CaCl_2对UV-B辐射下菠菜叶片光合特性的影响

以"丹麦旺盛菠菜"为材料,通过UV-B和CaCl2复合处理,测定光合色素含量、Hill反应活力、叶绿素荧光、MDA含量和抗氧化酶活性等参数,探讨了CaCl2对UV-B辐射下菠菜叶片电子传递链和光合膜酶保护系统的影响。结果表明,UV-B处理下,光合色素含量、chl/car、类囊体膜上PSII潜在活性(Fv/Fo)、光化学淬灭系数(qP)、非光化学淬灭系数(qN)、PSII光量子产量(ΦPSⅡ)、原初光能转化效率(Fv/Fm),以及Hill反应活力等降低,chla/chlb和MDA含量升高;喷洒CaCl2可不同程度缓解UV-B的伤害。不同处理下,POD、SOD和CAT活性的变化呈现补偿效应。UV-B强度与菠菜叶片PSII功能受损程度呈正相关,CaCl2则主要通过提高chlb含量、类囊体膜上的光量子产量和POD活性,以缓解伤害。重度UV-B辐射下,CaCl2使chlb含量显著提高可能是导致PSII捕光效率提高的重要因素。     

INTERACTIONS BETWEEN UV‐B EXPOSURE AND PHOSPHORUS NUTRITION. I. EFFECTS ON GROWTH,PHOSPHATE UPTAKE,AND CHLOROPHYLL FLUORESCENCE1

The interactive effects of P starvation and exposure to UV radiation on growth rates, quantum efficiency of PSII electron transport, and P‐uptake capacity of the chlorophyte microalga Dunaliella tertiolecta Butcher are presented. Ultraviolet radiation did not in itself cause marked changes in growth rate, though it did induce changes in the effective quantum yield of PSII. Depriving cells of phosphate resulted in significant changes in all parameters examined. The decline of growth rate and fluorescence parameters after P starvation was significantly faster in the presence of UV radiation. Ultraviolet radiation also stimulated the magnitude of the transient changes in chl fluorescence (nutrient‐induced fluorescence transient) exhibited by P‐starved cells after resupply of that nutrient.     

Effects of changes in ambient PAR and UV radiation on the nutritional quality of an Arctic diatom (Thalassiosira antarctica var. borealis)

Polyunsaturated fatty acids (PUFAs) are essential macromolecules that are synthesized by phytoplankton during spring bloom, and they play a key role in the Arctic food web. They are, however, considered to be sensitive to oxidation by UV radiation (280-400 nm). Changes in the food quality of primary producers may affect the transport of biomass and energy in the whole ecosystem. Using a common Arctic diatom, we looked at the effect of ambient and increased UV radiation on its nutritional quality, specifically, the fatty acid composition and elemental ratios. In May 2004, in the archipelago of Svalbard (79° N), a unialgal culture of Thalassiosira antarctica var. borealis was subjected to a 17-day experiment in outdoor aquaria. The diatoms were kept in semi-continuous culture (40 1) and exposed to three treatments with different levels of UV radiation: none (UV-shielded), ambient, and enhanced. Fatty acid composition, C:N:P ratios, photosynthetic pigment composition, optimum quantum yield of PSII, and cell numbers were analysed over the experimental period. An initial increase in PAR (photosynthetically active radiation, 400-700 nm) intensities profoundly affected the fatty acid composition and substantially inhibited the synthesis of PUFAs, but the relative amounts of PUFAs were not reduced by UV radiation. Enhanced UV radiation did, however, cause a significant reduction in optimum quantum yield of PSII and affected some fatty acids, mainly 18:0 and 16:1 n-7, during the first week of the experiment. Both ambient and enhanced UV radiation caused significantly lower C:P and N:P ratios. At the same time, these treatments elicited a higher relative content of the photoprotective pigments diadinoxanthin and diatoxanthin. After acclimation to the new light levels these effects faded off. Thus, brief periods with high light exposure may cause significant changes in photosynthetic activity and food quality, but the capacity for photo-acclimation seems high. The impact of UV radiation seems to be less important for food quality than that of PAR during a sudden rise in total light intensity.     

Modifications in photosystem II photochemistry in senescent leaves of maize plants

Analyses of CO2 exchange and chlorophyll fluorescence were carried out to assess photosynthetic performance during senescence of maize leaves. Senescent leaves displayed a significant decrease in CO2 assimilatory capacity accompanied by a decrease in stomatal conductance and an increase in intercellular CO2 concentration. The analyses of fluorescence quenching under steady-state photosynthesis showed that senescence resulted in an increase in non-photochemical quenching and a decrease in photo-chemical quenching. It also resulted in a decrease in the efficiency of excitation energy capture by open PSII reaction centres and the quantum yield of PSII electron transport, but had very little effect on the maximal efficiency of PSII photochemistry. The results determined from the fast fluorescence induction kinetics indicated an increase in the proportion of QB-non-reducing PSII reaction centres and a decrease in the rate of QA reduction in senescent leaves. Theoretical analyses of fluorescence parameters under steady-state photosynthesis suggest that the increase in the non-photochemical quenching was due to an increase in the rate constant to thermal dissipation of excitation energy by PSII and that the decrease in the quantum yield of PSII electron transport was associated with a decrease in the rate constant of PSII photochemistry. Based on these results, it is suggested that the decrease in the quantum yield of PSII electron transport in senescent leaves was down-regulated by an increase in the proportion of QB-non-reducing PSII reaction centres and in the non-photochemical quenching. The photosynthetic electron transport would thus match the decreased demand for ATP and NADPH in carbon assimilation which was inhibited significantly in senescent leaves.Key words: Chlorophyll fluorescence, gas exchange, maize (Zea mays L.), photochemical and non-photochemical quenching, photosystem II photochemistry.      

Oxyradicals and PSII activity in maize leaves in the absence of UV components of solar spectrum

The regulation of oxyradicals and PSII activity by UV-B (280-315 nm) and UV-A (315-400 nm) components were investigated in the leaves of maize [Zea mays L. var: HQPM.1]. The impact of ambient UV radiation on the production of superoxide (O2-) and hydroxyl (.OH) radicals were analysed in the leaves of 20-day-old plants. The amount of O2.- and .OH radicals and the radical scavenging activity were significantly higher in the leaves exposed to ambient UV radiation as compared to the leaves of the plants grown under UV exclusion filters. Smaller amount of oxyradicals in the leaves of UV excluded plants was accompanied by a substantial increase in quantum yield of electron transport (phi Eo), rate of electron transport (psi o) and performance index (PIABS), as indicated by chlorophyll a fluorescence transient. Although higher amounts of oxyradicals invoked higher activity of antioxidant enzymes like superoxide dismutase and peroxidase under ambient UV, they also imposed limitation on the photosynthetic efficiency of PSII. Exclusion of UV components (UV-B 280-315 nm; UV-A 315-400 nm) translated to enhanced photosynthesis, growth and biomass. Thus, solar UV components, especially in the tropical region, could be a major limiting factor in the photosynthetic efficiency of the crop plants.     

Photosynthetic performance of the Atlantic brown macroalgae, Cystoseira abies-marina, Dictyota dichotoma and Sargassum vulgare, measured in Gran Canaria on site

Photosynthetic performance was determined in three common Atlantic brown macroalgae, Cystoseira abies-marina, Dictyota dichotoma and Sargassum vulgare, in Gran Canaria, Canary Islands, on site. The photosynthetic quantum yield was measured with both a portable PAM instrument on site and a diving PAM under water in the habitat. In parallel, solar radiation was measured continuously above and under water by means of two three-channel dosimeters, ELDONET (Real Time Computer, M?hrendorf, Germany), in three wavelength ranges, UV-A, UV-B and PAR. The effective photosynthetic quantum yield decreased in all species in response to exposure to 15 min of solar radiation but recovered in the subsequent shade conditions within several hours. A 30-min exposure caused an even more profound photoinhibition from which the algae recovered only partially. Most of the effect was due to visible radiation, however, the UV wavelength range, and especially UV-B, considerably enhanced the decrease in photosynthetic quantum yield. In all species except Sargassum a significant photoinhibition was detected at their growth sites at high solar angles in the water column, measured with the diving PAM.     

紫外线-B辐射与3种植物幼苗的光合作用——光合作用对紫外线-B敏感性的比较

紫外线 - B辐射降低豌豆、大豆和黄瓜 3种植物幼苗的净光合作用速率 ( Pn)和表观量子效率 ( AQE) ,Pn和 AQE的减少幅度随紫外线 - B辐射时间的延长逐渐增大。 3种植物的光合作用对紫外线 - B辐射的敏感性依次为 :豌豆 >黄瓜 >大豆 ,光合作用光抑制的发生程度与光合作用对 UV- B辐射敏感性有内在联系。定量分析显示 ,光合作用光抑制的发生程度随UV- B辐射时间增加 ,是 UV- B辐射剂量的累积效应。     

Protective effect of nitric oxide against oxidative stress under ultraviolet-B radiation.

The response of bean leaves to UV-B radiation was extensively investigated. UV-B radiation caused increase of ion leakage, loss of chlorophyll, and decrease of the maximum efficiency of PSII photochemistry (Fv/Fm) and the quantum yield of PSII electron transport (PhiPSII) of bean leaves. H2O2 contents and the extent of thylakoid membrane protein oxidation increased, indicated by the decrease of thiol contents and the increase of carbonyl contents with the duration of UV-B radiation. Addition of sodium nitroprusside, a nitric oxide (NO) donor, can partially alleviate UV-B induced decrease of chlorophyll contents, Fv/Fm and PhiPSII. Moreover, the oxidative damage to the thylakoid membrane was alleviated by NO. The potassium salt of 2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide, a specific NO scavenger, arrested NO mediated protective effects against UV-B induced oxidative damage. Incubation of thylakoid membrane with increasing H2O2 concentrations showed a progressive enhancement in carbonyl contents. H2O2 contents were decreased in the presence of NO under UV-B radiation through increased activities of superoxide dismutases, ascorbate peroxidases, and catalases. Taken together, the results suggest that NO can effectively protect plants from UV-B damage mostly probably mediated by enhanced activities of antioxidant enzymes.     

Response of the photosynthetic apparatus of cotton (Gossypium hirsutum) to the onset of drought stress under field conditions studied by gas-exchange analysis and chlorophyll fluorescence imaging.

The functioning of the photosynthetic apparatus of cotton (Gossypium hirsutum) grown during the onset of water limitation was studied by gas-exchange and chlorophyll fluorescence to better understand the adaptation mechanisms of the photosynthetic apparatus to drought conditions. For this, cotton was grown in the field in Central Asia under well-irrigated and moderately drought-stressed conditions. The light and CO(2) responses of photosynthesis (A(G)), stomatal conductance (g(s)) and various chlorophyll fluorescence parameters were determined simultaneously. Furthermore, chlorophyll fluorescence images were taken from leaves to study the spatial pattern of photosystem II (PSII) efficiency and non-photochemical quenching parameters. Under low and moderate light intensity, the onset of drought stress caused an increase in the operating quantum efficiency of PSII photochemistry (varphi(PSII)) which indicated increased photorespiration since photosynthesis was hardly affected by water limitation. The increase in varphi(PSII) was caused by an increase of the efficiency of open PSII reaction centers (F(v)'/F(m)') and by a decrease of the basal non-photochemical quenching (varphi(NO)). Using a chlorophyll fluorescence imaging system a low spatial heterogeneity of varphi(PSII) was revealed under both irrigation treatments. The increased rate of photorespiration in plants during the onset of drought stress can be seen as an acclimation process to avoid an over-excitation of PSII under more severe drought conditions.     

Effect of iron,zinc and manganese shortage-induced change on photosynthetic pigments,some osmoregulators and chlorophyll fluorescence parameters in lettuce

Although the beneficial role of Fe, Zn, and Mn on many physiological and biochemical processes is well established, effects of each of these elements on chlorophyll (Chl) a fluorescence and photosynthetic pigment contents is not well studied. The objective of this study was to evaluate effects of Fe, Zn, and Mn deficiency in two lettuce cultivars. The parameters investigated could serve also as physiological and biochemical markers in order to identify stress-tolerant cultivars. Our results indicated that microelement shortage significantly decreased contents of photosynthetic pigments in both lettuce cultivars. Chl a fluorescence parameters including maximal quantum yield of PSII photochemistry and performance index decreased under micronutrient deficiency, while relative variable fluorescence at J-step and minimal fluorescence yield of the dark-adapted state increased under such conditions in both cultivars. Micronutrient deficiency also reduced all parameters of quantum yield and specific energy fluxes excluding quantum yield of energy dissipation, quantum yield of reduction of end electron acceptors at the PSI, and total performance index for the photochemical activity. Osmoregulators, such as proline, soluble sugar, and total phenols were enhanced in plants grown under micronutrient deficiency. Fe, Zn, and Mn deficiency led to a lesser production of dry mass. The Fe deficiency was more destructive than that of Zn and Mn on the efficiency of PSII in both lettuce cultivars. Our results suggest that the leaf lettuce, which showed a higher efficiency of PSII, electron transport, quantum yield, specific energy fluxes, and osmoregulators under micronutrient deficiency, was more tolerant to stress conditions than crisphead lettuce.     

Photoinhibition by visible and ultraviolet radiation in the red macroalga Porphyra umbilicalis grown in the laboratory

Photosynthetic oxygen production and PAM fluorescence measurements were used to follow photoinhibition in the red macroalga Porphyra umbilicalis. Exposure to simulated solar radiation caused inhibition of the effective photosynthetic quantum yield from which the thalli partially recovered in the shade in subsequent hours. There were no significant differences between samples exposed to unfiltered radiation and those exposed to radiation from which increasing portions of UV radiation had been removed indicating that the thalli are well adapted to current levels of solar PAR and UV radiation. This notion was supported by the finding of high concentrations of UV screening pigments which were even enhanced by exposure to increased UV radiation. However, when exposed to (only) UV radiation about 50% higher than that encountered by the organisms in their natural habitat, the photosynthetic yield decreased slowly and did not show any recovery even when the degree of inhibition did not exceed 10%.     

Photosynthetic and physiological responses of foxtail millet (<Emphasis Type="Italic">Setaria italica</Emphasis> L.) to low-light stress during grain-filling stage

Two foxtail millet (Setaria italica L.) varieties were subjected to different shading intensity treatments during a grain-filling stage in a field experiment in order to clarify physiological mechanisms of low-light effects on the yield. Our results showed that the grain fresh mass per panicle, yield, photosynthetic pigment contents, net photosynthetic rate, stomatal conductance, effective quantum yield of PSII photochemistry, and electron transport rate decreased with the increase of shading intensity, whereas the intercellular CO₂ concentration increased in both varieties. In addition, shading changed a double-peak diurnal variation of photosynthesis to a one-peak curve. In conclusion, the lower yield of foxtail millet was caused mainly by a reduction of grain mass assimilated, a decline in chlorophyll content, and the low photosynthetic rate due to low light during the grain-filling stage. Reduced light energy absorption and conversion, restricted electron transfer, and reduced stomatal conductance might cause the decrease in photosynthesis.     

外源硫对镉胁迫下马齿苋光合性状和矿质元素吸收的影响

从光合反应系统揭示外源硫(S)诱导马齿苋镉(Cd)耐受性的生理机制,为外源S缓解重金属毒害提供理论依据.采用营养液培养,研究外源S供体(NH4)2SO4对100 mg/L Cd胁迫下马齿苋叶片光合色素、光合特性、叶绿素荧光参数和矿质营养元素的影响.结果表明,Cd胁迫可显著降低马齿苋叶片中叶绿素a和叶绿素b含量;净光合速率、蒸腾速率、气孔导度均显著降低,而胞间二氧化碳浓度上升,表明非气孔因素是Cd胁迫诱导马齿苋光合抑制的主要因素;同时,PSⅡ实际光化学效率(ФPSII)、电子传递效率(J)、化学猝灭系数(qP)显著下降,而非化学猝灭系数(qN)显著上升,表明Cd胁迫影响马齿苋PSⅡ反应系统的正常运行.外施400 mg/L(NH4)2SO4显著提高马齿苋叶片叶绿素a含量、叶绿素b含量和叶绿素a/b比值,增强马齿苋叶片光合作用和PSⅡ原初光化学反应量子效率.对5种与光反应系统密切相关的矿质元素含量进行分析发现,Cd处理显著增加马齿苋叶片中的Ca和Fe含量,显著抑制马齿苋对Mg、Mn和Cu的吸收.Cd胁迫下马齿苋叶片的变黄与Mg、Mn的亏缺有关,而与Fe缺乏无关;添加外源S可显著提升马齿苋叶片中Ca、Mg、Fe、Cu和Mn含量,从而增强Cd胁迫下马齿苋叶片的PSII反应系统功能.     

The influence of antimycin A on pigment composition and functional activity of photosynthetic apparatus in <Emphasis Type="Italic">Triticum aestivum</Emphasis> L. under high temperature

The purpose of the current investigation was to evaluate the influence of antimycin A (AA) as an activator of the alternative respiratory pathway (AP) on photosynthetic pigment composition and functional activity of the photosynthetic apparatus of wheat seedlings (Triticum aestivum L.) under exposure to high temperature as well as their acclimation. Our results indicated that a significant decrease (44–74%) of photosynthetic pigment contents was caused by a long-term exposure to high temperature (42°C), while the short-term exposure resulted in 20–46% decline. However, a combined effect of AA and long-term high temperature reduced the total pigment contents by 28–41%. Our results demonstrated that the reduction of the chlorophyll a/b ratio was less significant under the combined effect of AA and high temperature than that under the stressful condition without AA. We observed that short-term and long-term high temperature modified PSII functionality of the first leaves in wheat seedlings, which was manifested by the low maximal quantum yield of PSII photochemistry, maximum fluorescence yield in the dark-adapted state, and by high minimum fluorescence yield in the dark-adapted state. The quantum yield of PSII photochemistry decreased rapidly by 16–24% under the combination of AA and high temperature. Overall, these results suggest that the activation of the alternative pathway, induced by AA, contributed to the stabilization of the photosynthetic apparatus in wheat seedlings under high temperature.     

Acclimation of Maximal Quantum Yield of Photosynthesis in the Brown Alga Alaria esculenta under High Light and UV Radiation

Abstract: Macroalgae of the upper sublittoral zone of Arctic coastal ecosystems are subjected to darkness or low light for several months during winter and have to withstand large changes in irradiance after the breakup of sea ice in the Arctic spring. Changes in photosynthetic response to high PAR (pho-tosynthetically active radiation) and UV-B radiation (UV-B) in the cold temperate brown alga A/aria esculenta were monitored with a PAM fluorometer to study photoinhibition, recovery and acclimation of maximal quantum yield of photochemistry. Plants collected in the field, as well as specimens raised in the laboratory, were exposed to various radiation conditions including different levels of PAR and UV radiation (UV-A + UV-B). Measurements of variable chlorophyll fluorescence of photosystem II revealed that the photosynthetic apparatus in A. esculenta was able to acclimate to the respective high light and UV treatments within several days. However, two different mechanisms of acclimation seem to be involved. Initially, the rate of recovery of maximal quantum yield increased after only a few exposures to high light or UV. Second, after several exposure cycles, the degree of inhibition was reduced. Data on fluorescence induction kinetics and quenching analysis showed that exposure to the respective UV radiation resulted in an increase of non-photochemical quenching, while effective quantum yield of photochemistry was hardly affected.     

高山植物美丽风毛菊PSII光化学效率和光合色素对短期增补UV-B辐射的响应

在中国科学院海北高寒草甸生态系统实验站的综合观测场, 于植物生长季的不同月份进行短期增补UV-B辐射的模拟试验, 研究了高山植物美丽风毛菊(Saussurea superba)的PSII光化学效率、光合色素和UV-B吸收物质对增强UV-B辐射的响应。结果表明, 尽管差异不显著, 暗适应3 min的PSII最大光化学量子效率在不同月份均有降低的趋势, 说明增强UV-B辐射能加剧光合机构的光抑制。不同月份短期增补UV-B辐射均引起光下PSII实际光化学量子效率和光化学猝灭系数的降低, 以及非光化学猝灭系数的增高, 表明增强UV-B辐射能降低叶片的光能捕获效率, 促进非光化学能量耗散过程。增补UV-B辐射后, 叶片光合色素的含量略有降低趋势, 可能与短时间内光合色素形成过程受抑制和光氧化程度的加剧, 以及叶片厚度的略微降低有关。UV-B吸收物质的含量在不同月份没有一致和较为显著的变化, 说明高原强UV-B环境下生存的美丽风毛菊叶表皮层中由类黄酮和衍生多酚类组成的内部紫外屏蔽物质相对稳定, 倾向于较少受增补UV-B辐射的影响。     

Investigating deleterious effects of ultraviolet (UV) radiations on wheat by a quick method

Increasing air pollution has resulted in ozone depletion, which has led to an increase in the amount of ultraviolet (UV) radiation reaching the surface of earth posing hazardous effects on the plant yield. The chlorophyll a fluorescence transients were recorded in vivo using plant efficiency analyzer and analyzed according to JIP test, which can monitor photosystem II (PSII) behavior. This study aims to evaluate the sensitivity of different components of PSII to UV radiations and the extent of damage caused when wheat plants were exposed to UV radiations for 2 and 4 h. It was observed that functional disconnection of light harvesting complexes from PSII complex, accumulation of inactive reaction center, inactivation of oxygen evolving complex, and thus the linear electron transport process (ET₀/CS), were drastically affected by UV radiations. This study will contribute to understanding of the basic photosynthetic mechanisms affected in crop plants due to increased UV radiations. The knowledge obtained will be relevant for environmental and plant scientists to plan strategies to cope with stresses.     

Response to ozone in two lettuce varieties on chlorophyll a fluorescence, photosynthetic pigments and lipid peroxidation.

The effect of different O3 concentrations on two lettuce (Lactuca sativa L.) varieties (Valladolid and Morella) was investigated through chlorophyll (Chl) a fluorescence parameters, photosynthetic pigments (Chl a, b and total carotenoid), lipid peroxidation and crop yield. Ozone fumigation caused: a decrease in maximum quantum yield of photosystem II (PSII) photochemistry (Fv/Fm) in mature leaves, a reduction in the non-cyclic electron flow (phiPSII) and a lower capacity to reoxidize the QA pool (qP). These reductions were significant in the Valladolid var. but not in the Morella var. A significant decrease in Chl a, b and in the total carotenoids was observed in the Valladolid var. but not in the Morella var. mainly under O3 fumigation conditions. We observed that the NPQ parameter did not increase in parallel to the qP reduction seen in the Valladolid var. O3 fumigation with respect to air charcoal filtered air conditions. This fact could be associated with a lower capacity for dissipation of non-radiative excess energy and it may be closely correlated with significant decreases in photosynthetic pigment concentration. A decrease in NPQ from air ozone-free to ozone fumigation in the Morella var. can be explained by the need to maintain the photochemical quenching under O3 stress. It may also be associated with a slight increase in photosynthetic pigments. The differences between the two varieties may indicate that the Valladolid var. is more susceptible to O3 damage.     

EFFECTS OF UV RADIATION ON A CHLOROPHYTE ALGA (SCENEDESMUS SP.) ISOLATED FROM THE FUMAROLE FIELDS OF MT. EREBUS,ANTARCTICA1

Acclimation to UV radiation (UVR) was examined in a unicellular chlorophyte isolated from fumarole fields adjacent to Mt. Erebus, Antarctica. Long‐term exposure to UVR (14 days) initially reduced the efficiency of photosynthetic energy conversion measured as the dark adapted quantum yield of PSII fluorescence (F_v/F_m) when compared with cultures not exposed to UVR. However, the UVR exposed cultures recovered to pre‐UVR exposure levels of photosynthetic efficiency by the end of the experimental period. Additionally, neither growth rates nor cell size were significantly affected by exposure to UVR. In contrast, the maximum rate of carbon fixation on a chl a basis was significantly reduced at the end of the experiment and was accompanied by a significant decrease in chl a concentrations. These results suggest a transient effect of UVR on primary photochemistry consistent with damage to PSII, possibly at the D1 protein, with more chronic effects on carbon fixation that did affect maximum photosynthetic capacity. Ultrastructural and molecular (18s rDNA) data show that this isolate from the fumoral fields of Mt. Erebus, Antarctica is a member of the genus Scenedesmus.     

田间大豆叶片成长过程中的光合特性及光破坏防御机制

田间大豆叶片在成长进程中光饱和光合速率持续提高,但气孔导度的增加明显滞后.尽管叶片在成长初期就具有较高的最大光化学效率,但是仍略低于发育成熟的叶片.随着叶片的成长,光下叶片光系统Ⅱ实际效率增加;非光化学猝灭下降.幼叶叶黄素总量与叶绿素之比较高,随着叶面积的增加该比值下降,在光下,幼叶的脱环氧化程度较高.因此认为大豆叶片成长初期就能够有效地进行光化学调节;在叶片生长过程中依赖叶黄素循环的热耗散机制迅速建立起来有效抵御强光的破坏.