香薷植物耐高铜毒害的机制及修复植物材料资源化利用前景
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摘要
重金属污染是土壤污染的主要类型之一。近年来,铜污染土壤的植物修复研究在国内掀起热潮。采用海州香薷开展铜污染土壤的研究,已从实验室水培、盆栽试验的生长特性反应和耐及解铜毒的生理生化反应,进展到室外大田修复污染土壤的示范工程及修复技术推广应用阶段。紫花香薷被报道为另一我国原生铜耐性/富集植物,在重金属复合污染土壤上,也有修复前景。本文以海州香薷和紫花香薷为材料,采用水培和大田实验,运用透射电镜(TEM)和能谱(EDX)技术、傅立叶红外(FTIR)技术、气质联用(GC/MS)技术、ICP分析技术等,研究了海州香薷和紫花香薷对铜胁迫的响应、高铜产生的细胞毒性及细胞壁组分变化、细胞壁及其组分参与解铜毒作用、多金属污染土壤的改良-植物修复效应、及海州香薷和紫花香薷地上部挥发油成分分析,阐明海州香薷和紫花香薷耐高铜毒害的机制及其修复材料资源化利用前景。主要研究结果总结如下:
1.水培试验表明,诸暨海州香薷耐铜能力远高于三门紫花香薷PFE1及WFE,九溪紫花香薷耐铜性最差。50μmol L~(-1)Cu处理时,诸暨海州香薷对铜的耐性指数为TI>100%,而PFE1、WFE和PFE2分别为65%、62%、58%;100μmol L~(-1)Cu处理时,诸暨海州香薷、PFE1、WFE和PFE2对铜的耐性指数(TI)分别下降为95%、62%、59%和42%。铜处理对紫花香薷的毒害甚于海州香薷,100μmol L~(-1)Cu处理时,诸暨海州香薷体内大量和微量元素变化不显著,而50μmol L~(-1)Cu处理时,PFE1、WFE和PFE2中K明显降低。就生物量而言,<100μmol L~(-1)Cu处理时,同样铜处理水平下,九溪紫花香薷的生物量明显高于诸暨海州香薷和三门紫花香薷,对铜的富集能力为:PFE2>WFE>PFE1>诸暨海州香薷。根和地上部生物富集因子均为:PFE2>WFE>PFE1>诸暨海州香薷。
2.TEM超微结构表明,矿山生态型海州香薷的根、茎及叶细胞在50μmol L~(-1)Cu处理时,没有显著的毒害;100μmol L~(-1)Cu处理时,海州香薷的根细胞器有明显受损,但茎及叶细胞器正常。而矿山生态型紫花香薷的根、茎及叶在50μmol L~(-1)Cu处理时,已发生明显的毒害。紫花香薷根部受到的毒害最严重,其次是茎、叶。250μmol L~(-1)Cu处理时,海州香薷的根及茎细胞壁显著增厚,而50μmol L~(-1)Cu处理时,紫花香薷的根及茎细胞壁明显增厚。250μmol L~(-1)Cu处理的海州香薷,以及50μmol L~(-1)Cu处理的紫花香薷的根及茎细胞壁增厚的现象,对植物抵抗重金属污染起着重要作用。
3.EDS能谱分析表明,100μmol L~(-1)Cu处理时,海州香薷的根、茎及叶细胞壁表面及细胞壁内,有大量的高密度电子体沉积,且其中Cu含量和Cu/C及Cu/O比值较高,尤其时海州香薷根细胞壁内的Cu/C及Cu/O比值最高。海州香薷的细胞壁,尤其是根细胞壁在富集和解高铜毒害中起着重要作用。50μmol L~(-1)Cu处理时,紫花香薷的根、茎及叶细胞壁上的Cu/C及Cu/O比值,要低于胞内高密度电子体中,对
Heavy metals contamination is the main type of soil contamination. Nowadays, more research work have been conducted on the phytoremediation of copper contaminated soil in China. Solution culture and pot experiment, field experiment have been done to investigate the physiological and biochemical response to copper toxicity, and its phytoremediation potential of copper from the contaminated soil by E. splendens. Elsholtzia argyi, another Cu-tolerant and accumulating plant species, also show the potential for the phytoremediation of heavy metal polluted soil. In this study, solution culture, pot culture and field experiment were used to do some investigations about both E. splendens and E. argyi: the response of both Elsholtzia plants to copper toxicity, Cu tolerance and accumulation in plant by transmission electron microscope (TEM) technique, cell wall nature and its complexation with copper by FTIR technique, sorption and desorption properties of copper to cell wall of both Elsholtzia plants, the phytoavailability of Cu, Pb, Zn and Cd in the multi-metal contaminated soil and rhizospheric soil of E. splendens as affected by soil amendments, and Cu, Pb, Zn and Cd removel by E. splendens, the changes of volatile oil components in shoot of both Elsholtzia plants under different Cu supplies both in solution culture and under naturally growing conditons by GC/MS technique, and their activities to resist Bacillus, Coccus, and so on. The main results are listed as follows:1. The response of both Elsholtzia plants to copper toxicity indicated that E. splendens and E. argyi naturally growing in Zhuji-mined soil, Sanmeng-mined soil, Fuyang contaminated soil, Huajiachi non-contaminated soil, and Jiuxi non- contaminated soil can grow well, with the concentrations of macro- and micro-nutrients in plant kept at the normal level, except for the plantt of E. argyi in the Fuyang contaminated soil, that the significantly constrained growth of plant and the increased levels of K, Mg and Ca were observed. Zhuji-E. splendens, Sanmen-E argyi (PFE1) ant its variation (WFE), Jiuxi-E. argyi (PFE2) were studied in response to copper toxicity. The results indicated that E. splendens can tolerate more copper than PFE1 and WFE, while PFE2 is more like Cu-sensitive plant species. 50 μmol L~(-1) Cu resulted in the tolerance index (TI) of Zhuji-E. splendens of >100%, while for PFE1, WFE and PFE2, it was 65%, 62%, 58%,
respectively. At 100 μmol L~(-1) Cu, TI of Zhuji-E splendens, PFE1, WFE and PFE2 were decreased to 95%, 62%, 59% and 42%, respectively. Copper caused more toxicty to E. argyi than to E. splendens. At 100 μmol L~(-1) Cu, no significant difference were noted for macronutrient and micronutrient in Zhuji-E. splendens, while for PFE1, WFE and PFE2, the considerably reduced K was observed at 50 μmol L~(-1) Cu. As for biomass, when suppiled at the same Cu supplies level (<100 μmol L~(-1) Cu), PFE2 has the greater shoot biomass than Zhuji E. splendens and PFE1. The total accumulating factor for copper followed PFE2>WFE> PFE1>E. splendens, and the bioaccumulation factor for root and shoot were also exhbited: PFE2>WFE>PFE1>E. splendens. As compared to E. argyi, Zhuji-E.splendens can mobilize more copper to its root than PFE1.2. The ultrastructure results indicated that the marked tocixity was observed at the root, stem, and leaf of Sanmen E. argyi when plant exposed to 50 μmol L~(-1) Cu, following the order of root > stem > leaf. While for E. splendens, it didn't markedly affated by 50 μmol L~(-1) Cu. The thickened cell wall was observed under copper toxicity of ≥ 50μmol L~(-1) for E. argyi and ≥ 250μmol L~(-1) E. splendens.3. EDS analyses indicate that, at 100μmol L~(-1) Cu, Much more electron density bodieswer found at the surface of cell wall and within cell wall of root, stem and leaf of E. splendens, with the highest level Cu and higheat Cu/C ratio and Cu/O ration. The cell wall of E. splendens, especially root cell wall, play more important role in accumulating and detoxifying high copper toxicity. At 50μmol L~(-1) Cu, the
1.水培试验表明,诸暨海州香薷耐铜能力远高于三门紫花香薷PFE1及WFE,九溪紫花香薷耐铜性最差。50μmol L~(-1)Cu处理时,诸暨海州香薷对铜的耐性指数为TI>100%,而PFE1、WFE和PFE2分别为65%、62%、58%;100μmol L~(-1)Cu处理时,诸暨海州香薷、PFE1、WFE和PFE2对铜的耐性指数(TI)分别下降为95%、62%、59%和42%。铜处理对紫花香薷的毒害甚于海州香薷,100μmol L~(-1)Cu处理时,诸暨海州香薷体内大量和微量元素变化不显著,而50μmol L~(-1)Cu处理时,PFE1、WFE和PFE2中K明显降低。就生物量而言,<100μmol L~(-1)Cu处理时,同样铜处理水平下,九溪紫花香薷的生物量明显高于诸暨海州香薷和三门紫花香薷,对铜的富集能力为:PFE2>WFE>PFE1>诸暨海州香薷。根和地上部生物富集因子均为:PFE2>WFE>PFE1>诸暨海州香薷。
2.TEM超微结构表明,矿山生态型海州香薷的根、茎及叶细胞在50μmol L~(-1)Cu处理时,没有显著的毒害;100μmol L~(-1)Cu处理时,海州香薷的根细胞器有明显受损,但茎及叶细胞器正常。而矿山生态型紫花香薷的根、茎及叶在50μmol L~(-1)Cu处理时,已发生明显的毒害。紫花香薷根部受到的毒害最严重,其次是茎、叶。250μmol L~(-1)Cu处理时,海州香薷的根及茎细胞壁显著增厚,而50μmol L~(-1)Cu处理时,紫花香薷的根及茎细胞壁明显增厚。250μmol L~(-1)Cu处理的海州香薷,以及50μmol L~(-1)Cu处理的紫花香薷的根及茎细胞壁增厚的现象,对植物抵抗重金属污染起着重要作用。
3.EDS能谱分析表明,100μmol L~(-1)Cu处理时,海州香薷的根、茎及叶细胞壁表面及细胞壁内,有大量的高密度电子体沉积,且其中Cu含量和Cu/C及Cu/O比值较高,尤其时海州香薷根细胞壁内的Cu/C及Cu/O比值最高。海州香薷的细胞壁,尤其是根细胞壁在富集和解高铜毒害中起着重要作用。50μmol L~(-1)Cu处理时,紫花香薷的根、茎及叶细胞壁上的Cu/C及Cu/O比值,要低于胞内高密度电子体中,对
Heavy metals contamination is the main type of soil contamination. Nowadays, more research work have been conducted on the phytoremediation of copper contaminated soil in China. Solution culture and pot experiment, field experiment have been done to investigate the physiological and biochemical response to copper toxicity, and its phytoremediation potential of copper from the contaminated soil by E. splendens. Elsholtzia argyi, another Cu-tolerant and accumulating plant species, also show the potential for the phytoremediation of heavy metal polluted soil. In this study, solution culture, pot culture and field experiment were used to do some investigations about both E. splendens and E. argyi: the response of both Elsholtzia plants to copper toxicity, Cu tolerance and accumulation in plant by transmission electron microscope (TEM) technique, cell wall nature and its complexation with copper by FTIR technique, sorption and desorption properties of copper to cell wall of both Elsholtzia plants, the phytoavailability of Cu, Pb, Zn and Cd in the multi-metal contaminated soil and rhizospheric soil of E. splendens as affected by soil amendments, and Cu, Pb, Zn and Cd removel by E. splendens, the changes of volatile oil components in shoot of both Elsholtzia plants under different Cu supplies both in solution culture and under naturally growing conditons by GC/MS technique, and their activities to resist Bacillus, Coccus, and so on. The main results are listed as follows:1. The response of both Elsholtzia plants to copper toxicity indicated that E. splendens and E. argyi naturally growing in Zhuji-mined soil, Sanmeng-mined soil, Fuyang contaminated soil, Huajiachi non-contaminated soil, and Jiuxi non- contaminated soil can grow well, with the concentrations of macro- and micro-nutrients in plant kept at the normal level, except for the plantt of E. argyi in the Fuyang contaminated soil, that the significantly constrained growth of plant and the increased levels of K, Mg and Ca were observed. Zhuji-E. splendens, Sanmen-E argyi (PFE1) ant its variation (WFE), Jiuxi-E. argyi (PFE2) were studied in response to copper toxicity. The results indicated that E. splendens can tolerate more copper than PFE1 and WFE, while PFE2 is more like Cu-sensitive plant species. 50 μmol L~(-1) Cu resulted in the tolerance index (TI) of Zhuji-E. splendens of >100%, while for PFE1, WFE and PFE2, it was 65%, 62%, 58%,
respectively. At 100 μmol L~(-1) Cu, TI of Zhuji-E splendens, PFE1, WFE and PFE2 were decreased to 95%, 62%, 59% and 42%, respectively. Copper caused more toxicty to E. argyi than to E. splendens. At 100 μmol L~(-1) Cu, no significant difference were noted for macronutrient and micronutrient in Zhuji-E. splendens, while for PFE1, WFE and PFE2, the considerably reduced K was observed at 50 μmol L~(-1) Cu. As for biomass, when suppiled at the same Cu supplies level (<100 μmol L~(-1) Cu), PFE2 has the greater shoot biomass than Zhuji E. splendens and PFE1. The total accumulating factor for copper followed PFE2>WFE> PFE1>E. splendens, and the bioaccumulation factor for root and shoot were also exhbited: PFE2>WFE>PFE1>E. splendens. As compared to E. argyi, Zhuji-E.splendens can mobilize more copper to its root than PFE1.2. The ultrastructure results indicated that the marked tocixity was observed at the root, stem, and leaf of Sanmen E. argyi when plant exposed to 50 μmol L~(-1) Cu, following the order of root > stem > leaf. While for E. splendens, it didn't markedly affated by 50 μmol L~(-1) Cu. The thickened cell wall was observed under copper toxicity of ≥ 50μmol L~(-1) for E. argyi and ≥ 250μmol L~(-1) E. splendens.3. EDS analyses indicate that, at 100μmol L~(-1) Cu, Much more electron density bodieswer found at the surface of cell wall and within cell wall of root, stem and leaf of E. splendens, with the highest level Cu and higheat Cu/C ratio and Cu/O ration. The cell wall of E. splendens, especially root cell wall, play more important role in accumulating and detoxifying high copper toxicity. At 50μmol L~(-1) Cu, the
引文
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