compared with ambient control. Calmodulin content also decreased after plants were treated with high supplemental UV-B for two weeks and medium UV-B treatment for four weeks. The results indicated that peroxidase and IAA oxidase activities in rice leaves were stimulated by supplemental UV-B, resulting in the destruction of IAA which in turn may cause inhibition of rice leaf growth. Although the mechanism is unclear, calmodulin is most likely involved in leaf growth.     

NaCI胁迫对UV-B辐射诱导的绿豆环丁烷嘧啶二聚体和紫外吸收物质含量变化的影响

将2个对UV-B敏感性不同的绿豆品种‘秦豆-20’和‘中绿-1’幼苗放在培养室内,进行0.4W/m~2 UV-B辐射和0.4%NaCl胁迫的单独或复合处理,研究了NaCl胁迫对UV-B辐射诱导的DNA伤害和修复的影响。结果显示:在NaCl胁迫下,(1)在光下抗UV-B的品种‘中绿-1’的环丁烷嘧啶二聚体(CPD)累积量降低,而敏感品种‘秦豆-20’的CPD累积量未发生变化;(2)两品种CPD形成量均比无NaCl胁迫时低;(3)抗UV-B品种DNA的光、暗修复能力均比无NaCl胁迫时高:(4)而敏感品种DNA的光修复能力比无NaCl胁迫时低、暗修复能力未发生变化。另外,CPD形成量与紫外吸收物含量间具有明显的负相关性。说明NaCl胁迫不仅影响2个绿豆品种幼苗的CPD形成量,而且影响DNA的光、暗修复能力,进而导致了CPD累积量发生变化,由此影响了幼苗的UV-B敏感性。结果也暗示CPD形成量的变化是由于紫外吸收物质含量的不同所导致的。     

Morphological and physiological responses of bean plants to supplemental UV radiation in a Mediterranean climate

During the last few decades many experiments have been performed to evaluate the responses of plants to enhanced solar UV-B radiation (280–320 nm) that may occur because of stratospheric ozone depletion; most of them were performed in controlled environment conditions where plants were exposed to low photosynthetically active radiation (PAR) levels and high UV-B irradiance. Since environmental radiative regimes can play a role in the response of plants to UV-B enhancement, it appears doubtful whether it is valid to extrapolate the results from these experiments to plants grown in natural conditions. The objective of this work was to evaluate the effects on physiology and morphology of a bean (Phaseolus vulgaris L.) cultivar Nano Bobis, exposed to supplemental UV radiation in the open-air. UV-B radiation was supplied by fluorescent lamps to simulate a 20% stratospheric ozone reduction. Three groups of plants were grown: control (no supplemental UV), UV-A treatment (supplementation in the UV-A band) and UV-B treatment (supplemental UV-B and UV-A radiation). Each group was replicated three times. After 33 days of treatment plants grown under UV-B treatment had lower biomass, leaf area and reduced leaf elongation compared to UV-A treatment. No significant differences were detected in photosynthetic parameters, photosynthetic pigments and UV-B absorbing compounds among the three groups of plants. However, plants exposed to UV-A treatment showed a sort of 'stimulation' of their growth when compared to the control. The results of this experiment showed that plants may be sensitive to UV-A radiation, thus it is difficult to evaluate the specific effects of UV-B (280–320 nm) radiation from fluorescent lamps and it is important to choose the appropriate control. Environmental conditions strongly affect plant response to UV radiation so further field studies are necessary to assess the interaction between UV-B exposure and meteorological variability.     

Epidermal transmittance and phenolic composition in leaves of atrazine-tolerant and atrazine-sensitive cultivars of Brassica napus grown under enhanced UV-B radiation

Experiments were conducted on the atrazine-tolerant mutant Stallion and the atrazine-sensitive cv. Paroll of Brassica napus L., which were grown under either visible light or with the addition of UV-B radiation (280–320 nm) for 15 days. The mutant has been shown to be sensitive to high levels of visible light as compared to the atrazine-sensitive cultivar and therefore we wished to determine plant response to UV-B radiation with respect to potential pigment changes, certain anatomical features, radiation penetration and partial photosynthesis. With regard to pigment changes, we were particularly interested in whether the compositional shift in flavonol pigments under enhanced UV-B radiation, previously suggested to favour increased antioxidant activity, is confined to the adaxial epidermis, which generally receives most UV-B radiation or whether the pigment shift is also inducible in the abaxial epidermis.As was to be expected, the penetration of UV-B radiation (310 nm) was lower in the UV-B-exposed plants, which was correlated with an increased amount of UV-screening pigments in the adaxial and abaxial epidermal layers. The main flavonoid glycosides showed the largest shift from kaempferol to quercetin as aglycone moiety in the adaxial epidermal layer. However, in the abaxial epidermal layer the hydroxycinnamic acid (HCA) derivatives and kaempferol glycosides were predominant. Penetration of 430 nm light was higher after UV-B exposure, and probably contributed to the fact that photosynthetic efficiency of photosystem II was unchanged or higher after UV-B exposure. UV-B radiation decreased leaf area in the atrazine-tolerant mutant only. Both cultivars showed an increased leaf thickness after UV-B exposure due to cell elongation mainly of the palisade tissue. This was especially evident in the mutant.     

UV-B辐射增强对两种不同抗性水稻叶片光合生理及超显微结构的影响

在UV-B辐射增强条件下,研究了两个不同水稻品种叶片光合作用系统的变化。结果表明：（1）UV-B辐射胁迫使两个水稻品种叶片总叶绿素含量,叶绿素a与叶绿素b（Chla／Chlb）比值下降,叶绿素a荧光诱导动力学参数改变,光系统Ⅱ活性受抑制,光合作用效率降低,其中Dular受抑制的程度较Lemont大。（2）利用扫描电镜（SEM）和透射电镜（TEM）进一步研究表明,UV-B辐射胁迫使水稻叶片气孔器受破坏,叶绿体结构变形,基粒片层排列稀疏紊乱,两个供试品种结构上受破坏的程度与它们光合生理受抑制的程度一致。（3）叶片边缘受破坏的程度较主脉两侧轻,这可能与硅质乳突密度较大有关。（4）两个供试品种叶片表面主脉两侧的硅质乳突数量及其受UV-B辐射影响的特性存在明显的差异,Lemont叶表面的乳突分布密度较大,且在UV-B辐射胁迫下有增加的趋势,而Dular则相反。这说明硅质体的累积特性可能是水稻对UV-B辐射胁迫的适应机制之一。     

不同小麦品种对UV-B辐射增强响应的生理特性差异

研究了大田条件下模拟增强UV-B辐射（500 KJ·m^-2,相当于昆明地区臭氧层减少20%）对10个小麦品种生理指标的影响以及小麦对UV B辐射响应的种内差异.结果表明,10个供试小麦品种中有6个品种的叶绿素含量显著下降,叶绿素a降低的程度大于叶绿素b,从而导致叶绿素a/b的比率下降.UV-B对小麦叶片内MDA和类黄酮的影响也具有种内差异,有5个品种的MDA含量显著上升, 2个品种的MDA含量显著下降;4个品种的类黄酮含量显著增加,2个品种的类黄酮含量显著减少.叶绿素和类黄酮含量变化与MDA含量均呈显著负相关关系,类黄酮与小麦UV-B抗性之间存在密切联系.