摘要
铜是世界范围内广泛分布的重金属之一,近年来随着工农业的发展和三废的排放,使得环境中的铜含量迅速增加,造成土壤环境中的铜污染日益严重。当植物生长在铜污染的区域中,植物细胞壁不仅仅起到隔绝铜污染的土壤环境与植物内部细胞的屏障作用,而且在植物对重金属铜的防御反应中发挥至关重要的作用。然而,关于植物细胞壁中的铜的积累、耐性、解毒及信号转导的细胞学机理有待进一步的研究。海州香薷(Elsholtzia splendens)是一种高铜耐性植物,其体内铜的积累量高达1000mg/kg的铜,可生存在高铜污染的区域中,是一种铜污染土壤修复的优选植物。因此,本文以海州香薷的细胞壁为研究对象,从亚细胞、蛋白质组学的水平上阐释了海州香薷细胞壁对铜吸收累积与耐性解毒的分子机理,并构建了细胞壁中压力胁迫的蛋白信号通路。本研究为细胞壁中的压力胁迫应答机制研究提供新的视野,并为土壤铜污染的植物修复提供理论依据和科学支撑。主要研究成果如下：
1、采用分级离心法分离细胞不同组份,明确了不同浓度铜胁迫下的铜离子在海州香薷根、茎、叶中亚细胞分布特征及其化学结合形态。细胞壁是Cu的主要积累位点(68%),其次为液泡(26.6%),只有少量的Cu分布在叶绿体等细胞器中。随着铜离子浓度的增加,亚细胞各个组分中的铜含量也相应的增加。铜离子在海州香薷体内尤其是根系中的形态分布和特征与其对铜离子的耐性有着密切的联系。铜离子在海州香薷各组织中结合的化学形态是不同的,在根中,铜离子主要以水溶性有机酸盐结合态铜或难溶性磷酸盐结合态铜的形式存在；在茎和叶中铜离子以难溶性磷酸盐结合态铜占绝对优势,其次为无机盐、蛋白结合态铜为主的形态存在。随着铜胁迫浓度的增加,铜离子由活动性较强的可溶性盐类,蛋白结合态铜向难溶的磷酸盐及草酸盐结合态转化。铜离子与低活性的磷酸盐及草酸盐结合被固定是海州香薷铜耐性和解毒的重要机制。聚类分析证实了铜离子在不同的亚细胞器官中具有不同的化学结合形态,各化学形态和亚细胞分布于铜的迁移能力及生物活性密切相关。
2、利用细胞壁化学改性结合吸附动力学试验和红外光谱学研究了海州香薷根细胞壁及其组份对铜离子的吸附动力学特征,以及它们吸附固定铜离子的功能基团。吸附动力学表明,海州香薷根细胞壁对Cu2+具有较高的亲和力,吸附300min后,根细胞壁的铜吸附量已接近于饱和水平,吸附量可以达到最大吸附量的90%左右,500mmin达到吸附饱和,饱和吸附量为5.85mg/g细胞壁。当细胞壁进行化学改性后,如果胶甲基化程度降低,及去除细胞壁中的纤维素后显著降低根细胞壁对铜的吸附量。红外光谱研究也表明,在海州香薷根细胞壁吸附Cu2+的过程中,羟基、羧基和氨基是Cu2+的主要结合位点。其中纤维素和半纤维素为Cu2+的结合提供了羧基官能团,果胶提供了羟基官能团,而细胞壁蛋白提供了氨基官能团等结合位点。细胞壁中果胶和纤维素是铜离子主要吸附结合部位,分别吸附了19.85%和25.48%的铜。此外,蛋白质与果胶,果胶和半纤维素之间存在协同作用共同吸附固定铜。根细胞壁及其各组份对铜离子具有较高的吸附固定能力是海州香薷根系Cu耐性的重要机制。
3、分析了细胞壁多糖的组成、含量和分布特征与海州香薷铜耐性之间的关系,测定了三种典型的细胞壁多糖指标(总糖、半乳糖醛酸及3-脱氧-D-甘露-2-辛酮糖酸)。细胞壁多糖含量变化与细胞解毒过程密切相关。海州香薷通过改变细胞壁组份如(果胶,半纤维素,纤维素)的含量增加对铜的耐受性。多糖为铜离子提供了较多的羧基结合位点,是细胞壁多糖解毒的重要机制。随着铜离子胁迫的增加,细胞壁多糖组分的含量发生变化。其中根,叶中总果胶,半纤维素总糖含量极显著增加(P<0.01),而茎中其含量显著减少,在根、茎、叶中纤维素总糖含量均显著降低(P<0.05)；而根、茎、叶中果胶及纤维素中的半乳糖醛酸含量略微增加,根、叶中半纤维素的半乳糖醛酸含量降低,同时,茎中半纤维素的半乳糖醛酸含量增加。根、茎、叶细胞壁果胶的KDO含量随着铜胁迫的增加均略微增加。植物通过这种行为把大量的铜聚集在细胞壁上,避免了对内部细胞器的胁迫,这也是海州香薷对铜离子胁迫解毒的一种生理响应,同时也揭示了细胞壁特别是细胞壁多糖在海州香薷对铜解毒方面发挥着重要的作用。
4、分离海州香薷细胞壁,优化细胞壁蛋白的提取方法,探究海州香薷细胞壁蛋白含量与细胞壁中铜积累量之间的相关性分析。结果显示：在根细胞壁中随着铜离子胁迫浓度的增加,根细胞壁中铜的积累量也是逐步增加的。当铜离子浓度为100μm0l/L时,细胞壁中的铜离子浓度达到最大值1.41mg/g细胞壁干重。根细胞壁蛋白随着铜胁迫浓度的升高先增加后减少,在50μmol/L铜胁迫下,根细胞壁蛋白含量达到最高值,26.85μg/g细胞壁干重。铜在根细胞壁中积累引起细胞壁对铜的一系列响应,包括基因的调控等,从而引起细胞壁蛋白含量的波动性变化。细胞壁蛋白含量变化的波动趋势与细胞壁铜积累量的波动趋势相同,由此说明细胞壁蛋白含量的变化与海州香薷耐铜性的表现趋于一致。在SDS-PAGE蛋白谱图中,根细胞壁蛋白有超过25个蛋白谱带被分离,大多落在标准分子量为10-170kD范围内,在50μmol/L铜胁迫下,根细胞壁蛋白在10-25kD范围内与对照相比条带明显增多。
5.iTRAQ技术结合LC-ESI-MS质谱技术分离鉴定了在50μmol/L铜胁迫48h后海州香薷根细胞壁蛋白表达情况。共479个表达蛋白被分离鉴定,其中55个蛋白显著差异表达,其中22个蛋白上调,33个蛋白下调。对根细胞壁差异表达蛋白进行GO分析,探究这些蛋白参与的生物学过程及分子功能,研究表明：23.24%参与了代谢过程,16.20%参与了细胞学过程及14.79%参与了应激反应；41.38%为结合蛋白,31.03%为催化活性蛋白及11.49%为转运蛋白。由此可见细胞壁差异表达蛋白具有多种分子功能,并参与了多种生物学过程。上调蛋白主要参与了抗氧化胁迫、细胞壁多糖修改及修饰及一些代谢过程,下调蛋白主要参与了信号转导、能量及蛋白合成等代谢过程。细胞壁蛋白是外界环境胁迫和细胞壁骨架间交换信息的媒介和桥梁,丰富的细胞壁内部应激相关信号通路及各种代谢过程相互作用共同构建了一个复杂的应激反应网络抵抗铜离子的胁迫。
Copper is a widespread heavy metal in the world, as a consequence of industrial and agricultural activities in recent years, which are released to the environment. The accumulation of Cu within soils is becoming a major environmental problem. When plant grew on the copper-contaminated areas, cell wall not just provided a physical barrier between copper-contaminated soils and the internal contents of plant cells, also played a crucial role in plant defense response to toxic metals. However, the cytological mechanism in cell wall of Cu uptake, accumulation, detoxification, tolerance and signals transduction is needed for further research. Elsholtzia splendens as a copper-tolerant plant can survive extremes in a high copper content area, which may serve as a good candidate for phytoremediation of copper-contaminated soils. It can accumulate over1000mg/kg of Cu in the body. In this study, we elucidated the mechanisms of Cu uptake, accumulation, tolerance and detoxification in E. splendens cell wall at the cytological and proteome levels for the first time and proposed a possible protein interaction network in response to copper stress of cell wall. Our research would give new insights into stress response in cell wall and could provide theoretical and scientific basis for phytoremediation of copper contaminated soil. The main results were summarized as follows:
1. Using of the grading centrifugation method, it was found that the subcellular distribution and the chemical form of copper ions in roots, stems and leaves under different copper concentration. The highest proportion of Cu was stored in the cell walls (68%) and vacuoles (26.6%), only a small amount of Cu was stored in cell organelles such as chloroplasts. With the increasing of Cu concentration, subcellular copper content of each component also increased. The distribution characteristic of Cu especially in roots was closely related to its tolerance. Meanwhile, Cu taken up by E. splendens existed in different chemical forms. In roots, the majority of Cu was in the forms of water-soluble and un-dissolved phosphate, while most of Cu was in the forms of un-dissolved phosphate, inorganic salts and proteins in stems and leaves. In addition, increasing Cu supply, the binding capacity of Cu to soluble salt or protein reduced but that to insoluble oxalate or un-dissolved phosphate increased. Cu ions integrated with low bio-available compounds, such as un-dissolved phosphate or oxalate which contributed to the metal tolerance and detoxification of E. splendens. Cluster analysis, based on copper abundance levels revealed that the correlations with the chemical forms and intracellular location which indicated the relationship between their migration activity and the toxicity.
2. Chemical modification of cell wall, adsorption dynamic experiments and the FTIR spectroscopy analysis were combined to explore adsorption kinetic characteristics and the functional groups of the cell wall and its components under copper stress. The adsorption dynamic experiments showed that the copper content adsorbed by the root cell wall was close to its saturation level after300minutes, which was about90%of the maximum adsorption capacity. We obtained a saturated adsorption capacity of copper after500minutes, which was5.85mg Cu/g cell wall. The proportion of copper adsorption by pectin and cellulose in the root cell wall were19.85%and25.48%, respectively. The copper adsorption by the root cell wall had been significantly reduced by the ammonia and cellulose treatments on the cell wall. The FTIR spectra also revealed that hydroxyl, carboxyl and amino group are the main binding sites of Cu2+by the root cell wall in the adsorption process. Cellulose and hemicellulose provide a combination of Cu2+with carboxyl functional groups, and pectin offers carboxyl groups while the cell wall proteins provide amino functional groups for binding sites. The results showed that the root cell wall and its components had high adsorption capacity for copper, which was an important mechanism of high copper tolerance by E. splendens.
3. The content, composition and distribution of cell wall polysaccharide of E. splendens and their relationship to copper tolerance were analyzied. Three typical biochemical indexes (sugar, galacturonic acid and Keto-deoxyoctulosonate) of cell wall polysaccharides were investigated. The results illustrated that the content of them was closely related to the toxicity degree. Changes of cell walls polysaccharide in cell walls fraction (such as pectin, cellulose, hemicellulose, etc.) decreased copper toxicity in plant. Expression of cell wall polysaccharides offered more carboxyl groups and provided more binding sites for copper ions. This behavior could be considered as a defense mechanism by E. splendens against heavy metal. With the increasing of Cu supply, various components in cell wall were fluctuated. The total sugar content of hemicellulose and pectin increased very significantly in roots and leaves (P<0.01), whereas that declined perceptibly in stems. The sugar content of cellulose all declined in roots, stems and leaves (P<0.05). The galacturonic acid content of pectin and cellulose increased slightly in roots, stems and leaves, while that of hemicellulose declined in roots, leaves and increased in stems. The Keto-deoxyoctulosonate (KDO) content of pectin was increased slightly by copper in roots, stems and leaves. Plants accumulated lots of copper ions in the cell wall to protect protoplast against copper toxicity, which was a physiological response by E. splendens. It also confirmed that cell wall, especially for cell wall polysaccharides played an important role in copper detoxification.
4. Optimization of the extraction methods of cell wall proteins and determined the relation on correlation of copper accumulation and cell wall proteins content in E. splendens. With the increasing of Cu supple, the copper accumulation in the root cell wall was also increased gradually. The content of copper in cell wall reached its highest level (1.41mg/g cell wall) under100μmol/L Cu. The cell wall proteins content in roots first increased then decreased with the increasing of Cu supply. The content of root cell wall protein reached its highest level (26.85μg/g dry weight of cell wall) on50μmol/L copper stress. Copper accumulation in the root cell wall caused a series of physiological response including gene regulation which presented the fluctuation in the content of cell wall proteins. The same fluctuation trend between the content of cell wall proteins and the copper accumulation indicated that cell wall proteins might participate in the copper tolerance or detoxification by the cell wall. In SDS-PAGE protein spectra, more than25protein bands from the root cell wall proteins were separated, most of them falled in the standard for10-170KD molecular weight range. With50μmol/L Cu stress, the root cell wall proteins within10to25KD increased obviously compared with the contrast stripe.
5. To gain a comprehensive understanding the response to Cu in plant cell wall. iTRAQ with LC-ESI-MS/MS approach were applied to identified differentially expressed cell wall proteins of E. splendens under Cu stress. A total of479proteins were identified, among them55proteins expressed significantly under48h-treatment at50μmol/L Cu. Functional classifications of the55different expressed proteins were classified based on the enrichment in two Gene Ontology (GO) categories:molecular function, and biological process. The top three categories of the identified proteins were those involved in metabolic processes (23.24%), cellular processes (16.20%), and response to stimuli (14.79%), following the top three abundant category was classified according to GO molecular annotation into binding (41.38%), catalytic activity (31.03%), and transport activity (11.49%) respectively. The observed diversity in biological function and biological processes of the identified cell wall proteins demonstrated that the response to Cu stress by the root cell wall of E. splendens was a complex process. Many physiological and biochemical characteristics were altered to counteract the adverse conditions. Among the22up-regulated proteins, most were related to the antioxidant defense pathway, cell wall polysaccharide modification and metabolisem process. However, the most highly down-regulated proteins in response to copper stress belonged to the category of signal, energy and protein synthesis. The communication between cytoskeleton and cell wall protein was one of the most characteristic features of cellular mechanism by which cells sense and respond to various extracellular signals effectively. Based on the abundant changes in these proteins, together with their putative functions, we proposed a possible protein interaction network associated with metal response in cell wall.
引文
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