Cadmium transport and tolerance in rice: perspectives for reducing grain cadmium accumulation

Cadmium (Cd) is a toxic heavy metal which harms human health. In Japan, a major source of human Cd-intake is rice grains and contamination of paddy soils by Cd and accumulation of Cd in rice grains are the serious agricultural issues. There also exist Cd contamination of rice and its toxicity in several populations in countries including China and Thailand. Understanding the Cd transport mechanisms in rice can be a basis for regulating rice Cd transport and accumulation by molecular engineering and marker-assisted breeding. Recently, a number of studies have revealed the behavior of Cd in rice, genetic diversity of Cd accumulation, quantitative trait loci controlling Cd accumulation and transporter molecules regulating Cd accumulation and distribution in rice. In this article, we summarize recent advances in the field and discuss perspectives to reduce grain Cd contents.     

水稻植株镉积累分配的差异

采用盆栽土培法，研究了高镉型与低镉型水稻品种植株镉积累、分布的差异。研究表明，两类型水稻品种根系与茎鞘镉含量差异不显著，糙米镉含量差异最大，达极显著水平。低积累型品种根系、叶片镉积累小于高镉型品种，但差异不显著，而其茎鞘镉积累量、茎鞘镉与植株镉总量比值都大于高镉型水稻品种。这可能是低镉型水稻品种糙米锅吉量极显著小于高镉型品种的主要原因。     

Effects of Exogenous 5-Aminolevulinic Acid and 24-Epibrassinolide on Cd Accumulation in Rice from Cd-Contaminated Soil

High grain-Cd-accumulating rice variety Yongyou 9 was planted in Cd-contaminated farmland in Taizhou City, Zhejiang Province, China to study the effects of 5-aminolevulinic acid (ALA) and 24-epibrassinolide (EBR) on Cd accumulation in brown rice. Results showed that the exogenous ALA and EBR had no significant effects on agronomic traits, soil pH and total Cd content in soil, but had some effects on the available Cd content in soil, and significantly influenced the Cd accumulation in the different parts of rice. Results also showed that 100 mg/L exogenous ALA significantly reduced the Cd accumulation in brown rice to blow the food safety standard (0.2 mg/kg), and also significantly reduced the Cd contents in the roots and culm of rice. However, 200 mg/L exogenous ALA treatment increased the Cd content in brown rice remarkably. In addition, 0.15 mg/L EBR treatment increased Cd accumulation in roots, culm, leaves and brown rice notably, whereas 0.30 mg/L exogenous EBR treatment reduced the Cd accumulation in brown rice properly, but it was not significant. Therefore, proper concentration of ALA can effectively reduce the Cd accumulation in brown rice, which can be used as an effective technical method for the safe production of rice in Cd polluted farmland.     

Cadmium Accumulation and Its Toxicity in Brittle Culm 1 （bcl）, a Fragile Rice Mutant

Cadmium （Cd） accumulation and toxicity in rice plants were characterized and identified by using brittle culm 1 （bcl）, a fragile rice mutant and its wild type （Shuangkezao, an indica rice） as materials by hydroponics. The low Cd level didn＇t obviously affect the growth parameters in both rice genotypes, but under high Cd levels （1.0 and 5.0 μmol/L）, the growth of both rice plants were substantially inhibited. Moreover, bcl tended to suffer more seriously from Cd toxicity than Shuangkezao. Cd accumulation in both rice plants increased with the increase of Cd levels. There was a significant difference in Cd accumulation between the two rice genotypes with constantly higher Cd concentration in bcl, which also accumulated more Cd at 0, 0.1, and 1.0 μmol/L Cd levels. The same case was found in the two rice plants grown on Cd-contaminated soil. This suggested that cell wall might play an important role in Cd accumulation in rice plants by the physiological mechanisms. The malondialdehyde （MDA） content, superoxide dismutase （SOD） and peroxidase （POD） activities in rice plants were affected differently under Cd treatments, and which implied that POD might play the main role in detoxifying active oxygen free radical. A significant difference in antioxidative system between the two rice genotypes was found with constantly higher MDA content, SOD and POD activities in bcl. In summary, bcl accumulated more Cd and appeared to be more sensitive to Cd stress compared with its wild type.     

Cd Toxicity and Accumulation in Rice Plants Vary with Soil Nitrogen Status and Their Genotypic Difference can be Partly Attributed to Nitrogen Uptake Capacity

Two indica rice genotypes, viz. Milyang 46 and Zhenshan 97B differing in Cd accumulation and tolerance were used as materials in a hydroponic system consisting of four Cd levels (0, 0.1, 1.0 and 5.0 μmol/L) and three N levels (23.2, 116.0 and 232.0 mg/L) to study the effects of nitrogen status and nitrogen uptake capacity on Cd accumulation and tolerance in rice plants. N-efficient rice genotype, Zhenshan 97B, accumulated less Cd and showed higher Cd tolerance than N-inefficient rice genotype, Milyang 46. There was consistency between nitrogen uptake capacity and Cd tolerance in rice plants. Increase of N level in solution slightly increased Cd concentration in shoots but significantly increased in roots of both genotypes. Compared with the control at low N level, Cd tolerance in both rice genotypes could be significantly enhanced under normal N level, but no significant difference was observed between the Cd tolerances under normal N (116.0 mg/L) and high N (232.0 mg/L) conditions. The result proved that genotypic differences in Cd accumulation and toxicity could be, at least in part, attributed to N uptake capacity in rice plants.     

晚粳稻籽粒中砷、镉、铬、镍、铅等重金属含量的品种和粒位效应

 选用籽粒着粒密度差异较大的密穗型品种秀水63和散穗型品种秀水11，种植于受重金属轻度污染的土壤，研究了晚粳稻籽粒中有毒重金属As、Cd、Cr、Ni、Pb含量的品种与粒位效应。结果表明，重金属含量的粒位间差异秀水63明显大于秀水11，As、Cd、Cr、Pb含量粒位间差异显著， Ni含量差异较小。穗上部籽粒As、Cd、Ni含量高于中部籽粒，穗下部籽粒最低，而Cr、Pb则正好相反。稻米中几种重金属含量与粒重的关系因重金属种类不同存在着差异，籽粒As、Cd、Ni含量与粒重呈正相关，而Cr和Pb含量与粒重呈负相关。秀水63和秀水11不同粒位的籽粒灌浆模式存在差异，下位粒灌浆速率较低，使粒重较上、中位粒低，导致粒重有较大的粒位差异，而这种差异也与穗部着粒密度有关，秀水63明显大于秀水11。籽粒As、Cd、Ni含量与最大灌浆速率（GRm）呈正相关，与到达最大灌浆速率时间（Tpoi）呈负相关，而Cr和Pb含量与以上两个籽粒灌浆特性参数分别呈负相关和正相关，说明籽粒As、Cd、Ni的积累与碳水化合物的积累具有相同或相近的模式，而Cr和Pb表现为不同的积累模式。     

杂交水稻不同器官镉浓度与累积量

以生长于盆钵中的两系杂交稻两优培九、扬两优6号和103S/郑粳2号及三系杂交稻丰优香占、K优818和汕优63为材料,移栽前盆土内加入镉（Cd）,使土壤Cd浓度为100 mg/kg,研究水稻植株不同器官Cd的浓度和累积量。与未加Cd的对照相比,Cd处理降低了两优培九和103S/郑粳2号的产量,其主要原因是每穗粒数减少,而对其余组合的产量无显著影响。水稻植株对Cd吸收及分配存在着基因型差异,以汕优63累积量最多,K优818累积量最少。水稻植株不同器官Cd的浓度和累积量的大小顺序为根>茎鞘>叶片。籽粒不同部位Cd的浓度大小顺序为糠层>颖壳>精米。精米中Cd累积量仅为谷粒Cd累积量的30%左右。在抽穗期和成熟期,同一器官Cd浓度及累积量呈显著或极显著正相关,但不同器官之间的相关不显著。在6个供试的杂交稻组合中,精米中Cd的浓度以扬两优6号最高,K优818最低。对不同杂交稻组合植株中Cd浓度和累积量差异的原因进行了讨论。     

杂交稻头季与再生季镉积累分配特性差异研究

【目的】探究杂交稻头季与再生季镉积累分配特性差异,为再生稻安全生产提供科学依据。【方法】以Y两优9918和甬优4149为材料,采用随机区组设计开展大田试验,比较头季与再生季产量与镉积累分配特性。【结果】1）甬优4149再生季产量显著低于头季,而Y两优9918表现相反;两品种再生季有效穗数、结实率显著高于头季,而千粒重显著低于头季;2）两品种头季成熟期根、茎、叶、穗镉含量均显著低于再生季,再生季糙米镉含量为0.13~0.17 mg/kg,显著高于头季;3）再生季各器官镉含量、镉积累量、日均镉积累速率、镉转移系数与富集系数均大于头季,Y两优9918与甬优4149再生季镉总积累量分别是头季的4.28和2.67倍,再生季糙米镉含量分别是头季的1.63和1.42倍;4）头季穗部镉主要来自灌浆中期-成熟期,而再生季主要来自齐穗前,镉积累最快阶段存在品种间差异;5）两品种稻桩镉积累量在再生季全生育期内表现累积趋势,但各生育阶段的表现存在品种间差异,Y两优9918以灌浆中期为界先降后升,甬优4149表现先降后升再降趋势。6）本研究条件下,Y两优9918头季产量低于甬优4149,但再生季产量表现相反,两品种全年产量差异不大;甬优4149器官镉含量、积累量、日均积累速率及富集系数一般高于Y两优9918。【结论】再生季镉超标风险大于头季,在镉污染稻作区应慎重发展再生稻,同时再生季降镉措施的应用应以齐穗前为重点。     

杂草稻和栽培稻氮代谢对镉胁迫反应的差异

在水培条件下,以杂草稻和粳稻越光为材料,研究了Cd胁迫对水稻氮吸收和氮代谢酶活性的影响。Cd处理设0、01、10和50 μmol/L 4个水平,以硝酸铵为氮源。结果表明,两种水稻的植株Cd含量和积累量均随溶液Cd处理水平的提高而显著增加,但增加程度两水稻材料存在着明显的差异,在较高Cd水平（10和50 μmol/L）下,杂草稻根系Cd含量和积累量低于越光,但地上部呈相反的趋势,表明杂草稻根系吸收的Cd转运至地上部的量高于越光,揭示出杂草稻地上部具有较强的解毒能力。植株氮含量、积累量以及氮代谢有关酶,如硝酸还原酶、谷草转氨酶和谷丙转氨酶等活性随着Cd处理水平的提高而下降,但Cd胁迫对这些参数的抑制程度两水稻材料之间存在着明显差异,总体上对杂草稻的抑制作用较弱。表明杂草稻对Cd的耐性要强于越光,且几种氮代谢酶活性变化与Cd耐性存在着一定的相关。     

镉胁迫对不同水稻基因型植株生长和抗氧化酶系统的影响

 以籽粒镉积累水平不同的两种品种（丙97252，低积累型；秀水63，高积累型）为材料，研究了镉胁迫对水稻植株生长和抗氧化酶系统的影响。采用水培试验，镉处理设0.0、0.1、1.0和5.0 μmol/L 4个水平。结果表明，镉胁迫抑制植株生长和叶绿素合成，改变植株丙二醛（MDA）含量和超氧物歧化酶（SOD）、过氧化氢酶（CAT）、过氧化物酶（POD）活性。在抗氧化酶活性上，根和地上部对镉胁迫的反应存在着差异。总体上，SOD、CAT和POD活性随镉水平的提高而减少，而MDA含量则表现相反。根和地上部MDA含量随着培养液中镉浓度提高而增加，且增加幅度秀水63明显大于丙97252。与对照相比，生长在5.0 μmol/L Cd处理下的植株， SOD活性在孕穗期下降46%~52%，在分蘖期仅下降13%~19%。高浓度镉胁迫下，两品种在MDA含量的增加幅度和叶绿素含量的降低幅度上表现不同，显示出它们对镉的耐性存在着差异。     

Variation among Rice Cultivars in Root Acidification and Its Relation to Cadmium Uptake

Cadmium (Cd) is one of the most toxic pollutants, which can easily enter into food chain through cereal. Excessive intake of Cd by human body may damage kidney and has several other chronic effects [1]. Though plants do not require Cd for growth and development, but the bioaccumulation index of Cd in plants is high and may exceed many essential elements [2]. Moreover, Cd may pose a risk to human and animal health at plant tissue concentrations that are not generally phytotoxic [3]. Therefore…     

不同水平Cd胁迫下低累积Cd水稻品种筛选

通过盆栽试验,研究了10个水稻品种在不同水平Cd胁迫下稻米对Cd的吸收累积情况。结果表明,水稻对Cd具有很强的吸收累积能力,但不同品种对Cd的吸收存在显著差异,相同条件下不同水稻品种稻米的Cd最高与最低含量差异值高达7.36 mg/kg;通过测定叶绿素相对含量（SPAD值）发现,随着添加的外源Cd浓度增加,水稻SPAD值呈现出先增加后减少的趋势,说明低浓度Cd可促使水稻生长,高浓度Cd则会抑制其生长;经聚类分析筛选出1个低累积Cd水稻品种——F优498,其稻米Cd含量仅为0.15 mg/kg（不添加外源Cd条件下）,低于国家食品卫生安全标准规定的限定值（0.20 mg/kg）。相关性研究发现,水稻普遍对土壤Cd有良好的吸收作用,稻米会随着土壤Cd含量的增加大量累积Cd,因此,筛选得到的Cd低累积水稻品种也只适合栽种于无污染或Cd轻度污染区域。     

Root Morphology and Anatomy Affect Cadmium Translocation and Accumulation in Rice

Paddy fields contaminated with cadmium (Cd) present decreased grain yield and produce Cd-contaminated grains. Screening for low-Cd-accumulating cultivars is a useful method to reduce the amount of Cd in the grains. The present study aimed to examine the roles of the root morphology and anatomy in Cd translocation and accumulation in rice plants. Twenty-two rice cultivars were used in the first experiment, after which two cultivars [Zixiangnuo (ZXN) and Jinyou T36 (JYT36)] were selected and used in subsequent experiments under hydroponic conditions. The results showed that there were significant differences in Cd concentrations in the shoots (ranging from 4 to 100 mg/kg) and the Cd translocation rates (shoot/root) (from 7% to 102%) among the 22 cultivars, and the shoot Cd concentration was significantly correlated with the Cd translocation rate of the 22 cultivars under 0.1 mg/L Cd treatment. Compared with cultivar ZXN, JYT36 had greater root Cd uptake and accumulation but lower shoot Cd accumulation and Cd translocation rate. The number of root tips per surface area of cultivar ZXN was greater than that of JYT36, while the average root diameter was lower than that of JYT36. Compared with ZXN, JYT36 had stronger apoplastic barriers, and the Casparian bands and suberin lamellae in the root endodermis and exodermis were closer to the root apex in both the control and Cd treatments, especially for suberin lamellae in the root exodermis with Cd treatments, with a difference of 25 mm. The results also showed that, compared with ZXN, JYT36 had greater percentages of Cd bound in cell walls and intracellular Cd but lower Cd concentrations in the apoplastic fluid under the Cd treatment. The results suggested that Cd translocation, rather than root Cd uptake, is a key process that determines Cd accumulation in the rice shoots. The root morphological and anatomical characteristics evidently affect Cd accumulation in the shoots by inhibiting Cd translocation, especially via the apoplastic pathway. It was possible to pre-screen low-Cd-accumulating rice cultivars on the basis of their root morphology, anatomical characteristics and Cd translocation rate at the seedling stage.     

Effect of Grain Position within a Panicle and Variety on As, Cd, Cr, Ni, Pb Concentrations in japonica Rice

Two japonica rice varieties, a compact panicle (CP) Xiushui 63 and a loose panicle (LP) Xiushui 11 were grown in a slightly contaminated paddy field, to determine the effect of grain positions within a panicle and variety on grain As, Cd, Cr, Ni and Pb concentrations. There was a significant difference in grain As, Cd, Cr, Ni and Pb concentrations among grain positions, and the extent of the difference was variety- and metal-specific; Xiushui 63 showed greater difference than Xiushui 11, and As, Cd, Cr and Pb being greater than Ni. The top grains (TG) of a panicle had higher As, Cd and Ni concentrations than the middle ones (MG), and the bottom ones (BG) contained the lowest. With regard to Cr and Pb, the case was opposite. Significantly positive correlations were found between grain weight and As, Cd, Ni concentrations, but there was a negative correlation between grain weight and Cr, and Pb concentrations. There was a remarkable variation in grain filling pattern among grains within a panicle, and between Xiushui 63 and Xiushui 11. In comparison with MG and TG, BG had the lowest grain filling rate and grain weight, leading to obvious difference in grain weight among grains within a panicle, and a greater difference for Xiushui 63 than for Xiushui 11. The regression analysis showed that grain As and Cd concentrations were positively correlated with the maximum grain filling rate (GRm),while negatively correlated with the required time for reaching the maximum grain filling rate (Tpol). Concerning Cr and Pb, the case was just reverse. It is suggested that As and Cd accumulation in grains might be accompanied by the carbohydrate accumulations, while Cr and Pb accumulation displayed a different pattern.     

Influences of Cadmium on Grain Mineral Nutrient Contents of Two Rice Genotypes Differing in Grain Cadmium Accumulation

A pot experiment was conducted to study the effects of Cd on grain Cd, K, P, Mg, Zn, Cu, Pb, Fe and Mn accumulation in two rice genotypes (Xiushui 63 and Xiushui 217) differing in grain Cd accumulation under four Cd levels, i.e. 0, 0.5, 2.5 and 12.5 mg/kg. Rice genotype greatly affected the grain K content, but not significantly for P, Mg, Zn, Cu, Pb, Fe and Mn contents. There were remarkable effects of additional Cd on the contents of P, Mg and Zn in grains, while not significant for K, Cu, Pb, Fe and Mn contents. No significant differences were found in the interaction of genotype by additional Cd on these nine element contents. The low grain Cd accumulation genotype Xiushui 217 had significantly higher grain K, Mg, Cu and Mn contents than the high grain Cd accumulation genotype Xiushui 63, but the case was opposite for Zn, Pb and Fe contents. It also showed that Cd addition levels significantly influenced the K, P, Mg, Zn, Cu, Pb, Fe and Mn contents in rice grains. Grain K, P, Mg, Zn, Fe and Mn contents reduced with the increasing rate of Cd addition.     

水稻镉吸收及其在各器官间转运积累的研究进展

随着土壤和大气沉降物中重金属污染的问题日益严峻,农作物重金属超标时有发生。土壤中的重金属难以降解,容易被植物根部吸收,在作物中积累;大气中的重金属沉降到叶片上,通过叶片吸收进入作物中,最终通过食物链被人类食用,危害人体健康。水稻是我国最重要的粮食作物,大米中存在镉超标问题。镉(Cd)是水稻的非必需元素,主要借助其他金属离子通道蛋白进入水稻根细胞,还可以通过叶片角质层的吸附内化和气孔的渗透作用进入水稻叶片。Cd依赖其他金属离子的转运体在水稻体内运输,Cd从土壤中吸收和植物内部的分配是一个动态过程,通过根吸收转运蛋白、根-地上部转运(木质部转运)和韧皮部的源-库转运(包括种子装载)驱动,实现器官间的转运分配。本文综述了水稻Cd的吸收与转运、器官间Cd的运输分配以及Cd向籽粒转运积累,以期为减少水稻籽粒中Cd含量的研究和解决我国南方部分种植地区Cd污染问题提供一些参考。     

OsNramp5基因变异影响水稻重要农艺性状的研究进展

筛选和培育镉(Cd)低积累水稻品种是解决稻米镉污染问题最经济、有效的办法。现有研究表明OsNramp5是介导水稻Cd吸收最重要的基因,其功能缺失后,水稻籽粒Cd含量极显著下降,但同时会影响水稻必需元素锰(Mn)的吸收,而在前人关于OsNramp5变异影响水稻生长发育的研究中,结论并不一致。系统了解OsNramp5基因变异对水稻重要农艺性状的影响有助于推动低Cd优质水稻新品种的培育。本文重点对OsNramp5基因变异对水稻中金属离子的含量,水稻的生长发育、产量性状及米质的影响进行了综述,以期为利用OsNramp5基因突变选育低Cd积累水稻品种提供参考。     

Genome-wide association study and candidate gene analysis of rice cadmium accumulation in grain in a diverse rice collection

Background

Cadmium (Cd) accumulation in rice followed by transfer to the food chain causes severe health problems in humans. Breeding of low Cd accumulation varieties is one of the most economical ways to solve the problem. However, information on the identity of rice germplasm with low Cd accumulation is limited, particularly in indica, and the genetic basis of Cd accumulation in rice is not well understood.

Results

Screening of 312 diverse rice accessions revealed that the grain Cd concentrations of these rice accessions ranged from 0.12 to 1.23?mg/kg, with 24 accessions less than 0.20?mg/kg. Three of the 24 accessions belong to indica. Japonica accumulated significantly less Cd than indica (p < 0.001), while tropical japonica accumulated significantly less Cd than temperate japonica (p < 0.01). GWAS in all accessions identified 14 QTLs for Cd accumulation, with 7 identified in indica and 7 identified in japonica subpopulations. No common QTL was identified between indica and japonica. The previously identified genes (OsHMA3, OsNRAMP1, and OsNRAMP5) from japonica were colocalized with QTLs identified in japonica instead of indica. Expression analysis of OsNRAMP2, the candidate gene of the novel QTL (qCd3–2) identified in the present study, demonstrated that OsNRAMP2 was mainly induced in the shoots of high Cd accumulation accessions after Cd treatment. Four amino acid differences were found in the open reading frame of OsNRAMP2 between high and low Cd accumulation accessions. The allele from low Cd accumulation accessions significantly increased the Cd sensitivity and accumulation in yeast. Subcellular localization analysis demonstrated OsNRAMP2 expressed in the tonoplast of rice protoplast.

Conclusion

The results suggest that grain Cd concentrations are significantly different among subgroups, with Cd concentrations decreasing from indica to temperate japonica to tropical japonica. However, considerable variations exist within subgroups. The fact that no common QTL was identified between indica and japonica implies that there is a different genetic basis for determining Cd accumulation between indica and japonica, or that some QTLs for Cd accumulation in rice are subspecies-specific. Through further integrated analysis, it is speculated that OsNRAMP2 could be a novel functional gene associated with Cd accumulation in rice.

     

水分管理调控水稻镉污染的研究与应用进展

我国稻田镉污染严重,稻米安全问题突出,发展能够有效调控水稻镉污染的产品与措施尤为迫切。稻田生态系统高度复杂,并易受多种因素影响,水稻镉污染调控难度大。大量研究表明,合理的水分管理不仅能够保障水稻正常的生长发育、产量和稻米品质,同时还能有效减少水稻镉积累。本文综述了水分对土壤理化性质、微生物动态变化以及植物生长发育的影响,阐述了水分管理降低水稻镉积累的主要机理,总结和展望了目前水分管理模式相关研究进展以及未来研究方向,旨在为实际生产上通过优化水分管理技术降低镉污染提供理论依据,更有效实现水稻安全生产。