镉形态分析与微区分布的质谱联用技术方法研究及其在印度芥菜耐镉机制中的应用
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摘要
超累积植物筛选是重金属污染土壤植物修复的基础和核心问题,同时也是污染环境植物修复的难点及前沿。明确植物抗重金属的生理和分子机理,从而提高植物在污染环境下的适应范围和生产能力,对于高效利用自然资源和进行污染环境的生物修复均具有重要意义。由于缺乏强有力的分析手段,植物体内重金属的迁移机制,解毒机制尚未得出一致结论。
本文在综述超积累植物对镉的耐性机制及其相关研究方法的基础上,选取镉富集植物印度芥菜为研究对象,重点开展镉的形态分析与微区分布方法研究,探讨植物螫合肽和区隔化在超积累植物对镉的耐受机制中的作用。综合运用高效液相色谱—电感耦合等离子体质谱(HPLC-ICP-MS)、超高效液相色谱—电喷雾质谱(UPLC-ESI-MS)和超高效液相色谱—飞行时间质谱(UPLC-QTOFMS)联用技术,建立了镉形态分析方法,重点解决了植物螫合肽(PCn)-Cd的定性分析难题,探讨植物螯合肽的形成机制及在植物体耐受与积累重金属中的作用。同时,借助激光烧蚀-电感耦合等离子体质谱(LA-ICP-MS)和透射电镜等手段,分析镉及其它六种元素在印度芥菜体内组织水平以及细胞水平上的分布特征,探讨各类组织/细胞对重金属的区隔化在植物体耐受耐镉机制中的作用。
本研究取得的主要研究结果如下：
1.通过温室水培试验研究了Cd胁迫时间和胁迫水平对印度芥菜生物量的影响以及Cd在植物体内的分布规律,从生物响应的角度探讨了Cd对印度芥菜的毒性及印度芥菜对Cd的耐性。印度芥菜对Cd的最大耐受量为1.0mg/L,超过此浓度植物的正常生长将受到影响,导致生物量下降。植物受胁迫48小时后出现中毒症状,生物量开始下降。随着胁迫时间和胁迫浓度增加,根、叶、茎中镉的浓度均显著增加,根部镉浓度远远大于茎和叶,茎大于叶。在考虑Cd对植物的危害时,胁迫时间和胁迫浓度两个因素要同时考虑。
2.利用UPLC-ESMSMS建立了PCn的分析方法,探讨了PCn稳定性及在耐镉机制中的作用,在Cd的强力诱导下,胁迫合成大量的PCn,从而能螯合更多的Cd,产生较强的耐Cd性。建立了PCn-Cd的UPLC-QTOFMS分析方法,检测出(PC2～PC5)-Cd形态。两种质谱联用技术为PCn-Cd的精确测定提供了强有力的分析工具,解决了由于Cd形态标准物质缺乏造成的SEC-HPLC-ICPMS定性难问题,也为研究PCn与其它金属络合物的分析方法提供了可能。
3.建立了SEC-HPLC-ICPMS测定Cd形态分析方法,在叶片和根部均检测到植物螯合肽(PC)3-Cd、植物螯合肽(PC)2-Cd、谷胱甘肽(GSH)-Cd,及半胱胺酸-Cd四种形态。在植物不同部位Cd存在形态不同,叶片中主要以GSH-Cd存在,而在根部主要以PC2-Cd为主。Cd形态极不稳定,样品处理采取液氮保护并-70℃保存,样品分析全流程用氮吹防氧化措施。研究结果证明,植物螯合肽的合成机制有两种,一是受Cd胁迫直接产生PCn,二是先形成GSH-Cd而后转化成PCn。
4.研究了胁迫浓度和胁迫时间对Cd形态分布的影响。可以推论：植物受Cd胁迫后,先储存在根部。随着胁迫浓度的增大,为了减轻Cd的毒害,先后形成GSH-Cd,PC2-Cd, PC3-Cd,并向叶部转移。同时,叶部受Cd胁迫产生的PCn也会传输到根部,结合更多的Cd,形成PCn-Cd,继续向叶部转移,从而使植物产生很强的Cd耐受性。另外,GSH与PCn结合Cd的能力也存在竞争。随着胁迫时间的增加,根和中GSH-Cd、PC2-Cd、PC3-Cd浓度先迅速增加,而后逐渐降低,96h降至最低。表明在植物耐镉机制中PCn起部分作用,受高浓度Cd胁迫时,其他机制可能会被激发。
5.建立了SEC-HPLC-ICP-MS同时测定印度芥菜中Cd、Cu、Zn形态分析的方法,在植物叶片和根部同时检测出三种元素的四种形态。在同一植物的不同部位,叶片中主要以GSH-Cd、Cys-Cu、 Cys-Zn为主,根部以PC2-Cd、PC2-Cu、Cys-Cu和Cys-Zn为主。另外,Cd、Cu、Zn在与谷胱甘肽(GSH)和半胱胺酸(Cys)的结合上有竞争。研究结论为一种植物同时修复Cd、Cu、Zn复合污染提供了可能。
6.借助LA-ICP-MS质谱技术对富集型印度芥菜茎、叶、叶柄中Cd、P、3、K、Ca、Cu和Zn七种元素的分布特征进行原位分析。结果表明,在茎、叶柄以及主叶脉等组织中,Cd优先分布于含木质部与韧皮部的维管组织中,表明Cd在运输过程中,不易向其他贮存组织扩散。在植物叶片中,Cd主要分布于含有大型液泡的叶肉细胞中,而在上下表皮中分布较低,表明液泡可能是植物存储镉的主要场所。其他元素与Cd的相关性分析结果表Cd与Ca具有相似的分布规律,可能与二者离子半径相近、化学性质相似有关。而K和Ca、P和S的分布呈显著正相关。说明重金属元素进入植株体内并被其吸收运输过程是伴随着植物对其他元素的吸收,且具有相似的运输机制。
7.扫描电镜结果证实胁迫浓度超过lmg/L后,细胞结构发生了变化。同时,细胞壁中镉浓度大于细胞液且远远大于细胞器,证明细胞区隔化在镉的解毒机制中具有重要的作用。
Cadmium (Cd) pollution in the soil is one of the most significant global environmental problems, as it will enter into human bodies through the accumulation of the food chain, and endangers the health of human being. Hyperaccumulator and accumulator species, which efficiently tolerate and accumulate heavy metals from the soil into shoots, have great promise in phytoremediation of contaminated environments. Understanding the mechanisms of metal tolerance and accumulation will provide insight into the identification and management of these hyperaccumulating species. Because of the analytical limit, there has not yet a common accepted conclusion on the migration mechanism of the heavy metal within the plant, as well as the detoxification mechanisms. Phytochelatins (PCn), characterized by the amino acid structure (y-Glu-Cys)n-Gly, where n ranges from2to11, are synthesized enzymatically by phytochelatin synthase (PCn) using glutathione as a substrate in the presence of many metals and metalloids.They are believed to play an important role in metal detoxification and tolerance. But few research on speciation analysis of PCn-Cd and its unstability on detoxification mechanism has been studied.
Therefore, in order to reveal migration and transformation pattern of heavy metals and investigate the relationship between speciation and tolerance of heavy metals, some studies on Indian Mustard which was demonstrated to tolerate and accumulate considerable amounts of cadmium were carried out. The aim was to establish methods for speciation and distribution analysis of cadmium, and obtain fundamental information on heavy metals accumulation and tolerance mechanisms by investigation of the metal localization and ligand abundance. Several modern hyphenated mass spectrometric techniques, such as high performance liquid chromatography-inductively coupled plasma mass spectrometry (HPLC-ICP-MS), ultra performance liquid chromatography-electrospray ilonization mass spectrometry (UPLC-ESI-MS), ultra performance liquid chromatography-time of flight mass spectrometry (UPLC-QTOFMS) and laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS) were used.
The main results include:
1. Exposure concentration of Cd above1.0mg/L will bring negative effects on the normal growth of the plant, and therefore result in the decrease of the biomass. Within48hours after the stimulation, the symptoms of intoxication can be observed in the plant. With the increase of the exposure time and concentration, the concentrations of cadmium in root, leaf and stem also increases. The concentration of cadmium in the root is far more than that in stem and leave, and and the lowest in in leave.
2. In this study, in vitro formed PCn-Cd complexes were characterized using UPLC-ESI-MSMS and UPLC-QTOFMS. These two methods have been demonstrated to be ideal and promising techniques for screening and characterizing peptide-metal complexes. It was demonstrated that both techniques had the ability to identify the formation of PCn-Cd complex.Furthermore, the increase of PC after Cd exposure demonstrated that Cd is the most efficient inducer of phytochelatin synthesis.
3. Method based on the coupling of size-exclusion HPLC with ICP-MS was used for speciation analysis of cadmium binding non-protein thiols in extracts of Indian mustard (Brassica Juncea). In the presence of Cd stress, PCn were induced, and Cd ions in extracts were associated with the induced PCn. So four different Cd species were detected in samples:PC3-Cd、PC2-Cd、GSH-Cd and Cys-Cd, which demonstrated that PCn were enzymatically synthesised from glutathione (GSH, Glu-Cys-Gly) by the constitutive enzyme PC synthase. The results indicated that GSH-Cd exists as the main specie in leaf samples while PC2-Cd is the main specie in root samples. A non-oxidizing (Nitrogen) preparation environment and a-70℃preservation condition were employed to avoid the oxidation of sulfhydryl groups.
4. Root and leaf samples were submitted to0(control),2.0,4.0,6.0,8.0,10.0mg L-1and harvested after1,2,3and4days exposure. Results indicated lower Cd stress levels were generally associated with higer PCn-Cd production. Higher stress induced the reduction of PCn-Cd, which proved that thiol based tolerance was only efficient in lower degrees of stress. Increasing exposure time also induced decreased PCn-Cd pordution indicated PCn may only had a partial role in metal resistance. Under continuous higher stress, plants maybe trigger other mechnisms to tolerate heavy metal toxicity. Our results suggested the concentration and time of exposure are important factors that must be taken into consideration when evaluating the true role of PCn in heavy metal detoxification.
5. Method based on the coupling of size-exclusion HPLC with ICP-MS was used for speciation analysis of Cd, Cu and Zn binding non-protein thiols in extracts of Indian mustard(Brassica Juncea). In the presence of Cd, Cu and Zn stress, PCn were induced, and metal ions in extracts were associated with the induced PCn. So four different species were detected in stem and root:PC3-Cd(Cu,Zn), PC2-Cd(Cu,Zn), GSH-Cd(Cu,Zn) and Cys-Cd(Cu,Zn). Moreover, results indicated the existence of complexation competition for GSH and cysteine between Cd, Cu and Zn.
6. Imaging of trace metal distribution in tissue sections by laser ablationinductively coupled plasma-mass spectrometry (LA-ICP-MS) is typically performed using spatial resolutions of25μm, including Cd, P,S, K,Ga,Gu-and Zn elements. The-results showed that Cd was preferentially distributed in vascular tissue. The high relationship between Ca and Cd distribution indicating a very similar sequestration of the two elements within Indian Mustard.
7. Method based on ICP-MS was used for subcellular distribution of cadmium in extracts of Indian mustard(Brassica Juncea). Brassica Juncea were submitted to0,0.5,1.0,3.0,5.0,10.0mg L-1and harvested after1,5,7,10and14days exposure. Cells were separated into three fractions:cell wall, soluble fraction and organelle containing fraction using differential centrifugation technique. Results indicated that higher subcellular cadmium in both leaf and root samples was found as exposure level and exposure time increase, but Cadmium concentration differs slightly under1.0mg/L exposure. Results of TEM under different exposure also demonstrate it. Subcellular fractionation of Cd-containing tissues indicated that about50-62%of the element was localized in cell walls and22-32%in soluble fraction, and the lowest in cellular organelles. It could be suggested that subcellular compartment played an important role in Cd detoxification.
本文在综述超积累植物对镉的耐性机制及其相关研究方法的基础上,选取镉富集植物印度芥菜为研究对象,重点开展镉的形态分析与微区分布方法研究,探讨植物螫合肽和区隔化在超积累植物对镉的耐受机制中的作用。综合运用高效液相色谱—电感耦合等离子体质谱(HPLC-ICP-MS)、超高效液相色谱—电喷雾质谱(UPLC-ESI-MS)和超高效液相色谱—飞行时间质谱(UPLC-QTOFMS)联用技术,建立了镉形态分析方法,重点解决了植物螫合肽(PCn)-Cd的定性分析难题,探讨植物螯合肽的形成机制及在植物体耐受与积累重金属中的作用。同时,借助激光烧蚀-电感耦合等离子体质谱(LA-ICP-MS)和透射电镜等手段,分析镉及其它六种元素在印度芥菜体内组织水平以及细胞水平上的分布特征,探讨各类组织/细胞对重金属的区隔化在植物体耐受耐镉机制中的作用。
本研究取得的主要研究结果如下：
1.通过温室水培试验研究了Cd胁迫时间和胁迫水平对印度芥菜生物量的影响以及Cd在植物体内的分布规律,从生物响应的角度探讨了Cd对印度芥菜的毒性及印度芥菜对Cd的耐性。印度芥菜对Cd的最大耐受量为1.0mg/L,超过此浓度植物的正常生长将受到影响,导致生物量下降。植物受胁迫48小时后出现中毒症状,生物量开始下降。随着胁迫时间和胁迫浓度增加,根、叶、茎中镉的浓度均显著增加,根部镉浓度远远大于茎和叶,茎大于叶。在考虑Cd对植物的危害时,胁迫时间和胁迫浓度两个因素要同时考虑。
2.利用UPLC-ESMSMS建立了PCn的分析方法,探讨了PCn稳定性及在耐镉机制中的作用,在Cd的强力诱导下,胁迫合成大量的PCn,从而能螯合更多的Cd,产生较强的耐Cd性。建立了PCn-Cd的UPLC-QTOFMS分析方法,检测出(PC2～PC5)-Cd形态。两种质谱联用技术为PCn-Cd的精确测定提供了强有力的分析工具,解决了由于Cd形态标准物质缺乏造成的SEC-HPLC-ICPMS定性难问题,也为研究PCn与其它金属络合物的分析方法提供了可能。
3.建立了SEC-HPLC-ICPMS测定Cd形态分析方法,在叶片和根部均检测到植物螯合肽(PC)3-Cd、植物螯合肽(PC)2-Cd、谷胱甘肽(GSH)-Cd,及半胱胺酸-Cd四种形态。在植物不同部位Cd存在形态不同,叶片中主要以GSH-Cd存在,而在根部主要以PC2-Cd为主。Cd形态极不稳定,样品处理采取液氮保护并-70℃保存,样品分析全流程用氮吹防氧化措施。研究结果证明,植物螯合肽的合成机制有两种,一是受Cd胁迫直接产生PCn,二是先形成GSH-Cd而后转化成PCn。
4.研究了胁迫浓度和胁迫时间对Cd形态分布的影响。可以推论：植物受Cd胁迫后,先储存在根部。随着胁迫浓度的增大,为了减轻Cd的毒害,先后形成GSH-Cd,PC2-Cd, PC3-Cd,并向叶部转移。同时,叶部受Cd胁迫产生的PCn也会传输到根部,结合更多的Cd,形成PCn-Cd,继续向叶部转移,从而使植物产生很强的Cd耐受性。另外,GSH与PCn结合Cd的能力也存在竞争。随着胁迫时间的增加,根和中GSH-Cd、PC2-Cd、PC3-Cd浓度先迅速增加,而后逐渐降低,96h降至最低。表明在植物耐镉机制中PCn起部分作用,受高浓度Cd胁迫时,其他机制可能会被激发。
5.建立了SEC-HPLC-ICP-MS同时测定印度芥菜中Cd、Cu、Zn形态分析的方法,在植物叶片和根部同时检测出三种元素的四种形态。在同一植物的不同部位,叶片中主要以GSH-Cd、Cys-Cu、 Cys-Zn为主,根部以PC2-Cd、PC2-Cu、Cys-Cu和Cys-Zn为主。另外,Cd、Cu、Zn在与谷胱甘肽(GSH)和半胱胺酸(Cys)的结合上有竞争。研究结论为一种植物同时修复Cd、Cu、Zn复合污染提供了可能。
6.借助LA-ICP-MS质谱技术对富集型印度芥菜茎、叶、叶柄中Cd、P、3、K、Ca、Cu和Zn七种元素的分布特征进行原位分析。结果表明,在茎、叶柄以及主叶脉等组织中,Cd优先分布于含木质部与韧皮部的维管组织中,表明Cd在运输过程中,不易向其他贮存组织扩散。在植物叶片中,Cd主要分布于含有大型液泡的叶肉细胞中,而在上下表皮中分布较低,表明液泡可能是植物存储镉的主要场所。其他元素与Cd的相关性分析结果表Cd与Ca具有相似的分布规律,可能与二者离子半径相近、化学性质相似有关。而K和Ca、P和S的分布呈显著正相关。说明重金属元素进入植株体内并被其吸收运输过程是伴随着植物对其他元素的吸收,且具有相似的运输机制。
7.扫描电镜结果证实胁迫浓度超过lmg/L后,细胞结构发生了变化。同时,细胞壁中镉浓度大于细胞液且远远大于细胞器,证明细胞区隔化在镉的解毒机制中具有重要的作用。
Cadmium (Cd) pollution in the soil is one of the most significant global environmental problems, as it will enter into human bodies through the accumulation of the food chain, and endangers the health of human being. Hyperaccumulator and accumulator species, which efficiently tolerate and accumulate heavy metals from the soil into shoots, have great promise in phytoremediation of contaminated environments. Understanding the mechanisms of metal tolerance and accumulation will provide insight into the identification and management of these hyperaccumulating species. Because of the analytical limit, there has not yet a common accepted conclusion on the migration mechanism of the heavy metal within the plant, as well as the detoxification mechanisms. Phytochelatins (PCn), characterized by the amino acid structure (y-Glu-Cys)n-Gly, where n ranges from2to11, are synthesized enzymatically by phytochelatin synthase (PCn) using glutathione as a substrate in the presence of many metals and metalloids.They are believed to play an important role in metal detoxification and tolerance. But few research on speciation analysis of PCn-Cd and its unstability on detoxification mechanism has been studied.
Therefore, in order to reveal migration and transformation pattern of heavy metals and investigate the relationship between speciation and tolerance of heavy metals, some studies on Indian Mustard which was demonstrated to tolerate and accumulate considerable amounts of cadmium were carried out. The aim was to establish methods for speciation and distribution analysis of cadmium, and obtain fundamental information on heavy metals accumulation and tolerance mechanisms by investigation of the metal localization and ligand abundance. Several modern hyphenated mass spectrometric techniques, such as high performance liquid chromatography-inductively coupled plasma mass spectrometry (HPLC-ICP-MS), ultra performance liquid chromatography-electrospray ilonization mass spectrometry (UPLC-ESI-MS), ultra performance liquid chromatography-time of flight mass spectrometry (UPLC-QTOFMS) and laser ablation inductively coupled plasma mass spectrometry (LA-ICP-MS) were used.
The main results include:
1. Exposure concentration of Cd above1.0mg/L will bring negative effects on the normal growth of the plant, and therefore result in the decrease of the biomass. Within48hours after the stimulation, the symptoms of intoxication can be observed in the plant. With the increase of the exposure time and concentration, the concentrations of cadmium in root, leaf and stem also increases. The concentration of cadmium in the root is far more than that in stem and leave, and and the lowest in in leave.
2. In this study, in vitro formed PCn-Cd complexes were characterized using UPLC-ESI-MSMS and UPLC-QTOFMS. These two methods have been demonstrated to be ideal and promising techniques for screening and characterizing peptide-metal complexes. It was demonstrated that both techniques had the ability to identify the formation of PCn-Cd complex.Furthermore, the increase of PC after Cd exposure demonstrated that Cd is the most efficient inducer of phytochelatin synthesis.
3. Method based on the coupling of size-exclusion HPLC with ICP-MS was used for speciation analysis of cadmium binding non-protein thiols in extracts of Indian mustard (Brassica Juncea). In the presence of Cd stress, PCn were induced, and Cd ions in extracts were associated with the induced PCn. So four different Cd species were detected in samples:PC3-Cd、PC2-Cd、GSH-Cd and Cys-Cd, which demonstrated that PCn were enzymatically synthesised from glutathione (GSH, Glu-Cys-Gly) by the constitutive enzyme PC synthase. The results indicated that GSH-Cd exists as the main specie in leaf samples while PC2-Cd is the main specie in root samples. A non-oxidizing (Nitrogen) preparation environment and a-70℃preservation condition were employed to avoid the oxidation of sulfhydryl groups.
4. Root and leaf samples were submitted to0(control),2.0,4.0,6.0,8.0,10.0mg L-1and harvested after1,2,3and4days exposure. Results indicated lower Cd stress levels were generally associated with higer PCn-Cd production. Higher stress induced the reduction of PCn-Cd, which proved that thiol based tolerance was only efficient in lower degrees of stress. Increasing exposure time also induced decreased PCn-Cd pordution indicated PCn may only had a partial role in metal resistance. Under continuous higher stress, plants maybe trigger other mechnisms to tolerate heavy metal toxicity. Our results suggested the concentration and time of exposure are important factors that must be taken into consideration when evaluating the true role of PCn in heavy metal detoxification.
5. Method based on the coupling of size-exclusion HPLC with ICP-MS was used for speciation analysis of Cd, Cu and Zn binding non-protein thiols in extracts of Indian mustard(Brassica Juncea). In the presence of Cd, Cu and Zn stress, PCn were induced, and metal ions in extracts were associated with the induced PCn. So four different species were detected in stem and root:PC3-Cd(Cu,Zn), PC2-Cd(Cu,Zn), GSH-Cd(Cu,Zn) and Cys-Cd(Cu,Zn). Moreover, results indicated the existence of complexation competition for GSH and cysteine between Cd, Cu and Zn.
6. Imaging of trace metal distribution in tissue sections by laser ablationinductively coupled plasma-mass spectrometry (LA-ICP-MS) is typically performed using spatial resolutions of25μm, including Cd, P,S, K,Ga,Gu-and Zn elements. The-results showed that Cd was preferentially distributed in vascular tissue. The high relationship between Ca and Cd distribution indicating a very similar sequestration of the two elements within Indian Mustard.
7. Method based on ICP-MS was used for subcellular distribution of cadmium in extracts of Indian mustard(Brassica Juncea). Brassica Juncea were submitted to0,0.5,1.0,3.0,5.0,10.0mg L-1and harvested after1,5,7,10and14days exposure. Cells were separated into three fractions:cell wall, soluble fraction and organelle containing fraction using differential centrifugation technique. Results indicated that higher subcellular cadmium in both leaf and root samples was found as exposure level and exposure time increase, but Cadmium concentration differs slightly under1.0mg/L exposure. Results of TEM under different exposure also demonstrate it. Subcellular fractionation of Cd-containing tissues indicated that about50-62%of the element was localized in cell walls and22-32%in soluble fraction, and the lowest in cellular organelles. It could be suggested that subcellular compartment played an important role in Cd detoxification.
引文
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