两种基因型水稻根际微域中重金属镉形态差异及其有效性研究
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摘要
水稻是我国最重要的粮食作物之一,但是由于近些年来农田重金属镉污染的日益加剧,所以稻米的食品安全问题越来越为人们所关注。根际环境作为有别于一般土体的特殊土壤环境,直接影响了重金属镉在土壤中的形态及其有效性,对研究重金属镉水稻污染具有重要意义。本论文围绕土壤-镉-水稻-微生物之间的相互关系,对镉胁迫下两种不同基因型水稻的生长及生理生化特性、水稻体内镉的迁移分配规律、镉在水稻毫米级根际微域中的形态变化、不同供铁条件下两个水稻品种的根系分泌物差异以及对镉吸收的影响等开展了研究,获得以下主要研究结果：
1.以我国东南地区水稻土黄斑田为供试土壤,籼稻四喜占(Sixizhan)和粳稻浙农54(Zhenong54)为供试水稻品种,采用多隔层三室根箱法开展盆栽试验(通过尼龙撩网插片的使用,把水稻根际土壤划分成根室,lmm、2mm、3mm、5mm近根际以及>5mm远根际等毫米级微域),通过改良的Tessier重金属分级提取法研究测定重金属镉在根际微域的形态变化(交换态、铁锰氧化态、有机质结合态、残留态)。实验结果表明,重金属镉各个形态含量,尤其是可交换态镉含量在根际微域中存在距离根系不同远近的差异,含量基本呈现根室、近根际<远根际的趋势；两个水稻品种在近根际不同毫米级微域以及根室中也存在着基因型的差异。就可交换态镉而言,浙农54的最低含量出现在近根际1或2mm处,而四喜占出现在根室,这与植物营养学中提到的营养因子根际梯度递减效应有所不同,说明重金属污染物的根际效应具有其特异性。相对应的是,两个水稻品种根际微域的土壤生物量碳含量趋势正好与土壤可交换态镉相反。由此可见,根室、近根际微域中有适宜的根系分泌物水平、根际微生物活性以及植物对交换态镉的吸收能力,这可能是诱导重金属镉形态变化的根本原因。
2.从阐明重金属镉在不同基因型水稻植株体内的生态行为角度出发,深入研究了重金属镉对两种不同基因型水稻生长抑制和抗氧化酶活性变化的差异,以及重金属镉在不同水稻品种体内迁移积累规律的差异。研究发现分蘖期镉胁迫下四喜占较浙农54受毒害严重,株高、干重以及POD和CAT酶的活性较无镉对照都有一定程度的下降,镉由根转移至茎、叶所占的比例较大；而至收获期,四喜占地上部分对镉的积累则显著减少。浙农54相较于四喜占有相反趋势,分蘖期时浙农54受毒害较轻,表现为株高、干重以及酶活性无明显抑制作用或高浓度时有抑制作用,且镉由根转移至地上部分的比例不大,而至收获期时,浙农54茎对锅的积累显著增加,最终导致浙农54籽粒含镉量较四喜占大。因此,不同品种或同一品种不同发育时期对镉胁迫的敏感性和耐受性不同,若仅研究镉胁迫对水稻某一时期的影响,结果有其局限性。
3.研究了缺铁和供铁条件下两种水稻品种根际分泌物的差异及其对植株吸收重金属镉的影响。结果表明,不同基因型水稻根际分泌物对供铁水平的响应差异与植株对镉的吸收有一定关系。两个水稻品种在缺铁和供铁条件下表现出的大致趋势相同：在缺铁处理中,水稻根系有机酸分泌的量较供铁处理都有显著的增加,植株对重金属镉的积累量也较供铁处理有显著的增加,但地上部分和根部干重较供铁处理有所下降。尽管趋势相同,但是两个水稻品种之间还是有一定的差异,其中四喜占缺铁处理时根系分泌物总量较浙农54缺铁处理根系分泌物总量大；四喜占根部和地上部对重金属镉的积累量均显著大于浙农54,加铁处理无显著差异；加铁处理的浙农54的地上部分和根部干重大于四喜占。
4.实验对镉胁迫下不同水稻基因型对镉、锌、钙、硫的吸收积累差异及三者与镉吸收积累之间的关系进行了研究。结果表明,锌、钙、硫对不同基因型水稻的生长及镉吸收有一定影响,并与不同生长阶段相关。不同处理中土壤镉的添加浓度极显著影响了两个品种水稻对镉的吸收和镉在地上部的积累(p<0.001),同时镉在地上部的积累还存在基因型和不同生长阶段的差异。整体而言,收获期时两个品种水稻地上部锌和硫含量较分蘖期都有减少,但收获期时两个品种地上部钙的含量却有所增加；加锌和加钙处理对水稻植株地上部干重有一定程度的增加,其中以分蘖期Teqing最为显著。
Rice (Oryza sativa L.) is one of the most important staple cereal crops in a large part of the world, especially in China. However, anthropogenic activities, such as agricultural practices, transportation, industrial activities and waste disposal have increased the concentrations of toxic elements such as cadmium (Cd) in agricultural soils. Cadmium has no metabolic role in living organisms, and is highly toxic and non-biodegradable. However it can be effectively absorbed and transported within rice plants. Thus, it can readily enter into the food chain, resulting in phytotoxicity and consequently causes serious threats to human health.
This dissertation aimed to understand the behavior of Cd speciation in rhizosphere microzone, determining the genetic difference between two rice cultivars in its growth, physiological-biochemical characteristics, absorption and accumulation regulation in the different rice tissues. The main experiments and conclusions are as follow:
1. To investigate rhizosphere effects on the chemical behavior of Cd, a glasshouse experiment was conducted in rhizobox, where two rice cultivars were grown in a soil spiked with cadmium at concentrations of3.9±0.5and8.3±0.5mg per kg soil until ripening stage. Chemical forms of cadmium with respect to root surface were then assessed using sequential extraction procedure (SEP). The results showed significant difference of Cd species, especially EXC-Cd (exchangeable-Cd) between two rice cultivars as affected by rice roots. The soil lowest EXC-Cd of Zhenong54appeared at near-rhizosphere area with a little difference under two Cd levels (lower Cd level showed the lowest EXC-Cd in1mm as higher Cd level showed in2mm) while Sixizhan had its lowest EXC-Cd at the root compartment. Both cultivars presented slight changes of FMO-Cd (Fe/Mn oxide-bound fraction) at ripening stage while the control treatments without plants had a significant increasing of FMO-Cd at ripening stage, indicating a transformation from less bioavailable form (FMO-Cd) to more bioavailable forms (EXC-Cd). Soil total microbial biomass carbon in the vicinity of the root surface had the opposite trends to some extent with EXC-Cd. Thus, the change of Cd speciation in the rhizosphere was mainly because of the suitable root exudates level, the microbial activities and the rice ability to absorb EXC-Cd in rhizosphere.
2. Plant Growth and physiological-biochemical characteristics under Cd pressure were investigated and the absorption regulation in the different rice tissues was studied. The results indicated that in tillering stage Sixizhan transferred more Cd from the root to the shoot, the height, dry weight and activities of Antioxidant enzyme system of Sixizhan were all decreased at some extent. Thus Cd caused more severe damage than Zhenong54in tillering stage. While in the harvesting stage, Sixizhan had less Cd accumulation in the shoots. However, Zhenong54had the just opposite trend Sixizhan had. Zhenong54had less Cd in the shoot and suffered less severe damage in the tillering stage, while Cd accumulation obviously increased in the harvesting stage, which finally caused larger grain Cd content in Zhenong54than in Sixizhan. Therefore, it is not appropriate to evaluate the effect of Cd pressure on rice just in one growth period because the Cd effects were different among different cultivars and among different growth periods.
3. In order to evaluate the root exudates difference between rice cultivars under different iron conditions and the exudates effect on Cd absorption by rice, a Hydroponic experiment was conducted under different Cd and Fe concentrations. The results showed that genetic-induced root exudates under different Fe conditions can finally affect the rice Cd absorption. The two rice cultivars indicated almost the same trend with or without Fe added:rice root exudates content (low molecular weight organic acid) was enlarged when Fe was not added, and the rice Cd accumulation was also increased whereas the dry weight of rice shoot and root were decreased. Despite those some trend, the two rice cultivars had their difference in different situations:with no Fe added, Sixizhan had larger root exudates amount than Zhenong54did, the Cd accumulation in its shoot and root was also obviously larger than Zhenong54s'; when Fe was added, the shoot and root dry weight of Zhenong54was larger than Sixizhan.
4. A green house experiment was conducted to study the different rice Cd absorption between different rice cultivars when exogenous Zn, Ca and S added. It was found that rice growth and plant Cd absorption difference between two rice cultivars were affected by exogenous Zn, Ca and S, and was changed with the rice growth period. Different added Cd levels obviously affected the rice Cd absorption and Cd distribution in different rice shoots (p<0.001). At mean time, Cd accumulation in rice shoots was varied between rice cultivars and rice growth period. Overall, the Zn and S contents in two rice cultivars decreased in harvesting stage, whereas the Ca contents increased; the Zn and Ca treatment increased rice shoot dry weight, especially in Teqing cultivar.
1.以我国东南地区水稻土黄斑田为供试土壤,籼稻四喜占(Sixizhan)和粳稻浙农54(Zhenong54)为供试水稻品种,采用多隔层三室根箱法开展盆栽试验(通过尼龙撩网插片的使用,把水稻根际土壤划分成根室,lmm、2mm、3mm、5mm近根际以及>5mm远根际等毫米级微域),通过改良的Tessier重金属分级提取法研究测定重金属镉在根际微域的形态变化(交换态、铁锰氧化态、有机质结合态、残留态)。实验结果表明,重金属镉各个形态含量,尤其是可交换态镉含量在根际微域中存在距离根系不同远近的差异,含量基本呈现根室、近根际<远根际的趋势；两个水稻品种在近根际不同毫米级微域以及根室中也存在着基因型的差异。就可交换态镉而言,浙农54的最低含量出现在近根际1或2mm处,而四喜占出现在根室,这与植物营养学中提到的营养因子根际梯度递减效应有所不同,说明重金属污染物的根际效应具有其特异性。相对应的是,两个水稻品种根际微域的土壤生物量碳含量趋势正好与土壤可交换态镉相反。由此可见,根室、近根际微域中有适宜的根系分泌物水平、根际微生物活性以及植物对交换态镉的吸收能力,这可能是诱导重金属镉形态变化的根本原因。
2.从阐明重金属镉在不同基因型水稻植株体内的生态行为角度出发,深入研究了重金属镉对两种不同基因型水稻生长抑制和抗氧化酶活性变化的差异,以及重金属镉在不同水稻品种体内迁移积累规律的差异。研究发现分蘖期镉胁迫下四喜占较浙农54受毒害严重,株高、干重以及POD和CAT酶的活性较无镉对照都有一定程度的下降,镉由根转移至茎、叶所占的比例较大；而至收获期,四喜占地上部分对镉的积累则显著减少。浙农54相较于四喜占有相反趋势,分蘖期时浙农54受毒害较轻,表现为株高、干重以及酶活性无明显抑制作用或高浓度时有抑制作用,且镉由根转移至地上部分的比例不大,而至收获期时,浙农54茎对锅的积累显著增加,最终导致浙农54籽粒含镉量较四喜占大。因此,不同品种或同一品种不同发育时期对镉胁迫的敏感性和耐受性不同,若仅研究镉胁迫对水稻某一时期的影响,结果有其局限性。
3.研究了缺铁和供铁条件下两种水稻品种根际分泌物的差异及其对植株吸收重金属镉的影响。结果表明,不同基因型水稻根际分泌物对供铁水平的响应差异与植株对镉的吸收有一定关系。两个水稻品种在缺铁和供铁条件下表现出的大致趋势相同：在缺铁处理中,水稻根系有机酸分泌的量较供铁处理都有显著的增加,植株对重金属镉的积累量也较供铁处理有显著的增加,但地上部分和根部干重较供铁处理有所下降。尽管趋势相同,但是两个水稻品种之间还是有一定的差异,其中四喜占缺铁处理时根系分泌物总量较浙农54缺铁处理根系分泌物总量大；四喜占根部和地上部对重金属镉的积累量均显著大于浙农54,加铁处理无显著差异；加铁处理的浙农54的地上部分和根部干重大于四喜占。
4.实验对镉胁迫下不同水稻基因型对镉、锌、钙、硫的吸收积累差异及三者与镉吸收积累之间的关系进行了研究。结果表明,锌、钙、硫对不同基因型水稻的生长及镉吸收有一定影响,并与不同生长阶段相关。不同处理中土壤镉的添加浓度极显著影响了两个品种水稻对镉的吸收和镉在地上部的积累(p<0.001),同时镉在地上部的积累还存在基因型和不同生长阶段的差异。整体而言,收获期时两个品种水稻地上部锌和硫含量较分蘖期都有减少,但收获期时两个品种地上部钙的含量却有所增加；加锌和加钙处理对水稻植株地上部干重有一定程度的增加,其中以分蘖期Teqing最为显著。
Rice (Oryza sativa L.) is one of the most important staple cereal crops in a large part of the world, especially in China. However, anthropogenic activities, such as agricultural practices, transportation, industrial activities and waste disposal have increased the concentrations of toxic elements such as cadmium (Cd) in agricultural soils. Cadmium has no metabolic role in living organisms, and is highly toxic and non-biodegradable. However it can be effectively absorbed and transported within rice plants. Thus, it can readily enter into the food chain, resulting in phytotoxicity and consequently causes serious threats to human health.
This dissertation aimed to understand the behavior of Cd speciation in rhizosphere microzone, determining the genetic difference between two rice cultivars in its growth, physiological-biochemical characteristics, absorption and accumulation regulation in the different rice tissues. The main experiments and conclusions are as follow:
1. To investigate rhizosphere effects on the chemical behavior of Cd, a glasshouse experiment was conducted in rhizobox, where two rice cultivars were grown in a soil spiked with cadmium at concentrations of3.9±0.5and8.3±0.5mg per kg soil until ripening stage. Chemical forms of cadmium with respect to root surface were then assessed using sequential extraction procedure (SEP). The results showed significant difference of Cd species, especially EXC-Cd (exchangeable-Cd) between two rice cultivars as affected by rice roots. The soil lowest EXC-Cd of Zhenong54appeared at near-rhizosphere area with a little difference under two Cd levels (lower Cd level showed the lowest EXC-Cd in1mm as higher Cd level showed in2mm) while Sixizhan had its lowest EXC-Cd at the root compartment. Both cultivars presented slight changes of FMO-Cd (Fe/Mn oxide-bound fraction) at ripening stage while the control treatments without plants had a significant increasing of FMO-Cd at ripening stage, indicating a transformation from less bioavailable form (FMO-Cd) to more bioavailable forms (EXC-Cd). Soil total microbial biomass carbon in the vicinity of the root surface had the opposite trends to some extent with EXC-Cd. Thus, the change of Cd speciation in the rhizosphere was mainly because of the suitable root exudates level, the microbial activities and the rice ability to absorb EXC-Cd in rhizosphere.
2. Plant Growth and physiological-biochemical characteristics under Cd pressure were investigated and the absorption regulation in the different rice tissues was studied. The results indicated that in tillering stage Sixizhan transferred more Cd from the root to the shoot, the height, dry weight and activities of Antioxidant enzyme system of Sixizhan were all decreased at some extent. Thus Cd caused more severe damage than Zhenong54in tillering stage. While in the harvesting stage, Sixizhan had less Cd accumulation in the shoots. However, Zhenong54had the just opposite trend Sixizhan had. Zhenong54had less Cd in the shoot and suffered less severe damage in the tillering stage, while Cd accumulation obviously increased in the harvesting stage, which finally caused larger grain Cd content in Zhenong54than in Sixizhan. Therefore, it is not appropriate to evaluate the effect of Cd pressure on rice just in one growth period because the Cd effects were different among different cultivars and among different growth periods.
3. In order to evaluate the root exudates difference between rice cultivars under different iron conditions and the exudates effect on Cd absorption by rice, a Hydroponic experiment was conducted under different Cd and Fe concentrations. The results showed that genetic-induced root exudates under different Fe conditions can finally affect the rice Cd absorption. The two rice cultivars indicated almost the same trend with or without Fe added:rice root exudates content (low molecular weight organic acid) was enlarged when Fe was not added, and the rice Cd accumulation was also increased whereas the dry weight of rice shoot and root were decreased. Despite those some trend, the two rice cultivars had their difference in different situations:with no Fe added, Sixizhan had larger root exudates amount than Zhenong54did, the Cd accumulation in its shoot and root was also obviously larger than Zhenong54s'; when Fe was added, the shoot and root dry weight of Zhenong54was larger than Sixizhan.
4. A green house experiment was conducted to study the different rice Cd absorption between different rice cultivars when exogenous Zn, Ca and S added. It was found that rice growth and plant Cd absorption difference between two rice cultivars were affected by exogenous Zn, Ca and S, and was changed with the rice growth period. Different added Cd levels obviously affected the rice Cd absorption and Cd distribution in different rice shoots (p<0.001). At mean time, Cd accumulation in rice shoots was varied between rice cultivars and rice growth period. Overall, the Zn and S contents in two rice cultivars decreased in harvesting stage, whereas the Ca contents increased; the Zn and Ca treatment increased rice shoot dry weight, especially in Teqing cultivar.
引文
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