外源微生物对小麦和蜈蚣草砷吸收转化的影响及机制研究
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摘要
根际微生物在重金属元素的生物地球化学循环中扮演着重要的角色。大量的研究揭示了植物-根际微生物之间的联合作用在植物养分利用和重金属吸收方面所起到的作用,但是相关的机制尚缺乏充分的探讨。本项目以砷超富集植物蜈蚣草(Pteris vittata L.)和粮食作物小麦(Triticum aestivum L.)为供试材料,研究了不同的外源微生物(砷酸还原菌：Ts33、Ts41；亚砷酸氧化菌：C13、D14)对上述两种植物吸收、积累和转化As的影响及其机制,研究结果对于开发有效的植物-微生物联合修复技术,及寻找抑制农作物吸收As、保障农产品品质安全的技术方法具有重要意义。主要研究结果如下：
1.采用石英砂培试验研究了砷酸还原菌(Ts33、Ts41)对小麦As吸收和价态变化的影响。研究结果表明：在低As处理浓度(≤10 mg·kg-1)时,除Ts41对小麦地上部的作用外,添加砷酸还原菌能够促进小麦对As的吸收,小麦地上部和地下部As浓度分别是不加菌处理的1.03-2.30倍和1.10-6.07倍。但是,随着As处理浓度的升高,添加砷酸还原菌能明显抑制小麦对As的吸收。与不加菌相比,添加砷酸还原菌小麦地上部、地下部As浓度分别减少64.04%-95.85%和2.58%-78.08%。As(Ⅲ)和As(Ⅴ)是As在小麦体内存在的两种主要价态。砷酸还原菌对小麦地上部As价态的影响不太,但是能显著提高小麦地下部和砂培营养液中As(Ⅲ)的百分含量,分别是不加菌处理的1.95和4.16倍。
2.研究了石英砂培条件下砷酸还原菌(Ts33、Ts41)对小麦抗As胁迫反应的影响。研究结果表明：低浓度As(<20 mg·kg-1)处理提高小麦叶片SOD、POD、CAT活性和GSH含量,而其中SOD活性在As处理浓度>10 mg·kg-1时就开始降低；高浓度As处理下(≥20 mg·kg-1),叶片SOD、POD、CAT活性和GSH含量降低。另外,添加砷酸还原菌(Ts33和Ts41)能显著增加叶片SOD、POD、CAT活性,同时降低MDA含量。添加Ts33和Ts41后,小麦叶片SOD、POD、CAT活性最大为不加菌对照的2.27、2.14、1.18倍,而MDA含量降低最大时其值是不加菌对照的38.29%。
3.收集和测定了添加砷酸还原菌(Ts33、Ts41)和As胁迫下小麦根系分泌物中的有机酸。研究结果表明：根系分泌有机酸的种类随As处理浓度的增加而增加。当As处理浓度为0 mg·kg-1时,小麦根系主要分泌的是草酸和乙酸；当As处理浓度为5 mg·kg-1时,增加分泌苹果酸；而As处理浓度≥10 mg·kg-1时,增加分泌酒石酸和柠檬酸。同时,As处理浓度的增加也促进根系分泌物分泌量的增加。其中,苹果酸、酒石酸和柠檬酸的增加达到显著差异,在As处理浓度为50 mg·kg-1时分别是As处理浓度为10 mg·kg-1时的5.46、1.06和1.97倍。另外,添加外源微生物会减少小麦根系分泌有机酸的种类。与不加菌处理相比,添加微生物处理的小麦根系分泌物中没有检测到酒石酸和柠檬酸。在微生物作用下,小麦根系分泌物的分泌量也有减少,其中苹果酸的分泌量显著减少。Ts41处理下小麦分泌的苹果酸量减少最多,只有不加菌处理的28.46%。
4.应用土壤盆栽试验研究了砷酸还原菌(Ts33、Ts41)和亚砷酸氧化菌(C13、D14)对蜈蚣草As吸收和价态的影响。研究结果表明：2类微生物都可以促进蜈蚣草对As的吸收。比如,在加菌处理5 d后,蜈蚣草地上部和地下部As浓度分别增加86.33%-187.13%和53.93%-108.82%。另外,砷酸还原菌能够将蜈蚣草体内As(Ⅲ)的百分含量提高2.99-49.01%；而亚砷酸氧化菌能够将蜈蚣草体内As(Ⅴ)的百分含量提高9.09-22.91%。
5.测定了砷酸还原菌(Ts33、Ts41)和亚砷酸氧化菌(C13、D14)作用下土壤的As含量和As价态。研究结果表明：添加微生物能够使根际As含量减少9.97%-18.64%。砷酸还原菌能够促进土壤中的As(Ⅴ)还原为As(Ⅲ),只加砷酸还原菌处理的土壤中As(Ⅲ)的百分含量为27.90%-30.87%。但是,蜈蚣草的根际活动会减弱砷酸还原菌对土壤中As的还原作用。种植蜈蚣草同时又添加砷酸还原菌Ts41的处理中,蜈蚣草根际土As(Ⅲ)的百分含量为只添加Ts41处理土的74.57%-80.86%。
6.收集和测定了在盆栽土壤中添加砷酸还原菌(Ts33、Ts41)和亚砷酸氧化菌(C13、D14)时蜈蚣草根系分泌物中的有机酸。研究结果表明：2类微生物作用对根系分泌物中有机酸组成的影响都不大,但是能够增加分泌的量。根系分泌物中有机酸主要为草酸、苹果酸、乙酸和琥珀酸,其中草酸的质量分数最高,超过了总酸的56%,是蜈蚣草根系分泌的低分子量有机酸的主要成分。草酸增加了22.74%-270.93%,苹果酸增加了31.06%-127.70%,乙酸增加了19.49%-80.12%。
Rhizosphere microorganisms play an important role in the biogechemical cycling of heavy metals. Many studies have revealed the synergetic effect between plant and rhizosphere microorganism in nutrient utilization and heavey metals uptake. However, the relevant mechanisms are still not clear.
In order to find an effective plant-microbial remediation technology for polluted soils, and to obtain a way to inhibit accumulation of pollutants in agricultural crops for the food safety, this research used an arsenic hyperaccumulator (Pteris vittata L.) and a food crop wheat (Triticum aestivum L.) as materials to study the influence of arsenate reducing bacteria (Ts33, Ts41) and arsenite oxization bacteria (C13, D14) on the uptake, utilization and speciation of arsenic and the primary mechanisms. The main results are summarized as follows:
1. The effects of arsenate reducing bacteria (Ts33, Ts41) on arsenic uptake and speciation in wheat were studied via the quartz pot experiment. The results showed that except for As uptaken by wheat shoots influenced by Ts41, addition of arsenate reducing bacteria promoted the As uptake of wheat with the increase of As concentrations when As concentration<= 10 mg·kg-1, being 1.03-2.30 times in shoots and 1.10-6.07 times in roots in comparison with the control without bacteria. However, with the increase of the concentration of As, the bacteria activities inhibited the As uptake of wheat. Compared to the control without bacteria addition, As concentrations in shoots and roots were decreased by 64.04%-95.85% and 2.58%-78.08% respectively. The As(Ⅲ) and As(Ⅴ) were the main species of arsenic in wheat. Arsenate reducing bacteria hardly had effects on the arsenic speciation in wheat shoots, but could increased As(Ⅲ) in wheat roots and sand culture nutrient solution, with the increase of 1.95 and 4.16 times over the control without bacteria addition.
2. The effects of arsenate reducing bacteria (Ts33, Ts41) on the arsenic stressed response of wheat under quartz pot growth condition were studied. The results showed that As treatments at the concentration of< 20 mg·kg-1 improved the activity of SOD, POD, CAT and the content of GSH in wheat leaves, but had the opposite effect at As>= 20 mg·kg-1. In addition, the bacteria (Ts33, Ts41) increased SOD, POD, CAT activity and decreased MDA content. The greatest activity of SOD, POD, CAT in wheat leaves with Ts33, Ts41 treatments were 2.27,2.14,1.18 times respectively over the control without bacteria, and the lowest MDA content was only 38.29% of the control without bacteria.
3. The organic acids in root exudates of wheat under arsenic stress influenced by arsenate reducing bacteria (Ts33, Ts41) were determined by HPLC techniques. The results showed that the increase of the concentration of As treatment raised the kind of wheat root exudates. The oxalic acid and acetic acid were the major components of wheat root exudates without As addition. However, malic acid, and tartaric acid/citric acid emerged when treated with 5 or 10 mg·kg-1 of As respectively. Meanwhile, the increase of As concentration in the treatments also raised the quantity of wheat root exudates. The amounts of malic acid, tartaric acid and citric acid increased signifcantly at the As concentration of 50 mg·kg-1, which were 5.46,1.06 and 1.97 times over the treatment with As concentration of 10 mg·kg-1. Addition of the bacteria reduced the types of organic acids in wheat root exudates. The tartaric acid and citric acid were not detected when adding the bacteria. In addition, the quantity of malic acid was significantly decreased by Ts33, Ts41 treatments, and the quantity of malic acid in Ts41 treatment reduced the most, which was by 28.46%% in comparison with the control without baterica addition.
4. A soil pot experiment was carried out to study the effects of arsenate reducing bacteria (Ts33, Ts41) and arsenite oxidizing bacteria (C13, D14) on As uptake and speciation of Pteris vittata. The results showed that addition of two kinds of bacteria increased As uptake by Pteris vittata, with the As content of shoots and roots of Pteris vittata increased by 86.33%-187.13% and 53.93%-108.82% respectively. In addition, arsenate reducing bacteria raised As(Ⅲ) percentages in Pteris vittata by 2.99%-49.01% compared with the control without bacteria addition. Meanwhile, the arsenite oxidizing bacteria raised As(Ⅴ) percentages by 9.09%-22.91%.
5. As content and speciation in soil was measured to understand the effects of arsenate reducing bacteria (Ts33, Ts41) and arsenite oxidizing bacteria (C13, D14). The results showed that addition of two kinds of bacteria decreased As contents in rhizosphere soils by 9.97%-18.64%. Arsenate reducing bacteria contributed to the reduction of As(Ⅴ) in soil, and the soil As(Ⅲ) percentages were 27.90%-30.87% when treated merely with the bacteria. However, the root activities of Pteris vittata inhibited the arsenate reduction by arsenate reducing bacteria. The As(Ⅲ) percentages in rhizosphere soil of the treatments with both Pteris vittata and Ts41 were 74.57%-80.86% of the treatments Ts41.
6. The determination of organic acids in root exudates of Pteris vittata influenced by arsenate reducing bacteria (Ts33, Ts41) and arsenite oxidizing bacteria (C13, D14). The results showed that bacteria addition had little impacts on the organic acid types of root exudates. Oxalic acid, acetic acid, malic acid and succinic acid were the mian organic acids in root exudates, and the concentration of oxalic acid was the highest, which was more than 56% of the total of these four kinds of organic acids. However, bacteria addition increased the exudating amount of organic acids, as the amount of oxalic acid, malic acid and acetic acid increased by 22.74%-270.93%,31.06%-127.70% and 19.49%-80.12% respectively.
1.采用石英砂培试验研究了砷酸还原菌(Ts33、Ts41)对小麦As吸收和价态变化的影响。研究结果表明：在低As处理浓度(≤10 mg·kg-1)时,除Ts41对小麦地上部的作用外,添加砷酸还原菌能够促进小麦对As的吸收,小麦地上部和地下部As浓度分别是不加菌处理的1.03-2.30倍和1.10-6.07倍。但是,随着As处理浓度的升高,添加砷酸还原菌能明显抑制小麦对As的吸收。与不加菌相比,添加砷酸还原菌小麦地上部、地下部As浓度分别减少64.04%-95.85%和2.58%-78.08%。As(Ⅲ)和As(Ⅴ)是As在小麦体内存在的两种主要价态。砷酸还原菌对小麦地上部As价态的影响不太,但是能显著提高小麦地下部和砂培营养液中As(Ⅲ)的百分含量,分别是不加菌处理的1.95和4.16倍。
2.研究了石英砂培条件下砷酸还原菌(Ts33、Ts41)对小麦抗As胁迫反应的影响。研究结果表明：低浓度As(<20 mg·kg-1)处理提高小麦叶片SOD、POD、CAT活性和GSH含量,而其中SOD活性在As处理浓度>10 mg·kg-1时就开始降低；高浓度As处理下(≥20 mg·kg-1),叶片SOD、POD、CAT活性和GSH含量降低。另外,添加砷酸还原菌(Ts33和Ts41)能显著增加叶片SOD、POD、CAT活性,同时降低MDA含量。添加Ts33和Ts41后,小麦叶片SOD、POD、CAT活性最大为不加菌对照的2.27、2.14、1.18倍,而MDA含量降低最大时其值是不加菌对照的38.29%。
3.收集和测定了添加砷酸还原菌(Ts33、Ts41)和As胁迫下小麦根系分泌物中的有机酸。研究结果表明：根系分泌有机酸的种类随As处理浓度的增加而增加。当As处理浓度为0 mg·kg-1时,小麦根系主要分泌的是草酸和乙酸；当As处理浓度为5 mg·kg-1时,增加分泌苹果酸；而As处理浓度≥10 mg·kg-1时,增加分泌酒石酸和柠檬酸。同时,As处理浓度的增加也促进根系分泌物分泌量的增加。其中,苹果酸、酒石酸和柠檬酸的增加达到显著差异,在As处理浓度为50 mg·kg-1时分别是As处理浓度为10 mg·kg-1时的5.46、1.06和1.97倍。另外,添加外源微生物会减少小麦根系分泌有机酸的种类。与不加菌处理相比,添加微生物处理的小麦根系分泌物中没有检测到酒石酸和柠檬酸。在微生物作用下,小麦根系分泌物的分泌量也有减少,其中苹果酸的分泌量显著减少。Ts41处理下小麦分泌的苹果酸量减少最多,只有不加菌处理的28.46%。
4.应用土壤盆栽试验研究了砷酸还原菌(Ts33、Ts41)和亚砷酸氧化菌(C13、D14)对蜈蚣草As吸收和价态的影响。研究结果表明：2类微生物都可以促进蜈蚣草对As的吸收。比如,在加菌处理5 d后,蜈蚣草地上部和地下部As浓度分别增加86.33%-187.13%和53.93%-108.82%。另外,砷酸还原菌能够将蜈蚣草体内As(Ⅲ)的百分含量提高2.99-49.01%；而亚砷酸氧化菌能够将蜈蚣草体内As(Ⅴ)的百分含量提高9.09-22.91%。
5.测定了砷酸还原菌(Ts33、Ts41)和亚砷酸氧化菌(C13、D14)作用下土壤的As含量和As价态。研究结果表明：添加微生物能够使根际As含量减少9.97%-18.64%。砷酸还原菌能够促进土壤中的As(Ⅴ)还原为As(Ⅲ),只加砷酸还原菌处理的土壤中As(Ⅲ)的百分含量为27.90%-30.87%。但是,蜈蚣草的根际活动会减弱砷酸还原菌对土壤中As的还原作用。种植蜈蚣草同时又添加砷酸还原菌Ts41的处理中,蜈蚣草根际土As(Ⅲ)的百分含量为只添加Ts41处理土的74.57%-80.86%。
6.收集和测定了在盆栽土壤中添加砷酸还原菌(Ts33、Ts41)和亚砷酸氧化菌(C13、D14)时蜈蚣草根系分泌物中的有机酸。研究结果表明：2类微生物作用对根系分泌物中有机酸组成的影响都不大,但是能够增加分泌的量。根系分泌物中有机酸主要为草酸、苹果酸、乙酸和琥珀酸,其中草酸的质量分数最高,超过了总酸的56%,是蜈蚣草根系分泌的低分子量有机酸的主要成分。草酸增加了22.74%-270.93%,苹果酸增加了31.06%-127.70%,乙酸增加了19.49%-80.12%。
Rhizosphere microorganisms play an important role in the biogechemical cycling of heavy metals. Many studies have revealed the synergetic effect between plant and rhizosphere microorganism in nutrient utilization and heavey metals uptake. However, the relevant mechanisms are still not clear.
In order to find an effective plant-microbial remediation technology for polluted soils, and to obtain a way to inhibit accumulation of pollutants in agricultural crops for the food safety, this research used an arsenic hyperaccumulator (Pteris vittata L.) and a food crop wheat (Triticum aestivum L.) as materials to study the influence of arsenate reducing bacteria (Ts33, Ts41) and arsenite oxization bacteria (C13, D14) on the uptake, utilization and speciation of arsenic and the primary mechanisms. The main results are summarized as follows:
1. The effects of arsenate reducing bacteria (Ts33, Ts41) on arsenic uptake and speciation in wheat were studied via the quartz pot experiment. The results showed that except for As uptaken by wheat shoots influenced by Ts41, addition of arsenate reducing bacteria promoted the As uptake of wheat with the increase of As concentrations when As concentration<= 10 mg·kg-1, being 1.03-2.30 times in shoots and 1.10-6.07 times in roots in comparison with the control without bacteria. However, with the increase of the concentration of As, the bacteria activities inhibited the As uptake of wheat. Compared to the control without bacteria addition, As concentrations in shoots and roots were decreased by 64.04%-95.85% and 2.58%-78.08% respectively. The As(Ⅲ) and As(Ⅴ) were the main species of arsenic in wheat. Arsenate reducing bacteria hardly had effects on the arsenic speciation in wheat shoots, but could increased As(Ⅲ) in wheat roots and sand culture nutrient solution, with the increase of 1.95 and 4.16 times over the control without bacteria addition.
2. The effects of arsenate reducing bacteria (Ts33, Ts41) on the arsenic stressed response of wheat under quartz pot growth condition were studied. The results showed that As treatments at the concentration of< 20 mg·kg-1 improved the activity of SOD, POD, CAT and the content of GSH in wheat leaves, but had the opposite effect at As>= 20 mg·kg-1. In addition, the bacteria (Ts33, Ts41) increased SOD, POD, CAT activity and decreased MDA content. The greatest activity of SOD, POD, CAT in wheat leaves with Ts33, Ts41 treatments were 2.27,2.14,1.18 times respectively over the control without bacteria, and the lowest MDA content was only 38.29% of the control without bacteria.
3. The organic acids in root exudates of wheat under arsenic stress influenced by arsenate reducing bacteria (Ts33, Ts41) were determined by HPLC techniques. The results showed that the increase of the concentration of As treatment raised the kind of wheat root exudates. The oxalic acid and acetic acid were the major components of wheat root exudates without As addition. However, malic acid, and tartaric acid/citric acid emerged when treated with 5 or 10 mg·kg-1 of As respectively. Meanwhile, the increase of As concentration in the treatments also raised the quantity of wheat root exudates. The amounts of malic acid, tartaric acid and citric acid increased signifcantly at the As concentration of 50 mg·kg-1, which were 5.46,1.06 and 1.97 times over the treatment with As concentration of 10 mg·kg-1. Addition of the bacteria reduced the types of organic acids in wheat root exudates. The tartaric acid and citric acid were not detected when adding the bacteria. In addition, the quantity of malic acid was significantly decreased by Ts33, Ts41 treatments, and the quantity of malic acid in Ts41 treatment reduced the most, which was by 28.46%% in comparison with the control without baterica addition.
4. A soil pot experiment was carried out to study the effects of arsenate reducing bacteria (Ts33, Ts41) and arsenite oxidizing bacteria (C13, D14) on As uptake and speciation of Pteris vittata. The results showed that addition of two kinds of bacteria increased As uptake by Pteris vittata, with the As content of shoots and roots of Pteris vittata increased by 86.33%-187.13% and 53.93%-108.82% respectively. In addition, arsenate reducing bacteria raised As(Ⅲ) percentages in Pteris vittata by 2.99%-49.01% compared with the control without bacteria addition. Meanwhile, the arsenite oxidizing bacteria raised As(Ⅴ) percentages by 9.09%-22.91%.
5. As content and speciation in soil was measured to understand the effects of arsenate reducing bacteria (Ts33, Ts41) and arsenite oxidizing bacteria (C13, D14). The results showed that addition of two kinds of bacteria decreased As contents in rhizosphere soils by 9.97%-18.64%. Arsenate reducing bacteria contributed to the reduction of As(Ⅴ) in soil, and the soil As(Ⅲ) percentages were 27.90%-30.87% when treated merely with the bacteria. However, the root activities of Pteris vittata inhibited the arsenate reduction by arsenate reducing bacteria. The As(Ⅲ) percentages in rhizosphere soil of the treatments with both Pteris vittata and Ts41 were 74.57%-80.86% of the treatments Ts41.
6. The determination of organic acids in root exudates of Pteris vittata influenced by arsenate reducing bacteria (Ts33, Ts41) and arsenite oxidizing bacteria (C13, D14). The results showed that bacteria addition had little impacts on the organic acid types of root exudates. Oxalic acid, acetic acid, malic acid and succinic acid were the mian organic acids in root exudates, and the concentration of oxalic acid was the highest, which was more than 56% of the total of these four kinds of organic acids. However, bacteria addition increased the exudating amount of organic acids, as the amount of oxalic acid, malic acid and acetic acid increased by 22.74%-270.93%,31.06%-127.70% and 19.49%-80.12% respectively.
引文
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