两优培九功能叶光反应特性和对低剂量UV-B辐射增强的光合响应机制
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摘要
水稻生殖生长期是夺取高产的主要阶段。其中,最上部3片功能叶光合产物占水稻经济产量的70%-80%,能为产量形成提供良好的基础。在在野外条件下,利用荧光动力学分析和生理生化研究技术,对生殖生长期超高产杂交稻两优培九和大面积推广稻汕优63功能叶原初反应、电子传递和光合磷酸化水平进行系统研究,比较光反应特性。结果表明：
(1)与汕优63相比,两优培九功能叶叶绿素含量高22.90%,光合功能期长约50%,叶绿素a/b比值高36.78%；
(2)光能吸收与分配方面,两优培九功能叶单位叶面积吸收的光能不占优势,但保持高吸收的稳定期长,有活性的反应中心数量多,热耗散的能量比例较少,进入电子传递链的能量高；
(3)荧光参数分析发现两优培九PSII供体侧、受体侧和反应中心性能优良,光能吸收、传递和转化为电能效率较高；
(4)叶绿体放氧活性、电子传递链和光合磷酸化活性均显著高于汕优63,表明两优培九电能转化为活跃化学能能力强。
两优培九正是在光反应的各个步骤中占有优势,拥有较高的光合效率和较长的光合功能期,奠定了其超高产的坚实基础。
空气污染导致平流层臭氧浓度下降,紫外线B到达地球表面。UV-B辐射的增加,会导致植物的生长发育发生一系列的变化,如影响植物形态结构包括株高、根冠比、叶片大小、生物产量等,对光系统产生伤害,引起基因表达和蛋白酶活性下降。
本文在两优培九光反应特性研究基础上,采用剂量为5.4kJ m-2d-1的低强度UV-B辐射胁迫水稻,系统地对生殖生长期功能叶光反应、碳同化、叶绿体超微结构和类囊体膜蛋白进行了详细研究,结果表明：
(1)光合生理指标显示两优培九上三片功能叶在剑叶露芽(DAE)15天后整体开始进入衰老期。与对照相比,UV-B处理后功能叶叶绿素含量较高,但净光合速率和气孔导度在剑叶露芽25天前显著下降。
(2)UV-B处理下,PSII在光能吸收、传递和转化上表现较好：单位面积光量子吸收(ABS/CSM)、光量子流捕获(TRo/CSM)、潜在电子传递能力(ETo/CSM)和最大量子产量(Fv/Fm)均增加,同时热耗散(DIo/CSM)较少。这可能是UV-B处理的植株PSII具有更高光化学活性的主要原因。
(3)虽然全电子链电子传递活性没有显著差异。但是,处理组功能叶Ca2+-ATP酶活性(Ca2+-ATPase)和光合磷酸化活性一直较低,说明碳同化力形成较少。UV-B处理功能叶中,暗反应关键酶Rubisco活性显著下降可能是光合效率较低的主要原因。
(4)淀粉粒聚集,脂质小滴和胞质小球增多是叶绿体受UV-B胁迫损伤的表现。但是,生殖生长后期处理组叶绿体膜裂解较晚,膜内结构较完整致密,叶绿素含量下降较慢和较多基质类囊体有序紧密排列,可能是UV-B处理下两优培九具有较好原初光反应活性的主要原因。叶绿体结构损伤程度和光合作用受抑制程度相一致。
(5)在UV-B辐射增强条件下,研究了功能叶抗氧化系统的变化。结果表明,UV-B辐射胁迫使功能叶抗氧化物酶超氧化物歧化酶(SOD)和过氧化氢酶(CAT)的活性先升后降再上升,过氧化物酶(POD)活性上升；抗氧化物质类还原型谷胱甘肽(GSH)含量和还原型抗坏血酸(ASA)含量提高,但增幅逐渐减少。功能叶活性氧(O2-)累积,丙二醛(MDA)相对含量先升高后下降,可溶性蛋白含量则相反。抗氧化系统变化幅度和光合生理指标变化基本一致。
(6)蛋白质组学研究发现两优培九功能叶类囊体膜对低强度UV-B的光合响应包括：(a)PSII的结构蛋白和重要组成蛋白未受损伤,同时与PSII关系密切的辅因子含量有所提高,表明PSII的光能吸收和转化能力得到加强。(b)ATP合酶各亚基蛋白含量的上升,说明胁迫促进了ATP的水解速率,以减缓过量光能积累造成的损伤。(c)调控碳在淀粉和蔗糖之间分配,参与卡尔文循环和糖酵解过程的关键酶蛋白含量显著下降,表明卡尔文循环和糖酵解循环受到了明显抑制。(d)胁迫响应蛋白的大量增加,开启了不同的抗逆信号传导途径。
The chloroplast light reaction characteristics were compared between super-high-yield hybrid rice Liangyoupeijiu and traditional hybrid rice Shanyou63in order to provide theoretical insights into physiological basis for high yield. Using fluorescence dynamics analysis and physiological and biochemical research techniques, in the field we systematically studied the primary response, electron transport chain and photophosphorylation during the reproductive period. The results showed that,
(1) As compared to Shanyou63, chlorophyll content in functional leaves of Liangyoupeijiu was22.90%higher, it had a longer photosynthetic function duration and chla/chlb ratio was36.78%higher;
(2) In the term of light absorption and distribution, there was no significant difference in light absorption per unit leaf area of functional leaves, but Liangyoupeijiu maintained a high and long stability period of light energy absorption capacity, the number of active reaction centers was more, the energy of heat dissipation was relatively fewer, the energy transferred into the electron transport chain was higher;
(3) Fluorescence analysis showed that structures and status of the body side, the receptor side and the reaction center in PSIIperformed better than Shanyou63, indicated that Liangyoupeijiu had a higher efficient in the transforming light energy into electric energy;
(4) In addition, oxygen evolution of the chloroplast, activities of electron transport chain and photophosphorylation were significantly higher, indicated that Liangyoupeijiu possessed some advantages in the energy converted into transforming electric energy into active chemical energy. For higher light energy absorption, transmission and conversion efficiency than Shanyou63, Liangyoupeijiu established a physicological basis of super-high-yield hybrid rice.
One of the most important aspects of global change is that of stratospheric ozone depletion resulting from air pollution and the resulting increase in UV-B radiation reaching the surface of the Earth. Enhanced UV-B radiation can lead to a series of changes in plant growth, such as to affect plant morphological structure including plant height, the ratio of root and shoot, leaf size and biological yield, to hurt light system, to cause expression of genes and activities of photosynthetic protein enzymes dropping.
In this paper, LYPJ was systematically studied the light reaction, carbon assimilation, chloroplast ultrastructure, the antioxidant system and thylakoid membrane proteins of the upper three functional leaves in detail during reproductive development based on the research light response characteristics of Liangyoupeijiu using the UV-B dose of5.4kJ m-2d-1to treat rice.The results were as follows:
(1) Photosynthesis physiological indices showed that the upper three functional leaves of LYP J entered into senescence approximately15days after flag leaf emergence (DAE). Compared with the control, the chlorophyll (Ch1) content was greater in UV-B treated leaves, but the net photosynthetic rate (PN) and stomatal conductance (gs) remained relatively stable at lower values before35DAE, which were significant differences.
(2) Increases in the absorption flux of photons per cross section (ABS/CSM), the phenomenological fluxes for trapping (TRo/CSM), the potential electron transport (ETo/CSm), the maximum quantum yield (Fv/Fm) and a decline in dissipation (DIo/CSM) might contribute to higher light energy absorption, transfer and the transformation of photosystem Ⅱ (PSII) in UV-B treated leaves.
(3) Besides, the results showed a not significant increase of the electron transport rate (ETR). However, Ca2+-ATPase and photophosphorylation activities of UV-B treated leaves maintained a relatively stable lower value, suggesting that the carbon assimilatory power was less. The significant decreased activity of ribulose-1,5-bisphosphate carboxylase (RuBPcase) was greatly associated with the decline in photosynthetic efficiency.
(4) Long term UV-B treatment delayed chloroplasts growth of flag leaves. Chloroplast membranes in UV-B treated flag leaves later swelled and disintegrated, and more stromal thylakoids were parallel to each other and were arranged in neat rows, which may be responsible for better manifest of the primary light reaction in UV-B treatment. It is likely that accumulation of starch and an increase in the number of lipid droplets and translucent plastoglobuli were results of inhibition of carbohydrate transport. Enhanced low intensity UV-B radiation tends to promote the absorption of light energy and the transport of electron, but result in the decrease of photophosphorylation and Rubisco activation. The extent of the damage to the chloroplast ultrastructure was consistent with the degree of inhibition of photosynthesis.
(5) The changes of antioxidant systems in functional leaves under enhancing ultraviolet-B radiation were investigated. The results showed that antioxidant enzymes superoxide dismutase (SOD) and catalase (CAT) activities increased signficiantly at first, then decreased, elevated at last. Peroxidase (POD) activities tended to increase; The contents of antioxidant substances ascorbic acid (ASA) and glutathione (GSH) increased under a long term UV-B stress, but amplitude was becoming smaller. The relative contents of hydrogen peroxide (O2-) and malon-dialhyde (MDA) increased at first, then slowly decreased. The reverse was true in soluble protein content. The results indicated that the change of the antioxidant systems was consistent with the performance of photosynthesis.
(6) Proteomics study showed that photosynthetic response mechanism of thylakoids membrane in functional leaves of LYPJ response to low intensity UV-B included the following aspects:(a) Structure and important constituent proteins of PSII were not damaged, at the same time cofactor content increased, that showed light energy absorption and transformation of the ability of PSII were strengthened.(b)Increases in content of ATPase subunits indicated the stress had promoted the hydrolysis rate of ATP in order to slow down light energy accumulation.(c) Key enzymes participated in the regulation in carbon distribution between starch and sucrose, the Calvin cycle and glycolysis, whose activities decreased significantly, that showed the Carbon assimilation was obviously inhibited.(d) A great increase of stress response proteins opened different signal transduction pathways.
(1)与汕优63相比,两优培九功能叶叶绿素含量高22.90%,光合功能期长约50%,叶绿素a/b比值高36.78%；
(2)光能吸收与分配方面,两优培九功能叶单位叶面积吸收的光能不占优势,但保持高吸收的稳定期长,有活性的反应中心数量多,热耗散的能量比例较少,进入电子传递链的能量高；
(3)荧光参数分析发现两优培九PSII供体侧、受体侧和反应中心性能优良,光能吸收、传递和转化为电能效率较高；
(4)叶绿体放氧活性、电子传递链和光合磷酸化活性均显著高于汕优63,表明两优培九电能转化为活跃化学能能力强。
两优培九正是在光反应的各个步骤中占有优势,拥有较高的光合效率和较长的光合功能期,奠定了其超高产的坚实基础。
空气污染导致平流层臭氧浓度下降,紫外线B到达地球表面。UV-B辐射的增加,会导致植物的生长发育发生一系列的变化,如影响植物形态结构包括株高、根冠比、叶片大小、生物产量等,对光系统产生伤害,引起基因表达和蛋白酶活性下降。
本文在两优培九光反应特性研究基础上,采用剂量为5.4kJ m-2d-1的低强度UV-B辐射胁迫水稻,系统地对生殖生长期功能叶光反应、碳同化、叶绿体超微结构和类囊体膜蛋白进行了详细研究,结果表明：
(1)光合生理指标显示两优培九上三片功能叶在剑叶露芽(DAE)15天后整体开始进入衰老期。与对照相比,UV-B处理后功能叶叶绿素含量较高,但净光合速率和气孔导度在剑叶露芽25天前显著下降。
(2)UV-B处理下,PSII在光能吸收、传递和转化上表现较好：单位面积光量子吸收(ABS/CSM)、光量子流捕获(TRo/CSM)、潜在电子传递能力(ETo/CSM)和最大量子产量(Fv/Fm)均增加,同时热耗散(DIo/CSM)较少。这可能是UV-B处理的植株PSII具有更高光化学活性的主要原因。
(3)虽然全电子链电子传递活性没有显著差异。但是,处理组功能叶Ca2+-ATP酶活性(Ca2+-ATPase)和光合磷酸化活性一直较低,说明碳同化力形成较少。UV-B处理功能叶中,暗反应关键酶Rubisco活性显著下降可能是光合效率较低的主要原因。
(4)淀粉粒聚集,脂质小滴和胞质小球增多是叶绿体受UV-B胁迫损伤的表现。但是,生殖生长后期处理组叶绿体膜裂解较晚,膜内结构较完整致密,叶绿素含量下降较慢和较多基质类囊体有序紧密排列,可能是UV-B处理下两优培九具有较好原初光反应活性的主要原因。叶绿体结构损伤程度和光合作用受抑制程度相一致。
(5)在UV-B辐射增强条件下,研究了功能叶抗氧化系统的变化。结果表明,UV-B辐射胁迫使功能叶抗氧化物酶超氧化物歧化酶(SOD)和过氧化氢酶(CAT)的活性先升后降再上升,过氧化物酶(POD)活性上升；抗氧化物质类还原型谷胱甘肽(GSH)含量和还原型抗坏血酸(ASA)含量提高,但增幅逐渐减少。功能叶活性氧(O2-)累积,丙二醛(MDA)相对含量先升高后下降,可溶性蛋白含量则相反。抗氧化系统变化幅度和光合生理指标变化基本一致。
(6)蛋白质组学研究发现两优培九功能叶类囊体膜对低强度UV-B的光合响应包括：(a)PSII的结构蛋白和重要组成蛋白未受损伤,同时与PSII关系密切的辅因子含量有所提高,表明PSII的光能吸收和转化能力得到加强。(b)ATP合酶各亚基蛋白含量的上升,说明胁迫促进了ATP的水解速率,以减缓过量光能积累造成的损伤。(c)调控碳在淀粉和蔗糖之间分配,参与卡尔文循环和糖酵解过程的关键酶蛋白含量显著下降,表明卡尔文循环和糖酵解循环受到了明显抑制。(d)胁迫响应蛋白的大量增加,开启了不同的抗逆信号传导途径。
The chloroplast light reaction characteristics were compared between super-high-yield hybrid rice Liangyoupeijiu and traditional hybrid rice Shanyou63in order to provide theoretical insights into physiological basis for high yield. Using fluorescence dynamics analysis and physiological and biochemical research techniques, in the field we systematically studied the primary response, electron transport chain and photophosphorylation during the reproductive period. The results showed that,
(1) As compared to Shanyou63, chlorophyll content in functional leaves of Liangyoupeijiu was22.90%higher, it had a longer photosynthetic function duration and chla/chlb ratio was36.78%higher;
(2) In the term of light absorption and distribution, there was no significant difference in light absorption per unit leaf area of functional leaves, but Liangyoupeijiu maintained a high and long stability period of light energy absorption capacity, the number of active reaction centers was more, the energy of heat dissipation was relatively fewer, the energy transferred into the electron transport chain was higher;
(3) Fluorescence analysis showed that structures and status of the body side, the receptor side and the reaction center in PSIIperformed better than Shanyou63, indicated that Liangyoupeijiu had a higher efficient in the transforming light energy into electric energy;
(4) In addition, oxygen evolution of the chloroplast, activities of electron transport chain and photophosphorylation were significantly higher, indicated that Liangyoupeijiu possessed some advantages in the energy converted into transforming electric energy into active chemical energy. For higher light energy absorption, transmission and conversion efficiency than Shanyou63, Liangyoupeijiu established a physicological basis of super-high-yield hybrid rice.
One of the most important aspects of global change is that of stratospheric ozone depletion resulting from air pollution and the resulting increase in UV-B radiation reaching the surface of the Earth. Enhanced UV-B radiation can lead to a series of changes in plant growth, such as to affect plant morphological structure including plant height, the ratio of root and shoot, leaf size and biological yield, to hurt light system, to cause expression of genes and activities of photosynthetic protein enzymes dropping.
In this paper, LYPJ was systematically studied the light reaction, carbon assimilation, chloroplast ultrastructure, the antioxidant system and thylakoid membrane proteins of the upper three functional leaves in detail during reproductive development based on the research light response characteristics of Liangyoupeijiu using the UV-B dose of5.4kJ m-2d-1to treat rice.The results were as follows:
(1) Photosynthesis physiological indices showed that the upper three functional leaves of LYP J entered into senescence approximately15days after flag leaf emergence (DAE). Compared with the control, the chlorophyll (Ch1) content was greater in UV-B treated leaves, but the net photosynthetic rate (PN) and stomatal conductance (gs) remained relatively stable at lower values before35DAE, which were significant differences.
(2) Increases in the absorption flux of photons per cross section (ABS/CSM), the phenomenological fluxes for trapping (TRo/CSM), the potential electron transport (ETo/CSm), the maximum quantum yield (Fv/Fm) and a decline in dissipation (DIo/CSM) might contribute to higher light energy absorption, transfer and the transformation of photosystem Ⅱ (PSII) in UV-B treated leaves.
(3) Besides, the results showed a not significant increase of the electron transport rate (ETR). However, Ca2+-ATPase and photophosphorylation activities of UV-B treated leaves maintained a relatively stable lower value, suggesting that the carbon assimilatory power was less. The significant decreased activity of ribulose-1,5-bisphosphate carboxylase (RuBPcase) was greatly associated with the decline in photosynthetic efficiency.
(4) Long term UV-B treatment delayed chloroplasts growth of flag leaves. Chloroplast membranes in UV-B treated flag leaves later swelled and disintegrated, and more stromal thylakoids were parallel to each other and were arranged in neat rows, which may be responsible for better manifest of the primary light reaction in UV-B treatment. It is likely that accumulation of starch and an increase in the number of lipid droplets and translucent plastoglobuli were results of inhibition of carbohydrate transport. Enhanced low intensity UV-B radiation tends to promote the absorption of light energy and the transport of electron, but result in the decrease of photophosphorylation and Rubisco activation. The extent of the damage to the chloroplast ultrastructure was consistent with the degree of inhibition of photosynthesis.
(5) The changes of antioxidant systems in functional leaves under enhancing ultraviolet-B radiation were investigated. The results showed that antioxidant enzymes superoxide dismutase (SOD) and catalase (CAT) activities increased signficiantly at first, then decreased, elevated at last. Peroxidase (POD) activities tended to increase; The contents of antioxidant substances ascorbic acid (ASA) and glutathione (GSH) increased under a long term UV-B stress, but amplitude was becoming smaller. The relative contents of hydrogen peroxide (O2-) and malon-dialhyde (MDA) increased at first, then slowly decreased. The reverse was true in soluble protein content. The results indicated that the change of the antioxidant systems was consistent with the performance of photosynthesis.
(6) Proteomics study showed that photosynthetic response mechanism of thylakoids membrane in functional leaves of LYPJ response to low intensity UV-B included the following aspects:(a) Structure and important constituent proteins of PSII were not damaged, at the same time cofactor content increased, that showed light energy absorption and transformation of the ability of PSII were strengthened.(b)Increases in content of ATPase subunits indicated the stress had promoted the hydrolysis rate of ATP in order to slow down light energy accumulation.(c) Key enzymes participated in the regulation in carbon distribution between starch and sucrose, the Calvin cycle and glycolysis, whose activities decreased significantly, that showed the Carbon assimilation was obviously inhibited.(d) A great increase of stress response proteins opened different signal transduction pathways.
引文
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