铜污染土壤的生物修复研究进展

随着工业化与农业化进程的加快,土壤重金属污染问题日益突出。铜(Cu)既是生命体生长发育的必需微量元素,也是重金属污染物之一。土壤中过量的Cu不仅会对植物产生毒害,而且能够通过食物链的富集作用,对人类健康造成严重威胁。生物修复技术作为治理重金属污染土壤的一种新型技术受到广泛关注。文中对生物修复的主要技术如植物修复、微生物修复、植物-微生物联合修复、动物修复等在治理Cu污染土壤方面的研究进展进行综述,以期为重金属污染土壤有效治理和可持续农业的发展提供理论依据。     

土壤重金属污染的植物修复

土壤重金属污染的危害范围广泛,使用传统的物理和化学修复方法成本高,对环境扰动大,而利用植物修复的效果较为明显,易于操作.本文论述了土壤重金属污染的单一植物、植物与微生物联合、植物与化学方法相结合.的修复方法,着重介绍了重金属超富'集植物的研究和植物体内螯合肽(PCs)的合成.生物螯合剂的应用及土壤重金属污染的动物、植物和微生物的联合修复将是未来研究的热点.     

重金属污染土壤中生物间相互作用及其协同修复应用

土壤是人类赖以生存的物质基础。我国土壤重金属污染状况不容乐观,给人类健康构成严重威胁。生物修复重金属污染土壤被广泛认为是可持续的修复技术,但当前仍存在修复效率不高的科学瓶颈问题。土壤中生活着丰富的微生物、植物和动物,且这些生物之间存在着复杂的相互作用,并且通过物质循环和能量传递形成了错综复杂的食物网联系。土壤生物间的相互作用能深刻影响土壤中污染物的迁移转化和生物修复的效率,多元生物协同的修复技术集合了单一生物修复方法的优势,具有强化生物修复效果的巨大潜力。文中综述了土壤中微生物-植物-动物之间的相互作用,及其对土壤重金属迁移转化和生物修复效果的影响,并对定向调控土壤食物网结构、提高重金属污染土壤的生物修复效果、建立基于食物网的多元生物协同修复技术进行了展望。     

基于文献计量的细菌-生物炭协同修复重金属污染土壤研究进展

为提高重金属污染土壤可持续修复效能,研究生物炭与细菌对重金属污染土壤的协同修复作用。基于文献计量学分析及重金属污染土壤修复背景,总结了细菌与生物炭对土壤重金属的稳定化特征及菌炭间的相互作用,分析了单一生物炭或细菌对重金属污染土壤修复的局限性,强调了细菌-生物炭协同修复技术的优势,阐述了细菌与生物炭主要通过离子交换、固定作用、氧化还原作用和迁移作用等重要机制有效修复重金属污染土壤,揭示了细菌-生物炭协同作用在重金属污染土壤修复中的巨大应用价值。文献计量学研究表明,生物炭与细菌对重金属污染土壤的协同修复已得到广泛关注。目前认为:生物炭与细菌的协同作用可有效改良土壤理化性质及提高土壤修复效率,也可促进植物生长及植物修复进程;生物炭对细菌影响具有双重性质,可促进细菌生长,也可能对细菌产生毒害;细菌可改变生物炭的理化性质,进而强化生物炭的重金属固定性能;细菌协同生物炭联合修复重金属污染土壤过程中,生物炭主导吸附和固定,细菌则发挥活化和解毒等功能;优化细菌-生物炭组合形式,发展混合细菌与多种类生物炭协同技术,是复合重金属污染土壤可持续修复亟待解决的重要问题;进一步揭示细菌与生物炭对重金属污染土壤的耦合作用及长效作用机制,规避生物炭生产和应用中的潜在生态健康风险,研发新型高效能细菌与生物炭复合体是细菌协同生物炭可持续修复重金属污染土壤应用领域面临的挑战。     

土壤汞污染生物修复技术研究进展

汞作为常温下唯一的一种液态金属,在环境中分布十分广泛,极易造成土壤污染。生物修复是一种用于土壤汞污染治理的重要修复技术,该技术因具有修复成本低、绿色环保等优点而受到重视。对现有的土壤汞污染生物修复技术进行了评述,主要涉及植物修复技术、微生物修复技术和动物修复技术的研究和应用情况:植物修复技术主要研究汞在植物体内代谢机理和规律,微生物修复技术研究侧重于利用外源微生物降解土壤中汞,动物修复技术研究较少,目前仅见有关蚯蚓富集汞的报道。现代生物修复技术已经发展成为一门包括土壤化学、植物生物学、生态学、土壤微生物学、分析化学及分子生物学等多学科融合性技术,借助这些学科力量可以对植物提取汞根际微界面过程和植物体内微界面过程、微生物和动物汞富集机制有更深刻的认识。综述最后对目前土壤汞污染生物修复技术主要研究方向进行总结和归纳,明确了汞污染生物修复领域存在的几大问题以及研究方向。     

植物内生细菌修复重金属污染土壤作用机制研究进展

内生细菌生活在植物组织内部,长期以来与宿主植物形成了紧密的共生关系。内生细菌在重金属吸收、耐受和解毒方面具有优良的特性,为修复重金属污染土壤提供了有效的新方案。综述了内生细菌强化植物修复重金属污染土壤的作用机制,包括内生细菌通过产生植物生长调节激素,分泌ACC脱氨酶和几丁质酶等,促进宿主植物在重金属胁迫条件下的生长;通过改变重金属的生物有效性/毒性,减轻植物重金属毒害;通过与植物形成联合修复体,加强植物抗重金属毒性的能力。分析了近几年超富集植物内生细菌多样性及其影响因素,探讨了联合修复过程中影响内生细菌强化修复效果的主要因素,包括内生细菌的来源、活性和环境胁迫等各种生物因素和非生物因素,并对内生细菌与植物联合修复的研究方向进行展望,涉及内生细菌自身存活原因和如何耐受重金属的机制研究,植物内生细菌的行为动力学和代谢,以及内生细菌、植物及土壤之间的生态互作效应等,以期推动内生细菌大规模应用于植物修复重金属污染土壤。     

重金属污染的植物修复及相关分子机制

随着近代工业的发展,土壤重金属污染问题日益严重。重金属即使在极低浓度下仍然可以对人畜造成健康上的威胁,因此迫切需要有效的修复方法对土壤进行修复。生物修复,特别是植物修复目前已经成为重金属污染修复的重要手段之一,了解相关植物的重金属解毒和积累分子机制是提高修复效率、解决重金属污染问题的基础。文中以土壤修复方式为起点,结合植物吸收积累重金属以及解毒的相关分子机制研究,探讨了植物修复的发展现状以及趋势。     

功能内生菌强化超积累植物修复重金属污染土壤的研究进展

土壤重金属污染已成为威胁人类健康和经济可持续发展的重大环境问题。内生菌-超积累植物联合修复是近年来发展起来的一种重金属污染土壤的生物修复技术,不仅能促进植物生长、提高其重金属抗性,同时,还可以改变重金属的形态和迁移率,提高植物修复的效率,具有广阔的应用前景。现概述植物内生菌(plant growth promoting endophyte, PGPE)的特征、种类以及促进超积累植物生长的作用机理,综述近年来国内外有关功能内生菌协同超积累植物修复重金属污染土壤的应用现状和研究进展,同时展望内生菌协同超积累植物修复土壤重金属的研究思路,旨在为今后土壤重金属污染治理提供新的思路和理论依据。     

重金属污染评估及其生物健康效应

重金属污染是目前全球面临的最为严峻的环境问题之一,不仅引发了粮食安全危机,还直接危害着动物和人类健康.土壤线虫作为陆地环境中重金属污染的重要指示生物,不仅会对重金属的毒害产生复杂的防御机制,其肠道微生物还被证明可以协助宿主抵御重金属胁迫.本文系统梳理了国内外重金属的污染现状及其评估方法,以线虫为模型概述了重金属对土壤动物行为、生理、生化三个方面的影响,总结了动物对重金属暴露的应激响应,主要包括细胞壁和细胞膜的屏障作用、活性物质改变、金属硫蛋白表达以及非编码RNA调节等四个层面,提出了肠道微生物的组成和多样性作为指示重金属毒性的新方法,归纳了长期重金属暴露导致的生物适应性进化,包括重金属超积累基因、抗性基因、解毒基因的出现和富集等,进而量化环境胁迫引起的动物遗传变异在基因组水平上的响应,比如基因的抗性突变和抗性基因的协同作用.最后,本文对重金属生态毒理学的研究进行了展望,指出未来研究需着眼于当下的生态环境和人类健康,探索土壤动物肠道微生物的生态作用,深入探究重金属毒性发生发展以及生物解毒的潜在机制.     

溶磷微生物在钝化和植物修复重金属污染土壤中的作用

钝化和植物修复是重金属污染土壤修复的重要技术手段,而溶磷微生物可进一步增强钝化和植物修复重金属污染土壤的作用。介绍了钝化和植物修复重金属污染土壤的基本原理,总结了溶磷微生物对土壤中难溶性磷酸盐的溶解、利用磷酸盐钝化修复重金属污染土壤、溶磷微生物对磷酸盐钝化修复的强化以及溶磷微生物强化植物修复重金属污染土壤的研究进展,探讨了溶磷微生物对重金属的抗性及其溶磷机理、溶磷微生物对磷酸盐钝化修复重金属污染土壤的强化作用机理以及溶磷微生物强化植物修复重金属污染土壤的作用机理。旨在为生物修复重金属污染土壤研究提供一定的理论依据和技术支撑。     

铜离子稳态平衡分子机理研究进展

铜离子是生物体不可缺少的微量元素。作位酶的辅助因子,铜离子驱动着包括细胞呼吸、神经递质的传递、铁离子的摄取和抵抗氧化应激在内的重要生理过程。然而,过量时,铜离子是有害的,能损坏像DNA、蛋白质和脂肪这样的生物分子。正因为如此,生物体必须平衡细胞体内铜离子的水平。铜离子稳态平衡相关的遗传缺陷是造成Menke和Wilson疾病的原因。铜离子也被发现与癌症和神经退行性疾病有关。对酵母和其他生物体的研究发现,存在铜离子的摄取、分送、储存、排泄和抵抗毒性水平铜离子的专一机制。调控这些专一机制的铜离子信号分子是细胞平衡铜这个必不可少却又有害的离子的关键。     

Structure of the Ctr1 copper trans'PORE'ter reveals novel architecture

Copper is essential for biological processes such as free radical detoxification, mitochondrial respiration and iron metabolism. A central player in copper homeostasis is the high-affinity integral plasma membrane copper transporter Ctr1. However, the precise mechanisms by which Ctr1 functions are not known. Here, we highlight an important breakthrough in our understanding of how Ctr1 facilitates Cu(I) movement across membranes: the publication of structural details for human Ctr1 obtained from 2D crystallography and electron microscopy.     

农田土壤中铜的主要输入途径及其污染风险控制

综述农田土壤中畜禽粪便、杀菌剂、再生水和化肥等途径对铜的输入及其污染风险,以土壤理化性质为切入点概述外源铜进入土壤后的吸附-解吸和移动性特点,探讨不同作物类型和土地利用方式对农产品中铜含量和对通过膳食途径铜健康风险的影响,并针对实际生产提出规避农田土壤铜污染风险的有效措施.     

Molecular basis of active copper resistance mechanisms in Gram-negative bacteria

Copper is a metallic element that is crucial for cell metabolism; however, in extended concentrations, it is toxic for all living organisms. The dual nature of copper has forced organisms, including bacteria, to keep a tight hold on cellular copper content. This challenge has led to the evolution of complex mechanisms that on one hand enable them to deliver the essential element and on the other to protect cells against its toxicity. Such mechanisms have been found in both eukaryotic and prokaryotic cells. In bacteria a number of different systems such as extra- and intracellular sequestration, enzymatic detoxification, and metal removal from the cell enabling them to survive in the presence of high concentration of copper have been identified. Gram-negative bacteria, due to their additional compartment, need to deal with both cytoplasmic and periplasmic copper. Therefore, these bacteria have evolved intricate and precisely regulated systems which interact with each other. In this review the active mechanisms of copper resistance at their molecular level are discussed.     

Effects of copper on mammalian cell components

Both deficiency and excess of copper induce toxic effects on mammalian cell systems in vivo and in vitro. The effects can be related to the affinities of Cu(II) ions for specific cell components. The nucleus is a potential site for temporary Cu storage while primary targets for free Cu(II) ions are the thiol groups which reduce the ions to Cu(I). Cu(II) ions show a high affinity for nucleic acids, binding with DNA both at intrastrand and interstrand levels, possibly through intercalation between GC pairs. The ability to chelate Cu(II) ions is seen to be of the order: purine greater than purine ribonucleotides greater than purine ribonucleoside greater than pyrimidine ribonucleotides. Copper is an integral part of enzyme activation and enters into the molecular structure of several proteins, like ceruloplasmin. Cu(II) ion is a potential mutagenic agent as seen by its property of inducing infidelity in DNA synthesis in vitro. Teratogenic activities of copper have been reported but carcinogenicity is not yet confirmed. Copper is an essential component of chromatin and is known to accumulate preferentially in the heterochromatic regions. External application of higher doses, however, induces both clastogenic effects and spindle disturbances. In certain forms, inorganic copper enhances the clastogenic activity of other agents. The most widely studied human genetic maladies linked with copper metabolism are Menkes' and Wilson's diseases. Several mutations are known which influence Cu homeostasis in mammals. Such mutations in mice have been used extensively for biochemical studies.     

Phytochelatin is not a primary factor in determining copper tolerance

Phytochelatin (PC) is involved in the detoxification of harmful, non-essential heavy metals and the homeostasis of essential heavy metals in plants. Its synthesis can be induced by either cadmium (Cd) or copper (Cu), and can form stable complexes with either element. This might suggest that PC has an important role in determining plant tolerance to both. However, this is not clearly apparent, as evidenced by a PC-deficient and Cd-sensitiveArabidopsis mutant (cad1-3) that shows no significant increase in its sensitivity to copper. Therefore, we investigated whether the mechanism for Cu tolerance differed from that for Cd by analyzing copper sensitivity in Cd-tolerant transgenics and Cd-sensitive mutants ofArabidopsis. Cadmium-tolerant transgenic plants that over-expressedA. thaliana phytochelatin synthase 1 (AtPCS1) were not tolerant of copper stress, thereby supporting the hypothesis that PC is not primarily involved in this tolerance mechanism. We also investigated Cu tolerance incad2-1, a Cd-sensitive and glutathione (GSH)-deficientArabidopsis mutant. Paradoxically,cad2-1 was more resistant to copper stress than were wild-type plants. This was likely due to the high level of cysteine present in that mutant. However, when the growth medium was supplemented with cysteine, the wild types also exhibited copper tolerance. Moreover,Saccharomyces cerevisiae that expressedAtPCS1 showed tolerance to Cd but hypersensitivity to Cu. All these results indicate that PC is not a major factor in determining copper tolerance in plants.     

Proteomic analysis of copper stress responses in the roots of two rice (Oryza sativa L.) varieties differing in Cu tolerance

Background and aims

Copper (Cu) is an essential micronutrient required for growth and development of plants. However, excess Cu is toxic to plants. To understand the mechanisms involved in copper stress response, a proteomic approach was used to investigate the differences in Cu stress-induced protein expression between a Cu-tolerant variety (B1139) and a Cu-sensitive one (B1195) of rice.

Methods

Rice seedlings were exposed to 8 μM Cu for 3 days, with plants grown in the normal nutrient solution containing 0.32 μM Cu serving as the control. Proteins were extracted from the roots and separated by two-dimensional PAGE. Thirty four proteins were identified using MALDI-TOF mass spectrometry.

Results

Thirty-four protein spots were found to be differently expressed in the Cu-stressed roots in at least one variety of rice, including those involved in antioxidative defense, redox regulation, stress response, sulfur and glutathione (GSH) metabolism, carbohydrate metabolism, signal transduction, and some other proteins with various functions. Nine proteins, including putative cysteine synthase, probable serine acetyltransferase 3, L-ascorbate peroxidase 1, putative glutathione S-transferase 2, and thioredoxin-like 3-3, exhibited a greater increase in response to Cu stress in the Cu-tolerant variety B1139 compared with the Cu–sensitive variety B1195.

Conclusion

The majority of the proteins showing differential expression in response to Cu exposure are involved in the redox regulation, and sulfur and GSH metabolism, suggesting that these proteins, together with antioxidant enzymes, play an important role in the detoxification of excess Cu and maintaining cellular homeostasis.     

Leaf proteome characterization in the context of physiological and morphological changes in response to copper stress in sorghum

Copper (Cu) is an essential micronutrient required for normal growth and development of plants; however, at elevated concentrations in soil, copper is also generally considered to be one of the most toxic metals to plant cells due to its inhibitory effects against many physiological and biochemical processes. In spite of its potential physiological and economical significance, molecular mechanisms under Cu stress has so far been grossly overlooked in sorghum. To explore the molecular alterations that occur in response to copper stress, the present study was performed in ten-day-old Cu-exposed leaves of sorghum seedlings. The growth characteristics were markedly inhibited, and ionic alterations were prominently observed in the leaves when the seedlings were exposed to different concentrations (0, 100, and 150 µM) of CuSO₄. Using two-dimensional gels with silver staining, 643 differentially expressed protein spots (≥1.5-fold) were identified as either significantly increased or reduced in abundance. Of these spots, a total of 24 protein spots (≥1.5-fold) from Cu-exposed sorghum leaves were successfully analyzed by MALDI-TOF-TOF mass spectrometry. Of the 24 differentially expressed proteins from Cu-exposed sorghum leaves, 13 proteins were up-regulated, and 11 proteins were down-regulated. The abundance of most identified protein species, which function in carbohydrate metabolism, stress defense and protein translation, was significantly enhanced, while that of another protein species involved in energy metabolism, photosynthesis and growth and development were severely reduced. The resulting differences in protein expression patterns together with related morpho-physiological processes suggested that these results could help to elucidate plant adaptation to Cu stress and provide insights into the molecular mechanisms of Cu responses in C₄ plants.     

Comparative genomic analyses of copper transporters and cuproproteomes reveal evolutionary dynamics of copper utilization and its link to oxygen

Copper is an essential trace element in many organisms and is utilized in all domains of life. It is often used as a cofactor of redox proteins, but is also a toxic metal ion. Intracellular copper must be carefully handled to prevent the formation of reactive oxygen species which pose a threat to DNA, lipids, and proteins. In this work, we examined patterns of copper utilization in prokaryotes by analyzing the occurrence of copper transporters and copper-containing proteins. Many organisms, including those that lack copper-dependent proteins, had copper exporters, likely to protect against copper ions that inadvertently enter the cell. We found that copper use is widespread among prokaryotes, but also identified several phyla that lack cuproproteins. This is in contrast to the use of other trace elements, such as selenium, which shows more scattered and reduced usage, yet larger selenoproteomes. Copper transporters had different patterns of occurrence than cuproproteins, suggesting that the pathways of copper utilization and copper detoxification are independent of each other. We present evidence that organisms living in oxygen-rich environments utilize copper, whereas the majority of anaerobic organisms do not. In addition, among copper users, cuproproteomes of aerobic organisms were larger than those of anaerobic organisms. Prokaryotic cuproproteomes were small and dominated by a single protein, cytochrome c oxidase. The data are consistent with the idea that proteins evolved to utilize copper following the oxygenation of the Earth.