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.     

Biochar and alternate partial root-zone irrigation greatly enhance the effectiveness of mulberry in remediating lead-contaminated soils

土壤铅污染日益严重,植物修复是一种环保的污染土壤修复技术。本文旨在研究四种土壤铅污染水平下,添加生物碳和分根区交替 灌溉(Alternative Partial Root-zone Irrigation,APRI)对桑树幼苗的生长、铅适应性和铅积累的影响。我们以生物碳(添加与不添加生物碳)、灌溉方式(APRI 与常规灌溉)和土壤铅水平(0、50、200 和800 mg kg⁻¹ Pb)为三因素实施了温室试验。通过测定桑树幼苗的生长性状、渗透物质代谢、抗氧化酶活性、铅的积累和转运等参数,探讨了不同处理对桑树生长发育的影响。结果表明,桑树对土壤铅污染有较强的适应能力;生物碳和APRI 在不同土壤铅水平上协同提高了生物量和吸收根表面积。桑树通过调节谷胱甘肽 (GSH)、脯氨酸代谢和过氧化物酶(POD)活性,增加了渗透和抗氧化调节能力,进而提高了对重度铅污染土壤(800 mg kg⁻¹)的抗性。桑苗中的铅离子主要集中在根中,与土壤铅浓度具有剂量效应。土壤铅、生物碳和ARPI的交互作用影响了叶片和根系中铅的浓度、转运和生物富集系数。综上所述,在桑树栽培中结合外源生物碳和APRI可有效地用于修复土壤铅污染。     

超积累植物的金属配位体及其在植物修复中的应用

综述了超积累植物体内金属配位体（包括植物螯合肽、植物金属硫蛋白、有机酸和氨基酸）的生物合成,参与的植物体内金属的吸收、运输、积累的解毒过程的生理及分子机制,并对金属配位体在植物修复中的应用作了评述。     

铅对山西省路域优势草本植物生长的影响及铅累积特征

采用温室盆栽试验,研究了不同浓度铅(0、500、1000、1500mg.kg-1)对14种山西省路域优势草本植物生长的影响及其铅吸收积累特征.结果表明:在14种草本植物中,随着铅浓度的增大,反枝苋和高丹草表现出明显的中毒症状,其他12种植物的株高和生物量与对照相比均无显著降低,表现出对铅污染具有一定的耐受性;藜和新麦草植株的地上部铅含量最低,各浓度铅处理下平均值分别为12.70和11.33mg.kg-1,地上部与根的铅含量比(S/R)最低,分别为0.12和0.10,表明二者为低积累植物,可用于铅污染土壤的植被恢复;红叶苋和绿叶苋植株地上部的铅迁移量最高,1500mg.kg-1铅处理下每百株铅迁移总量分别为53.37和45.29mg,可作为修复铅污染土壤的先锋植物.     

铅污染现状及其修复机理研究进展

介绍了铅污染的现状、修复机理及存在的问题。其中重点阐述了国内在寻找铅的超累积植物的研究进展,国外研究者对铅在植物体内吸收、运输和储存机理方面的研究现状,同时对已发现的与铅污染有关的基因以及转基因技术的应用情况进行了简单介绍,并探讨了提高铅污染修复效率的一些方法。     

伴矿景天植物螯合肽合酶基因的克隆及功能分析

重金属超积累植物由于长期生长在高浓度的重金属环境中,使得经由植物螯合肽（phytochelatins, PCs）解毒途径来应对重金属毒害代价高昂。我们从Zn/Cd超积累植物伴矿景天（Sedum plumbizincicola）中克隆了植物螯合肽合酶（phytochelatin synthase, PCS）基因SepPCS。该基因在裂殖酵母和拟南芥中表达后都具有PCS活性,而且能够互补它们的PCs缺失突变体的Cd敏感表型。SepPCS在伴矿景天中的表达受到高浓度Cd处理的诱导。与其近亲非超积累生态型东南景天（S. alfredii）相比,虽然伴矿景天地上部PCs与Cd的摩尔比远低于东南景天,但是在高浓度Cd处理时PCs含量以及PCs与Cd的摩尔比急剧增加。我们推测在伴矿景天应对Cd毒害的过程中, PCs起到一定的作用,并且在高浓度Cd胁迫时地上部PCs依赖的解毒作用有所加强。     

一种新的铅富集植物--富集生态型东南景天

对浙江一古老铅锌矿区的土壤和植物种群进行调查后发现一种新的具有耐铅特性和铅富集能力的植物--景天科景天属东南景天(Sedum alfredii Hance),称为铅富集生态型植物.进一步比较和分析了不同浓度硝酸铅处理对富集和非富集生态型东南景天的生长及其对铅的吸收特性的影响.结果表明, 320 mg Pb/L处理对富集生态型的地上部分生长无显著影响,而非富集生态型在20 mg Pb/L时即出现受害症状.富集和非富集生态型的地上部分铅含量、根系铅含量以及单株铅积累速率均随处理浓度的增加而表现出先升后降的变化趋势.其中富集生态型的地上部分铅含量、根系铅含量以及单株铅积累速率最高可达到514 mg/kg、13 922 mg/kg和8.62 μg/plant/d,分别是非富集生态型的2.27、2.62和7.16倍.由于具有生长速度快和高积累铅的能力,从植物修复的观点来说,东南景天铅富集生态型在铅污染土壤的修复方面具有巨大的潜力.     

植物螯合肽及其功能

植物螯合肽（phytochelatin,PC）是一类富含Cys、由PC合酶以GSH为底物催化合成的小分子多肽,能通过Cys的-SH络合重金属.研究PC的合成机理及其重金属解毒机制、研究PC合酶和PC合酶基因的表达模式及其功能对于运用植物修复技术治理重金属污染的土壤和水体具有重要意义.     

一种新的铅富集植物——富集生态型东南景天

对浙江一古老铅锌矿区的土壤和植物种群进行调查后发现一种新的具有耐铅特性和铅富集能力的植物———景天科景天属东南景天 (SedumalfrediiHance) ,称为铅富集生态型植物。进一步比较和分析了不同浓度硝酸铅处理对富集和非富集生态型东南景天的生长及其对铅的吸收特性的影响。结果表明 ,32 0mgPb/L处理对富集生态型的地上部分生长无显著影响 ,而非富集生态型在 2 0mgPb/L时即出现受害症状。富集和非富集生态型的地上部分铅含量、根系铅含量以及单株铅积累速率均随处理浓度的增加而表现出先升后降的变化趋势。其中富集生态型的地上部分铅含量、根系铅含量以及单株铅积累速率最高可达到 5 14mg/kg、1392 2mg/kg和 8.6 2 μg/plant/d ,分别是非富集生态型的 2 .2 7、2 .6 2和 7.16倍。由于具有生长速度快和高积累铅的能力 ,从植物修复的观点来说 ,东南景天铅富集生态型在铅污染土壤的修复方面具有巨大的潜力。     

抗坏血酸过氧化物酶活性测定的探讨

抗坏血酸过氧化物酶（ascorbateperoxi－dase,EC1.11.1.11,ASP）是以抗坏血酸为电子供体的专一性强的过氧化物酶［3,8］,主要存在于植物叶绿体和胞浆中［3,7］,一般用愈创木酚作为电子供体测定酚特异性过氧化物酶（PPOD）的方法不能测出其大部分活性。它和超氧化物歧化酶（SOD）、过氧化氨酶（CAT）、单脱氢抗坏血酸还原酶（MDARD）、双脱氢抗坏血酸还原酶（DHARD）以及谷联甘肽还原酶（GR）一起构成了活性氧清除系统中的酶系统,旦ASP目前被认为是叶绿体中清除H。0。的关键酶「’,‘j。迄今为止,国内还没有文献具体报…     

Mechanisms of cadmium detoxification in cattail (Typha angustifolia L.)

Cadmium (Cd) is a widespread heavy metal pollutant and environmental and human health hazard, which may be partially resolved using green and cost-effective phytoremediation techniques. However, the efficiency of phytoremediation is often limited by the small biomass of Cd-hyperaccumulator plants. Although cattail (Typha angustifolia L.) is tolerant of heavy metals and has a high biomass, there is little information available on its detoxification mechanisms for heavy metals, especially Cd. In the present study we investigated the tolerance of cattail to Cd and mechanisms involved in its Cd detoxification. Our results show that: (a) cattail is tolerant of Cd; (b) the root Casparian band, cell wall, vacuole, glutathione (GSH), and glutathione peroxidase (GPX) play important roles in Cd detoxification; and (c) mechanisms of Cd detoxification differ in leaf cell cytoplasm (mainly a GSH-related antioxidant defense system) and root cell cytoplasm (mainly a GSH-related chelation system). In summary, cattail possesses multiple detoxification mechanisms for Cd and is a promising species for phytoremediation of Cd-polluted environments.     

Effects of Cadmium,Chromium and Lead on Growth,Metal Uptake and Antioxidative Capacity in <Emphasis Type="Italic">Typha angustifolia</Emphasis>

This study investigates the modulation of antioxidant defence system of Typha angustifolia after 30 days exposure of 1 mM chromium (Cr), cadmium (Cd), or lead (Pb). T. angustifolia showed high tolerance to heavy metal toxicity with no visual toxic symptom when exposed to metal stress, and Cd/Pb addition also increased plant height and biomass especially in Pb treatment. Along with increased Cr, Cd, and Pb uptake in metal treatments, there was enhanced uptake of plant nutrients including Ca and Fe, and Zn in Pb treatment. A significant increase in malondialdehyde (MDA) content and superoxide dismutase (SOD) and peroxidase (POD) activities were recorded in plants subjected to Cr, Cd, or Pb stress. Furthermore, Pb stress also improved catalase (CAT), ascorbate peroxidase (APX), and glutathione peroxidase (GPX) activities; whereas Cr stress depressed APX and GPX. The results indicate that enzymatic antioxidants and Ca/Fe uptake were important for heavy metal detoxification in T. angustifolia, stimulated antioxidative enzymes, and Ca, Fe, and Zn uptake could partially explain its hyper-Pb tolerance.     

Involvement of glutathione metabolism in Eichhornia crassipes tolerance to arsenic

• Aquatic macrophytes are potentially useful for phytoremediation programmes in environments contaminated by arsenic (As). Biochemical and physiological modification analyses in different plant parts are important to understand As tolerance mechanisms.
• The objective was to evaluate glutathione metabolism in leaves and roots of Eichhornia crassipes (Mart.) Solms treated to As. Specimens of E. crassipes were cultured for 3 days in Clark's nutrient solution containing 7 μm As. The enzymes ATP sulphurylase (ATPS), glutathione reductase (GR), glutathione peroxidase (GSH‐Px), glutathione sulphotransferase (GST) and γ‐glutamylcysteine synthetase (γ‐ECS) activity, glutathione content, total protein and non‐protein thiols were evaluated.
• The ATPS activity increased in roots. GR activity in leaves and GSH‐Px in roots were lower. GST activity was higher in roots and lower in leaves, and γ‐ECS activity was higher in leaves. Glutathione levels were lower, total thiol levels were higher and non‐protein levels did not change in E. crassipes leaves and roots. Exposure to As increased enzyme activity involved with sulphur metabolism, such as ATPS. Higher GR activity and lower GSH‐Px indicate increased glutathione conjugation to As due to increased GSH availability. The higher GST activity indicates its participation in As detoxification and accumulation through As GSH conjugation. Changes in glutathione and thiol levels suggest high phytochelatin synthesis.
• In conclusion, the increments in ATPS, GR, GST and γ‐ECS activity indicate that these enzymes are involved in GSH metabolism and are part of the E. crassipes As detoxification mechanism.

     

Heavy metal-induced oxidative damage, defense reactions, and detoxification mechanisms in plants

Heavy metal (HMs) contamination is widespread globally due to anthropogenic, technogenic, and geogenic activities. The HMs exposure could lead to multiple toxic effects in plants by inducing reactive oxygen species (ROS), which inhibit most cellular processes at various levels of metabolism. ROS being highly unstable could play dual role (1) damaging cellular components and (2) act as an important secondary messenger for inducing plant defense system. Cells are equipped with enzymatic and non-enzymatic defense mechanisms to counteract this damage. Some are constitutive and others that are activated only when a stress-specific signal is perceived. Enzymatic scavengers of ROS include superoxide dismutase, catalase, glutathione reductase, and peroxidase, while non-enzymatic antioxidants are glutathione, ascorbic acid, α-tocopherol, flavonoids, anthocyanins, carotenoids, and organic acids. The intracellular and extracellular chelation mechanisms of HMs are associated with organic acids such as citric, malic and oxalic acid, etc. The important mechanism of detoxification includes metal complexation with glutathione, amino acids, synthesis of phytochelatins and sequestration into the vacuoles. Excessive stresses induce a cascade, MAPK (mitogen-activated protein kinase) pathway and synthesis of metal-detoxifying ligands. Metal detoxification through MAPK cascade and synthesis of metal-detoxifying ligands will be of considerable interest in the field of plant biotechnology. Further, the photoprotective roles of pigments of xanthophylls cycle under HMs stress were also discussed.     

Dualities in plant tolerance to pollutants and their uptake and translocation to the upper plant parts

There is a duality in plant tolerance to pollutants and its response to the pollutants’ stress.On the one hand some plants, (hyper)tolerant to heavy metals, are able to hyperaccumulate these metals in shoots, which could be beneficial for phytoremediation purposes to clean-up soil and water. On the other hand tolerant food crops, exposed to heavy metals in their growth medium, may be dangerous as carriers of toxic metals in the food chain leading to food toxicity. There is an additional duality in plant tolerance to heavy metals and that is in food crops that are tolerant and/or hyperaccumulators, which could be used on one hand for phytoremediation, under controlled conditions and on the other hand for food fortification with essential metals.Similarly, plants are also exposed to a large number of xenobiotic organic pollutants. Because they generally cannot avoid these compounds, plants take up, translocate, metabolize and detoxify many of them. There is a large variability in tolerance (defence) mechanisms against organic pollutants among plant species. This includes production of reductants but also scavenger molecules like ascorbate and glutathione and expression of the P-450 defence system, and superfamilies of the enzymes glutathione- and glucosyl-transferases. Again, with view to organic pollutants, plant detoxification mechanisms might well protect the plant itself, but produce compounds with some deleterious potential for other organisms.In this review we discuss these dualities on the basis of examples of agricultural and ‘wild’ species exposed to metal contaminants (mainly Cd) and organic pollutants. Differences in uptake and translocation of various pollutants and their consequences will be considered. We will separately outline the effects of the organic and non-organic pollutants on the internal metabolism and the detoxification mechanisms and try to indicate the differences between both types of pollutants. Finally the consequences and solutions of these dualities in plant tolerance to pollutants will be discussed.     

重金属污染的转基因植物修复——原理与应用

污染环境的植物修复技术具有成本低、不造成二次污染等优点。从自然界中寻找用于污染环境修复的超富积植物不仅难度大 ,而且受生物量、生长周期以及地理环境等因素的限制。近几年迅速发展起来的通过转基因植物进行污染环境的修复技术显示了广阔的应用前景。外源基因在植物的高效表达可以提高植物吸收、运输、降解污染物的能力以及修复的效率 ,并可以作为研究不同污染物修复机理的实验系统。以转基因植物修复几种主要的重金属污染为例 ,介绍了转基因植物修复的原理、现状及存在问题 ,并探讨了提高转基因植物修复效率的一些方法 。     

大气湿沉降油菜体内铅的形态累积及氧化胁迫效应

大气沉降是叶菜类农作物可食部位重金属铅（Pb）累积的主要来源，但大气湿沉降下铅在油菜（Brassica chinensis L.）体内的形态累积特征及氧化胁迫效应，尚不十分清晰。通过模拟大气湿沉降铅胁迫，研究油菜体内重金属Pb的累积、亚细胞分布、化学形态特征及油菜抗氧化响应。结果表明，油菜可食部位Pb含量为1.72-6.35 mg/kg，超出标准（GB 2762-2017）4.73-20.16倍。Pb大量分布在油菜茎和叶的细胞壁中（52.14%-86.99%），以草酸盐沉淀的形式存在（20.07%-59.27%），这可能会导致Pb在可食部位的大量累积（>95%）。细胞壁的固持作用和Pb主要以草酸盐和不溶性磷酸盐存在，可能是油菜重要的解毒耐性机制之一。湿沉降铅胁迫可以增加叶和茎的丙二醛（MDA）含量，造成细胞的氧化损伤，抑制叶绿素的合成。油菜可以通过提高抗氧化酶（SOD、POD、CAT）活性（7.73%-346.91%），增加抗氧化剂（AsA和GSH）和可溶性物质（可溶性糖、可溶性蛋白）含量（9.11%-279.59%），来有效应对湿沉降Pb胁迫。抗氧化酶在叶中变化最大，过氧化物酶（POD）和过氧化氢酶（CAT）活性分别上升49.41%-91.62%和123.42%-346.91%。抗氧化剂则在根中变化最显著，抗坏血酸（AsA）和谷胱甘肽（GSH）含量分别上升了134.15%-182.93%和238.78%-279.59%。可溶性糖、可溶性蛋白、POD、CAT和GSH可能是油菜叶能缓解湿沉降Pb氧化胁迫的主要因素之一（P<0.05）。研究结果可加深对大气沉降下Pb在叶菜体内累积分布行为的理解，并为大气重金属胁迫地区农作物重金属风险评估提供一定的理论依据。     

Detoxification of herbicides in Phragmites australis

Unintentional loss of herbicides into drainage ditches, shores or other waterbodies may cause large problems in farmland. Therefore strategies for the phytoremediation of agrochemicals and especially herbicides have become a topic of great interest in many agricultural areas. However, in order to establish effective biological pollution control, information on the detoxification capacity of riparian plants and aquatic macrophytes (e.g., Phragmites australis) is important to build up effective buffer stripes. We determined the detoxification capacity of Phragmites australis roots and leaves for the conjugation of agrochemicals to glutathione by assaying the model substrate CDNB as well as the herbicides fenoxaprop-P, propachlor, pethoxamid and terbuthylazine. Specific GST activities were always higher in the rhizomes (6.78 +/- 0.88 microkat/mg protein for CDNB) than in leaves (1.08 +/- 0.21 microkat/mg protein). The detoxification capacity is distributed across an array of GST isoforms. In summary, Phragmites australis seems to be efficient in herbicide detoxification and a good candidate for phytoremediation of effluents from agricultural sites.