根区亚低温胁迫下外源GABA对山定子根系抗氧化功能的影响
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摘要
以当年生山定子实生苗为试材,采用温室盆栽土培试验,模拟根区亚低温(5℃)条件,设置亚低温(L)、亚低温+外源γ氨基丁酸(LG)和亚低温+氨己烯酸(LV)处理,分别于处理后0、12、24、48、96和144h后剪取白色幼嫩根系,测定山定子根系内源γ-氨基丁酸(GABA)含量、渗透调节物质含量、活性氧含量和抗氧化酶活性等指标,研究根区亚低温下GABA对山定子根系抗氧化系统的调节效应。结果显示:(1)根区亚低温下,山定子根系内源GABA含量略高于对照,渗透调节物质也不同程度积累,超氧阴离子(O_2~-·)含量、过氧化氢(H_2O_2)含量显著升高,膜脂过氧化程度加深。(2)外源施加GABA进一步促进了内源GABA的积累,明显增加根系中可溶性糖、可溶性蛋白和脯氨酸(Pro)等渗透调节物质含量,显著提高根系的超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、抗坏血酸过氧化物酶(APX)等酶活性,最终使得山定子根系中H_2O_2等的含量降低,膜脂过氧化程度缓解。(3)GABA专一性抑制剂VGB处理后的效果与外源GABA处理结果相反。研究表明,GABA代谢对山定子根系抵御亚低温胁迫引发的氧化胁迫有积极作用,外源GABA具有通过诱导内源GABA代谢提高根系抗氧化能力的生理效应。
The potted seedlings of Malus baccata were used as materials in the treatments of simulated suboptimal root-zone temperature(5 ℃)in culture room.The treatments included the suboptimal root-zone temperature(L),L +exogenous GABA(LG)and L + VGB(LV),the white roots were cut at 0,12,24,48,96,and 144 hafter treatment,respectively.We measured the contents of endogenous GABA,osmotic adjustment substances and reactive oxygen species,the activities of antioxidant enzymes to investigate the regulating effect of GABA on the antioxidant system in roots under suboptimal root-zone temperature.The results showed that:(1)the endogenous GABA content in the roots of Malus baccataincreased under the L treatment,osmotic adjustment substances accumulated in different degrees.And as a consequence,the contents of superoxide anion (O_2~-) and hydrogen peroxide(H_2O_2)increased significantly.The peroxidation in roots was deepened.(2)Exogenous GABA increased the contents of endogenous GABA in roots,soluble sugar,soluble protein and proline(Pro)in the roots,significantly enhanced the activities of superoxide dismutase(SOD),catalase(CAT)and ascorbate peroxidase(APX)in roots,and effectively reduced the contents of H_2O_2 and O_2~-,which meant exogenous GABA could significantly alleviate the degree of membrane lipid peroxidation.(3)The effect of VGB,which is the specific inhibitor of GABA(LV),treatment was opposite to that of LG treatment.Those results indicated that the GABA metabolism had a positive effect on the defense of the roots to the oxidative stress triggered by suboptimal rootzone temperature,and exogenous GABA had the effect of enhancing the antioxidant capacity of the roots through regulating endogenous GABA metabolism.
The potted seedlings of Malus baccata were used as materials in the treatments of simulated suboptimal root-zone temperature(5 ℃)in culture room.The treatments included the suboptimal root-zone temperature(L),L +exogenous GABA(LG)and L + VGB(LV),the white roots were cut at 0,12,24,48,96,and 144 hafter treatment,respectively.We measured the contents of endogenous GABA,osmotic adjustment substances and reactive oxygen species,the activities of antioxidant enzymes to investigate the regulating effect of GABA on the antioxidant system in roots under suboptimal root-zone temperature.The results showed that:(1)the endogenous GABA content in the roots of Malus baccataincreased under the L treatment,osmotic adjustment substances accumulated in different degrees.And as a consequence,the contents of superoxide anion (O_2~-) and hydrogen peroxide(H_2O_2)increased significantly.The peroxidation in roots was deepened.(2)Exogenous GABA increased the contents of endogenous GABA in roots,soluble sugar,soluble protein and proline(Pro)in the roots,significantly enhanced the activities of superoxide dismutase(SOD),catalase(CAT)and ascorbate peroxidase(APX)in roots,and effectively reduced the contents of H_2O_2 and O_2~-,which meant exogenous GABA could significantly alleviate the degree of membrane lipid peroxidation.(3)The effect of VGB,which is the specific inhibitor of GABA(LV),treatment was opposite to that of LG treatment.Those results indicated that the GABA metabolism had a positive effect on the defense of the roots to the oxidative stress triggered by suboptimal rootzone temperature,and exogenous GABA had the effect of enhancing the antioxidant capacity of the roots through regulating endogenous GABA metabolism.
引文
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[5]罗黄颖,高洪波,夏庆平,等.γ-氨基丁酸对盐胁迫下番茄活性氧代谢及叶绿素荧光参数的影响[J].中国农业科学,2011,44(4):753-761.LUO H Y,GAO H B,XIA Q P,et al.Effects of exogenous GABA on reactive oxygen species metabolism and chlorophyll fluorescence parameters in tomato under NaCl stress[J].Scientia Agricultura Sinica,2011,44(4):753-761.
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[9]MITTLER R,VANDERAUWERA S,GOLLERY M,et al.Reactive oxygen gene network of plants[J].Trends in Plant Science,2004,9(10):490-498.
[10]马金虎,邢国芳,杨小环,等.外源EBR和NO信号对低温胁迫下玉米种胚抗氧化系统和低温响应基因表达的影响[J].应用生态学报,2015,26(5):1 411-1 418.MA J H,XING G F,YANG X H,et al.Effects of exogenous EBR and NO signal on antioxidant system and low response gene expression under cold stress on maize embryo[J].Chinese Journal of Applied Ecology,2015,26(5):1 411-1 418.
[11]张帆,郁继华,颉建明,等.外源ALA和Spd对低温弱光下辣椒幼苗光合作用及抗氧化系统的影响[J].中国农业科学,2013,46(11):2 298-2 306.ZHANG F,YU J H,XIE J M,et al.Influence of exogenous ALA and spd on photosynthesis and antioxidant system of low temperature and poor light affected pepper seedlings[J].Scientia Agricultura Sinica,2013,46(11):2 298-2 306.
[12]向丽霞,胡立盼,胡晓辉,等.外源γ-氨基丁酸调控甜瓜叶绿体活性氧代谢应对短期盐碱胁迫[J].应用生态学报,2015,26(12):3 746-3 752.XIANG L X,HU L P,HU X H,et al.Response of reactive oxygen metabolism in melon chloroplasts to short-term salinity-alkalinity stress regulated by exogenousγ-aminobutyric acid[J].Chinese Journal of Applied Ecology,2015,26(12):3 746-3 752.
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[14]BAI T H,LI C Y,MA F W,et al.Responses of growth and antioxidant system to root-zone hypoxia stress in two Malus species[J].Plant and Soil,2010,327(1-2):95-105.
[15]SHAH K,KUMAR R G,VERMA S,et al.Effect of cadmium on lipid peroxidation,superoxide anion generation and activities of antioxidant enzymes in growing rice seedlings[J].Plant Science,2001,161(6):1 135-1 144.
[16]王爱国,罗广华,邵从本,等.大豆种子超氧物歧化酶的研究[J].植物生理学报,1983,(1):77-84.WANG A G,LUO G H,SHAO C B,et al.A study on the superoxide dismutase of soybean seeds[J].Journal of Plant Physiology,1983,(1):77-84.
[17]杨兰芳,庞静,彭小兰,等.紫外分光光度法测定植物过氧化氢酶活性[J].现代农业科技,2009,(20):364-366.YANG L F,PANG J,PENG X L,et al.Measurement of catalase activity in plants by ultraviolet spectrophotometry[J].Modern Agricultural Science and Technology,2009,(20):364-366.
[18]邹琦.植物生理学实验指导[M].北京:中国农业出版社,2000.
[19]SHELP B J,ZAREI A.Subcellular compartmentation of 4-aminobutyrate(GABA)metabolism in Arabidopsis:An update[J].Plant Signaling&Behavior,2017,12(5):e1322244.
[20]张蒙,王秀峰,魏珉,等.叶绿酸铁对硝酸盐胁迫下黄瓜幼苗渗透调节物质及抗氧化酶活性的影响[J].中国蔬菜,2014,(3):14-18.ZHANG M,WANG X F,WEI M,et al.Effect of chlorophyllin-iron on osmoregulation substance and activities of antioxidant enzymes of cucumber seedling under nitrate stress[J].China Vegetables,2014,(3):14-18.
[21]GIL R,BOSCAIU M,LULL C,et al.Are soluble carbohydrates ecologically relevant for salt tolerance in halophytes?[J].Functional Plant Biology,2013,40(9):805-818.
[22]JANMOHAMMADI M,ZOLLA L,RINALDUCCI S.Low temperature tolerance in plants:changes at the protein level[J].Phytochemistry,2015,117:76-89.
[23]刘杜玲,彭少兵,张博勇,等.超敏蛋白对低温胁迫下核桃生理特性的影响[J].果树学报,2017,34(6):698-705.LIU D L,PENG S B,ZHANG B Y,et al.Effect of harpin on physiological characteristics of walnut under low temperature stress[J].Journal of Fruit Science,2017,34(6):698-705
[24]李桂荣,朱自果,马俊伟,等.低温胁迫对几种无核葡萄品种抗寒生理指标的影响[J].西北林学院学报,2015,30(5):75-78,291.LI G R,ZHU Z G,MA J W,et al.Effect of low temperature stress on physiological indices of seedless grape species[J].Journal of Northwest Forestry University,2015,30(5):75-78,291.
[25]张悦,周琳,张会慧,等.低温胁迫对蓝莓枝条呼吸作用及生理生化指标的影响[J].经济林研究,2016,34(2):12-18.ZHANG Y,ZHOU L,ZHANG H H,et al.Effects of low temperature stress on respiration,physiological and biochemical parameters of blueberry branches[J].Nonwood Forest Research,2016,34(2):12-18.
[26]陈镇,李永强,陈文荣,等.温度对短柄樱桃花芽萌发及若干生理指标的影响[J].浙江师范大学学报(自然科学版),2010,33(2):210-215.CHEN Z,LI Y Q,CHEN W R,et al.Effects of temperatures on the flower bud germination and some physiological indexes of Cerasus pseudocerasus[J].Journal of Zhejiang Normal University(Nat.Sci),2010,33(2):210-215.
[27]欧欢,王绪春,王振磊,等.自然越冬过程中扁桃枝条生理特性变化与抗寒性评价[J].新疆农业科学,2017,54(10):1 785-1 795.OU H,WANG X C,WANG Z L,et al.Changes of physiological indexes and comprehensive evaluation of cold resistance of almond during natural overwintering[J].Xinjiang Agricultural Sciences,2017,54(10):1 785-1 795.
[28]SZABADOS L,SACOURE A.Proline:a multifunctional amino acid[J].Trends in Plant Science,2010,15(2):89-97.
[29]REJEB K B,ABDELLY C,SACOURE A.How reactive oxygen species and proline face stress together[J].Plant Physiology and Biochemistry,2014,80:278-284.
[30]FARNESE F S,MENEZES-SILVA P E,GUSMAN G S,et al.When bad guys become good ones:the key role of reactive oxygen species and nitric oxide in the plant responses to abiotic stress[J].Frontiers in Plant Science,2016,7:471.
[31]梁杨杨,吴晓蕾,李敬蕊,等.盐碱胁迫下GABA对番茄幼苗活性氧代谢及光合性能的影响[J].河北农业大学学报,2016,39(3):7-13.LIANG Y Y,WU X L,LI J R,et al.Effects of exogenous GABA on reactive oxygen species metabolism and photosynthesis characteristics of tomatoes under mixed NaCl and NaHCO3stress[J].Journal of Agricultural University of Hebei,2016,39(3):7-13.
[32]高洪波,郭世荣.外源γ-氨基丁酸对营养液低氧胁迫下网纹甜瓜幼苗抗氧化酶活性和活性氧含量的影响[J].植物生理与分子生物学学报,2004,(6):651-659.GAO H B,GUO S R.Effects of exogenousγ-aminobutyric acid on antioxidant enzyme activities and reactive oxygen species in muskmelon seedlings under hypoxia stress[J].Journal of Plant Physiology and Molecular Biology,2004,(6):651-659.
[33]田小磊,吴晓岚,李云,等.盐胁迫条件下γ-氨基丁酸对玉米幼苗SOD、POD及CAT活性的影响[J].实验生物学报,2005,(1):77-81.TIAN X L,WU X L,LI Y,et al.Effects ofγ-aminobutyric acid on activities of SOD,POD and CAT in maize seedlings under salt stress[J].Journal of Experimental Biology,2005,(1):77-81.
[34]夏庆平,高洪波,李敬蕊.γ-氨基丁酸(GABA)对低氧胁迫下甜瓜幼苗光合作用和叶绿素荧光参数的影响[J].应用生态学报,2011,22(4):999-1 006.XIA Q P,GAO H B,LI J R.Effects ofγ-aminobutyric acid on the photosynthesis and chlorophyll fluorescence parameters of muskmelon seedlings under hypoxia stress[J].Chinese Journal of Applied Ecology,2011,22(4):999-1 006.
[2]SHELP B J,BOWN A W,MCLEAN M D.Metabolism and functions of gamma-aminobutyric acid[J].Trends in Plant Science,1999,4(11):446-452.
[3]SHELP B J,BOZZO G G,TROBACHER C P,et al.Strategies and tools for studying the metabolism and function ofγ-aminobutyrate in plants.I.Pathway structure[J].Botany,2012,90(8):651-668
[4]VERGARA R,PARADA F,PREZ F J.Is GABA-shunt functional in endodormant grapevine buds under respiratory stress?[J].Plant Growth Regulation,2013,71(3):253-260.
[5]罗黄颖,高洪波,夏庆平,等.γ-氨基丁酸对盐胁迫下番茄活性氧代谢及叶绿素荧光参数的影响[J].中国农业科学,2011,44(4):753-761.LUO H Y,GAO H B,XIA Q P,et al.Effects of exogenous GABA on reactive oxygen species metabolism and chlorophyll fluorescence parameters in tomato under NaCl stress[J].Scientia Agricultura Sinica,2011,44(4):753-761.
[6]FAIT A,FROMM H,WALTER D,et al.Highway or byway:the metabolic role of the GABA shunt in plants[J].Trends in Plant Science,2008,13(1):14-19.
[7]TROSSAT C,RATHINASABAPATHI B,HANSON A D.Transgenically expressed betaine aldehyde dehydrogenase efficiently catalyzes oxidation of dimethylsulfoniopropionaldehyde and[omega]-aminoaldehydes[J].Plant Physiology,1997,113(4):1 457-1 461.
[8]LIU C L,ZHAO L,YU G H.The dominant glutamic acid metabolic flux to produceγ-amino butyric acid over proline in Nicotiana tabacum leaves under water stress relates to its significant role in antioxidant activity[J].Journal of Integrative Plant Biology,2011,53(8):608-618.
[9]MITTLER R,VANDERAUWERA S,GOLLERY M,et al.Reactive oxygen gene network of plants[J].Trends in Plant Science,2004,9(10):490-498.
[10]马金虎,邢国芳,杨小环,等.外源EBR和NO信号对低温胁迫下玉米种胚抗氧化系统和低温响应基因表达的影响[J].应用生态学报,2015,26(5):1 411-1 418.MA J H,XING G F,YANG X H,et al.Effects of exogenous EBR and NO signal on antioxidant system and low response gene expression under cold stress on maize embryo[J].Chinese Journal of Applied Ecology,2015,26(5):1 411-1 418.
[11]张帆,郁继华,颉建明,等.外源ALA和Spd对低温弱光下辣椒幼苗光合作用及抗氧化系统的影响[J].中国农业科学,2013,46(11):2 298-2 306.ZHANG F,YU J H,XIE J M,et al.Influence of exogenous ALA and spd on photosynthesis and antioxidant system of low temperature and poor light affected pepper seedlings[J].Scientia Agricultura Sinica,2013,46(11):2 298-2 306.
[12]向丽霞,胡立盼,胡晓辉,等.外源γ-氨基丁酸调控甜瓜叶绿体活性氧代谢应对短期盐碱胁迫[J].应用生态学报,2015,26(12):3 746-3 752.XIANG L X,HU L P,HU X H,et al.Response of reactive oxygen metabolism in melon chloroplasts to short-term salinity-alkalinity stress regulated by exogenousγ-aminobutyric acid[J].Chinese Journal of Applied Ecology,2015,26(12):3 746-3 752.
[13]王爱国,罗广华.植物的超氧物自由基与羟胺反应的定量关系[J].植物生理学通讯,1990,(6):55-57.WANG A G,LUO G H.Quantitative relation between the reaction of hydroxylamine and superoxide anion radicals in plants[J].Plant Physiology Communications,1990,(6):55-57.
[14]BAI T H,LI C Y,MA F W,et al.Responses of growth and antioxidant system to root-zone hypoxia stress in two Malus species[J].Plant and Soil,2010,327(1-2):95-105.
[15]SHAH K,KUMAR R G,VERMA S,et al.Effect of cadmium on lipid peroxidation,superoxide anion generation and activities of antioxidant enzymes in growing rice seedlings[J].Plant Science,2001,161(6):1 135-1 144.
[16]王爱国,罗广华,邵从本,等.大豆种子超氧物歧化酶的研究[J].植物生理学报,1983,(1):77-84.WANG A G,LUO G H,SHAO C B,et al.A study on the superoxide dismutase of soybean seeds[J].Journal of Plant Physiology,1983,(1):77-84.
[17]杨兰芳,庞静,彭小兰,等.紫外分光光度法测定植物过氧化氢酶活性[J].现代农业科技,2009,(20):364-366.YANG L F,PANG J,PENG X L,et al.Measurement of catalase activity in plants by ultraviolet spectrophotometry[J].Modern Agricultural Science and Technology,2009,(20):364-366.
[18]邹琦.植物生理学实验指导[M].北京:中国农业出版社,2000.
[19]SHELP B J,ZAREI A.Subcellular compartmentation of 4-aminobutyrate(GABA)metabolism in Arabidopsis:An update[J].Plant Signaling&Behavior,2017,12(5):e1322244.
[20]张蒙,王秀峰,魏珉,等.叶绿酸铁对硝酸盐胁迫下黄瓜幼苗渗透调节物质及抗氧化酶活性的影响[J].中国蔬菜,2014,(3):14-18.ZHANG M,WANG X F,WEI M,et al.Effect of chlorophyllin-iron on osmoregulation substance and activities of antioxidant enzymes of cucumber seedling under nitrate stress[J].China Vegetables,2014,(3):14-18.
[21]GIL R,BOSCAIU M,LULL C,et al.Are soluble carbohydrates ecologically relevant for salt tolerance in halophytes?[J].Functional Plant Biology,2013,40(9):805-818.
[22]JANMOHAMMADI M,ZOLLA L,RINALDUCCI S.Low temperature tolerance in plants:changes at the protein level[J].Phytochemistry,2015,117:76-89.
[23]刘杜玲,彭少兵,张博勇,等.超敏蛋白对低温胁迫下核桃生理特性的影响[J].果树学报,2017,34(6):698-705.LIU D L,PENG S B,ZHANG B Y,et al.Effect of harpin on physiological characteristics of walnut under low temperature stress[J].Journal of Fruit Science,2017,34(6):698-705
[24]李桂荣,朱自果,马俊伟,等.低温胁迫对几种无核葡萄品种抗寒生理指标的影响[J].西北林学院学报,2015,30(5):75-78,291.LI G R,ZHU Z G,MA J W,et al.Effect of low temperature stress on physiological indices of seedless grape species[J].Journal of Northwest Forestry University,2015,30(5):75-78,291.
[25]张悦,周琳,张会慧,等.低温胁迫对蓝莓枝条呼吸作用及生理生化指标的影响[J].经济林研究,2016,34(2):12-18.ZHANG Y,ZHOU L,ZHANG H H,et al.Effects of low temperature stress on respiration,physiological and biochemical parameters of blueberry branches[J].Nonwood Forest Research,2016,34(2):12-18.
[26]陈镇,李永强,陈文荣,等.温度对短柄樱桃花芽萌发及若干生理指标的影响[J].浙江师范大学学报(自然科学版),2010,33(2):210-215.CHEN Z,LI Y Q,CHEN W R,et al.Effects of temperatures on the flower bud germination and some physiological indexes of Cerasus pseudocerasus[J].Journal of Zhejiang Normal University(Nat.Sci),2010,33(2):210-215.
[27]欧欢,王绪春,王振磊,等.自然越冬过程中扁桃枝条生理特性变化与抗寒性评价[J].新疆农业科学,2017,54(10):1 785-1 795.OU H,WANG X C,WANG Z L,et al.Changes of physiological indexes and comprehensive evaluation of cold resistance of almond during natural overwintering[J].Xinjiang Agricultural Sciences,2017,54(10):1 785-1 795.
[28]SZABADOS L,SACOURE A.Proline:a multifunctional amino acid[J].Trends in Plant Science,2010,15(2):89-97.
[29]REJEB K B,ABDELLY C,SACOURE A.How reactive oxygen species and proline face stress together[J].Plant Physiology and Biochemistry,2014,80:278-284.
[30]FARNESE F S,MENEZES-SILVA P E,GUSMAN G S,et al.When bad guys become good ones:the key role of reactive oxygen species and nitric oxide in the plant responses to abiotic stress[J].Frontiers in Plant Science,2016,7:471.
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