随着城市化、工业化发展的脚步加快，土壤重金属污染成为国内外普遍关注的环境问题，并且已经严重威胁到地球生态系统的稳定和人类生命的安全，因此，对土壤重金属治理刻不容缓。本研究在介绍土壤重金属污染的现状的基础上，综述了土壤重金属对植物各方面的影响，概述了土壤重金属的植物修复机理以及土壤重金属的植物修复研究进展，提出了今后的研究方向：未来应挑选出超富集植物，为土壤重金属污染植物修复机理的研究工作奠定基础；重点研究微生物对植物修复的研究效果；加强对木本植物的研究，特别是耐性强的乡土植物修复土壤重金属的能力研究；对超富集植物凋落后的安全回收、合理利用进行研究。

This article reviews the responses of plant roots to elevated rhizosphere cadmium (Cd) concentrations. Cadmium enters plants from the soil solution. It traverses the root through symplasmic or apoplasmic pathways before entering the xylem and being translocated to the shoot. Leaf Cd concentrations in excess of 5-10 μg g(-1) dry matter are toxic to most plants, and plants have evolved mechanisms to limit Cd translocation to the shoot. Cadmium movement through the root symplasm is thought to be restricted by the production of phytochelatins and the sequestration of Cd-chelates in vacuoles. Apoplasmic movement of Cd to the xylem can be restricted by the development of the exodermis, endodermis, and other extracellular barriers. Increasing rhizosphere Cd concentrations increase Cd accumulation in the plant, especially in the root. The presence of Cd in the rhizosphere inhibits root elongation and influences root anatomy. Cadmium concentrations are greater in the root apoplasm than in the root symplasm, and tissue Cd concentrations decrease from peripheral to inner root tissues. This article reviews current knowledge of the proteins involved in the transport of Cd across root cell membranes and its detoxification through sequestration in root vacuoles. It describes the development of apoplastic barriers to Cd movement to the xylem and highlights recent experiments indicating that their maturation is accelerated by high Cd concentrations in their immediate locality. It concludes that accelerated maturation of the endodermis in response to local Cd availability is of functional significance in protecting the shoot from excessive Cd loads.

芥菜（Brassica juncea）对多种重金属具有富集能力。重金属ATP酶（heavy metal transporting ATPase，HMA）在植物转运重金属过程中具有重要的作用。基于基因组和转录组数据，对芥菜HMA家族基因成员进行了全基因组鉴定。芥菜基因组中包含27个HMA基因，其编码的蛋白质中有6个不稳定指数大于40，有22个为疏水性蛋白；这些基因聚类为P-1B-1、P-1B-2和P-1B-4等3个亚家族；27个HMA蛋白均为膜蛋白，都具有E1-E2 ATPase和hydrolase结构域；芥菜HMA蛋白都含有8 ~ 15个保守基序，不同亚族因具有独特的保守基序结构从而转运重金属的种类不同；在芥菜HMA基因启动子区域中与生长发育有关的顺式作用元件TGA和与逆境响应相关的顺式作用元件ABRE、MBS、LTR、TC广泛分布；在镉胁迫下，BjuA033764、BjuB019118、BjuB035256、BjuB045293等基因在叶片中表达量明显上调，BjuA003596、BjuB040613、BjuA025022等基因在根系中表达量明显上调，说明其参与了芥菜对镉胁迫的响应。

Tn7 insertion mutagenesis has been used to facilitate the generation of a physical (restriction endonuclease) and genetic map of the IncM plasmid, R831b. The only selectable phenotypes carried by this 90-kb conjugative plasmid are resistances to inorganic mercury [Hg(II)] and to organomercury compounds. Mutants in the Hgr locus of R831b complemented previously described mutants in the mer operon of the IncFII plasmid R100, indicating functional homology of the locus in each of these different plasmids. However, the R831b Hgr locus is not notably similar in restriction site pattern to either the mer operon of R100 or the mercury resistance transposon, Tn501. Although the enzymes they encode are co-ordinately regulated, the Omr locus of R831b maps approx. 13.5 kb away from the Hgr locus. Three insertions which affect neither phenotype lie between the Hgr and Omr loci; thus, the loci are separated both physically and genetically. One mutant was obtained which tentatively identifies the position of the Tra locus of R831b as adjacent to the Hgr locus.

The mercury resistance (mer) operon of the gram-negative transposon Tn21 encodes not only a mercuric reductase and regulatory genes but also two inner membrane proteins (MerT and MerC) and a periplasmic protein (MerP). Although the merT, merP, and merC genes have been implicated in Hg(II) transport, the individual roles of these genes have not been established. We created in vitro precise deletion and frameshift mutations that eliminated each of the genes singly and in combination. Our results show that both merT and merP are required for Hg(II) binding but that merC is not. Both merT and merP are required for full expression of Hg(II) resistance, but loss of merP is less deleterious than loss of merT. Furthermore, mutations eliminating both merT and merP decrease resistance more than the single mutations do. In contrast, mutating merC had no effect on Hg(II) resistance. Both the merT and merP mutations increase the threshold Hg(II) concentration for induction of merA-lacZ transcriptional fusions and cause an increase in the maximal expression level. In contrast, the merC mutation had little effect on the threshold inducing concentration of Hg(II) but decreased the level of expression. Our results show that merT and merP alone are sufficient to specify a mercury transport system. The role of merC remains obscure.

Methylmercury is a highly toxic, organic derivative found in mercury-polluted wetlands and coastal sediments worldwide. Though commonly present at low concentrations in the substrate, methylmercury can biomagnify to concentrations that poison predatory animals and humans. In the interest of developing an in situ detoxification strategy, a model plant system was transformed with bacterial genes (merA for mercuric reductase and merB for organomercurial lyase) for an organic mercury detoxification pathway. Arabidopsis thaliana plants expressing both genes grow on 50-fold higher methylmercury concentrations than wild-type plants and up to 10-fold higher concentrations than plants that express merB alone. An in vivo assay demonstrated that both transgenes are required for plants to detoxify organic mercury by converting it to volatile and much less toxic elemental mercury.

Heavy metal pollution often occurs together with organic contaminants. Brassinosteroids (BRs) induce plant tolerance to several abiotic stresses, including phenanthrene (PHE) and cadmium (Cd) stress. However, the role of BRs in PHE+Cd co-contamination-induced stress amelioration is unknown. Here, the interactive effects of PHE, Cd, and 24-epibrassinolide (EBR; a biologically active BR) were investigated in tomato plants. The application of Cd (100 µM) alone was more phytotoxic than PHE applied alone (100 µM); however, their combined application resulted in slightly improved photosynthetic activity and pigment content compared with Cd alone after a 40 d exposure. Accumulation of reactive oxygen species and membrane lipid peroxidation were induced by PHE and/or Cd; however, the differences in effect were insignificant between Cd and PHE+Cd. The foliar application of EBR (0.1 µM) to PHE- and/or Cd-stressed plants alleviated photosynthetic inhibition and oxidative stress by causing enhancement of the activity of the enzymes and related transcript levels of the antioxidant system, secondary metabolism, and the xenobiotic detoxification system. Additionally, PHE and/or Cd residues were significantly decreased in both the leaves and roots after application of EBR, more specifically in PHE+Cd-stressed plants when treated with EBR, indicating a possible improvement in detoxification of these pollutants. The findings thus suggest a potential interaction of EBR and PHE for Cd stress alleviation. These results advocate a positive role for EBR in reducing pollutant residues for food safety and also strengthening phytoremediation.

Environmental stress, including heavy metal stress, can cause oxidative damage to plants. Up-regulation of the antioxidant defense system induced by proline and glycinebetaine (betaine) alleviates the damaging effects of oxidative stress in plants. Here, we investigated the protective effects of exogenously applied proline and betaine on growth, accumulation of proline and betaine, lipid peroxidation and activity of antioxidant enzymes in cultured tobacco Bright Yellow-2 (BY-2) cells exposed to cadmium (Cd) stress. Cadmium stress (at 100 microM Cd) caused a significant inhibition of the growth of BY-2 cells, and both proline and betaine significantly mitigated this inhibition. In addition, the mitigating effect of proline was more pronounced than that of betaine. Cadmium stress leads to an accumulation of Cd and endogenous proline in cultured cells, increased lipid peroxidation and peroxidase (POX) activity, and decreased activity of superoxide dismutase (SOD) and catalase (CAT). Exogenous application of proline resulted in a decrease in lipid peroxidation and an increase in SOD and CAT activities without reducing Cd contents under Cd stress, while application of betaine resulted in a decrease in lipid peroxidation and an increase in CAT activity with reducing Cd accumulation. Furthermore, exogenous proline and betaine intensified the accumulation of proline and betaine in Cd-stressed BY-2 cells, respectively. The present study suggests that proline and betaine confer tolerance to Cd stress in tobacco BY-2 cells by different mechanisms.

植物激素可打破重金属超富集植物的种子休眠、促进发芽和快速生长,从而提高其富集重金属的效率。虽然人工合成的植物生长调节剂在促进超富集植物生长和提高重金属富集方面已取得很多成果,但大多是盆栽试验,大田试验甚少。文章在课题组前期室内盆栽试验筛选出的吲哚乙酸（IAA）和激动素（KT）提高砷（As）超富集植物蜈蚣草（Pteris vittata L.）砷提取效率的最佳配比（IAA?KT=25 mg∙L^-1?20 mg∙L^-1）基础上,在云南省个旧市大屯镇重金属污染农田分45个小区、喷施2次激素开展IAA和KT配合施用对As超富集植物蜈蚣草和镉（Cd）超富集植物龙葵（Solanum nigrum L.）超富集As和Cd的影响和机理研究。结果表明,大田条件下IAA和KT配合施用能够促进2种超富集植物快速生长,在含Cd为3.12 mg∙kg^-1、含砷As 98.7 mg∙kg^-1的农田土壤上,与未施用植物激素的对照相比,25 mg∙L^-1 IAA和20 mg∙L^-1 KT配合施用后,龙葵和蜈蚣草的株高、鲜物质量、地上部和地下部Cd/As含量、Cd/As转运系数和富集系数均显著增加,且龙葵对Cd、蜈蚣草对As的提取效率最高可分别达7.52%和6.06%。第2次喷施激素后,单加KT和激素配合施用条件下,龙葵和蜈蚣草间作时两种植物叶片过氧化物酶（POD）活性均显著增加。逐步回归分析结果表明,龙葵对As、蜈蚣草对Cd的提取效率均与叶片POD活性成显著正相关。因此,叶片保持较高的POD活性对两种超富集植物对Cd和As的提取具有重要意义。

陆地生态系统的地上、地下是相互联系的。植物与土壤微生物作为陆地生态系统中的重要组成部分, 它们之间的相互作用是生态系统地上、地下结合的重要纽带。该文首先介绍了植物在养分循环中对营养元素的吸收、积累和归还等作用, 阐述了土壤微生物对养分有效性及土壤质量具有重要的作用。其次, 重点综述了植物与土壤微生物之间相互依存、相互竞争的关系。植物通过其凋落物与分泌物为土壤微生物提供营养, 土壤微生物作为分解者提供植物可吸收的营养元素, 比如共生体菌根真菌即可使植物根与土壤真菌达到互惠。然而, 植物的养分吸收与微生物的养分固持同时存在, 因而两者之间存在对养分的竞争。通过植物多样性对土壤微生物多样性的影响分析, 以及土壤微生物直接或间接作用于植物多样性和生产力的分析,探讨了植物物种多样性与土壤微生物多样性之间的内在联系。针对当前植物与土壤微生物对养分循环的调控机制的争论, 提出植物凋落物是调节植物与土壤微生物养分循环的良好媒介, 植物与土壤微生物的共同作用对维持整个生态系统的稳定性具有重要意义。也指出了目前在陆地生态系统地上、地下研究中存在的不足和亟待解决的问题。

Microbes isolated from hyperaccumulating plants have been reported to be effective in achieving higher phytoextraction efficiency. The plant growth-promoting bacteria (PGPB) SaMR12 from the cadmium (Cd)/zinc hyperaccumulator Sedum alfredii Hance could promote the growth of a non-host plant, oilseed rape, under Cd stress. However, the effect of SaMR12 on Brasscia juncea antioxidative response under Cd exposure was still unclear.A hydroponic experiment was conducted to study the effects of Sphingomonas SaMR12 on its non-host plant Brassica juncea (L.) Czern. under four different Cd treatments. The results showed that SaMR12 could colonize and aggregate in the roots and then move to the shoots. SaMR12 inoculation promoted plant growth by up to 71% in aboveground biomass and 81% in root biomass over that of the non-inoculated plants. SaMR12-inoculated plants significantly enhanced root Cd accumulation in the 10 and 20 μM Cd treatments, with 1.72- and 0.86-fold increases, respectively, over that of the non-inoculated plants. SaMR12 inoculation not only decreased shoot hydrogen peroxide (HO) content by up to 38% and malondialdehyde (MDA) content by up to 60% but also reduced proline content by 7-30% in shoots and 17-32% in roots compared to the levels in non-inoculated plants. Additionally, SaMR12 inoculation promoted the activities of superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), and ascorbate peroxidase (APX) and facilitated the relative gene expression levels of dehydroascorbate reductase (DHAR) and glutathione reductase (GR) involved in the glutathione (GSH)-ascorbic acid (AsA) cycle.The results demonstrated that, under Cd stress, SaMR12 inoculation could activate the antioxidative response of B. juncea by decreasing the concentrations of HO, MDA and proline, increasing the activities of antioxidative enzymes, and regulating the GSH-AsA cycle. These results provide a theoretical foundation for the potential application of hyperaccumulator endophytic bacteria as remediating agents to improve heavy metal tolerance within non-host plant species, which could further improve phytoextraction efficiency.

Copper is a micronutrient required for living organisms, but is potentially toxic in excess. EDDS enhances the phytoextraction of many metals, but the underlying mechanism is fully unclear. Exposure of 200 μM Cu2+ for 3 days resulted in rice seedling growth inhibition, accompanied by a decrease in plasma membrane H+-ATPase activity, and an increase in relative electrolyte leakage ratios, indicating that maintaining of membrane structure integrity is crucial in acclimation of plants to heavy metal stress. In addition, the chlorophyll and carotenoid content was markedly decreased and the level of the mRNA of Cytochrome P450 gene, OsHMA9, the sulfate transporter gene, and the metallothionein-like protein gene was observed to increase in response to Cu stress. Cu treatment also induced a global epigenetic response which is associated with cell nucleus condensation. These physiological, genetic, and epigenetic responses of rice seedlings to excess copper were modified by the addition of EDDS, suggesting that the supply of EDDS in medium containing a high concentration of Cu ions could enhance plant tolerance potential to excess Cu toxicity through alleviating Cu-induced poisonous effects at various levels.

Most studies on chelate-induced phytoextraction have focused on EDTA-mediated Pb phytoextraction. But EDTA and the formed EDTA-Pb complexes have low biodegradability and high solubility in soil, resulting in an elevated risk of adverse environmental effects. EDDS is an easily biodegradable chelating agent that has recently been proposed as an environmentally sound alternative to EDTA. Consequently, a greenhouse experiment, using a completely randomized factorial design with four replications, was carried out to compare the potential of EDTA and EDDS for chelate-induced Pb phytoextraction with Cynara cardunculus, as well as to investigate the toxicity of these two chelates to both cardoon plants and soil microorganisms. The effects of chelate addition on soil microbial communities were studied through the determination of a variety of biological indicators of soil quality such as soil enzyme activities, basal and substrate-induced respiration, potentially mineralizable nitrogen, and community level physiological profiles. EDTA was much more efficient than EDDS for the enhancement of root Pb uptake and root-to-shoot Pb translocation. In a soil polluted with 5000 mg Pb kg(-1), as a result of the addition of 1 g EDTA kg(-1) soil, a value of 1332 mg Pb kg(-1) DW shoot was obtained. EDDS application resulted in a shoot Pb accumulation of only 310 mg kg(-1)DW. Plants treated with EDDS showed lower values of biomass than those treated with EDTA. EDDS proved to be rapidly degraded, and less toxic to the soil microbial community in control non-polluted soils. Pb-polluted EDDS-treated soils showed significantly higher values of basal and substrate-induced respiration than those treated with EDTA. Although EDDS had a lower capacity to enhance Pb phytoextraction than EDTA, it has the advantage of rapid biodegradation.

Pb is one of the most widespread and metal pollutants in soil. It is generally concentrated in surface layers with only a minor portion of the total metal found in soil solution. Phytoextraction has been proposed as an inexpensive, sustainable, in situ plant-based technology that makes use of natural hyperaccumulators as well as high biomass producing crops to help rehabilitate soils contaminated with heavy metals without destructive effects on soil properties. The success of phytoextraction is determined by the amount of biomass, concentration of heavy metals in plant, and bioavailable fraction of heavy metals in the rooting medium. In general, metal hyperaccumulators are low biomass, slow growing plant species that are highly metal specific. For some metals such as Pb, there are no hyperaccumulator plant species known to date. Although high biomass-yielding non-hyperaccumulator plants lack an inherent ability to accumulate unusual concentrations of Pb, soil application of chelating agents such as EDTA has been proposed to enhance the metal concentration in above-ground harvestable plant parts through enhancing the metal solubility and translocation from roots to shoots. Leaching of metals due to enhanced mobility during EDTA-assisted phytoextraction has been demonstrated as one of the potential hazards associated with this technology. Due to environmental persistence of EDTA in combination with its strong chelating abilities, the scientific community is moving away from the use of EDTA in phytoextraction and is turning to less aggressive alternative strategies such as the use of organic acids or more degradable APCAs (aminopolycarboxylic acids). We have therefore arrived at a point in phytoremediation research history in which we need to distance ourselves from EDTA as a proposed soil amendment within the context of phytoextraction. However, valuable lessons are to be learned from over a decade of EDTA-assisted phytoremediation research when considering the implementation of more degradable alternatives in assisted phytoextraction practices.

近30年来，过量施用磷肥导致土壤磷素累积继而引起水体富营养化等问题备受关注。植物根系分泌的低分子量有机酸能活化土壤积累态磷，提高土壤磷素有效性，已成为研究热点问题之一。本文结合国内外已有研究，从低分子量有机酸的类型、添加浓度、土壤类型和土壤磷素水平等方面总结了低分子量有机酸活化土壤无机磷与有机磷的效果，并通过对比土壤磷活化试验前后各形态磷的变化探讨了低分子量有机酸对土壤磷的活化机制。低分子量有机酸对土壤无机磷的活化主要是促进了土壤中有效性低的无机磷形态向有效性较高的形态转化，而低分子量有机酸对土壤有机磷的活化结论尚不一致，活化机制也不明确，仍需进一步研究。未来研究应关注低分子量有机酸与磷肥之间的协同增效机制，并进一步探索低分子量有机酸对土壤磷素（特别是有机磷）的活化机制。