NaCl胁迫对西伯利亚白刺根系生长及K~+/Na~+平衡的影响
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摘要
【目的】探讨西伯利亚白刺根系生长特征及根系中K~+/Na~+平衡与NaCl胁迫的关系。【方法】以西伯利亚白刺水培幼苗为材料,研究0(对照),100,200,300,400 mmol/L NaCl处理对其植株生物量积累、分配,根系形态特征及根系中K~+、Na~+、Cl~-积累情况的影响。【结果】随着NaCl浓度的升高,西伯利亚白刺幼苗根、茎、叶、冠层生物量和根冠比均呈先升高后下降的趋势,其中最大值均出现在200 mmol/L NaCl处理下,而400 mmol/L NaCl处理明显降低了西伯利亚白刺幼苗根、茎、叶、冠层生物量。随着NaCl浓度的升高,西伯利亚白刺幼苗根长、平均直径、表面积及根尖数均呈先升高后下降的趋势,其中最大值均出现在200 mmol/L NaCl处理下,300 mmol/L NaCl处理下西伯利亚白刺根系平均直径和表面积较对照略有增加,但差异不显著,而400 mmol/L NaCl处理下根系生长受到显著抑制。不同浓度NaCl处理下,西伯利亚白刺幼苗根系对Na~+、Cl~-和K~+的吸收增加,但K~+/Na~+随着NaCl浓度的升高逐渐降低。【结论】低浓度(≤200 mmol/L) NaCl处理下,西伯利亚白刺可以通过增加根系生物量、总长度、表面积、根尖数促进根系生长,并通过增加根系离子含量提高根系渗透势差来适应盐胁迫;高盐(400 mmol/L NaCl)处理下则通过提高根系对K~+的选择性吸收来降低根系中K~+/Na~+,从而降低单盐毒害作用,提高其耐盐性。
【Objective】 This study investigated the role of salinity in root growth characteristics and K~+/Na~+ balance of Nitraria sibirica Pall. seedlings.【Method】 The plant biomass accumulation and distribution,root morphology characteristics as well as K~+,Na~+ and Cl~- accumulation in root of N. sibirica seedlings under different NaCl levels(0(CK),100,200,300 and 400 mmol/L) were investigated through hydroponic experiment in greenhouse.【Result】 The biomass accumulations in root,stem and leaf,and root/sheet ratio of N. sibirica increased first and then decreased with the increase of NaCl concentration,and the maximum values were at 200 mmol/L NaCl treatment.The 400 mmol/L NaCl treatment significantly reduced the biomass accumulations in root,stem and leaf.With the increase of NaCl concentration,total length,surface area,average diameter and root tip number of N.sibirica roots increased first and then decreased,and the maximum values were at 200 mmol/L NaCl treatment.Compared with the control,the surface area and average diameter of roots were slightly increased in the 300 mmol/L NaCl treatment without significant difference.The growth of N.sibirica roots were significantly inhibited by 400 mmol/L NaCl treatment. With the increase of NaCl concentration,Na~+,Cl~- and K~+ contents in roots of N. sibirica were increased compared with control,but K~+/Na~+ ratio decreased.【Conclusion】 Low concentration(≤200 mmol/L) of NaCl significantly enhanced root growth of N. sibirica possibly via increasing biomass accumulation,total length,active absorbing area,tip number of roots and adapted salt stress by increasing ion content and osmotic potential difference.High concentration(400 mmol/L) of NaCl significantly inhibited root absorption unit via decreasing biomass accumulation,total length and tip number of roots,and decreasing K~+/Na~+ in roots.Thus,high concentration can reduce toxic effect of single salt and maintain relative balance of ions.
【Objective】 This study investigated the role of salinity in root growth characteristics and K~+/Na~+ balance of Nitraria sibirica Pall. seedlings.【Method】 The plant biomass accumulation and distribution,root morphology characteristics as well as K~+,Na~+ and Cl~- accumulation in root of N. sibirica seedlings under different NaCl levels(0(CK),100,200,300 and 400 mmol/L) were investigated through hydroponic experiment in greenhouse.【Result】 The biomass accumulations in root,stem and leaf,and root/sheet ratio of N. sibirica increased first and then decreased with the increase of NaCl concentration,and the maximum values were at 200 mmol/L NaCl treatment.The 400 mmol/L NaCl treatment significantly reduced the biomass accumulations in root,stem and leaf.With the increase of NaCl concentration,total length,surface area,average diameter and root tip number of N.sibirica roots increased first and then decreased,and the maximum values were at 200 mmol/L NaCl treatment.Compared with the control,the surface area and average diameter of roots were slightly increased in the 300 mmol/L NaCl treatment without significant difference.The growth of N.sibirica roots were significantly inhibited by 400 mmol/L NaCl treatment. With the increase of NaCl concentration,Na~+,Cl~- and K~+ contents in roots of N. sibirica were increased compared with control,but K~+/Na~+ ratio decreased.【Conclusion】 Low concentration(≤200 mmol/L) of NaCl significantly enhanced root growth of N. sibirica possibly via increasing biomass accumulation,total length,active absorbing area,tip number of roots and adapted salt stress by increasing ion content and osmotic potential difference.High concentration(400 mmol/L) of NaCl significantly inhibited root absorption unit via decreasing biomass accumulation,total length and tip number of roots,and decreasing K~+/Na~+ in roots.Thus,high concentration can reduce toxic effect of single salt and maintain relative balance of ions.
引文
[1]Munns R,Tester M.Mechanisms of salinity tolerance[J].Annu Rev Plant Biol,2008,59:651-681.
[2]张建锋,张旭东,周金星,等.世界盐碱地资源及其改良利用的基本措施[J].水土保持研究,2005,12(6):28-30.Zhang J F,Zhang X D,Zhou J X,et al.Word resource of saline soil and main amelioration measures[J].Research of Soil and Water Conservation,2005,12(6):28-30.
[3]刘正祥,张华新,杨秀艳,等.NaCl胁迫下沙枣幼苗生长和阳离子吸收、运输与分配特性[J].生态学报,2014,34(2):326-336.Liu Z X,Zhang H X,Yang X Y,et al.Growth,and cationic absorption,transportation and allocation of Elaeagnus angustifolia seedlings under NaCl stress[J].Acta Ecologica Sinica,2014,34(2):326-336.
[4]刘静,王庆祥.NaCl和NaCO3胁迫对玉米幼苗根系的影响[J].杂粮作物,2010,30(1):19-21.Liu J,Wang Q X.Effects of NaCl and NaCO3stress on root system of maize seedlings[J].Rain Fed Crops,2010,30(1):19-21.
[5]Giehl R F H,Gruber B D,von Wirén N.It’s time to make changes:modulation of root system architecture by nutrient signals[J].Journal of Experimental Botany,2014,65(3):769-778.
[6]Kellermeier F,Armengaud P,Seditas T J,et al.Analysis of the root system architecture of Arabidopsis provides a quantitative readout of crosstalk between nutritional signals[J].Plant Cell,2014,26(4):1480-1496.
[7]Galvanampudia C S,Testerink C.Salt stress signals shape the plant root[J].Current Opinion in Plant Biology,2011,14(3):296-302.
[8]冯中涛,王殿,袁芳,等.真盐生植物盐地碱蓬根系边缘细胞在耐盐中的作用初探[J].植物生理学报,2011,47(10):976-982.Feng Z T,Wang D,Yuan F,et al.Preliminary study on the role of root border cells in salt tolerance of euhalophytte Suaeda salsa L[J].Plant Physiology Journal,2011,47(10):976-982.
[9]郭建荣,郑聪聪,李艳迪,等.NaCl处理对真盐生植物盐地碱蓬根系特征及活力的影响[J].植物生理学报,2017,53(1):63-70.Guo J R,Zheng C C,Li Y D,et al.Effects of NaCl treatment on root system characteristics and activity of the euhalophyte Suaeda salsa[J].Plant Physiology Journal,2017,53(1):63-70.
[10]赵可夫,冯立田.中国盐生植物资源[M].北京:科学出版社,2001:198-199.Zhao K F,Feng L T.Halophyte resources of China[M].Beijing:Science Press,2001:198-199.
[11]史滟滪,杨静慧,左凤月,等.盐胁迫对3种白刺种子萌发的影响及其耐盐性比较[J].河南农业科学,2014,43(9):124-128.Shi Y Y,Yang J H,Zuo F Y,et al.Effects of salt stress on gremination of three species of Nitraria and their salt tolerance analysis[J].Journal of Henan Agricultural Sciences,2014,43(9):124-128.
[12]武香,倪建伟,张华新,等.盐胁迫对3种白刺渗透调节物质的影响[J].东北林业大学学报,2012,40(1):44-47.Wu X,Ni J W,Zhang H X,et al.Effects of salt stress on osmotic adjustment substances in three species of Nitraria[J].Journal of Northeast Forestry University,2012,40(1):44-47.
[13]萨日娜,陈贵林.外源亚精胺对盐胁迫下白刺幼苗叶片抗氧化酶系统的影响[J].西北植物学报,2013,33(2):352-356.Sa R N,Chen G L.Effect of exogenous spermidine on antioxidant enzyme system in leaves of Nitraria sibirica Pall.seedlings under salt stress[J].Acta Botanica Boreali-Occidentalia Sinica,2013,33(2):352-356.
[14]李焕勇,唐晓倩,杨秀艳,等.NaCl处理对西伯利亚白刺幼苗中矿质元素含量的影响[J].植物生理学报,2017,53(12):2125-2136.Li H Y,Tang X Q,Yang X Y,et al.Effects of NaCl stress on mineral element contents in Nitraria sibirica seedlings[J].Plant Physiology Journal,2017,53(12):2125-2136.
[15]Li H Y,Tang X Q,Zhu J F,et al.De Novo transcriptome characterization,gene expression profiling and ionic responses of Nitraria sibirica Pall.under salt stress[J].Forests,2017,8(6):211-231.
[16]Song J,Shi G W,Gao B,et al.Waterlogging and salinity effects on two Suaeda salsa populations[J].Physiologia Plantarum,2011,141(4):343-351.
[17]Rejili M,Vadel A M,Guetet A,et al.Effect of NaCl on the growth and the ionic balance K+/Na+of two populations of Lotus creticus(L.)(Papilionaceae)[J].South African Journal of Botany,2007,73(4):623-631.
[18]Khayat P N,Jamaatiesomarin S,Zabihiemahmoodabad R,et al.Screening of salt tolerance canola cultivars(Brassica napus L.)using physiological markers[J].World Applied Sciences Journal,2010,10(7):817-820.
[19]刘友良,毛才良,汪良驹.植物耐盐性研究进展[J].植物生理学通讯,1987,38(4):87-90.Liu Y L,Mao C L,Wang L J.Recent progress in studies on salinity tolerance in plants[J].Plant Physiology Communications,1987,38(4):87-90.
[20]闫道良,余婷,徐菊芳,等.盐胁迫对海滨锦葵生长及Na+、K+离子积累的影响[J].生态环境学报,2013,22(1):105-109.Yan D L,Yu T,Xu J F,et al.Effect of salt stress on the growth and accumulation of Na+,K+in seedlings of Kosteletzkya virginica[J].Ecology and Environmental Sciences,2013,22(1):105-109.
[21]唐晓倩,刘广全.NaCl胁迫对圆柏幼苗生长和离子吸收及分配的影响[J].西北植物学报,2017,37(7):1372-1380.Tang X Q,Liu G Q.Effects of NaCl stress on the growth,ion absorption and distribution of Juniperus chinensis seedlings[J].Acta Botanica Boreali-Occidentalia Sinica,2017,37(7):1372-1380.
[22]王树凤,胡韵雪,孙海菁,等.盐胁迫对2种栎树苗期生长和根系生长发育的影响[J].生态学报,2014,34(4):1021-1029.Wang S F,Hu Y X,Sun H J,et al.Effects of salt stress on growth and root development of two oak seedlings[J].Acta Ecologica Sinica,2014,34(4):1021-1029.
[23]Ali A,Tucker T C,Thompson T L,et al.Effects of salinity and mixed ammonium and nitrate nutrition on the growth and nitrogen utilization of barley[J].Journal of Agronomy&Crop Science,2010,186(4):223-228.
[24]邢建宏,潘德灼,谭芳林,等.NaCl胁迫对秋茄根系渗透调节物质含量的影响[J].生态环境学报,2017,26(11):1865-1871.Xing J H,Pan D Z,Tan F L,et al.Effects of NaCl stress on the osmotic substance contents in Kandelia candel roots[J].Ecology and Environmental Sciences,2017,26(11):1865-1871.
[25]韩志平,郭世荣,郑瑞娜,等.盐胁迫对小型西瓜幼苗体内离子分布的影响[J].植物营养与肥料学报,2013,19(4):908-917.Han Z P,Guo S R,Zheng R N,et al.Effect of salinity on distribution of ions on mini-watermelon seedlings[J].Journal of Plant Nutrition and Fertilizer,2013,19(4):908-917.
[26]孙健.胡杨响应盐胁迫与离子平衡调控信号网络研究[D].北京:北京林业大学,2011.Sun J.Signaling network in the perception of salt stress and ionic homeostasis regulation in Populus euphratica[D].Beijing:Beijing Forestry University,2011.
[27]Horie T,Schroeder J I.Sodium transporters in plants.Diverse genes and physiological functions[J].Plant Physiology,2004,136(1):2457-2462.
[28]苏丹,殷小琳,董智,等.白榆无性系生长特性及离子分布对NaCl胁迫的响应[J].北京林业大学学报,2017,39(5):48-57.Su D,Yin X L,Dong Z,et al.Response in growth and ionic distribution of elm clones under NaCl stress[J].Journal of Beijing Forestry University,2017,39(5):48-57.
[29]张海燕.盐胁迫下盐地碱蓬体内无机离子含量分布特点的研究[J].西北植物学报,2002,22(1):129-135.Zhang H Y.A study on the characters of content of inorganic ions in salt-stressed Suaeda salsa[J].Acta Botanica BorealiOccidentalia Sinica,2002,22(1):129-135.
[30]杨成龙,段瑞军,李瑞梅,等.盐生植物海马齿耐盐的生理特性[J].生态学报,2010,30(17):4617-4627.Yang C L,Duan R J,Li R M,et al.The physiological characteristics of salt-tolerance in Sesuviumportulacastrum L[J].Acta Ecologica Sinica,2010,30(17):4617-4627.
[2]张建锋,张旭东,周金星,等.世界盐碱地资源及其改良利用的基本措施[J].水土保持研究,2005,12(6):28-30.Zhang J F,Zhang X D,Zhou J X,et al.Word resource of saline soil and main amelioration measures[J].Research of Soil and Water Conservation,2005,12(6):28-30.
[3]刘正祥,张华新,杨秀艳,等.NaCl胁迫下沙枣幼苗生长和阳离子吸收、运输与分配特性[J].生态学报,2014,34(2):326-336.Liu Z X,Zhang H X,Yang X Y,et al.Growth,and cationic absorption,transportation and allocation of Elaeagnus angustifolia seedlings under NaCl stress[J].Acta Ecologica Sinica,2014,34(2):326-336.
[4]刘静,王庆祥.NaCl和NaCO3胁迫对玉米幼苗根系的影响[J].杂粮作物,2010,30(1):19-21.Liu J,Wang Q X.Effects of NaCl and NaCO3stress on root system of maize seedlings[J].Rain Fed Crops,2010,30(1):19-21.
[5]Giehl R F H,Gruber B D,von Wirén N.It’s time to make changes:modulation of root system architecture by nutrient signals[J].Journal of Experimental Botany,2014,65(3):769-778.
[6]Kellermeier F,Armengaud P,Seditas T J,et al.Analysis of the root system architecture of Arabidopsis provides a quantitative readout of crosstalk between nutritional signals[J].Plant Cell,2014,26(4):1480-1496.
[7]Galvanampudia C S,Testerink C.Salt stress signals shape the plant root[J].Current Opinion in Plant Biology,2011,14(3):296-302.
[8]冯中涛,王殿,袁芳,等.真盐生植物盐地碱蓬根系边缘细胞在耐盐中的作用初探[J].植物生理学报,2011,47(10):976-982.Feng Z T,Wang D,Yuan F,et al.Preliminary study on the role of root border cells in salt tolerance of euhalophytte Suaeda salsa L[J].Plant Physiology Journal,2011,47(10):976-982.
[9]郭建荣,郑聪聪,李艳迪,等.NaCl处理对真盐生植物盐地碱蓬根系特征及活力的影响[J].植物生理学报,2017,53(1):63-70.Guo J R,Zheng C C,Li Y D,et al.Effects of NaCl treatment on root system characteristics and activity of the euhalophyte Suaeda salsa[J].Plant Physiology Journal,2017,53(1):63-70.
[10]赵可夫,冯立田.中国盐生植物资源[M].北京:科学出版社,2001:198-199.Zhao K F,Feng L T.Halophyte resources of China[M].Beijing:Science Press,2001:198-199.
[11]史滟滪,杨静慧,左凤月,等.盐胁迫对3种白刺种子萌发的影响及其耐盐性比较[J].河南农业科学,2014,43(9):124-128.Shi Y Y,Yang J H,Zuo F Y,et al.Effects of salt stress on gremination of three species of Nitraria and their salt tolerance analysis[J].Journal of Henan Agricultural Sciences,2014,43(9):124-128.
[12]武香,倪建伟,张华新,等.盐胁迫对3种白刺渗透调节物质的影响[J].东北林业大学学报,2012,40(1):44-47.Wu X,Ni J W,Zhang H X,et al.Effects of salt stress on osmotic adjustment substances in three species of Nitraria[J].Journal of Northeast Forestry University,2012,40(1):44-47.
[13]萨日娜,陈贵林.外源亚精胺对盐胁迫下白刺幼苗叶片抗氧化酶系统的影响[J].西北植物学报,2013,33(2):352-356.Sa R N,Chen G L.Effect of exogenous spermidine on antioxidant enzyme system in leaves of Nitraria sibirica Pall.seedlings under salt stress[J].Acta Botanica Boreali-Occidentalia Sinica,2013,33(2):352-356.
[14]李焕勇,唐晓倩,杨秀艳,等.NaCl处理对西伯利亚白刺幼苗中矿质元素含量的影响[J].植物生理学报,2017,53(12):2125-2136.Li H Y,Tang X Q,Yang X Y,et al.Effects of NaCl stress on mineral element contents in Nitraria sibirica seedlings[J].Plant Physiology Journal,2017,53(12):2125-2136.
[15]Li H Y,Tang X Q,Zhu J F,et al.De Novo transcriptome characterization,gene expression profiling and ionic responses of Nitraria sibirica Pall.under salt stress[J].Forests,2017,8(6):211-231.
[16]Song J,Shi G W,Gao B,et al.Waterlogging and salinity effects on two Suaeda salsa populations[J].Physiologia Plantarum,2011,141(4):343-351.
[17]Rejili M,Vadel A M,Guetet A,et al.Effect of NaCl on the growth and the ionic balance K+/Na+of two populations of Lotus creticus(L.)(Papilionaceae)[J].South African Journal of Botany,2007,73(4):623-631.
[18]Khayat P N,Jamaatiesomarin S,Zabihiemahmoodabad R,et al.Screening of salt tolerance canola cultivars(Brassica napus L.)using physiological markers[J].World Applied Sciences Journal,2010,10(7):817-820.
[19]刘友良,毛才良,汪良驹.植物耐盐性研究进展[J].植物生理学通讯,1987,38(4):87-90.Liu Y L,Mao C L,Wang L J.Recent progress in studies on salinity tolerance in plants[J].Plant Physiology Communications,1987,38(4):87-90.
[20]闫道良,余婷,徐菊芳,等.盐胁迫对海滨锦葵生长及Na+、K+离子积累的影响[J].生态环境学报,2013,22(1):105-109.Yan D L,Yu T,Xu J F,et al.Effect of salt stress on the growth and accumulation of Na+,K+in seedlings of Kosteletzkya virginica[J].Ecology and Environmental Sciences,2013,22(1):105-109.
[21]唐晓倩,刘广全.NaCl胁迫对圆柏幼苗生长和离子吸收及分配的影响[J].西北植物学报,2017,37(7):1372-1380.Tang X Q,Liu G Q.Effects of NaCl stress on the growth,ion absorption and distribution of Juniperus chinensis seedlings[J].Acta Botanica Boreali-Occidentalia Sinica,2017,37(7):1372-1380.
[22]王树凤,胡韵雪,孙海菁,等.盐胁迫对2种栎树苗期生长和根系生长发育的影响[J].生态学报,2014,34(4):1021-1029.Wang S F,Hu Y X,Sun H J,et al.Effects of salt stress on growth and root development of two oak seedlings[J].Acta Ecologica Sinica,2014,34(4):1021-1029.
[23]Ali A,Tucker T C,Thompson T L,et al.Effects of salinity and mixed ammonium and nitrate nutrition on the growth and nitrogen utilization of barley[J].Journal of Agronomy&Crop Science,2010,186(4):223-228.
[24]邢建宏,潘德灼,谭芳林,等.NaCl胁迫对秋茄根系渗透调节物质含量的影响[J].生态环境学报,2017,26(11):1865-1871.Xing J H,Pan D Z,Tan F L,et al.Effects of NaCl stress on the osmotic substance contents in Kandelia candel roots[J].Ecology and Environmental Sciences,2017,26(11):1865-1871.
[25]韩志平,郭世荣,郑瑞娜,等.盐胁迫对小型西瓜幼苗体内离子分布的影响[J].植物营养与肥料学报,2013,19(4):908-917.Han Z P,Guo S R,Zheng R N,et al.Effect of salinity on distribution of ions on mini-watermelon seedlings[J].Journal of Plant Nutrition and Fertilizer,2013,19(4):908-917.
[26]孙健.胡杨响应盐胁迫与离子平衡调控信号网络研究[D].北京:北京林业大学,2011.Sun J.Signaling network in the perception of salt stress and ionic homeostasis regulation in Populus euphratica[D].Beijing:Beijing Forestry University,2011.
[27]Horie T,Schroeder J I.Sodium transporters in plants.Diverse genes and physiological functions[J].Plant Physiology,2004,136(1):2457-2462.
[28]苏丹,殷小琳,董智,等.白榆无性系生长特性及离子分布对NaCl胁迫的响应[J].北京林业大学学报,2017,39(5):48-57.Su D,Yin X L,Dong Z,et al.Response in growth and ionic distribution of elm clones under NaCl stress[J].Journal of Beijing Forestry University,2017,39(5):48-57.
[29]张海燕.盐胁迫下盐地碱蓬体内无机离子含量分布特点的研究[J].西北植物学报,2002,22(1):129-135.Zhang H Y.A study on the characters of content of inorganic ions in salt-stressed Suaeda salsa[J].Acta Botanica BorealiOccidentalia Sinica,2002,22(1):129-135.
[30]杨成龙,段瑞军,李瑞梅,等.盐生植物海马齿耐盐的生理特性[J].生态学报,2010,30(17):4617-4627.Yang C L,Duan R J,Li R M,et al.The physiological characteristics of salt-tolerance in Sesuviumportulacastrum L[J].Acta Ecologica Sinica,2010,30(17):4617-4627.