重金属污染的转基因植物修复——原理与应用

污染环境的植物修复技术具有成本低、不造成二次污染等优点。从自然界中寻找用于污染环境修复的超富积植物不仅难度大 ,而且受生物量、生长周期以及地理环境等因素的限制。近几年迅速发展起来的通过转基因植物进行污染环境的修复技术显示了广阔的应用前景。外源基因在植物的高效表达可以提高植物吸收、运输、降解污染物的能力以及修复的效率 ,并可以作为研究不同污染物修复机理的实验系统。以转基因植物修复几种主要的重金属污染为例 ,介绍了转基因植物修复的原理、现状及存在问题 ,并探讨了提高转基因植物修复效率的一些方法 。     

EDTA-Enhanced Phytoremediation of Heavy Metals: A Review

The increase in heavy metal terrestrial ecosystems’ contamination through anthropogenic activities is a widespread and serious global problem due to their various environmental and human implications. For these reasons, several techniques, including phytoremediation of heavy metals, have been extensively studied. In spite of significant recent advancement, ethylene diamine tetraacetic acid (EDTA)-enhanced heavy metal phytoextraction as well as related ecological risks are still topical and remain an important area of research. In fact, EDTA favors the solubilization of metals and metalloids in soils, and was therefore extensively studied during the last two decades in order to improve phytoextraction efficiency and reduce treatment duration. This review highlights the recent findings (2010–2012) and mechanisms behind EDTA-enhanced (1) solubilization of heavy metals in soil, (2) mobilization/transport of soluble metals towards plant root zone, and (3) metal absorption by plant roots and translocation towards aerial parts. The review also presents potential risks associated with EDTA-enhanced phytoextraction: (1) environmental persistence of EDTA and/or metal-EDTA complex; (2) potential toxicity of EDTA and/or metal-EDTA complex to plants; and (3) leaching and contamination of groundwater. Moreover, field-scale cost of EDTA-enhanced remediation and the role of EDTA in time required for heavy metal remediation is discussed.     

重金属诱导细胞凋亡的分子机制

重金属诱导的细胞凋亡是一个十分复杂的过程,不同种类的重金属以及同类重金属离子的不同价态所诱导的凋亡效应及其分子机制不尽相同。目前的研究表明,重金属与DNA形成加合物而导致DNA损伤可能是引发细胞凋亡的重要步骤：多种重金属能通过激活内质网、线粒体钙通道,使Ca^2 释放进入细胞质而引发凋亡;重金属还能使细胞中ROS升高,在直接导致DNA损伤的同时,启动与线粒体相关的细胞凋亡信号通路。此外,ROS还能通过MAPKs增强JNK介导的：FasL和Fas表达,最终使caspase-3和caspase-7激活,从而促进凋亡的发生。重金属诱导细胞凋亡还涉及一系列重要基因和蛋白质的参与,包括促进凋亡的Src家族酪氨酸激酶、bax、fas和p53等基因及相关蛋白,抑制凋亡的Sp1锌指转录因子、bcl-2和myc等基因及相关蛋白。部分重金属如镉、锌等对细胞凋亡具有诱导和拮抗双重效应,其中拮抗效应主要是通过与自由钙离子协同进行的,而诱导效应则可能是通过调节caspase-3活性而实现的。     

Chelate-Enhanced Phytoremediation of Soils Polluted with Heavy Metals

In general, hyperaccumulators are low biomass, slow-growing plants. High biomass non-hyperaccumulator plants by themselves are not a valid alternative for phytoextraction as they also have many limitations, such as small root uptake and little root-to-shoot translocation. In this context, chemically-induced phytoextraction (based on the fact that the application of certain chemicals, mostly chelating agents, to the soil significantly enhances metal accumulation by plants) has been proposed as an alternative for the cleaning up of metal polluted soils. But chelate-induced phytoextraction increases the risk of adverse environmental effects due to metal mobilization during extended periods of time. In order to minimize the phytotoxicity and environmental problems associated with the use of chelating agents, nowadays, research is being carried out on the gradual application of small doses of the chelating agent during the growth period. However, EDTA utilization in the future will most likely be limited to ex situconditions where control of the leachates can be achieved. There are other mobilizing agents which are much less harmful to the environment such as citric acid, NTA, and particularly EDDS. Research should also be aimed towards more innovative agronomic practices. Environmentally safe methods of chelate-induced phytoextraction must be developed before steps towards further development and commercialization of this remediation technology are taken. Most importantly, more applied projects in this field are needed to clarify the real potential and risks of this technology.     

Phytoremediation of Heavy and Transition Metals Aided by Legume-Rhizobia Symbiosis

Legumes are important for nitrogen cycling in the environment and agriculture due to the ability of nitrogen fixation by rhizobia. In this review, we introduce an important and potential role of legume-rhizobia symbiosis in aiding phytoremediation of some metal contaminated soils as various legumes have been found to be the dominant plant species in metal contaminated areas. Resistant rhizobia used for phytoremediation could act on metals directly by chelation, precipitation, transformation, biosorption and accumulation. Moreover, the plant growth promoting (PGP) traits of rhizobia including nitrogen fixation, phosphorus solubilization, phytohormone synthesis, siderophore release, and production of ACC deaminase and the volatile compounds of acetoin and 2, 3-butanediol may facilitate legume growth while lessening metal toxicity. The benefits of using legumes inoculated with naturally resistant rhizobia or recombinant rhizobia with enhanced resistance, as well as co-inoculation with other plant growth promoting bacteria (PGPB) are discussed. However, the legume-rhizobia symbiosis appears to be sensitive to metals, and the effect of metal toxicity on the interaction between legumes and rhizobia is not clear. Therefore, to obtain the maximum benefits from legumes assisted by rhizobia for phytoremediation of metals, it is critical to have a good understanding of interactions between PGP traits, the symbiotic plant-rhizobia relationship and metals.     

EDTA-Facilitated Phytoremediation of Soil with Heavy Metals from Sewage Sludge

The objective of this research was to determine the effect of the chelate EDTA (ethylenediaminetetraacetic acid), which is used in phytoremediation, on plant availability of heavy metals in liquid sewage sludge applied to soil. Sunflower (Helianthus annuus L.) was grown under greenhouse conditions in a commercial potting soil; the tetrasodium salt of EDTA (EDTA Na₄) was added at a rate of 1 g kg^-1 to half the pots. Immediately after seeds were planted, half of the pots with each soil (with or without EDTA) were irrigated with 60 ml sludge, and half were irrigated with 60 ml tap water. For the subsequent five irrigations, plants in soil with EDTA received either sludge or tap water containing 0.5 g EDTA Na₄ per 1000 ml, and plants in soil without EDTA received sludge or tap water without EDTA. Of the four heavy metals whose extractable concentrations in the soil were measured (Cu, Fe, Mn, and Zn), only Zn had a higher concentration in sludge-treated soil with EDTA compared to sludge-treated soil without EDTA. The concentrations of Fe, Cu, and Mn were similar in sludge-treated soil with and without EDTA. Of the three heavy metals whose total concentrations in the soil were measured (Cd, Pb, Cr), Pb (<10 mg kg^-1) and Cd (< 1 mg kg^-1) were below detection limits, and Cr was unaffected by treatment. The concentration of all measured elements in plants (Cd, Cu, Fe, Zn, Pb) was higher than the concentrations measured in the soil. With no EDTA, sludge-treated plants had a higher concentration of the five heavy elements than plants grown without sludge. Cadmium was lower in sludge-treated plants with EDTA than plants with EDTA and no sludge. After treatment with EDTA, the concentrations of Cu, Fe, and Zn were similar in plants with and without sludge. Lead was higher in plants with EDTA than plants without EDTA, showing that EDTA can facilitate phytoremediation of soil with Pb from sewage sludge.     

重金属污染环境的植物修复及其分子机制

重金属污染物的排放和扩散造成了日益严重的环境污染。如何消除环境中的重金属污染物已成为国际性难题。近年来,植物修复技术的出现和快速发展为我们展示了一条新的有效途径：即利用植物对重金属化合物的吸收、富集和转化能力去除土壤和水体中残存的重金属污染物。文章简要介绍了重金属污染物与植物修复的关系和植物修复的生理机制,重点总结了重金属污染环境的植物修复在分子生物学方面所取得的研究进展,包括有机汞裂解酶基因merB、汞离子还原酶基因merA和金属硫蛋白基因MT等的生物学功能及其在植物修复上的应用,展望了植物修复研究工作的发展方向,并针对汞污染提出了一套修复方案。     

微生物抗重金属的生理机制

虽然环境中高浓度的重金属对微生物有毒害作用,但一些微生物表现出抗重金属的特性,在细胞结构、生理代谢和遗传水平方面形成了不同的抗性机制.主要从生物吸附、胞外沉淀、生物转化、生物累积和外排作用等5个方面阐述了微生物对重金属抗性的生理和分子机制,为重金属污染环境的生物修复提供理论依据.     

强化植物修复重金属污染土壤的策略及其机制

重金属对生态环境、农业生产、人类健康等诸多方面造成重要危害。植物修复因其具有经济有效、绿色生态等优点,已经成为土壤重金属污染修复研究领域的热点。由于植物重金属毒害、修复耗时过长等因素致使植物修复技术受限于研究阶段而不能广泛应用于实践。采用科学合理的强化措施提高植物修复的效率可能是解决该矛盾的关键之一。讨论了根瘤菌、丛枝菌根真菌、溶磷微生物和内生真菌构建的微生物-植物共生系统在强化植物修复过程中的具体应用;概述了EDTA、EDDS等螯合剂在改变土壤中重金属可溶态,促进重金属从土壤向植株转运的重要作用;介绍了植物中编码金属转运蛋白、金属硫蛋白、植物螯合肽等与重金属转运和代谢相关的基因在植物修复领域的实际应用;归纳了上述强化策略主要机制为微生物促进植物生长、缓解重金属植物毒性以及提高了土壤中重金属生物利用度,从而促进重金属在富集植物中积累和植物生物量的增加;最后总结并展望了植物修复强化技术在今后研究的重点及存在的问题。综述植物修复技术采用的主要强化策略及其机制,旨在为利用植物修复技术治理土壤重金属污染提供重要参考。     

Phytoremediation of Metals: A Numerical Analysis

A finite element code was used for investigating the effect of some relevant characteristics of a phytoremediation project (crop type and density, presence of an irrigation system, soil capping and root depth). The evolution of the plume of contamination of Cd²⁺, Pb²⁺, and Zn²⁺ was simulated taking into account reactive transport and root processes. The plant contaminant uptake model was previously calibrated using data from greenhouse experiments. The simulations adopted pedological and climatological data representative of a sub-tropical environment. Although the results obtained were specific for the proposed scenario, it was observed that, for more mobile contaminants, poor water conditions favor stabilization but inhibit plant extraction. Otherwise an irrigation system that decreases crop water stress had an opposite effect. For less mobile contaminants, the remediation process did not have appreciable advantages. Despite its simplifying assumptions, particularly about contaminant sorption in the soil and plant system, the numerical analysis provided useful insight for the phytoextraction process important in view of field experiments.     

植物修复的生物学机制

随着现代化工业的发展,全球向土壤和环境中排放的重金属逐年增加。重金属污染已日益成为威胁人类健康和影响人类生活质量的严重环境问题和社会问题。这一问题可部分通过植物修复技术得以解决。植物修复技术是依据植物从环境中积累重金属元素和化合物的能力及其将这些有毒物质在植物体内代谢成无毒生物小分子的能力而建立的新的生物技术。本篇综述主要论及利用植物修复技术解决重金属污染的生物学机制。     

Phytoremediation of Heavy Metals in Contaminated Water and Soil Using Miscanthus sp. Goedae-Uksae 1

The aim of this study is to characterize the heavy metal phytoremediation potential of Miscanthus sp. Goedae-Uksae 1, a hybrid, perennial, bio-energy crop developed in South Korea. Six different metals (As, Cu, Pb, Ni, Cd, and Zn) were used for the study. The hybrid grass effectively absorbed all the metals from contaminated soil. The maximum removal was observed for As (97.7%), and minimum removal was observed for Zn (42.9%). Similarly, Goedae-Uksae 1 absorbed all the metals from contaminated water except As. Cd, Pb, and Zn were completely (100%) removed from contaminated water samples. Generally, the concentration of metals in roots was several folds higher than in shoots. Initial concentration of metals highly influenced the phytoremediation rate. The results of the bioconcentration factor, translocation factor, and enrichment coefficient tests indicate that Goedae-Uksae 1 could be used for phytoremediation in a marginally contaminated ecosystem.     

Physiological Effects of Heavy Metals on the Moss Rhytidiadelphus squarrosus

Experiments on the effect of Cd, Cu, Ni, Pb and Zn, suppliedas single salt solutions between 10 µM and 0·1M, on the moss Rhytidiadelphus squarrosus showed little alterationto respiratory rates but reduced photosynthetic rates, and somemembrane damage as assessed by K leakage. Two distinct morphologicalforms of the moss showed different responses to supplied heavymetal. Storage of material, after 30 mins exposure to heavymetals, resulted in a further decrease in the photosyntheticrate. Expressing the photosynthetic decline relative to thetotal heavy metal recovered from the moss showed a similar pattern.Transfer of metal from extracellular exchange sites into theprotoplast was also demonstrated with storage after exposure.An approximately linear relationship was demonstrated betweenphotosynthetic decline and intracellular heavy-metal concentration,irrespective of the duration of exposure or morphological natureof the material used. Photosynthetic decline on storage is concludedto be a response to additional metal stress rather than a progressivedeterioration of the physiological process. Cadmium, copper, lead, nickel, zinc, photosynthesis, respiration, membrane damage, intracellular metals, Rhytidiadelphus squarrosus (Hedw.) Warnst     

Phytoremediation of Metals Using Transgenic Plants

An ideal plant for environmental cleanup can be envisioned as one with high biomass production, combined with superior capacity for pollutant tolerance, accumulation, and/or degradation, depending on the type of pollutant and the phytoremediation technology of choice. With the use of genetic engineering, it is feasible to manipulate a plant's capacity to tolerate, accumulate, and/or metabolize pollutants, and thus to create the ideal plant for environmental cleanup. In this review, we focus on the design and creation of transgenic plants for phytoremediation of metals. Plant properties important for metal phytoremediation are metal tolerance and accumulation, which are determined by metal uptake, root-shoot translocation, intracellular sequestration, chemical modification, and general stress resistance. If we know which molecular mechanisms are involved in these tolerance and accumulation processes, and which genes control these mechanisms, we can manipulate them to our advantage. This review aims to give a succinct overview of plant metal tolerance and accumulation mechanisms, and to identify possible strategies for genetic engineering of plants for metal phytoremediation. An overview is presented of what has been achieved so far regarding the manipulation of plant metal metabolism. In fact, both enhanced metal tolerance and accumulation have been achieved by overproducing metal chelating molecules (citrate, phytochelatins, metallothioneins, phytosiderophores, ferritin) or by the overexpression of metal transporter proteins. Mercury volatilization and tolerance was achieved by introduction of a bacterial pathway. The typical increase in metal accumulation as the result of these genetic engineering approaches is 2-to 3-fold more metal per plant, which could potentially enhance phytoremediation efficiency by the same factor. As for the applicability of these transgenics for environmental cleanup, results from lab and greenhouse studies look promising for several of these transgenics, but field studies will be the ultimate test to establish their phytoremediation potential, their competitiveness, and risks associated with their use.     

Hyperaccumulator Oilcake Manure as an Alternative for Chelate-Induced Phytoremediation of Heavy Metals Contaminated Alluvial Soils

The ability of hyperaccumulator oilcake manure as compared to chelates was investigated by growing Calendula officinalis L for phytoremediation of cadmium and lead contaminated alluvial soil. The combinatorial treatment T₆ [2.5 g kg^?1oilcake manure + 5 mmol kg^?1 EDDS] caused maximum cadmium accumulation in root, shoot and flower up to 5.46, 4.74 and 1.37 mg kg^?1and lead accumulation up to 16.11, 13.44 and 3.17 mg kg^?1, respectively at Naini dump site, Allahabad (S₃). The treatment showed maximum remediation efficiency for Cd (RR = 0.676%) and Pb (RR = 0.202%) at Mumfordganj contaminated site (S₂). However, the above parameters were also observed at par with the treatment T₅ [2.5 g kg^?1oilcake manure +2 g kg^?1 humic acid]. Applied EDDS altered chlorophyll–a, chlorophyll–b, and carotene contents of plants while application of oilcake manure enhanced their contents in plant by 3.73–8.65%, 5.81–17.65%, and 7.04–17.19%, respectively. The authors conclude that Calendula officinalis L has potential to be safely grown in moderately Cd and Pb-contaminated soils and application of hyperaccumulator oilcake manure boosts the photosynthetic pigments of the plant, leading to enhanced clean-up of the cadmium and lead-contaminated soils. Hence, the hyperaccumulator oilcake manure should be preferred over chelates for sustainable phytoremediation through soil-plant rhizospheric process.     

植物UV-B生理效应的分子机制研究进展

中波紫外线UV-B（280～320nm）是植物必需的太阳光线的组成部分,具有明显的双重效应：一方面UV-B在强度较高时,就触发产生大量活性氧对DNA、蛋白质以及生物膜等造成伤害,同时植物通过抗氧化系统对其作出防御反应以减轻伤害;另一方面,低强度的UV-B是植物生长发育的光信号因子之一,经由UVR8等光受体介导中、低、极低强度的UV-B信号,可能通过几个分子途径控制相关基因的表达,分别对植物的UV-B保护基因表达、形态建成、昼夜节律、生长发育等进行调控。目前对UVR8介导的低强度UV-B信号转导的分子机制研究相对深入。在本文中,将对UV-B生理效应分子机制的最新研究进展作一个比较全面的介绍。     

Molecular and Physiological Mechanisms of Flooding Avoidance and Tolerance in Rice

Submergence is one of the major constraints in rice production. The main factor limiting rice survival during submergence is oxygen deprivation. To cope with flooding conditions, rice has developed two survival strategies: either rapid elongation of the submerged tissues to keep up with the rising water level or no elongation to save carbohydrate resources for maintenance of energy production under submerged and concomitant hypoxic conditions. The survival strategies used by rice have been studied quite extensively and the role of several phytohormones in the elongation response has been established. The mechanisms of submergence tolerance include metabolic changes, for instance, the shift to an ethanolic fermentation pathway, reduced elongation growth to save carbohydrates and energy for maintenance processes, and protective antioxidant systems. Current molecular technology can provide tools for the understanding of mechanisms developed by rice to survive submergence. In addition, cloning of genes related to submergence tolerance might open new ways to genetic improvement of this crop.     

细菌抗生素和重金属协同选择抗性机制研究进展

随着各类抗生素和新型复合金属材料的不断开发和使用,环境中抗生素和重金属离子协同污染的机率不断提高,对环境选择最为敏感的细菌通过自身的进化和发展形成了二者协同选择的抗性机制,如协同抗性、交叉抗性、协同调控和生物膜诱导机制。     

植物种子寿命的生理及分子机制研究进展

种子寿命是衡量种子质量高低的关键指标之一,直接关系到种子萌发、萌发后幼苗的生长发育以及作物产量高低。种子寿命的调控是一个复杂的生物学过程,影响种子寿命的因素包括环境因素、种子自身的结构、营养成分组成及含量以及调控种子寿命相关的关键基因。研究储藏过程中种子生理生化指标的变化,以及相应关键基因的生物学功能,掌握调控种子寿命的生理及分子机制,对于减少种子内部能量消耗,进一步延长种子寿命具有重要意义。该文综述了近年来国内外有关调控种子寿命的生理及分子机制,重点阐述了调控种子寿命的相关关键基因的研究进展,并讨论了各种外部因素对种子寿命的调控机理。