水稻砷、镉、铅低含量基因型筛选及体内转运特征
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摘要
筛选低重金属含量水稻(Oryza sativa L.)品种或减少水稻对种植土壤中重金属的吸收是控制稻米重金属含量最直接有效的办法。而水稻籽粒中重金属的含量与植株对重金属元素的吸收、运输和积累有关,因此研究重金属在水稻体内的吸收与转运特征,尤其是向经济器官的迁移规律具有重要科学意义。本论文采用电感耦合等离子体质谱(ICP-MS)法,测定了78份水稻品种的糙米及粳稻品种浙优12号不同生长期地上部不同器官的重金属元素砷(As)、镉(Cd)、铅(Pb)含量。筛选水稻As、Cd、Pb低含量基因型,探索As、Cd、Pb在水稻体内的转运特征。结果摘要如下：
1.本研究测定了78份水稻品种的糙米中As、Cd、Pb的含量。同时比较其中的20份品种在3个试验点的糙米As、Cd、Pb含量。结果表明：水稻籽粒对As、Cd、Pb的积累存在着明显的品种差异,其中Cd和Pb积累的基因型明显较As积累的基因型丰富；相关性分析表明,糙米中Cd和Pb含量存在极显著相关性(P<0.01)；筛选得到Cd、Pb含量低的基因型5个,Cd、As含量低的基因型1个,As、Pb含量低的基因型2个,Cd、Pb、As含量均低的基因型1个；土壤As、Cd、Pb含量对糙米As、Cd、Pb含量有较明显的影响,且三者对土壤环境的反应也各不相同。
2.以粳稻品种浙优12号为研究材料,通过水培与大田种植,分析水稻主要生长期(苗期、分蘖期、抽穗期、乳熟期、蜡熟期、完熟期)植株地上部不同器官中As、Cd、Pb含量变化。结果表明：水稻植株叶片和叶鞘中As、Cd、Pb含量大致呈现“U”字型变化过程,苗期最高、抽穗期最低；进入生殖生长期,三个元素在水稻的生长后期表现出不同的转运特点,但在最终的贮藏器官(籽实)中又表现相同的趋势；完熟期,水稻植株地上部As含量由高到低的顺序为：茎秆>叶片>叶鞘>穗轴>颖壳>糙米,Cd含量顺序为茎秆>叶片>叶鞘>糙米>穗轴>颖壳,Pb含量顺序为叶片>叶鞘>茎秆>穗轴>颖壳>糙米,显示出水稻重金属含量自下而上递减的趋势；在大田栽培和水培条件下,Pb含量规律为叶片>叶鞘>茎秆>穗轴>颖壳>糙米,地上部分叶片是Pb的主要集中器官,叶中含量最高,而糙米中富集最低,说明水稻吸收Pb后地上部分主要富集在茎叶中,向籽粒转移较少；As、Cd、Pb三种元素间的含量在水稻植株地上部各器官的关系较一致表现为As>Pb>Cd。
3.对21份水稻品种在3个试验点的糙米As含量进行分析,结果表明：在3个土壤As含量由低至高的试验点中,21份水稻品种糙米As含量平均值也呈现出由低到高的趋势；分析水稻As含量对不同种植环境的反应,多数品种对不同试验点环境表现的敏感性不一致,但有4个品种表现稳定,如明恢70都表现为迟钝型,明恢86表现为敏感型,明恢72和明恢81表现为中间型。分析粳稻品种浙优12号在15个栽培点中土壤As含量、穗茎节As含量、糙米As含量的相关性,结果表明：As含量高低顺序依次为土壤>糙米>穗茎节；土壤As含量与糙米As含量的相关性不显著,土壤As含量与穗茎节As含量存在不显著负相关,穗茎节As含量与糙米As含量之间呈极显著正相关。
Selection and breeding low heavy metal accumulation varieties or decreasing their absorption from growth soil were the most immediate and effective ways to control heavy metal concentration in rice(Oryza sativa L.). As the concentration of heavy metal in the kernels were related to their absorption, transportation and accumulation in rice plant, it was important to research the characteristics of absorption and transportation in rice, especially the migrating regularity of heavy metal to economy organ. In present paper, inductively coupled plasma-mass spectrometry(ICP-MS) was used to detect arsenic(As), cadmium(Cd) and lead(Pb) concentration of brown rice in 78 varieties, and of aboveground organs in different growth stages of variety Zheyou 12. The major amid was to screening rice genotypes with low As, Cd, Pb concentration and to explore regularity of their transportation in rice. Results are as follows:
1. The As, Cd, Pb concentration of brown rice of 78 varieties was determined. Simultaneously, As, Cd, Pb concentration of brown rice of 20 varieties, wich were grown in three field trials with different As, Cd, Pb concentration was compared. The results were showed that there is remarkable difference in accumulation of As, Cd, Pb in brown rice among varieties, and there are more polymorphisms in genotypes of Cd and Pb concentration than that of As. Correlation analyses showed that Cd concentration in brown rice have most positively correlation with Pb concentration(P<0.01). Out of 78 varieties, we obtained five genotypes of low Cd concentration and low Pb concentration in brown rice, one genotype of low Cd+As, two genotypes of low As+Pb, only one genotype of low As+Cd+Pb. t Test showed that environments have significant influence on As, Cd, Pb concentration in brown rice, and certain were at significant level. As,Cd and Pb expressed different responses to growth environments.
2. The concentrations of As, Cd, Pb in aboveground organs at different growth stages, including seedling stage, tillering stage, heading stage, milk stage, dough stage and full ripe stage, were investigated using variety Zheyou 12 by experiments of water culture and field. Results showed that changes of As, Cd, Pb concentration in leaf and leaf sheath from seedling stage to full ripe stage present a U-shaped curve, the highest point at seedling stage and the lowest at heading stage. At the reproductive growth stage, the three elements showed different characteristics of transportation in different reproductive organs, but showed similar trend in kernels. Ultimately, the rank from high to low As concentration in aboveground organs was stalk>leaf>leaf sheath>spike stalk> glume>brown rice, and rank of Cd concentrations was stalk>leaf>leaf sheath>brown rice >spike stalk>glume, rank of Pb concentrations was leaf>leaf sheath>stalk>spike stalk> glume>brown rice, these results supported the regularity of decrease successively from root to top for heavy metal distribution in plant. Comparison with culture of water and field, the rank of Pb concentrations in different organs was the same as two experiments. Pb was higher concentration in leaf, and the lowest in brown rice, suggested that stalk and leaf of aboveground organs were the major parts for Pb accumulation in rice, and less transfered into the grain. Concentrations of As, Cd, Pb in the same aboveground organs in different growth stages in rice plants was As>Pb>Cd.
3. Total As concentration of brown rice was investigated for 21 varieties grown in three field trials. Results showed that averages of total As concentration of brown rice express from high to low according to the total As concentration of growth soil. Analyzing the reaction of varieties to growth environments, it showed that most varieties exhibited different reaction to growth environment, but there were four stable varieties, for example, Minghui 70 consistently exhibited insensitive reaction, Minghui 86 exhibited sensitive reaction, Minghui 72 and Minghui 81 fell into intermediate category. Total brown rice, culm and growth soil environment of variety Zheyou 12 grown from fifteen tested field trials were measured, the average As concentration were soil> brown rice> culm in turn. Correlation analyses showed that As concentration of growth soil is no significant correlation to that of brown rice, and no negatively correlation to culm As concentration, but As concentration of brown rice has most positively correlation to that of culm.
1.本研究测定了78份水稻品种的糙米中As、Cd、Pb的含量。同时比较其中的20份品种在3个试验点的糙米As、Cd、Pb含量。结果表明：水稻籽粒对As、Cd、Pb的积累存在着明显的品种差异,其中Cd和Pb积累的基因型明显较As积累的基因型丰富；相关性分析表明,糙米中Cd和Pb含量存在极显著相关性(P<0.01)；筛选得到Cd、Pb含量低的基因型5个,Cd、As含量低的基因型1个,As、Pb含量低的基因型2个,Cd、Pb、As含量均低的基因型1个；土壤As、Cd、Pb含量对糙米As、Cd、Pb含量有较明显的影响,且三者对土壤环境的反应也各不相同。
2.以粳稻品种浙优12号为研究材料,通过水培与大田种植,分析水稻主要生长期(苗期、分蘖期、抽穗期、乳熟期、蜡熟期、完熟期)植株地上部不同器官中As、Cd、Pb含量变化。结果表明：水稻植株叶片和叶鞘中As、Cd、Pb含量大致呈现“U”字型变化过程,苗期最高、抽穗期最低；进入生殖生长期,三个元素在水稻的生长后期表现出不同的转运特点,但在最终的贮藏器官(籽实)中又表现相同的趋势；完熟期,水稻植株地上部As含量由高到低的顺序为：茎秆>叶片>叶鞘>穗轴>颖壳>糙米,Cd含量顺序为茎秆>叶片>叶鞘>糙米>穗轴>颖壳,Pb含量顺序为叶片>叶鞘>茎秆>穗轴>颖壳>糙米,显示出水稻重金属含量自下而上递减的趋势；在大田栽培和水培条件下,Pb含量规律为叶片>叶鞘>茎秆>穗轴>颖壳>糙米,地上部分叶片是Pb的主要集中器官,叶中含量最高,而糙米中富集最低,说明水稻吸收Pb后地上部分主要富集在茎叶中,向籽粒转移较少；As、Cd、Pb三种元素间的含量在水稻植株地上部各器官的关系较一致表现为As>Pb>Cd。
3.对21份水稻品种在3个试验点的糙米As含量进行分析,结果表明：在3个土壤As含量由低至高的试验点中,21份水稻品种糙米As含量平均值也呈现出由低到高的趋势；分析水稻As含量对不同种植环境的反应,多数品种对不同试验点环境表现的敏感性不一致,但有4个品种表现稳定,如明恢70都表现为迟钝型,明恢86表现为敏感型,明恢72和明恢81表现为中间型。分析粳稻品种浙优12号在15个栽培点中土壤As含量、穗茎节As含量、糙米As含量的相关性,结果表明：As含量高低顺序依次为土壤>糙米>穗茎节；土壤As含量与糙米As含量的相关性不显著,土壤As含量与穗茎节As含量存在不显著负相关,穗茎节As含量与糙米As含量之间呈极显著正相关。
Selection and breeding low heavy metal accumulation varieties or decreasing their absorption from growth soil were the most immediate and effective ways to control heavy metal concentration in rice(Oryza sativa L.). As the concentration of heavy metal in the kernels were related to their absorption, transportation and accumulation in rice plant, it was important to research the characteristics of absorption and transportation in rice, especially the migrating regularity of heavy metal to economy organ. In present paper, inductively coupled plasma-mass spectrometry(ICP-MS) was used to detect arsenic(As), cadmium(Cd) and lead(Pb) concentration of brown rice in 78 varieties, and of aboveground organs in different growth stages of variety Zheyou 12. The major amid was to screening rice genotypes with low As, Cd, Pb concentration and to explore regularity of their transportation in rice. Results are as follows:
1. The As, Cd, Pb concentration of brown rice of 78 varieties was determined. Simultaneously, As, Cd, Pb concentration of brown rice of 20 varieties, wich were grown in three field trials with different As, Cd, Pb concentration was compared. The results were showed that there is remarkable difference in accumulation of As, Cd, Pb in brown rice among varieties, and there are more polymorphisms in genotypes of Cd and Pb concentration than that of As. Correlation analyses showed that Cd concentration in brown rice have most positively correlation with Pb concentration(P<0.01). Out of 78 varieties, we obtained five genotypes of low Cd concentration and low Pb concentration in brown rice, one genotype of low Cd+As, two genotypes of low As+Pb, only one genotype of low As+Cd+Pb. t Test showed that environments have significant influence on As, Cd, Pb concentration in brown rice, and certain were at significant level. As,Cd and Pb expressed different responses to growth environments.
2. The concentrations of As, Cd, Pb in aboveground organs at different growth stages, including seedling stage, tillering stage, heading stage, milk stage, dough stage and full ripe stage, were investigated using variety Zheyou 12 by experiments of water culture and field. Results showed that changes of As, Cd, Pb concentration in leaf and leaf sheath from seedling stage to full ripe stage present a U-shaped curve, the highest point at seedling stage and the lowest at heading stage. At the reproductive growth stage, the three elements showed different characteristics of transportation in different reproductive organs, but showed similar trend in kernels. Ultimately, the rank from high to low As concentration in aboveground organs was stalk>leaf>leaf sheath>spike stalk> glume>brown rice, and rank of Cd concentrations was stalk>leaf>leaf sheath>brown rice >spike stalk>glume, rank of Pb concentrations was leaf>leaf sheath>stalk>spike stalk> glume>brown rice, these results supported the regularity of decrease successively from root to top for heavy metal distribution in plant. Comparison with culture of water and field, the rank of Pb concentrations in different organs was the same as two experiments. Pb was higher concentration in leaf, and the lowest in brown rice, suggested that stalk and leaf of aboveground organs were the major parts for Pb accumulation in rice, and less transfered into the grain. Concentrations of As, Cd, Pb in the same aboveground organs in different growth stages in rice plants was As>Pb>Cd.
3. Total As concentration of brown rice was investigated for 21 varieties grown in three field trials. Results showed that averages of total As concentration of brown rice express from high to low according to the total As concentration of growth soil. Analyzing the reaction of varieties to growth environments, it showed that most varieties exhibited different reaction to growth environment, but there were four stable varieties, for example, Minghui 70 consistently exhibited insensitive reaction, Minghui 86 exhibited sensitive reaction, Minghui 72 and Minghui 81 fell into intermediate category. Total brown rice, culm and growth soil environment of variety Zheyou 12 grown from fifteen tested field trials were measured, the average As concentration were soil> brown rice> culm in turn. Correlation analyses showed that As concentration of growth soil is no significant correlation to that of brown rice, and no negatively correlation to culm As concentration, but As concentration of brown rice has most positively correlation to that of culm.
引文
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