硒对植物吸收转运镉影响机制的研究进展

硒是人和动物必需的微量营养元素之一,在植物中可抵御体内自由基伤害、提高作物产量和质量,并且可以有效缓解植物重金属的胁迫和积累。综述了在不同植物中硒对镉吸收转运影响的研究进展,探讨了硒缓解镉胁迫的机制,以期为采取措施降低农产品镉污染提供参考。     

植物吸收、转运和积累镉的机理研究进展

重金属镉(Cd)虽然不是植物生长的必需矿质元素,但依然能被植物吸收。且部分植物具有富集镉的特点,从而导致农产品镉含量超标,并通过食物链危害人类健康。研究植物吸收、转运和积累Cd的机理,对于培育低镉作物品种、降低农产品镉含量,以及选育超富集镉植物,修复镉污染土壤具有重要意义。从影响植物吸收Cd的因子,植物吸收、转运和积累Cd的机理以及植物拒Cd和富集Cd的分子机制等方面进行综述,以期为低镉作物的研究以及Cd污染土壤的综合治理提供一些参考。     

不同镉污染土壤下水稻镉富集与转运效率

选取适合湖南种植的48个水稻品种(系)在中轻度和重度镉污染土壤上进行盆栽试验,通过分析水稻籽粒产量、各器官富集系数、转运系数及植株净化效率,探明不同镉污染土壤对水稻镉富集、转运及净化效率的影响。结果表明:中轻度和重度镉污染土壤下糙米Cd含量、籽粒产量分别为0.010~0.048 mg·kg~(-1)、96.63~152.55 g·盆~(-1)和0.105~0.476mg·kg~(-1)、101.10~165.12 g·盆~(-1),重度镉污染土壤明显促进糙米镉含量增加,但对籽粒产量影响不大;中轻度和重度镉污染土壤下糙米镉富集效率和产量均不存在显著相关性;依据产量和糙米镉富集效率进行聚类分析,将所有品种在中轻度镉污染土壤下分为低镉低产组、低镉高产组;重度镉污染土壤分为高镉低产组、高镉高产组;两种镉污染土壤水稻各器官镉富集大小顺序均为糙米叶茎秆根系,转运效率则为土-根茎-叶茎-糙米根-茎,土壤镉主要转运至根部并富集于根部;水稻镉富集及转运效率受产量因素影响较小,但受土壤镉浓度影响较大;重度镉污染土壤明显增强水稻各器官镉富集和根-糙米各环节镉转运效率,较中轻度镉污染土壤增幅显著;水稻根部的净化效率远高于地上部分。     

硅对镉胁迫下红树白骨壤酚类代谢的影响

采用水培方式研究硅对重金属镉胁迫下白骨壤幼苗酚类代谢影响,探讨硅对白骨壤耐受重金属胁迫的影响,结果表明,低浓度镉胁迫(Cd 0.5 mg·L-1)下,随着外源硅的增加,叶片和根系丙二醛含量降低,但Si 50 mg·L-1处理中各器官总酚含量最高,硅对各器官单宁含量影响不显著;高浓度镉(Cd 5 mg·L-1)导致白骨壤叶片和根系膜脂过氧化,使得丙二醛含量显著高于对照,外源硅的添加显著降低了白骨壤叶片和根系丙二醛积累,Si 100mg·L-1显著增加了各器官中总酚含量和单宁含量,根系丙二醛含量与根系总酚含量呈显著负相关,叶片丙二醛含量与叶片单宁含量呈显著负相关,表明硅显著刺激高镉胁迫下白骨壤植物组织中酚类物质代谢,增加根系和叶片的抗氧化性,进而增加白骨壤对镉的耐受性。     

镉不同积累型作物品种对镉吸收、转运和积累特性的研究进展

农田土壤镉污染问题日趋严重, 利用可食部位低积累镉的农作物品种成为保障食品安全的有效策略之一。镉在作物可食部位的累积与镉的吸收转运有关。分析了根系对镉的吸收、镉向地上部的运输和分配及镉在细胞内的分布和结合形态与作物积累镉的关系, 指出作物品种间的镉积累差异受以上多种因素的调控, 但不同作物之间影响镉积累的主要因素存在差异。同时提出在对低镉积累作物品种的筛选和机制研究方面还有很多工作需要开展。     

水稻对天然富硒土壤硒的吸收及转运

以杂交晚稻“皖稻205”为试验材料,采用盆栽试验方法,研究了不同硒含量（0.5、1.0、1.5 mg·kg^-1）土壤对水稻产量、硒的吸收、转运和分配的影响.结果表明： 土壤硒含量≤1.5 mg·kg^-1对水稻产量无明显影响;水稻根系、茎叶和籽粒中的硒含量均随着土壤硒含量增加而增加,并呈现根系＞茎叶＞籽粒的特点,籽粒各组成部分的硒含量呈现米糠＞精米＞稻壳的特点;水稻根系能从富硒土壤中富集硒,根系硒的吸收系数达1.86,而硒向籽粒的转运和积累则相对恒定,转运系数为0.53～0.59;土壤硒含量为0.5～1.0 mg·kg^-1所产的富硒大米（0.15～0.20 mg Se·kg^-1）,可满足人体60～80 μg·d^-1硒的需要量,而土壤硒含量≥1.5 mg·kg^-1所产大米硒含量达到0.319 mg·kg^-1,超出粮食硒含量安全标准.综合产量和籽粒硒含量的表现,在富硒土壤（0.5～1.0 mg·kg^-1）上直接生产富硒大米,而不需要添加外源硒,既降低了生产成本,又避免了可能造成的水土污染.     

硒镉处理对萝卜硒镉吸收的影响及其交互作用

采用盆栽试验的方法,研究不同含量的硒、镉单一和复合作用对萝卜生长及其对硒、镉吸收的影响。结果表明:适量的硒(2.5~5.0 mg·kg-1)可以促进萝卜的生长,表现为萝卜发芽率提高43%、叶片SPAD值提高22%、生物量增加50%;镉单一污染时,镉显著抑制萝卜的生长(P0.05);土壤中硒的浓度低于1.5 mg·kg-1时对镉含量5.0 mg·kg-1具有拮抗作用,同时可以促进萝卜的生长;萝卜中硒的浓度随着土壤中硒的添加量的增加而增加,对硒的富集效果明显,且地下部分对硒的吸收能力大于地上部分;土壤中硒浓度5.0 mg·kg-1时,5.0 mg·kg-1镉的存在不影响萝卜对硒的吸收;当土壤中硒的浓度为5.0~10.0 mg·kg-1时,镉的存在虽然影响萝卜对硒的吸收,但萝卜中硒的含量仍处于稳定积累状态;而当土壤中硒浓度≥10.0 mg·kg-1时,镉的存在会抑制萝卜对硒的吸收,且这种作用表现为地上部分强于地下部分;硒能显著降低萝卜对镉的吸收,且这种作用表现为地下部分强于地上部分。因此,在轻度镉污染的土壤中种植富硒萝卜,既能生产营养安全的富硒食品,又可实现污染农田的再利用。     

镉锌复合污染对龙葵苗期生长和镉锌累积特性的影响

采用营养液培养法研究了苗期龙葵(Solanumnigrum L.)对镉锌复合污染的反应及其对镉锌的吸收和积累特性.结果表明:(1)高浓度镉(200μmol·L-1)、锌(500μmol·L-1)胁迫使苗期龙葵植株生长受到严重抑制,但低浓度镉锌复合处理的部分植株生物量甚至超过了对照.(2)添加锌对苗期龙葵地上部镉积累有显著的影响(P<0.05);当锌浓度为100μmol·L-1时,镉积累量达到峰值,其中的100Cd/100Znμmol·L-1处理组合的镉积累量为所有处理中的最大值(地上部积累量高达171.31μg·plant-1);当锌浓度增加到500μmol·L-1时,植株体内镉积累量则呈现下降趋势.(3)添加镉对苗期龙葵地上部锌积累也有显著的影响(P<0.05);当镉浓度达50μmol·L-1时,根、茎、叶中锌积累量均达到最高值;当镉浓度为200μmol·L-1时,各处理单株地上部锌积累量最低.(4)从植株整体的镉、锌积累量来看,苗期龙葵地上部镉积累量最高达根系的14.67倍,而其锌积累量最高达根系的13.59倍.研究发现,龙葵幼苗根系、茎和叶中镉、锌含量随营养液中镉、锌浓度的增加而显著增加(P<0.05);添加锌在一定程度上缓解了镉对苗期龙葵的毒害效应,低浓度锌还可以提高植株对镉的吸收和积累;添加镉在一定程度上增强了植株对锌的耐受性和吸收富集能力;植株地上部的镉、锌积累量明显高于根系.     

霞多丽苗木中镉的积累、亚细胞分布及化学存在形态

以一年生盆栽葡萄品种霞多丽(Vitis vinifera L.cv.Chardonnay)自根苗及SO4砧嫁接苗为试材,采用CdCl2和CaCl2根部灌入的方法,研究了镉在霞多丽植株内的亚细胞分布和存在形态,以及外源添加氯化钙对植株镉吸收的影响.结果表明:大部分镉积累在霞多丽自根苗及嫁接苗的地下部器官;4 mmol·L-1浓度的CdCl2处理下,镉在自根苗根及根颈部分的积累量占77.1％,在叶片中的积累量占1.4％,而嫁接苗中镉在嫁接口以下部分的积累量高达93.9％,叶片中的积累量仅占0.1％;5 mmol·L-1外源钙缓解了植株对镉的吸收积累,而10mmol ·L-1外源钙则显著增加了植株对镉的吸收积累.镉在根系和叶片中的亚细胞分布规律为细胞壁＞可溶性部分＞细胞器,且在细胞壁中积累50％以上;镉在根系中主要以氯化钠提取态存在,其次为乙酸提取态,去离子水提取态含量最少.随着镉处理浓度的增加,各提取态含量有所变化.     

硒对镉胁迫下大球盖菇菌丝生长的影响

选用大球盖菇Stropharia rugosoannulata为试验材料,分析在镉胁迫处理下（2mg/L）添加不同浓度硒（0、6、15、20mg/L）对大球盖菇菌丝生长的影响,初步研究了施加硒对镉胁迫下大球盖菇菌丝生长抑制的缓解效应和作用机制。结果表明：镉对大球盖菇生长速度和生物量具有明显抑制作用,镉造成了大球盖菇渗透调节物质、过氧化氢和丙二醛含量显著增加;添加硒后,镉对大球盖菇菌丝的生长速度、生物量抑制作用均有所改善,且增加了渗透调节物质,促进镉胁迫下大球盖菇渗透调节能力;硒诱导抗氧化酶的活性上调,提高活性氧清除能力,降低了菌丝中丙二醛和过氧化氢的含量,缓解细胞膜过氧化程度,增强了大球盖菇对镉的耐受性。本研究重点探讨外源添加硒对镉胁迫下大球盖菇菌丝生长、生理特性及抗氧化胁迫能力的影响,为进一步研究硒在食用菌抗镉栽培中的应用奠定了一定的理论基础。     

Variation in Cadmium Tolerance and Accumulation and Their Relationship in Wheat Recombinant Inbred Lines at Seedling Stage

In order to identify the variation of cadmium (Cd) tolerance and accumulation in wheat (Triticum aestivum L.), a study was conducted in hydroponic culture with or without Cd using recombinant inbred lines (RILs) consisting of 103 RILs derived from a cross of Chuan 35050 × Shannong 483 at seedling stage. The parameters of shoot height, secondary roots numbers, tiller numbers, shoot dry weights, root dry weights, and maximum efficiency of photosystem II photochemistry under dark-adopted conditions were measured. Cd-tolerant indexes were then calculated as relative the above traits under Cd stress to those under the control. Cd concentration in shoot or root was determined and Cd accumulation and translocation were calculated. Based on the Cd-tolerant indexes, membership function analysis was used to determine the variation of the above parameters. The results showed a continuous distribution among the RILs and then the RILs were divided into five groups according to their tolerance. Lines 76 and 17 were considered as the most Cd-tolerant lines while lines 103 and 51 were as the most Cd-sensitive lines. Meanwhile, lines 38 and 79 were with minimum Cd translocation ratio while lines 88 and 53 were with maximum Cd translocation ratio, respectively. The relationship between Cd tolerance and accumulation was not significant, indicating Cd tolerance and accumulation may be independent traits in the RILs. Thus, lines with high Cd tolerance and less Cd accumulation could be selected for wheat breeding.     

Identification of Barley Varieties Tolerant to Cadmium Toxicity

Two successive hydroponic experiments were carried out to identify barley varieties tolerant to Cd toxicity via examining Soil–Plant Analyses Development (SPAD) value, plant height, leaves and tillers per plant, root number and volume, and biomass accumulation. The results showed that SPAD values (chlorophyll meter readings), plant height, leaf number, root number and volume, and biomass accumulation of shoot/root were significantly reduced in the plants grown in 20 μM Cd nutrient solution compared with control, and the uptake and translocation of Zn, Mn, and Cu was also strictly hindered. Furthermore, there was a highly significant difference in the reduction in these growth parameters among varieties, and varieties “Weisuobuzhi” and “Jipi 1” showed the least reduction both in the two experiments, suggesting their high tolerance to Cd toxicity, while “Dong 17” and “Suyinmai 2” with the greatest reduction and the toxicity symptoms appeared rapidly and severely, denoting as Cd-sensitive varieties. Significant variety difference in Cd concentration was also found, with Weisuobuzhi containing the highest and Jipi 1 the lowest Cd concentration in shoots.     

Cadmium tolerant and sensitive wheat lines: their differences in pollutant accumulation,cell damage,and autophagy

Cadmium (Cd) is a major abiotic stressor that affects plant growth and reduces the productivity of field crops. Here, we examined the ultrastructural, physiological, and molecular changes in three wheat cultivars [Sumai 3, Jingdong 8 (JD 8), and Nannong 9918 (9918)] in response to different concentrations of Cd (0, 10, 50, and 100 μM) in 1/4 Hoagland nutrient solution. The results showed that JD 8 contained the lowest shoot Cd content and the highest root Cd content among the three cultivars at higher Cd concentrations and so JD 8 was proposed to be a relatively Cd-tolerant cultivar. Next, the stress responses of JD 8 and 9918 were compared. Cadmium reduced root growth and size and number of the leaves, inhibited root hair development, and promoted leaf cell death. The result of trypan blue staining showed that the dead leaf cells induced by Cd stress gradually emerged in the xylem, supporting the hypothesis that cell death could restrict Cd transport. The Cd-induced deterioration of the leaf ultrastructure led to the complete disorganization of the chloroplasts, which had lower amounts of transitory starch and an increased number of osmiophilic granules compared to those in the untreated controls. Autophagy-related genes and autophagy in the leaves were induced by Cd stress. At the same concentration and Cd treatment time, the Cd-tolerant genotype JD 8 exhibited less toxic symptoms compared to the Cd-sensitive genotype 9918. The results of this study provide insights into the ultrastructural and physiological damages induced by Cd stress, which may help in selecting Cd-tolerant wheat cultivars.     

Comparisons of cadmium tolerance and accumulation at seedling stage in wheat varieties grown in different decades in China

Cadmium (Cd) tolerance and accumulation in wheat varieties were investigated at seedling stage under a controlled environmental condition. The 46 leading wheat varieties cultivated from the 1950s to 2000s in China were treated at the three-leaf stage with a 50 μM CdCl₂ solution for 24 days. Growth and photosynthesis parameters were measured and the Cd-tolerance index (ratio of a given parameter under Cd treatment to that of the control) was determined. Cd accumulation in shoots and roots and Cd translocation were also determined. It was found that Cd tolerance and accumulation of these wheat varieties varied over the different decades. Principal components analysis (PCA) showed that wheat varieties in 1950s and 1980s were tolerant while varieties from the last decade were sensitive to Cd stress. Wheat varieties in 1960s and 1970s were particularly tolerant to Cd stress for the parameters of shoot height, secondary root numbers, net photosynthesis and transpiration rate while the varieties in the 1990s were sensitive to Cd stress for shoot dry weight and root dry weight. Comparing each decade to the average Cd translocation ratio from the roots to the shoots for the whole period, the varieties from the 1950s and 1960s had a higher translocation ratio, while varieties in the 1970s were below that average. Varieties from the 1980s to 2000s showed an average translocation ratio. Using cluster analysis (CA), Shannongfu 63, Yangmai 1 and Yangmai 158 were the most Cd-tolerant varieties in which Cd translocation ratio were low, and Yumai 18 and Huaimai 20 were the most Cd-sensitive varieties in which Cd translocation ratio were high. The results indicating that wheat varieties from different decades were different in Cd tolerance and accumulation, and could be useful for breeding wheat for Cd stress tolerance.     

水稻镉积累特性的生理和分子机制研究概述

我国的稻米镉超标对国民身体健康造成严重威胁, 而选育低镉积累的水稻(Oryza sativa)品种是降低稻米镉含量行之有效的策略, 因此有必要了解水稻对镉的积累特性及其生理过程和相关功能基因。该文概述了镉在水稻根部的吸收、木质部中的装载与运输、茎节中的分配、叶片中再分配以及籽粒镉积累等过程的生理和分子机制研究进展, 以期为低镉水稻的选育和安全生产提供理论参考。     

镉胁迫对不同甘蓝基因型光合特性和养分吸收的影响

以2个耐镉(Cd)性不同的甘蓝品种为材料,研究了不同Cd浓度(0、20、50、100μmol.L-1)对甘蓝植株生长、叶片光合特性和养分吸收的影响.结果表明:Cd敏感品种在低浓度Cd(20μmol.L-1)处理下生长受到明显抑制,叶片净光合速率(Pn)、气孔导度(Gs)、PSⅡ光化学效率(Fv/Fm)、PSⅡ光合电子传递量子效率(ΦPSⅡ)及地上部、根系干质量显著降低;Cd耐性品种在高浓度Cd(50和100μmol.L-1)处理下生长和光合特性受到显著影响;Cd胁迫降低了甘蓝叶片叶绿素a和b含量,尤其对叶绿素a的影响较大,进而抑制了叶片光合能力.Cd胁迫显著降低了植株对Mn的吸收,抑制了Mg和Fe从根部向地上部的转运,且Cd敏感品种受抑制幅度更大;Cd胁迫促进了Cd耐性品种对P和S的吸收,而Cd敏感品种相反.因此,Cd胁迫下甘蓝敏感品种叶片Mn、Fe、Mg、S和P含量的降低是影响其叶片光合作用,进而抑制植株生长的重要生理原因.     

Comparative expression analysis of heavy metal ATPase subfamily genes between Cd-tolerant and Cd-sensitive turnip landraces

The heavy metal ATPase(HMA)subfamily is mainly involved in heavy metal(HM)tolerance and transport in plants,but an understanding of the definite roles and mechanisms of most HMA members are still limited.In the present study,we identified 14 candidate HMA genes named BrrHMAl—BrrHMA8 from the turnip genome and analyzed the phylogeny,gene structure,chromosome distribution,and conserved domains and motifs of HMAs in turnip(Brassica rapa var.rapa).According to our phylogenetic tree,the BrrHMAs are divided into a Zn/Cd/Co/Pb subclass and Cu/Ag subclass.The BrrHMA members show similar structural characteristics within subclasses.To explore the roles of BrrHMAs in turnip,we compared the gene sequences and expression patterns of the BrrHMA genes between a Cd-tolerant landrace and a Cd-sensitive landrace.Most BrrHMA genes showed similar spatial expression patterns in both Cd-tolerant and Cd-sensitive turnip landraces;some BrrHMA genes,however,were differentially expressed in specific tissue in Cd-tolerant and Cd-sensitive turnip.Specifically,BrrHMA genes in the Zn/Cd/Co/Pb subclass shared the same coding sequence but were differentially expressed in Cd-tolerant and Cd-sensitive turnip landraces under Cd stress.Our findings suggest that the stable expression and up-regulated expression of BrrHMA Zn/Cd/Co/Pb subclass genes under Cd stress may contribute to the higher Cd tolerance of turnip landraces.     

Genotype-Dependent Effect of Exogenous Nitric Oxide on Cd-induced Changes in Antioxidative Metabolism, Ultrastructure, and Photosynthetic Performance in Barley Seedlings (Hordeum vulgare)

A greenhouse hydroponic experiment was performed using Cd-sensitive (cv. Dong 17) and Cd-tolerant (Weisuobuzhi) barley seedlings to evaluate how different genotypes responded to cadmium (Cd) toxicity in the presence of sodium nitroprusside (SNP), a nitric oxide (NO) donor. Results showed that 5 μM Cd increased the accumulation of O₂^•−, H₂O₂, and malondialdehyde (MDA) but reduced plant height, chlorophyll content, net photosynthetic rate (P _n), and biomass, with a much more severe response in the Cd-sensitive genotype. Antioxidant enzyme activities increased significantly under Cd stress in the roots of the tolerant genotype, whereas in leaves of the sensitive genotype, superoxide dismutase (SOD) and ascorbate peroxide (APX), especially cytosol ascorbate peroxidase (cAPX), decreased after 5–15 days Cd exposure. Moreover, Cd induces NO synthesis by stimulating nitrate reductase and nitric oxide synthetase-like enzymes in roots/leaves. A Cd-induced NO transient increase in roots of the Cd-tolerant genotype might partly contribute to its Cd tolerance. Exogenous NO dramatically alleviated Cd toxicity, markedly diminished Cd-induced reactive oxygen species (ROS) and MDA accumulation, ameliorated Cd-induced damage to leaf/root ultrastructure, and increased chlorophyll content and P _n. External NO counteracted the pattern of alterations in certain antioxidant enzymes induced by Cd; for example, it significantly elevated the depressed SOD, APX, and catalase (CAT) activities in the Cd-sensitive genotype after 10- and 15-day treatments. Furthermore, NO significantly increased stromal APX and Mn-SOD activities in both genotypes and upregulated Cd-induced decrease in cAPX activity and gene expression of root/leaf cAPX and leaf CAT1 in the Cd-sensitive genotype. These data suggest that under Cd stress, NO, as a potent antioxidant, protects barley seedlings against oxidative damage by directly and indirectly scavenging ROS and helps to maintain stability and integrity of the subcellular structure.     

Phytochelatins in Cadmium-Sensitive and Cadmium-Tolerant Silene vulgaris (Chain Length Distribution and Sulfide Incorporation)

In response to a range of Cd concentrations, the root tips of Cd-tolerant plants of Silene vulgaris exhibit a lower rate of PC production accompanied by a lower rate of longer chain PC synthesis than those of Cd-sensitive plants. At the same Cd exposure level, stable PC-Cd complexes are more rapidly formed in the roots of Cd-sensitive plants than in those of tolerant plants. At an equal PC concentration in the roots, the PC composition and the amount of sulfide incorporated per unit of PC-thiol is the same in both populations. Although these compounds might play some role in mechanisms that contribute to Cd detoxification, the ability to produce these compounds in greater amounts is not, itself, the mechanism that produces increased Cd tolerance in tolerant S. vulgaris plants.     

In Vitro Evolution and Preliminary Characterization of a Cadmium-Resistant Population of Chlamydomonas reinhardtii

A cadmium-tolerant population of Chlamydomonas reinhardtii was derived from a Cd-sensitive cell wall-deficient strain by long-term selection in liquid culture. A comparison of Cd-sensitive and Cd-tolerant cells revealed that Cd tolerance was due to genetically determined alterations of metabolism rather than to increased efficiency of a detoxification system.