水稻铬胁迫耐性的遗传分析与还原型谷胱甘肽缓解铬毒害的机理研究
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摘要
铬(Cr)是一种过渡金属元素,在土壤、水体和大气中均有分布,具有多种氧化态,以三价和六价最为稳定。随着环境污染的加剧,农田十壤的铬含量逐渐提高,对农业生产与农产品安全造成了严重威胁。本研究以亲本在铬积累存在明显差异的杂交组合(ZYQ8/JX17)构建的一个水稻DH群体为材料,确定与铬耐性相关的数量性状位点和鉴定耐铬性差异显著的株系：进而以耐铬株系117和铬敏感株系41为材料,通过外源添加还原型谷胱甘肽(reduced Glutathione, GSH)的方式,研究其对铬形态、吸收、转运、累积和分布的影响,并从抗氧化系统、细胞活性和超微结构、巯基化合物、水分与养分吸收以及有机酸分泌等方面了解水稻铬毒害的缓解作用和相关机理。主要研究结果如下：
1铬胁迫下水稻DH群体各株系的生长表现和铬耐性相关性状的QTL定位
以一个水稻双单倍体(Double Haploid, DH)群体为材料,利用数量性状位点(Quantitative Trait Loci, QTL)定位分析,研究了铬胁迫下不同株系的生长和铬含量差异。结果表明,铬处理下,亲本ZYQ8和JX17的株高、根长、叶绿素含量和地上部干重均降低,而根干重稍有增加；群体的平均株高、根长和地上部干重均表现下降,而平均根干重和叶绿素含量影响较小。水稻铬含量根显著高于地上部,且群体各株系间差异显著。鉴定到多个与铬耐性相关的QTL位点,但它们只能解释不到10%的表型变异；鉴定到两个与铬含量相关的QTL位点,它们均位于第10号染色体的同一遗传区间,可解释近11%的表型变异。另外,锌含量与铬含量呈正相关,但相关的QTL位点只能解释很小的含量变异。
2水稻DH群体不同铬耐性株系的鉴定
利用权重分析法对铬胁迫下水稻DH群体各株系的生长表现进行了研究,从中鉴定到铬耐性差异显著的株系。在考察的生长性状中,铬胁迫对地上部干重的影响最大,对叶绿素含量的影响最小。株系117和101的加权综合分值最高,41和49的分值最低。根部铬含量株系117显著大于41,而根部铬累积量两株系恰好相反。铬胁迫显著抑制两亲本及群体各株系的锌含量,亲本的地上部锌含量下降较多,而根部含量受影响较小。由此可以认为,株系117和101的耐铬性较强,而株系41和49耐铬性较弱。
3外源谷胱甘肽对水稻铬毒害的缓解效应与最适浓度筛选
研究了外源添加GSH对缓解水稻铬毒害的效应。结果显示,在无铬胁迫条件下,外源GSH并不影响水稻生长和叶绿素含量。在铬胁迫培养液下,外源添加GSH具有缓解铬胁迫抑制水稻生长的作用,且在一定范围内随着GSH浓度的增加,缓解效果增强,在铬离子与GSH浓度比为1：3或1：5时效果较好,当两者的浓度比超过1：10时,水稻根系生长受到抑制。外源GSH提高水稻根的铬含量,但降低地上部的铬含量。结果表明,适宜浓度的GSH可缓解水稻的铬毒害,铬离子与GSH的浓度比为1:3(100μM Cr6+/300μM GSH)，具有最佳的缓解作用。
4GSH缓解铬胁迫对水稻抗氧化系统、细胞活性和超微结构的影响
从抗氧化系统、细胞活性和亚细胞水平上研究了水稻DH株系117和41对铬胁迫的响应和外源GSH的缓解作用。结果表明,铬胁迫下,水稻DH群体的两株系(117和41)的光合色素含量显著下降,而丙二醛含量显著上升；外源添加GSH后,以上两种性状均得到明显恢复。铬胁迫影响抗氧化酶活性,且影响程度因株系和组织而异,株系117的抗氧化能力强于株系41。外源添加GSH缓解铬毒害对抗氧化酶活性的影响。铬胁迫严重影响水稻根尖细胞活性,而外源添加GSH可使这种影响明显减小。铬胁迫导致水稻叶肉细胞叶绿体肿胀变形,基粒数目减少、片层结构松散；根系细胞结构变形,胞质变疏,液泡体积剧增和核仁消失；外源添加GSH后,以上水稻超微结构均得到不同程度的恢复。
5GSH对铬胁迫下水稻巯基化合物含量、水分和养分吸收以及根系有机酸分泌的影响
研究了铬胁迫下水稻DH群体的两个铬胁迫耐性不同的株系(117和41)的水分和养分吸收、植株含巯基化合物含量和根系有机酸分泌的影响,并分析了外源添加GSH对上述性状在铬胁迫下的缓解作用。结果显示,GSH显著改善铬胁迫下水稻对水分的吸收,并明显减轻铁(Fe)和锰(Mn)的吸收抑制。铬胁迫下根部锌含量增加,说明铬损伤了根细胞,使其失去完整性和选择性。铬胁迫影响水稻根际有机酸分泌,且株系间差异显著；外源添加GSH增加根系分泌乙酸和苹果酸,且耐性株系117的增加幅度明显大于敏感株系41,启示出GSH对铬毒害的缓解作用与这两种有机酸分泌有关。本研究未检测到植物螯合肽(Phytochelatins, PCs),但发现铬胁迫影响水稻体内GSH和半胱氨酸(Cys)水平,并在叶片中发现了一种与PC3出峰时间相近的物质,值得进一步对此进行化学鉴定。
6GSH对水稻根际铬形态及其吸收、转运、累积和分布的影响
研究了外源GSH对铬离子形态变化以及在水稻组织、细胞、亚细胞水平分布的影响。结果表明,添加GSH显著降低培养液中的Cr6+含量,启示出GSH可能具有改变铬离子形态的作用。水稻根系对铬离子的吸收存在时间效应,外源添加GSH后,水稻根的铬含量增加,而地上部的铬含量减少,从而间接证明铬的吸收和转运并非以Cr6+为主。水稻根系表皮、皮层以及维管束组织中均有铬的分布,而外源添加GSH后,使铬离子主要分布在根表,证明表皮细胞能固定和区隔一部分铬离子,从而缓解其对植物的毒性。铬主要累积在细胞壁上,其次在可溶性成分(以液泡为主)中,说明细胞壁和液泡是水稻体内Cr的主要分布器官。外源添加GSH后,地上部各亚细胞组分以及根部可溶性成分中的铬离子含量下降,细胞壁中的铬含量变化较小,说明GSH抑制植物对铬的转运,但对吸收无影响。
As a kind of transition metal element, Chromium (Cr) is widely located in soil, water and air. It has several oxidation states with+3and+6being the most stable forms. Recently, Cr concentration in agricultural land increase dramatically for the aggravation of environment pollution. Therefore, current experiments were carried out to find the quantitative trait loci (QTLs) related to the Cr tolerance by using a rice DH population developed a cross of ZYQ8/JX17with difference in Cr-tolerance, and to identify the lines with significantly different Cr-tolerance. Further more, alleviating effect and mechanism of exogenous GSH to the Cr toxicity on antioxidative system, cell viability and ultra-structure, sulfhydryl compound concentration, water-uptake, nutrition balance and organic acid excretion were also studied by using the Cr-tolerant DH line117and Cr-sensitive line41. The major results are summarized as follows:
1QTL mapping of growth traits and Cr-tolerance of a rice DH population under Cr stress
The results showed the differences caused by Cr stress on the growth and the Cr concentration among the lines of a rice DH population as well as the two parents. For the parents ZYQ8and JX17, plant height (PH), root length (RL), chlorophyll content (SPAD value) and shoot dry weight (SDW) were all inhibited under Cr stress, while root dry weight (RDW) was slightly increased. For the DH population, PH, RL and SDW were also decreased, while RDW and SPAD values were weakly affected. Cr concentration in roots was much higher than in shoot and varied significantly among DH lines. QTL analysis identified several loci related to the growth traits under Cr stress, while each one explained less than10%of the phenotypic variation. Two QTLs linked to the Cr concentration and translocation was also found, each one explained more than10%of the variation. Additionally, positive correlation existed between Zn and Cr concentrations, so QTL mapping was also conducted.
2Identification of different Cr-tolerance lines in a rice DH population
Current experiment studied the growth variation of all DH lines by using the weighted analysis under Cr tress, then identified the lines with high and low Cr-tolerance. Among the observed growth traits, SDW was most significantly affected by Cr stress, while SPAD value was less affected. The integrated scores of the lines117and101were the highest, while the lines41and49were the lowest. Cr concentration in roots of the line117was significantly higher than that of the line41, but Cr accumulation in root (RCC) was just opposite for the two lines. Cr stress affected the zinc concentration among DH lines and parents, with shoot Zn concentration (SZC) being reduced much more than root Zn concentration (RZC) for the two parents. Hence, it may be suggested that the lines117and101were high Cr tolerant, while the lines41and49were Cr sensitive.
3Alleviating effect of exogenous GSH on Cr toxicity to rice plants and the identification of the optimal GSH concentration
The two experiments were conducted to investigate the alleviating effect of exogenous GSH on Cr toxicity. The results showed that addition of GSH into the culture solution, without Cr addition, did not affect growth and chlorophyll content of rice seedlings, while GSH addition could greatly alleviate the growth inhibition caused by Cr exposure. The greatest alleviating effect occurred when the concentration ratio of Cr6+and GSH was1:3or1:5, while root elongation was significantly inhibited when the ratio was up to1:10. GSH addition increased Cr concentration in roots and reduced it in shoots, while Cr concentration in both roots and shoots was reduced when the ratio was1:10. It may be suggested that suitable GSH addition in culture solution may alleviate Cr toxicity to rice.
4The influence of Cr stress on anti-oxidative system, cell viability and ultra structure, and its alleviation by GSH
The response of rice DH lines117and41to Cr stress and its alleviation by exogenous GSH were examined. The results showed that Cr stress reduced the concentration of photosynthetic pigment, and increased MDA content, affected activities of anti-oxidative enzymes. The difference could be found in the extent of the influence between lines, with117having higher activities of oxidative enzymes under Cr stress. Exogenous GSH alleviated Cr toxicity, being characterized by higher anti-oxidative capacity In addition, GSH improved cell viability in rice roots relative to the Cr treatment alone. Cr stress caused chloroplast deformation and chloroplast lamellae loose in mesophyll cells, and cell structure disintegration, matrix thinning, vacuole larger and nucleolus disappearing in root tip cells, while GSH addition alleviated these changes of cell ultra-structure.
5Alleviation of exogenous GSH on Cr toxicity to thiol-compound synthesis, water and nutrient uptake, and organic acid secretion in rice seedlings
The present study showed Cr toxicity to thiol-compound synthesis (concentration), water and nutrient uptake, and organic acid secretion in the two rice DH lines117and41, while addition of exogenous GSH in the culture solution could alleviate these toxicity caused by Cr stress. GSH ameliorated Cr inhibition on the uptake of water, Fe and Mn. Cr affected the pattern of organic acid secretion in rice roots, however the difference existed between lines117and41. Addition of GSH dramatically increased level of acetic acid and malic acid, with line117having more increase than line41, and there was a close association between the two organic acid contents and alleviation of GSH to Cr toxicity. Although no PCs was found in this study, it was clearly showed that Cr stress affected GSH and Cys levels in rice seedlings. Moreover, the compound with the similar retention time as PC3was identified and waits for further identification.
6Effect of GSH on Cr form in rice rhizosphere and its absorption, translocation and accumulation
This experiment determined the effect of GSH on Cr form in rice rhizosphere and its distribution in rice tissular, cellular and subcellular levels. Addition of GSH caused a dramatic decrease of Cr6+in the cultural solution. Meanwhile, GSH addition increased Cr concentration in roots, but reduced it in shoots, suggesting that most Cr was not absorbed in the form of Cr6+. In the treatment of100pM Cr6+, Cr was mainly located in epidermis, cortices and vascular bundles of rice roots, but in the treatment of100μM Cr6+and300μM GSH, Cr was mainly found in epidermis, indicating that epidermic cells can alleviate Cr toxicity to the plants. The results at sub-cellular level showed that Cr was mainly presented on cell walls, followed by soluble composition. Exogenous GSH reduced Cr concentration in all subcellular components of shoots, and in the soluble composition of roots, while no effect was found on cell walls. It may be suggested that GSH inhibited Cr translocation, but did not affect Cr uptake.
1铬胁迫下水稻DH群体各株系的生长表现和铬耐性相关性状的QTL定位
以一个水稻双单倍体(Double Haploid, DH)群体为材料,利用数量性状位点(Quantitative Trait Loci, QTL)定位分析,研究了铬胁迫下不同株系的生长和铬含量差异。结果表明,铬处理下,亲本ZYQ8和JX17的株高、根长、叶绿素含量和地上部干重均降低,而根干重稍有增加；群体的平均株高、根长和地上部干重均表现下降,而平均根干重和叶绿素含量影响较小。水稻铬含量根显著高于地上部,且群体各株系间差异显著。鉴定到多个与铬耐性相关的QTL位点,但它们只能解释不到10%的表型变异；鉴定到两个与铬含量相关的QTL位点,它们均位于第10号染色体的同一遗传区间,可解释近11%的表型变异。另外,锌含量与铬含量呈正相关,但相关的QTL位点只能解释很小的含量变异。
2水稻DH群体不同铬耐性株系的鉴定
利用权重分析法对铬胁迫下水稻DH群体各株系的生长表现进行了研究,从中鉴定到铬耐性差异显著的株系。在考察的生长性状中,铬胁迫对地上部干重的影响最大,对叶绿素含量的影响最小。株系117和101的加权综合分值最高,41和49的分值最低。根部铬含量株系117显著大于41,而根部铬累积量两株系恰好相反。铬胁迫显著抑制两亲本及群体各株系的锌含量,亲本的地上部锌含量下降较多,而根部含量受影响较小。由此可以认为,株系117和101的耐铬性较强,而株系41和49耐铬性较弱。
3外源谷胱甘肽对水稻铬毒害的缓解效应与最适浓度筛选
研究了外源添加GSH对缓解水稻铬毒害的效应。结果显示,在无铬胁迫条件下,外源GSH并不影响水稻生长和叶绿素含量。在铬胁迫培养液下,外源添加GSH具有缓解铬胁迫抑制水稻生长的作用,且在一定范围内随着GSH浓度的增加,缓解效果增强,在铬离子与GSH浓度比为1：3或1：5时效果较好,当两者的浓度比超过1：10时,水稻根系生长受到抑制。外源GSH提高水稻根的铬含量,但降低地上部的铬含量。结果表明,适宜浓度的GSH可缓解水稻的铬毒害,铬离子与GSH的浓度比为1:3(100μM Cr6+/300μM GSH)，具有最佳的缓解作用。
4GSH缓解铬胁迫对水稻抗氧化系统、细胞活性和超微结构的影响
从抗氧化系统、细胞活性和亚细胞水平上研究了水稻DH株系117和41对铬胁迫的响应和外源GSH的缓解作用。结果表明,铬胁迫下,水稻DH群体的两株系(117和41)的光合色素含量显著下降,而丙二醛含量显著上升；外源添加GSH后,以上两种性状均得到明显恢复。铬胁迫影响抗氧化酶活性,且影响程度因株系和组织而异,株系117的抗氧化能力强于株系41。外源添加GSH缓解铬毒害对抗氧化酶活性的影响。铬胁迫严重影响水稻根尖细胞活性,而外源添加GSH可使这种影响明显减小。铬胁迫导致水稻叶肉细胞叶绿体肿胀变形,基粒数目减少、片层结构松散；根系细胞结构变形,胞质变疏,液泡体积剧增和核仁消失；外源添加GSH后,以上水稻超微结构均得到不同程度的恢复。
5GSH对铬胁迫下水稻巯基化合物含量、水分和养分吸收以及根系有机酸分泌的影响
研究了铬胁迫下水稻DH群体的两个铬胁迫耐性不同的株系(117和41)的水分和养分吸收、植株含巯基化合物含量和根系有机酸分泌的影响,并分析了外源添加GSH对上述性状在铬胁迫下的缓解作用。结果显示,GSH显著改善铬胁迫下水稻对水分的吸收,并明显减轻铁(Fe)和锰(Mn)的吸收抑制。铬胁迫下根部锌含量增加,说明铬损伤了根细胞,使其失去完整性和选择性。铬胁迫影响水稻根际有机酸分泌,且株系间差异显著；外源添加GSH增加根系分泌乙酸和苹果酸,且耐性株系117的增加幅度明显大于敏感株系41,启示出GSH对铬毒害的缓解作用与这两种有机酸分泌有关。本研究未检测到植物螯合肽(Phytochelatins, PCs),但发现铬胁迫影响水稻体内GSH和半胱氨酸(Cys)水平,并在叶片中发现了一种与PC3出峰时间相近的物质,值得进一步对此进行化学鉴定。
6GSH对水稻根际铬形态及其吸收、转运、累积和分布的影响
研究了外源GSH对铬离子形态变化以及在水稻组织、细胞、亚细胞水平分布的影响。结果表明,添加GSH显著降低培养液中的Cr6+含量,启示出GSH可能具有改变铬离子形态的作用。水稻根系对铬离子的吸收存在时间效应,外源添加GSH后,水稻根的铬含量增加,而地上部的铬含量减少,从而间接证明铬的吸收和转运并非以Cr6+为主。水稻根系表皮、皮层以及维管束组织中均有铬的分布,而外源添加GSH后,使铬离子主要分布在根表,证明表皮细胞能固定和区隔一部分铬离子,从而缓解其对植物的毒性。铬主要累积在细胞壁上,其次在可溶性成分(以液泡为主)中,说明细胞壁和液泡是水稻体内Cr的主要分布器官。外源添加GSH后,地上部各亚细胞组分以及根部可溶性成分中的铬离子含量下降,细胞壁中的铬含量变化较小,说明GSH抑制植物对铬的转运,但对吸收无影响。
As a kind of transition metal element, Chromium (Cr) is widely located in soil, water and air. It has several oxidation states with+3and+6being the most stable forms. Recently, Cr concentration in agricultural land increase dramatically for the aggravation of environment pollution. Therefore, current experiments were carried out to find the quantitative trait loci (QTLs) related to the Cr tolerance by using a rice DH population developed a cross of ZYQ8/JX17with difference in Cr-tolerance, and to identify the lines with significantly different Cr-tolerance. Further more, alleviating effect and mechanism of exogenous GSH to the Cr toxicity on antioxidative system, cell viability and ultra-structure, sulfhydryl compound concentration, water-uptake, nutrition balance and organic acid excretion were also studied by using the Cr-tolerant DH line117and Cr-sensitive line41. The major results are summarized as follows:
1QTL mapping of growth traits and Cr-tolerance of a rice DH population under Cr stress
The results showed the differences caused by Cr stress on the growth and the Cr concentration among the lines of a rice DH population as well as the two parents. For the parents ZYQ8and JX17, plant height (PH), root length (RL), chlorophyll content (SPAD value) and shoot dry weight (SDW) were all inhibited under Cr stress, while root dry weight (RDW) was slightly increased. For the DH population, PH, RL and SDW were also decreased, while RDW and SPAD values were weakly affected. Cr concentration in roots was much higher than in shoot and varied significantly among DH lines. QTL analysis identified several loci related to the growth traits under Cr stress, while each one explained less than10%of the phenotypic variation. Two QTLs linked to the Cr concentration and translocation was also found, each one explained more than10%of the variation. Additionally, positive correlation existed between Zn and Cr concentrations, so QTL mapping was also conducted.
2Identification of different Cr-tolerance lines in a rice DH population
Current experiment studied the growth variation of all DH lines by using the weighted analysis under Cr tress, then identified the lines with high and low Cr-tolerance. Among the observed growth traits, SDW was most significantly affected by Cr stress, while SPAD value was less affected. The integrated scores of the lines117and101were the highest, while the lines41and49were the lowest. Cr concentration in roots of the line117was significantly higher than that of the line41, but Cr accumulation in root (RCC) was just opposite for the two lines. Cr stress affected the zinc concentration among DH lines and parents, with shoot Zn concentration (SZC) being reduced much more than root Zn concentration (RZC) for the two parents. Hence, it may be suggested that the lines117and101were high Cr tolerant, while the lines41and49were Cr sensitive.
3Alleviating effect of exogenous GSH on Cr toxicity to rice plants and the identification of the optimal GSH concentration
The two experiments were conducted to investigate the alleviating effect of exogenous GSH on Cr toxicity. The results showed that addition of GSH into the culture solution, without Cr addition, did not affect growth and chlorophyll content of rice seedlings, while GSH addition could greatly alleviate the growth inhibition caused by Cr exposure. The greatest alleviating effect occurred when the concentration ratio of Cr6+and GSH was1:3or1:5, while root elongation was significantly inhibited when the ratio was up to1:10. GSH addition increased Cr concentration in roots and reduced it in shoots, while Cr concentration in both roots and shoots was reduced when the ratio was1:10. It may be suggested that suitable GSH addition in culture solution may alleviate Cr toxicity to rice.
4The influence of Cr stress on anti-oxidative system, cell viability and ultra structure, and its alleviation by GSH
The response of rice DH lines117and41to Cr stress and its alleviation by exogenous GSH were examined. The results showed that Cr stress reduced the concentration of photosynthetic pigment, and increased MDA content, affected activities of anti-oxidative enzymes. The difference could be found in the extent of the influence between lines, with117having higher activities of oxidative enzymes under Cr stress. Exogenous GSH alleviated Cr toxicity, being characterized by higher anti-oxidative capacity In addition, GSH improved cell viability in rice roots relative to the Cr treatment alone. Cr stress caused chloroplast deformation and chloroplast lamellae loose in mesophyll cells, and cell structure disintegration, matrix thinning, vacuole larger and nucleolus disappearing in root tip cells, while GSH addition alleviated these changes of cell ultra-structure.
5Alleviation of exogenous GSH on Cr toxicity to thiol-compound synthesis, water and nutrient uptake, and organic acid secretion in rice seedlings
The present study showed Cr toxicity to thiol-compound synthesis (concentration), water and nutrient uptake, and organic acid secretion in the two rice DH lines117and41, while addition of exogenous GSH in the culture solution could alleviate these toxicity caused by Cr stress. GSH ameliorated Cr inhibition on the uptake of water, Fe and Mn. Cr affected the pattern of organic acid secretion in rice roots, however the difference existed between lines117and41. Addition of GSH dramatically increased level of acetic acid and malic acid, with line117having more increase than line41, and there was a close association between the two organic acid contents and alleviation of GSH to Cr toxicity. Although no PCs was found in this study, it was clearly showed that Cr stress affected GSH and Cys levels in rice seedlings. Moreover, the compound with the similar retention time as PC3was identified and waits for further identification.
6Effect of GSH on Cr form in rice rhizosphere and its absorption, translocation and accumulation
This experiment determined the effect of GSH on Cr form in rice rhizosphere and its distribution in rice tissular, cellular and subcellular levels. Addition of GSH caused a dramatic decrease of Cr6+in the cultural solution. Meanwhile, GSH addition increased Cr concentration in roots, but reduced it in shoots, suggesting that most Cr was not absorbed in the form of Cr6+. In the treatment of100pM Cr6+, Cr was mainly located in epidermis, cortices and vascular bundles of rice roots, but in the treatment of100μM Cr6+and300μM GSH, Cr was mainly found in epidermis, indicating that epidermic cells can alleviate Cr toxicity to the plants. The results at sub-cellular level showed that Cr was mainly presented on cell walls, followed by soluble composition. Exogenous GSH reduced Cr concentration in all subcellular components of shoots, and in the soluble composition of roots, while no effect was found on cell walls. It may be suggested that GSH inhibited Cr translocation, but did not affect Cr uptake.
引文
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