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。     

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。     

植物对铅的吸收、转运、累积和解毒机制研究进展

揭示植物对铅的吸收、转运、累积和解毒的分子机制,可以明晰农作物吸收铅的关键过程,阻控铅在粮食、蔬菜中的积累,降低重金属的食用风险;也可以阐明某些铅超积累植物的耐性与解毒机制,分离并克隆铅超积累的功能基因,培育高效的铅污染土壤修复植物.本文从铅进入植物的两个重要途径(叶片的吸附与吸收以及根系的吸收与转运)出发,系统总结讨论了植物对铅的吸收、转运、累积和分布的研究进展;采用胞外至胞内的空间顺序,分别从植物根系分泌物的解毒、细胞壁的固定和动态响应、细胞质膜的选择透过性作用、液泡的区隔化作用以及金属有机配体的螯合等方面论述植物铅耐性和解毒的分子机理,并在此基础上提出存在的问题和今后研究的重点.     

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。     

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。     

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．     

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.     

重金属超富集植物东南景天毛状根的诱导

为建立重金属超富集植物东南景天(Sedum alfredii)的毛状根诱导体系,采用发根农杆菌(Agrobacterium rhizogenes)A4侵染叶片,研究了预培养时间、侵染时间和共培养时间对毛状根诱导率的影响。结果表明,东南景天叶片外植体的预培养时间为48 h、农杆菌侵染时间为6 min、共培养时间为48 h是适宜的毛状根诱导时间,毛状根的诱导率可达85%。PCR检测表明诱导的毛状根中存在rol B基因片段。这是东南景天首次建立用发根农杆菌诱导毛状根体系。     

海岸带不同林龄木麻黄对重金属的吸收与富集作用

为研究不同林龄木麻黄对重金属的吸收与富集作用,用ICP-MS测定了福建东山县国有防护林场10、16、20、30年生木麻黄小枝、细根、凋落物及林地土壤中的5种重金属(Cr、Cu、Zn、Cd和Pb)含量。结果表明,木麻黄对不同重金属的吸收能力不同,从高到低为ZnCrPbCuCd;木麻黄细根中Cr和Cu、Pb,Fe和Cu,Zn和Cd具有明显的协同作用;各林龄木麻黄对重金属均具有一定的耐性,对Cu、Zn、Cd、Pb有很好的富集作用;20年生木麻黄的重金属年积累量最高;说明木麻黄对重金属的净化作用具有极大的潜力,20年生木麻黄作用最为明显。     

Heavy metal uptake and chemical changes in the rhizosphere of Thlaspi caerulescens and Thlaspi ochroleucum grown in contaminated soils

Thlaspi caerulescens (J. and C. Presl) and Thlaspi ochroleucum (Boiss. ex Heldr) were grown in three different soils containing moderate to high amounts of heavy metals in a pot experiment, using a rhizobag technique. T. caerulescens accumulated significantly more Zn in the shoots than T. ochroleucum. The concentrations of Zn in the shoots of T. caerulescens ranged from 3100 to 8100 mg kg^-1 dry matter (DM), but only from 800 to 1600 mg kg^-1 DM in T. ochroleucum. Total uptake of Zn in the shoots of T. caerulescens was about 5 times that of T. ochroleucum. In contrast, the differences between the two species in the uptake of Cd, Cr, Cu, Ni and Pb were generally small. Concentrations of mobile Zn (extractable with 1M NH₄NO₃) in the rhizosphere and non-rhizosphere soils decreased considerably after growth of both plants, but the decreases were greater with T. caerulescens than with T. ochroleucum. The decreases in the mobile fraction accounted for less than 10% of the total Zn uptake by T. caerulescens indicating that this species was effective in mobilising Zn from less soluble fractions in the soils. The rhizosphere soils tended to have higher concentrations of mobile Zn than the non-rhizosphere soils, probably because of the lower pH in the rhizosphere. The pH in the rhizosphere soils was 0.2-0.4 units lower than that in the non-rhizosphere soils at the end of the experiment. However, there were no significant differences between the two species in the degree of rhizosphere acidification. The results suggest that T. caerulescens has potential for removing Zn from moderately to highly contaminated soils, but that this ability was not related to the pH changes in the rhizosphere.     

Cadmium tolerance and antioxidative defenses in hairy roots of the cadmium hyperaccumulator,Thlaspi caerulescens

Plant species capable of hyperaccumulating heavy metals are of considerable interest for phytoremediation and phytomining. This work aims to identify the role of antioxidative metabolism in heavy metal tolerance in the Cd hyperaccumulator, Thlaspi caerulescens. Hairy roots of T. caerulescens and the non-hyperaccumulator, Nicotiana tabacum (tobacco), were used to test the effects of high Cd environments. In the absence of Cd, endogenous activities of catalase were two to three orders of magnitude higher in T. caerulescens than in N. tabacum. T. caerulescens roots also contained significantly higher endogenous superoxide dismutase activity and glutathione concentrations. Exposure to 20 ppm (178 microM) Cd prevented growth of N. tabacum roots and increased hydrogen peroxide (H(2)O(2)) levels by a factor of five relative to cultures without Cd. In contrast, growth was maintained in T. caerulescens, and H(2)O(2) concentrations were controlled to low, nontoxic levels in association with a strong catalase induction response. Treatment of roots with the glutathione synthesis inhibitor, buthionine sulfoximine (BSO), exacerbated H(2)O(2) accumulation in Cd-treated N. tabacum, but had a relatively minor effect on H(2)O(2) levels and did not reduce Cd tolerance in T. caerulescens. Lipid peroxidation was increased by Cd treatment in both the hyperaccumulator and non-hyperaccumulator roots. This work demonstrates that metal-induced oxidative stress occurs in hyperaccumulator tissues even though growth is unaffected by the presence of heavy metals. It also suggests that superior antioxidative defenses, particularly catalase activity, may play an important role in the hyperaccumulator phenotype of T. caerulescens.     

Cadmium accumulation in populations of Thlaspi caerulescens and Thlaspi goesingense

The capacity to accumulate cadmium (Cd) and zinc (Zn) was compared in Thlaspi goesingense and four populations of Thlaspi caerulescens . Two populations of T. caerulescens were grown in hydroponics at five concentrations of Cd. In addition, plants were grown in pots containing compost in which three different concentrations of Cd and two concentrations of Zn were added. A field trial was conducted to compare Zn and Cd uptake by three populations of T. caerulescens on nine selected plots of the Woburn Market Garden Experiment (UK) which had been contaminated to different degrees with heavy metals owing to past applications of sewage sludge. Results show that the four populations of T. caerulescens had the same ability to hyperaccumulate Zn but were significantly different in terms of Cd accumulation. Two populations of T. caerulescens from Southern France accumulated much more Cd than the populations from Prayon (Belgium) and Whitesike (UK). Generally, uptake of Cd was not decreased by increased concentrations of Zn in the substrate. These results indicate that the mechanisms of Cd and Zn hyperaccumulation are not identical in this species. This is the first report of hyperaccumulation of Cd by T. goesingense , but the growth of this species was markedly reduced by the large concentrations of Zn in the substrate. Future work should focus on the differences between Cd and Zn uptake in hyperaccumulator plants at the species and population level.     

Uptake and transport of zinc in the hyperaccumulator Thlaspi caerulescens and the non-hyperaccumulator Thlaspi ochroleucum

Growth and zinc uptake of the hyperaccumulator species Thlaspi caerulescens J. & C. Presl and the non-hyperaccumulator species Thlaspi ochroleucum Boiss. & Heldr. were compared in solution culture experiments. T. caerulescens was able to tolerate 500 mmol m^?3 (32.5 g m^?3) Zn in solution without growth reduction, and up to 1000 mmol m^?3 (65 g m^?3) Zn without showing visible toxic symptoms but with a 25% decrease in dry matter (DM) yield. Up to 28 g kg^?1 of Zn in shoot DM was obtained in healthy plants of T. caerulescens. In contrast, T. ochroleucum suffered severe phytotoxicity at 500 mmol m^?3 Zn. Marked differences were shown in Zn uptake, distribution and redistribution between the two species. T. caerulescens had much higher concentrations of Zn in the shoots, whereas T. ochroleucum accumulated higher concentrations of Zn in the roots. When an external supply of 500 mmol m^?3 Zn was withheld, 89% of the Zn accumulated previously in the roots of T. caerulescens was transported to the shoots over a 33 d period, whereas in T. ochroleucum only 32% was transported. T. caerulescens was shown to have a greater internal requirement for Zn than other plants. Increasing the supply of Zn from 1 to 10 mmol m^?3 gave a 19% increase in the total DM of this species. liven the shoots from the 1 mmol m^?3 Zn treatment which showed Zn deficiency contained 10 times greater Zn concentrations than the widely reported critical value for Zn deficiency to occur in many other plant species. The results obtained suggest that strongly expressed constitutive sequestration mechanisms exist in the hyperaccumulator T. caerulescens, which detoxify the large amount of Zn present in shoot tissues and decrease its physiological availability in the cytosol. Both T. caerulescens and T. ochroleucum had constitutively high concentrations of malate in shoots, which were little affected by different Zn treatments. Although malate may play a role in Zn chelation because of the high concentrations present, it cannot explain the species specificity of Zn tolerance and hyperaccumulation.     

Isolation,characterization, and identification of bacteria associated with the zinc hyperaccumulator Thlaspi caerulescens subsp. calaminaria

We investigated bacterial populations associated with the Zn hyperaccumulator Thlaspi caerulescens subsp. calaminaria grown in a soil collected from an abandoned Zn-Pb mine and smelter in Plombières, Belgium. The bacterial population of the nonrhizospheric soil consisted of typical soil bacteria, some exhibiting multiple heavy-metal resistance characteristics that often are associated with polluted substrates: 7.8% and 4% of the population survived in the presence of elevated levels of Zn (1 mM) and Cd (0.8 mM), respectively. For the bacterial population isolated from the rhizosphere, the comparable survival rates were 88 and 78%. This observation indicates a selective enrichment of the metal-resistant strains due to an increased availability of the metals in soils near the roots compared with nonrhizospheric soil. The endophytic inhabitants of the roots and shoots were isolated, identified, and characterized. Although similar endophytic species were isolated from both compartments, those from the rhizoplane and roots showed lower resistance to Zn and Cd than the endophytic bacteria isolated from the shoots. In addition, root endophytic bacteria had additional requirements. Contrary to the rootresiding inhabitants, the shoot represented a niche rich in metal-resistant bacteria and even seemed to contain species that were exclusively abundant there. These differences in the characteristics of the bacterial microflora associated with T. caerulescens might possibly reflect altered metal speciation in the different soils and plant compartments studied.