硅提高水稻对高锌胁迫抗性的生理与分子机理
摘要
水稻是我国的主要粮食作物,在我国的种植面积已达到4.5亿亩,每年生产的稻谷产量占我国粮食总产的44%左右。近年来,随着工业“三废”的排放,城市生活污水、垃圾以及不合理施用含有锌的农药和化肥,土壤锌污染对水稻造成的毒害作用日益严重。硅肥对植物的生长和发育是有益的,特别是在土壤重金属污染方面,目前有关Si缓解重金属锌毒害的研究仍处于初步阶段,其具体机制仍需进一步研究。
因此本报告采用水培试验,以根系耐性指数为指标,从19个水稻品种中筛选出2个受高锌胁迫后有显著差异的品种,TY-167(耐锌型)和FYY-326(锌敏感型)。利用这两个品种为试验材料,研究了硅对高锌胁迫下水稻的生长、元素吸收、运输及分布、光合作用、抗氧化体统的影响；利用敏感型品种为材料,利用高通量测序技术进一步分析了高锌胁迫下加硅和不加硅处理对水稻基因表达图谱；利用实时定量PCR技术,研究了高锌胁迫下加硅和不加硅处理对有关光合作用相关基因的调控。主要结果如下：
一、硅对锌胁迫下水稻根系抗氧化系统和过氧化伤害的缓解效应
高锌胁迫下,显著抑制了植物超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)活性的变化,诱导了两个水稻品种根部过氧化氢(H202)和丙二醛(MDA)含量的迅速增加,高锌胁迫诱导了植物细胞死亡,导致根系质膜完整性遭到破坏,和脂质过氧化伤害。加硅后上述指标都发生相反的趋势变化,加Si缓解了质膜完整性破坏和质脂过氧化伤害。这种缓解效应在耐性品种TY-167根系中比敏感型品种FYY-326品种根系显著。结果都表示TY-167根系与FYY-326相比不易于遭受Zn胁迫诱导的脂膜过氧化伤害。
二、硅对锌胁迫下水稻根系形态参数变化及超微结构的影响
锌在两个水稻品种地上部和地下部含量随着营养液中锌浓度升高而增加,高锌胁迫下,根表总面积、总根长和总根尖数量根系形态参数的在高锌胁迫下这些参数显著降低,根尖超微结构发生变化,细胞壁呈现不规则的增厚,出现质壁分离现象,细胞器融合,模糊不清,部分细胞液泡化程度提高,抑制了水稻地上部和地下部的生物量,但是加硅处理后,减少水稻对地上部对锌的吸收和抑制锌从地下部-地上部迁移,根系形态参数都显著增加,细胞壁与高锌胁迫相比变得较为清晰,质壁分离不显著,抵消了锌胁迫对植物生长的抑制作用,提高了水稻的生物量。
三、硅对锌胁迫下水稻元素相对含量与分布的影响
高锌胁迫下加硅或不加硅,锌主要累积在水稻根系,高锌处理导致水稻K、Ca、Mn、Fe、Cu相对含量的变化,影响体内K、Ca、Mn、Fe、Cu元素的相对吸收和运输,水稻体内这些元素的相对含量平衡遭到破坏,加硅处理降低Zn在水稻体内地上部的积累和分布,增加了地下部对Zn的积累,减少了锌从地下部向地上部迁移；抑制了某些元素在水稻体内的不平衡移动,减缓Zn造成的毒害。
四、硅对锌胁迫下水稻叶片毒害的缓解效应
高锌胁迫下,两水稻叶片变黄,从老叶逐渐向新叶转移,特别是敏感型FYY-326,降低了SOD、CAT和APX酶活性,诱导了两个水稻叶片H2O2、MDA、抗坏血酸(AsA)、谷胱甘肽(GSH)和非蛋白巯基(NPT)含量的迅速增加。加硅后除AsA、GSH和NPT含量仍表现进一步增加的趋势外,其它指标都发生相反的趋势变化。说明加硅缓解了锌诱导的水稻叶片毒害,而且这种缓解效应在耐锌型品种TY-167叶片中比敏感型品种FYY-326叶片显著。
五、硅对锌胁迫下水稻光合作用、荧光参数及叶绿体超微结构的影响
高锌胁迫下,两个水稻品种的净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)、水分利用率(WUE)、叶绿素含量和叶绿素荧光光谱中Fv/Fm比值都呈下降的趋势,细胞间隙CO2浓度(Ci)在高锌处理下增加。叶绿体的基质片层结构破坏,片层方向紊乱,加硅处理后上述指标都表现出相反的趋势,而且使叶绿体结构也变的完整,基质片层排列较为有序。说明加硅处理提高了水稻的光合效率,改善了叶绿体超微结构的变化。而这种缓解效应在耐锌型品种中表现的效果更加显著。
六、硅对锌胁迫下水稻基因表达图谱的影响
硅在正常(非胁迫)条件下能使56798个水稻基因中的2077个基因出现表达差异,其中101个基因表达上调,另外1976个基因表达下调；重金属胁迫72小时后,能诱导3888个基因出现差异,其中1920个基因表达上调,另外1968个基因表达下调；高锌胁迫下,硅使873个基因表达差异,其中504个基因表达上调,369个基因表达下调；硅存在下,高锌胁迫使4976个基因表达差异,其中4115个基因表达上调,861个基因表达下调。这些差异表达的基因涉及到代谢调节、离子转运、信号传导、转录调节和逆境应答等方面。高通量测序的结果表明,硅在正常(非胁迫)条件下对植物生长代谢具有明显作用,至少对于水稻这类单子叶植物是必需的,当植物受到重金属胁迫时,硅能调控植物本身产生更高效或更同步的关键基因减轻重金属对植物的伤害。
七、硅对高锌胁迫下与水稻光合作用相关基因表达的影响
高锌胁迫下,无论加硅与否,HemA、Os03g36540、PsaH、PetC和PsbY基因的相对表达均上升；但是在高锌胁迫初期硅能更早更快提高这些基因的相对表达,增长幅度也显著高于不施硅处理。高锌胁迫下Os02g49870基因的相对表达变化不大,施硅处理后显著提高了其相对表达。说明在高锌胁迫下,硅积极参与了与水稻光合作用相关的基因的激活和调控,从而提高了水稻的光合作用。
Rice is the main food crop in China, with the planting area reached30million ha. The annual rice yield accounts for about44%of the total grain output. In recent years, soil contamination with heavy metal Zinc (Zn) has become more and more serious because of the impacts of industrial wastes, sewage sludge, compost, agrochemicals, and mine tailings, leading to toxicity of excess Zn to rice growth. Silicon (Si) has been proven to be beneficial for healthy growth and development of many plant species and plays an important role in enhancing plant resistance against heavy metal, however, the mechanisms underpinning Si-enhanced metal tolerance remain poorly understood and thus need further investigation.
Therefore, hydroponics experiments were carried out in this study. Two rice cultivars:i.e. TY-167, a Zn-resistance cultivar and FYY-326, a Zn-sensitive cultivar, were selected among19rice genotypes based on the root tolerance index to investigate the effects of Si on plant growth, elements uptake, transport, distribution, photosynthetic parameters and antioxidative defenses of both cultivars under Zn stress; Using high-throughput sequencing, we performed a comprehensive analysis of the influence of Zn on gene expression of the sensitive rice (FYY-326) with or without Si. The effect of Si on regulating photosynthesis-related genes of the sensitive rice under high Zn stress with or without Si was conducted using real-time quantitative PCR approach. The main results are presented as follows:
1. Effect of Si on antioxidant defense capacity and oxidative of rice in roots under Zn stress
Superoxide dismutase (SOD), catalase (CAT) and asorbate peroxidases (APX) activities were decreased by high Zn stress; the contents of hydrogen peroxide (H2O2) and malondialdehyde (MDA) were increased by high Zn treatment, which were all counteracted by addition of Si. Furthermore, such effects were confirmed by using histochemical staining methods. High Zn stress induced the loss of plasma membrane integrity and the damage of membranes due to lipid peroxidation, which was alleviated by Si. The alleviative effects of Si on Zn toxicity were more significant in the Zn-resistance cultivar than in the Zn-sensitive cultivar.
2. Effect of Si on root morphology and ultrastructure of rice in roots under Zn stress
Zn concentration in both shoots and roots of the two cultivars increased in response to an altered Zn supply in the nutrient solution. Total root surface area (TRSA), total root length (TRL) and total root tip amount (TRTA) were decreased under Zn stress, and transmission electron microscope (TEM) analysis showed that serious untrastructural damage was observed in both cultivar roots treated with Zn. Plasmolysis, concentrated cytoplasm, ambiguity of organelle were all symptoms of root cell damage under Zn stress. Shoot and root growth of both cultivars was negatively affected by a root environment with2mM Zn ion concentrations. However, addition of Si significantly decreased root-to-shoot Zn transport and increased the parameters of root morphology. And exogenous Si alleviated the plasmolysis of root cell, counteracted the inhibitory effect of Zn on plant growth and increased the biomass of rice.
3. Effect of Si on relative elements concentration and distribution in rice under Zn stress
High Zn stress dramatically increased the relative Zn concentration with Zn concentration being significantly higher in roots than in shoots irrespective of cultivar used and silicon added. The balance of K, Ca, Mn, Fe and Cu in shoots and roots of both cultivars tested was disturbed by high Zn stress. However, Si significantly decreased Zn uptake and root-to-shoot transport, improved the unbalanced distribution of some elements in rice, and alleviated Zn toxicity to rice.
4. Effect of Si on antioxidant defence capacity in rice plants under Zn stress
The symptoms of Zn toxicity were typically manifested as a yellow colour on the lower leaves starting from the tips and spreading toward the bases of the leaves, which became severer as the experiment continued. Severer leafy symptoms of Zn toxicity were observed in the Zn-sensitive cultivar (FYY-326) than in the Zn-resistant cultivar (TY-167). Under Zn stress, the activities of SOD, CAT, APX decreased, but the contents of H2O2, MDA, ascorbic acid (AsA), glutathione (GSH) and non-protein thiols (NPT) except AsA, GSH and NPT were increased by Zn treatments, which were all counteracted by addition of Si. The toxicity symptoms of Zn were alleviated by addition of Si. The alleviative effects of Si on Zn toxicity were more significant in the Zn-tolerant cultivar than in the Zn-sensitive cultivar.
5. Effect of Si on photosynthetic and fluorescence parameters and chloroplast ultrastructure in rice under Zn stress
Net photosynthetic rate (Pn), stomatal conductance (Gs), transpiration rate (Tr), water use efficiency (WUE), chlorophyll cocntent, the value of Fv/Fm of the two rice cultivars tested were all decreased under high Zn stress, while intercellular CO2concentration (Ci) was increased. Chloroplast structure was swollen observably with chloroplast granae being destroyed heavily under Zn-stress, which was all counteracted by addition of Si. The results showed that addition of Si improved the photosynthetic efficiency and alleviated the chloroplast ultrastructure in Zn-stressed rice. However, the alleviation of Si on above-mentioned parameters was smaller in Zn-sensitive plant cultivar than in Zn-resistant plant.
6. Effect of Si on expression of genes in shoots of rice under Zn stress
We identified2077genes out of56798rice genes which were differentially expressed in the Si-amended plants in the treatments without high Zn added (Si/CK), with101genes up-regulated and1976genes down-regulated. Among3888differentially-expressed genes in rice plants treated with high Zn (Zn/CK),1920genes were up-regulated and1968genes were down-regulated. In the high Zn-treated Si+plants,873genes were differentially expressed, compared with the high Zn-treated Si-plants (Si+Zn/Zn), with504genes up-regulated and369genes down-regulated. In high Zn-treated plants amended with Si,4976genes were differentially expressed, compared with the normal Zn-treated plants amended with Si (Si+Zn/Si), with4115genes up-regulated and861genes down-regulated. These differentially-expressed genes included metabolic regulation, ion transport, transcription and adversity response, etc. On the basis of analyzing expression profile of rice gens, it can be concluded that Si can exhibit obvious impacts on growth and development of plants, especially in rice plants under non-stressed condition, and can regulate natural resistance mechanisms to produce more efficient and timely key genes to alleviate damage caused by high Zn stress.
7. Effect of Si on genes responsible for photosynthesis of rice under Zn stress
The gene expression levels of HemA, Os03g36540, PsaH, PetC and PsbY all increased in plants treated with high Zn with or without Si. However, Si could increase expression of these genes earlier and faster in plants under high Zn stress in the early stages, and the increasing extent of expression was higher in Si+plants than in Si-plants. The gene expression of Os02g49870was not remarkable under high Zn stress, but was significantly increased by addition of Si. The results show that Si activated and regulated some photosynthesis-related genes in response to high Zn stress in rice, and consequently increased the photosynthesis.
因此本报告采用水培试验,以根系耐性指数为指标,从19个水稻品种中筛选出2个受高锌胁迫后有显著差异的品种,TY-167(耐锌型)和FYY-326(锌敏感型)。利用这两个品种为试验材料,研究了硅对高锌胁迫下水稻的生长、元素吸收、运输及分布、光合作用、抗氧化体统的影响；利用敏感型品种为材料,利用高通量测序技术进一步分析了高锌胁迫下加硅和不加硅处理对水稻基因表达图谱；利用实时定量PCR技术,研究了高锌胁迫下加硅和不加硅处理对有关光合作用相关基因的调控。主要结果如下：
一、硅对锌胁迫下水稻根系抗氧化系统和过氧化伤害的缓解效应
高锌胁迫下,显著抑制了植物超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)活性的变化,诱导了两个水稻品种根部过氧化氢(H202)和丙二醛(MDA)含量的迅速增加,高锌胁迫诱导了植物细胞死亡,导致根系质膜完整性遭到破坏,和脂质过氧化伤害。加硅后上述指标都发生相反的趋势变化,加Si缓解了质膜完整性破坏和质脂过氧化伤害。这种缓解效应在耐性品种TY-167根系中比敏感型品种FYY-326品种根系显著。结果都表示TY-167根系与FYY-326相比不易于遭受Zn胁迫诱导的脂膜过氧化伤害。
二、硅对锌胁迫下水稻根系形态参数变化及超微结构的影响
锌在两个水稻品种地上部和地下部含量随着营养液中锌浓度升高而增加,高锌胁迫下,根表总面积、总根长和总根尖数量根系形态参数的在高锌胁迫下这些参数显著降低,根尖超微结构发生变化,细胞壁呈现不规则的增厚,出现质壁分离现象,细胞器融合,模糊不清,部分细胞液泡化程度提高,抑制了水稻地上部和地下部的生物量,但是加硅处理后,减少水稻对地上部对锌的吸收和抑制锌从地下部-地上部迁移,根系形态参数都显著增加,细胞壁与高锌胁迫相比变得较为清晰,质壁分离不显著,抵消了锌胁迫对植物生长的抑制作用,提高了水稻的生物量。
三、硅对锌胁迫下水稻元素相对含量与分布的影响
高锌胁迫下加硅或不加硅,锌主要累积在水稻根系,高锌处理导致水稻K、Ca、Mn、Fe、Cu相对含量的变化,影响体内K、Ca、Mn、Fe、Cu元素的相对吸收和运输,水稻体内这些元素的相对含量平衡遭到破坏,加硅处理降低Zn在水稻体内地上部的积累和分布,增加了地下部对Zn的积累,减少了锌从地下部向地上部迁移；抑制了某些元素在水稻体内的不平衡移动,减缓Zn造成的毒害。
四、硅对锌胁迫下水稻叶片毒害的缓解效应
高锌胁迫下,两水稻叶片变黄,从老叶逐渐向新叶转移,特别是敏感型FYY-326,降低了SOD、CAT和APX酶活性,诱导了两个水稻叶片H2O2、MDA、抗坏血酸(AsA)、谷胱甘肽(GSH)和非蛋白巯基(NPT)含量的迅速增加。加硅后除AsA、GSH和NPT含量仍表现进一步增加的趋势外,其它指标都发生相反的趋势变化。说明加硅缓解了锌诱导的水稻叶片毒害,而且这种缓解效应在耐锌型品种TY-167叶片中比敏感型品种FYY-326叶片显著。
五、硅对锌胁迫下水稻光合作用、荧光参数及叶绿体超微结构的影响
高锌胁迫下,两个水稻品种的净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)、水分利用率(WUE)、叶绿素含量和叶绿素荧光光谱中Fv/Fm比值都呈下降的趋势,细胞间隙CO2浓度(Ci)在高锌处理下增加。叶绿体的基质片层结构破坏,片层方向紊乱,加硅处理后上述指标都表现出相反的趋势,而且使叶绿体结构也变的完整,基质片层排列较为有序。说明加硅处理提高了水稻的光合效率,改善了叶绿体超微结构的变化。而这种缓解效应在耐锌型品种中表现的效果更加显著。
六、硅对锌胁迫下水稻基因表达图谱的影响
硅在正常(非胁迫)条件下能使56798个水稻基因中的2077个基因出现表达差异,其中101个基因表达上调,另外1976个基因表达下调；重金属胁迫72小时后,能诱导3888个基因出现差异,其中1920个基因表达上调,另外1968个基因表达下调；高锌胁迫下,硅使873个基因表达差异,其中504个基因表达上调,369个基因表达下调；硅存在下,高锌胁迫使4976个基因表达差异,其中4115个基因表达上调,861个基因表达下调。这些差异表达的基因涉及到代谢调节、离子转运、信号传导、转录调节和逆境应答等方面。高通量测序的结果表明,硅在正常(非胁迫)条件下对植物生长代谢具有明显作用,至少对于水稻这类单子叶植物是必需的,当植物受到重金属胁迫时,硅能调控植物本身产生更高效或更同步的关键基因减轻重金属对植物的伤害。
七、硅对高锌胁迫下与水稻光合作用相关基因表达的影响
高锌胁迫下,无论加硅与否,HemA、Os03g36540、PsaH、PetC和PsbY基因的相对表达均上升；但是在高锌胁迫初期硅能更早更快提高这些基因的相对表达,增长幅度也显著高于不施硅处理。高锌胁迫下Os02g49870基因的相对表达变化不大,施硅处理后显著提高了其相对表达。说明在高锌胁迫下,硅积极参与了与水稻光合作用相关的基因的激活和调控,从而提高了水稻的光合作用。
Rice is the main food crop in China, with the planting area reached30million ha. The annual rice yield accounts for about44%of the total grain output. In recent years, soil contamination with heavy metal Zinc (Zn) has become more and more serious because of the impacts of industrial wastes, sewage sludge, compost, agrochemicals, and mine tailings, leading to toxicity of excess Zn to rice growth. Silicon (Si) has been proven to be beneficial for healthy growth and development of many plant species and plays an important role in enhancing plant resistance against heavy metal, however, the mechanisms underpinning Si-enhanced metal tolerance remain poorly understood and thus need further investigation.
Therefore, hydroponics experiments were carried out in this study. Two rice cultivars:i.e. TY-167, a Zn-resistance cultivar and FYY-326, a Zn-sensitive cultivar, were selected among19rice genotypes based on the root tolerance index to investigate the effects of Si on plant growth, elements uptake, transport, distribution, photosynthetic parameters and antioxidative defenses of both cultivars under Zn stress; Using high-throughput sequencing, we performed a comprehensive analysis of the influence of Zn on gene expression of the sensitive rice (FYY-326) with or without Si. The effect of Si on regulating photosynthesis-related genes of the sensitive rice under high Zn stress with or without Si was conducted using real-time quantitative PCR approach. The main results are presented as follows:
1. Effect of Si on antioxidant defense capacity and oxidative of rice in roots under Zn stress
Superoxide dismutase (SOD), catalase (CAT) and asorbate peroxidases (APX) activities were decreased by high Zn stress; the contents of hydrogen peroxide (H2O2) and malondialdehyde (MDA) were increased by high Zn treatment, which were all counteracted by addition of Si. Furthermore, such effects were confirmed by using histochemical staining methods. High Zn stress induced the loss of plasma membrane integrity and the damage of membranes due to lipid peroxidation, which was alleviated by Si. The alleviative effects of Si on Zn toxicity were more significant in the Zn-resistance cultivar than in the Zn-sensitive cultivar.
2. Effect of Si on root morphology and ultrastructure of rice in roots under Zn stress
Zn concentration in both shoots and roots of the two cultivars increased in response to an altered Zn supply in the nutrient solution. Total root surface area (TRSA), total root length (TRL) and total root tip amount (TRTA) were decreased under Zn stress, and transmission electron microscope (TEM) analysis showed that serious untrastructural damage was observed in both cultivar roots treated with Zn. Plasmolysis, concentrated cytoplasm, ambiguity of organelle were all symptoms of root cell damage under Zn stress. Shoot and root growth of both cultivars was negatively affected by a root environment with2mM Zn ion concentrations. However, addition of Si significantly decreased root-to-shoot Zn transport and increased the parameters of root morphology. And exogenous Si alleviated the plasmolysis of root cell, counteracted the inhibitory effect of Zn on plant growth and increased the biomass of rice.
3. Effect of Si on relative elements concentration and distribution in rice under Zn stress
High Zn stress dramatically increased the relative Zn concentration with Zn concentration being significantly higher in roots than in shoots irrespective of cultivar used and silicon added. The balance of K, Ca, Mn, Fe and Cu in shoots and roots of both cultivars tested was disturbed by high Zn stress. However, Si significantly decreased Zn uptake and root-to-shoot transport, improved the unbalanced distribution of some elements in rice, and alleviated Zn toxicity to rice.
4. Effect of Si on antioxidant defence capacity in rice plants under Zn stress
The symptoms of Zn toxicity were typically manifested as a yellow colour on the lower leaves starting from the tips and spreading toward the bases of the leaves, which became severer as the experiment continued. Severer leafy symptoms of Zn toxicity were observed in the Zn-sensitive cultivar (FYY-326) than in the Zn-resistant cultivar (TY-167). Under Zn stress, the activities of SOD, CAT, APX decreased, but the contents of H2O2, MDA, ascorbic acid (AsA), glutathione (GSH) and non-protein thiols (NPT) except AsA, GSH and NPT were increased by Zn treatments, which were all counteracted by addition of Si. The toxicity symptoms of Zn were alleviated by addition of Si. The alleviative effects of Si on Zn toxicity were more significant in the Zn-tolerant cultivar than in the Zn-sensitive cultivar.
5. Effect of Si on photosynthetic and fluorescence parameters and chloroplast ultrastructure in rice under Zn stress
Net photosynthetic rate (Pn), stomatal conductance (Gs), transpiration rate (Tr), water use efficiency (WUE), chlorophyll cocntent, the value of Fv/Fm of the two rice cultivars tested were all decreased under high Zn stress, while intercellular CO2concentration (Ci) was increased. Chloroplast structure was swollen observably with chloroplast granae being destroyed heavily under Zn-stress, which was all counteracted by addition of Si. The results showed that addition of Si improved the photosynthetic efficiency and alleviated the chloroplast ultrastructure in Zn-stressed rice. However, the alleviation of Si on above-mentioned parameters was smaller in Zn-sensitive plant cultivar than in Zn-resistant plant.
6. Effect of Si on expression of genes in shoots of rice under Zn stress
We identified2077genes out of56798rice genes which were differentially expressed in the Si-amended plants in the treatments without high Zn added (Si/CK), with101genes up-regulated and1976genes down-regulated. Among3888differentially-expressed genes in rice plants treated with high Zn (Zn/CK),1920genes were up-regulated and1968genes were down-regulated. In the high Zn-treated Si+plants,873genes were differentially expressed, compared with the high Zn-treated Si-plants (Si+Zn/Zn), with504genes up-regulated and369genes down-regulated. In high Zn-treated plants amended with Si,4976genes were differentially expressed, compared with the normal Zn-treated plants amended with Si (Si+Zn/Si), with4115genes up-regulated and861genes down-regulated. These differentially-expressed genes included metabolic regulation, ion transport, transcription and adversity response, etc. On the basis of analyzing expression profile of rice gens, it can be concluded that Si can exhibit obvious impacts on growth and development of plants, especially in rice plants under non-stressed condition, and can regulate natural resistance mechanisms to produce more efficient and timely key genes to alleviate damage caused by high Zn stress.
7. Effect of Si on genes responsible for photosynthesis of rice under Zn stress
The gene expression levels of HemA, Os03g36540, PsaH, PetC and PsbY all increased in plants treated with high Zn with or without Si. However, Si could increase expression of these genes earlier and faster in plants under high Zn stress in the early stages, and the increasing extent of expression was higher in Si+plants than in Si-plants. The gene expression of Os02g49870was not remarkable under high Zn stress, but was significantly increased by addition of Si. The results show that Si activated and regulated some photosynthesis-related genes in response to high Zn stress in rice, and consequently increased the photosynthesis.
引文
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