一氧化氮对水稻幼苗镉毒害的缓解效应及其生理机制
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摘要
重金属镉(cadmium, Cd)是最主要的环境污染物之一,它会严重影响农作物的产量和品质。Cd对水稻的污染具有隐蔽性和危险性,它可以通过食物链给人民健康带来风险和危害。一氧化氮(nitricoxide, NO)是近年来备受关注的信号分子,它在植物信号转导及抗逆过程中发挥着重要作用。目前有关NO缓解植物重金属毒害的研究越来越多,但NO缓解植物重金属毒害的生理和分子机制鲜为人知。探讨外源NO对水稻Cd毒害的影响具有重要的理论和实践指导意义。
本研究以秀水63为试验材料,采用营养液培养的方法研究了外源NO对水稻Cd毒害的缓解作用,并从水稻生长、氧化胁迫、光合特性、矿质元素吸收、细胞超显微结构和蛋白组学等角度探讨了外源NO缓解水稻Cd毒害的生理及分子机制。主要研究结果如下：
(1)低浓度NO供体硝普钠(sodium nitroprusside, SNP,≤0.2mM)能够缓解Cd胁迫对水稻的毒害,而高浓度的SNP会抑制水稻的生长。0.005mM SNP能够缓解0.1mM Cd胁迫对水稻的毒害作用；0.1mM SNP能够缓解0.2mM Cd胁迫对水稻的毒害作用。具体表现为：外源NO能够缓解Cd毒害对水稻株高、根长和干重的抑制作用。同时也反映出外源NO缓解水稻Cd毒害具有浓度效应。
(2)外施NO能显著减少Cd诱导的膜脂过氧化程度和HO2的含量,还能降低水稻植株SOD、POD、APX、GR活性和ASA的含量,促进CAT活性的升高和GSH的含量。表明NO通过调节水稻受到的氧化胁迫来缓解Cd对水稻的毒害作用。
(3)外源NO能够增加Cd毒害下水稻叶片叶绿素的含量,提高叶片净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)、叶绿素最大荧光(Fm)、PSⅡ原初光能转换效率(Fv/Fm)、(?)合电子传递量子效率(ΦPSⅡ),降低叶片胞间CO2浓度(Ci),表明NO参与调节植物的光合作用,抑制Cd毒害对光系统损伤,降低热能耗散,缓解了Cd对水稻的毒害。
(4)外源NO可以使Cd毒害下水稻叶片叶绿体基粒类囊体片层结构变得密集,淀粉粒增加,液泡变小,液泡中沉积更多的电子富集颗粒,水稻Cd中毒现象得到一定缓解。
(5)外源NO降低了Cd毒害下水稻地上部Cd的含量,显著提高了水稻地上部Fe、Zn、Mn、Cu的含量以及根中K、Mg、Zn、Mn、Cu的含量,显著降低了根中Ca的含量。表明外源NO可能通过维持细胞质离子稳态平衡来缓解Cd对水稻生长发育的抑制作用。
(6)外源NO处理的Cd毒害下水稻叶片和根中成功鉴定出41个蛋白质。其中,水稻叶片中磷酸甘油酸激酶、3-磷酸甘油醛脱氢酶、ATP合酶、NADP-苹果酸、氨甲基转移酶蛋白均下调表达；单脱氢抗坏血酸还原酶、磷酸甘油酸变位酶和铁氧还蛋白-亚硝酸盐还原酶上调表达。水稻根中5-甲基四氢叶酸-高半胱氨酸S-甲基转移酶、热激蛋白、ATP合酶和蔗糖-二磷酸尿核苷葡萄糖基转移酶蛋白表达上调；果糖激酶、异黄酮还原酶和半胱氨酸结合β裂解酶蛋白表达下调。表明,NO参与了水稻光合作用、碳代谢、氮代谢、能量代谢、氧化胁迫、信号转导和细胞分裂等途径的调节。
Cadmium (Cd) is one of the most deleterious elements that pollute environment. Cd can be absorbed and accumulated easily by plants. Excess Cd typically causes direct or indirect inhibition of physiological processes, such as respiration, transpiration, photosynthesis, oxidative stress, cell elongation, nitrogen metabolism and mineral nutrition, resulting in growth retardation, leaf chlorosis and low biomass in plants. Cd can be absorbed and accumulated by rice without obvious deleterious symptoms, which gives rise to a threat on human health by food chain. Therefore, high accumulation of Cd in rice is a potential risk for both animals and human. As a crucial gaseous signaling molecule in plants, Nitric oxide (NO) plays a significant role in modulating several physiological and biochemical functions. In recent decades, an increasing number of articles have reported the effect of exogenous NO in alleviating heavy metal toxicity in different plants, although the physiological processes and mechanisms of NO in alleviating heavy metal toxicity are still far from clear. Therefore, it is necessary and meaningful to investigate the mechanisms of NO in alleviating Cd toxicity in rice, it will also provide significant theory basis to control the accumulation of Cd in rice seeds.
Japonica rice Xiushui63was used in the studies, a hydroponic experiment was conducted to investigate the effect of exogenous sodium nitroprusside (SNP), a NO donor, on alleviating the toxicity of Cd to rice. In order to elucidate the mechanisms of mitigate effect of NO application on Cd toxicity, growth parameters, reactive oxygen species, photosynthesis, mineral nutrition, ultrastruture and proteomic were researched. The main results were summarized as follows:
(1) Low concentrations of SNP (≤0.2mM) could alleviate the toxicity of Cd to rice, high concentrations of SNP deteriorated the toxicity of Cd to rice plants. A hydroponic experiment was conducted to investigate the effects of exogenous sodium nitroprusside (SNP), a NO donor, on growth in rice plants treated with0.1mM Cd and0.2mM Cd. The results concluded that the application of0.005mM SNP significantly alleviated0.1mM Cd-induced inhibition in rice seedlings growth, as plant height, root length, shoot weight and root weight. The application of0.1mM SNP significantly alleviated0.2mM Cd stress in rice growth. There is SNP concentration effect on alleviating Cd stress in rice plants.
(2) Exogenous NO significantly reduced the Cd-induced lipid peroxidation, hydrogen peroxide (H2O2) content and ascorbic acid (ASA) content in rice plants. The addition of NO reversed (only partially) the Cd-induced increase in activities of antioxidant enzymes-superoxide dismutase (SOD), peroxidase (POD), ascorbate peroxidase (APX) and glutathione reductase (GR). The addition of NO with Cd increased catalase (CAT) activity and glutathione (GSH) content. The exogenous application of NO could be advantageous against Cd toxicity, and could confer tolerance to heavy metal stress in rice plants.
(3) Exogenous NO significantly increased the chlorophyll content of rice leaves under Cd stress. NO significantly inhibited the decreases of net photosynthetic rate (Pn), transpiration rate (Tr), stomatal conductance (Gs), maximum fluorescence (Fm), PSII intrinsic photochemical efficiency (Fv/Fm), quantum yield of PSII (OPSII), and also inhibited the increase of intercellular CO2concentration (Ci). The results indicated that NO could modulate photosynthesis in plant, reduce energy dissipation and increase plant resistance by enhancing photosynthesis.
(4) Exogenous NO made chloroplasts become better or in relatively normal shape with parallel pattern of lamellae. Addition of NO with Cd, the number of vacuoles was increased, but with much smaller size, compared with Cd alone treatment. Moreover, more EDG were deposited in vacuoles.
(5) Exogenous NO decreased the Cd content in rice leaves, increased Cd content in root roots. NO significantly increased iron (Fe), zinc (Zn), manganese (Mn), copper (Cu) contents in rice shoot and potassium (K), magnesium (Mg), zinc (Zn), copper (Cu) contents in rice root; significantly decreased calcium (Ca) contents in rice roots induced by Cd stress. The results indicated that NO could maintain the mineral nutrition balance to mitigate Cd stress in rice plants.
(6) In rice leaves and roots,41proteins were identified under SNP treatment. Phosphoglycerate kinase, Glyceraldehyde-3-phosphate dehydrogenase B, ATP synthase, NADP dependent malic enzyme, and aminomethyl transferase in rice leaves were down-regulated expressed, while monodehydroascorbate reductase, phosphoglycerate mutase and ferredoxin-nitrite reductase were up-regulated expressed. In rice roots,5-methyltetrahydropteroyltriglutamate-homocysteine methyltransferase, heat shock protein, ATP synthase, and sucrose-UDP glucosyltransferase1were up-regulated expressed, Fructokinase II, isoflavone reductase and cysteine conjugate betalyase were down-regulated expressed. The results indicated that these proteins which participated in photosynthesis, carbohydrate metabolism, nitrogen metabolism, oxidative phosphorylation, oxidative stress responses, signal transductions and cell division.
本研究以秀水63为试验材料,采用营养液培养的方法研究了外源NO对水稻Cd毒害的缓解作用,并从水稻生长、氧化胁迫、光合特性、矿质元素吸收、细胞超显微结构和蛋白组学等角度探讨了外源NO缓解水稻Cd毒害的生理及分子机制。主要研究结果如下：
(1)低浓度NO供体硝普钠(sodium nitroprusside, SNP,≤0.2mM)能够缓解Cd胁迫对水稻的毒害,而高浓度的SNP会抑制水稻的生长。0.005mM SNP能够缓解0.1mM Cd胁迫对水稻的毒害作用；0.1mM SNP能够缓解0.2mM Cd胁迫对水稻的毒害作用。具体表现为：外源NO能够缓解Cd毒害对水稻株高、根长和干重的抑制作用。同时也反映出外源NO缓解水稻Cd毒害具有浓度效应。
(2)外施NO能显著减少Cd诱导的膜脂过氧化程度和HO2的含量,还能降低水稻植株SOD、POD、APX、GR活性和ASA的含量,促进CAT活性的升高和GSH的含量。表明NO通过调节水稻受到的氧化胁迫来缓解Cd对水稻的毒害作用。
(3)外源NO能够增加Cd毒害下水稻叶片叶绿素的含量,提高叶片净光合速率(Pn)、蒸腾速率(Tr)、气孔导度(Gs)、叶绿素最大荧光(Fm)、PSⅡ原初光能转换效率(Fv/Fm)、(?)合电子传递量子效率(ΦPSⅡ),降低叶片胞间CO2浓度(Ci),表明NO参与调节植物的光合作用,抑制Cd毒害对光系统损伤,降低热能耗散,缓解了Cd对水稻的毒害。
(4)外源NO可以使Cd毒害下水稻叶片叶绿体基粒类囊体片层结构变得密集,淀粉粒增加,液泡变小,液泡中沉积更多的电子富集颗粒,水稻Cd中毒现象得到一定缓解。
(5)外源NO降低了Cd毒害下水稻地上部Cd的含量,显著提高了水稻地上部Fe、Zn、Mn、Cu的含量以及根中K、Mg、Zn、Mn、Cu的含量,显著降低了根中Ca的含量。表明外源NO可能通过维持细胞质离子稳态平衡来缓解Cd对水稻生长发育的抑制作用。
(6)外源NO处理的Cd毒害下水稻叶片和根中成功鉴定出41个蛋白质。其中,水稻叶片中磷酸甘油酸激酶、3-磷酸甘油醛脱氢酶、ATP合酶、NADP-苹果酸、氨甲基转移酶蛋白均下调表达；单脱氢抗坏血酸还原酶、磷酸甘油酸变位酶和铁氧还蛋白-亚硝酸盐还原酶上调表达。水稻根中5-甲基四氢叶酸-高半胱氨酸S-甲基转移酶、热激蛋白、ATP合酶和蔗糖-二磷酸尿核苷葡萄糖基转移酶蛋白表达上调；果糖激酶、异黄酮还原酶和半胱氨酸结合β裂解酶蛋白表达下调。表明,NO参与了水稻光合作用、碳代谢、氮代谢、能量代谢、氧化胁迫、信号转导和细胞分裂等途径的调节。
Cadmium (Cd) is one of the most deleterious elements that pollute environment. Cd can be absorbed and accumulated easily by plants. Excess Cd typically causes direct or indirect inhibition of physiological processes, such as respiration, transpiration, photosynthesis, oxidative stress, cell elongation, nitrogen metabolism and mineral nutrition, resulting in growth retardation, leaf chlorosis and low biomass in plants. Cd can be absorbed and accumulated by rice without obvious deleterious symptoms, which gives rise to a threat on human health by food chain. Therefore, high accumulation of Cd in rice is a potential risk for both animals and human. As a crucial gaseous signaling molecule in plants, Nitric oxide (NO) plays a significant role in modulating several physiological and biochemical functions. In recent decades, an increasing number of articles have reported the effect of exogenous NO in alleviating heavy metal toxicity in different plants, although the physiological processes and mechanisms of NO in alleviating heavy metal toxicity are still far from clear. Therefore, it is necessary and meaningful to investigate the mechanisms of NO in alleviating Cd toxicity in rice, it will also provide significant theory basis to control the accumulation of Cd in rice seeds.
Japonica rice Xiushui63was used in the studies, a hydroponic experiment was conducted to investigate the effect of exogenous sodium nitroprusside (SNP), a NO donor, on alleviating the toxicity of Cd to rice. In order to elucidate the mechanisms of mitigate effect of NO application on Cd toxicity, growth parameters, reactive oxygen species, photosynthesis, mineral nutrition, ultrastruture and proteomic were researched. The main results were summarized as follows:
(1) Low concentrations of SNP (≤0.2mM) could alleviate the toxicity of Cd to rice, high concentrations of SNP deteriorated the toxicity of Cd to rice plants. A hydroponic experiment was conducted to investigate the effects of exogenous sodium nitroprusside (SNP), a NO donor, on growth in rice plants treated with0.1mM Cd and0.2mM Cd. The results concluded that the application of0.005mM SNP significantly alleviated0.1mM Cd-induced inhibition in rice seedlings growth, as plant height, root length, shoot weight and root weight. The application of0.1mM SNP significantly alleviated0.2mM Cd stress in rice growth. There is SNP concentration effect on alleviating Cd stress in rice plants.
(2) Exogenous NO significantly reduced the Cd-induced lipid peroxidation, hydrogen peroxide (H2O2) content and ascorbic acid (ASA) content in rice plants. The addition of NO reversed (only partially) the Cd-induced increase in activities of antioxidant enzymes-superoxide dismutase (SOD), peroxidase (POD), ascorbate peroxidase (APX) and glutathione reductase (GR). The addition of NO with Cd increased catalase (CAT) activity and glutathione (GSH) content. The exogenous application of NO could be advantageous against Cd toxicity, and could confer tolerance to heavy metal stress in rice plants.
(3) Exogenous NO significantly increased the chlorophyll content of rice leaves under Cd stress. NO significantly inhibited the decreases of net photosynthetic rate (Pn), transpiration rate (Tr), stomatal conductance (Gs), maximum fluorescence (Fm), PSII intrinsic photochemical efficiency (Fv/Fm), quantum yield of PSII (OPSII), and also inhibited the increase of intercellular CO2concentration (Ci). The results indicated that NO could modulate photosynthesis in plant, reduce energy dissipation and increase plant resistance by enhancing photosynthesis.
(4) Exogenous NO made chloroplasts become better or in relatively normal shape with parallel pattern of lamellae. Addition of NO with Cd, the number of vacuoles was increased, but with much smaller size, compared with Cd alone treatment. Moreover, more EDG were deposited in vacuoles.
(5) Exogenous NO decreased the Cd content in rice leaves, increased Cd content in root roots. NO significantly increased iron (Fe), zinc (Zn), manganese (Mn), copper (Cu) contents in rice shoot and potassium (K), magnesium (Mg), zinc (Zn), copper (Cu) contents in rice root; significantly decreased calcium (Ca) contents in rice roots induced by Cd stress. The results indicated that NO could maintain the mineral nutrition balance to mitigate Cd stress in rice plants.
(6) In rice leaves and roots,41proteins were identified under SNP treatment. Phosphoglycerate kinase, Glyceraldehyde-3-phosphate dehydrogenase B, ATP synthase, NADP dependent malic enzyme, and aminomethyl transferase in rice leaves were down-regulated expressed, while monodehydroascorbate reductase, phosphoglycerate mutase and ferredoxin-nitrite reductase were up-regulated expressed. In rice roots,5-methyltetrahydropteroyltriglutamate-homocysteine methyltransferase, heat shock protein, ATP synthase, and sucrose-UDP glucosyltransferase1were up-regulated expressed, Fructokinase II, isoflavone reductase and cysteine conjugate betalyase were down-regulated expressed. The results indicated that these proteins which participated in photosynthesis, carbohydrate metabolism, nitrogen metabolism, oxidative phosphorylation, oxidative stress responses, signal transductions and cell division.
引文
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