不同红光与远红光对盐胁迫下番茄叶绿素合成的影响
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摘要
以番茄"MoneyMaker"植株为试材,采用营养液水培方法,在盐胁迫下设置不同红光(R)与远红光(FR)处理,T_1(R∶FR=7.4)、T_2(R∶FR=1.2)、T_3(R∶FR=0.8),以正常营养液栽培(R∶FR=7.4)为对照,研究了在盐胁迫条件下不同红光与远红光比值(R∶FR)光环境对番茄生长指标、叶绿素及其前体物质含量和气体交换参数的影响,探讨低R∶FR光环境对盐胁迫下叶绿素合成途径的影响。结果表明:在盐胁迫下,番茄植株的干质量、鲜质量、叶绿素a(Chl a)含量、叶绿素b(Chl b)含量、净光合速率(Pn)、气孔导度(Gs)、蒸腾速率(Tr)和胞间CO_2浓度(Ci)均显著下降,叶绿素含量下降导致光合能力下降,进一步造成干物质积累减少;在盐胁迫下,降低番茄植株光环境中的R∶FR值,干物质量、气体交换参数和叶绿素含量均显著上升,且在T_3处理下显著高于T_2处理。盐胁迫下番茄叶片中的5-氨基乙酰丙酸(ALA)、胆色素原(PBG)和尿卟啉Ⅲ(UroⅢ)的含量显著升高,而原卟啉Ⅸ(ProtoⅨ)、镁原叶琳Ⅸ(Mg-protoⅨ)和原叶绿素酸酯(Pchl)显著降低。低R∶FR处理降低了ALA、PBG、UroⅢ的积累量,增加了ProtoⅨ、Mg-protoⅨ、Pchl的含量,同时降低了Chl a/b的值,T_2和T_3处理中的差异不明显。表明低R∶FR值光环境通过促进叶绿素合成缓解了番茄受到的盐胁迫,R∶FR值为0.8时效果最佳,盐胁迫下番茄叶绿素合成受阻的起始点在ProtoⅨ的合成过程,低R∶FR值能缓解叶绿素合成受阻碍的程度,R∶FR值为1.2和0.8对叶绿素合成的促进没有显著差异。
In this study,three groups different red light(R),far-red light(FR)were set under NaCl stress treatment respectively:T_1(R∶FR=7.4);T_2(R∶FR=1.2);T_3(R∶FR=0.8)was compared with normal nutrient solution culture(R∶FR=7.4),effect of different red and far-red ratios(R∶FR)on the growth paramaters,chlorophyll biosynthesis precursors content and gas exchange parameters of tomato under NaCl stress was studied by hydroponics using‘MoneyMaker',to explore the effect of low R∶FR value on chlorophyll biosynthesis pathway under NaCl stress.The results showed that,fresh weight,dry weight,contents of chlorophyll a(Chl a)and chlorophyll b(Chl b),net photosynthetic rate(Pn),stomatal conductance(Gs),transpiration rate(Tr)and intercellular CO_2 concentration(Ci)of tomato seedlings decreased significantly under conditions of salinity.The decrease of chlorophyll content resulted in the decrease of photosynthetic capacity,which further resulted in the decrease of dry mass accumulation of tomato.Subjected to a lower R∶FR value,tomato seedlings under conditions of salinity increased fresh weight,dry weight,chlorophyll content and gas exchange parameters significantly,which in T_3 treatment significantly higher than T_2 treatment5-aminolevulinic acid(ALA),porphobilinogen(PBG)and uroporphyrin Ⅲ(UroⅢ)contents of tomato leaves increased significantly under conditions of salinity,but protoporphyrinⅨ(ProtoⅨ),magnesium protoporphyrinⅨ(Mg-ProtoⅨ)and protochlorophyllide(Pchl)contents decreased significantly.The accumulation of ALA,PBG,UroⅢ decreased significantly when tomato leaves were subjected to a lower R∶FR value,but ProtoⅨ,Mg-protoⅨ,Pchl contents increased significantly,meanwhile the value of Chl a/b decreased significantly,the differences of T_2 and T_3treatment were not significant.These results suggested that the light environment with low R∶FR value alleviated the NaCl stress of tomato seedling by promoting chlorophyll biosynthesis,the effects were the best when the R∶FR value was 0.8.Under NaCl stress,the starting point blocked the chlorophyll synthesis was in ProtoⅨsynthesis process,while low R∶FR value alleviate the obstacles of chlorophyll biosynthesis.There was no difference on promoting chlorophyll biosynthesis between R∶FR value of 1.2 and 0.8.
In this study,three groups different red light(R),far-red light(FR)were set under NaCl stress treatment respectively:T_1(R∶FR=7.4);T_2(R∶FR=1.2);T_3(R∶FR=0.8)was compared with normal nutrient solution culture(R∶FR=7.4),effect of different red and far-red ratios(R∶FR)on the growth paramaters,chlorophyll biosynthesis precursors content and gas exchange parameters of tomato under NaCl stress was studied by hydroponics using‘MoneyMaker',to explore the effect of low R∶FR value on chlorophyll biosynthesis pathway under NaCl stress.The results showed that,fresh weight,dry weight,contents of chlorophyll a(Chl a)and chlorophyll b(Chl b),net photosynthetic rate(Pn),stomatal conductance(Gs),transpiration rate(Tr)and intercellular CO_2 concentration(Ci)of tomato seedlings decreased significantly under conditions of salinity.The decrease of chlorophyll content resulted in the decrease of photosynthetic capacity,which further resulted in the decrease of dry mass accumulation of tomato.Subjected to a lower R∶FR value,tomato seedlings under conditions of salinity increased fresh weight,dry weight,chlorophyll content and gas exchange parameters significantly,which in T_3 treatment significantly higher than T_2 treatment5-aminolevulinic acid(ALA),porphobilinogen(PBG)and uroporphyrin Ⅲ(UroⅢ)contents of tomato leaves increased significantly under conditions of salinity,but protoporphyrinⅨ(ProtoⅨ),magnesium protoporphyrinⅨ(Mg-ProtoⅨ)and protochlorophyllide(Pchl)contents decreased significantly.The accumulation of ALA,PBG,UroⅢ decreased significantly when tomato leaves were subjected to a lower R∶FR value,but ProtoⅨ,Mg-protoⅨ,Pchl contents increased significantly,meanwhile the value of Chl a/b decreased significantly,the differences of T_2 and T_3treatment were not significant.These results suggested that the light environment with low R∶FR value alleviated the NaCl stress of tomato seedling by promoting chlorophyll biosynthesis,the effects were the best when the R∶FR value was 0.8.Under NaCl stress,the starting point blocked the chlorophyll synthesis was in ProtoⅨsynthesis process,while low R∶FR value alleviate the obstacles of chlorophyll biosynthesis.There was no difference on promoting chlorophyll biosynthesis between R∶FR value of 1.2 and 0.8.
引文
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[8]郝树芹,刘世琦,张自坤,等.西葫芦银叶病发病叶片叶绿素代谢及其荧光特性[J].园艺学报,2009,36(6):879-884.
[9]孙锦,李娟,郭世荣,等.不同菠菜品种对海水胁迫的生理响应差异[J].西北植物学报,2008,28(4):737-744.
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[14]SHEERIN D J,MENON C,ZUR O S,et al.Light-activated phytochrome A and B interact with members of the SPA family to promote photomorphogenesis in Arabidopsis by reorganizing the COP1/SPA complex[J].The Plant Cell,2015,27(1):189-201.
[15]黄薪历,于捷,艾楷棋,等.红光和远红光对番茄生长发育的影响[J].西北农林科技大学学报(自然科学版),2018(3):111-118,127.
[16]张新平.盐胁迫对番茄幼苗生长发育的影响研究[J].安徽农业科学,2007,35(19):5713-5714.
[17]XIE X Z,XUE Y J,ZHOU J J,et al.Phytochromes regulate SA and JA signaling pathways in rice and are required for developmentally controlled resistance to Magnaporthe grisea[J].Molecular Plant,2011(4):688-696.
[18]XIE Y J,XU S,HAN B,et al.Evidence of Arabidopsis salt acclimation induced by up-regulation of HY1 and the regulatory role of RbohD-derived reactive oxygen species synthesis[J].The Plant Journal,2011,66(2):280-292.
[19]FRANKLIN K A,TOLEDO-ORTIZ G,PYOTT D E,et al.Interaction of light and temperature signalling[J].Journal of Experimental Botany,2014,65(11):2859-2871.
[20]BOCCALANDRO H E,RUGNONE M L,MORENO J E,et al.Phytochrome B enhances photosynthesis at the expense of water-use efficiency in Arabidopsis[J].Plant Physiology,2009,150(2):1083-1092.
[21]高俊凤.植物生理学实验指导[M].北京:高等教育出版社,2006:74-77.
[22]RICHARD A T.Biochemical spectroscopy[M].London:Adam Hilger Ltd,1975:327-330.
[23]BOUORAD I.Porphyrin synthesis[M]//DARON H H,CUNSALUNS I C.Methods in enzymology.New York:Academic Press,1962:885-895.
[24]HODUINS R,van HUYSTEE R B.Rapid simultaneous estimation of protoporphyrin and Mg-protophyrins in higher plants[J].Journal Plant Physiology,1986,125(3-4):311-323.
[25]LINCOLN T,EDUARDO Z.Plant physiology[M].4edition.Oxford:Sinauer Associates,2010:99.
[26]何磊,陆兆华,管博,等.盐碱胁迫对两种高粱种子萌发及幼苗生长的影响[J].西北植物学报,2012,32(2):362-369.
[27]COCKBURN W,WHITELAM G,BROAD A,et al.The participation of phytochrome in the signal transduction pathway of salt stress responses in Mesembryanthemum crystallinum L.[J].Journal of Experimental Botany,1996,47(5):647-653.
[28]INDORF M,CORDERO J,NEUHAUS G,et al.Salt tolerance(STO),a stress-related protein,has a major role in light signalling[J].The Plant Journal,2007,51(4):563-574.
[29]MCKENZIE J,WEISER C,BURKE M.Effects of red and far red light on the initiation of cold acclimation in Cornus stolonifera Michx[J].Plant Physiology,1974,53(6):783-789.
[30]SLOCOMBE S,WHITELAM G,COCKBURN W.Investigation of phosphoeno/pyruvate carboxylase(PEPCase)in Mesembryanthemum crystallinum L.in C3 and CAM photosynthetic states[J].Plant,Cell&Environment,1993,16(4):403-411.
[31]ESTK Z,BLANKENSHIP R E.Molecular mechanism of photosynthesis[J].Photosynthetica,2002,40(1):12.
[32]REINBOTHE S,REINBOTHE C,APEL K,et al.Evolution of chlorophyll biosynthesis:The challenge to survive photooxidation[J].Cell,1996,86:703-705.
[33]MATRINGE M,CAMADRO J M,LABBE P,et al.Protoporphyrinogen oxidase as a molecular target for diphenyl ether herbicides[J].Biochem J,1989,260:231-235.
[34]MOCK H P,KEETMAN U,KRUSE E,et al.Defense responses to tetrapyrrole-induced oxidative stress in transgenic plants with reduced uroporphyrinogen decarboxylase or coproporphyrinogen oxidase activity[J].Plant Physiol,1998,116:107-116.
[35]孙锦,贾永霞,郭世荣,等.海水胁迫对菠菜(Spinacia olerancea L.)叶绿体活性氧和叶绿素代谢的影响[J].生态学报,2009,29(8):4361-4370.
[36]王颖,郭世荣,束胜,等.外源亚精胺对盐胁迫下菠菜叶绿素合成前体含量的影响[J].西北植物学报,2015,35(10):2026-2034.
[37]陈新斌,孙锦,郭世荣,等.海水胁迫对菠菜叶绿素代谢的影响[J].西北植物学报,2012,32(9):1781-1787.
[38]吕明,刘海衡,毛虎德,等.芥菜型油菜黄化突变体叶片叶绿素合成代谢变化[J].西北植物学报,2010,311(11):2177-2183.
[39]张丽,徐志然,胡晓辉,等.叶面喷施亚精胺对盐碱胁迫下番茄幼苗生长及其叶绿素合成前体含量的影响[J].西北植物学报,2015,35(1):125-130.
[40]喻敏,胡承孝,王运华.低温条件下钼对冬小麦叶绿素合成前体的影响[J].中国农业科学,2006,39(4):702-708.
[41]BELYAEVA O B,LITVIN F F.Pathways of formation of pigment forms at the terminal photobiochemical stage of chlorophyll biosynthesis[J].Biochemistry,2009,74(13):1535-1544.
[42]REINBOTHE S,REINBOTHE C.The regulation of enzymes involved in chlorophyll biosynthesis[J].Eur J Biochem,1996,237:323-343.
[43]白伟岚,任建武,苏雪痕.八种植物耐阴性比较研究[J].北京林业大学学报,1999,21(3):46-51.
[44]杨清,艾沙江·买买提,刘国杰,等.DA-6对草莓叶片叶绿素合成前体物质及关键酶酶活性的影响[J].中国农业大学学报,2012,17(4):86-90.
[2] LAKSHMI A,RAMANJULU S,VEERANJANEYULU K,et al.Effect of NaCl on photosynthesis parameters in two cultivars of mulberry[J].Photosynthetica,1996,32(2):285-289.
[3] MISRA A N,SAHU S M,MISRA M,et al.Sodium chloride induced changes in leaf growth,and pigment and protein contents in two rice cultivars[J].Biologia Plantarum,1997,39(2):257-262.
[4] ZHU J K.Salt and drought stress signal transduction in plants[J].Annual Review Plant Biology,2002,53:247-273.
[5]史典义,刘忠香,金危危.植物叶绿素合成、分解代谢及信号调控[J].遗传,2009,31(7):698-704.
[6] BEALE S I.Green genes gleaned[J].Trends Plant Sci,2005(10):309-312.
[7] TANAKA A,TANAKA R.Chlorophyll metabolism[J].Current Opinion in Plant Biology,2006,9(3):248-255.
[8]郝树芹,刘世琦,张自坤,等.西葫芦银叶病发病叶片叶绿素代谢及其荧光特性[J].园艺学报,2009,36(6):879-884.
[9]孙锦,李娟,郭世荣,等.不同菠菜品种对海水胁迫的生理响应差异[J].西北植物学报,2008,28(4):737-744.
[10]SAIRAM R K,SRIVASTAVA G C.Changes in antioxidant activity in sub-cellular fractions of tolerant and susceptible wheat genotypes in response to long term salt stress[J].Plant Science,2002,162(6):897-904.
[11]CARVALHO R F,TAKAKI M,AZEVEDO R A.Plant pigments:The many faces of light perception[J].Acta Physiologiae Plantarum,2011,33(2):241-248.
[12]HOLMES M,SMITH H.The function of phytochrome in the natural environment I.Characterization of daylight for studies in photomorphogenesis and photoperiodism[J].Photochemistry and Photobiology,1977,25(6):533-538.
[13]POSSART A,XU T,PAIK I,et al.Characterization of phytochrome interacting factors from the moss physcomitrella patens illustrates conservation of phytochrome signaling modules in land plants[J].The Plant Cell,2017,29(2):310-330.
[14]SHEERIN D J,MENON C,ZUR O S,et al.Light-activated phytochrome A and B interact with members of the SPA family to promote photomorphogenesis in Arabidopsis by reorganizing the COP1/SPA complex[J].The Plant Cell,2015,27(1):189-201.
[15]黄薪历,于捷,艾楷棋,等.红光和远红光对番茄生长发育的影响[J].西北农林科技大学学报(自然科学版),2018(3):111-118,127.
[16]张新平.盐胁迫对番茄幼苗生长发育的影响研究[J].安徽农业科学,2007,35(19):5713-5714.
[17]XIE X Z,XUE Y J,ZHOU J J,et al.Phytochromes regulate SA and JA signaling pathways in rice and are required for developmentally controlled resistance to Magnaporthe grisea[J].Molecular Plant,2011(4):688-696.
[18]XIE Y J,XU S,HAN B,et al.Evidence of Arabidopsis salt acclimation induced by up-regulation of HY1 and the regulatory role of RbohD-derived reactive oxygen species synthesis[J].The Plant Journal,2011,66(2):280-292.
[19]FRANKLIN K A,TOLEDO-ORTIZ G,PYOTT D E,et al.Interaction of light and temperature signalling[J].Journal of Experimental Botany,2014,65(11):2859-2871.
[20]BOCCALANDRO H E,RUGNONE M L,MORENO J E,et al.Phytochrome B enhances photosynthesis at the expense of water-use efficiency in Arabidopsis[J].Plant Physiology,2009,150(2):1083-1092.
[21]高俊凤.植物生理学实验指导[M].北京:高等教育出版社,2006:74-77.
[22]RICHARD A T.Biochemical spectroscopy[M].London:Adam Hilger Ltd,1975:327-330.
[23]BOUORAD I.Porphyrin synthesis[M]//DARON H H,CUNSALUNS I C.Methods in enzymology.New York:Academic Press,1962:885-895.
[24]HODUINS R,van HUYSTEE R B.Rapid simultaneous estimation of protoporphyrin and Mg-protophyrins in higher plants[J].Journal Plant Physiology,1986,125(3-4):311-323.
[25]LINCOLN T,EDUARDO Z.Plant physiology[M].4edition.Oxford:Sinauer Associates,2010:99.
[26]何磊,陆兆华,管博,等.盐碱胁迫对两种高粱种子萌发及幼苗生长的影响[J].西北植物学报,2012,32(2):362-369.
[27]COCKBURN W,WHITELAM G,BROAD A,et al.The participation of phytochrome in the signal transduction pathway of salt stress responses in Mesembryanthemum crystallinum L.[J].Journal of Experimental Botany,1996,47(5):647-653.
[28]INDORF M,CORDERO J,NEUHAUS G,et al.Salt tolerance(STO),a stress-related protein,has a major role in light signalling[J].The Plant Journal,2007,51(4):563-574.
[29]MCKENZIE J,WEISER C,BURKE M.Effects of red and far red light on the initiation of cold acclimation in Cornus stolonifera Michx[J].Plant Physiology,1974,53(6):783-789.
[30]SLOCOMBE S,WHITELAM G,COCKBURN W.Investigation of phosphoeno/pyruvate carboxylase(PEPCase)in Mesembryanthemum crystallinum L.in C3 and CAM photosynthetic states[J].Plant,Cell&Environment,1993,16(4):403-411.
[31]ESTK Z,BLANKENSHIP R E.Molecular mechanism of photosynthesis[J].Photosynthetica,2002,40(1):12.
[32]REINBOTHE S,REINBOTHE C,APEL K,et al.Evolution of chlorophyll biosynthesis:The challenge to survive photooxidation[J].Cell,1996,86:703-705.
[33]MATRINGE M,CAMADRO J M,LABBE P,et al.Protoporphyrinogen oxidase as a molecular target for diphenyl ether herbicides[J].Biochem J,1989,260:231-235.
[34]MOCK H P,KEETMAN U,KRUSE E,et al.Defense responses to tetrapyrrole-induced oxidative stress in transgenic plants with reduced uroporphyrinogen decarboxylase or coproporphyrinogen oxidase activity[J].Plant Physiol,1998,116:107-116.
[35]孙锦,贾永霞,郭世荣,等.海水胁迫对菠菜(Spinacia olerancea L.)叶绿体活性氧和叶绿素代谢的影响[J].生态学报,2009,29(8):4361-4370.
[36]王颖,郭世荣,束胜,等.外源亚精胺对盐胁迫下菠菜叶绿素合成前体含量的影响[J].西北植物学报,2015,35(10):2026-2034.
[37]陈新斌,孙锦,郭世荣,等.海水胁迫对菠菜叶绿素代谢的影响[J].西北植物学报,2012,32(9):1781-1787.
[38]吕明,刘海衡,毛虎德,等.芥菜型油菜黄化突变体叶片叶绿素合成代谢变化[J].西北植物学报,2010,311(11):2177-2183.
[39]张丽,徐志然,胡晓辉,等.叶面喷施亚精胺对盐碱胁迫下番茄幼苗生长及其叶绿素合成前体含量的影响[J].西北植物学报,2015,35(1):125-130.
[40]喻敏,胡承孝,王运华.低温条件下钼对冬小麦叶绿素合成前体的影响[J].中国农业科学,2006,39(4):702-708.
[41]BELYAEVA O B,LITVIN F F.Pathways of formation of pigment forms at the terminal photobiochemical stage of chlorophyll biosynthesis[J].Biochemistry,2009,74(13):1535-1544.
[42]REINBOTHE S,REINBOTHE C.The regulation of enzymes involved in chlorophyll biosynthesis[J].Eur J Biochem,1996,237:323-343.
[43]白伟岚,任建武,苏雪痕.八种植物耐阴性比较研究[J].北京林业大学学报,1999,21(3):46-51.
[44]杨清,艾沙江·买买提,刘国杰,等.DA-6对草莓叶片叶绿素合成前体物质及关键酶酶活性的影响[J].中国农业大学学报,2012,17(4):86-90.