超积累植物美洲商陆锰累积与耐性机制研究
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摘要
美洲商陆(Phytolacca americana Linn.)是近年来颇受关注的锰超积累植物,然其锰耐性机制仍不清楚。本论文通过实验室人工营养液培养体系,采用电镜结合能谱、超速离心亚细胞组分分离、化学形态提取、高效液相色谱、晶体X射线衍射等方法观察不同锰处理下美洲商陆表观胁迫响应,检测植物对锰的吸收与累积以及锰在植物组织细胞中的分布,分析美洲商陆锰耐受可能涉及因素与锰累积与解毒的关系,综合研究美洲商陆锰耐性的可能作用机制。当前研究进一步促进了植物对锰耐性机理的理解,为实际的锰毒土壤上的作物生产改良、酸性土壤或相关重金属污染土壤的植物修复等提供一定的理论支持与科学依据。取得的主要研究结果与结论如下:
不同锰处理培养观察显示,美洲商陆对锰具有极强的耐性。5000μM以下锰处理,美洲商陆生长良好,无明显锰毒性症状,而更高浓度锰处理,植物地上部,特别是叶片呈现出典型的锰中毒特征。光合作用检测表明,美洲商陆并非嗜锰植物,过量锰不能促进植物代谢与生长。较高浓度锰连续胁迫下,美洲商陆根部依锰处理浓度高低先后析出含锰晶体,表明美洲商陆对锰存在一定的排斥机制,可能与根际沉淀作用相关。电镜观察发现,锰对美洲商陆细胞毒性作用较弱。扫描电镜下美洲商陆细胞内观察到两类晶体:草酸钙异胞体与含锰晶体。叶片含锰晶体能谱扫描显示,锰可能主要与羟基、羧基、磷酸根等含氧基团结合。检测美洲商陆可能涉及锰耐性的相关作用物质,发现该植物草酸与巯基含量较高。初步研究结果表明,美洲商陆锰耐性可能与草酸、钙、巯基及磷酸根等因素密切相关。
美洲商陆可以耐受和累积高浓度的锰,将过量的锰累积于叶细胞可溶性组分内,而在细胞器与细胞膜组分的过量累积则导致锰植物毒性。液泡成分包含于可溶性组分内,作为锰累积的关键场所,同时也是植物储存草酸的细胞器。不同锰处理植物,美洲商陆可溶性组分内总可以提供过量的草酸络合过量累积的锰,使之转化为无植物毒性的化学形态。XRD检测证实可溶性组分中锰以草酸盐形式存在。研究表明,锰的液泡区室化作用及草酸的络合作用很可能是美洲商陆锰超积累与耐受最重要的生物机制。
比较美洲商陆钙和锰的累积发现,钙和锰主要都以草酸盐形式储存于植物叶片中。其中,钙对锰的累积具有非常显著的影响,但美洲商陆叶片中草酸钙晶体并未发现对锰的解毒起到作用。在美洲商陆叶中含锰晶体的发现揭示锰可能被草酸盐络合,也或与磷酸根结合。美洲商陆钙和锰以如此相近的形式累积,提示参考植物对钙的调控机制的认识,深入比较研究锰的吸收、转运与累积,将有助于进一步理解植物对锰动态平衡的维持与植物对锰的解毒机制。
较高浓度锰连续处理美洲商陆,植物根部随着锰处理浓度的高低先后析出磷酸锰晶体。含磷与缺磷实验发现,磷酸盐可促进美洲商陆锰的累积并明显减缓锰的毒性效应。电镜与能谱检测发现,含磷锰处理的美洲商陆叶与根内存在锰的磷酸盐晶体,而缺磷处理组植物组织中没有出现。特别是美洲商陆根的皮层与表皮,存在明显的锰磷酸盐晶体。当前研究结果表明,磷酸盐在美洲商陆锰的超积累与耐性中扮演着重要角色:一方面,磷酸根在植物体内通过固定作用将锰沉积,降低锰毒性;另一方面,磷酸根可通过根际沉淀作用将锰以磷酸锰晶体形式累积于根表或表层细胞内,从而阻滞植物对锰的进一步吸收。
研究美洲商陆对不同硫锰作用下的植物锰的累积与总巯基类物质(TSH)含量变化情况,以及分析两者之间对美洲商陆相互作用的关系,结果表明,不同的硫锰处理对美洲商陆TSH含量以及植物锰耐性与累积均具有一定影响。其中,美洲商陆经不同的锰硫处理,其根组织锰的累积量与TSH含量呈现良好的正相关关系。研究结果表明,根中巯基类物质对于美洲商陆锰的耐性与超累积具有重要作用。鉴于美洲商陆极强的生长适应性以及对多种重金属耐受与超积累的特性,根中巯基类物质可能发挥着关键作用,为植物的正常生长和免受外界金属离子的损伤建立重要的保护机制。
Pokeweed (Phytolacca americana Linn.) has drawn much attention for its hyperaccumulation of manganese (Mn) in recent years. However, the mechanisms of its Mn resistance are still not well clarified. The present study carried out series of experiments, including hydroponic culture, photosynthesis detection, scanning electronic microscopy (SEM) and transmission electronic microscopy (TEM), energy dispersive analysis of X-rays (EDX), atom absorption spectrometer (AAS), subcellular fractioning, high performance liquid chromatography (HPLC), extraction of different chemical forms, and phase analysis of X-ray diffraction (XRD), to investigate the responses of pokeweeds treated with different levels of Mn, detect Mn uptake, accumulation and subcellular distribution in pokeweed, analyze the correlation between the possible factors involved with Mn resistance and Mn accumulation and detoxification, and further explore the potential mechanisms of Mn resistance in the plant. This research could advance our understanding of Mn resistance in plant, and provide theoretical evidence and scientific directions for the correction of Mn toxicity to crops and phytoremediation of acid soils or relative heavy metal-contaminated soils. The major results obtained are as follows.
Pokeweed showed great tolerance to Mn, and it grew well with no discernable Mn phytotoxicity under different Mn treatment. Detection of its photosynthesis found that pokeweed is not a Mn-philic plant, and that Mn can not enhance its metabolism and growth. Mn-containing crystals were found to appear time sequently on the roots of pokeweeds treated with different concentrations of Mn, which suggested that pokeweed could exclude Mn through rhizodeposition. Observation with electronic microscopy found that Mn had minor toxicity to pokeweed cells in which, meanwhile, two kinds of crystals were found, calcium oxalate crystals and Mn-containing crystals. EDX detection showed Mn could be complexed with hydroxyl, carboxyl or phosphate groups. Pokeweed was also found to have a high content of oxalic acid and thiols. The primary study indicated that Mn resistance in pokeweed could be closely related to factors, including oxalic acid, calcium, thiols and phosphate.
The plant was found to accumulate excess Mn in the leaves, mainly in the water soluble fraction and over 80% existing in a water-soluble chemical form, while accumulating excess Mn in the fraction of cellular organelles and membrane was observed to cause phytotoxicity. In addition, pokeweed was found with intrinsically high oxalate content. In all cases, there were sufficient levels of oxalate to chelate Mn in leaf water extracts throughout different levels of Mn. Phase analysis of X-ray diffraction detected the oxalate-Mn chelate complexes in the freeze-dried soluble fraction of pokeweed leaves. The results suggest that pokeweed accumulates excess Mn in the soluble fraction of leaf cells, most likely in vacuoles, in which the detoxification of Mn could be achieved by chelation with oxalate.
Ca has an important impact on divalent metal accumulation in plants, and hence, the accumulation of Ca and Mn and their interaction in pokeweed were investigated. Exogenous Ca was observed to have a distinctive impact on the Mn phytotoxicity and accumulation in pokeweed, but exogenous Mn had little influence on the accumulation of Ca. Both Ca and Mn accumulated in pokeweed were detected to be mainly in the form of oxalate. Investigation with SEM and TEM found there were two kinds of crystals in the leaves, Ca oxalate crystals and Mn-containing crystals. Further detection showed that there was no inclusion of Mn inside the Ca oxalate crystals, and that other elements, such as C, 0 and P, were present in the Mn-containing crystals. These results suggest that Ca oxalate crystals in pokeweed have no direct effect on the detoxification of Mn. In addition, the finding of element P and 0 in the Mn-containing crystals indicates that excess Mn could be deposited by phosphate, which could contribute to Mn accumulation and detoxification in pokeweed.
Mn phosphate crystals were observed to appear on the root surface, and phosphate was investigated to have a distinctive effect on the Mn accumulation and toxicity in pokeweed. The finding of Mn-containing crystals in the leaf and root indicated that Mn could be precipitated by phosphate, and the crystals in the tissue of root cortex, epidermis, root hair and root surface showed that pokeweed could exclude excess Mn by rhizodiposition with phosphate. The results suggest that phosphate plays a very important role in Mn tolerance and exclusion in pokeweed.
Investigation of Mn accumulation and total sulfhydryl (TSH) and their interaction in pokeweeds treated with different levels of Mn and sulphate found that treatment with different concentrations of sulphate and Mn had an impact on the content of TSH, Mn accumulation and tolerance. In particular, Mn accumulation showed a much positive correlation to the content of TSH in the root. The results suggest that sulfhydryl compounds in the root act as an important role in the Mn hyperaccumulation and resistance in pokeweed. Considering the great adaptability and multiple tolerances to heavy metals, thiols in the pokeweed root could play a vital role which protects the plant from the damages from heavy metal ions.
不同锰处理培养观察显示,美洲商陆对锰具有极强的耐性。5000μM以下锰处理,美洲商陆生长良好,无明显锰毒性症状,而更高浓度锰处理,植物地上部,特别是叶片呈现出典型的锰中毒特征。光合作用检测表明,美洲商陆并非嗜锰植物,过量锰不能促进植物代谢与生长。较高浓度锰连续胁迫下,美洲商陆根部依锰处理浓度高低先后析出含锰晶体,表明美洲商陆对锰存在一定的排斥机制,可能与根际沉淀作用相关。电镜观察发现,锰对美洲商陆细胞毒性作用较弱。扫描电镜下美洲商陆细胞内观察到两类晶体:草酸钙异胞体与含锰晶体。叶片含锰晶体能谱扫描显示,锰可能主要与羟基、羧基、磷酸根等含氧基团结合。检测美洲商陆可能涉及锰耐性的相关作用物质,发现该植物草酸与巯基含量较高。初步研究结果表明,美洲商陆锰耐性可能与草酸、钙、巯基及磷酸根等因素密切相关。
美洲商陆可以耐受和累积高浓度的锰,将过量的锰累积于叶细胞可溶性组分内,而在细胞器与细胞膜组分的过量累积则导致锰植物毒性。液泡成分包含于可溶性组分内,作为锰累积的关键场所,同时也是植物储存草酸的细胞器。不同锰处理植物,美洲商陆可溶性组分内总可以提供过量的草酸络合过量累积的锰,使之转化为无植物毒性的化学形态。XRD检测证实可溶性组分中锰以草酸盐形式存在。研究表明,锰的液泡区室化作用及草酸的络合作用很可能是美洲商陆锰超积累与耐受最重要的生物机制。
比较美洲商陆钙和锰的累积发现,钙和锰主要都以草酸盐形式储存于植物叶片中。其中,钙对锰的累积具有非常显著的影响,但美洲商陆叶片中草酸钙晶体并未发现对锰的解毒起到作用。在美洲商陆叶中含锰晶体的发现揭示锰可能被草酸盐络合,也或与磷酸根结合。美洲商陆钙和锰以如此相近的形式累积,提示参考植物对钙的调控机制的认识,深入比较研究锰的吸收、转运与累积,将有助于进一步理解植物对锰动态平衡的维持与植物对锰的解毒机制。
较高浓度锰连续处理美洲商陆,植物根部随着锰处理浓度的高低先后析出磷酸锰晶体。含磷与缺磷实验发现,磷酸盐可促进美洲商陆锰的累积并明显减缓锰的毒性效应。电镜与能谱检测发现,含磷锰处理的美洲商陆叶与根内存在锰的磷酸盐晶体,而缺磷处理组植物组织中没有出现。特别是美洲商陆根的皮层与表皮,存在明显的锰磷酸盐晶体。当前研究结果表明,磷酸盐在美洲商陆锰的超积累与耐性中扮演着重要角色:一方面,磷酸根在植物体内通过固定作用将锰沉积,降低锰毒性;另一方面,磷酸根可通过根际沉淀作用将锰以磷酸锰晶体形式累积于根表或表层细胞内,从而阻滞植物对锰的进一步吸收。
研究美洲商陆对不同硫锰作用下的植物锰的累积与总巯基类物质(TSH)含量变化情况,以及分析两者之间对美洲商陆相互作用的关系,结果表明,不同的硫锰处理对美洲商陆TSH含量以及植物锰耐性与累积均具有一定影响。其中,美洲商陆经不同的锰硫处理,其根组织锰的累积量与TSH含量呈现良好的正相关关系。研究结果表明,根中巯基类物质对于美洲商陆锰的耐性与超累积具有重要作用。鉴于美洲商陆极强的生长适应性以及对多种重金属耐受与超积累的特性,根中巯基类物质可能发挥着关键作用,为植物的正常生长和免受外界金属离子的损伤建立重要的保护机制。
Pokeweed (Phytolacca americana Linn.) has drawn much attention for its hyperaccumulation of manganese (Mn) in recent years. However, the mechanisms of its Mn resistance are still not well clarified. The present study carried out series of experiments, including hydroponic culture, photosynthesis detection, scanning electronic microscopy (SEM) and transmission electronic microscopy (TEM), energy dispersive analysis of X-rays (EDX), atom absorption spectrometer (AAS), subcellular fractioning, high performance liquid chromatography (HPLC), extraction of different chemical forms, and phase analysis of X-ray diffraction (XRD), to investigate the responses of pokeweeds treated with different levels of Mn, detect Mn uptake, accumulation and subcellular distribution in pokeweed, analyze the correlation between the possible factors involved with Mn resistance and Mn accumulation and detoxification, and further explore the potential mechanisms of Mn resistance in the plant. This research could advance our understanding of Mn resistance in plant, and provide theoretical evidence and scientific directions for the correction of Mn toxicity to crops and phytoremediation of acid soils or relative heavy metal-contaminated soils. The major results obtained are as follows.
Pokeweed showed great tolerance to Mn, and it grew well with no discernable Mn phytotoxicity under different Mn treatment. Detection of its photosynthesis found that pokeweed is not a Mn-philic plant, and that Mn can not enhance its metabolism and growth. Mn-containing crystals were found to appear time sequently on the roots of pokeweeds treated with different concentrations of Mn, which suggested that pokeweed could exclude Mn through rhizodeposition. Observation with electronic microscopy found that Mn had minor toxicity to pokeweed cells in which, meanwhile, two kinds of crystals were found, calcium oxalate crystals and Mn-containing crystals. EDX detection showed Mn could be complexed with hydroxyl, carboxyl or phosphate groups. Pokeweed was also found to have a high content of oxalic acid and thiols. The primary study indicated that Mn resistance in pokeweed could be closely related to factors, including oxalic acid, calcium, thiols and phosphate.
The plant was found to accumulate excess Mn in the leaves, mainly in the water soluble fraction and over 80% existing in a water-soluble chemical form, while accumulating excess Mn in the fraction of cellular organelles and membrane was observed to cause phytotoxicity. In addition, pokeweed was found with intrinsically high oxalate content. In all cases, there were sufficient levels of oxalate to chelate Mn in leaf water extracts throughout different levels of Mn. Phase analysis of X-ray diffraction detected the oxalate-Mn chelate complexes in the freeze-dried soluble fraction of pokeweed leaves. The results suggest that pokeweed accumulates excess Mn in the soluble fraction of leaf cells, most likely in vacuoles, in which the detoxification of Mn could be achieved by chelation with oxalate.
Ca has an important impact on divalent metal accumulation in plants, and hence, the accumulation of Ca and Mn and their interaction in pokeweed were investigated. Exogenous Ca was observed to have a distinctive impact on the Mn phytotoxicity and accumulation in pokeweed, but exogenous Mn had little influence on the accumulation of Ca. Both Ca and Mn accumulated in pokeweed were detected to be mainly in the form of oxalate. Investigation with SEM and TEM found there were two kinds of crystals in the leaves, Ca oxalate crystals and Mn-containing crystals. Further detection showed that there was no inclusion of Mn inside the Ca oxalate crystals, and that other elements, such as C, 0 and P, were present in the Mn-containing crystals. These results suggest that Ca oxalate crystals in pokeweed have no direct effect on the detoxification of Mn. In addition, the finding of element P and 0 in the Mn-containing crystals indicates that excess Mn could be deposited by phosphate, which could contribute to Mn accumulation and detoxification in pokeweed.
Mn phosphate crystals were observed to appear on the root surface, and phosphate was investigated to have a distinctive effect on the Mn accumulation and toxicity in pokeweed. The finding of Mn-containing crystals in the leaf and root indicated that Mn could be precipitated by phosphate, and the crystals in the tissue of root cortex, epidermis, root hair and root surface showed that pokeweed could exclude excess Mn by rhizodiposition with phosphate. The results suggest that phosphate plays a very important role in Mn tolerance and exclusion in pokeweed.
Investigation of Mn accumulation and total sulfhydryl (TSH) and their interaction in pokeweeds treated with different levels of Mn and sulphate found that treatment with different concentrations of sulphate and Mn had an impact on the content of TSH, Mn accumulation and tolerance. In particular, Mn accumulation showed a much positive correlation to the content of TSH in the root. The results suggest that sulfhydryl compounds in the root act as an important role in the Mn hyperaccumulation and resistance in pokeweed. Considering the great adaptability and multiple tolerances to heavy metals, thiols in the pokeweed root could play a vital role which protects the plant from the damages from heavy metal ions.
引文
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