水稻钾离子通道OsAKT1及其调控因子参与水稻钾吸收的实验证据
摘要
钾(K+)是植物生长发育所必需的三大营养元素之一,在植物的生长发育过程中起着十分重要的作用。已有研究表明,植物通过钾离子通道和钾离子转运体蛋白从外界吸收K+,并在体内进行运输和分配。对于模式植物拟南芥,目前已有多个钾通道和钾转运体的生理功能得到了详细报道。AKT1是拟南芥根中最重要的钾离子通道之一,也是介导拟南芥根中K+吸收的主要途径之一,其活性受到上游多种调节因子的控制。然而,在作物中,虽然已经克隆了部分钾通道和转运体,但是这些钾通道和钾转运体在作物体内的生理功能仍然不清楚。本论文工作以水稻为材料,主要研究水稻钾离子通道OsAKT1的生理功能及其分子调控机制。
氨基酸序列比对分析发现,水稻OsAKT1与拟南芥AKT1具有较高的序列相似性,预示它们可能具有类似的生理功能。将OsAKT1转入酵母钾吸收缺陷型菌株后,能够恢复缺陷型菌株在低钾培养基上的生长,说明OsAKT1具有K+转运活性。将OsAKT1转入拟南芥akt1突变体后,可以恢复akt1突变体冠部发黄的低钾敏感表型,说明OsAKT1在植物体内可能具有K+吸收功能。对水稻osakt1突变体进行低钾水培表型检测,结果显示,与野生型相比,osakt1突变体的生长受到显著抑制,叶片出现明显缺钾褐斑,植株K+含量显著下降。钾吸收动力学实验结果显示,osakt1突变体的K+吸收速率显著低于野生型,说明OsAKT1的功能缺失造成水稻K+吸收能力的下降,从而导致植株出现低钾敏感表型。土培实验还表明,osaktl突变体的抽穗期和灌浆期都滞后于野生型,并且结实也受到严重影响,产量显著降低。由此可知,OsAKT1作为一个钾离子通道,在水稻根部K+吸收过程中起着重要作用。
本论文工作还对OsAKT1的调控因子进行了筛选和功能验证。拟南芥中的研究结果显示,AKT1活性受到钙感受器CBL1/9和蛋白激酶CIPK23的调控。由此推测OsAKT1的活性也可能受到水稻OsCBL1和OsCIPK23的调控。表达检测结果显示,OsCBL1和OsCIPK23在水稻根部均有表达,并且不同程度的受低钾诱导表达。酵母双杂交实验结果显示,OsCIPK23能够与OsAKT1互作,而OsCBLl又能够与OsCIPK23互作。OsCIPK23可以恢复拟南芥Iks1(cipk23)突变体的低钾敏感表型,同时OsCBLl可以恢复拟南芥cbll cbl9双突变体的低钾敏感表型。说明水稻OsCBL1和OsCIPK23可能具有与拟南芥CBLl和CIPK23类似的生理功能。此外,表型检测结果还显示,水稻OsCIPK23RNAi株系在低钾水培处理下也表现出与osakt1突变体类似的低钾敏感表型,说明OsCBL1和OsCIPK23应该与OsAKT1处于同一调控通路,参与调控OsAKT1介导的水稻K+吸收。
本论文研究结果表明,OsAKT1介导水稻根细胞从土壤中吸收K+,从而控制水稻的钾营养性状,最终影响水稻的产量和品质。OsAKT1的活性受到钙感受器蛋白OsCBL1和蛋白激酶OsCIPK23的调控。在水稻中也存在和拟南芥中类似的钾营养分子调控通路。这些研究结果将为我们利用分子生物学手段改良水稻品质,提高水稻的K+吸收利用效率提供有力的理论依据。
Potassium (K+) is the most abundant monovalent cation in plant cells, and plays crucial roles in plant growth and development. Potassium in plants is mainly absorbed from soil, which is mediated by K+channels and K+transporters. In the model plant Arabidopsis, K+channel AKT1is identified as the important component involving in root K+uptake, whose activity is modulated by many different regulators. For the crops, some K+channels and K transporters were also cloned, however, their physiological functions in crops are still unclear. In this dissertation work, we analyzed the physiological function of K+channel OsAKT1in rice. In addition, the regulators of OsAKT1were screened, whose physiological functions were also verified in rice.
Rice OsAKT1is the homolog of Shaker potassium channel AKT1from Arabidopsis. The amino acid sequence analysis showed that OsAKT1and AKT1are highly conserved, which indicated their similar physiological functions in plants. OsAKT1could complement the growth of yeast K+defect mutant strain R5421suggesting the K+transport activity of OsAKT1.In addition, OsAKT1could rescue the low-K+-sensitive phenotype of Arabidopsis aktl mutant. The phenotype analysis showed that the rice osaktl mutant plants displayed the obvious low-K+-sensitive phenotype compared with the wild-type rice plants. The osaktl mutant showed growth inhibition, leaf brown spots and reduction of K1content. The results of K--depletion experiments showed that the K+uptake rate in osaktl mutant plants was obviously slower than that in wild-type plants. These results demonstrated that the loss-of-function of OsAKT1led to the reduction of K+uptake in osaktl mutant, which caused the growth inhibition and K+-deficient symptom in mutant plants. Furthermore, the growth of osaktl mutant plants was inhibited during the whole development stages. The heading and grain-filling stages were delayed in osaktl mutant, and the grain yield was also impaired. These data demonstrate that the K+channel OsAKT1plays a crucial role in rice root K+uptake.
Furthermore, we tried to screen the regulators of OsAKT1and verified the physiological functions of these regulators. According to the investigation of AKT1in Arabidopsis, the rice calcium sensor OsCBLl and protein kinase OsCIPK23are proposed as the regulators of OsAKT1. OsCBL1and OsCIPK23were both expressed in rice roots, and especially OsCIPK23could be induced by low-K+treatment. Yeast two-hybrid results showed that OsCIPK23could interact with OsAKT1as well as OsCBL1. Phenotype analysis indicated that OsCIPK23and OsCBL1could rescue the low-K'-sensitive phenotype of Arabidopsis Iksl (cipk23) and cbll cbl9mutants, respectively. Moreover, OsCIPK23RNAi rice plants displayed the similar low-K+-sensitive phenotype as osaktl mutant. These results indicated that OsAKT1-mediated K+uptake in rice roots is modulated by OsCBLl-OsCIPK23complex.
All the data in this dissertation demonstrate that K+channel OsAKT1is an important component mediated K+uptake in rice roots. The activity of OsAKT1is regulated by the calcium sensor OsCBLl and protein kinase OsCIPK23. These results will provide the theoretical basis that would be used to improve the rice K+utilization efficiency via molecular biology methods.
氨基酸序列比对分析发现,水稻OsAKT1与拟南芥AKT1具有较高的序列相似性,预示它们可能具有类似的生理功能。将OsAKT1转入酵母钾吸收缺陷型菌株后,能够恢复缺陷型菌株在低钾培养基上的生长,说明OsAKT1具有K+转运活性。将OsAKT1转入拟南芥akt1突变体后,可以恢复akt1突变体冠部发黄的低钾敏感表型,说明OsAKT1在植物体内可能具有K+吸收功能。对水稻osakt1突变体进行低钾水培表型检测,结果显示,与野生型相比,osakt1突变体的生长受到显著抑制,叶片出现明显缺钾褐斑,植株K+含量显著下降。钾吸收动力学实验结果显示,osakt1突变体的K+吸收速率显著低于野生型,说明OsAKT1的功能缺失造成水稻K+吸收能力的下降,从而导致植株出现低钾敏感表型。土培实验还表明,osaktl突变体的抽穗期和灌浆期都滞后于野生型,并且结实也受到严重影响,产量显著降低。由此可知,OsAKT1作为一个钾离子通道,在水稻根部K+吸收过程中起着重要作用。
本论文工作还对OsAKT1的调控因子进行了筛选和功能验证。拟南芥中的研究结果显示,AKT1活性受到钙感受器CBL1/9和蛋白激酶CIPK23的调控。由此推测OsAKT1的活性也可能受到水稻OsCBL1和OsCIPK23的调控。表达检测结果显示,OsCBL1和OsCIPK23在水稻根部均有表达,并且不同程度的受低钾诱导表达。酵母双杂交实验结果显示,OsCIPK23能够与OsAKT1互作,而OsCBLl又能够与OsCIPK23互作。OsCIPK23可以恢复拟南芥Iks1(cipk23)突变体的低钾敏感表型,同时OsCBLl可以恢复拟南芥cbll cbl9双突变体的低钾敏感表型。说明水稻OsCBL1和OsCIPK23可能具有与拟南芥CBLl和CIPK23类似的生理功能。此外,表型检测结果还显示,水稻OsCIPK23RNAi株系在低钾水培处理下也表现出与osakt1突变体类似的低钾敏感表型,说明OsCBL1和OsCIPK23应该与OsAKT1处于同一调控通路,参与调控OsAKT1介导的水稻K+吸收。
本论文研究结果表明,OsAKT1介导水稻根细胞从土壤中吸收K+,从而控制水稻的钾营养性状,最终影响水稻的产量和品质。OsAKT1的活性受到钙感受器蛋白OsCBL1和蛋白激酶OsCIPK23的调控。在水稻中也存在和拟南芥中类似的钾营养分子调控通路。这些研究结果将为我们利用分子生物学手段改良水稻品质,提高水稻的K+吸收利用效率提供有力的理论依据。
Potassium (K+) is the most abundant monovalent cation in plant cells, and plays crucial roles in plant growth and development. Potassium in plants is mainly absorbed from soil, which is mediated by K+channels and K+transporters. In the model plant Arabidopsis, K+channel AKT1is identified as the important component involving in root K+uptake, whose activity is modulated by many different regulators. For the crops, some K+channels and K transporters were also cloned, however, their physiological functions in crops are still unclear. In this dissertation work, we analyzed the physiological function of K+channel OsAKT1in rice. In addition, the regulators of OsAKT1were screened, whose physiological functions were also verified in rice.
Rice OsAKT1is the homolog of Shaker potassium channel AKT1from Arabidopsis. The amino acid sequence analysis showed that OsAKT1and AKT1are highly conserved, which indicated their similar physiological functions in plants. OsAKT1could complement the growth of yeast K+defect mutant strain R5421suggesting the K+transport activity of OsAKT1.In addition, OsAKT1could rescue the low-K+-sensitive phenotype of Arabidopsis aktl mutant. The phenotype analysis showed that the rice osaktl mutant plants displayed the obvious low-K+-sensitive phenotype compared with the wild-type rice plants. The osaktl mutant showed growth inhibition, leaf brown spots and reduction of K1content. The results of K--depletion experiments showed that the K+uptake rate in osaktl mutant plants was obviously slower than that in wild-type plants. These results demonstrated that the loss-of-function of OsAKT1led to the reduction of K+uptake in osaktl mutant, which caused the growth inhibition and K+-deficient symptom in mutant plants. Furthermore, the growth of osaktl mutant plants was inhibited during the whole development stages. The heading and grain-filling stages were delayed in osaktl mutant, and the grain yield was also impaired. These data demonstrate that the K+channel OsAKT1plays a crucial role in rice root K+uptake.
Furthermore, we tried to screen the regulators of OsAKT1and verified the physiological functions of these regulators. According to the investigation of AKT1in Arabidopsis, the rice calcium sensor OsCBLl and protein kinase OsCIPK23are proposed as the regulators of OsAKT1. OsCBL1and OsCIPK23were both expressed in rice roots, and especially OsCIPK23could be induced by low-K+treatment. Yeast two-hybrid results showed that OsCIPK23could interact with OsAKT1as well as OsCBL1. Phenotype analysis indicated that OsCIPK23and OsCBL1could rescue the low-K'-sensitive phenotype of Arabidopsis Iksl (cipk23) and cbll cbl9mutants, respectively. Moreover, OsCIPK23RNAi rice plants displayed the similar low-K+-sensitive phenotype as osaktl mutant. These results indicated that OsAKT1-mediated K+uptake in rice roots is modulated by OsCBLl-OsCIPK23complex.
All the data in this dissertation demonstrate that K+channel OsAKT1is an important component mediated K+uptake in rice roots. The activity of OsAKT1is regulated by the calcium sensor OsCBLl and protein kinase OsCIPK23. These results will provide the theoretical basis that would be used to improve the rice K+utilization efficiency via molecular biology methods.
引文
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