盐碱胁迫对转Lc-CDPK基因水稻抗氧化酶活性及基因表达的影响
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摘要
【目的】研究盐碱胁迫对转羊草钙依赖蛋白激酶基因(Lc-CDPK)水稻根、叶片抗氧化酶活性及基因表达的影响。【方法】以转Lc-CDPK水稻为试验材料,通过qRT-PCR技术,测定根、叶片中Lc-CDPK基因的表达,确定该基因表达最高时的盐碱胁迫条件,以此条件处理幼苗,并以非转基因水稻为对照,分别测定其根、叶片中甜菜碱、脯氨酸、甘露醇、山梨醇、蔗糖等渗透调节物质含量和超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)、谷胱甘肽还原酶(GR)、谷胱甘肽过氧化物酶(GPX)等抗氧化酶活性及相关基因的表达。【结果】200 mmol/L NaCl-Na_2CO_3胁迫24 h时,转基因水稻根和叶片中Lc-CDPK基因表达的综合水平最高,此时转基因水稻和非转基因水稻(对照)的渗透调节物质含量和抗氧化酶活性有显著差异,转基因水稻根中脯氨酸、甘露醇和山梨醇含量分别较对照增加116.0%,88.9%和115.4%(P<0.01),GR活性较对照提高9.6%(P<0.05);叶片中脯氨酸含量较对照增加129.2%(P<0.01),CAT、POD、GR和GPX活性分别较对照提高52.1%,27.5%,163.5%和46.7%。叶片中抗氧化酶基因相对表达显著增强,CAT、GPX基因相对表达量分别是对照的3.96和2.73倍(P<0.01),根中仅GR基因相对表达增强,为对照的1.6倍(P<0.01)。相关分析表明,根、叶片抗氧化酶活性与其基因相对表达量呈正相关。【结论】推测Lc-CDPK基因可能参与调控盐碱胁迫过程,能增强植株渗透调节物质的合成能力及抗氧化酶活性,提高水稻对盐碱胁迫的耐受性。
【Objective】 The effects of saline-alkali stress on antioxidant enzyme activities and gene expression in roots and leaves of transgenic rice with Leymus chinensis Ca~(2+)-dependent protein kinase gene(Lc-CDPK) were analyzed.【Method】 Using qRT-PCR,the most appropriate treatment conditions for highest gene expression in roots and leaves of transgenic rice were clarified.After saline-alkali stress treatments,the contents of betaine,proline,mannitol,sorbitol and sucrose were measured and the activities and gene expression of antioxidant enzymes including superoxide dismutase(SOD),peroxidase(POD),catalase(CAT),glutathione reductase(GR) and glutathione peroxidase(GPX) in seedlings were assayed.【Result】 The expression levels of Lc-CDPK in roots and leaves of transgenic rice were the highest 24 h after treatment by 200 mmol/L NaCl-Na_2CO_3.Under this stress condition,difference between transgenic rice and no-transgenic rice(CK) was significant in contents of osmotic regulation solutes and activities of antioxidant enzymes.The contents of proline, mannitol and sorbitol were increased by 116.0%,88.9% and 115.4% comparing CK(P<0.01),and GR activity was increased by 9.6%(P<0.05).The proline content in leaves was increased by 129.2%(P<0.01),and the activities of CAT,POD,GR and GPX was 52.1%,27.5%,163.5% and 46.7% higher than those of CK in leaves.The relative expression levels and antioxidant enzyme activities in leaves of the transgenic rice were enhanced,and relative expression levels of CAT,GPX of transgenic rice were 3.96 and 2.73 times of CK(P<0.01).Relative expression levels of GR were only increased in roots,which was 1.6 times of CK(P<0.01).Correlation analysis indicated that the gene expression was positively correlated with activities of antioxidant enzymes in roots and leaves.【Conclusion】 This study indicated that the Lc-CDPK involved in regulation of saline-alkali stress response,which increased the accumulation of osmotic regulators and activities of antioxidant enzymes,and improved the tolerance to saline-alkali stress.
【Objective】 The effects of saline-alkali stress on antioxidant enzyme activities and gene expression in roots and leaves of transgenic rice with Leymus chinensis Ca~(2+)-dependent protein kinase gene(Lc-CDPK) were analyzed.【Method】 Using qRT-PCR,the most appropriate treatment conditions for highest gene expression in roots and leaves of transgenic rice were clarified.After saline-alkali stress treatments,the contents of betaine,proline,mannitol,sorbitol and sucrose were measured and the activities and gene expression of antioxidant enzymes including superoxide dismutase(SOD),peroxidase(POD),catalase(CAT),glutathione reductase(GR) and glutathione peroxidase(GPX) in seedlings were assayed.【Result】 The expression levels of Lc-CDPK in roots and leaves of transgenic rice were the highest 24 h after treatment by 200 mmol/L NaCl-Na_2CO_3.Under this stress condition,difference between transgenic rice and no-transgenic rice(CK) was significant in contents of osmotic regulation solutes and activities of antioxidant enzymes.The contents of proline, mannitol and sorbitol were increased by 116.0%,88.9% and 115.4% comparing CK(P<0.01),and GR activity was increased by 9.6%(P<0.05).The proline content in leaves was increased by 129.2%(P<0.01),and the activities of CAT,POD,GR and GPX was 52.1%,27.5%,163.5% and 46.7% higher than those of CK in leaves.The relative expression levels and antioxidant enzyme activities in leaves of the transgenic rice were enhanced,and relative expression levels of CAT,GPX of transgenic rice were 3.96 and 2.73 times of CK(P<0.01).Relative expression levels of GR were only increased in roots,which was 1.6 times of CK(P<0.01).Correlation analysis indicated that the gene expression was positively correlated with activities of antioxidant enzymes in roots and leaves.【Conclusion】 This study indicated that the Lc-CDPK involved in regulation of saline-alkali stress response,which increased the accumulation of osmotic regulators and activities of antioxidant enzymes,and improved the tolerance to saline-alkali stress.
引文
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[2] 王云贺,王志春,杨帆,等.不同改良物质对苏打碱土盐碱度及水稻生长的影响 [J].华南农业大学学报,2013,34(4):445-449. Wang Y H,Wang Z C,Yang F,et al.Effect of different ameliorate materials on soda alkaline-alkalinity and rice growth [J].Journal of South China Agricultural University,2013,34(4):445-449.
[3] Chen Y X,Zhou X J,Chang S,et al.Calcium-dependent protein kinases 21 phosphorylates 14-3-3 proteins in response to ABA signaling and salt stress in rice [J].Biochem Bioph Res Co,2017,493(4):1450-1456.
[4] Cheng S H,Willmann M R,Chen H C,et al.Calcium signaling through protein kinases:the Arabidopsis calcium-dependent protein kinase gene family [J].Plant Physiol,2002,129(2):469-485.
[5] Asano T,Tanakan,Yang G,et al.Genome-wide identification of the rice calcium-dependent proteinkinase and its closely related kinase gene family:comprehensive analysis of the CDPKs gene family in rice [J].Plant Cell Physiol,2005,46(2):356-366.
[6] 李月树,祝廷成.羊草种群地上生物量形成规律的探讨 [J].植物生态学与地植物学丛刊,1983,7(4):289-298. Li Y S,Zhu T C.The regularities of formation of aboveground biomass of Leymus chinensis' population [J].Acta Phytoecologica et Geobotanice Sinica,1983,7(4):289-298.
[7] 崔喜艳,张继伟,秦思余,等.羊草钙依赖蛋白激酶基因生物信息学及表达特性分析 [J].中国草地学报,2015,37(3):11-18. Cui X Y,Zhang J W,Qin S Y,et al.Analysis of bioinformatics and expression pattern of calcium-dependent protein kinase gene in Leymus chinensis [J].Chinese Journal of Grassland,2015,37(3):11-18.
[8] Sheen J.Ca2+-dependent protein kinases and stress signal transduction in plants [J].Science,1996,274(5294):1900-1902.
[9] 郑炳松,朱诚,金松恒.高级植物生理学 [M].杭州:浙江大学出版社,2011:222-223. Zheng B S,Zhu C,Jin S H.Advanced plant physiology [M].Hangzhou:Zhejiang University Press,2011:222-223.
[10] 麻浩,王爽,周亚丽. 植物中钙依赖蛋白激酶的研究进展[J].南京农业大学学报,2017,40(4):565-572. Ma H,Wang S,Zhou Y L.Research progress of calcium-dependent protein kinases in plants [J].Journal of Nanjing Agricultural University,2017,40(4):565-572.
[11] 王佺珍,刘倩,高娅妮,等.植物对盐碱胁迫的响应机制研究进展 [J].生态学报,2017,37(16):5565-5577. Wang Q Z,Liu Q,Gao Y N,et al.Review on the mechanisms of the response to salinity-alkalinity stress in plants [J].Acta Ecologica Sinica,2017,37(16):5565-5577.
[12] 陈芸,郑勇,刘霞,等.过量表达棉花CBF2基因提高转基因拟南芥抗旱耐盐能力 [J].植物科学学报,2016,34(6):888-900. Chen Y,Zheng Y,Liu X,et al.Overexpression of the cotton CBF2 gene enhances salt and drought tolerance in Arabidopsis thaliana [J].Plant Science Journal,2016,34(6):888-900.
[13] 刘文瑜,杨发荣,黄杰,等.NaCl 胁迫对藜麦幼苗生长和抗氧化酶活性的影响 [J].西北植物学报,2017,37(9):1797-1804. Liu W Y,Yang F R,Huang J,et al.Response of seedling growth and the activities of antioxidant enzymes of Chenopodium quinoato salt stress [J].Acta Bot Boreal-Occident Sin,2017,37(9):1797-1804.
[14] Andrès C G,Leandro A S,Luiz G A,et al.Vermicompost humic acids modulate the accumulation and metabolism of ROS in rice plants [J].J Plant Physiol,2016,192(15):56-63.
[15] Shirasawa K,Takabe T,Takabe T,et al.Accumulation of glycinebetainein rice plant that overexpress choline monooxygenase from spinach and evaluation of their tolerance to abiotic stress [J].Annals of Botany,2006,98(3):565-571.
[16] Wang Y,Gao P,Huang M,et al.Effects of high temperature on the activity and expression of antioxidative enzymes in rice flag leaves during the flowering stage [J].Plant Science Journal,2015,33(3):355-361.
[17] Hu T,Li H Y,Zhang X Z,et al.Toxic effect of NaCl on ion metabolism,antioxidative enzymes and gene expression of perennial ryegrass [J].Ecotoxicology and Environmental Safety,2011,74(7):2050-2056.
[18] 丁艳芬.玉米ZmCPK11在ABA诱导的抗氧化防护中的功能分析 [D].南京:南京农业大学,2012. Ding Y F.Function analysis of ZmCPK11 in the ABA-induced antioxidant defense in maize leave [D].Nanjing:Nanjing Agricultural University,2012.
[19] 张庆华.枳两个基因ptrABF和ptrCDPK转化枳的抗逆功能及其作用机制解析 [D].武汉:华中农业大学,2015. Zhang Q H.Function characterization of stress-response and elucidation of mechanism underling the enhanced stress tolerance of ptrABF and ptrCDPK in Poncirus trifoliata [D].Wuhan:Central China Agricultural University,2015.
[20] Yoshida S,Forno D A,Cock J H,et al.Laboratory manual for physiological studies of rice [M].Manila:Int Rice Res Inst,1976.
[21] 汤章城.现代植物生理学实验指南 [M].北京:科技出版社,1999:127,303-304. Tang Z C.Modern plant physiology experiment [M].Beijing: Science Press,1999:127,303-304.
[22] 梁振明.分光光度法测定山梨醇含量 [J].中国食品添加剂,2007(1):184-185,183. Liang Z M.Determination of sorbitol content by spectrophotometry [J].China Food Additives,2007(1):184-185,183.
[23] 焦锐,邹翔,王英.分光光度法测定海产藻类中甘露醇的含量 [J].连云港师范高等专科学校学报,2016(1):106-108. Jiao R,Zou X,Wang Y.Determination of mannitol in sea algaes by spectrophotometry [J].Journal of Lianyungang Normal College,2016(1):106-108.
[24] 李合生.植物生理生化实验原理与技术 [M].北京:高等教育出版社,2000:164-168. Li H S.Principles and techniques of plant physiological biochemical experiment [M].Beijing:Higher Education Press,2000:164-168.
[25] Khatun S,Ali M B,Hahn E, et al.Copper toxicity in Withania somnifera:growth and antioxidant enzymes responses of in vitro grown plants [J].Environmental and Experimental Botany,2008,64(3):279-285.
[26] Hodges D M,Andrews C J,Johnson D A,et al.Antioxidant enzyme responses to chilling stress in differentially sensitive inbred maize lines [J].Journal of Experimental Botany,1997,48(310):1105-1113.
[27] Livak K J,Schmitttgen T D. Analysis of relative gene expression data using real-time quantitative PCR and the 2-ΔΔCTmethod [J].Methods,2001,25(4):402-408.
[28] Fridovich I.Superoxide dismutases [J].Annu Rev Biochem,1975,44:147-159.
[29] Bai L P,Sui F G,Ge T D,et al.Effect of soil drought stress on leaf water status,membrane permeability and enzymatic antioxidant system of maize [J].Pedosphere,2006,16(3):326-332.
[30] Campos M K F,Carvalho K,Souza F S,et al.Drought tolerance and antioxidant enzymatic activity in transgenic ‘Swingle' citrumelo plants over-accumulating proline [J].Environ Exp Bot,2011,72:242-250.
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