杂交水稻对土壤Cd、Zn吸收与Cd耐性的基因型差异
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摘要
随着农田Cd污染的不断加剧,Cd在土壤-作物-食物中的活跃迁移对人类健康已构成严重威胁,是已知威胁最严重的农田污染物之一。我国南方主要水稻产区的偏酸、低有机碳和Cd污染日益严峻土壤环境因素和人为因素的叠加使控制水稻,尤其是在确保粮食安全前提下广泛推广种植的具有强吸收养分能力的高产杂交稻对土壤Cd的吸收极为复杂,使我们必须正视稻米Cd污染及其严峻的食物安全风险挑战。植物的Cd耐性以及Cd在植株不同部位器官中的积累与分布存在着显著的种间和品种间差异。因此,筛选籽粒低Cd积累品种是进行稻米安全生产的重要途径。同时,Cd对植物造成的毒害作用以及植物对Cd的吸收积累与耐性强度的基因型差异机理尚未明确,限制了作物包括水稻的安全生产。本研究通过对我国南方广泛种植的110个杂交水稻品种进行大田低Cd品种筛选的大田试验研究了杂交水稻对土壤中Cd、Zn吸收积累的基因型差异,筛选出部分低Cd品种,并在此基础上利用筛选出的籽粒高Cd、低Cd品种在采自江西鹰潭的典型红壤上进行全生育期的水稻盆栽试验,更进一步地明确了杂交水稻Cd、Zn的吸收与籽粒积累规律,并运用透射电镜观察、差速离心法等深入探讨了Cd耐性机理的品种差异,如外加Cd下品种细胞超微结构、抗氧化酶系统清除、亚细胞结构、PCs螯合等方面的差异机制,主要研究结果如下:
一、杂交水稻对土壤中Cd、Zn吸收积累的基因型差异
通过110个杂交水稻品种籽粒对水稻土中Cd、Zn的吸收积累情况的研究表明,供试杂交水稻籽粒Cd、Zn含量的变化范围分别介于0.004-0.057、10.25-30.06 mg·kg(-1)之间,指示不同水稻品种对同一土壤中Cd、Zn吸收及其在籽粒中积累存在显著差异。本试验还观察到相对于普通杂交稻而言,超级稻品种具有籽粒高Cd低Zn的现象。并且用籽粒Cd和籽粒Cd/Zn关系做图,供试杂交稻的斜率高于常规稻(已报道),表明水稻吸Cd同时对Zn的排斥作用强度依基因型而异,杂交稻吸Cd排Zn的作用更明显。采用国家营养协会推荐的每日膳食中营养素参考摄入量(RNI)衡量,对就地消费者而言存在Cd暴露和“Zn饥饿”的潜在食物安全风险,尤其对南方酸性土壤地区食用超级稻品种更甚。因此,在土壤重金属Cd污染地区,必须密切注意高Cd/Zn品种的大规模栽培,而筛选高Zn低Cd的水稻品种进行品质育种是可能的。
以大田筛选的籽粒镉积累水平不同的两品种(J196,低积累型,品种C;中浙优1号,高积累型,品种A)为材料,采用添加Cd(2.5 mg·kg~(-1))和不添加Cd处理在红壤性水稻土进行盆栽试验研究了杂交水稻在成熟期对土壤中Cd、Zn的吸收分配及籽粒Cd、Zn积累特点。结果表明,杂交水稻对Cd的吸收及籽粒Cd、Zn积累存在品种基因型差异。外源Cd的添加使水稻籽粒Cd积累成倍提高。本试验结果还显示,在供试土壤条件下尤其是污染条件下,高Cd品种对Cd的吸收与籽粒积累十分强烈,就地消费人群的籽粒Cd暴露风险水平达到数倍于人体安全临界摄入剂量水平,Zn存在相对缺乏的食物安全风险。高Cd品种具有较强的籽粒Cd分配和较弱的籽粒Zn转运能力,而加Cd处理下,低Cd品种对Cd、Zn的根部、茎叶滞留能力更强,这也正是高籽粒含量品种籽粒Cd积累强而Cd/Zn高的植株因素。
二、外加Cd下水稻Cd耐性机制
以大田筛选的籽粒镉积累水平不同的两品种(J196,低积累型,品种C;中浙优1号,高积累型,品种A)为材料,在红壤性水稻土上采用盆栽实验,镉处理设0,2.5,5,10,25 mg·kg~(-1)5个水平,在第一吸Cd高峰苗期研究了外加cd处理对水稻抗氧化酶系统、NPT含量的影响变化,结果表明:高Cd水稻品种中浙优1号的耐Cd性强于低Cd水稻品种J196,表现为前者体内的MDA含量低于后者,而SOD、CAT、POD酶活性高于J196;在Cd胁迫下,耐性强的品种中浙优1号的SOD活力、CAT活力随Cd处理浓度增加先增后降,在10mg·kg~(-1)时达到峰值,POD活力则为先降后增,在Cd浓度为5 mg·kg~(-1)时达到峰值;而低Cd品种随着外加Cd增长,3种酶活性在25 mg·kg~(-1)达到峰值,除SOD比对照降低外,CAT和POD分较对照升高38.07%和62.10%,表明本试验中,外加Cd胁迫下,两品种中三种抗氧化酶保护作用不同,高Cd品种CAT的保护性最强,低Cd品种POD的保护作用最强。外加Cd胁迫则显著诱导根内合成NPT-SH,外加Cd及品种差异对水稻体内Cd的吸收具有不同程度的影响,根中NPT-SH的诱导量与水稻根系内Cd的含量变化一致。外加Cd处理下,J196根系内NPT-SH含量均高于中浙优1号,从细胞水平上表明和中浙优1号相比,J196可能具有从根系向茎叶的高转运能力。
镉处理设0,5,25mg·kg~(-1)3个水平,以差速离心法在第二吸Cd高峰抽穗期研究了外加Cd处理对水稻亚细胞结构的影响变化,结果表明,外加Cd下,两品种根系细胞的Cd大部分存在于细胞壁中,占52.6-83.2%,少部分存在于细胞可溶部分(细胞质),其镉含量分布的百分率为3.0-10.1%,且根系细胞壁中J196的分布率高于中浙优1号。表明J196的根系细胞壁的固定Cd能力较强;和对照相比,外加Cd促使中浙优1号根系细胞内可溶物向地上部移动,且高Cd处理下,Cd在中浙优1号细胞内溶物的分配率增高程度大于J196,增加了前者叶片中Cd由韧皮部运往籽粒的可能性,从而在亚细胞水平上推测了品种中浙优1号为高转运品种的可能机制。
镉处理设0,25mg·kg~(-1) 2个水平,用透射电镜观察的方法在第二吸Cd高峰抽穗期研究了外加Cd处理对水稻超微结构的影响变化,结果表明Cd处理下两品种的部分根细胞有Cd沉积现象,但不同品种的沉积部位表现不同,高Cd品种主要在液泡边缘而低Cd品种则在细胞壁和细胞间隙。表明不同水稻品种根系的耐Cd机制不同,前者主要表现液泡区室化,而低Cd品种则表现为细胞壁上有特定沉积。和对照相比,外加Cd对茎叶细胞的超微结构造成伤害,主要表现在使叶绿体上淀粉粒消失,叶绿体空泡化,部分线粒体出现肿胀现象,低Cd品种的受害程度强于高Cd品种A,同时也表明水稻叶肉细胞器中叶绿体和线粒体是对Cd胁迫较为敏感的细胞器。
杂交水稻在籽粒Cd、Zn积累,抗氧化酶清除、细胞超微结构等耐性机制存在显著的基因型差异,因此,在高产水稻育种中必须考虑高耐Cd性、低籽粒Cd/Zn比水稻品种的选育,在推广高产杂交水稻根据其Cd耐性及籽粒Cd、Zn积累特性考虑土壤-品种的合理布局并在食物结构调整上配合膳食结构优化以确保就地人群Cd暴露风险的消除及对Zn的营养摄入。
In China,Cd concentration in croplands and subsequent accumulation in crops has been increased as a result of soil polltuion due to human activities such as mining,emission from industry and application of sewage sludges,which is posing potential public health risk for human through dietary intake.Among the major staple crops,rice(Oryza sativa L.) is the particular food crop with high Cd uptake and accumulation in grains.Rice is produced predominantly in South China where the soils are either high in Cd mobility or deficient in Zn or both.However,for meeting the demand by the increasing population with less available arable lands,high-yielding hybrid rice or super rice cultivars with high Cd affinity are under extension,which may raise serious potential health concerns for Chinese subsistence diet farmers.However,there had been much evidences of differences in Cd accumulation in plants and plant tolerance to between rice species and genotypes.
For food security,it is imperative to breed cultivars with low Cd accumulation in grains.Meanwhile,Cd tolerance and low Cd accumulation in grains may be used as a strategy for low Cd rice production.However,the mechanisms involved in Cd tolerance and grain accumulation has been not clearly demonstrated.In the dissertation,an experiment of 110 cultivars including super rice and common hybrid rice cultivars was grown on a single paddy soil(Fluventic Stagnic Anthrosol) with a neutral reaction and low in total Cd in 2006 in order to depict the difference in grain Cd and Zn between rice cultivars.Furthermore,a pot experiment with 2 selected cultivars Zhongzheyou 1(high grain Cd accumulation) and J196(high grain Cd accumulation) of hybrid rice was conducted for the whole growing period using acidic red earth derived paddy soil,a Typical Hapludults which was collected from Yingtan City,Jiangxi Province.The up-take, accumulation and distribution of Cd and Zn in plant organs of hybrid rice and the Cd partitioning at sub-cell level were analysed for genotype tolerance of Cd using transmission electron microscopy observation and differential characterization of the antioxidative enzyme elimination,cell ultrastructure,subcellular structure and PCs chelating in response to Cd stress.The main results were as follows:
1.Genotypic difference on Cd,Zn absorption and accumulation of hybrid rice
Grain Cd and Zn concentrations of the totally 110 cultivars in the field experiment were determined with GFAAS and AAS respectively.Wider variation of grain Cd was found in a range of 0.004-0.057 mg·kg~(-1) than of Zn in a range of 10.25-30.06 mg·kg~(-1) among the cultivars.Higher Cd but lower Zn concentration in grains of super rice cultivars was observed than of the common hybrid ones.Although a significantly negative linear correlation of grain Cd with grain Zn was found for both super rice and common hybrid cultivars,with much higher slope for the hybrid cultivars than the reported non-hybrid cultivars.Using the RNI value recommended by CFSG,calculated potential risk of food Cd exposure with 'Zn hungry' through subsistence diet intake was prominent with all the studied 110 hybrid rice cultivars,possessing high potential health problems for rice production in South China using the super rice cultivars.Thus,breeding of genotypes of rice cultivars with low grain Cd and low Cd/Zn ratio is needed for rice production in soils of South China where high Cd bioavailability is prevalent with acidic red soils.
In the pot experiment of Cd and Zn uptake and portioning in organs in maturity with Red earth derived paddy soil was analysed.Cd was spiked at 0(as the control) and 2.5 mg·kg~(-1) of soil,respectively.Two cultivars of hybrid rice,Zhongzheyou 1(high grain Cd cultivar) and J196(low grain Cd cultivar),were harvested and the biomass,Cd and Zn contents,of different organs were determined.Significant genotype differences in Cd,Zn uptake and grain partitioning was observed.A phenomenon of intense Cd accumulation in grains by super rice and,thus,imposing a very high Cd exposure and Zn depletion risk(as several times as the acceptable daily intake,ADI) to subsistence-diet farmers was demonstrated.The low Cd cultivar J196 tended to hamper the up-taken Cd in root and stem while the super rice Zhongzheyou 1 promoted higher partitioning to grain.Furthermore, stronger Cd but weaker Zn transfering to grains by the high grain Cd accumulation cultivars than the low one could be considered as a major plant factor controlling high Cd uptake and accumulation,and the high Cd/Zn ratio in rice grain.
2.Genotypie difference in Cd tolerance
The genotypic difference in the changes of three antioxidative enzymes(SOD,POD and CAT) activities and the MDA contents under cd spiking were characterized.The results showed Cd stress altered malondialdehyde(MDA) content and activities of superoxide dismutase(SOD),calase(CAT) and peroxidase(POD) in rice plant.The response to Cd stress differed in terms of MDA contents and antioxidant enzyme activity.Generally,the activities of SOD,CAT firstly ascend then droped with increased Cd level,in contrast to that of POD.While MDA contents was found higher in cultivar J196,but SOD,CAT,POD tend to increase in Zhongzheyou 1.The higher MDA and smaller antioxidant enzyme activity were found in J196,indicating that J196 was more sensitive to Cd stress than Zhongzheyou 1.
The effect of Cd-spiking treatment on Cd content and nonprotein thiols(NPT) production in rice was also studied using the pot experiment The study demonstrated a phenomenon that Cd stress had significant inhibitory effects on NPT overproduction.The Cd uptake of rice was affected by the Cd-spiking treatment and the difference between genotypes.Furthermore,the Cd content was enhanced under Cd spiking,coinciding with the enhancement of NPT level in root.Root NPT contents of J196 was higher than those of Zhongzheyou-1,suggested that J196 has the higher Cd transfer potential from root to stem compared to Zhongzheyou-1.
In an effort to understand the process of Cd bioaccumulation,subcelluar partitioning of Cd were characterized with subcelluar distribution of Cd in roots and leaves of rice.The results showed that there was a genotypic difference in cell Cd concentrations between two cultivars.Cd concentrations in different plant parts were as follows:root＞leaf;Cd was mainly distributed in the cell wall,about 52.6-83.2%,and less distributed in the fractions of cell soluble component with distribution rate of 3.0-10.1%.The cell wall distribution of Cd by J196 was higher than Zhongzheyou-1,showing a stronger cell wall compartmentation of Cd in J196 than in Zhongzheyou-1;Under Cd-spiking,Cd transfer from root cell soluble component transfering to shoot via xylem by Zhongzheyou-1 was promoted.The increased Cd distribution in cell soluble component by Zhongzheyou-1 compared to by J196 may be account for the lower Cd in stem and more Cd traslocation from shoot to grain via phloem.
To examine the rice(Oryza sativa L.) genotypic difference in the alternation of cell ultrastructure,comparative electronic microscopic observation was undertaken of cells from the two hybrid rice cultivars(Zhongzheyou 1,high grain Cd accumulation and J196,low grain Cd accumulation) under Cd spiked at 0 and 25 mg·kg~(-1).Under Cd spiking, Cd was presnet in binding to cell wall and accumulating in vacuole in rice root cell of both cultivars.The leaf cell ultrastructure of two rice cultivars presented the similar symptom: Chloroplasts and mitochondria were swelling up;and thylakoids vacuolized apparently. However,Cd toxicity symptom existed in the rice cultivar-J196,making mitochondrion mesenchyme dissolve and mitochondria was on the verge of disorganization.Tolerance of cell ultrastructure of rice cultivar-Zhongzheyou 1 was stronger than that of J196.And it demonstrated a phenomenon of mitochondria and chloroplast of rice mesophyll cells,which were the most sensitive organelles.
In summary,the significant genotypic difference in the Cd and Zn accumulation and distribution of hybrid rice and Cd tolerance mechanisms existed between the two cultivars. Therefore,it is possible to develop the cultivars with high tolerance and low grain Cd/Zn accumulation in rice breeding forh high yield.It is also suggested the practical measures should be taken such as rational soil-cultivar combination and optimization dietary structure for controling the human Cd food exposure and guarantee Zn nutrition intake by diet.
一、杂交水稻对土壤中Cd、Zn吸收积累的基因型差异
通过110个杂交水稻品种籽粒对水稻土中Cd、Zn的吸收积累情况的研究表明,供试杂交水稻籽粒Cd、Zn含量的变化范围分别介于0.004-0.057、10.25-30.06 mg·kg(-1)之间,指示不同水稻品种对同一土壤中Cd、Zn吸收及其在籽粒中积累存在显著差异。本试验还观察到相对于普通杂交稻而言,超级稻品种具有籽粒高Cd低Zn的现象。并且用籽粒Cd和籽粒Cd/Zn关系做图,供试杂交稻的斜率高于常规稻(已报道),表明水稻吸Cd同时对Zn的排斥作用强度依基因型而异,杂交稻吸Cd排Zn的作用更明显。采用国家营养协会推荐的每日膳食中营养素参考摄入量(RNI)衡量,对就地消费者而言存在Cd暴露和“Zn饥饿”的潜在食物安全风险,尤其对南方酸性土壤地区食用超级稻品种更甚。因此,在土壤重金属Cd污染地区,必须密切注意高Cd/Zn品种的大规模栽培,而筛选高Zn低Cd的水稻品种进行品质育种是可能的。
以大田筛选的籽粒镉积累水平不同的两品种(J196,低积累型,品种C;中浙优1号,高积累型,品种A)为材料,采用添加Cd(2.5 mg·kg~(-1))和不添加Cd处理在红壤性水稻土进行盆栽试验研究了杂交水稻在成熟期对土壤中Cd、Zn的吸收分配及籽粒Cd、Zn积累特点。结果表明,杂交水稻对Cd的吸收及籽粒Cd、Zn积累存在品种基因型差异。外源Cd的添加使水稻籽粒Cd积累成倍提高。本试验结果还显示,在供试土壤条件下尤其是污染条件下,高Cd品种对Cd的吸收与籽粒积累十分强烈,就地消费人群的籽粒Cd暴露风险水平达到数倍于人体安全临界摄入剂量水平,Zn存在相对缺乏的食物安全风险。高Cd品种具有较强的籽粒Cd分配和较弱的籽粒Zn转运能力,而加Cd处理下,低Cd品种对Cd、Zn的根部、茎叶滞留能力更强,这也正是高籽粒含量品种籽粒Cd积累强而Cd/Zn高的植株因素。
二、外加Cd下水稻Cd耐性机制
以大田筛选的籽粒镉积累水平不同的两品种(J196,低积累型,品种C;中浙优1号,高积累型,品种A)为材料,在红壤性水稻土上采用盆栽实验,镉处理设0,2.5,5,10,25 mg·kg~(-1)5个水平,在第一吸Cd高峰苗期研究了外加cd处理对水稻抗氧化酶系统、NPT含量的影响变化,结果表明:高Cd水稻品种中浙优1号的耐Cd性强于低Cd水稻品种J196,表现为前者体内的MDA含量低于后者,而SOD、CAT、POD酶活性高于J196;在Cd胁迫下,耐性强的品种中浙优1号的SOD活力、CAT活力随Cd处理浓度增加先增后降,在10mg·kg~(-1)时达到峰值,POD活力则为先降后增,在Cd浓度为5 mg·kg~(-1)时达到峰值;而低Cd品种随着外加Cd增长,3种酶活性在25 mg·kg~(-1)达到峰值,除SOD比对照降低外,CAT和POD分较对照升高38.07%和62.10%,表明本试验中,外加Cd胁迫下,两品种中三种抗氧化酶保护作用不同,高Cd品种CAT的保护性最强,低Cd品种POD的保护作用最强。外加Cd胁迫则显著诱导根内合成NPT-SH,外加Cd及品种差异对水稻体内Cd的吸收具有不同程度的影响,根中NPT-SH的诱导量与水稻根系内Cd的含量变化一致。外加Cd处理下,J196根系内NPT-SH含量均高于中浙优1号,从细胞水平上表明和中浙优1号相比,J196可能具有从根系向茎叶的高转运能力。
镉处理设0,5,25mg·kg~(-1)3个水平,以差速离心法在第二吸Cd高峰抽穗期研究了外加Cd处理对水稻亚细胞结构的影响变化,结果表明,外加Cd下,两品种根系细胞的Cd大部分存在于细胞壁中,占52.6-83.2%,少部分存在于细胞可溶部分(细胞质),其镉含量分布的百分率为3.0-10.1%,且根系细胞壁中J196的分布率高于中浙优1号。表明J196的根系细胞壁的固定Cd能力较强;和对照相比,外加Cd促使中浙优1号根系细胞内可溶物向地上部移动,且高Cd处理下,Cd在中浙优1号细胞内溶物的分配率增高程度大于J196,增加了前者叶片中Cd由韧皮部运往籽粒的可能性,从而在亚细胞水平上推测了品种中浙优1号为高转运品种的可能机制。
镉处理设0,25mg·kg~(-1) 2个水平,用透射电镜观察的方法在第二吸Cd高峰抽穗期研究了外加Cd处理对水稻超微结构的影响变化,结果表明Cd处理下两品种的部分根细胞有Cd沉积现象,但不同品种的沉积部位表现不同,高Cd品种主要在液泡边缘而低Cd品种则在细胞壁和细胞间隙。表明不同水稻品种根系的耐Cd机制不同,前者主要表现液泡区室化,而低Cd品种则表现为细胞壁上有特定沉积。和对照相比,外加Cd对茎叶细胞的超微结构造成伤害,主要表现在使叶绿体上淀粉粒消失,叶绿体空泡化,部分线粒体出现肿胀现象,低Cd品种的受害程度强于高Cd品种A,同时也表明水稻叶肉细胞器中叶绿体和线粒体是对Cd胁迫较为敏感的细胞器。
杂交水稻在籽粒Cd、Zn积累,抗氧化酶清除、细胞超微结构等耐性机制存在显著的基因型差异,因此,在高产水稻育种中必须考虑高耐Cd性、低籽粒Cd/Zn比水稻品种的选育,在推广高产杂交水稻根据其Cd耐性及籽粒Cd、Zn积累特性考虑土壤-品种的合理布局并在食物结构调整上配合膳食结构优化以确保就地人群Cd暴露风险的消除及对Zn的营养摄入。
In China,Cd concentration in croplands and subsequent accumulation in crops has been increased as a result of soil polltuion due to human activities such as mining,emission from industry and application of sewage sludges,which is posing potential public health risk for human through dietary intake.Among the major staple crops,rice(Oryza sativa L.) is the particular food crop with high Cd uptake and accumulation in grains.Rice is produced predominantly in South China where the soils are either high in Cd mobility or deficient in Zn or both.However,for meeting the demand by the increasing population with less available arable lands,high-yielding hybrid rice or super rice cultivars with high Cd affinity are under extension,which may raise serious potential health concerns for Chinese subsistence diet farmers.However,there had been much evidences of differences in Cd accumulation in plants and plant tolerance to between rice species and genotypes.
For food security,it is imperative to breed cultivars with low Cd accumulation in grains.Meanwhile,Cd tolerance and low Cd accumulation in grains may be used as a strategy for low Cd rice production.However,the mechanisms involved in Cd tolerance and grain accumulation has been not clearly demonstrated.In the dissertation,an experiment of 110 cultivars including super rice and common hybrid rice cultivars was grown on a single paddy soil(Fluventic Stagnic Anthrosol) with a neutral reaction and low in total Cd in 2006 in order to depict the difference in grain Cd and Zn between rice cultivars.Furthermore,a pot experiment with 2 selected cultivars Zhongzheyou 1(high grain Cd accumulation) and J196(high grain Cd accumulation) of hybrid rice was conducted for the whole growing period using acidic red earth derived paddy soil,a Typical Hapludults which was collected from Yingtan City,Jiangxi Province.The up-take, accumulation and distribution of Cd and Zn in plant organs of hybrid rice and the Cd partitioning at sub-cell level were analysed for genotype tolerance of Cd using transmission electron microscopy observation and differential characterization of the antioxidative enzyme elimination,cell ultrastructure,subcellular structure and PCs chelating in response to Cd stress.The main results were as follows:
1.Genotypic difference on Cd,Zn absorption and accumulation of hybrid rice
Grain Cd and Zn concentrations of the totally 110 cultivars in the field experiment were determined with GFAAS and AAS respectively.Wider variation of grain Cd was found in a range of 0.004-0.057 mg·kg~(-1) than of Zn in a range of 10.25-30.06 mg·kg~(-1) among the cultivars.Higher Cd but lower Zn concentration in grains of super rice cultivars was observed than of the common hybrid ones.Although a significantly negative linear correlation of grain Cd with grain Zn was found for both super rice and common hybrid cultivars,with much higher slope for the hybrid cultivars than the reported non-hybrid cultivars.Using the RNI value recommended by CFSG,calculated potential risk of food Cd exposure with 'Zn hungry' through subsistence diet intake was prominent with all the studied 110 hybrid rice cultivars,possessing high potential health problems for rice production in South China using the super rice cultivars.Thus,breeding of genotypes of rice cultivars with low grain Cd and low Cd/Zn ratio is needed for rice production in soils of South China where high Cd bioavailability is prevalent with acidic red soils.
In the pot experiment of Cd and Zn uptake and portioning in organs in maturity with Red earth derived paddy soil was analysed.Cd was spiked at 0(as the control) and 2.5 mg·kg~(-1) of soil,respectively.Two cultivars of hybrid rice,Zhongzheyou 1(high grain Cd cultivar) and J196(low grain Cd cultivar),were harvested and the biomass,Cd and Zn contents,of different organs were determined.Significant genotype differences in Cd,Zn uptake and grain partitioning was observed.A phenomenon of intense Cd accumulation in grains by super rice and,thus,imposing a very high Cd exposure and Zn depletion risk(as several times as the acceptable daily intake,ADI) to subsistence-diet farmers was demonstrated.The low Cd cultivar J196 tended to hamper the up-taken Cd in root and stem while the super rice Zhongzheyou 1 promoted higher partitioning to grain.Furthermore, stronger Cd but weaker Zn transfering to grains by the high grain Cd accumulation cultivars than the low one could be considered as a major plant factor controlling high Cd uptake and accumulation,and the high Cd/Zn ratio in rice grain.
2.Genotypie difference in Cd tolerance
The genotypic difference in the changes of three antioxidative enzymes(SOD,POD and CAT) activities and the MDA contents under cd spiking were characterized.The results showed Cd stress altered malondialdehyde(MDA) content and activities of superoxide dismutase(SOD),calase(CAT) and peroxidase(POD) in rice plant.The response to Cd stress differed in terms of MDA contents and antioxidant enzyme activity.Generally,the activities of SOD,CAT firstly ascend then droped with increased Cd level,in contrast to that of POD.While MDA contents was found higher in cultivar J196,but SOD,CAT,POD tend to increase in Zhongzheyou 1.The higher MDA and smaller antioxidant enzyme activity were found in J196,indicating that J196 was more sensitive to Cd stress than Zhongzheyou 1.
The effect of Cd-spiking treatment on Cd content and nonprotein thiols(NPT) production in rice was also studied using the pot experiment The study demonstrated a phenomenon that Cd stress had significant inhibitory effects on NPT overproduction.The Cd uptake of rice was affected by the Cd-spiking treatment and the difference between genotypes.Furthermore,the Cd content was enhanced under Cd spiking,coinciding with the enhancement of NPT level in root.Root NPT contents of J196 was higher than those of Zhongzheyou-1,suggested that J196 has the higher Cd transfer potential from root to stem compared to Zhongzheyou-1.
In an effort to understand the process of Cd bioaccumulation,subcelluar partitioning of Cd were characterized with subcelluar distribution of Cd in roots and leaves of rice.The results showed that there was a genotypic difference in cell Cd concentrations between two cultivars.Cd concentrations in different plant parts were as follows:root＞leaf;Cd was mainly distributed in the cell wall,about 52.6-83.2%,and less distributed in the fractions of cell soluble component with distribution rate of 3.0-10.1%.The cell wall distribution of Cd by J196 was higher than Zhongzheyou-1,showing a stronger cell wall compartmentation of Cd in J196 than in Zhongzheyou-1;Under Cd-spiking,Cd transfer from root cell soluble component transfering to shoot via xylem by Zhongzheyou-1 was promoted.The increased Cd distribution in cell soluble component by Zhongzheyou-1 compared to by J196 may be account for the lower Cd in stem and more Cd traslocation from shoot to grain via phloem.
To examine the rice(Oryza sativa L.) genotypic difference in the alternation of cell ultrastructure,comparative electronic microscopic observation was undertaken of cells from the two hybrid rice cultivars(Zhongzheyou 1,high grain Cd accumulation and J196,low grain Cd accumulation) under Cd spiked at 0 and 25 mg·kg~(-1).Under Cd spiking, Cd was presnet in binding to cell wall and accumulating in vacuole in rice root cell of both cultivars.The leaf cell ultrastructure of two rice cultivars presented the similar symptom: Chloroplasts and mitochondria were swelling up;and thylakoids vacuolized apparently. However,Cd toxicity symptom existed in the rice cultivar-J196,making mitochondrion mesenchyme dissolve and mitochondria was on the verge of disorganization.Tolerance of cell ultrastructure of rice cultivar-Zhongzheyou 1 was stronger than that of J196.And it demonstrated a phenomenon of mitochondria and chloroplast of rice mesophyll cells,which were the most sensitive organelles.
In summary,the significant genotypic difference in the Cd and Zn accumulation and distribution of hybrid rice and Cd tolerance mechanisms existed between the two cultivars. Therefore,it is possible to develop the cultivars with high tolerance and low grain Cd/Zn accumulation in rice breeding forh high yield.It is also suggested the practical measures should be taken such as rational soil-cultivar combination and optimization dietary structure for controling the human Cd food exposure and guarantee Zn nutrition intake by diet.
引文
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