汞诱导蕨类植物和印度芥菜氧化胁迫反应及其污染土壤的植物修复作用研究
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摘要
汞污染对农业生态环境和人类健康的危害日趋严重,应用植物修复技术对包括汞在内的重金属污染进行治理是目前的研究热点。本文通过研究特定植物对水体、土壤和空气中汞的吸收与积累,以及相应条件下植物的生理生化变化,对汞污染的植物修复技术进行了初探。
凤尾蕨和波士顿蕨分别是已知的砷超积累品种和砷敏感品种,鉴于超积累植物在抵抗重金属胁迫方面的某些共同特征,我们用Hg2+溶液处理两种蕨类植物的根部,研究植物对溶液中Hg2+的吸收和积累,以及由此诱导的地上部生理毒害和氧化胁迫反应。两种蕨类植物根系都能积累大量汞,但向地上部的转运效率较低。凤尾蕨地上部和根系对汞的吸收和积累量显著大于波士顿蕨。汞胁迫造成凤尾蕨地上部产生可视的萎焉、缺绿等胁迫症状,电子显微镜观察也显示出叶片维管束细胞的收缩和叶绿体的降解；而波士顿蕨则没有显示出上述明显的毒害症状。进一步的研究表明汞胁迫诱导凤尾蕨地上部H2O2和TBARS含量显著增加,而抗氧化酶SOD、CAT和GR的活性没有显著变化。波士顿蕨则不同,汞胁迫诱导了其地上部SOD、CAT和GR的活性显著增加,从而维持了H2O2和TBARS的水平保持不变。ASA-DHA的分析表明,相较于凤尾蕨,汞胁迫通过诱导波士顿蕨地上部ASA/(ASA+DHA)比值的显著上升,调节细胞内的氧化还原状态,更加有效的抵御环境胁迫。与波士顿蕨相比,凤尾蕨在汞胁迫下表现出较重的植物毒害和氧化胁迫反应,这可能与凤尾蕨体内积累更多汞有关。
印度芥菜是用于研究重金属植物修复的良好材料,其生长量大,能够积累多种重金属元素,但目前还没有发现其在汞方面的研究。我们用Hg2+溶液处理印度芥菜两个品种(阔叶青和斯坦丁芥)的根部。发现两个品种根部都能积累大量汞,并且能将部分汞转移到地上部,转移效率高于蕨类。在高浓度汞处理下,阔叶青地上部和根系汞的积累量高于斯坦丁芥。两个印度芥菜品种地上部在高浓度汞胁迫下都表现出萎焉、缺绿、生长受抑制等胁迫症状。阔叶青和斯坦丁芥在汞胁迫下地上部的氧化胁迫反应无显著差异。汞胁迫诱导两者地上部TBARS的增加和H2O2的减少。抗氧化酶CAT和POD的活性显著上升。线性分析表明,CAT和POD活性与H2O2含量呈显著负相关。说明汞胁迫下,印度芥菜可能通过调节CAT和POD活性来避免过量H2O2造成的毒害。
结合以上研究结果,我们选用三种植物材料(印度芥菜、凤尾蕨、旱叶草)进行了汞污染土壤和空气的植物修复研究。通过栽培基质加HgCl2溶液处理、新鲜汞污染土壤处理和多年汞污染土壤处理三个循序渐进的实验。凤尾蕨在没有出现明显胁迫症状的情况下,其根系对不同类型汞污染土壤(HgCl2、Hg(NO3)2、HgS)中的汞都具有较强的吸收能力(根中最高汞积累达1774 mg/kg),而且能够将部分汞转移到地上部(地上部最高汞积累量达290 mg/kg),说明凤尾蕨在汞污染土壤的植物提取和植物固定方面有一定潜力。针对汞具有挥发性,能够污染大气的特点。我们还设计了气室实验,发现凤尾蕨和印度芥菜地上部对空气中的汞有一定的吸收能力,而且向根系的转运量很少,在治理汞污染空气方面具有一定潜力。
将遥感技术应用于监测植物的生理状态是目前研究环境污染检测的热点。我们在应用凤尾蕨进行土壤汞污染的植物修复过程中,发现汞胁迫引起了植物内部结构和生理状态的改变,而植物在可见光和近红外区域的反射光谱对于这些改变反应非常敏感。本研究通过分析凤尾蕨地上部在汞胁迫下的反射光谱变化,确立了部分反射光谱指标与汞胁迫下植物生理状态之间的相关性。NDVⅠ、R1110/R810、WⅠ的变化分别能够反映汞胁迫引起的凤尾蕨地上部生物量下降、叶片内部细胞结构的改变、相对含水量下降。R550的增加则反应出汞胁迫导致的叶绿素含量下降。这些结果为遥感技术应用于汞污染植物修复的长期监测提供了理论和应用基础。
Mercury (Hg) contamination has evocked great concerns worldwide because of its harmful effect on agricultural ecosystem and environment. Phytoremediation is a nearly perfect strategy for controlling soil heavy metal contamination. However, no Hg hyperaccumulator has been found thus far. Despite of this, great efforts have been made to explore physicological response of selected plant speices to Hg stress as well as their ability to accumulate Hg from Hg-contaminated soils, air and waters.
Pteris vittata, a fern, is a well-known arsinic-hyperaccumulating plant, while its counterpart Nephrolepis exaltata is arsinic-sensitive. We made comparative investigation on the Hg accumulation and its phytotoxicity in the test plants. The roots of both ferns accumulated a large amount of Hg, but exhibited limited Hg translocation to shoots. Pteris vittata accumulated more Hg in shoots and roots than Nephrolepis exaltata. Exposure to high level of Hg induced visible toxic symptoms in the shoots of Pteris vittata, e.g. withering and chlorosis. Electronic microscopy analysis revealed the shrinkage of vascular cells and breakdown of chloroplasts in Pteris vittata leaves under Hg exposure. In contrast, no visible toxicity and foliar structural changes were observed in Nephrolepis exaltata under Hg stress. Hg exposure induced substaintial accumulation of H2O2 and TBARS in the shoots of Pteris vittata. But the activities of anti-oxidative enzymes such as SOD, CAT, and GR remained relatively unchanged. For Nephrolepis exaltata, Hg exposure significantly incresed the activities of these anti-oxidative enzymes, which play critical roles in removing excess H2O2 and consequently suppressed TBARS production under Hg stress. Additionally, Nephrolepis exaltata had a higher level of ASA/(ASA+DHA) ratio than Pteris vittata. Overall, the results indicated that Nephrolepis exaltata appeared to be more tolerant to Hg than Pteris vittata with regard to their response to Hg-induced oxidative stress.
Indian mustard is believed to be a potential candidate plant for phtoremediation of heavy metal-contaminated soils. Two major commercial cultivars of Indian mustard (Brassica juncea), Broad leaf and Long-stading, were used in the study. Mature plants accumulated more Hg than seedlings. The roots of the two cultivars accumulated high level of Hg from root-bathing solutions. Shoots accumulated higher amounts of Hg than other plants as previously reported, indicating the significant translocation of Hg from roots to shoots. Content of Hg in Broad leaf was higher than that of Long-standing at the external concentrations of Hg within 12.2-16.7 mg/L. However, at such levels, both cultivars showed chlorosis and stunted growth of roots. In gerenal, Hg exposure significantly increased the activities of CAT, POD, and SOD, but decreased the concentratios of H2O2 in the shoots of Indian mustard. Liner analysis showed negative correlations between H2O2 concentrations and the activities of CAT and POD, which implyed that CAT and POD probably played important roles in protecting the plants from H2O2 toxicity under Hg stress.
We screened and investigated the potential of three plants (Pteris vittata, Brassica juncea, and Polypogon monspeliensis) in phytoextraction of Hg-contaminated soil. Three independent experiments were performed in turn. (1) Pot study with potting mix and HgCl2 solution. (2) Pot study with freshly HgCl2-spiked soil. (3) Experiment using aged soil contaminated with different Hg sources. Both shoots and roots of Pteris vittata could accumulate large amount of Hg from many kinds of Hg-contaminated soils (such as HgCl2, Hg(NO3)2, or HgS) without any visible stress symptoms. Therefore, Pteris vittata was a possible candidate for Hg phytoextraction and phytostabilization. Elemental Hg can evaporate into air because of its special physical properties. So we designed a chamber experiment to study the foliar Hg uptake by plants. Shoots of both Pteris vittata and Brassica juncea accumulated Hg at the concentratios of 63-105 mg/kg from Hg-contaminated air. So both Pteris vittata and Brassica juncea probably have potential in remediating Hg-contaminated air.
The application of remote sensing in monitoring plant physiological status has been growing concerns. We found that relative spectral reflectances in visible region (390-780 nm) and near infrared region (NIR,800-1300 nm) were sensitive to leaf structure and physiological status in Pteris vittata during the phytoremediation of Hg-contaminated soil. We selected several high spectral resolution reflectance vegetation indices to evaluate the change of spectral reflectance in Pteris vittata. We also estabilished the correlations between NDVI and biomass, R1110/R810 and leaf structure, WI and relative water content, respectively, in the shoots of Pteris vittata grown in Hg-contaminated soil. Hg stress induced significant increase in relative reflectance around 680 nm corresponding to the breakdown of chloroplast observed in TEM photographs in Pteris vittata. These results provided some theoretical evidences for the application of remote sensing in monitoring plant physiological status during the phytoremediation of Hg-contaminated soil.
凤尾蕨和波士顿蕨分别是已知的砷超积累品种和砷敏感品种,鉴于超积累植物在抵抗重金属胁迫方面的某些共同特征,我们用Hg2+溶液处理两种蕨类植物的根部,研究植物对溶液中Hg2+的吸收和积累,以及由此诱导的地上部生理毒害和氧化胁迫反应。两种蕨类植物根系都能积累大量汞,但向地上部的转运效率较低。凤尾蕨地上部和根系对汞的吸收和积累量显著大于波士顿蕨。汞胁迫造成凤尾蕨地上部产生可视的萎焉、缺绿等胁迫症状,电子显微镜观察也显示出叶片维管束细胞的收缩和叶绿体的降解；而波士顿蕨则没有显示出上述明显的毒害症状。进一步的研究表明汞胁迫诱导凤尾蕨地上部H2O2和TBARS含量显著增加,而抗氧化酶SOD、CAT和GR的活性没有显著变化。波士顿蕨则不同,汞胁迫诱导了其地上部SOD、CAT和GR的活性显著增加,从而维持了H2O2和TBARS的水平保持不变。ASA-DHA的分析表明,相较于凤尾蕨,汞胁迫通过诱导波士顿蕨地上部ASA/(ASA+DHA)比值的显著上升,调节细胞内的氧化还原状态,更加有效的抵御环境胁迫。与波士顿蕨相比,凤尾蕨在汞胁迫下表现出较重的植物毒害和氧化胁迫反应,这可能与凤尾蕨体内积累更多汞有关。
印度芥菜是用于研究重金属植物修复的良好材料,其生长量大,能够积累多种重金属元素,但目前还没有发现其在汞方面的研究。我们用Hg2+溶液处理印度芥菜两个品种(阔叶青和斯坦丁芥)的根部。发现两个品种根部都能积累大量汞,并且能将部分汞转移到地上部,转移效率高于蕨类。在高浓度汞处理下,阔叶青地上部和根系汞的积累量高于斯坦丁芥。两个印度芥菜品种地上部在高浓度汞胁迫下都表现出萎焉、缺绿、生长受抑制等胁迫症状。阔叶青和斯坦丁芥在汞胁迫下地上部的氧化胁迫反应无显著差异。汞胁迫诱导两者地上部TBARS的增加和H2O2的减少。抗氧化酶CAT和POD的活性显著上升。线性分析表明,CAT和POD活性与H2O2含量呈显著负相关。说明汞胁迫下,印度芥菜可能通过调节CAT和POD活性来避免过量H2O2造成的毒害。
结合以上研究结果,我们选用三种植物材料(印度芥菜、凤尾蕨、旱叶草)进行了汞污染土壤和空气的植物修复研究。通过栽培基质加HgCl2溶液处理、新鲜汞污染土壤处理和多年汞污染土壤处理三个循序渐进的实验。凤尾蕨在没有出现明显胁迫症状的情况下,其根系对不同类型汞污染土壤(HgCl2、Hg(NO3)2、HgS)中的汞都具有较强的吸收能力(根中最高汞积累达1774 mg/kg),而且能够将部分汞转移到地上部(地上部最高汞积累量达290 mg/kg),说明凤尾蕨在汞污染土壤的植物提取和植物固定方面有一定潜力。针对汞具有挥发性,能够污染大气的特点。我们还设计了气室实验,发现凤尾蕨和印度芥菜地上部对空气中的汞有一定的吸收能力,而且向根系的转运量很少,在治理汞污染空气方面具有一定潜力。
将遥感技术应用于监测植物的生理状态是目前研究环境污染检测的热点。我们在应用凤尾蕨进行土壤汞污染的植物修复过程中,发现汞胁迫引起了植物内部结构和生理状态的改变,而植物在可见光和近红外区域的反射光谱对于这些改变反应非常敏感。本研究通过分析凤尾蕨地上部在汞胁迫下的反射光谱变化,确立了部分反射光谱指标与汞胁迫下植物生理状态之间的相关性。NDVⅠ、R1110/R810、WⅠ的变化分别能够反映汞胁迫引起的凤尾蕨地上部生物量下降、叶片内部细胞结构的改变、相对含水量下降。R550的增加则反应出汞胁迫导致的叶绿素含量下降。这些结果为遥感技术应用于汞污染植物修复的长期监测提供了理论和应用基础。
Mercury (Hg) contamination has evocked great concerns worldwide because of its harmful effect on agricultural ecosystem and environment. Phytoremediation is a nearly perfect strategy for controlling soil heavy metal contamination. However, no Hg hyperaccumulator has been found thus far. Despite of this, great efforts have been made to explore physicological response of selected plant speices to Hg stress as well as their ability to accumulate Hg from Hg-contaminated soils, air and waters.
Pteris vittata, a fern, is a well-known arsinic-hyperaccumulating plant, while its counterpart Nephrolepis exaltata is arsinic-sensitive. We made comparative investigation on the Hg accumulation and its phytotoxicity in the test plants. The roots of both ferns accumulated a large amount of Hg, but exhibited limited Hg translocation to shoots. Pteris vittata accumulated more Hg in shoots and roots than Nephrolepis exaltata. Exposure to high level of Hg induced visible toxic symptoms in the shoots of Pteris vittata, e.g. withering and chlorosis. Electronic microscopy analysis revealed the shrinkage of vascular cells and breakdown of chloroplasts in Pteris vittata leaves under Hg exposure. In contrast, no visible toxicity and foliar structural changes were observed in Nephrolepis exaltata under Hg stress. Hg exposure induced substaintial accumulation of H2O2 and TBARS in the shoots of Pteris vittata. But the activities of anti-oxidative enzymes such as SOD, CAT, and GR remained relatively unchanged. For Nephrolepis exaltata, Hg exposure significantly incresed the activities of these anti-oxidative enzymes, which play critical roles in removing excess H2O2 and consequently suppressed TBARS production under Hg stress. Additionally, Nephrolepis exaltata had a higher level of ASA/(ASA+DHA) ratio than Pteris vittata. Overall, the results indicated that Nephrolepis exaltata appeared to be more tolerant to Hg than Pteris vittata with regard to their response to Hg-induced oxidative stress.
Indian mustard is believed to be a potential candidate plant for phtoremediation of heavy metal-contaminated soils. Two major commercial cultivars of Indian mustard (Brassica juncea), Broad leaf and Long-stading, were used in the study. Mature plants accumulated more Hg than seedlings. The roots of the two cultivars accumulated high level of Hg from root-bathing solutions. Shoots accumulated higher amounts of Hg than other plants as previously reported, indicating the significant translocation of Hg from roots to shoots. Content of Hg in Broad leaf was higher than that of Long-standing at the external concentrations of Hg within 12.2-16.7 mg/L. However, at such levels, both cultivars showed chlorosis and stunted growth of roots. In gerenal, Hg exposure significantly increased the activities of CAT, POD, and SOD, but decreased the concentratios of H2O2 in the shoots of Indian mustard. Liner analysis showed negative correlations between H2O2 concentrations and the activities of CAT and POD, which implyed that CAT and POD probably played important roles in protecting the plants from H2O2 toxicity under Hg stress.
We screened and investigated the potential of three plants (Pteris vittata, Brassica juncea, and Polypogon monspeliensis) in phytoextraction of Hg-contaminated soil. Three independent experiments were performed in turn. (1) Pot study with potting mix and HgCl2 solution. (2) Pot study with freshly HgCl2-spiked soil. (3) Experiment using aged soil contaminated with different Hg sources. Both shoots and roots of Pteris vittata could accumulate large amount of Hg from many kinds of Hg-contaminated soils (such as HgCl2, Hg(NO3)2, or HgS) without any visible stress symptoms. Therefore, Pteris vittata was a possible candidate for Hg phytoextraction and phytostabilization. Elemental Hg can evaporate into air because of its special physical properties. So we designed a chamber experiment to study the foliar Hg uptake by plants. Shoots of both Pteris vittata and Brassica juncea accumulated Hg at the concentratios of 63-105 mg/kg from Hg-contaminated air. So both Pteris vittata and Brassica juncea probably have potential in remediating Hg-contaminated air.
The application of remote sensing in monitoring plant physiological status has been growing concerns. We found that relative spectral reflectances in visible region (390-780 nm) and near infrared region (NIR,800-1300 nm) were sensitive to leaf structure and physiological status in Pteris vittata during the phytoremediation of Hg-contaminated soil. We selected several high spectral resolution reflectance vegetation indices to evaluate the change of spectral reflectance in Pteris vittata. We also estabilished the correlations between NDVI and biomass, R1110/R810 and leaf structure, WI and relative water content, respectively, in the shoots of Pteris vittata grown in Hg-contaminated soil. Hg stress induced significant increase in relative reflectance around 680 nm corresponding to the breakdown of chloroplast observed in TEM photographs in Pteris vittata. These results provided some theoretical evidences for the application of remote sensing in monitoring plant physiological status during the phytoremediation of Hg-contaminated soil.
引文
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