锰毒对酸柚光合作用及抗氧化系统的影响
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摘要
柑橘是亚热带常绿果树,种植于湿润、半湿润的热带、亚热带和温带地区,多为酸性或强酸性的红壤、黄壤或砖红壤,容易受到锰(Mn)毒的危害。目前,Mn毒对柑橘光合作用和抗氧化系统的研究甚少。
每隔一天用含2μM(对照)和500μM(Mn毒)MnSO4的营养液浇盆栽(基质为砂)酸柚[Citrus grandis (L.) Osbeck]实生苗,17周后调查Mn毒对酸柚实生苗生长、叶片CO2同化、核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)、碳水化合物和光合电子传递,以及根系和叶片抗氧化系统的影响,旨在明确Mn毒所导致的叶片CO2同化下降的机制,证实Mn毒引起的活性氧代谢的变化在根系和叶片中不同的假说。
锰毒显著增加根、茎和叶的Mn含量,但绝大部分Mn仍保留在根部。锰毒下酸柚实生幼苗根/冠比增大,因为Mn毒对地上部生长的影响比对根系生长的影响大。Mn毒减少酸柚叶片CO2同化和气孔导度,增加其胞间CO2浓度,但并不影响其叶绿素(Chl)含量;Mn毒叶片Rubisco初始活性和总活性下降的幅度均小于CO2同化下降的幅度。锰毒叶片果糖、葡萄糖、淀粉和总的非结构性碳水化合物含量与对照无显著差异,但其蔗糖含量则比对照高。Mn毒叶片OJIP曲线的O点上升,P点下降,并出现正的ΔL-和K-带。Mn毒降低暗适应叶片最大光量子产额(Fv/Fm)和总的性能指数(PItot, abs),增加其I点的相对可变荧光(VI)和热耗散能。当用单位蛋白质表示时,Mn毒叶片有较高的单脱氢抗坏血酸还原酶(MDAR)、谷胱甘肽还原酶(GR)、超氧物岐化酶(SOD)、过氧化氢酶(CAT)和愈创木酚过氧化物酶(GPX)活性,较高的抗氧化物质含量和较低的脱氢抗坏血酸还原酶(DHAR),而Mn毒根系有相似或较低的抗氧化酶活性和抗氧化物质含量。锰毒并不影响酸柚叶片和根系丙二醛(MDA)含量。
总之,Mn毒损伤了从光系统II(PSII)供体侧到光系统I(PSI)末端电子受体还原的整个光合电子传递链,从而限制还原力的产生,最终导致CO2同化下降。锰毒叶片抗氧化系统和热耗散均被上调,而在Mn毒处理的根系中其抗氧化系统并没有被上调,但仍保持高的抗氧化能力。Mn毒叶片和根系抗氧化系统均可为它们提供足够的保护使其免遭氧化伤害。
Citrus belongs to evergreen subtropical fruit trees and is cultivated in humid and sub-humid of tropical, sub-tropical, and temperate regions of the world mainly on acidic or highly acidic soils such as red soil, yellow soil and latosol and is liable to suffer from manganese (Mn)-toxicity.
Sour pummelo [Citrus grandis (L.) Osbeck] seedling grown in pots containing sands were fertigated for 17 weeks with nutrient solution containing 2μM (control) or 500μM (Mn-toxicity) MnSO4. Thereafter, the effects of Mn-toxicity on growth, CO2 assimilation, Ribulose-1,5- bisphosphate carboxylase/oxygenase (Rubisco), non-structural carbohydrates and photosynthetic electron transport in leaves, and antioxidant systems in roots and leaves of sour pummelo seedlings. The objective of this study were to understand the mechanisms by which Mn-toxicity leads to a decrease in CO2 assimilation and to test the hypothesis that Mn-toxicity-induced changes in antioxidant systems differr between roots and leaves.
Mn-toxicity decreased CO2 assimilation and stomatal conductance, increased intercellular CO2 concentration, but did not affect chlorophyll (Chl) content. Both initial and total Rubisco activity in Mn-toxicity leaves decreased to a lesser extent than CO2 assimilation. Contents of glucose, fructose, starch and total nonstructural carbohydrates did not differ between Mn-toxicity leaves and controls, while sucrose content was higher in the former. Chl a fluorescence (OJIP) transients from Mn-toxicity leaves showed increased O-step and decreased P-step, accompanied by postivieΔL- and K-bands. Mn-toxicity decreased maximum quantum yield of primary photochemistry (Fv/Fm) and total performance index (PItot, abs), but increased relative variable fluorescence at I-steps (VI) and energy dissipation. On a protein basis, Mn-toxicity leaves displayed higher activities of monodehydroascorbate reductase (MDAR), glutathione reductase (GR), superoxide dismutase (SOD), catalase (CAT) and guaiacol peroxidase (GPX) and contents of antioxidants, similar ascorbate peroxidase (APX) activities and lower dehydroascorbate reductase (DHAR) activities; while Mn-toxicity roots had similar or lower activities of antioxidant enzymes and contents of antioxidants. Mn-toxicity did not affect malondialdehyde (MDA) content of roots and leaves.
In conclusion, Mn-toxicity impaired the whole photosynthetic electron transport chain from the donor side of photosystem II (PSII) up to the reduction of end acceptors of PSI, thus limiting the production of reducing equivalents, and hence the rate of CO2 assimilation. Both the energy dissipation and the antioxidant systems were enhanced in Mn-toxicity leaves, while the antioxidant systems in Mn-toxicity roots were not enhanced, but still remained high activity. Antioxidant systems in Mn-toxicity roots and leaves provided sufficient protection to them against oxidative damage.
每隔一天用含2μM(对照)和500μM(Mn毒)MnSO4的营养液浇盆栽(基质为砂)酸柚[Citrus grandis (L.) Osbeck]实生苗,17周后调查Mn毒对酸柚实生苗生长、叶片CO2同化、核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)、碳水化合物和光合电子传递,以及根系和叶片抗氧化系统的影响,旨在明确Mn毒所导致的叶片CO2同化下降的机制,证实Mn毒引起的活性氧代谢的变化在根系和叶片中不同的假说。
锰毒显著增加根、茎和叶的Mn含量,但绝大部分Mn仍保留在根部。锰毒下酸柚实生幼苗根/冠比增大,因为Mn毒对地上部生长的影响比对根系生长的影响大。Mn毒减少酸柚叶片CO2同化和气孔导度,增加其胞间CO2浓度,但并不影响其叶绿素(Chl)含量;Mn毒叶片Rubisco初始活性和总活性下降的幅度均小于CO2同化下降的幅度。锰毒叶片果糖、葡萄糖、淀粉和总的非结构性碳水化合物含量与对照无显著差异,但其蔗糖含量则比对照高。Mn毒叶片OJIP曲线的O点上升,P点下降,并出现正的ΔL-和K-带。Mn毒降低暗适应叶片最大光量子产额(Fv/Fm)和总的性能指数(PItot, abs),增加其I点的相对可变荧光(VI)和热耗散能。当用单位蛋白质表示时,Mn毒叶片有较高的单脱氢抗坏血酸还原酶(MDAR)、谷胱甘肽还原酶(GR)、超氧物岐化酶(SOD)、过氧化氢酶(CAT)和愈创木酚过氧化物酶(GPX)活性,较高的抗氧化物质含量和较低的脱氢抗坏血酸还原酶(DHAR),而Mn毒根系有相似或较低的抗氧化酶活性和抗氧化物质含量。锰毒并不影响酸柚叶片和根系丙二醛(MDA)含量。
总之,Mn毒损伤了从光系统II(PSII)供体侧到光系统I(PSI)末端电子受体还原的整个光合电子传递链,从而限制还原力的产生,最终导致CO2同化下降。锰毒叶片抗氧化系统和热耗散均被上调,而在Mn毒处理的根系中其抗氧化系统并没有被上调,但仍保持高的抗氧化能力。Mn毒叶片和根系抗氧化系统均可为它们提供足够的保护使其免遭氧化伤害。
Citrus belongs to evergreen subtropical fruit trees and is cultivated in humid and sub-humid of tropical, sub-tropical, and temperate regions of the world mainly on acidic or highly acidic soils such as red soil, yellow soil and latosol and is liable to suffer from manganese (Mn)-toxicity.
Sour pummelo [Citrus grandis (L.) Osbeck] seedling grown in pots containing sands were fertigated for 17 weeks with nutrient solution containing 2μM (control) or 500μM (Mn-toxicity) MnSO4. Thereafter, the effects of Mn-toxicity on growth, CO2 assimilation, Ribulose-1,5- bisphosphate carboxylase/oxygenase (Rubisco), non-structural carbohydrates and photosynthetic electron transport in leaves, and antioxidant systems in roots and leaves of sour pummelo seedlings. The objective of this study were to understand the mechanisms by which Mn-toxicity leads to a decrease in CO2 assimilation and to test the hypothesis that Mn-toxicity-induced changes in antioxidant systems differr between roots and leaves.
Mn-toxicity decreased CO2 assimilation and stomatal conductance, increased intercellular CO2 concentration, but did not affect chlorophyll (Chl) content. Both initial and total Rubisco activity in Mn-toxicity leaves decreased to a lesser extent than CO2 assimilation. Contents of glucose, fructose, starch and total nonstructural carbohydrates did not differ between Mn-toxicity leaves and controls, while sucrose content was higher in the former. Chl a fluorescence (OJIP) transients from Mn-toxicity leaves showed increased O-step and decreased P-step, accompanied by postivieΔL- and K-bands. Mn-toxicity decreased maximum quantum yield of primary photochemistry (Fv/Fm) and total performance index (PItot, abs), but increased relative variable fluorescence at I-steps (VI) and energy dissipation. On a protein basis, Mn-toxicity leaves displayed higher activities of monodehydroascorbate reductase (MDAR), glutathione reductase (GR), superoxide dismutase (SOD), catalase (CAT) and guaiacol peroxidase (GPX) and contents of antioxidants, similar ascorbate peroxidase (APX) activities and lower dehydroascorbate reductase (DHAR) activities; while Mn-toxicity roots had similar or lower activities of antioxidant enzymes and contents of antioxidants. Mn-toxicity did not affect malondialdehyde (MDA) content of roots and leaves.
In conclusion, Mn-toxicity impaired the whole photosynthetic electron transport chain from the donor side of photosystem II (PSII) up to the reduction of end acceptors of PSI, thus limiting the production of reducing equivalents, and hence the rate of CO2 assimilation. Both the energy dissipation and the antioxidant systems were enhanced in Mn-toxicity leaves, while the antioxidant systems in Mn-toxicity roots were not enhanced, but still remained high activity. Antioxidant systems in Mn-toxicity roots and leaves provided sufficient protection to them against oxidative damage.
引文
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