Nicotianamine: mediator of transport of iron and heavy metals in the phloem?

Recent work has demonstrated that minerals in plants are circulated between root and shoot. This occurs during the whole life time and renders possible response to changing environmental conditions. This mineral circulation occurs through intensive solute exchange between xylem and phloem in roots, stems, and leaves. The transport form of heavy metals such as iron, manganes, zinc and copper in the phloem, whether ionic or chelated, is unclear in most cases.
The unusual amino acid nicotianamine (NA) is ubiquitous throughout the plant kingdom. It is a chelator of several divalent transition metals. Its physiological role was investigated with the tomato mutant chloronerva, the only known NA-free multicellular plant. The mutant also exhibits disturbances of its iron metabolism and that of other heavy metals. This leads, among others, to a typical intercostal chlorosis and progressive iron accumulation in the leaves. From the heavy metal chelating properties of NA and from the phenotype of the mutant chloronerva it is concluded that NA is needed for normal distribution of heavy metals in young growing tissues fed via the phloem. This function could be fulfilled by mediating phloem loading or unloading of heavy metals as well as by preventing their precipitation in the alkaline phloem sap. An attempt is made to explain the chloronerva phenotype in the light of the phloem transport hypothesis of chelated iron.     

The isochorismate pathway is negatively regulated by salicylic acid signaling in O<Subscript>3</Subscript>-exposed <Emphasis Type="Italic">Arabidopsis</Emphasis>

Phytochelatins (PCs) are heavy metal binding peptides that play an important role in sequestration and detoxification of heavy metals in plants. In this study, our goal was to develop transgenic plants with increased tolerance for and accumulation of heavy metals from soil by expressing an Arabidopsis thaliana AtPCS1 gene, encoding phytochelatin synthase (PCS), in Indian mustard (Brassica juncea L.). A 35S promoter fused to a FLAG–tagged AtPCS1 cDNA was expressed in Indian mustard, and transgenic lines, designated pc lines, were evaluated for tolerance to and accumulation of Cd and Zn. Transgenic plants with moderate AtPCS1 expression levels showed significantly higher tolerance to Cd and Zn stress, but accumulated significantly less Cd and Zn than wild type plants in both shoot and root tissues. However, transgenic plants with highest expression of the transgene did not exhibit enhanced Cd and Zn tolerance. Shoots of Cd-treated pc plants had significantly higher levels of phytochelatins and thiols than wild-type plants. Significantly lower concentrations of gluthatione in Cd-treated shoot and root tissues of transgenic plants were observed. Moderate expression levels of phytochelatin synthase improved the ability of Indian mustard to tolerate certain levels of heavy metals, but at the same time did not increase the accumulation potential for Cd and Zn.     

Cadmium and nickel accumulation in rice plants. Effects on mineral nutrition and possible interactions of abscisic and gibberellic acids

Rice plants accumulate high quantities of Cd and Ni when grown for 10 days in a medium containing these heavy metals. Accompanying Cd and Ni uptake, a decrease in shoot and root length was observed, though dry matter accumulation was not affected accordingly. Metal treatments also induced a decrease in K, Ca and Mg contents in the plants, particularly in the shoots, indicating that Cd and Ni interfered not only with nutrient uptake but also with nutrient distribution into the different plant parts. Addition of abscisic acid (ABA) or gibberellic acid (GA₃) to the external solution could not overcome the depressing effects of the metals on nutrient acquisition, and even induced a further decrease of Ca content in Ni-treated plants. Both hormones also reduced, significantly, heavy metal incorporation into the plants. Additionally, hormonal applications affected the transport of Cd and Ni to the shoots, resulting in a higher percentage of the metals taken up remaining in the roots.     

Accumulation and distribution of iron, cadmium, lead and nickel in cucumber plants grown in hydroponics containing two different chelated iron supplies

Cucumber plants grown in hydroponics containing 10 μM Cd(II), Ni(II) and Pb(II), and iron supplied as Fe(III) EDTA or Fe(III) citrate in identical concentrations, were investigated by total-reflection X-ray fluorescence spectrometry with special emphasis on the determination of iron accumulation and distribution within the different plant compartments (root, stem, cotyledon and leaves). The extent of Cd, Ni and Pb accumulation and distribution were also determined. Generally, iron and heavy-metal contaminant accumulation was higher when Fe(III) citrate was used. The accumulation of nickel and lead was higher by about 20% and 100%, respectively, if the iron supply was Fe(III) citrate. The accumulation of Cd was similar. In the case of Fe(III) citrate, the total amounts of Fe taken up were similar in the control and heavy-metal-treated plants (27-31 μmol/plant). Further, the amounts of iron transported from the root towards the shoot of the control, lead- and nickel-contaminated plants were independent of the iron(III) form. Although Fe mobility could be characterized as being low, its distribution within the shoot was not significantly affected by the heavy metals investigated.     

Impact of two iron(III) chelators on the iron, cadmium, lead and nickel accumulation in poplar grown under heavy metal stress in hydroponics

Poplar (Populus jacquemontiana var. glauca cv. Kopeczkii) was grown in hydroponics containing 10 μM Cd(II), Ni(II) or Pb(II), and Fe as Fe(III) EDTA or Fe(III) citrate in identical concentrations. The present study was designed to compare the accumulation and distribution of Fe, Cd, Ni and Pb within the different plant compartments. Generally, Fe and heavy-metal accumulation were higher by factor 2-7 and 1.6-3.3, respectively, when Fe(III) citrate was used. Iron transport towards the shoot depended on the Fe(III) chelate and, generally, on the heavy metal used. Lead was accumulated only in the root. The amounts of Fe and heavy metals accumulated by poplar were very similar to those of cucumber grown in an identical way, indicating strong Fe uptake regulation of these two Strategy I plants: a cultivar and a woody plant. The Strategy I Fe uptake mechanism (i.e. reducing Fe(III) followed by Fe(II) uptake), together with the Fe(III) chelate form in the nutrient solution had significant effects on Fe and heavy metal uptake. Poplar appears to show phytoremediation potential for Cd and Ni, as their transport towards the shoot was characterized by 51-54% and 26-48% depending on the Fe(III) supply in the nutrient solution.     

Molecular mechanisms of heavy metal hyperaccumulation and phytoremediation

A relatively small group of hyperaccumulator plants is capable of sequestering heavy metals in their shoot tissues at high concentrations. In recent years, major scientific progress has been made in understanding the physiological mechanisms of metal uptake and transport in these plants. However, relatively little is known about the molecular bases of hyperaccumulation. In this paper, current progresses on understanding cellular/molecular mechanisms of metal tolerance/hyperaccumulation by plants are reviewed. The major processes involved in hyperaccumulation of trace metals from the soil to the shoots by hyperaccumulators include: (a) bioactivation of metals in the rhizosphere through root–microbe interaction; (b) enhanced uptake by metal transporters in the plasma membranes; (c) detoxification of metals by distributing to the apoplasts like binding to cell walls and chelation of metals in the cytoplasm with various ligands, such as phytochelatins, metallothioneins, metal-binding proteins; (d) sequestration of metals into the vacuole by tonoplast-located transporters. The growing application of molecular-genetic technologies led to the well understanding of mechanisms of heavy metal tolerance/accumulation in plants, and subsequently many transgenic plants with increased resistance and uptake of heavy metals were developed for the purpose of phytoremediation. Once the rate-limiting steps for uptake, translocation, and detoxification of metals in hyperaccumulating plants are identified, more informed construction of transgenic plants would result in improved applicability of the phytoremediation technology.     

Heavy metal accumulation in iron plaque and growth of rice plants upon exposure to single and combined contamination by copper, cadmium and lead

A pot experiment was conducted to evaluate the bioaccumulation of heavy metals and growth response of rice plants after exposure to single and combined contamination by Cu, Cd and Pb. The results showed that the biomass production was not significantly affected by either single or combined treatment of Cu, Cd and Pb. Adding Cu (Cd, or Pb) separately all increased concentrations of the respective element in root and shoot (p < 0.001). In the combined contamination, Pb promoted both root and shoot absorption of Cu and Cd (p < 0.001), and Cu affected Cd and Pb absorption in the root, but Pb concentrations in both root and shoot were not affected by Cd application. The formation of iron plaques varied obviously with soil types. Heavy metal accumulation in iron plaques was induced by the three elements (p < 0.001). Furthermore, the three heavy metals exhibited an interactive relationship as measured by the Cu, Cd, Pb and Fe concentrations in root surface iron plaques. The iron plaques partially inhibited transfer of Pb to root and shoot, but no such effect was observed for Cu and Cd. This research indicates that the interaction among different heavy metal elements is very complex. It is very important to have a clear understanding on the associated mechanism and the consequential impact on plant growth.     

Heavy metal accumulation in iron plaque and growth of rice plants upon exposure to single and combined contamination by copper, cadmium and lead

A pot experiment was conducted to evaluate the bioaccumulation of heavy metals and growth response of rice plants after exposure to single and combined contamination by Cu, Cd and Pb. The results showed that the biomass production was not significantly affected by either single or combined treatment of Cu, Cd and Pb. Adding Cu (Cd, or Pb) separately all increased concentrations of the respective element in root and shoot (p < 0.001). In the combined contamination, Pb promoted both root and shoot absorption of Cu and Cd (p < 0.001), and Cu affected Cd and Pb absorption in the root, but Pb concentrations in both root and shoot were not affected by Cd application. The formation of iron plaques varied obviously with soil types. Heavy metal accumulation in iron plaques was induced by the three elements (p < 0.001). Furthermore, the three heavy metals exhibited an interactive relationship as measured by the Cu, Cd, Pb and Fe concentrations in root surface iron plaques. The iron plaques partially inhibited transfer of Pb to root and shoot, but no such effect was observed for Cu and Cd. This research indicates that the interaction among different heavy metal elements is very complex. It is very important to have a clear understanding on the associated mechanism and the consequential impact on plant growth.     

Remediation of lead and cadmium-contaminated soils

The research was designated to study the ability of plants to bio-accumulate, translocate and remove the heavy metals, lead and cadmium from contaminated soil. The herbal plant ryegrass, Lolium multiflorum was investigated as a bio-accumulator plant for these metals. The translocation of these heavy metals in the herbal plant was compared considering root to shoot transport and redistribution of metals in the root and shoot system. The trace metal contents from root and shoot parts were determined using atomic absorption spectrometer. The results showed that the percent of lead and cadmium transferred to ryegrass plant were averaged as 51.39, and 74.57%, respectively, while those remained in the soil were averaged as 48.61 and 25.43% following 60 days of treatment. The soil-plant transfer index in root and shoot system of ryegrass was found to be 0.32 and 0.20 for lead, and 0.50 and 0.25 for cadmium. These findings indicated that the herbal plant ryegrass, Lolium multiflorum is a good accumulator for cadmium than lead. The soil-plant transfer factor (the conc. of heavy metal in plant to the conc. in soil) indicated that the mechanism of soil remedy using the investigated plant is phytoextraction where the amounts of heavy metals transferred by plant roots into the above ground portions were higher than that remained in the soil. The method offers green technology solution for the contamination problem since it is effective technology with minimal impact on the environment and can be easily used for soil remedy.     

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。     

Phytochelatins as biomarkers for heavy metal stress in maize (Zea mays L.) and wheat (Triticum aestivum L.): combined effects of copper and cadmium

Heavy metal contaminated soils often show increased levels of more than one metal, e.g. copper (Cu), cadmium (Cd), zinc (Zn), lead (Pb) or nickel (Ni). In case such soils are used for crop production, prediction of yield reduction or quality decline due to heavy metals in the soil is inadequate when based only on chemical soil analysis. The use of biomarkers such as phytochelatins (PC), non-protein thiols specifically induced in plants upon exposure to heavy metals, may be an additional tool or diagnostic criterion in heavy metal research and in practice. In the present work, Cu and Cd uptake and induction of PC synthesis are studied with hydroponically grown maize and wheat plants exposed to mixtures of the two metals. We observed a close positive relationship between the concentrations of Cd and PC in the plant shoot material. A decreased shoot concentration of Cd after addition of Cu, due to metal competition at common root absorption sites, coincided with lower shoot PC levels. Also differences in metal uptake and xylary metal transport among the two plant species were reflected in corresponding differences in PC concentration. The observed direct relationship between shoot PC concentration and the degree of metal-induced growth inhibition makes the use of PC promising for the purpose tested for.     

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。     

矿区绿化树木对镉和锌的吸收与分布

研究了南京某矿区11种树木的重金属吸收和分布特征.结果表明：树木对重金属的吸收和富集因树木种类、部位及重金属种类的不同而异.法国冬青对Cd的含量、富集和转运系数均高于其它树种,可以植物萃取方式修复土壤污染.Cd主要积累在树木根部,树木不同器官对Cd积累能力的总趋势为根＞叶及枝＞树皮＞树干;而Zn更多地积累在树木地上部（叶和枝）而非根部.11种树木对Cd和Zn的富集系数均<0.2.树木对Cd和Zn的转运系数差异显著,总体上是Zn>Cd.     

Effects of cadmium, nickel and lead on growth, chlorophyll content and proteins of weeds

The effects of Cd, Ni and Pb on the growth, chlorophyll (Chl) and protein contents, and content of proteases of potted weed plants Cyperus difformis, Chenopodium ambrosioides and Digitaria sanguinolis were determined. The three heavy metals inhibited the shoot growth but were less suppresive to root growth. They also lowered leaf Chl content. The changes in root and shoot protein and proteases contents of weeds were interrelated. The heavy metal additions to soil increased their contents in both roots and shoots, several times more in roots than in shoots. This revised version was published online in July 2006 with corrections to the Cover Date.     

Eco-service enhancement in peri-urban area of coal mining city of Huaibei in East China

After 50 years of coal mining, Huaibei Mine, located at 50 km southeast of Xuzhou City in East China, has grown to a middle-size city of 600,000 people from a small village of 2000 farmers. The Zhahe Valley, with 400 km² of a built-up area and more than 100 km² of subsided peri-urban wetland at the city center, is surrounded by eight exhausted old mines and communities. In cooperation with the local city government, an ecological landuse change assessment and eco-city planning project has been carried out with a focus on the assessment, restoration and enhancement of the wetland as an eco-service to the community. The assessment includes relationships to Green House Gas emissions and heat island effects, as well as measures for a livable, workable, affordable and sustainable human settlement development through industrial transition, landscape design and capacity building. This paper will briefly introduce the main ecological approaches and results of the assessment, including measures such as changing the car-dominated transportation network to a rail-dominated network, transforming the coal-oriented high-carbon industry to a service-oriented low-carbon industry, the C-shape urban form to an O-shape with a green–blue core at the center, and the fragmentized collapsed land to integrative eco-service land.     

植物重金属转运蛋白研究进展

土壤中的有毒重金属不仅对植物有害,也可通过食物链危害人类和动物的健康.重金属转运蛋白在植物吸收、抵抗重金属的复杂机制中起着关键作用.植物重金属转运蛋白分为吸收蛋白和排出蛋白,其中,吸收蛋白转运必需重金属进入细胞,同时也会因为必需重金属的缺乏或离子之间的竞争而运载有毒重金属;排出蛋白是一类解毒蛋白,可将过量的或有毒的重金属逆向转运出细胞,或区室化于液泡中.目前,细胞内多种重金属转运蛋白基因的转录水平与重金属离子积累之间的联系已被揭示,并分离克隆出诸多相关蛋白家族成员.本文综述了近年来发现并鉴定的主要重金属转运蛋白的金属亲和性、器官表达特异性及细胞内定位等的研究进展.     

Phytoremediation of Cd, Cr, Cu, Mn, Fe, Ni, Pb and Zn from aqueous solution using Phragmites cummunis, Typha angustifolia and Cyperus esculentus

A comparative bioaccumulation pattern and ultra structural changes were studied in Phragmites cummunis, Typha angustifolia and Cyperus esculentus in mixed metals solution of cadmium (Cd), chromium (Cr), copper (Cu), iron (Fe), manganese (Mn), nickel (Ni), lead (Pb) and zinc (Zn). P. cummunis was observed to be a shoot accumulator for Cr, Fe, Mn, Ni, Pb, and Zn. However, T. angustifolia was found to be a root accumulator for Cd, Cr, Cu, Fe, Ni and Pb. In addition, C. esculentus also accumulated most of the tested heavy metals in the roots, while Mn and Fe were translocated up to leaves. Further, the long term metal treatment showed maximum accumulation of all heavy metals in P. cummunis followed by T. angustifolia and C. esculentus. Among heavy metals, Fe was accumulated maximum, i.e., >1000 microg g(-1) by all three plants. Simultaneously, the adverse effects on biochemical parameters were noted earlier in C. esculentus than T. angustifolia and P. cummunis. Ultra structural observation showed the cellular changes in wetland plants after longer exposure. Results revealed that P. cummunis and T. angustifolia had more potential for tested metals than C. esculentus. This study established that these wetland plants could be used for heavy metals phytoremediation from metal containing industrial wastewater.     

Heavy metals in white lupin: uptake, root-to-shoot transfer and redistribution within the plant

The translocation of manganese (Mn), nickel (Ni), cobalt (Co), zinc (Zn) and cadmium (Cd) in white lupin (Lupinus albus cv. Amiga) was compared considering root-to-shoot transport, and redistribution in the root system and in the shoot, as well as the content at different stages of cluster roots and in other roots. To investigate the redistribution of these heavy metals, lupin plants were labelled via the root for 24 h with radionuclides and subsequently grown hydroponically for several weeks. 54Mn, 63Ni and 65Zn were transported via the xylem to the shoot. 63Ni and 65Zn were redistributed afterwards via the phloem from older to younger leaves, while 54Mn remained in the oldest leaves. A strong retention in the root was observed for 57Co and 109Cd. Cluster roots contained higher concentrations of all heavy metals than noncluster roots. Concentrations were generally higher at the beginning of cluster root development (juvenile and immature stages). Mature cluster roots also contained high levels of 54Mn and 57Co, but only reduced concentrations of 63Ni, 65Zn and 109Cd.     

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.