铅（Pb~(2+)）胁迫下荇菜和菹草无菌苗的生理生化特性与容忍机制
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摘要
在常见污染物中,铅是对植物产生毒性和频繁遭受的毒害金属之一。目前,铅由于污染频率高,毒性强,以及具有导致人体疾病的潜质,已被有毒物质与疾病登记联合机构报告为第二大最有害物质,仅次于砷(ATSDR2003).因此,很有必要全面地理解铅对植物的毒害效应和植物对于铅胁迫的耐性机理。在本文中,以荇菜和菹草的无菌苗作为实验材料,旨在研究铅污染对各项生理生化参数的影响,从而进一步探索植物体在铅胁迫下所采取的适应策略。结果如下：
(1)以期探究铅的毒理效应,本文研究了不同浓度的铅离子(Pb)对荇菜无菌苗叶片内铅的积累、营养元素、丙二醛(MDA)含量、超氧阴离子(O2·-)的产生、过氧化氢(H202)含量、抗氧化酶活性、可溶性蛋白和光合色素以及叶绿体超微结构的影响。结果表明：叶片内在铅的积累以浓度依赖的方式增加。营养元素(Ca, K, Fe, Mn和Mo)的平衡也出现紊乱。丙二醛含量和超氧阴离子产生速率逐步增加,H202含量在低铅浓度12.5μM时大量增加后,却出现了下降的趋势。过氧化物酶(POD)和过氧化氢酶(CAT)的活性交替增加,而超氧化物歧化酶(SOD)活性逐渐下降。同时,研究表明铅的浓度与可溶性蛋白质和光合色素之间呈显著的负相关。此外,铅处理还导致了叶绿体结构出现明显的氧化损伤。总之,研究结果支持了荇菜在铅胁迫下产生氧化应激这一假设。此外,营养元素的紊乱和叶绿体超微结构的破坏也充分表明细胞功能出现了紊乱。
(2)为了研究潜在的抗氧化防御机制,不同浓度的铅对荇菜无菌苗多胺(PAs)、各种硫醇、维生素C和E以及脯氨酸含量的影响在5天毒害后进行了分析。总体腐胺(Put)、亚精胺(Spd)和精胺(Spm)的水平显著下降,而(Spd+Spm)/Put的比值随铅浓度的加大呈先增加后降低的趋势。游离态(Free)、高氯酸可溶共轭(PS-conjugated)和高氯酸不溶性结合(PIS-bound)多胺的变化趋势与总体多胺趋势一致。另外,在高浓度铅诱导下,还原型谷胱甘肽(GSH)、非蛋白巯基(NP-SH)、植物络合素(PCs)和维生素C的含量上升。维生素E则没有任何显著的变化,不受铅的影响。脯氨酸含量随着铅浓度的升高起初下降,之后出现了上升。下降的Put和上升的GSF、NP-SH、PCs,维生素C和脯氨酸含量与抗氧化效率密切关联,它们对于荇菜在铅胁迫下的适应机制中发挥着至关重要的作用。
(3)为了研究菹草无菌苗在铅胁迫下抗氧化系统的干预机制,不同浓度的铅对O2·-、H2O2、光合色素、MDA、可溶性蛋白、抗氧化酶、各种硫醇、维生素C和E以及总抗氧化能力(T-AOC)的影响进行了分析。铅离子对菹草产生了氧化应激和毒性作用,包括O2·-和H2O2过量生成、光合色素的减少、MDA和可溶性蛋白含量略有增加。随着溶液中铅浓度的加大,CAT活性逐渐增加,而POD活性逐渐下降。NP-SH和PCs则在低浓度铅下诱导增加,之后却在较高浓度下降低,而GSH含量略有升高。维生素C的合成随着铅的浓度增加显著地提高。然而,SOD活性和维生素E变化不明显。因此,T-AOC在适度的铅溶液(25和50μM)下经过短暂的上升之后迅速下降。研究结果表明,菹草无菌苗体内的抗氧化剂干预(CAT、GSH、NP-SH、PCs和维生素C)能赋予植物减弱铅诱导的氧化应激。
(4)为了研究潜在的耐受机制,本文研究了菹草的无菌苗在不同浓度铅离子下亚细胞水平的铅积累和矿质元素、多胺(PAs)和脯氨酸代谢的影响。铅的积累呈明显的浓度依赖性。亚细胞分布表明,大多数的Pb被存储在细胞壁,以细胞壁>可溶性部分>细胞器的降序排列。营养元素分析表明,总的P,Mg,Na和Zn的水平上升和总Fe水平下降,总Ca呈先增加后下降的趋势。此外,营养元素吸收的加强主要是由于细胞壁组分中的元素增加,而在可溶性组分和细胞器的元素则呈下降趋势。铅胁迫还导致总体的Put大幅度下降,而总体的Spd、Spm和(Spd+Spm)/Put的比值逐渐上升,但随后出现下降。铅对游离态、束缚态和结合态的多胺含量的变化与总体多胺的趋势保持一致,除了束缚态Spm显著增加。精氨酸脱羧酶(ADC)和鸟氨酸脱羧酶(ODC)活性略微升高,二胺氧化酶(DAO)活性则升高后快速下降；另外,由于铅离子对r-谷氨酰激酶(GK)活性的调控,脯氨酸含量先上升后下降。与此同时,鸟氨酸-d-氨基转移酶(OAT)的活性逐渐降低,而脯氨酸脱氢酶(PDH)活性变化不明显。因此,菹草对铅胁迫的耐受性机制基于细胞壁的区域化以及其组分矿质元素的促进、多胺含量的调整以及脯氨酸的诱导。
总而言之,本文从铅的积累、铅的毒害效应和铅的耐受机制三个角度进行研究,涉及的实验内容包括总体Pb的富集、Pb的亚细胞分布、Pb在叶绿体上的定位、活性氧、叶绿素分解和叶绿体结构的破坏、质膜的过氧化程度和可溶性蛋白的分解、矿质元素的紊乱、细胞壁的区域化作用、抗氧化酶、巯基、维生素、离子的竞争、脯氨酸和多胺含量的变化,一方面筛选了水体重金属铅污染条件下灵敏的监测性指标,另一方面探讨了植物本身内在抗氧化物质的抵抗机理,为全面理解植物对铅的响应、铅的耐性机制和水体环境修复的应用提供了有价值的理论基础。
Among common pollutants that affect plants, lead is toxic and frequently encountered. Currently, lead (Pb) was reported as being the second most hazardous substance, after arsenic, based on the frequency of occurrence, toxicity, and the potential for human exposure by the Agency for Toxic Substances and Disease Registry (ATSDR2003). Therefore, a comprehensive understanding of phytotoxic and plant resistance mechanisms exposed to various concentrations of lead (Pb) was required. In the present study, steriled seedlings of Nymphoides peltata and Potamogeton crispus were chosed as the experimental materials, aiming to study the effects of Pb contamination on physiological and biochemical parameters, and explore possible strategies adopted by plants under Pb stress. The results are as follow:
(1) The effects of increasing concentrations of lead on Pb accumulation and its influence on nutrient elements, malondialdehyde (MDA) content, generation of superoxide anion (O2·-), hydrogen peroxide (H2O2) content, antioxidant enzymes activities, soluble protein and photosynthetic pigment, as well as chloroplast ultrastructure in steriled seedlings of Nymphoides peltata (S. G. Gmel.) Kuntze were investigated in order to understand Pb-induced toxicity. The accumulation of Pb was found to increase in a concentration-dependent manner. Nutrient elements (Ca, K, Fe, Mn and Mo) were also affected. MDA content and O2generation rate increased progressively, while H2O2content first boosted up at a low Pb concentration of12.5μM but then declined. Guaiacol peroxides (POD) and catalase (CAT) activities increased alternately, while superoxide dismutase (SOD) activity gradually fell. Negative correlations were found between Pb and soluble protein and photosynthetic pigment. Moreover, Pb exposure resulted in significant damage of chloroplasts. Taken together, these findings supported the hypothesis that N. peltatum underwent oxidative stress induced by Pb. In addition, both the disorder of nutrient elements and the damage to the ultrastructure of chloroplasts were indicative of general disarray in the cellular functions exerted by Pb.
(2) In order to study potential antioxidant defense mechanisms, the effects of increasing concentrations of lead (Pb) on polyamines (PAs), various thiols, vitamins C and E and proline contents in sterilized seedlings of Nymphoides peltata (S.G. mel.) Kuntze were investigated after5days exposure. The levels of total putrescine (Put), spermidine (Spd), and spermine (Spm) decreased significantly, while the ratio of (Spd+Spm)/Put first increased but then declined as the concentration of Pb increased. The trends for free, perchloric acid soluble conjugated (PS-conjugated) and perchloric acid insoluble bound (PIS-bound) PAs were similar to the trend seen for total PAs. Moreover, reduced glutathione (GSH), non-protein thiols (NP-SH), phytochelatins (PCs) and vitamin C were induced at high Pb concentrations. No significant change was observed in vitamin E. An initial decline in proline content was followed by an increase as the Pb concentration rose. The reduced level of Put and elevated contents of GSH, NP-SH, PCs, vitamin C and proline were found to be associated with antioxidant efficiency, which supports the hypothesis that they could play a significant role in the adaptation mechanisms of N. peltatum under Pb stress.
(3) To study antioxidant intervention compensates oxidative stress in sterilized seedlings of Potamogeton crispus L. exposed to lead (Pb) stress, the effects of increasing Pb concentration on generation of superoxide anion (O2·-), hydrogen peroxide (H2O2), photosynthetic pigment, malondialdehyde (MDA) content, soluble protein, antioxidant enzymes activities, various thiols, vitamins C and E as well as total antioxidant capacity (T-AOC) were investigated. The oxidative stress and toxic effect caused by Pb were evident by the production of excess O2·-and H2O2, the reduction in photosynthetic pigments and slightly increase in malondialddehyde (MDA) and soluble protein contents. Catalase (CAT) activity increased progressively while Guaiacol peroxidase (POD) activity gradually fell. Non-protein thiols (NP-SH) and phytochelatins (PCs) were induced at lower concentration followed by decline, while a mild increase was observed in reduced glutathione (GSH) content. Vitamin C was synthesized to significant levels as the concentration of Pb increased. However, no significant changes were observed in superoxide dismutase (SOD) activity and vitamin E. Thus, T-AOC rapidly declined after a transitory increase at moderate exposure conditions (25and50uM). Our results demonstrated that antioxidant intervention (CAT, GSH, NP-SH, PCs and vitamin C) was able to confer a protective effect against the Pb-induced oxidative stress in sterilized seedlings of P. crispus.
(4) In order to study potential tolerance mechanisms, the effects of increasing concentrations of lead (Pb) on Pb accumulation and nutrients at the subcellular level, polyamines (PAs) and proline metabolism in sterilized seedlings of Potamogeton crispus L. were investigated after5days exposure. The accumulation of Pb was found to increase in a concentration-dependent manner. Subcellular distribution demonstrated that most of Pb was stored in the cell wall, followed the descending order of cell wall> soluble fraction>organelle fraction. Analysis of nutrient elements showed that the levels of total P, Mg, Na and Zn rose and the level of total Fe fell; total Ca increased initially only to decline later. Moreover, the increased nutrients in cell wall fraction were responsible for enhancement of nutrients absorption whereas those in soluble and organelle fraction declined. Pb stress also induced a considerable decrease in total putrescine (Put), while the levels of total spermidine (Spd), spermine (Spm) and (Spd+Spm)/Put ratio increased progressively but then declined. The trends for free, perchloric acid soluble conjugated (PS-conjugated) and perchloric acid insoluble bound (PIS-bound) PAs were similar to the trend seen for total PAs, except that PIS-bound Spm increased significantly. The activities of arginine decarboxylase (ADC) and ornithine decarboxylase (ODC) rose gradually with an increase in Pb concentration, while diamine oxidase (DAO) initially increased but then declined. In addition, an initial increase in proline content was followed by a decline as the Pb concentration rose, due to the increase of r-glutamyl kinase (GK) activity. Meanwhile, ornithine-d-aminotransferase (OAT) activity gradually reduced, while no significant change was observed in proline dehydrogenase (PDH) activity. Therefore, our results indicated that the tolerance of P. crispus to Pb stress was based on cell wall compartmentalization combined with promotion of nutrients, alterations of PAs and induction of proline.
In summary, this paper studied from three angles (bioaccumulation, toxic effects and tolerance mechanisms of lead), and the experiments involved multi-changes in the overall Pb content, subcellular distribution of Pb, Pb positioned in chloroplast, levels of reactive oxygen species (ROS), decomposition of chlorophyll and destruction of chloroplast structure, peroxidation of membrane and soluble proteins, disorders of mineral elements, regionalization of cell wall, antioxidant enzyme, thiol, vitamins, ions competition, proline and PAs contents. On one hand, sensitive monitoring indicators could serve as useful and indicative biomarkers in toxicological tests with Pb in aquatic environment. On the other hand, our results explored the intrinsic resistance mechanism of antioxidants to Pb, providing valuable theoretical basis for a comprehensive understanding of plant response, tolerance systems and restoration of Pb-contaminated water environment.
(1)以期探究铅的毒理效应,本文研究了不同浓度的铅离子(Pb)对荇菜无菌苗叶片内铅的积累、营养元素、丙二醛(MDA)含量、超氧阴离子(O2·-)的产生、过氧化氢(H202)含量、抗氧化酶活性、可溶性蛋白和光合色素以及叶绿体超微结构的影响。结果表明：叶片内在铅的积累以浓度依赖的方式增加。营养元素(Ca, K, Fe, Mn和Mo)的平衡也出现紊乱。丙二醛含量和超氧阴离子产生速率逐步增加,H202含量在低铅浓度12.5μM时大量增加后,却出现了下降的趋势。过氧化物酶(POD)和过氧化氢酶(CAT)的活性交替增加,而超氧化物歧化酶(SOD)活性逐渐下降。同时,研究表明铅的浓度与可溶性蛋白质和光合色素之间呈显著的负相关。此外,铅处理还导致了叶绿体结构出现明显的氧化损伤。总之,研究结果支持了荇菜在铅胁迫下产生氧化应激这一假设。此外,营养元素的紊乱和叶绿体超微结构的破坏也充分表明细胞功能出现了紊乱。
(2)为了研究潜在的抗氧化防御机制,不同浓度的铅对荇菜无菌苗多胺(PAs)、各种硫醇、维生素C和E以及脯氨酸含量的影响在5天毒害后进行了分析。总体腐胺(Put)、亚精胺(Spd)和精胺(Spm)的水平显著下降,而(Spd+Spm)/Put的比值随铅浓度的加大呈先增加后降低的趋势。游离态(Free)、高氯酸可溶共轭(PS-conjugated)和高氯酸不溶性结合(PIS-bound)多胺的变化趋势与总体多胺趋势一致。另外,在高浓度铅诱导下,还原型谷胱甘肽(GSH)、非蛋白巯基(NP-SH)、植物络合素(PCs)和维生素C的含量上升。维生素E则没有任何显著的变化,不受铅的影响。脯氨酸含量随着铅浓度的升高起初下降,之后出现了上升。下降的Put和上升的GSF、NP-SH、PCs,维生素C和脯氨酸含量与抗氧化效率密切关联,它们对于荇菜在铅胁迫下的适应机制中发挥着至关重要的作用。
(3)为了研究菹草无菌苗在铅胁迫下抗氧化系统的干预机制,不同浓度的铅对O2·-、H2O2、光合色素、MDA、可溶性蛋白、抗氧化酶、各种硫醇、维生素C和E以及总抗氧化能力(T-AOC)的影响进行了分析。铅离子对菹草产生了氧化应激和毒性作用,包括O2·-和H2O2过量生成、光合色素的减少、MDA和可溶性蛋白含量略有增加。随着溶液中铅浓度的加大,CAT活性逐渐增加,而POD活性逐渐下降。NP-SH和PCs则在低浓度铅下诱导增加,之后却在较高浓度下降低,而GSH含量略有升高。维生素C的合成随着铅的浓度增加显著地提高。然而,SOD活性和维生素E变化不明显。因此,T-AOC在适度的铅溶液(25和50μM)下经过短暂的上升之后迅速下降。研究结果表明,菹草无菌苗体内的抗氧化剂干预(CAT、GSH、NP-SH、PCs和维生素C)能赋予植物减弱铅诱导的氧化应激。
(4)为了研究潜在的耐受机制,本文研究了菹草的无菌苗在不同浓度铅离子下亚细胞水平的铅积累和矿质元素、多胺(PAs)和脯氨酸代谢的影响。铅的积累呈明显的浓度依赖性。亚细胞分布表明,大多数的Pb被存储在细胞壁,以细胞壁>可溶性部分>细胞器的降序排列。营养元素分析表明,总的P,Mg,Na和Zn的水平上升和总Fe水平下降,总Ca呈先增加后下降的趋势。此外,营养元素吸收的加强主要是由于细胞壁组分中的元素增加,而在可溶性组分和细胞器的元素则呈下降趋势。铅胁迫还导致总体的Put大幅度下降,而总体的Spd、Spm和(Spd+Spm)/Put的比值逐渐上升,但随后出现下降。铅对游离态、束缚态和结合态的多胺含量的变化与总体多胺的趋势保持一致,除了束缚态Spm显著增加。精氨酸脱羧酶(ADC)和鸟氨酸脱羧酶(ODC)活性略微升高,二胺氧化酶(DAO)活性则升高后快速下降；另外,由于铅离子对r-谷氨酰激酶(GK)活性的调控,脯氨酸含量先上升后下降。与此同时,鸟氨酸-d-氨基转移酶(OAT)的活性逐渐降低,而脯氨酸脱氢酶(PDH)活性变化不明显。因此,菹草对铅胁迫的耐受性机制基于细胞壁的区域化以及其组分矿质元素的促进、多胺含量的调整以及脯氨酸的诱导。
总而言之,本文从铅的积累、铅的毒害效应和铅的耐受机制三个角度进行研究,涉及的实验内容包括总体Pb的富集、Pb的亚细胞分布、Pb在叶绿体上的定位、活性氧、叶绿素分解和叶绿体结构的破坏、质膜的过氧化程度和可溶性蛋白的分解、矿质元素的紊乱、细胞壁的区域化作用、抗氧化酶、巯基、维生素、离子的竞争、脯氨酸和多胺含量的变化,一方面筛选了水体重金属铅污染条件下灵敏的监测性指标,另一方面探讨了植物本身内在抗氧化物质的抵抗机理,为全面理解植物对铅的响应、铅的耐性机制和水体环境修复的应用提供了有价值的理论基础。
Among common pollutants that affect plants, lead is toxic and frequently encountered. Currently, lead (Pb) was reported as being the second most hazardous substance, after arsenic, based on the frequency of occurrence, toxicity, and the potential for human exposure by the Agency for Toxic Substances and Disease Registry (ATSDR2003). Therefore, a comprehensive understanding of phytotoxic and plant resistance mechanisms exposed to various concentrations of lead (Pb) was required. In the present study, steriled seedlings of Nymphoides peltata and Potamogeton crispus were chosed as the experimental materials, aiming to study the effects of Pb contamination on physiological and biochemical parameters, and explore possible strategies adopted by plants under Pb stress. The results are as follow:
(1) The effects of increasing concentrations of lead on Pb accumulation and its influence on nutrient elements, malondialdehyde (MDA) content, generation of superoxide anion (O2·-), hydrogen peroxide (H2O2) content, antioxidant enzymes activities, soluble protein and photosynthetic pigment, as well as chloroplast ultrastructure in steriled seedlings of Nymphoides peltata (S. G. Gmel.) Kuntze were investigated in order to understand Pb-induced toxicity. The accumulation of Pb was found to increase in a concentration-dependent manner. Nutrient elements (Ca, K, Fe, Mn and Mo) were also affected. MDA content and O2generation rate increased progressively, while H2O2content first boosted up at a low Pb concentration of12.5μM but then declined. Guaiacol peroxides (POD) and catalase (CAT) activities increased alternately, while superoxide dismutase (SOD) activity gradually fell. Negative correlations were found between Pb and soluble protein and photosynthetic pigment. Moreover, Pb exposure resulted in significant damage of chloroplasts. Taken together, these findings supported the hypothesis that N. peltatum underwent oxidative stress induced by Pb. In addition, both the disorder of nutrient elements and the damage to the ultrastructure of chloroplasts were indicative of general disarray in the cellular functions exerted by Pb.
(2) In order to study potential antioxidant defense mechanisms, the effects of increasing concentrations of lead (Pb) on polyamines (PAs), various thiols, vitamins C and E and proline contents in sterilized seedlings of Nymphoides peltata (S.G. mel.) Kuntze were investigated after5days exposure. The levels of total putrescine (Put), spermidine (Spd), and spermine (Spm) decreased significantly, while the ratio of (Spd+Spm)/Put first increased but then declined as the concentration of Pb increased. The trends for free, perchloric acid soluble conjugated (PS-conjugated) and perchloric acid insoluble bound (PIS-bound) PAs were similar to the trend seen for total PAs. Moreover, reduced glutathione (GSH), non-protein thiols (NP-SH), phytochelatins (PCs) and vitamin C were induced at high Pb concentrations. No significant change was observed in vitamin E. An initial decline in proline content was followed by an increase as the Pb concentration rose. The reduced level of Put and elevated contents of GSH, NP-SH, PCs, vitamin C and proline were found to be associated with antioxidant efficiency, which supports the hypothesis that they could play a significant role in the adaptation mechanisms of N. peltatum under Pb stress.
(3) To study antioxidant intervention compensates oxidative stress in sterilized seedlings of Potamogeton crispus L. exposed to lead (Pb) stress, the effects of increasing Pb concentration on generation of superoxide anion (O2·-), hydrogen peroxide (H2O2), photosynthetic pigment, malondialdehyde (MDA) content, soluble protein, antioxidant enzymes activities, various thiols, vitamins C and E as well as total antioxidant capacity (T-AOC) were investigated. The oxidative stress and toxic effect caused by Pb were evident by the production of excess O2·-and H2O2, the reduction in photosynthetic pigments and slightly increase in malondialddehyde (MDA) and soluble protein contents. Catalase (CAT) activity increased progressively while Guaiacol peroxidase (POD) activity gradually fell. Non-protein thiols (NP-SH) and phytochelatins (PCs) were induced at lower concentration followed by decline, while a mild increase was observed in reduced glutathione (GSH) content. Vitamin C was synthesized to significant levels as the concentration of Pb increased. However, no significant changes were observed in superoxide dismutase (SOD) activity and vitamin E. Thus, T-AOC rapidly declined after a transitory increase at moderate exposure conditions (25and50uM). Our results demonstrated that antioxidant intervention (CAT, GSH, NP-SH, PCs and vitamin C) was able to confer a protective effect against the Pb-induced oxidative stress in sterilized seedlings of P. crispus.
(4) In order to study potential tolerance mechanisms, the effects of increasing concentrations of lead (Pb) on Pb accumulation and nutrients at the subcellular level, polyamines (PAs) and proline metabolism in sterilized seedlings of Potamogeton crispus L. were investigated after5days exposure. The accumulation of Pb was found to increase in a concentration-dependent manner. Subcellular distribution demonstrated that most of Pb was stored in the cell wall, followed the descending order of cell wall> soluble fraction>organelle fraction. Analysis of nutrient elements showed that the levels of total P, Mg, Na and Zn rose and the level of total Fe fell; total Ca increased initially only to decline later. Moreover, the increased nutrients in cell wall fraction were responsible for enhancement of nutrients absorption whereas those in soluble and organelle fraction declined. Pb stress also induced a considerable decrease in total putrescine (Put), while the levels of total spermidine (Spd), spermine (Spm) and (Spd+Spm)/Put ratio increased progressively but then declined. The trends for free, perchloric acid soluble conjugated (PS-conjugated) and perchloric acid insoluble bound (PIS-bound) PAs were similar to the trend seen for total PAs, except that PIS-bound Spm increased significantly. The activities of arginine decarboxylase (ADC) and ornithine decarboxylase (ODC) rose gradually with an increase in Pb concentration, while diamine oxidase (DAO) initially increased but then declined. In addition, an initial increase in proline content was followed by a decline as the Pb concentration rose, due to the increase of r-glutamyl kinase (GK) activity. Meanwhile, ornithine-d-aminotransferase (OAT) activity gradually reduced, while no significant change was observed in proline dehydrogenase (PDH) activity. Therefore, our results indicated that the tolerance of P. crispus to Pb stress was based on cell wall compartmentalization combined with promotion of nutrients, alterations of PAs and induction of proline.
In summary, this paper studied from three angles (bioaccumulation, toxic effects and tolerance mechanisms of lead), and the experiments involved multi-changes in the overall Pb content, subcellular distribution of Pb, Pb positioned in chloroplast, levels of reactive oxygen species (ROS), decomposition of chlorophyll and destruction of chloroplast structure, peroxidation of membrane and soluble proteins, disorders of mineral elements, regionalization of cell wall, antioxidant enzyme, thiol, vitamins, ions competition, proline and PAs contents. On one hand, sensitive monitoring indicators could serve as useful and indicative biomarkers in toxicological tests with Pb in aquatic environment. On the other hand, our results explored the intrinsic resistance mechanism of antioxidants to Pb, providing valuable theoretical basis for a comprehensive understanding of plant response, tolerance systems and restoration of Pb-contaminated water environment.
引文
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