混合盐碱胁迫对燕麦幼苗矿质离子吸收和光合特性的影响
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摘要
为探讨燕麦新品种定莜6号在低、高浓度盐碱胁迫下的生理响应机制,采用盆栽试验研究了25 mmol·L-1和75 mmol·L-1混合盐碱胁迫对幼苗生长、矿质离子吸收和光合特性的影响。结果表明,25 mmol·L-1混合盐碱胁迫10 d并未引起燕麦幼苗干重的明显改变,但75 mmol·L-1混合盐碱胁迫显著降低了幼苗干重。25 mmol·L-1混合盐碱胁迫下,燕麦根系和地上部K+/Na+、Ca2+/Na+和Mg2+/Na+显著降低,根系选择吸收K+、Ca2+、Mg2+及由根系向地上部运输K+、Ca2+的能力明显增强,但由根系向地上部运输Mg2+的能力下降,从而使地上部K+/Na+、Ca2+/Na+高于根系,而Mg2+/Na+低于根系;75 mmol·L-1混合盐碱胁迫下的上述变化大于25 mmol·L-1。25 mmol·L-1混合盐碱胁迫使燕麦幼苗叶片总叶绿素含量下降,叶绿素a/b和类胡萝卜素含量提高,而净光合速率(Pn)、气孔导度(Gs)、胞间CO2浓度(Ci)、蒸腾速率(Tr)和气孔限制值(Ls)变化不大;与25 mmol·L-1混合盐碱胁迫相比,75 mmol·L-1混合盐碱胁迫下燕麦叶片总叶绿素和类胡萝卜素含量、Pn、Gs、Tr、Ls明显下降,Ci显著提高,而叶绿素a/b无明显差异。25 mmol·L-1混合盐碱胁迫下,燕麦幼苗叶片最大光化学效率(Fv/Fm)和非光化学猝灭系数(NPQ)明显下降,保护性热耗散(ΦNPQ)显著提高,而初始荧光(Fo)、实际光化学效率(ΦPSII)、光化学猝灭系数(q L)、非调节性能量耗散(ΦNO)、光系统I和光系统II之间激发能分配的不平衡性(β/α-1)和叶绿体Hill反应活性无显著变化;与25 mmol·L-1混合盐碱胁迫相比,75 mmol·L-1混合盐碱胁迫下Fo、ΦNO、β/α-1显著提高,Fv/Fm、ΦPSII、q L、NPQ、ΦNPQ和叶绿体Hill反应活性明显下降。上述结果表明调控矿质离子吸收和运输以保持地上部K+/Na+、Ca2+/Na+高于根系是燕麦适应盐碱的重要机制,高浓度盐碱胁迫造成PSⅡ反应中心受损是燕麦Pn降低的主要因素,而过剩光能耗散是保护光合机构的重要途径。
In order to reveal the physiological adaptation of a new variety of oat Dingyou 6 to saline-alkali stress,oat seedlings were exposed to 0,25,75 mmol·L-1of complex saline-alkali stress( molar ratio of Na Cl∶ Na2SO4∶Na HCO3∶ Na2CO3= 12∶ 8∶ 9∶ 1) in a pot experiment,and then the growth,mineral ions absorption and photosynthetic performance of oat seedlings were measured after the stress treatment for 10 days. The results showed that the dry weight of oat seedlings were not affected under 25 mmol·L-1complex saline-alkali stress,while the dry weight of oat seedlings were significantly decreased under 75 mmol·L-1complex saline-alkali stress for 10 days. Under 25 mmol·L-1complex saline-alkali stress,the K+/Na+,Ca2 +/Na+and Mg2 +/Na+in both roots and shoots of oat seedlings were decreased,and the contents of K+,Ca2 +and Mg2 +selective absorbed by roots from outside and the transport of K+and Ca2 +from roots to shoots were increased. Moreover,the Mg2 +transport was inhibited,resulting in higher K+/Na+andCa2 +/Na+but lower Mg2 +/Na+of shoots than that of roots. Compared to 25 mmol·L-1complex saline-alkali stress,the above changes further increased under 75 mmol·L-1complex saline-alkali stress significantly. Under 25 mmol·L-1complex saline-alkali stress,the content of chlorophyll( a + b) in leaves of oat seedlings was significantly decreased,while chlorophyll a/b ratio and carotenoid content were increased significantly,net photosynthetic rate( Pn),stomatal conductance( Gs),intercellular CO2concentration( Ci),transpiration rate( Tr) and stomatal limitation value( Ls) had no significant changes; Compared with 25 mmol ·L-1complex saline-alkali stress,75 mmol ·L-1complex saline-alkali stress significantly reduced the contents of chlorophyll( a + b) and carotenoid,Pn,Gs,Tr,and Ls,but with no significant difference in the chlorophyll a/b ratio. Under 25 mmol·L-1complex saline-alkali stress,the maximum fluorescence efficiency( Fv/Fm) and non-photochemical quenching coefficient( NPQ) in leaves of oat seedlings were significantly decreased,while regulated thermal energy dissipation( ΦNPQ) was increased significantly,the minimal fluorescence( F0),actual photochemical efficiency( ΦPSII),photochemical quenching coefficient( q L),non-regulatory energy dissipation( ΦNO),the deviation from full balance between PSI and PSII( β/α-1),and Hill reaction activity had no significant changes. Compared with 25 mmol ·L-1complex saline-alkali stress,75 mmol ·L-1complex saline-alkali stress significantly improved the F0,ΦNo,β/α-1,while decreased Fv/Fm,ΦPSII,q L,NPQ,ΦNPQand Hill reaction activity. Collectively,these results indicated that it was an important mechanism of oat seedlings adapt to the saline-alkali stress that the high values of K+/Na+and Ca2 +/Na+in shoots were maintained by regulating the uptake and transportation of mineral ions,the structure damage in PSII reaction center was main cause for decreasing of photosynthetic rate,dissipation of excessive light energy might be a critical approach for protecting photosynthetic structure in oat seedlings under high concentration of complex saline-alkali stress.
In order to reveal the physiological adaptation of a new variety of oat Dingyou 6 to saline-alkali stress,oat seedlings were exposed to 0,25,75 mmol·L-1of complex saline-alkali stress( molar ratio of Na Cl∶ Na2SO4∶Na HCO3∶ Na2CO3= 12∶ 8∶ 9∶ 1) in a pot experiment,and then the growth,mineral ions absorption and photosynthetic performance of oat seedlings were measured after the stress treatment for 10 days. The results showed that the dry weight of oat seedlings were not affected under 25 mmol·L-1complex saline-alkali stress,while the dry weight of oat seedlings were significantly decreased under 75 mmol·L-1complex saline-alkali stress for 10 days. Under 25 mmol·L-1complex saline-alkali stress,the K+/Na+,Ca2 +/Na+and Mg2 +/Na+in both roots and shoots of oat seedlings were decreased,and the contents of K+,Ca2 +and Mg2 +selective absorbed by roots from outside and the transport of K+and Ca2 +from roots to shoots were increased. Moreover,the Mg2 +transport was inhibited,resulting in higher K+/Na+andCa2 +/Na+but lower Mg2 +/Na+of shoots than that of roots. Compared to 25 mmol·L-1complex saline-alkali stress,the above changes further increased under 75 mmol·L-1complex saline-alkali stress significantly. Under 25 mmol·L-1complex saline-alkali stress,the content of chlorophyll( a + b) in leaves of oat seedlings was significantly decreased,while chlorophyll a/b ratio and carotenoid content were increased significantly,net photosynthetic rate( Pn),stomatal conductance( Gs),intercellular CO2concentration( Ci),transpiration rate( Tr) and stomatal limitation value( Ls) had no significant changes; Compared with 25 mmol ·L-1complex saline-alkali stress,75 mmol ·L-1complex saline-alkali stress significantly reduced the contents of chlorophyll( a + b) and carotenoid,Pn,Gs,Tr,and Ls,but with no significant difference in the chlorophyll a/b ratio. Under 25 mmol·L-1complex saline-alkali stress,the maximum fluorescence efficiency( Fv/Fm) and non-photochemical quenching coefficient( NPQ) in leaves of oat seedlings were significantly decreased,while regulated thermal energy dissipation( ΦNPQ) was increased significantly,the minimal fluorescence( F0),actual photochemical efficiency( ΦPSII),photochemical quenching coefficient( q L),non-regulatory energy dissipation( ΦNO),the deviation from full balance between PSI and PSII( β/α-1),and Hill reaction activity had no significant changes. Compared with 25 mmol ·L-1complex saline-alkali stress,75 mmol ·L-1complex saline-alkali stress significantly improved the F0,ΦNo,β/α-1,while decreased Fv/Fm,ΦPSII,q L,NPQ,ΦNPQand Hill reaction activity. Collectively,these results indicated that it was an important mechanism of oat seedlings adapt to the saline-alkali stress that the high values of K+/Na+and Ca2 +/Na+in shoots were maintained by regulating the uptake and transportation of mineral ions,the structure damage in PSII reaction center was main cause for decreasing of photosynthetic rate,dissipation of excessive light energy might be a critical approach for protecting photosynthetic structure in oat seedlings under high concentration of complex saline-alkali stress.
引文
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[5]薛延丰,刘兆普.不同浓度Na Cl和Na2CO3处理对菊芋幼苗光合及叶绿素荧光的影响[J].植物生态学报,2008,32(1):161-167.
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[7]闫永庆,王文杰,朱虹,等.混合盐碱胁迫对青山杨渗透调节物质及活性氧代谢的影响[J].应用生态学报,2009,20(9):2085-2091.
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[10]彭远英,颜红海,郭来春,等.燕麦属不同倍性种质资源抗旱性状评价及筛选[J].生态学报,2011,31(9):2478-2491.
[11]吴娜,卜洪震,曾昭海,等.灌溉定额对夏播裸燕麦产量和品质的影响[J].草业学报,2010,19(5):204-209.
[12]吴斌,张宗文.燕麦葡聚糖合酶基因As CSLH的克隆及特征分析[J].作物学报,2011,37(4):723-728.
[13]王波,张金才,宋凤斌,等.燕麦对盐碱胁迫的生理响应[J].水土保持学报,2007,21(3):86-89.
[14]王波,张金才,宋凤斌,等.盐碱胁迫对燕麦光合特性的影响[J].中国农学通报,2007,23(5):235-238.
[15]萨如拉,刘景辉,刘伟,等.盐碱胁迫对燕麦幼苗Na+、K+含量及产量的影响[J].西北农业学报,2014,23(3):50-54.
[16]Flowers T J,Yeo A R.Ion relations of salt tolerance[C]//Baker D A,Hall J L.Solute transport in plant cells and tissues.New York:John Wiley&Sons,1988:399-412.
[17]高俊凤.植物生理学实验指导[M].北京:高等教育出版社,2006:17-19.
[18]Schreiber U,Schliwa U,Bilger W.Continuous recording of photochemical and non-photochemical chlorophyll fluorescence quenching with a new type of modulation fluorometer[J].Photosynthesis Research,1986,10(1/2):51-62.
[19]Genty B,Briantais J M,Baker N R.The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence[J].Biochimica et Biophysica Acta-General Subjects,1989,990(1):87-92.
[20]Branun G,Malkin S.Regulation of the imbalance in light excitation between photosystemⅡand photosystemⅠby cations and by the energized state of the thylakoid membrance[J].Biochimica et Biophysica Acta,1990,1017(1):79-90.
[21]叶济宇,钱月琴.希尔反应的分光光度计法测定[C]//薛应龙,夏镇澳.植物生理学实验手册.上海:上海科学技术出版社,1985:92-107.
[22]Alshammary S F,Qian Y L,Wallner S J.Growth response of four turfgrass species to salinity[J].Agricultural Water Management,2004,66(2):97-111.
[23]王殿,袁芳,王宝山,等.能源植物杂交狼尾草对Na Cl胁迫的响应及其耐盐阈值[J].植物生态学报,2012,36(6):572-577.
[24]陆嘉惠,吕新,梁永超,等.新疆胀果甘草幼苗耐盐性及对Na Cl胁迫的离子响应[J].植物生态学报,2013,37(9):839-850.
[25]高奔,宋杰,刘金萍,等.盐胁迫对不同生境盐地碱蓬光合及离子积累的影响[J].植物生态学报,2010,34(6):671-677.
[26]Farquhar G D,Sharkey T D.Stomatal conductance and photosynthesis[J].Annual Review of Plant Physiology,1982,33(3):317-345.
[27]Foyer C H,Noctor G.Oxygen processing in photosynthesis:regulation and signaling[J].New Phytologist,2000,146(3):359-388.
[28]尹海龙,田长彦.氮调控对盐环境下甜菜功能叶光系统II荧光特性的影响[J].植物生态学报,2013,37(2):122-131.
[29]苏秀荣,王秀峰,杨凤娟,等.硝酸根胁迫对黄瓜幼苗叶片光合速率、PSⅡ光化学效率及光能分配的影响[J].应用生态学报,2007,18(7):1441-1446.
[30]Gilmore A M.Mechanistic aspects of xanthophyll cycle-dependent photoprotection in higher plant chloroplasts and leaves[J].Physiology Plant,1997,99(1):197-209.
[31]Horton P,Hague A.Studies on the induction of chlorophyll fluorescence in isolated barley protoplasts.IV.Resolution of nonphotochemical quenching[J].Biochimica et Biophysica Acta,1988,932(1):107-115.
[32]赵滢,艾军,王振兴,等.外源NO对Na Cl胁迫下山葡萄叶片叶绿素荧光和抗氧化酶活性的影响[J].核农学报,2013,27(6):867-872.
[2]范远,任长忠,李品芳,等.盐碱胁迫下燕麦生长及阳离子吸收特征[J].应用生态学报,2011,22(11):2875-2882.
[3]杨春武,李长有,尹红娟,等.小冰麦(Triticum aestivum-Agropyron intermedium)对盐胁迫和碱胁迫的生理响应[J].作物学报,2007,33(8):1255-1261.
[4]颜宏,赵伟,盛艳敏,等.碱胁迫对羊草和向日葵的影响[J].应用生态学报,2005,16(8):1497-1501.
[5]薛延丰,刘兆普.不同浓度Na Cl和Na2CO3处理对菊芋幼苗光合及叶绿素荧光的影响[J].植物生态学报,2008,32(1):161-167.
[6]Gong B,Zhang C,Li X,et al.Identification of Na Cl and Na HCO3stress responsive proteins in tomato roots using i TRAQ-based analysis[J].Biochemistry Biophysics and Reserach Communications,2014,446(1):417-22.
[7]闫永庆,王文杰,朱虹,等.混合盐碱胁迫对青山杨渗透调节物质及活性氧代谢的影响[J].应用生态学报,2009,20(9):2085-2091.
[8]闫永庆,刘兴亮,王,等.白刺对不同浓度混合盐碱胁迫的生理响应[J].植物生态学报,2010,34(10):1213-1219.
[9]林叶春,曾昭海,任长忠,等.局部根区灌溉对裸燕麦光合特征曲线及叶绿素荧光特性的影响[J].作物学报,2012,38(6):1062-1070.
[10]彭远英,颜红海,郭来春,等.燕麦属不同倍性种质资源抗旱性状评价及筛选[J].生态学报,2011,31(9):2478-2491.
[11]吴娜,卜洪震,曾昭海,等.灌溉定额对夏播裸燕麦产量和品质的影响[J].草业学报,2010,19(5):204-209.
[12]吴斌,张宗文.燕麦葡聚糖合酶基因As CSLH的克隆及特征分析[J].作物学报,2011,37(4):723-728.
[13]王波,张金才,宋凤斌,等.燕麦对盐碱胁迫的生理响应[J].水土保持学报,2007,21(3):86-89.
[14]王波,张金才,宋凤斌,等.盐碱胁迫对燕麦光合特性的影响[J].中国农学通报,2007,23(5):235-238.
[15]萨如拉,刘景辉,刘伟,等.盐碱胁迫对燕麦幼苗Na+、K+含量及产量的影响[J].西北农业学报,2014,23(3):50-54.
[16]Flowers T J,Yeo A R.Ion relations of salt tolerance[C]//Baker D A,Hall J L.Solute transport in plant cells and tissues.New York:John Wiley&Sons,1988:399-412.
[17]高俊凤.植物生理学实验指导[M].北京:高等教育出版社,2006:17-19.
[18]Schreiber U,Schliwa U,Bilger W.Continuous recording of photochemical and non-photochemical chlorophyll fluorescence quenching with a new type of modulation fluorometer[J].Photosynthesis Research,1986,10(1/2):51-62.
[19]Genty B,Briantais J M,Baker N R.The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence[J].Biochimica et Biophysica Acta-General Subjects,1989,990(1):87-92.
[20]Branun G,Malkin S.Regulation of the imbalance in light excitation between photosystemⅡand photosystemⅠby cations and by the energized state of the thylakoid membrance[J].Biochimica et Biophysica Acta,1990,1017(1):79-90.
[21]叶济宇,钱月琴.希尔反应的分光光度计法测定[C]//薛应龙,夏镇澳.植物生理学实验手册.上海:上海科学技术出版社,1985:92-107.
[22]Alshammary S F,Qian Y L,Wallner S J.Growth response of four turfgrass species to salinity[J].Agricultural Water Management,2004,66(2):97-111.
[23]王殿,袁芳,王宝山,等.能源植物杂交狼尾草对Na Cl胁迫的响应及其耐盐阈值[J].植物生态学报,2012,36(6):572-577.
[24]陆嘉惠,吕新,梁永超,等.新疆胀果甘草幼苗耐盐性及对Na Cl胁迫的离子响应[J].植物生态学报,2013,37(9):839-850.
[25]高奔,宋杰,刘金萍,等.盐胁迫对不同生境盐地碱蓬光合及离子积累的影响[J].植物生态学报,2010,34(6):671-677.
[26]Farquhar G D,Sharkey T D.Stomatal conductance and photosynthesis[J].Annual Review of Plant Physiology,1982,33(3):317-345.
[27]Foyer C H,Noctor G.Oxygen processing in photosynthesis:regulation and signaling[J].New Phytologist,2000,146(3):359-388.
[28]尹海龙,田长彦.氮调控对盐环境下甜菜功能叶光系统II荧光特性的影响[J].植物生态学报,2013,37(2):122-131.
[29]苏秀荣,王秀峰,杨凤娟,等.硝酸根胁迫对黄瓜幼苗叶片光合速率、PSⅡ光化学效率及光能分配的影响[J].应用生态学报,2007,18(7):1441-1446.
[30]Gilmore A M.Mechanistic aspects of xanthophyll cycle-dependent photoprotection in higher plant chloroplasts and leaves[J].Physiology Plant,1997,99(1):197-209.
[31]Horton P,Hague A.Studies on the induction of chlorophyll fluorescence in isolated barley protoplasts.IV.Resolution of nonphotochemical quenching[J].Biochimica et Biophysica Acta,1988,932(1):107-115.
[32]赵滢,艾军,王振兴,等.外源NO对Na Cl胁迫下山葡萄叶片叶绿素荧光和抗氧化酶活性的影响[J].核农学报,2013,27(6):867-872.