低温胁迫下交替呼吸途径对当归幼苗叶绿素荧光活性的影响
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摘要
以岷当归幼苗为材料,通过温室盆栽试验研究了在低温(4℃和-7℃)胁迫下交替呼吸途径对当归幼苗叶片叶绿素含量和荧光特性的影响,探讨交替呼吸途径在当归响应低温逆境过程中的作用。结果表明,随着胁迫温度的降低和处理时间的延长,当归叶片的叶绿素含量、实际光化学效率[Y(Ⅱ)]、光合电子传递速率(ETR)及光化学淬灭系数(qL)逐渐降低,而非光化学猝灭系数(NPQ)升高;同时,低温胁迫也导致当归幼苗叶片交替呼吸容量显著升高;在低温胁迫下,经交替呼吸途径抑制剂[1 mmol·L~(-1)的水杨基氧肟酸(SHAM)]预处理的当归幼苗叶片叶绿素含量、Y(Ⅱ)、ETR及qL进一步显著下降,而NPQ进一步显著升高,且温度越低升降幅度越大。研究发现,低温胁迫对当归幼苗叶片叶绿素的合成以及PSⅡ的光化学性能产生了显著抑制,低温胁迫下交替呼吸途径对当归幼苗叶绿素合成以及PSⅡ的光化学性能具有一定的保护作用。
We studied the effects of alternate respiration pathway on chlorophyll content and fluorescence characteristics of Angelica sinensis seedlings by pot experiment in greenhouse to explore the role of alternate respiratory pathway in the response of A. sinensis under low temperature(4 ℃ and-7 ℃). It was found that, with the decrease of temperature and the increase of low temperature time, the chlorophyll content, actual photochemical efficiency [Y(Ⅱ)], photosynthetic electron transfer rate(ETR), and photochemical quenching coefficient(qL) of A. sinensis decreased, while the non-photochemical quenching coefficient(NPQ) was increased. Meanwhile, low temperature stress also resulted in a significant increase in the alternate respiratory capacity of the A. sinensis seedlings. Under low temperature stress, compared with the A. sinensis seedlings without chemical treatment, the chlorophyll content, Y(Ⅱ), ETR and qL of A. sinensis seedlings pretreated with the inhibitor of the alternate respiratory pathway [1 mmol·L~(-1) salicylhydroxamic acid(SHAM)] decreased further, but NPQ increased significantly, and the lower the temperature was, the greater the range of rise and fall was. These results indicate that low temperature stress significantly inhibited the chlorophyll synthesis and photochemical properties in the leaves of A. sinensis seedlings, and the alternate respiratory pathway has protective effects on PSⅡ photochemistry and chlorophyll bio-synthesis of A. sinensis seedlings under low temperature stress.
We studied the effects of alternate respiration pathway on chlorophyll content and fluorescence characteristics of Angelica sinensis seedlings by pot experiment in greenhouse to explore the role of alternate respiratory pathway in the response of A. sinensis under low temperature(4 ℃ and-7 ℃). It was found that, with the decrease of temperature and the increase of low temperature time, the chlorophyll content, actual photochemical efficiency [Y(Ⅱ)], photosynthetic electron transfer rate(ETR), and photochemical quenching coefficient(qL) of A. sinensis decreased, while the non-photochemical quenching coefficient(NPQ) was increased. Meanwhile, low temperature stress also resulted in a significant increase in the alternate respiratory capacity of the A. sinensis seedlings. Under low temperature stress, compared with the A. sinensis seedlings without chemical treatment, the chlorophyll content, Y(Ⅱ), ETR and qL of A. sinensis seedlings pretreated with the inhibitor of the alternate respiratory pathway [1 mmol·L~(-1) salicylhydroxamic acid(SHAM)] decreased further, but NPQ increased significantly, and the lower the temperature was, the greater the range of rise and fall was. These results indicate that low temperature stress significantly inhibited the chlorophyll synthesis and photochemical properties in the leaves of A. sinensis seedlings, and the alternate respiratory pathway has protective effects on PSⅡ photochemistry and chlorophyll bio-synthesis of A. sinensis seedlings under low temperature stress.
引文
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[8] 刘剑光,肖松华,关巧娟,等.盐胁迫对棉种萌发及幼苗生长的影响[J].江苏农业科学,2010,(5):124-125.LIU J G,XIAO S H,GUAN Q J,et al.Effects of salt stress on seed germination and seedling growth of cotton[J].Jiangsu Agricultural Science,2010,(5):124-125.
[9] 谢佳佳,芦丽娜,石岱龙,等.交替呼吸途径对CuCl2胁迫下菜豆叶片光系统 Ⅱ 的保护作用[J].植物研究,2018,(1):75-80.XIE J J,LU L N,SHI D L,et al.Protective effects of alternative respiratory pathway on photosystemⅡin Phaseolus vulgaris under CuCl2 stress[J].Bulletin of Botanical Research,2018,(1):75-80.
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[11] LEOPOLD A C.Respiratory transition during seed germination[J].Plant Physiology,1976,57(2):274-276.
[12] LIANG H G,LU C S.A comparative study of CN-resistant respiration in different cultures of tobacco callus[J].Plant Physiology,1984,75(3):876-878.
[13] MEEUSE B J D.Thermogenic respiration in aroids[J].Annual Review of Plant Physiology,2003,26(1):117-126.
[14] 马建忠,梁厚果.萌发绿豆子叶衰老期间的呼吸作用与线粒体功能的改变[J].植物学通报,1991,8:45-50.MA J Z,LIANG H G.Changfs of the respiration and mitochondrial functions during senescence of cotyledons of mung bean seedlings[J].Chinese Bulletin of Botany,1991,8:45-50.
[15] 张年辉,韦振泉,何军贤,等.小麦幼苗叶片抗氰呼吸对轻度水分胁迫的响应[J].西北植物学报,2001,21(1):21-25.ZHANG N H,WEI Z Q,HE J X,et al.Responses of the cynanide-resistant respiration to mild water stress in wheat leaves[J].Acta Botanica Boreali-Occidentalia Sinica,2001,21(1):21-25.
[16] DAY D A,WHELAN J,MILLLAR A H,et al.Regulation of the alternative oxidase in plants and fungi[J].Australian Journal of Plant Physiology,1995,22(3):497-509.
[17] FENG H Q,SUN K,LI M Q,et al.The expression,function and regulation of mitochondrial alternative oxidase under biotic stresses[J].Molecular Plant Pathology,2010,11(3):429-440.
[18] BARTOLI C G,GOMEZ F,GERGOFF G,et al.Up-regulation of the mitochondrial alternative oxidase pathway enhances photosynthetic electron transport under drought conditions.[J].Journal of Experimental Botany,2005,56(415):1 269-1 276.
[19] DINAKAR C,VISHWAKARMA A,RAGHAVENDRA A S,et al.Alternative oxidase pathway optimizes photosynthesis during osmotic and temperature stress by regulating cellular ROS,malate valve and antioxidative systems[J].Frontiers in Plant Science,2016,7:68.doi.org/10.3389/fpls.2016.00068.
[20] SMITH C A,MELINO V J,SWEETMAN C,et al.Manipulation of alternative oxidase can influence salt tolerance in Arabidopsis thaliana[J].Physiologia Plantarum,2010,137(4):459-472.
[21] MILLENAAR F F,LAMBERS H.The alternative oxidase:in vivo regulation and function[J].Plant Biology,2010,5(1):2-15.
[22] GUILLAUME T,EDOUARD B F,ALINE M,et al.Respiratory carbon fluxes in leaves[J].Current Opinion in Plant Biology,2012,15(3):308-314.
[23] 冯汉青,管东东,焦青松,等.光照下菜豆叶片抗氰呼吸与光合作用关系的分析[J].植物生态学报,2015,39(1):104-109.FENG H Q,GUAN D D,JIAO Q S,et al.Analysis of the relationship between cyanide-resistant respiration and photosynthesis under light in Phaseolus vulgaris leaves[J].Chinese Journal of Plant Ecology,2015,39(1):104-109.
[24] ZHANG L T,ZHANG Z S,GAO H Y,et al.Mitochondrial alternative oxidase pathway protects plants against photoinhibition by alleviating inhibition of the repair of photodamaged PSⅡ through preventing formation of reactive oxygen species in Rumex,K-1 leaves[J].Physiologia Plantarum,2011,143(4):396-407.
[25] 徐秀玉,孙山,金立桥,等.干旱胁迫下平邑甜茶叶片交替呼吸途径上调对光破坏的防御作用[J].植物生理学报,2015,(12):2 119-2 125.XU X Y,SUN S,JIN L Q,et al.Up-regulation of the mitochondrial alternative oxidase pathway enhances photoprotection in Malus hupehensis leaves under drought stress[J].Plant Physiology Journal,2015,(12):2 119-2 125.
[26] ZHANG L T,GAO H Y,ZHANG Z S,et al.Multiple effects of inhibition of mitochondrial alternative oxidase pathway on photosynthetic apparatus in Rumex,K-1 leaves[J].Biologia Plantarum,2012,56(2):365-368.
[27] FLEXAS J,ESCALONA J M,MEDRANO H.Water stress induces different levels of photosynthesis and electron transport rate regulation in grapevines [J].Plant Cell & Environment,1999,22(1):39-48.
[28] HAZRATI S,TAHMASEBI-SARVESTANI Z,MODARRES-SANAVY S A,et al.Effects of water stress and light intensity on chlorophyll fluorescence parameters and pigments of Aloe vera L [J].Plant Physiology & Biochemistry Ppb,2016,106:141-148.
[29] ARNON D I.Copper enzymes in isolated chloroplasts.Polyphenoloxidase in Beta vulgaris [J].Plant Physiology,1949,24(1):1-15.
[30] BINGHHAM I J,FARRAR J F.Activity and capacity of respiratory pathways in barley roots deprived of inorganic nutrients[J].Plant Physiology & Biochemistry,1989,27(6):847-854.
[31] 芦丽娜,谢佳佳,王庆文,等.NaCl胁迫下交替呼吸途径对叶绿素含量及其荧光特性的影响[J].西北植物学报,2017,37(6):1 175-1 181.LU L N,XIE J J,WANG Q W,et al.Effect of alternative respiratory pathway on chlorophyll content and chlorophyll fluorescence characteristics under NaCl stress[J].Acta Botanica Boreali-Occidentalia Sinica,2017,37(6):1 175-1 181.
[32] 王利军,马履一,王爽,等.水盐胁迫对沙枣幼苗叶绿素荧光参数和色素含量的影响[J].西北农业学报,2010,19(12):122-127.WANG L J,MA L Y,WANG S,et al.Effects of water and salt stress on chlorophyll fluorescence parameters and pigment contents of Elaeagnus angustifolia L.seedlings[J].Acta Agriculturae Boreali-Occidentalis Sinica,2010,19(12):122-127.
[33] BAKER N R,ROSENQVIST E.Applications of chlorophyll fluorescence can improve crop production strategies:an examination of future possibilities[J].Journal of Experimental Botany,2004,55(403):1 607-1 621.
[34] CALATAYUD A,ROCA D,MARTINEZ P F.Spatial-temporal variations in rose leaves under water stress conditions studied by chlorophyll fluorescence imaging [J].Plant Physiology and Biochemistry (Paris),2006,44(10):564-573.
[35] 杨扬,王晓燕,王江,等.物种多样性对植物生长与土壤镉污染修复的影响[J].环境科学学报,2016,36(6):2 103-2 113.YANG Y,WANG X Y,WANG J,et al.Effects of species diversity on plant growth and remediation of Cd contamination in soil[J].Acta Scientiae Circumstantiae,2016,36(6):2 103-2 113.
[36] ASHRAF M,HARRIS P J C.Photosynthesis under stressful environments:An overview [J].Photosynthetica,2013,51(2):163-190.
[37] ZHANG D W,YUAN S,XU F,et al.Light intensity affects chlorophyll synthesis during greening process by metabolite signal from mitochondrial alternative oxidase in Arabidopsis[J].Plant Cell and Environment,2016,39(1):12-25.
[38] ZHANG D W,FEI X U,ZHANG Z W,et al.Effects of light on cyanide-resistant respiration and alternative oxidase function in Arabidopsis seedlings[J].Plant Cell & Environment,2010,33(12):2 121-2 131.
[2] FANG L,XIAO X F,LIU C X,et al.Recent advance in studies on Angelica sinensis[J].Chinese Herbal Medicines,2012,4(1):12-25.
[3] 刘学周,康天兰.当归栽培新技术研究综述[J].甘肃农业科技,2016,(11):62-66.LIU X Z,KANG T L.Research summary of cultivation new technology of Angelica sinensis[J].Gansu Agricultural Science and Technology,2016,(11):62-66.
[4] 张强,王润元,邓振镛,等.中国西北干旱气候变化对农业与生态影响及对策[M].北京:气象出版社,2012.
[5] 武延安,蔺海明,赵贵宾,等.遮光对当归栽培的效应 [J].中药材,2008,31(3):334 -336.WU Y A,LIN H M,ZHAO G B,et al.Effect of shading on cultivation of Angelica sinensis[J].Journal of Chinese Medicinal Materials,2008,31(3):334 -336.
[6] KOPSELL D A,LEFSRUD M G,KOPSELL D E,et al.Air temperature affects biomass and carotenoid pigment accumulation in Kale and Spinach grown in a controlled environment [J].Hortscience A Publication of the American Society for Horticultural Science,2005,40(7):2 026 - 2 030.
[7] HIKOSAKA K,ISHIKAWA K,BORJIGIDAI A,et al.Temperature acclimation of photosynthesis:mechanisms involved in the changes in temperature dependence of photosynthetic rate [J].Journal of Experimental Botany,2006,57(2):291-302.
[8] 刘剑光,肖松华,关巧娟,等.盐胁迫对棉种萌发及幼苗生长的影响[J].江苏农业科学,2010,(5):124-125.LIU J G,XIAO S H,GUAN Q J,et al.Effects of salt stress on seed germination and seedling growth of cotton[J].Jiangsu Agricultural Science,2010,(5):124-125.
[9] 谢佳佳,芦丽娜,石岱龙,等.交替呼吸途径对CuCl2胁迫下菜豆叶片光系统 Ⅱ 的保护作用[J].植物研究,2018,(1):75-80.XIE J J,LU L N,SHI D L,et al.Protective effects of alternative respiratory pathway on photosystemⅡin Phaseolus vulgaris under CuCl2 stress[J].Bulletin of Botanical Research,2018,(1):75-80.
[10] VANLERBERGHE G C,MCINTOSH L.Alternative oxidase:from gene to function[J].Annual Review of Plant Physiology and Plant Molecular Biology,1997,48(1):703-734.
[11] LEOPOLD A C.Respiratory transition during seed germination[J].Plant Physiology,1976,57(2):274-276.
[12] LIANG H G,LU C S.A comparative study of CN-resistant respiration in different cultures of tobacco callus[J].Plant Physiology,1984,75(3):876-878.
[13] MEEUSE B J D.Thermogenic respiration in aroids[J].Annual Review of Plant Physiology,2003,26(1):117-126.
[14] 马建忠,梁厚果.萌发绿豆子叶衰老期间的呼吸作用与线粒体功能的改变[J].植物学通报,1991,8:45-50.MA J Z,LIANG H G.Changfs of the respiration and mitochondrial functions during senescence of cotyledons of mung bean seedlings[J].Chinese Bulletin of Botany,1991,8:45-50.
[15] 张年辉,韦振泉,何军贤,等.小麦幼苗叶片抗氰呼吸对轻度水分胁迫的响应[J].西北植物学报,2001,21(1):21-25.ZHANG N H,WEI Z Q,HE J X,et al.Responses of the cynanide-resistant respiration to mild water stress in wheat leaves[J].Acta Botanica Boreali-Occidentalia Sinica,2001,21(1):21-25.
[16] DAY D A,WHELAN J,MILLLAR A H,et al.Regulation of the alternative oxidase in plants and fungi[J].Australian Journal of Plant Physiology,1995,22(3):497-509.
[17] FENG H Q,SUN K,LI M Q,et al.The expression,function and regulation of mitochondrial alternative oxidase under biotic stresses[J].Molecular Plant Pathology,2010,11(3):429-440.
[18] BARTOLI C G,GOMEZ F,GERGOFF G,et al.Up-regulation of the mitochondrial alternative oxidase pathway enhances photosynthetic electron transport under drought conditions.[J].Journal of Experimental Botany,2005,56(415):1 269-1 276.
[19] DINAKAR C,VISHWAKARMA A,RAGHAVENDRA A S,et al.Alternative oxidase pathway optimizes photosynthesis during osmotic and temperature stress by regulating cellular ROS,malate valve and antioxidative systems[J].Frontiers in Plant Science,2016,7:68.doi.org/10.3389/fpls.2016.00068.
[20] SMITH C A,MELINO V J,SWEETMAN C,et al.Manipulation of alternative oxidase can influence salt tolerance in Arabidopsis thaliana[J].Physiologia Plantarum,2010,137(4):459-472.
[21] MILLENAAR F F,LAMBERS H.The alternative oxidase:in vivo regulation and function[J].Plant Biology,2010,5(1):2-15.
[22] GUILLAUME T,EDOUARD B F,ALINE M,et al.Respiratory carbon fluxes in leaves[J].Current Opinion in Plant Biology,2012,15(3):308-314.
[23] 冯汉青,管东东,焦青松,等.光照下菜豆叶片抗氰呼吸与光合作用关系的分析[J].植物生态学报,2015,39(1):104-109.FENG H Q,GUAN D D,JIAO Q S,et al.Analysis of the relationship between cyanide-resistant respiration and photosynthesis under light in Phaseolus vulgaris leaves[J].Chinese Journal of Plant Ecology,2015,39(1):104-109.
[24] ZHANG L T,ZHANG Z S,GAO H Y,et al.Mitochondrial alternative oxidase pathway protects plants against photoinhibition by alleviating inhibition of the repair of photodamaged PSⅡ through preventing formation of reactive oxygen species in Rumex,K-1 leaves[J].Physiologia Plantarum,2011,143(4):396-407.
[25] 徐秀玉,孙山,金立桥,等.干旱胁迫下平邑甜茶叶片交替呼吸途径上调对光破坏的防御作用[J].植物生理学报,2015,(12):2 119-2 125.XU X Y,SUN S,JIN L Q,et al.Up-regulation of the mitochondrial alternative oxidase pathway enhances photoprotection in Malus hupehensis leaves under drought stress[J].Plant Physiology Journal,2015,(12):2 119-2 125.
[26] ZHANG L T,GAO H Y,ZHANG Z S,et al.Multiple effects of inhibition of mitochondrial alternative oxidase pathway on photosynthetic apparatus in Rumex,K-1 leaves[J].Biologia Plantarum,2012,56(2):365-368.
[27] FLEXAS J,ESCALONA J M,MEDRANO H.Water stress induces different levels of photosynthesis and electron transport rate regulation in grapevines [J].Plant Cell & Environment,1999,22(1):39-48.
[28] HAZRATI S,TAHMASEBI-SARVESTANI Z,MODARRES-SANAVY S A,et al.Effects of water stress and light intensity on chlorophyll fluorescence parameters and pigments of Aloe vera L [J].Plant Physiology & Biochemistry Ppb,2016,106:141-148.
[29] ARNON D I.Copper enzymes in isolated chloroplasts.Polyphenoloxidase in Beta vulgaris [J].Plant Physiology,1949,24(1):1-15.
[30] BINGHHAM I J,FARRAR J F.Activity and capacity of respiratory pathways in barley roots deprived of inorganic nutrients[J].Plant Physiology & Biochemistry,1989,27(6):847-854.
[31] 芦丽娜,谢佳佳,王庆文,等.NaCl胁迫下交替呼吸途径对叶绿素含量及其荧光特性的影响[J].西北植物学报,2017,37(6):1 175-1 181.LU L N,XIE J J,WANG Q W,et al.Effect of alternative respiratory pathway on chlorophyll content and chlorophyll fluorescence characteristics under NaCl stress[J].Acta Botanica Boreali-Occidentalia Sinica,2017,37(6):1 175-1 181.
[32] 王利军,马履一,王爽,等.水盐胁迫对沙枣幼苗叶绿素荧光参数和色素含量的影响[J].西北农业学报,2010,19(12):122-127.WANG L J,MA L Y,WANG S,et al.Effects of water and salt stress on chlorophyll fluorescence parameters and pigment contents of Elaeagnus angustifolia L.seedlings[J].Acta Agriculturae Boreali-Occidentalis Sinica,2010,19(12):122-127.
[33] BAKER N R,ROSENQVIST E.Applications of chlorophyll fluorescence can improve crop production strategies:an examination of future possibilities[J].Journal of Experimental Botany,2004,55(403):1 607-1 621.
[34] CALATAYUD A,ROCA D,MARTINEZ P F.Spatial-temporal variations in rose leaves under water stress conditions studied by chlorophyll fluorescence imaging [J].Plant Physiology and Biochemistry (Paris),2006,44(10):564-573.
[35] 杨扬,王晓燕,王江,等.物种多样性对植物生长与土壤镉污染修复的影响[J].环境科学学报,2016,36(6):2 103-2 113.YANG Y,WANG X Y,WANG J,et al.Effects of species diversity on plant growth and remediation of Cd contamination in soil[J].Acta Scientiae Circumstantiae,2016,36(6):2 103-2 113.
[36] ASHRAF M,HARRIS P J C.Photosynthesis under stressful environments:An overview [J].Photosynthetica,2013,51(2):163-190.
[37] ZHANG D W,YUAN S,XU F,et al.Light intensity affects chlorophyll synthesis during greening process by metabolite signal from mitochondrial alternative oxidase in Arabidopsis[J].Plant Cell and Environment,2016,39(1):12-25.
[38] ZHANG D W,FEI X U,ZHANG Z W,et al.Effects of light on cyanide-resistant respiration and alternative oxidase function in Arabidopsis seedlings[J].Plant Cell & Environment,2010,33(12):2 121-2 131.