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。     

超富集植物遏蓝菜对重金属吸收、运输和累积的机制

遏蓝菜Thlaspi caerulescens可以在其地上部累积大量重金属如锌、镉等,是公认的超富集植物。由于该植物生物量小,不宜直接用于重金属污染的土壤植物修复,而被广泛作为一种模式植物来进行重金属富集机制研究。遏蓝菜对重金属离子的累积大致经过螯合剂解毒、地上部长距离运输以及在液泡中的储存等生理过程。已经发现的植物体内的金属螯合剂——有机酸、氨基酸、植物络合素(PCs)、金属硫蛋白(MT)和尼克烟酰胺NA等,区室化以及长距离运输相关的转运蛋白——ZIP(ZRT/IRTlike protein)、CDF(Cation diffusion facilitator)、Nramp(Natural resistance and macrophage protein)和HMA(Heavy metal ATPase)等家族,以上各种基因、多肽与蛋白等共同参与了植物对金属累积与耐受过程并发挥各自重要的作用。以下主要介绍了遏蓝菜重金属超富集相关的基因、多肽和蛋白,以及它们在重金属螯合作用和运输过程中的功能。     

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．     

重金属超富集植物的研究与利用

阐述了超富集植物的定义、特性及研究历史和现状,介绍了国内外应用超富集植物修复重金属污染土壤的实例,指出了超富集植物研究存在的问题和发展方向。     

几种重金属(Pb、Zn、Cd、Cu)的超富集植物种类及增强植物修复措施研究进展

近年来土壤重金属污染问题越来越严重, 植物修复技术以其安全、廉价的特点正成为研究和开发的热点, 国内外对利用超富集植物来修复土壤重金属污染的研究已有大量报道。对超富集植物概念的提出及超富集植物吸收富集重金属的机理进行了归纳总结, 主要就铅、锌、镉和铜四种重金属超富集植物的相关研究进展进行了分类、归纳与总结, 同时还对增强植物修复效果的措施进行了探讨, 以期为进一步合理有效利用植物修复土壤主要重金属污染提供一定的参考依据。     

超富集植物重金属吸收转运机制的研究进展

简要概述了近年来重金属超富集植物在重金属吸收、转运和耐毒机制的一些进展．以及抗生物胁迫的可能机制。     

废弃重金属的植物修复与再利用研究进展

介绍了重金属污染的植物修复方法,通过分析超富集植物对重金属的耐受机制,对超富集植物回收利用重金属进行了总结,最后讨论并展望了重金属植物修复未来的研究方向和热点。     

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.     

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。     

赣南钨矿区土壤重金属含量与植物富集特征

对赣州大余县境内四大国有钨矿(西华山、荡坪、漂塘、下垄)的尾砂库区土壤和植物重金属含量进行了分析,结果表明:尾砂库区土壤受到重金属Zn、Cd、Mo、Cu、Pb与W的污染,而且Cd和Mo含量较高;在4个尾砂库区中,下垄矿区尾砂库的重金属污染比其他3个尾砂库严重.4个尾砂库区共出现了53种植物,隶属31科52属,这些植物重金属富集系数的高低顺序为Zn＞Cd＞Mo＞Cu＞Pb＞W.另外,植物不同的耐性机制使它们对重金属的富集表现出不同特性,芒箕、龙葵、酸模等植物地上部富集较多重金属,可用于污染土壤的植物修复;乌毛蕨、梵天花和狗脊蕨等在根部富集较多重金属,可用于植物固化技术;狗尾草、鬼针草、白苏富集极少重金属,可作为矿区废弃地植被重建的先锋物种.     

Molecular physiology of aluminum toxicity and tolerance in plants

Aluminum being the third most abundant metal in the earth’s crust poses a serious threat to crop productivity in acid soils, which comprise almost half of the arable land. This review travels across time and updates research done on aluminum stress in plants. In its phytotoxic forms, aluminum affects root growth by acting in the root apical zone, resulting in growth inhibition in a very short time at micromolar concentrations. The mechanisms of aluminum toxicity in plants may proceed by growth inhibition, callose accumulation, cytoskeletal distortion, disturbance of plasma membrane surface charge, and H⁺-ATPase activity, lipid peroxidation of membranes, production of reactive oxygen species in cytosol and mitochondria, respiratory dysfunction, opening of mitochondrial permeability transition pores, collapsing of inner mitochondrial membrane potential, activation of mitochondrial protease, and induction of nuclear apoptosis, resulting ultimately in programmed cell death. In contrast, the mechanism of tolerance involves the exudation of organic acid anions, complexation of aluminum with organic acids, and subsequent detoxification. Many oxidative stress genes and other metabolically important genes have also been found to be induced under aluminum stress and overexpression analyses have also shown some plants to develop some degree of tolerance. In the future, researchers in the area of aluminum research should investigate more basic mechanisms of aluminum toxicity and discover and study more aluminum-responsive genes that confer resistance against this toxic metal, to ensure food security for ever-increasing human populations in the future.     

铅胁迫下植物抗性机制的研究进展

综述了近10多年来国内外研究重金属铅对植物的伤害机理,以及植物对铅的耐性机制的研究进展.首先从细胞分裂、细胞膜透性、光合作用和抗氧化酶系统等生理生化方面讨论了铅对植物的伤害机理,其次分析了铅离子跨膜运输、根系分泌物、金属配位体、铅离子区域化分布与植物抗性的关系.     

Cadmium absorption and transportation pathways in plants

Controlling the uptake, transport, translocation, and accumulation of excessive amounts of cadmium from polluted environments is critical for plants and, consequently, humans with regard to food safety. Plants adopt various cellular and molecular mechanisms to minimize Cd toxicity. Upon exposure to Cd, plants initially implement avoidance strategies, such as production of organic acids, chelation, and sequestration, to prevent metal access to root cells. Nevertheless, Cd can be transported through the roots, stems, and leaves via apoplastic and symplastic pathways. These processes have been controlled by specific sites at the root surface and root cortex, in cells responsible for loading the root xylem, at the transition between the vascular systems of the root and the shoot, and in connecting tissues and cells at the stem. Although resistance to heavy metal cadmium can be achieved by either avoidance or tolerance, genetic basis to tolerance is therefore implied, in that these mechanisms are heritable attributes of tolerant mutants or genotypes.     

植物耐重金属机理研究进展

由于工业“三废”和机动车尾气的排放、污水灌溉及农药、除草剂和化肥的使用,严重地污染了土壤、水质和大气,其中土壤中的重金属（Ｈｇ、Ｃｄ、Ａｓ、Ｃｕ和Ａｌ）污染更为严重［１］。重金属在植物根、茎、叶及籽粒中的大量累积,不仅严重地影响植物的生长和发育［１～...     

Direct measurement of aluminum uptake and distribution in single cells of Chara corallina

Quantitative information on the uptake and distribution of Al at the cellular level is required to understand mechanisms of Al toxicity, but direct measurement of uptake across the plasma membrane has remained elusive. We measured rates of Al transport across membranes in single cells of Chara corallina using the rare (26)Al isotope, an emerging technology (accelerator mass spectrometry), and a surgical technique for isolating subcellular compartments. Accumulation of Al in the cell wall dominated total uptake (71-318 microgram m(-2) min(-1)), although transport across the plasma membrane was detectable (71-540 ng m(-2) min(-1)) within 30 min of exposure. Transport across the tonoplast was initially negligible, but accelerated to rates approximating uptake across the plasma membrane. The avacuolate protoplasm showed signs of saturation after 60 min, but continued movement across the plasma membrane was supported by sequestration in the vacuole. Saturation of all compartments was observed after 12 to 24 h. Accumulation of Al in the cell wall reflected variation in [Al(3+)] induced by changes in Al supply or complexing ligands, but was unaffected by pH. In contrast, transport across the plasma membrane peaked at pH 4.3 and increased when [Al(3+)] was reduced by complexing ligands. Cold temperature (4 degrees C) reduced accumulation in the cell wall and protoplasm, whereas 2,4-dinitrophenol and m-chlorocarbonylcyanidephenyl hydrazone increased membrane transport by 12- to 13-fold. Our data suggest that the cell wall is the major site of Al accumulation. Nonetheless, membrane transport occurs within minutes of exposure and is supported by subsequent sequestration in the vacuole. The rapid delivery of Al to the protoplasm suggests that intracellular lesions may be possible.     

湿地植物根形态结构和泌氧与盐和重金属吸收、积累、耐性关系的研究进展

目前大面积湿地面临着重金属污染和盐渍化问题。利用湿地植物修复这些受损生态系统和提高海水稻的产量、减少毒性金属元素在稻米中的积累是当前面临的重要任务。湿地植物(包括水稻)已发展出各种策略和机制来耐受不同的环境胁迫,它们的根系发育具有可塑性,如根形态和解剖结构会随外界条件的变化而变化,这些变化直接影响其对环境胁迫的适应性能。近年来,对湿地植物根形态和结构、泌氧与其对盐、重金属的吸收、积累和耐性之间的关系方面进行了一些重要研究。本文分别对湿地植物根系形态、质外体屏障、通气组织和泌氧与其对盐和重金属吸收、积累和耐性的关系等方面的研究进展进行了综述,并对该领域未来的发展方向作了展望。     

Recognition and transport of ferric enterobactin in Escherichia coli.

The specificity of the outer membrane protein receptor for ferric enterobactin transport in Escherichia coli and the mechanism of enterobactin-mediated transport of ferric ions across the outer membrane have been studied. Transport kinetic and inhibition studies with ferric enterobactin and synthetic structural analogs have mapped the parts of the molecule important for receptor binding. The ferric complex of the synthetic structural analog of enterobactin, 1,3,5-N,N',N'-tris-(2,3-dihydroxybenzoyl)triaminomethylbenzene (MECAM), was transported with the same maximum velocity as was ferric enterobactin. A double-label transport assay with [59Fe, 3H]MECAM showed that the ligand and the metal are transported across the outer membrane at an identical rate. Under the growth conditions used, large fractions of the transported complexes were available for exchange across the outer membrane when a large excess of extracellular complex was added to the cell suspension; at least 60% of the internalized [59Fe]enterobactin exchanged with extracellular [55Fe]enterobactin. Internalized [59Fe, 3H]MECAM was released from the cell as the intact complex when either unlabeled Fe-MECAM or Fe-enterobactin was added extracellularly. The results suggest a mechanism of active transport of unmodified coordination complex across the outer membrane with possible accumulation in the periplasm.     

The rice OsNAC6 transcription factor orchestrates multiple molecular mechanisms involving root structural adaptions and nicotianamine biosynthesis for drought tolerance

Drought has a serious impact on agriculture worldwide. A plant's ability to adapt to rhizosphere drought stress requires reprogramming of root growth and development. Although physiological studies have documented the root adaption for tolerance to the drought stress, underlying molecular mechanisms is still incomplete, which is essential for crop engineering. Here, we identified OsNAC6‐mediated root structural adaptations, including increased root number and root diameter, which enhanced drought tolerance. Multiyear drought field tests demonstrated that the grain yield of OsNAC6 root‐specific overexpressing transgenic rice lines was less affected by drought stress than were nontransgenic controls. Genome‐wide analyses of loss‐ and gain‐of‐function mutants revealed that OsNAC6 up‐regulates the expression of direct target genes involved in membrane modification, nicotianamine (NA) biosynthesis, glutathione relocation, 3′‐phophoadenosine 5′‐phosphosulphate accumulation and glycosylation, which represent multiple drought tolerance pathways. Moreover, overexpression of NICOTIANAMINE SYNTHASE genes, direct targets of OsNAC6, promoted the accumulation of the metal chelator NA and, consequently, drought tolerance. Collectively, OsNAC6 orchestrates novel molecular drought tolerance mechanisms and has potential for the biotechnological development of high‐yielding crops under water‐limiting conditions.     

MTP1-dependent Zn sequestration into shoot vacuoles suggests dual roles in Zn tolerance and accumulation in Zn-hyperaccumulating plants

The integral membrane protein Thlaspi goesingense metal tolerance protein 1 (TgMTP1) has been suggested to play an important role in Zn hyperaccumulation in T. goesingense . Here, we show that the TgMTP1 protein is accumulated to high levels at the vacuolar membrane in shoot tissue of T. goesingense . TgMTP1 is likely to act in the transport of Zn into the vacuole, enhancing both Zn accumulation and tolerance. By specifically expressing TgMTP1 in Arabidopsis thaliana shoots, we show that TgMTP1, localized at the vacuolar membrane, can drive the enhanced shoot accumulation of Zn by initiating a systemic Zn deficiency response. The systematic response includes increased expression of Zn transporters ( ZIP3 , ZIP4 , ZIP5 and ZIP9 ) in both shoot and root tissue. Furthermore, shoot-specific accumulation of TgMTP1 at the vacuolar membrane also leads to increased resistance to Zn in A. thaliana , probably through enhanced Zn compartmentalization in the vacuole. Such evidence leads to the conclusion that the high levels of TgMTP1 at the vacuolar membrane in shoot tissue of the Zn hyperaccumulator T. goesingense play a role in both Zn tolerance and enhanced Zn uptake and accumulation, via the activation of a systemic Zn deficiency response.     

The soybean NRAMP homologue,GmDMT1, is a symbiotic divalent metal transporter capable of ferrous iron transport

Iron is an important nutrient in N2-fixing legume root nodules. Iron supplied to the nodule is used by the plant for the synthesis of leghemoglobin, while in the bacteroid fraction, it is used as an essential cofactor for the bacterial N2-fixing enzyme, nitrogenase, and iron-containing proteins of the electron transport chain. The supply of iron to the bacteroids requires initial transport across the plant-derived peribacteroid membrane, which physically separates bacteroids from the infected plant cell cytosol. In this study, we have identified Glycine max divalent metal transporter 1 (GmDmt1), a soybean homologue of the NRAMP/Dmt1 family of divalent metal ion transporters. GmDmt1 shows enhanced expression in soybean root nodules and is most highly expressed at the onset of nitrogen fixation in developing nodules. Antibodies raised against a partial fragment of GmDmt1 confirmed its presence on the peribacteroid membrane (PBM) of soybean root nodules. GmDmt1 was able to both rescue growth and enhance 55Fe(II) uptake in the ferrous iron transport deficient yeast strain (fet3fet4). The results indicate that GmDmt1 is a nodule-enhanced transporter capable of ferrous iron transport across the PBM of soybean root nodules. Its role in nodule iron homeostasis to support bacterial nitrogen fixation is discussed.