海州香薷Cu耐性与积累机制研究
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摘要
耐性／超积累植物重金属耐性与积累机制是土壤重金属污染及其修复领域的一项重要基础研究。各国学者针对不同植物种类和不同重金属污染物，从生态学、植物生理学、分子生物学等角度开展了广泛探讨。本文根据前人研究成果，以Cu耐性植物海州香薷为研究对象、重金属Cu为目标污染物，深入分析了海州香薷Cu耐性和积累机制。另外，本文在方法学方面获得一定突破，成功建立了生物软组织LA-ICP-MS定量成像分析法。研究成果主要包括：
明确了海州香薷细胞壁特别是根细胞壁对Cu吸附固定的分子机制。海州香薷细胞壁对Cu具有较强的吸附能力，且根细胞壁比地上部细胞壁对Cu具有更高的亲和力，能够吸附更多的Cu。Fischer酯化反应、氨基甲基化反应等表明，细胞壁上羧基、氨基和羟基是Cu的主要结合位点，这些位点存在于纤维素、木质素中，而果胶质则对Cu的吸附贡献不大。同步辐射X射线吸收光谱研究表明，细胞壁结合Cu的分子形态类似于Cu-组氨酸、Cu-草酸和水合态Cu~(2+)。
发现游离组氨酸和半胱氨酸在Cu耐性和积累中的作用。Cu胁迫引起了海州香薷根系中谷氨酸和苏氨酸含量显著下降，是根系游离氨基酸总量的下降的主要原因。但是，Cu胁迫显著提高了海州香薷根、茎和叶中游离组氨酸和半胱氨酸的合成，它们的含量分别与Cu处理浓度和组织中Cu含量成显著正相关关系。特别是茎中组氨酸含量在高浓度Cu胁迫后升高12倍，推测其可能参与了Cu从根系向地上部的转运。组氨酸是海州香薷根、茎、叶中Cu的一个重要配体，参与Cu的螯合解毒。
发现Cu胁迫后海州香薷根系产生的含Cu蛋白。采用SDS-PAGE和双向电泳技术，对Cu胁迫后海州香薷根系蛋白进行分离，发现Cu胁迫使得根系中的12个蛋白点出现或上调，17个蛋白点消失或下调。采用LA-ICP-MS扫描凝胶电泳分离的蛋白点(或条带)，用~(107)Ag~+作为内标，能够清晰表征蛋白点的分布和相对含量；发现Cu胁迫后产生的含Cu蛋白，以及一些含Zn蛋白；其他元素如Mg、K、Mn、P、S等在凝胶上的分布也予以表征。这些Cu含量较高的蛋白可能是对Cu胁迫的响应，参与Cu的螯合，对海州香薷Cu耐性和积累起着积极的作用。
成功建立LA-ICP-MS对生物软组织中元素分布定位的定量成像技术，构建了新的定量化策略，该技术能够清晰表征海州香薷叶片中Cu、K、Mg、Mn、P、S、B等元素的分布定位。通过构建实验室标准参比物质，各元素的分布成像能够实现定量化；另外，采用~(13)C~+作为内标，以校正水分及样品不均一性对离子强度的影响，本研究发展了成像LA-ICP-MS技术的定量化过程，并使之成为生物软组织中元素分布定位定量研究的新手段和新方法。结合高丰度同位素(89.2％~(65)Cu)示踪实验表明，短期Cu处理使得海州香薷新生叶片中Cu迅速积累，而成熟和年老叶片中Cu的积累相对缓慢。~(65)Cu／~(63)Cu同位素比的变化表明，在新叶中，Cu通过叶脉首先输送到底部叶缘和叶尖。Cu处理引起新生叶片中K、Mg、Mn、P和S的升高，而B受到Cu处理的影响并不显著。
One of the most important issues in the field of environment engineering is theremediation of heavy metal-polluted soils. Some plants are known as hyperaccumulatoror tolerant plants which are able to accumulate considerable amount of heavy metalssuch as Ni, Zn, Cd or Cu in their roots or shoots. Therefore, the mechanisms fortolerance and accumulation of heavy metals in these plants are of interest for thescientists in the field of environmental, plant and biological sciences all over the world.From the ecological, plant physiological and molecular biological points of view,respectively, scientists are devoting themselves in revealing the mysteries of the specialabilities in plants.
Elsholtzia splendens is one of these plants, which is reported as a Cu-tolerant plant,usually found in copper mining areas of southern China. The Cu concentration in theleaves can reach 1000 mg kg~(-1) in solution cultivation. The plant is considered to be anappropriate choice for phytoextracting the highly concentrated Cu inmetal-contaminated soils. Corresponding research is being undertaken with respect tothe mechanisms for Cu tolerance and accumulation. In the present study, furtherinvestigation on the Cu tolerance and accumulation was conducted with respect to cellwall precipitation of Cu, ligands involved in Cu chelation and elemental distributionwithin the plant leaves. On the other hand, new quantification strategies on imagingmass spectrometry using laser ablation inductively coupled plasma mass spectrometry(LA-ICP-MS) of essential and toxic elements in biological tissues were developed tostudy the uptake of heavy but also nutrient elements.
It was illustrated that the precipitation of Cu on cell walls was one of the importantmechanisms for Cu tolerance and accumulation. E. splendens had high absorptioncapacity for Cu on the cell walls. More Cu was absorbed on the root cell walls than onthe shoot cell walls. Esterification of carboxyl groups and methylation of amino groupsof the cell walls resulted in reduction in the amount of Cu absorbed by about 64% and25%, respectively. Cu was bound with the carboxyl, amino and hydroxyl groups in thecell walls revealed by Fourier transform infrared spectroscopy. Synchrotron radiationX-ray absorption spectroscopy was used to investigate the Cu species on the cell walls.The percentages of potential Cu ligands were estimated by fitting the XANES spectra with linear combinations of the references'. Cu in cell walls was bound withoxalate-like ligands, histidine-like ligands, as well as aqueous Cu.
Free histidine and cysteine were found to be potential ligands chelating with Cu inE. splendens. Sixteen free amino acids were. analyzed to investigate the correlationsbetween free amino acids and Cu stress. The synthesis of free histidine and cysteine wasinduced by high Cu content in the nutrient solution. The concentrations of both aminoacids were significantly correlated with Cu concentrations in the nutrient solution and intissues of the plant. The two amino acids were involved in the chelation of Cu in theplant. Particularly, the histidine concentration in stems after 500μM Cu treatment waseleven times more than it in the stems without treatment, indicating that free histidineprobably participated in the transport of Cu from the roots to the shoots.
Cu- and Zn-containing proteins in the roots of E. splendens were investigated bymetaliomic studies using 2D gel electrophoresis (2D-PAGE) for protein separation andLA-ICP-MS for screening of metal-containing proteins. Twelve proteins were inducedor up-regulated, while another seventeen proteins were disappeared or down-regulatedby Cu treatment. LA-ICP-MS was an effective technique to imaging metals andnon-metals in protein spots on gels after electrophoresis with silver staining, ~(107)Ag~+ wasused to indicate the location and the relative content of the proteins. SeveralCu-containing proteins were found, as well as Zn-, Mn- or Mg-containing proteins werepresent in the gels. Further studies with respect to protein identification and theirfunctions should be carried out.
New quantification strategies on imaging mass spectrometry using LA-ICP-MS ofessential and toxic elements in biological tissues were developed to study the uptake ofheavy but also nutrient elements. In the present work, it was demonstrated thatLA-ICP-MS is a powerful analytical technique for directly imaging both macro- andmicronutrients in the plant leaves without sample preparation, and the resolution of thistechnique is sufficient for all the elements measured (Cu, K, Mg, Mn, P, S and B). Themethod was able to quantify the analytical data with synthetic laboratory standardcalibration. The established quantitative imaging LA-ICP-MS method will provide aneasy and effective tool in the field of the phytosphere with respect to distribution ofelements, and will open up a new window onto in vivo ionomic imaging of plant tissues.LA-ICP-MS was successfully applied together with tracer experiments using enriched~(65)Cu isotope tracer to investigate the accumulation of Cu and other essential elements of interest (K, Mg, Mn, P, S and B) in the leaves of E. splendens after ~(65)Cu treatment. Cuaccumulation and the effect of Cu stress on essential element uptake in the newlyformed leaves can be clearly observed by quantitative imaging, while no significantchanges were found in the fully grown leaves due to the short-term treatment. The~(65)Cu/~(63)Cu isotope ratios measured by LA-ICP-MS increased after ~(65)Cu treatment. Theresults indicated the path of Cu uptake via the petiole and main veins of the leaves.Together with tracer experiments, quantitative imaging LA-ICP-MS was successfullyapplied for studying the uptake and accumulation of toxicological and essentialelements in plant leaves.
明确了海州香薷细胞壁特别是根细胞壁对Cu吸附固定的分子机制。海州香薷细胞壁对Cu具有较强的吸附能力，且根细胞壁比地上部细胞壁对Cu具有更高的亲和力，能够吸附更多的Cu。Fischer酯化反应、氨基甲基化反应等表明，细胞壁上羧基、氨基和羟基是Cu的主要结合位点，这些位点存在于纤维素、木质素中，而果胶质则对Cu的吸附贡献不大。同步辐射X射线吸收光谱研究表明，细胞壁结合Cu的分子形态类似于Cu-组氨酸、Cu-草酸和水合态Cu~(2+)。
发现游离组氨酸和半胱氨酸在Cu耐性和积累中的作用。Cu胁迫引起了海州香薷根系中谷氨酸和苏氨酸含量显著下降，是根系游离氨基酸总量的下降的主要原因。但是，Cu胁迫显著提高了海州香薷根、茎和叶中游离组氨酸和半胱氨酸的合成，它们的含量分别与Cu处理浓度和组织中Cu含量成显著正相关关系。特别是茎中组氨酸含量在高浓度Cu胁迫后升高12倍，推测其可能参与了Cu从根系向地上部的转运。组氨酸是海州香薷根、茎、叶中Cu的一个重要配体，参与Cu的螯合解毒。
发现Cu胁迫后海州香薷根系产生的含Cu蛋白。采用SDS-PAGE和双向电泳技术，对Cu胁迫后海州香薷根系蛋白进行分离，发现Cu胁迫使得根系中的12个蛋白点出现或上调，17个蛋白点消失或下调。采用LA-ICP-MS扫描凝胶电泳分离的蛋白点(或条带)，用~(107)Ag~+作为内标，能够清晰表征蛋白点的分布和相对含量；发现Cu胁迫后产生的含Cu蛋白，以及一些含Zn蛋白；其他元素如Mg、K、Mn、P、S等在凝胶上的分布也予以表征。这些Cu含量较高的蛋白可能是对Cu胁迫的响应，参与Cu的螯合，对海州香薷Cu耐性和积累起着积极的作用。
成功建立LA-ICP-MS对生物软组织中元素分布定位的定量成像技术，构建了新的定量化策略，该技术能够清晰表征海州香薷叶片中Cu、K、Mg、Mn、P、S、B等元素的分布定位。通过构建实验室标准参比物质，各元素的分布成像能够实现定量化；另外，采用~(13)C~+作为内标，以校正水分及样品不均一性对离子强度的影响，本研究发展了成像LA-ICP-MS技术的定量化过程，并使之成为生物软组织中元素分布定位定量研究的新手段和新方法。结合高丰度同位素(89.2％~(65)Cu)示踪实验表明，短期Cu处理使得海州香薷新生叶片中Cu迅速积累，而成熟和年老叶片中Cu的积累相对缓慢。~(65)Cu／~(63)Cu同位素比的变化表明，在新叶中，Cu通过叶脉首先输送到底部叶缘和叶尖。Cu处理引起新生叶片中K、Mg、Mn、P和S的升高，而B受到Cu处理的影响并不显著。
One of the most important issues in the field of environment engineering is theremediation of heavy metal-polluted soils. Some plants are known as hyperaccumulatoror tolerant plants which are able to accumulate considerable amount of heavy metalssuch as Ni, Zn, Cd or Cu in their roots or shoots. Therefore, the mechanisms fortolerance and accumulation of heavy metals in these plants are of interest for thescientists in the field of environmental, plant and biological sciences all over the world.From the ecological, plant physiological and molecular biological points of view,respectively, scientists are devoting themselves in revealing the mysteries of the specialabilities in plants.
Elsholtzia splendens is one of these plants, which is reported as a Cu-tolerant plant,usually found in copper mining areas of southern China. The Cu concentration in theleaves can reach 1000 mg kg~(-1) in solution cultivation. The plant is considered to be anappropriate choice for phytoextracting the highly concentrated Cu inmetal-contaminated soils. Corresponding research is being undertaken with respect tothe mechanisms for Cu tolerance and accumulation. In the present study, furtherinvestigation on the Cu tolerance and accumulation was conducted with respect to cellwall precipitation of Cu, ligands involved in Cu chelation and elemental distributionwithin the plant leaves. On the other hand, new quantification strategies on imagingmass spectrometry using laser ablation inductively coupled plasma mass spectrometry(LA-ICP-MS) of essential and toxic elements in biological tissues were developed tostudy the uptake of heavy but also nutrient elements.
It was illustrated that the precipitation of Cu on cell walls was one of the importantmechanisms for Cu tolerance and accumulation. E. splendens had high absorptioncapacity for Cu on the cell walls. More Cu was absorbed on the root cell walls than onthe shoot cell walls. Esterification of carboxyl groups and methylation of amino groupsof the cell walls resulted in reduction in the amount of Cu absorbed by about 64% and25%, respectively. Cu was bound with the carboxyl, amino and hydroxyl groups in thecell walls revealed by Fourier transform infrared spectroscopy. Synchrotron radiationX-ray absorption spectroscopy was used to investigate the Cu species on the cell walls.The percentages of potential Cu ligands were estimated by fitting the XANES spectra with linear combinations of the references'. Cu in cell walls was bound withoxalate-like ligands, histidine-like ligands, as well as aqueous Cu.
Free histidine and cysteine were found to be potential ligands chelating with Cu inE. splendens. Sixteen free amino acids were. analyzed to investigate the correlationsbetween free amino acids and Cu stress. The synthesis of free histidine and cysteine wasinduced by high Cu content in the nutrient solution. The concentrations of both aminoacids were significantly correlated with Cu concentrations in the nutrient solution and intissues of the plant. The two amino acids were involved in the chelation of Cu in theplant. Particularly, the histidine concentration in stems after 500μM Cu treatment waseleven times more than it in the stems without treatment, indicating that free histidineprobably participated in the transport of Cu from the roots to the shoots.
Cu- and Zn-containing proteins in the roots of E. splendens were investigated bymetaliomic studies using 2D gel electrophoresis (2D-PAGE) for protein separation andLA-ICP-MS for screening of metal-containing proteins. Twelve proteins were inducedor up-regulated, while another seventeen proteins were disappeared or down-regulatedby Cu treatment. LA-ICP-MS was an effective technique to imaging metals andnon-metals in protein spots on gels after electrophoresis with silver staining, ~(107)Ag~+ wasused to indicate the location and the relative content of the proteins. SeveralCu-containing proteins were found, as well as Zn-, Mn- or Mg-containing proteins werepresent in the gels. Further studies with respect to protein identification and theirfunctions should be carried out.
New quantification strategies on imaging mass spectrometry using LA-ICP-MS ofessential and toxic elements in biological tissues were developed to study the uptake ofheavy but also nutrient elements. In the present work, it was demonstrated thatLA-ICP-MS is a powerful analytical technique for directly imaging both macro- andmicronutrients in the plant leaves without sample preparation, and the resolution of thistechnique is sufficient for all the elements measured (Cu, K, Mg, Mn, P, S and B). Themethod was able to quantify the analytical data with synthetic laboratory standardcalibration. The established quantitative imaging LA-ICP-MS method will provide aneasy and effective tool in the field of the phytosphere with respect to distribution ofelements, and will open up a new window onto in vivo ionomic imaging of plant tissues.LA-ICP-MS was successfully applied together with tracer experiments using enriched~(65)Cu isotope tracer to investigate the accumulation of Cu and other essential elements of interest (K, Mg, Mn, P, S and B) in the leaves of E. splendens after ~(65)Cu treatment. Cuaccumulation and the effect of Cu stress on essential element uptake in the newlyformed leaves can be clearly observed by quantitative imaging, while no significantchanges were found in the fully grown leaves due to the short-term treatment. The~(65)Cu/~(63)Cu isotope ratios measured by LA-ICP-MS increased after ~(65)Cu treatment. Theresults indicated the path of Cu uptake via the petiole and main veins of the leaves.Together with tracer experiments, quantitative imaging LA-ICP-MS was successfullyapplied for studying the uptake and accumulation of toxicological and essentialelements in plant leaves.
引文
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