薦草及其根际对模拟湿地中芘污染的降解机制
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摘要
本文以黄浦江湿地的典型植物藨草为修复植物，以芘为目标污染物，通过模拟实验考察了芘污染对藨草生长参数和酶活性的影响、芘污染及藨草种植对土壤酶活性和微生物群落结构的影响，阐明了藨草对芘污染的修复效果。并结合蛋白质组学的研究方法，揭示了藨草对芘污染响应的分子机制。主要研究结果与结论如下：
（1）藨草的种植能够提高芘的去除率，80天藨草处理后去除率可高达70%。芘的毒性效应在一定程度上阻碍了藨草的生长。芘浓度在10~40mg·kg-1之间时，能够降低藨草体内过氧化氢酶（CAT）活性，之后随芘浓度升高，CAT活性也随之升高。藨草生长早期，随芘浓度升高，秆过氧化物酶（POD）活性先受到抑制，而后被激发；而生长茂盛期，则经历激发—抑制—激发的过程。随着芘浓度的升高，藨草超氧化物歧化酶（SOD）的活性逐渐下降。藨草秆CAT的活性大于根部，而POD和SOD的活性则是根部大于秆。在藨草生长的三个不同阶段，三种酶活性均表现出显著性差异。
（2）芘对土壤CAT的影响表现为低浓度促进，高浓度抑制，且在藨草生长茂盛期和繁殖期，根际土壤和非根际土壤CAT活性远远高于空白土壤。在藨草繁殖期，芘对POD的活性有一定的抑制作用。在5~20mg kg-1的芘污染浓度范围内，藨草生长茂盛期的空白土壤和非根际土壤以及藨草繁殖期的根际土壤多酚氧化酶（PPO）活性与芘浓度呈显著正相关。芘浓度与土壤脱氢酶（DHA）活性显著负相关，且根际和非根际土壤中DHA活性高于未种植物的空白土壤。藨草生长阶段是四种土壤酶活性的主要影响因素，芘浓度及其与藨草生长阶段的交互作用对四种土壤酶活性的影响则与酶的种类和土壤的处理方式有关。
（3）较高浓度的芘能够减少某些细菌、真菌和放线菌的数量，但是能在一定程度上增加芘降解菌的数量。土壤pH值和有机质是影响微生物生长的重要参数。
（4）受芘污染后，土壤微生物的生物量均有不同程度的降低，而藨草的种植缓解了这种负面效应。不同微生物对芘污染的耐受性不同，细菌比真菌的耐受性强，革兰氏阳性菌比革兰氏阴性菌的耐受性强。藨草的引入可以刺激真菌、革兰氏阴性菌和好氧菌的生长。另外，主成分分析（PCA）显示不同芘浓度下微生物群落结构存在一定的差异。
（5）经芘处理后，藨草秆共发现37个具有显著差异的蛋白点，其中13个蛋白点的表达量是下调的，24个蛋白点的表达量是上调的；根部共发现55个具有显著差异的蛋白点，其中36个蛋白点的表达量是下调的，19个蛋白点的表达量是上调的。根据蛋白点表达量差异分析结果，将秆和根部差异较大的蛋白点进行质谱鉴定，结果在秆和根部样品中分别有17个和7个蛋白质得到了可靠的鉴定。
（6）将鉴定出的蛋白质进行功能分类，秆差异蛋白质包括：光合作用、防御、能量和物质代谢、辅酶代谢和蛋白质代谢相关蛋白质，其中参与光合作用的蛋白质占鉴定出的蛋白质的70%以上。而根部的蛋白质包括：胁迫反应、防御、能量和物质代谢以及蛋白质修饰相关的蛋白。芘胁迫严重影响了藨草秆的光合作用，同时对秆和根部的多个代谢过程、蛋白质合成及信号传递等产生影响。
Scirpus triqueter, a dominant species in wetland of Huangpu River, was firstly used asphytoremediation plant. Pyrene was used as objective contaminant. The objectives of thisstudy were:(1) to estimate the dissipation ratios of pyrene in the presence of S. triqueter;(2)to ascertain the growth parameters and physiological response of S. triqueter after pyrenetreatment;(3) to study the changes of the soil enzymes in different parts of the soils afterpyrene treatment at the presence of S. triqueter;(4) to investigate the response of themicrobial communities to pyrene and S. triqueter;(5) finally, to elucidate the molecularmechanism of S. triqueter in response to pyrene by using proteomic method. The majorfindings are summarized as follow:
(1) The dissipation ratios of pyrene were enhanced at the presence of S. triqueter, thedissipation ratio was up to70%after80-day treatment. Pyrene contaminated soil inhibited thegrowth of S. triqueter.10-40mg·kg–1of pyrene could reduce activities of catalase (CAT), andthen the activities would increase with increasing pyrene concentration. At the early growthstage of S. triqueter, the peroxidase (POD) activity of shoot was inhibited at first, and thenwas stimulated with increasing pyrene concentration; however, the activity experienced theprocess of stimulation-inhibition-stimulation at the vigorous stage. The activity of superoxidedismutase (SOD) declined with increasing pyrene concentration. Activity of CAT in shootwas higher than that in root, while the activities of POD and SOD in root were higher thanthose in shoot. Enzyme activities exhibited significant differences in different growth stages.
(2) The CAT activities of soils were promoted at lower pyrene concentrations and wereinhibited at higher pyrene concentrations; CAT activities in the rhizospheric soils increasedgreatly compared to those in the bulk and unplanted soils at the vigorous and reproductivestage of S. triqueter. The activities of POD in soils were inhibited by prene at the reproductivestage of S. triqueter. When the concentration range of pyrene was from5to20mg·kg–1, thepolyphenoloxidase (PPO) activities of bulk and unplanted soils at the vigorous stage andthose of rhizospheric soils at the reproductive stage were significantly positively related topyrene. However, the dehydrogenase (DHA) activities of soils were significantly negativelyrelated to pyrene; moreover, PPO activities in the rhizospheric soils were higher than those in the bulk and unplanted soils. The growth stage of S. triqueter was the main impact factor tothe activities of soil enzymes; the effects of pyrene concentration and the interactionsbetween growth stages and pyrene concentration on the activities of soil enzymes dependedon the type of enzyme and soil handling.
(3) High pyrene concentration reduced the populations of some bacteria, fungi andactinomyces, but increased pyrene-tolerant bacteria abundance. It is important that S.triqueter is favorable for the growth of microorganisms. The pH and organic matter of soilwere important parameters to the growth of microorganism.
(4) The biomass of soil microbes declined after pyrene treatment, however, the biomassincreased at the presence of S. triqueter. Bacteria were more tolerant to pyrene than fungi;while Gram-positive bacteria showed a greater tolerance than Gram-negative bacteria. Theintroduction of S. triqueter was benefit to fungi, Gram-negative bacteria and aerobic bacteria.Additionally, there was discrimination in microbial community structure under the stress ofpyrene from the principal component analysis (PCA).
(5) After treated by pyrene, a total of37protein spots from culm showed significantchanges, among which, the expression levels of13protein spots were downregulated,24protein spots were upregulated. A total of55protein spots from root showed significantchanges, and36protein spots were downregulated,19protein spots were upregulated.Protein spots which were significantly regulated by pyrene were identified by massspectrometry. A total of17and7proteins were successfully identified in shoot and root,respectively.
(6) The identified proteins were grouped according to functional categories. Proteins inshoot included: photosynthesis related protein, defense related protein, energy and materialmetabolism related protein, coenzyme metabolism related protein and protein metabolismrelated protein. Proteins involved in photosynthesis accounted for more than70%of theidentified proteins of shoot. The proteins in root included: stress response related proteins,defense related protein, energy and material metabolism related protein and proteinmodification related protein. Pyrene affected the photosynthesis of S. triqueter seriously,simultaneously, had an impact on the multiple metabolic processes, protein synthesis andsignal transduction.
（1）藨草的种植能够提高芘的去除率，80天藨草处理后去除率可高达70%。芘的毒性效应在一定程度上阻碍了藨草的生长。芘浓度在10~40mg·kg-1之间时，能够降低藨草体内过氧化氢酶（CAT）活性，之后随芘浓度升高，CAT活性也随之升高。藨草生长早期，随芘浓度升高，秆过氧化物酶（POD）活性先受到抑制，而后被激发；而生长茂盛期，则经历激发—抑制—激发的过程。随着芘浓度的升高，藨草超氧化物歧化酶（SOD）的活性逐渐下降。藨草秆CAT的活性大于根部，而POD和SOD的活性则是根部大于秆。在藨草生长的三个不同阶段，三种酶活性均表现出显著性差异。
（2）芘对土壤CAT的影响表现为低浓度促进，高浓度抑制，且在藨草生长茂盛期和繁殖期，根际土壤和非根际土壤CAT活性远远高于空白土壤。在藨草繁殖期，芘对POD的活性有一定的抑制作用。在5~20mg kg-1的芘污染浓度范围内，藨草生长茂盛期的空白土壤和非根际土壤以及藨草繁殖期的根际土壤多酚氧化酶（PPO）活性与芘浓度呈显著正相关。芘浓度与土壤脱氢酶（DHA）活性显著负相关，且根际和非根际土壤中DHA活性高于未种植物的空白土壤。藨草生长阶段是四种土壤酶活性的主要影响因素，芘浓度及其与藨草生长阶段的交互作用对四种土壤酶活性的影响则与酶的种类和土壤的处理方式有关。
（3）较高浓度的芘能够减少某些细菌、真菌和放线菌的数量，但是能在一定程度上增加芘降解菌的数量。土壤pH值和有机质是影响微生物生长的重要参数。
（4）受芘污染后，土壤微生物的生物量均有不同程度的降低，而藨草的种植缓解了这种负面效应。不同微生物对芘污染的耐受性不同，细菌比真菌的耐受性强，革兰氏阳性菌比革兰氏阴性菌的耐受性强。藨草的引入可以刺激真菌、革兰氏阴性菌和好氧菌的生长。另外，主成分分析（PCA）显示不同芘浓度下微生物群落结构存在一定的差异。
（5）经芘处理后，藨草秆共发现37个具有显著差异的蛋白点，其中13个蛋白点的表达量是下调的，24个蛋白点的表达量是上调的；根部共发现55个具有显著差异的蛋白点，其中36个蛋白点的表达量是下调的，19个蛋白点的表达量是上调的。根据蛋白点表达量差异分析结果，将秆和根部差异较大的蛋白点进行质谱鉴定，结果在秆和根部样品中分别有17个和7个蛋白质得到了可靠的鉴定。
（6）将鉴定出的蛋白质进行功能分类，秆差异蛋白质包括：光合作用、防御、能量和物质代谢、辅酶代谢和蛋白质代谢相关蛋白质，其中参与光合作用的蛋白质占鉴定出的蛋白质的70%以上。而根部的蛋白质包括：胁迫反应、防御、能量和物质代谢以及蛋白质修饰相关的蛋白。芘胁迫严重影响了藨草秆的光合作用，同时对秆和根部的多个代谢过程、蛋白质合成及信号传递等产生影响。
Scirpus triqueter, a dominant species in wetland of Huangpu River, was firstly used asphytoremediation plant. Pyrene was used as objective contaminant. The objectives of thisstudy were:(1) to estimate the dissipation ratios of pyrene in the presence of S. triqueter;(2)to ascertain the growth parameters and physiological response of S. triqueter after pyrenetreatment;(3) to study the changes of the soil enzymes in different parts of the soils afterpyrene treatment at the presence of S. triqueter;(4) to investigate the response of themicrobial communities to pyrene and S. triqueter;(5) finally, to elucidate the molecularmechanism of S. triqueter in response to pyrene by using proteomic method. The majorfindings are summarized as follow:
(1) The dissipation ratios of pyrene were enhanced at the presence of S. triqueter, thedissipation ratio was up to70%after80-day treatment. Pyrene contaminated soil inhibited thegrowth of S. triqueter.10-40mg·kg–1of pyrene could reduce activities of catalase (CAT), andthen the activities would increase with increasing pyrene concentration. At the early growthstage of S. triqueter, the peroxidase (POD) activity of shoot was inhibited at first, and thenwas stimulated with increasing pyrene concentration; however, the activity experienced theprocess of stimulation-inhibition-stimulation at the vigorous stage. The activity of superoxidedismutase (SOD) declined with increasing pyrene concentration. Activity of CAT in shootwas higher than that in root, while the activities of POD and SOD in root were higher thanthose in shoot. Enzyme activities exhibited significant differences in different growth stages.
(2) The CAT activities of soils were promoted at lower pyrene concentrations and wereinhibited at higher pyrene concentrations; CAT activities in the rhizospheric soils increasedgreatly compared to those in the bulk and unplanted soils at the vigorous and reproductivestage of S. triqueter. The activities of POD in soils were inhibited by prene at the reproductivestage of S. triqueter. When the concentration range of pyrene was from5to20mg·kg–1, thepolyphenoloxidase (PPO) activities of bulk and unplanted soils at the vigorous stage andthose of rhizospheric soils at the reproductive stage were significantly positively related topyrene. However, the dehydrogenase (DHA) activities of soils were significantly negativelyrelated to pyrene; moreover, PPO activities in the rhizospheric soils were higher than those in the bulk and unplanted soils. The growth stage of S. triqueter was the main impact factor tothe activities of soil enzymes; the effects of pyrene concentration and the interactionsbetween growth stages and pyrene concentration on the activities of soil enzymes dependedon the type of enzyme and soil handling.
(3) High pyrene concentration reduced the populations of some bacteria, fungi andactinomyces, but increased pyrene-tolerant bacteria abundance. It is important that S.triqueter is favorable for the growth of microorganisms. The pH and organic matter of soilwere important parameters to the growth of microorganism.
(4) The biomass of soil microbes declined after pyrene treatment, however, the biomassincreased at the presence of S. triqueter. Bacteria were more tolerant to pyrene than fungi;while Gram-positive bacteria showed a greater tolerance than Gram-negative bacteria. Theintroduction of S. triqueter was benefit to fungi, Gram-negative bacteria and aerobic bacteria.Additionally, there was discrimination in microbial community structure under the stress ofpyrene from the principal component analysis (PCA).
(5) After treated by pyrene, a total of37protein spots from culm showed significantchanges, among which, the expression levels of13protein spots were downregulated,24protein spots were upregulated. A total of55protein spots from root showed significantchanges, and36protein spots were downregulated,19protein spots were upregulated.Protein spots which were significantly regulated by pyrene were identified by massspectrometry. A total of17and7proteins were successfully identified in shoot and root,respectively.
(6) The identified proteins were grouped according to functional categories. Proteins inshoot included: photosynthesis related protein, defense related protein, energy and materialmetabolism related protein, coenzyme metabolism related protein and protein metabolismrelated protein. Proteins involved in photosynthesis accounted for more than70%of theidentified proteins of shoot. The proteins in root included: stress response related proteins,defense related protein, energy and material metabolism related protein and proteinmodification related protein. Pyrene affected the photosynthesis of S. triqueter seriously,simultaneously, had an impact on the multiple metabolic processes, protein synthesis andsignal transduction.
引文
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