外施褪黑素对盐胁迫下番茄幼苗光合功能的影响
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摘要
褪黑素是植物响应多种非生物胁迫的一种重要的保护因子。为了明确褪黑素对盐胁迫下番茄叶片光合机构功能的影响,以番茄品种‘辽园多丽’为实验材料,利用150mmol·L~(-1)NaCl水培方式做盐处理,并设150μmol·L~(-1)的褪黑素预处理组,研究外施褪黑素对盐胁迫下番茄叶片PSII的影响。结果表明:盐胁迫导致番茄叶片叶绿素含量降低、光合作用减弱,PSII活性的降低主要表现为PSII电子供体侧OEC的活性降低及PSII电子受体侧电子由QA向QB的传递过程受阻,特别是PSII电子供体侧对盐胁迫的敏感程度明显大于受体侧,PSII中D1和Psb O蛋白均发生降解,表明番茄叶片PSII反应供体侧和受体侧受到严重损伤; PSII反应中心对光能的捕获及利用能力降低,单位面积有活性反应中心的数量和吸收的光能用于电子传递的能量比例均有不同程度的下降;外施褪黑素使叶绿素含量和光合速率的降低幅度明显减小,褪黑素通过平衡PSII的供体侧、受体侧和反应中心的电子传递来缓解盐胁迫对番茄PSII造成的伤害,提高番茄的耐盐性。
Melatonin( MT) is an important factor protecting plants from a variety of abiotic stresses. To explore the effect of MT application on photosynthetic function of Solanum lycopersicum L.cv. We used 150 mmol · L-1NaCl as salt stress and 150 μmol · L-1MT as pretreatment to explore the effect of MT on photosystem II in tomato( Liaoyuandouli) leaves under salt stress.The results showed that salt stress reduced chlorophyll content,photosynthetic function. The decrease of PSII activity was mainly due to the reduction in the activity of oxygen-evolving complex( OEC) on donor side of PSII and the blockage of electron transfer from QAto QBon receptor side of PSII. The donor side of PSII was more sensitive to salt stress than the receptor side of PSII. In addition,the ability of PSII reaction center to harvest and utilize light energy,the concentration of reaction centers( expressed as per unit leaf area) and the ratio of excitation energy to absorbed energy were all decreased. However,MT application significantly reduced the magnitude of chlorophyll content reduction and photosynthetic rate. MT application can alleviate the damage on tomato caused by salt stress through balancing the electron transfer on donor side,receptor side and reaction center of PSII.
Melatonin( MT) is an important factor protecting plants from a variety of abiotic stresses. To explore the effect of MT application on photosynthetic function of Solanum lycopersicum L.cv. We used 150 mmol · L-1NaCl as salt stress and 150 μmol · L-1MT as pretreatment to explore the effect of MT on photosystem II in tomato( Liaoyuandouli) leaves under salt stress.The results showed that salt stress reduced chlorophyll content,photosynthetic function. The decrease of PSII activity was mainly due to the reduction in the activity of oxygen-evolving complex( OEC) on donor side of PSII and the blockage of electron transfer from QAto QBon receptor side of PSII. The donor side of PSII was more sensitive to salt stress than the receptor side of PSII. In addition,the ability of PSII reaction center to harvest and utilize light energy,the concentration of reaction centers( expressed as per unit leaf area) and the ratio of excitation energy to absorbed energy were all decreased. However,MT application significantly reduced the magnitude of chlorophyll content reduction and photosynthetic rate. MT application can alleviate the damage on tomato caused by salt stress through balancing the electron transfer on donor side,receptor side and reaction center of PSII.
引文
金立桥,车兴凯,张子山,等.2015.高温、强光下黄瓜叶片PSII供体侧和受体侧的伤害程度与快速荧光参数Wk变化的关系.植物生理学报,51(6):969-976.
李鹏民,高辉远,Strasser J.2005.快速叶绿素荧光诱导动力学分析在光合作用研究中的应用.植物生理与分子生物学学报,31(6):559-566.
孟凡娟,庞洪影,王建中,等.2011.Na Cl和Na2SO4胁迫下两种刺槐叶肉细胞叶绿体超微结构.生态学报,31(3):734-741.
许楠,龙静泓,张文石,等.2018.淹水胁迫对乡土风箱果和引种紫叶风箱果光合特性的影响.生态学杂志,37(6):1880-1888.
许大全.2002.光合作用效率.上海:上海科学技术出版社.
张会慧,张秀丽,李鑫,等.2012.Na Cl和Na2CO3胁迫对桑树幼苗生长和光合特性的影响.应用生态学报,23(3):625-631.
Afreen F,Zobayed SM,Kozai T.2006.Melatonin in Glycyrrhiza uralensis:Response of plant roots to spectral quality of light and UV-B radiation.Journal of Pineal Research,41:108-115.
Al-Taweel K,Iwaki T,Yabuta Y,et al.2007.A bacterial transgene for catalase protects translation of D1 protein during exposure of salt-stressed tobacco leaves to strong light.Plant Physiology,145:258-265.
Appenroth KJ,St9ckel J,Srivastava A,et al.2001.Multiple effect of chromate on the photosynthetic apparatus of Spirorlela polyrhiza probed by OJIP chlorophyll a fluorescence measurements.Environmental Pollution,115:49-64.
Arnao M,Hernandez-Ruiz J.2009.Protective effect of melatonin against chlorophyll degradation during the senescence of barley leaves.Journal of Pineal Research,46:58-63.
Arnao MB,Hernandez-Ruiz J.2015.Functions of melatonin in plants:A review.Journal of Pineal Research,59:133-150.
Chao L,Liang M,Cong C,et al.2016.Exogenous melatonin improved potassium content in Malus under different stress conditions.Journal of Pineal Research,61:218-229.
Chen S,Dai X,Qiang S,et al.2010.Effect of a nonhost-selective toxin from Alternaria alternata on chloroplast-electron transfer activity in Eupatorium adenophorum.Plant Pathology,54:671-677.
Dietz KJ,Pfannschmidt T.2011.Novel regulators in photosynthetic redox control of plant metabolism and gene expression.Plant Physiology,155:1477-1485.
Fan X,Zhang Z,Gao H,et al.2014.Photoinhibition-like damage to the photosynthetic apparatus in plant leaves induced by submergence treatment in the dark.PLo S ONE,9:e89067.
Galano A,Tan DX,Reiter RJ.2011.Melatonin as a natural ally against oxidative stress:A physicochemical examination.Journal of Pineal Research,51:1-16.
Haldimann P,Strasser RJ.1999.Effects of anaerobiosis as probed by the polyphasic chlorophyll a fluorescence rise kinetic in pea(Pisum sativum L.).Photosynthesis Research,62:67-83.
Han Y,Wang S,Zhao N,et al.2016.Exogenous abscisic acid alleviates cadmium toxicity by restricting Cd2+influx in Populus euphratica cells.Journal of Plant Growth Regulation,35:827-837.
Kang K,Lee K,Park S,et al.2010.Enhanced production of melatonin by ectopic overexpression of human serotonin N-acetyltransferase plays a role in cold resistance in transgenic rice seedlings.Journal of Pineal Research,49:176-182.
Kudoh H,Sonoike K.2002.Irreversible damage to photosystem I by chilling in the light:Cause of the degradation of chlorophyll after returning to normal growth temperature.Planta,215:541-548.
Li P,Cheng L,Gao H,et al.2009.Heterogeneous behavior of PSII in soybean(Glycine max)leaves with identical PSIIphotochemistry efficiency under different high temperature treatments.Journal of Plant Physiology,166:1607-1615.
Lu Y,Lei JQ,Zeng FJ,et al.2016.Effect of Na Cl-induced changes in growth,photosynthetic characteristics,water status and enzymatic antioxidant system of Calligonum caput-medusae seedlings.Photosynthetica,55:1-12.
Manchester LC,Coto-Montes A,Boga JA,et al.2015.Melatonin:An ancient molecule that makes oxygen metabolically tolerable.Journal of Pineal Research,59:403-419.
Misra AN,Srivastava A,Strasser RJ.2001.Utilization of fast chlorophyll a,fluorescence technique in assessing the salt/ion sensitivity of mung bean and Brassica seedlings.Journal of Plant Physiology,158:1173-1181.
Ort DR,Melis A.2011.Optimizing antenna size to maximize photosynthetic efficiency.Plant Physiology,155:79-85.
Ort DR,Merchant SS,Alric J,et al.2015.Redesigning photosynthesis to sustainably meet global food and bioenergy demand.Proceedings of the National Academy of Science of the United States of America,112:8529-8536.
Porra RJ.2002.The chequered history of the development and use of simultaneous equations for the accurate determination of chlorophylls a and b.Photosynthesis Research,73:149-156.
Posmyk MM,Kuran H,Marciniak K,et al.2008.Presowing seed treatment with melatonin protects red cabbage seedlings against toxic copper ion concentrations.Journal of Pineal Research,45:24-31.
Rana M,Mark T.2008.Mechanisms of salinity tolerance.Annual Review of Plant Biology,59:651-681.
Sharkey TD,Bernacchi CJ,Farquhar GD,et al.2007.Fitting photosynthetic carbon dioxide response curves for C3 leaves.Plant,Cell and Environment,30:1035-1040.
Shi H,Chan Z.2014.The cysteine2/histidine2-type transcription factor ZINC FINGER OF ARABIDOPSIS THALIANA6-activated C-REPEAT-BINDING FACTOR pathway is essential for melatonin-mediated freezing stress resistance in Arabidopsis.Journal of Pineal Research,57:185-191.
Shi Z,Jiang Y,Han X,et al.2017.Sl PIN1 regulates auxin efflux to affect flower abscission process.Scientific Reports,7:14919.
Srivastava A,GuisséB,Greppin H,et al.1997.Regulation of antenna structure and electron transport in photosystem II of Pisum sativum under elevated temperature probed by the fast polyphasic chlorophyll a,fluorescence transient:OKJIP.Biochimicaet Biophysica Acta:Bioenergetics,1320:95-106.
Strasser BJ,Strasser RJ.1995.Measuring fast fluorescence transients to address environmental questions:The JIP-test//Mathis P,ed.Photosynthesis:From Light to Biosphere.Dordrecht:Kluwer Academic Publishers:977-980.
Strasser BJ.1997.Donor side capacity of photosystem II probed by chlorophyll a fluorescence transients.Photosynthesis Research,52:147-155.
Strasser RJ,Tsimillimichael M,Srivastava A.2004.Analysis of the chlorophyll a fluorescence transient.Chlorophyll a Fluorescence:A Signature of Photosynthesis.Dordrecht:Springer:321-362.
Takahashi S,Murata N.2008.How do environmental stresses accelerate photoinhibition.Trends in Plant Science,13:178-182.
Tal O,Haim A,Harel O,et al.2011.Melatonin as an antioxidant and its semi-lunar rhythm in green macroalga Ulva sp.Journal of Experimental Botany,62:1903-1910.
Wang L,Zhao Y,Reiter RJ,et al.2014.Changes in melatonin levels in transgenic‘Micro-Tom’tomato overexpressing ovine AANAT and ovine HIOMT genes.Journal of Pineal Research,56:134-142.
Wang LY,Liu JL,Wang WX,et al.2016.Exogenous melatonin improves growth and photosynthetic capacity of cucumber under salinity-induced stress.Photosynthetica,54:19-27.
Wang P,Yin L,Liang D,et al.2012.Delayed senescence of apple leaves by exogenous melatonin treatment:Toward regulating the ascorbate-glutathione cycle.Journal of Pineal Research,53:11-20.
Yin ZP,Ren J,Zhou LJ,et al.2017.Water deficit mechanisms in perennial shrubs Cerasus humilis leaves revealed by physiological and proteomic analyses.Proteome Science,15:9.
Zhan GM,Li RJ,Hu ZY,et al.2014.Cosuppression of RBCS3B in Arabidopsis leads to severe photoinhibition caused by ROS accumulation.Plant Cell Reports,33:1091.
Zhang D,Ren L,Chen GQ,et al.2015a.ROS-induced oxidative stress and apoptosis-like event directly affect the cell viability of cryopreserved embryogenic callus in Agapanthus praecox.Plant Cell Reports,34:1499.
Zhang HH,Zhong HX,Sui X,et al.2016.Adaptive changes in chlorophyll content and photosynthetic features to low light in Physocarpus amurensis Maxim and Physocarpus opulifolius“Diabolo”.Peer J,4:e2125
Zhang N,Sun Q,Zhang H,et al.2015b.Roles of melatonin in abiotic stress resistance in plants.Journal of Experimental Botany,66:647-656.
Zhang N,Zhao B,Zhang HJ,et al.2013.Melatonin promotes water-stress tolerance,lateral root formation,and seed germination in cucumber(Cucumis sativus L.).Journal of Pineal Research,54:15-23.
Zhou X,Zhao H,Cao K,et al.2016.Beneficial roles of melatonin on redox regulation of photosynthetic electron transport and synthesis of D1 protein in tomato seedlings under salt stress.Frontiers in Plant Science,7:1823.
Zhu JK.2002.Salt and drought stress signal transduction in plants.Annual Review of Plant Biology,53:247-273.
Zushi K,Matsuzoe N.2017.Using of chlorophyll a fluorescence OJIP transients for sensing salt stress in the leaves and fruits of tomato.Scientia Horticulturae,219:216-221.
李鹏民,高辉远,Strasser J.2005.快速叶绿素荧光诱导动力学分析在光合作用研究中的应用.植物生理与分子生物学学报,31(6):559-566.
孟凡娟,庞洪影,王建中,等.2011.Na Cl和Na2SO4胁迫下两种刺槐叶肉细胞叶绿体超微结构.生态学报,31(3):734-741.
许楠,龙静泓,张文石,等.2018.淹水胁迫对乡土风箱果和引种紫叶风箱果光合特性的影响.生态学杂志,37(6):1880-1888.
许大全.2002.光合作用效率.上海:上海科学技术出版社.
张会慧,张秀丽,李鑫,等.2012.Na Cl和Na2CO3胁迫对桑树幼苗生长和光合特性的影响.应用生态学报,23(3):625-631.
Afreen F,Zobayed SM,Kozai T.2006.Melatonin in Glycyrrhiza uralensis:Response of plant roots to spectral quality of light and UV-B radiation.Journal of Pineal Research,41:108-115.
Al-Taweel K,Iwaki T,Yabuta Y,et al.2007.A bacterial transgene for catalase protects translation of D1 protein during exposure of salt-stressed tobacco leaves to strong light.Plant Physiology,145:258-265.
Appenroth KJ,St9ckel J,Srivastava A,et al.2001.Multiple effect of chromate on the photosynthetic apparatus of Spirorlela polyrhiza probed by OJIP chlorophyll a fluorescence measurements.Environmental Pollution,115:49-64.
Arnao M,Hernandez-Ruiz J.2009.Protective effect of melatonin against chlorophyll degradation during the senescence of barley leaves.Journal of Pineal Research,46:58-63.
Arnao MB,Hernandez-Ruiz J.2015.Functions of melatonin in plants:A review.Journal of Pineal Research,59:133-150.
Chao L,Liang M,Cong C,et al.2016.Exogenous melatonin improved potassium content in Malus under different stress conditions.Journal of Pineal Research,61:218-229.
Chen S,Dai X,Qiang S,et al.2010.Effect of a nonhost-selective toxin from Alternaria alternata on chloroplast-electron transfer activity in Eupatorium adenophorum.Plant Pathology,54:671-677.
Dietz KJ,Pfannschmidt T.2011.Novel regulators in photosynthetic redox control of plant metabolism and gene expression.Plant Physiology,155:1477-1485.
Fan X,Zhang Z,Gao H,et al.2014.Photoinhibition-like damage to the photosynthetic apparatus in plant leaves induced by submergence treatment in the dark.PLo S ONE,9:e89067.
Galano A,Tan DX,Reiter RJ.2011.Melatonin as a natural ally against oxidative stress:A physicochemical examination.Journal of Pineal Research,51:1-16.
Haldimann P,Strasser RJ.1999.Effects of anaerobiosis as probed by the polyphasic chlorophyll a fluorescence rise kinetic in pea(Pisum sativum L.).Photosynthesis Research,62:67-83.
Han Y,Wang S,Zhao N,et al.2016.Exogenous abscisic acid alleviates cadmium toxicity by restricting Cd2+influx in Populus euphratica cells.Journal of Plant Growth Regulation,35:827-837.
Kang K,Lee K,Park S,et al.2010.Enhanced production of melatonin by ectopic overexpression of human serotonin N-acetyltransferase plays a role in cold resistance in transgenic rice seedlings.Journal of Pineal Research,49:176-182.
Kudoh H,Sonoike K.2002.Irreversible damage to photosystem I by chilling in the light:Cause of the degradation of chlorophyll after returning to normal growth temperature.Planta,215:541-548.
Li P,Cheng L,Gao H,et al.2009.Heterogeneous behavior of PSII in soybean(Glycine max)leaves with identical PSIIphotochemistry efficiency under different high temperature treatments.Journal of Plant Physiology,166:1607-1615.
Lu Y,Lei JQ,Zeng FJ,et al.2016.Effect of Na Cl-induced changes in growth,photosynthetic characteristics,water status and enzymatic antioxidant system of Calligonum caput-medusae seedlings.Photosynthetica,55:1-12.
Manchester LC,Coto-Montes A,Boga JA,et al.2015.Melatonin:An ancient molecule that makes oxygen metabolically tolerable.Journal of Pineal Research,59:403-419.
Misra AN,Srivastava A,Strasser RJ.2001.Utilization of fast chlorophyll a,fluorescence technique in assessing the salt/ion sensitivity of mung bean and Brassica seedlings.Journal of Plant Physiology,158:1173-1181.
Ort DR,Melis A.2011.Optimizing antenna size to maximize photosynthetic efficiency.Plant Physiology,155:79-85.
Ort DR,Merchant SS,Alric J,et al.2015.Redesigning photosynthesis to sustainably meet global food and bioenergy demand.Proceedings of the National Academy of Science of the United States of America,112:8529-8536.
Porra RJ.2002.The chequered history of the development and use of simultaneous equations for the accurate determination of chlorophylls a and b.Photosynthesis Research,73:149-156.
Posmyk MM,Kuran H,Marciniak K,et al.2008.Presowing seed treatment with melatonin protects red cabbage seedlings against toxic copper ion concentrations.Journal of Pineal Research,45:24-31.
Rana M,Mark T.2008.Mechanisms of salinity tolerance.Annual Review of Plant Biology,59:651-681.
Sharkey TD,Bernacchi CJ,Farquhar GD,et al.2007.Fitting photosynthetic carbon dioxide response curves for C3 leaves.Plant,Cell and Environment,30:1035-1040.
Shi H,Chan Z.2014.The cysteine2/histidine2-type transcription factor ZINC FINGER OF ARABIDOPSIS THALIANA6-activated C-REPEAT-BINDING FACTOR pathway is essential for melatonin-mediated freezing stress resistance in Arabidopsis.Journal of Pineal Research,57:185-191.
Shi Z,Jiang Y,Han X,et al.2017.Sl PIN1 regulates auxin efflux to affect flower abscission process.Scientific Reports,7:14919.
Srivastava A,GuisséB,Greppin H,et al.1997.Regulation of antenna structure and electron transport in photosystem II of Pisum sativum under elevated temperature probed by the fast polyphasic chlorophyll a,fluorescence transient:OKJIP.Biochimicaet Biophysica Acta:Bioenergetics,1320:95-106.
Strasser BJ,Strasser RJ.1995.Measuring fast fluorescence transients to address environmental questions:The JIP-test//Mathis P,ed.Photosynthesis:From Light to Biosphere.Dordrecht:Kluwer Academic Publishers:977-980.
Strasser BJ.1997.Donor side capacity of photosystem II probed by chlorophyll a fluorescence transients.Photosynthesis Research,52:147-155.
Strasser RJ,Tsimillimichael M,Srivastava A.2004.Analysis of the chlorophyll a fluorescence transient.Chlorophyll a Fluorescence:A Signature of Photosynthesis.Dordrecht:Springer:321-362.
Takahashi S,Murata N.2008.How do environmental stresses accelerate photoinhibition.Trends in Plant Science,13:178-182.
Tal O,Haim A,Harel O,et al.2011.Melatonin as an antioxidant and its semi-lunar rhythm in green macroalga Ulva sp.Journal of Experimental Botany,62:1903-1910.
Wang L,Zhao Y,Reiter RJ,et al.2014.Changes in melatonin levels in transgenic‘Micro-Tom’tomato overexpressing ovine AANAT and ovine HIOMT genes.Journal of Pineal Research,56:134-142.
Wang LY,Liu JL,Wang WX,et al.2016.Exogenous melatonin improves growth and photosynthetic capacity of cucumber under salinity-induced stress.Photosynthetica,54:19-27.
Wang P,Yin L,Liang D,et al.2012.Delayed senescence of apple leaves by exogenous melatonin treatment:Toward regulating the ascorbate-glutathione cycle.Journal of Pineal Research,53:11-20.
Yin ZP,Ren J,Zhou LJ,et al.2017.Water deficit mechanisms in perennial shrubs Cerasus humilis leaves revealed by physiological and proteomic analyses.Proteome Science,15:9.
Zhan GM,Li RJ,Hu ZY,et al.2014.Cosuppression of RBCS3B in Arabidopsis leads to severe photoinhibition caused by ROS accumulation.Plant Cell Reports,33:1091.
Zhang D,Ren L,Chen GQ,et al.2015a.ROS-induced oxidative stress and apoptosis-like event directly affect the cell viability of cryopreserved embryogenic callus in Agapanthus praecox.Plant Cell Reports,34:1499.
Zhang HH,Zhong HX,Sui X,et al.2016.Adaptive changes in chlorophyll content and photosynthetic features to low light in Physocarpus amurensis Maxim and Physocarpus opulifolius“Diabolo”.Peer J,4:e2125
Zhang N,Sun Q,Zhang H,et al.2015b.Roles of melatonin in abiotic stress resistance in plants.Journal of Experimental Botany,66:647-656.
Zhang N,Zhao B,Zhang HJ,et al.2013.Melatonin promotes water-stress tolerance,lateral root formation,and seed germination in cucumber(Cucumis sativus L.).Journal of Pineal Research,54:15-23.
Zhou X,Zhao H,Cao K,et al.2016.Beneficial roles of melatonin on redox regulation of photosynthetic electron transport and synthesis of D1 protein in tomato seedlings under salt stress.Frontiers in Plant Science,7:1823.
Zhu JK.2002.Salt and drought stress signal transduction in plants.Annual Review of Plant Biology,53:247-273.
Zushi K,Matsuzoe N.2017.Using of chlorophyll a fluorescence OJIP transients for sensing salt stress in the leaves and fruits of tomato.Scientia Horticulturae,219:216-221.