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。     

植物重金属超富集机理研究进展

植物超富集重金属机理主要涉及植物对金属离子高的吸收、运输能力,区域化作用及螯合作用等方面,其中跨膜运载蛋白的表达、调控对重金属超富集这一特性起了关键作用。金属阳离子运载蛋白家族主要包括CDF家族、NRAMP家族和ZIP家族等,在超富集植物中已克隆出多个家族的金属运载蛋白基因,这些基因的过量表达对重金属在细胞中的运输、分布和富集及提高植物的抗性方面发挥了重要作用。综述了近年来研究重金属超富集植物吸收、转运和贮存Zn、Ni、Cd等重金属的生理和分子机制所取得的主要进展。     

植物超积累重金属的生理机制研究进展

土壤重金属污染已经成为一个全球性问题。重金属超积累植物在修复土壤重金属污染中具有重要的应用前景。重金属超积累植物通常具备三个基本特征,即：根系具有从土壤中吸收重金属的强大能力、能从根到地上部分高效转运重金属、在叶片中能解毒和隔离大量重金属。本文总结了重金属超积累植物吸收、转运、隔离和解毒重金属的生理机制研究进展,以期为进一步阐明植物超积累重金属的机制及其在植物修复中的应用提供参考。     

Effect of arbuscular mycorrhizal fungal inoculation on heavy metal accumulation of maize grown in a naturally contaminated soil

A pot culture experiment was carried out to study heavy metal (HM) phytoaccumulation from soil contaminated with Cu, Zn, Pb, and Cd by maize (Zea mays L.) inoculated with arbuscular mycorrhizal (AM) fungi (AMF). Two AM fungal inocula--MI containing only one AM fungal strain (Glomus caledonium 90036) and MII consisting of Gigaspora margarita ZJ37, Gigaspora decipens ZJ38, Scutellospora gilmori ZJ39, Acaulospora spp., and Glomus spp.--were applied to the soil under unsterilized conditions. The control received no mycorrhizal inoculation. The maize plants were harvested after 10 wk of growth. MI-treated plants had higher mycorrhizal colonization than MII-treated plants. Both MI and MII increased P concentrations in roots, but not in shoots. Neither MI nor MII had significant effects on shoot or root dry weight (DW). Compared with the control, shoot Cu, Zn, Pb, and Cd concentrations were decreased by MI but increased by MII. Cu, Zn, Pb, and Cd uptake into shoots and roots all increased in MII-treated plants, while in MI-treated plants Cu, Zn, and Pb uptake into shoots and Cd uptake into roots decreased but Cu, Zn, and Pb uptake into roots and Cd into shoots increased. MII was more effective than MI in promoting HM extraction efficiencies. The results indicate that MII can benefit HMphytoextraction and, therefore, show potential in the phytoremediation of HM-contaminated soils.     

超富集植物吸收富集重金属的生理和分子生物学机制

与普通植物相比,超富集植物在地上部富集大量重金属离子的情况下可以正常生长,其富集重金属的机理已经成为当前植物逆境生理研究的热点领域．尤其是近两年,随着分子生物学等现代技术手段的引人,关于重金属离子富集机理的研究取得了一定进展．通过与酵母突变株功能互补克隆到了多条编码微量元素转运蛋白的全长cDNA;也从分子水平上研究了谷胱甘肽、植物螯合素、金属硫蛋白、有机酸或氨基酸等含巯基物质与重金属富集之间的可能关系．本文从植物生理和分子生物学角度简要评述超富集植物对重金属元素的吸收、富集、整合及区室化的机制．     

Effect of simultaneous establishment of Sedum alfredii and Zea mays on heavy metal accumulation in plants

Land application of biosolids to improve agricultural productivity is a cost-effective approach for resource recovery. Unfortunately, municipal biosolids often contain high concentrations of heavy metals, including zinc and copper. In this study, a co-cropping technique was investigated using a known zinc hyperaccumulator, Sedum alfredii with a grain crop, Zea mays. After a 3-mo growth trial, the results indicate that when Z. mays is co-cropped with S. alfredii, heavy metals accumulated in the grains were significantly reduced when compared to monoculture cropping. Co-cropping improved the growth of both plant species. In addition, the biosolids maintained stable pH, N-P-K concentrations, germination potential, and water content after the plant treatment, regardless of the plant species used in the trial. In conclusion, co-cropping with hyperaccumulators may be an effective approach to reducing the risk of contaminant uptake in edible crops.     

超富集植物对重金属耐受和富集机制的研究进展

超富集植物对重金属耐受和富集机制的研究成为近年来植物逆境生理研究的热点,在简要总结细胞壁沉淀、重金属螯合效应、酶活性机制和细胞区室化作用的基础上,概述了超富集植物对重金属的耐受机制,讨论了重金属跨根细胞质膜运输,共质体内运输、木质部运输和跨叶细胞膜运输的富集过程。     

水体重金属污染植物修复研究进展

植物修复技术作为一种废水处理新技术,具有投资少、效果好、产出高、环境效益好等优点,现已得到迅速发展及广泛应用。介绍了水生超富集植物的3种生态类型,造成水体污染重金属元素类型,植物应答重金属污染的检测方法,并对植物修复水体重金属污染的发展前景提出展望。     

Phospholipid transfer protein deficiency ameliorates diet-induced hypercholesterolemia and inflammation in mice

Phospholipid transfer protein (PLTP) facilitates the transfer of phospholipids from triglyceride-rich lipoproteins into HDL. PLTP has been shown to be an important factor in lipoprotein metabolism and atherogenesis. Here, we report that chronic high-fat, high-cholesterol diet feeding markedly increased plasma cholesterol levels in C57BL/6 mice. PLTP deficiency attenuated diet-induced hypercholesterolemia by dramatically reducing apolipoprotein E-rich lipoproteins (-88%) and, to a lesser extent, LDL (-40%) and HDL (-35%). Increased biliary cholesterol secretion, indicated by increased hepatic ABCG5/ABCG8 gene expression, and decreased intestinal cholesterol absorption may contribute to the lower plasma cholesterol in PLTP-deficient mice. The expression of proinflammatory genes (intercellular adhesion molecule-1 and vascular cell adhesion molecule-1) is reduced in aorta of PLTP knockout mice compared with wild-type mice fed either a chow or a high-cholesterol diet. Furthermore, plasma interleukin-6 levels are significantly lower in PLTP-deficient mice, indicating reduced systemic inflammation. These data suggest that PLTP appears to play a proatherogenic role in diet-induced hyperlipidemic mice.     

粤东铅锌尾矿三种优势植物对重金属的吸收和富集特性研究

为探讨铅锌矿废弃地优势植物在重金属污染土壤植物修复中的应用潜力,利用野外采样分析法,从粤东梅县丙村铅锌尾矿区采集其三种优势植物类芦、黄荆、盐肤木的根、茎、叶和土壤样品,测定和分析Pb、Zn、Cu、Cd四种重金属含量.结果表明:该矿区土壤污染严重,Pb、Zn、Cd含量远超土壤环境质量的三级标准,Cu超出二级标准;根际土壤和非根际土壤重金属含量均为Pb＞Zn＞Cu＞Cd,但根际土壤的重金属含量显著低于非根际土壤;这三种植物对Pb、Zn、Cu的转移系数大于1.0,对Cu的富集系数最高,Pb最小,但对四种重金属的富集系数均小于1.0,均未达到超富集植物临界含量标准.三种植物为该矿区的优势植物,说明它们对土壤的重金属污染有很强的耐性,虽然并非典型的超富集植物,但对污染土壤仍有较好的修复效果.     

Heterogeneity of high density lipoprotein generated by ABCA1 and ABCA7

The assembly of HDL by helical apolipoprotein and cellular lipid was studied using HEK293 cells to which ecdysone-inducible human ABCA1 or human ABCA7 was transfected. Expression of both ABCA1 and ABCA7 was induced linearly proportional to ponasterone A concentration in the medium. In the experimental conditions used, the ABC protein expression levels limited the rate of lipid release when the apolipoprotein concentration was high, and the apolipoprotein concentration was rate-limiting when the ABC protein expression levels were high. When ABCA1 expression increased in conditions in which it was rate-limiting, relative cholesterol content to phospholipid increased in the HDL produced. In contrast, it was constant when ABCA7 expression increased. To investigate the background mechanism, the HDL particles were analyzed by density gradient ultracentrifugation and high performance lipid chromatography. The ABCA1-mediated reaction produced two distinct HDLs, large cholesterol-rich and small cholesterol-poor particles, and the ABCA7-mediated reaction generated mostly small cholesterol-poor particles. The increase of HDL assembly with the increase of ABCA1 expression was predominant in large cholesterol-rich particles, whereas only small cholesterol-poor HDL increased as ABCA7 expression increased. We conclude that ABCA1 generates cholesterol-rich and cholesterol-poor HDL and that the former is more prominently dependent on the increase of ABCA1 expression. ABCA7 produces this HDL subfraction only as a very minor component.     

Structure and metal binding properties of ZnuA, a periplasmic zinc transporter from Escherichia coli

ZnuA is the periplasmic Zn²⁺-binding protein associated with the high-affinity ATP-binding cassette ZnuABC transporter from Escherichia coli. Although several structures of ZnuA and its homologs have been determined, details regarding metal ion stoichiometry, affinity, and specificity as well as the mechanism of metal uptake and transfer remain unclear. The crystal structures of E. coli ZnuA (Eco-ZnuA) in the apo, Zn²⁺-bound, and Co²⁺-bound forms have been determined. ZnZnuA binds at least two metal ions. The first, observed previously in other structures, is coordinated tetrahedrally by Glu59, His60, His143, and His207. Replacement of Zn²⁺ with Co²⁺ results in almost identical coordination geometry at this site. The second metal binding site involves His224 and several yet to be identified residues from the His-rich loop that is unique to Zn²⁺ periplasmic metal binding receptors. Electron paramagnetic resonance and X-ray absorption spectroscopic data on CoZnuA provide additional insight into possible residues involved in this second site. The second site is also detected by metal analysis and circular dichroism (CD) titrations. Eco-ZnuA binds Zn²⁺ (estimated K _d < 20 nM), Co²⁺, Ni²⁺, Cu²⁺, Cu⁺, and Cd²⁺, but not Mn²⁺. Finally, conformational changes upon metal binding observed in the crystal structures together with fluorescence and CD data indicate that only Zn²⁺ substantially stabilizes ZnuA and might facilitate recognition of ZnuB and subsequent metal transfer. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

构树的污泥适应性及养分和重金属吸收累积特征

构树是我国重要速生经济树种,具有适应性强、生物量大和重金属富集能力强等优点,而污泥中含有大量养分和重金属,在污泥中种植构树有望同时实现污泥生态修复和构树资源生产。本研究通过盆栽试验,分析在对照(赤红壤)、50%污泥(污泥、赤红壤混合基质,重量比各50%)和100%污泥基质中构树生长及不同部位(根、茎、叶)养分和重金属吸收累积特征,并通过主成分分析和隶属度函数对吸收累积能力进行综合评价。结果表明: 构树在50%和100%污泥中均可正常生长且株高、生物量显著高于对照,在100%污泥中长势最好,质量指数(1.02)分别是对照和50%污泥处理的4.3和2.4倍。50%和100%污泥处理构树各部位N含量和茎P含量显著高于对照,100%污泥处理构树茎、叶K含量显著低于对照。构树对Cu、Zn、Pb、Cd、Ni的吸收部位以根为主,根系重金属含量与污泥比例呈正相关,叶Pb、Cd含量符合《饲料卫生标准》(GB 13078—2017)。构树对Cd的吸收累积效果好于其他重金属元素。与对照相比,50%和100%污泥处理构树根部Zn、Pb、Cd滞留率显著提高(57.8%~85.8%),100%污泥处理构树根部Cu、Ni滞留率显著提高(67.5%和74.8%)。污泥处理全株养分和重金属累积量均显著大于对照,其中100%污泥处理显著大于50%污泥处理。与50%污泥处理相比,100%污泥处理构树各部位及全株养分和重金属累积量大幅提高。不同处理下构树污泥适应性和元素吸收累积的综合评价得分为100%污泥(0.848)>50%污泥(0.344)>对照(0.080)。构树对污泥具有良好的适应性,在纯污泥中能够正常生长并具有较强的吸收累积养分和重金属能力,可在修复污泥的同时进行构树资源生产。     

Ezetimibe normalizes metabolic defects in mice lacking ABCG5 and ABCG8

The ATP binding cassette transporters ABCG5 (G5) and ABCG8 (G8) limit the accumulation of neutral sterols by restricting sterol uptake from the intestine and promoting sterol excretion into bile. Humans and mice lacking G5 and G8 (G5G8-/-) accumulate plant sterols in the blood and tissues. However, despite impaired biliary cholesterol secretion, plasma and liver cholesterol levels are lower in G5G8-/- mice than in wild-type littermates. To determine whether the observed changes in hepatic sterol metabolism were a direct result of decreased biliary sterol secretion or a metabolic consequence of the accumulation of dietary noncholesterol sterols, we treated G5G8-/- mice with ezetimibe, a drug that reduces the absorption of both plant- and animal-derived sterols. Ezetimibe feeding for 1 month sharply decreased sterol absorption and plasma levels of sitosterol and campesterol but increased cholesterol in both the plasma (from 60.4 to 75.2 mg/dl) and the liver (from 1.1 to 1.87 mg/g) of the ezetimibe-treated G5G8-/- mice. Paradoxically, the increase in hepatic cholesterol was associated with an increase in mRNA levels of HMG-CoA reductase and synthase. Together, these results indicate that pharmacological blockade of sterol absorption can ameliorate the deleterious metabolic effects of plant sterols even in the absence of G5 and G8.     

重金属胁迫对毛竹种子萌发及其富集效应的影响

以毛竹种子为供试材料,研究4种重金属(Pb2+、Zn2+、Cu2+、Cd2+)胁迫对毛竹种子萌发的影响,并考察重金属在毛竹幼苗各组织部分的富集情况。结果表明:(1)Pb2+和Cd2+对毛竹种子的发芽率、发芽势、发芽指数及活力指数有抑制作用,低浓度下Cu2+和Zn2+对毛竹种子的发芽势、发芽率、发芽指数等指标有促进作用,高浓度则显著抑制;当浓度达到1600μmol/L时Cd2+对种子萌发的抑制效果明显强于其他3种元素;(2)选取根尖数、根表面积、根体积、根系总长4个根系形态指标发现,低浓度处理下Pb2+、Zn2+对根系生长有促进作用,而Cu2+和Cd2+起到明显的抑制作用;(3)处理10d后,种子萌发幼苗地上部对Pb2+、Zn2+、Cu2+、Cd2+的含量最高可达6810.51、1387.77、951.77、429.33 mg/kg,转移系数Zn2+Cd2+Pb2+Cu2+。综上,系统揭示了毛竹种子在重金属胁迫下的萌发和富集情况,为今后的土培、大田试验提供了有益的参考,也为将毛竹作为植物修复材料加以研究开启了新的研究视角,具有重要的研究价值。     

Enhancement of heavy metal tolerance and accumulation efficiency by expressing Arabidopsis ATP sulfurylase gene in alfalfa

Abstract

Transgenic alfalfa (Medicago sativa L.) plants overexpressing the Arabidopsis ATP sulfurylase gene were generated using Agrobacterium-mediated genetic transformation to enhance their heavy metal accumulation efficiency. The ATP sulfurylase gene was cloned from Arabidopsis, following exposure to vanadium (V) and lead (Pb), and transferred into an Agrobacterium tumefaciens binary vector. This was co-cultivated with leaf explants of the alfalfa genotype Regen SY. Co-cultivated leaf explants were cultured on callus and somatic embryo induction medium, followed by regeneration medium for regenerating complete transgenic plants. The transgenic nature of the plants was confirmed using PCR and southern hybridization. The expression of Arabidopsis ATP sulfurylase gene in the transgenic plants was evaluated through RT-PCR. The selected transgenic lines showed increased tolerance to a mixture of five heavy metals and also demonstrated enhanced metal uptake ability under controlled conditions. The transgenic lines were fertile and did not exhibit any apparent morphological abnormality. The results of this study indicated an effective approach to improve the heavy metal accumulation ability of alfalfa plants which can then be used for the remediation of contaminated soil in arid regions.     

重金属超富集植物及植物修复技术研究进展

植物修复技术（Phytoremediation)是近年来发展起来的一种主要用于清除土壤重金属污染的绿色生态技术,重金属超富集植物（hyperaccumulator)及植物修复技术是当前学术界研究的热点领域,目前虽已有Cd、Co、Cr、Cu、Mn、Ni、Pb、Zn等超富集植物发现的报道,但尚无一例报道来自于中国,中国具有广袤的国土面积、丰富的植物类型和多种（处）古老的矿山开采与冶炼场所,在中国开展超富集植物的寻找,研究与开发工作,将会有重要突破,并具有重要的理论与实践意义,本文拟就国内外在这一领域的研究进展作一简要综述。     

ATP binding cassette transporter A1 - key roles in cellular lipid transport and atherosclerosis

ATP-binding cassette transporter A1 (ABCA1) was recently recognized as the mutant molecule responsible for Tangier disease with low HDL levels, accumulation of cholesteryl esters in tissues, and increased risk of cardiovascular disease. Extensive studies for the past 2 years have recognized the critical role of ABCA1 in cholesterol and phospholipid trafficking. Since the removal of cholesterol from tissues is a key step in the prevention of atherosclerosis, significant attention has been focused on this molecule. Natural ABCA1 mutations in Tangier disease (TD) patients and WHAM chickens together with induced mutation in ABCA1 knock-out mice unequivocally established the important role of ABCA1 in maintaining circulating HDL levels and promoting cholesterol efflux from the arterial wall. Mice lacking ABCA1 showed similar phenotypes observed in Tangier disease patients with low levels of HDL. Further understanding of the roles of ABCA1 in lipid transport and atherosclerosis became clear from studies with ABCA1 transgenic mice. These mice showed enhanced cholesterol efflux from macrophages and reduced atherosclerotic lesion formation. The promoter of the ABCA1 gene has been mapped to a large extent, with the exception of cAMP response element. The present review summarizes recent developments on the role of ABCA1 in cholesterol efflux and prevention of atherosclerosis. Given the antiatherogenic properties of ABCA1, this molecule can serve as an appropriate target for developing drugs to treat individuals with low levels of HDL.     

Phytochelatins and heavy metal tolerance

The induction and heavy metal binding properties of phytochelatins in heavy metal tolerant (Silene vulgaris) and sensitive (tomato) cell cultures, in water cultures of these plants and in Silene vulgaris grown on a medieval copper mining dump were investigated. Application of heavy metals to cell suspension cultures and whole plants of Silene vulgaris and tomato induces the formation of heavy metal–phytochelatin-complexes with Cu and Cd and the binding of Zn and Pb to lower molecular weight substances. The binding of heavy metal ions to phytochelatins seems to play only a transient role in the heavy metal detoxification, because the Cd- and Cu-complexes disappear in the roots of water cultures of Silene vulgaris between 7 and 14 days after heavy metal exposition. Free heavy metal ions were not detectable in the extracts of all investigated plants and cell cultures. Silene vulgaris plants grown under natural conditions on a mining dump synthesize low molecular weight heavy metal binding compounds only and show no complexation of heavy metal ions to phytochelatins. The induction of phytochelatins is a general answer of higher plants to heavy metal exposition, but only some of the heavy metal ions are able to form stable complexes with phytochelatins. The investigation of tolerant plants from the copper mining dump shows that phytochelatins are not responsible for the development of the heavy metal tolerant phenotypes.     

重金属污染土壤的湿地生物修复技术

生物修复技术是近年来发展起来的一种有前途的污染治理技术,重金属的湿地植物稳定技术和植物萃取技术等植物修复技术是一研究热点;同时,针对微生物对重金属的生物积累、生物转化及生物修复作一分析,重金属的生物修复生理机制及其提高富集效率的条件作一综述,以期推动国内这一国际热点领域的研究。