CO2浓度升高对苦草(Vallisneria natans)叶绿素荧光特性的影响

为了探究大气CO₂升高对沉水植物光合生理的影响,利用便携式植物效率分析仪（Handy PEA）,在无损的情况下测定不同CO₂浓度处理下的苦草（Vallisneria natans）叶绿素荧光诱导曲线,并采用JIP-test分析方法分析数据,研究CO₂浓度对苦草叶片叶绿素荧光特性的影响。结果表明在实验进行60 d后,与对照相比,高CO₂浓度处理下的苦草叶片PSⅡ反应中心受体侧荧光参数V_j、M_o显著升高,S_m、ψ_o、φ_Eo显著降低,叶片电子传递能力减弱;K相相对可变荧光W_k显著提高,PSⅡ反应中心供体侧放氧复合体OEC受到伤害;ABS/RC、DI_o/RC、TR_o/RC、DI_o/CS_o显著升高,ET_o/RC、RE_o/RC、ET_o/CS_o、RE_o/CS_o显著降低,苦草叶片用于热耗散的能量显著增加,导致用于电子传递及传递到电子链末端的能量显著减少;性能参数F_v/F_m、PI_abs显著降低,苦草叶片PSⅡ潜在活性和光合作用原初反应过程受到抑制。以上结果表明,在长期高CO₂浓度处理下,苦草叶片光合机构功能受到抑制,PSⅡ反应中心活性降低,光合功能下调,发生光适应现象。     

Chlorophyll Fluorescence Analysis of Cyanobacterial Photosynthesis and Acclimation

Cyanobacteria are ecologically important photosynthetic prokaryotes that also serve as popular model organisms for studies of photosynthesis and gene regulation. Both molecular and ecological studies of cyanobacteria benefit from real-time information on photosynthesis and acclimation. Monitoring in vivo chlorophyll fluorescence can provide noninvasive measures of photosynthetic physiology in a wide range of cyanobacteria and cyanolichens and requires only small samples. Cyanobacterial fluorescence patterns are distinct from those of plants, because of key structural and functional properties of cyanobacteria. These include significant fluorescence emission from the light-harvesting phycobiliproteins; large and rapid changes in fluorescence yield (state transitions) which depend on metabolic and environmental conditions; and flexible, overlapping respiratory and photosynthetic electron transport chains. The fluorescence parameters F_V/F_M, F_V′/F_M′,q_p,q_N, NPQ, and PS II were originally developed to extract information from the fluorescence signals of higher plants. In this review, we consider how the special properties of cyanobacteria can be accommodated and used to extract biologically useful information from cyanobacterial in vivo chlorophyll fluorescence signals. We describe how the pattern of fluorescence yield versus light intensity can be used to predict the acclimated light level for a cyanobacterial population, giving information valuable for both laboratory and field studies of acclimation processes. The size of the change in fluorescence yield during dark-to-light transitions can provide information on respiration and the iron status of the cyanobacteria. Finally, fluorescence parameters can be used to estimate the electron transport rate at the acclimated growth light intensity.     

Variable Chlorophyll a Fluorescence and CO(2) Uptake in Water-Stressed White Spruce Seedlings

Regulation of photosynthetic activity can contribute to the prevention of photodamage in stress resistant plants during exposure to drought or low temperatures. Responses to increasing levels of water stress were examined in seedlings of the stress resistant forest conifer, white spruce (Picea glauca [Moench.] Voss). Some seedlings were grown under aseptic in vitro conditions and others in pots. In relatively resistant in vivo seedlings, photosynthetic activities changed slowly in response to increasing water stress. Highly sensitive in vitro seedlings responded to water deficits similarly to in vivo seedlings but over a much shorter time scale. Fluorescence, CO₂ exchange, and stomatal conductance data reported here suggest possible mechanisms for the regulation of photochemical activity in these plants.     

Chlorophyll a Fluorescence and Photosynthetic and Growth Responses of Pinus radiata to Phosphorus Deficiency, Drought Stress, and High CO(2)

Needles from phosphorus deficient seedlings of Pinus radiata D. Don grown for 8 weeks at either 330 or 660 microliters CO₂ per liter displayed chlorophyll a fluorescence induction kinetics characteristic of structural changes within the thylakoid chloroplast membrane, i.e. constant yield fluorescence (F_O) was increased and induced fluorescence ([F_P-F_I]/F_O) was reduced. The effect was greatest in the undroughted plants grown at 660 μl CO₂ L⁻¹. By week 22 at 330 μl CO₂ L⁻¹ acclimation to P deficiency had occurred as shown by the similarity in the fluorescence characteristics and maximum rates of photosynthesis of the needles from the two P treatments. However, acclimation did not occur in the plants grown at 660 μl CO₂ L⁻¹. The light saturated rate of photosynthesis of needles with adequate P was higher at 660 μl CO₂ L⁻¹ than at 330 μl CO₂ L⁻¹, whereas photosynthesis of P deficient plants showed no increase when grown at the higher CO₂ concentration. The average growth increase due to CO₂ enrichment was 14% in P deficient plants and 32% when P was adequate. In drought stressed plants grown at 330 μl CO₂ L⁻¹, there was a reduction in the maximal rate of quenching of fluorescence (R_Q) after the major peak. Constant yield fluorescence was unaffected but induced fluorescence was lower. These results indicate that electron flow subsequent to photosystem II was affected by drought stress. At 660 μl CO₂ L⁻¹ this response was eliminated showing that CO₂ enrichment improved the ability of the seedlings to acclimate to drought stress. The average growth increase with CO₂ enrichment was 37% in drought stressed plants and 19% in unstressed plants.     

植物对开放式CO2 浓度增高(FACE)的响应与适应研究进展

开放式CO2浓度增高(FACE)系统是近年研究植物对高CO2浓度响应和适应的新手段,它比以往密闭和半密闭系统对实验植物生长环境的干扰少.利用FACE系统进行研究更有助于正确地预测未来大气CO2浓度增高对植物的影响.该文结合作者的研究工作简要评介了FACE系统与以往密闭和半密闭式CO2浓度增高实验系统的不同之处以及近年来利用FACE系统所作的最新研究进展.     

冬小麦光合作用对开放式空气CO2 浓度增高(FACE)的非气孔适应

在同样CO2浓度下测定时,开放式空气CO2浓度增高(FACE,580 μmol CO2 /mol)条件下生长的冬小麦叶片的净光合速率、气孔导度和羧化效率都显著低于普通空气(380 μmol CO2 /mol)中生长的对照叶片.与此相一致,FACE叶片的可溶性蛋白、二磷酸核酮糖羧化酶/加氧酶(Rubisco)和Rubisco活化酶含量也都显著低于对照叶片.这些结果表明,在根系生长不受限制的田间条件下,冬小麦叶片的光合作用对高浓度CO2产生了适应现象,其主要原因可能是碳同化的关键酶Rubisco等含量的降低.     

Elevated CO2 Effects during Leaf Ontogeny (A New Perspective on Acclimation)

For many plants growth in elevated CO2 leads to reduced rates of photosynthesis. To examine the role that leaf ontogeny plays in the acclimation response, we monitored photosynthesis and some related parameters at short intervals throughout the ontogenetic development of tobacco (Nicotiana tabacum L.) leaves under ambient (350 [mu]L L-1)- and high (950 [mu]L L-1)-CO2 conditions. The pattern of photosynthetic rate over time was similar between the two treatments and consistent with the expected pattern for a typical dicot leaf. However, the photosynthesis pattern in high-CO2-grown tobacco was shifted temporally to an earlier maximum and subsequent senescent decline. Ribulose-1,5-biphosphate carboxylase/oxygenase activity appeared to be the main factor regulating photosynthetic rates in both treatments. Therefore, we propose a new model for interpreting the acclimation response. Lowered photosynthetic rates observed during acclimation appear to be the result of a shift in the timing of the normal photosynthetic stages of leaf ontogeny to an earlier onset of the natural decline in photosynthetic rates associated with senescence.     

Photosynthetic Gas Exchange and Chlorophyll a Fluorescence in Salicornia brachiata (Roxb.) Under Osmotic Stress

Osmotic stress negatively affects the photosynthetic efficiency and cause a significant loss of crop productivity. Salicornia brachiata (Roxb.) is a eu-halophyte. We hereby report on photosynthetic gas exchange and chlorophyll fluorescence in S. brachiata under sodium chloride (NaCl), seawater and polyethylene glycol (PEG) induced osmotic stress. It grows luxuriantly and exhibited a higher tolerance index and better accumulation of organic solutes under 100% strength of seawater (32.5 ppt) and 0.5 M NaCl salinity. It exhibited comparatively better gas exchange, stomatal conductance, PSII photochemistry and electron transfer under 100% strength of seawater salinity. Higher chlorophyll a/b ratio under stress conditions indicated a lower ratio of PSII to PSI and balanced excitation of PSI and PSII in S. brachiata resulting in efficient photosynthetic processes. The lower total chlorophyll/carotenoids ratio and higher non-photochemical quenching indicated the photo-protection and safer dissipation of heat energy in S. brachiata under stress. The 100% strength of seawater and 0.5 M NaCl salinity in S. brachiata did not cause significant changes in antenna size, connectivity between PSII reaction centres (RCs) and reduction of electrons on PSII donor side. The 20% PEG induced the inactivation of RCs and cause damage to PSII RCs in S. brachiata thus reduced the electron transfer from QA^– to QB pool-sized and activity of water-splitting complex. Higher φ(P₀) and F_V/F_M in S. brachiata under seawater salinity indicated a comparatively better quantum yield of primary photochemistry. The higher PI_Total in S. brachiata under 100% strength of seawater and 0.5 M NaCl stress indicated a better energy flux reaching to PSII RCs, electron transport and performance of RCs. The higher strengths of osmotic stress cause reduction in the quantum yield of PSII electron transport and capturing efficiency of excitation energy by open PSII RCs in S. brachiata.

Graphic Abstract

     

Chlorophyll a Fluorescence Transients in Mesophyll and Guard Cells : MODULATION OF GUARD CELL PHOTOPHOSPHORYLATION BY CO(2)

Chlorophyll fluorescence transients from mesophyll and guard cell chloroplasts of variegated leaves from Chlorophytum comosum were compared using high resolution fluorescence spectroscopy. Like their mesophyll counterparts, guard cell chloroplasts showed the OPS fluorescence transient indicating the operation of the linear electron transport and the possible generation of NADPH in these organelles. They also showed a slow fluorescence yield decrease, equivalent to the MT transition in mesophyll, suggesting the formation of the high energy state and photophosphorylation. Unlike the mesophyll chloroplasts, the fluorescence from guard cell chloroplasts lacked the increment of the SM transition, indicating that the two types of chloroplasts have some metabolic differences. The presence of CO₂ (supplied as bicarbonate, pH 6.7) specifically inhibited the MT-equivalent transition while its absence accelerated it. These observations constitute the first specific evidence of a guard cell chloroplast response to CO₂. Control of photosynthetic ATP levels in the guard cell cytoplasm by CO₂ may provide a mechanism regulating the availability of high energy equivalents at the guard cell plasmalemma, thus affecting stomatal opening.     

Chlorophyll a Fluorescence Predicts Total Photosynthetic Electron Flow to CO(2) or NO(3)/NO(2) under Transient Conditions

A model which predicts total photosynthetic electron flow from a linear regression of the relationship between corrected steady-state quantum yield and nonphotochemical quenching (E Weis, JA Berry [1987] Biochem Biophys Acta 894: 198-208) was formulated for N-limited cells of the green alga Selenastrum minutum. Unlike other models based on net CO₂ fixation, our model is based on total photosynthetic electron flow measured as gross O₂ evolution. This allowed for the prediction of total photosynthetic electron flow from water to both CO₂ fixation and NO₃⁻/NO₂⁻ reduction. The linear regression equation predicting electron flow is of the form: J = I · Q_q[0.4777-0.3282 Q_NP] (where J = gross photosynthetic electron flow, I = incident PAR, Q_q = photochemical quenching, Q_NP = nonphotochemical quenching). During steady-state photosynthesis, over a range of irradiance, the model predicted a photosynthetic light saturation curve which was well correlated with that observed. Although developed under steady-state conditions, the model was tested during nonsteady-state photosynthesis induced by transient nitrogen assimilation. The model predicted transient rates of gross O₂ evolution which were in excellent agreement with the rates observed under a variety of conditions regardless of whether CO₂ or NO₃⁻/NO₂⁻ served as the physiological electron acceptor. The fluorescence transients resulting from ammonium and nitrate assimilation are discussed with respect to metabolic demands for reductant and ATP.     

Chlorophyll a Fluorescence Yield as a Monitor of Both Active CO(2) and HCO(3) Transport by the Cyanobacterium Synechococcus UTEX 625

Simultaneous measurements have been made of inorganic carbon accumulation (by mass spectrometry) and chlorophyll a fluorescence yield of the cyanobacterium Synechococcus UTEX 625. The accumulation of inorganic carbon by the cells was accompanied by a substantial quenching of chlorophyll a fluorescence. The quenching occurred even when CO₂ fixation was inhibited by iodoacetamide and whether the accumulation of inorganic carbon resulted from either active CO₂ or HCO₃⁻ transport. Measurement of chlorophyll a fluorescence yield of cyanobacteria may prove to be a rapid and convenient means of screening for mutants of inorganic carbon accumulation.     

Photosynthetic Acclimation to Elevated CO2 in Relation to Leaf Saccharide Constituents in Wheat and Sunflower

Wheat (T. durum cvs. HD 4502 and B 449, T. aestivum cvs. Kalyansona and Kundan) and sunflower (Helianthus annuus L. cv. Morden) were grown under atmospheric (360±10 cm³ m^–3, AC) and elevated CO₂ (650±50 cm³ m^–3, EC) concentration in open top chambers for entire period of growth and development till maturity. Leaf net photosynthetic rate (P _N) of EC-grown plants of wheat measured at EC was significantly decreased in comparison with AC-plants of wheat measured at EC. Sunflower, however, showed no significant depression in P _N in EC-plants. There was a decrease in ribulose-1,5-bisphosphate carboxylase (RuBPC) activity, its activation state and amount in EC-plants of wheat, whereas no significant decrease was observed in sunflower. The above different acclimation to EC in wheat and sunflower was related with saccharide constituents accumulated in the leaves. Under EC, sunflower accumulated in the leaves more starch, whereas wheat accumulated more sugars.     

Simultaneous Measurements of Steady State Chlorophyll a Fluorescence and CO(2) Assimilation in Leaves: The Relationship between Fluorescence and Photosynthesis in C(3) and C(4) Plants

Rates of CO₂ assimilation and steady state chlorophyll a fluorescence were measured simultaneously at different intercellular partial pressures of CO₂ in attached cotton (Gossypium hirsutum L. cv Deltapine 16) leaves at 25°C. Electron transport activity for CO₂ assimilation plus photorespiration was calculated for these experiments. Under light saturating (1750 microeinsteins per square meter per second) and light limiting (700 microeinsteins per square meter per second) conditions there was a good correlation between fluorescence and the calculated electron transport activity at 19 and 200 millibars O₂, and between fluorescence and rates of CO₂ assimilation at 19 millibars but not 200 millibars O₂. The values of fluorescence measured at about 220 microbars intercellular CO₂ were not greatly affected by increasing O₂ from 19 to 800 millibars. Fluorescence increased with light intensity at any one intercellular CO₂ partial pressure. But the values obtained for fluorescence, expressed as a ratio of the maximum fluorescence obtained in DCMU-treated tissue, over the same range of CO₂ partial pressure at 500 microeinsteins per square meter per second were similar to those obtained at 1000 and 2000 microeinsteins per square meter per second. There were two phases in the observed correlation between fluorescence and calculated electron transport activity: an initial inverse relationship at low CO₂ partial pressures which reversed to a positive correlation at higher values of CO₂ partial pressures. Similar results were observed in the C₃ species Helianthus annuus L., Phaseolus vulgaris L., and Brassica chinensis. In all C₄ species (Zea mays L., Sorghum bicolor L., Panicum maximum Jacq., Amaranthus edulis Speg., and Echinochloa frumentacea [Roxb.] Link) examined changes in fluorescence were directly correlated with changes in CO₂ assimilation rates. The nature and the extent to which Q (primary quencher) and high-energy state (q_E) quenching function in determining the steady state fluorescence obtained during photosynthesis in leaves is discussed.     

Ethylene Contamination of CO(2) Cylinders: Effects on Plant Growth in CO(2) Enrichment Studies

The mechanical extensibilities of stage IVb Phycomyces were measured before and after a humidified wind stimulus. We find that when the humidity of the wind is greater than that of the ambient air, there is an increase in the mechanical extensibility of the cell wall. We also find that a step decrease in wind humidity results in a decrease in the mechanical extensibility of the cell wall.     

渗透胁迫及信号处理对海南粗榧叶绿素荧光特性的影响

为了探究甘露醇、氯化钠的渗透胁迫和脱落酸、茉莉酸甲酯、乙烯利、水杨酸等信号物质对海南粗榧(Cephalotaxus hainanensis)叶片叶绿素荧光特性的影响,本研究以海南粗榧新鲜叶片为试材,分别选用甘露醇、氯化钠、脱落酸、茉莉酸甲酯、乙烯利、水杨酸处理海南粗榧叶片,测定PSⅡ实际量子产量Y(Ⅱ)、非光化学淬灭系数(NvPQ)、光化学淬灭系数(qP)等叶绿素荧光参数.结果表明:甘露醇处理和氯化钠处理在短时间内可以造成这些参数的波动,然而在24 h后,这些参数趋于平稳,大多数可以恢复到对照水平,但渗透胁迫处理后的Y(Ⅱ)值显著降低,NPQ值却显著升高.在脱落酸、茉莉酸甲酯、乙烯利、水杨酸等信号物质处理后,叶绿素荧光参数多呈现出和甘露醇处理或氯化钠处理相似的趋势,表明了渗透胁迫可能通过不同的信号转导通路影响叶绿素荧光特性,其中水杨酸信号通路可能参与了叶片实际光能转换效率适应调节.     

土壤干旱下不同抗旱性小麦品种的渗透调节和渗透调节物质

在土壤干旱下抗旱性强的小麦品种的渗透调节能力大于抗旱性弱的品种。土壤缓慢干旱时,几种主要渗透调节物质增加的先后次序为:Pro,K~ ,然后是可溶性糖和其他游离氨基酸,后两者几乎平行增加、在土壤干旱下的渗透调节物质中,有机溶质为可溶性糖、其他游离氨基酸、Pro和Mal;无机离子为K~ ,Ca~(2 ),Mg~(2 )等。对渗透调节的相对贡献率为:K~ >可溶性糖>其他游离氨基酸>Ca~(2 )>Mg~(2 )>Pro。六种物质总和对渗透调节的相对贡献率为:轻度干旱56％～70％;中度干旱70％～79％;严重干旱79％～88％。     

Photosynthesis and Growth of Water Hyacinth under CO(2) Enrichment

Water hyacinth (Eichhornia crassipes [Mart.] Solms) plants were grown in environmental chambers at ambient and enriched CO₂ levels (330 and 600 microliters CO₂ per liter). Daughter plants (ramets) produced in the enriched CO₂ gained 39% greater dry weight than those at ambient CO₂, but the original mother plants did not. The CO₂ enrichment increased the number of leaves per ramet and leaf area index, but did not significantly increase leaf size or the number of ramets formed. Flower production was increased 147%. The elevated CO₂ increased the net photosynthetic rate of the mother plants by 40%, but this was not maintained as the plants acclimated to the higher CO₂ level. After 14 days at the elevated CO₂, leaf resistance increased and transpiration decreased, especially from the adaxial leaf surface. After 4 weeks in elevated as compared to ambient CO₂, ribulose bisphosphate carboxylase activity was 40% less, soluble protein content 49% less, and chlorophyll content 26% less; whereas starch content was 40% greater. Although at a given CO₂ level the enriched CO₂ plants had only half the net photosynthetic rate of their counterparts grown at ambient CO₂, they showed similar internal CO₂ concentrations. This suggested that the decreased supply of CO₂ to the mesophyll, as a result of the increased stomatal resistance, was counterbalanced by a decreased utilization of CO₂. Photorespiration and dark respiration were lower, such that the CO₂ compensation point was not altered. The photosynthetic light and CO₂ saturation points were not greatly changed, nor was the O₂ inhibition of photosynthesis (measured at 330 microliters CO₂ per liter). It appears that with CO₂ enrichment the temporary increase in net photosynthesis produced larger ramets. After acclimation, the greater total ramet leaf area more than compensated for the lower net photosynthetic rate on a unit leaf area basis, and resulted in a sustained improvement in dry weight gain.     

Acclimation of Photosynthesis to Elevated CO(2) in Five C(3) Species

The effect of long-term (weeks to months) CO₂ enhancement on (a) the gas-exchange characteristics, (b) the content and activation state of ribulose-1,5-bisphosphate carboxylase (rubisco), and (c) leaf nitrogen, chlorophyll, and dry weight per area were studied in five C₃ species (Chenopodium album, Phaseolus vulgaris, Solanum tuberosum, Solanum melongena, and Brassica oleracea) grown at CO₂ partial pressures of 300 or 900 to 1000 microbars. Long-term exposure to elevated CO₂ affected the CO₂ response of photosynthesis in one of three ways: (a) the initial slope of the CO₂ response was unaffected, but the photosynthetic rate at high CO₂ increased (S. tuberosum); (b) the initial slope decreased but the CO₂-saturated rate of photosynthesis was little affected (C. album, P. vulgaris); (c) both the initial slope and the CO₂-saturated rate of photosynthesis decreased (B. oleracea, S. melongena). In all five species, growth at high CO₂ increased the extent to which photosynthesis was stimulated following a decrease in the partial pressure of O₂ or an increase in measurement CO₂ above 600 microbars. This stimulation indicates that a limitation on photosynthesis by the capacity to regenerate orthophosphate was reduced or absent after acclimation to high CO₂. Leaf nitrogen per area either increased (S. tuberosum, S. melongena) or was little changed by CO₂ enhancement. The content of rubisco was lower in only two of the five species, yet its activation state was 19% to 48% lower in all five species following long-term exposure to high CO₂. These results indicate that during growth in CO₂-enriched air, leaf rubisco content remains in excess of that required to support the observed photosynthetic rates.     

Changes in Osmotic Pressure and Mucilage during Low-Temperature Acclimation of Opuntia ficus-indica

Opuntia ficus-indica, a Crassulacean acid metabolism plant cultivated for its fruits and cladodes, was used to examine chemical and physiological events accompanying low-temperature acclimation. Changes in osmotic pressure, water content, low molecular weight solutes, and extracellular mucilage were monitored in the photosynthetic chlorenchyma and the water-storage parenchyma when plants maintained at day/night air temperatures of 30/20°C were shifted to 10/0°C. An increase in osmotic pressure of 0.13 megapascal occurred after 13 days at 10/0°C. Synthesis of glucose, fructose, and glycerol accounted for most of the observed increase in osmotic pressure during the low-temperature acclimation. Extracellular mucilage and the relative apoplastic water content increased by 24 and 10%, respectively, during exposure to low temperatures. These increases apparently favor the extracellular nucleation of ice closer to the equilibrium freezing temperature for plants at 10/0°C, which could make the cellular dehydration more gradual and less damaging. Nuclear magnetic resonance studies helped elucidate the cellular processes during ice formation, such as those revealed by changes in the relaxation times of two water fractions in the chlorenchyma. The latter results suggested a restricted mobility of intracellular water and an increased mobility of extracellular water for plants at 10/0°C compared with those at 30/20°C. Increased mobility of extracellular water could facilitate extracellular ice growth and thus delay the potentially lethal intracellular freezing during low-temperature acclimation.