不同盐分水平对柽柳扦插苗根系生长及生理特性的影响
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摘要
柽柳(Tamarix chinensis)作为黄河三角洲湿地的优势种之一,对盐渍化环境具有较好的适应性。根系作为植物直接感受盐分变化的器官,其对盐分条件的响应特征对研究植物生存、湿地恢复、土壤改良等具有重要意义。以一年生柽柳插穗为研究材料,采用水培法,设置6个盐分水平:CK(空白对照)、0.4%、0.8%、1.2%、2.4%、3.6%进行试验,测定柽柳生根率、根寿命、根系生物量等生长指标及根中ATP合成酶和过氧化氢酶(CAT)活性及脱落酸(ABA)含量,研究柽柳根系生长及生理特性对不同盐分水平的响应。结果表明:(1)柽柳适合扦插繁殖的培养溶液含盐量低于0.8%;含盐量超过0.8%后,扦插生根率显著降低,根系寿命减短。(2)柽柳根长生长随含盐量升高呈下降趋势;主根数随含盐量升高先增加,至含盐量超过1.2%后逐渐减少。(3)柽柳可通过调整生物量的分配模式来适应盐环境,低盐时地上部生物量高于地下,高盐时根系生物量比例逐渐增加,但生物量仍低于地上部分。(4)ATP合成酶活性、CAT活性在含盐量低于0.8%时增加;含盐量超过0.8%时,活性降低;脱落酸(ABA)含量随含盐量增加先增加,含盐量超过1.2%时减少。
Tamarix chinensis is a dominant shrub in the Yellow River Delta region. Salt glands occur on the leaves of T.chinensis,which can effectively exclude salt from the plant. However,due to the effects of global climate change and human over-exploitation of seawater resources,soil salinization and degradation has occurred in this region. The growth of T.chinensis has gradually decreased under conditions of high soil osmotic pressure,poor aeration and moisture,low nutrient availability,and high soil salinity. However,the roots can directly sense soil salinity changes,which mean that the effect of differing salinity on the eco-physiological characteristics of roots of T. chinensis cuttings is very important for the ecological restoration of coastal wetlands. In the present study,we studied the effects of different salinities( 0.0,0.4%,0.8%,1.2%,2.4%,3. 6% NaCl solutions) on the eco-physiological characteristics of roots of 1-year old T. chinensis. To explore the response of T. chinensis root growth and eco-physiological characteristics to different salinity levels,the rooting rate of T.chinensis cuttings,root lifespan,root number,root length,aboveground biomass,root biomass,hydrogen peroxide enzyme( CAT) activity,adenosine triphosphate( ATP) synthase,and the content of abscisic acid were determined. The results revealed that( 1) the rooting rate was approximately 83% in the 0.8% Na Cl treatment,and the rooting rate and life of theroot decreased significantly when the salinity increased to greater than 0.8%;( 2) the root length gradually decreased with increasing salinity,while the root quantity initially increased and then decreased with increasing salinity once the salinity was greater than 1. 2%;( 3) examining the biomass allocation pattern of T. chinensis adjustment to acclimate to salinity conditions,the aboveground biomass was higher than root biomass at lower salinity and the proportion of root biomass increased gradually; however,was still less than the aboveground biomass; and( 4) the activity of ATP synthase and CAT increased when salinity was less than 0.8%,and the activities decreased when the salt content was greater than 0.8%. In addition,the content of abscisic acid increased with increasing salinity and then decreased when the salinity increased to greater than 1.2%.
Tamarix chinensis is a dominant shrub in the Yellow River Delta region. Salt glands occur on the leaves of T.chinensis,which can effectively exclude salt from the plant. However,due to the effects of global climate change and human over-exploitation of seawater resources,soil salinization and degradation has occurred in this region. The growth of T.chinensis has gradually decreased under conditions of high soil osmotic pressure,poor aeration and moisture,low nutrient availability,and high soil salinity. However,the roots can directly sense soil salinity changes,which mean that the effect of differing salinity on the eco-physiological characteristics of roots of T. chinensis cuttings is very important for the ecological restoration of coastal wetlands. In the present study,we studied the effects of different salinities( 0.0,0.4%,0.8%,1.2%,2.4%,3. 6% NaCl solutions) on the eco-physiological characteristics of roots of 1-year old T. chinensis. To explore the response of T. chinensis root growth and eco-physiological characteristics to different salinity levels,the rooting rate of T.chinensis cuttings,root lifespan,root number,root length,aboveground biomass,root biomass,hydrogen peroxide enzyme( CAT) activity,adenosine triphosphate( ATP) synthase,and the content of abscisic acid were determined. The results revealed that( 1) the rooting rate was approximately 83% in the 0.8% Na Cl treatment,and the rooting rate and life of theroot decreased significantly when the salinity increased to greater than 0.8%;( 2) the root length gradually decreased with increasing salinity,while the root quantity initially increased and then decreased with increasing salinity once the salinity was greater than 1. 2%;( 3) examining the biomass allocation pattern of T. chinensis adjustment to acclimate to salinity conditions,the aboveground biomass was higher than root biomass at lower salinity and the proportion of root biomass increased gradually; however,was still less than the aboveground biomass; and( 4) the activity of ATP synthase and CAT increased when salinity was less than 0.8%,and the activities decreased when the salt content was greater than 0.8%. In addition,the content of abscisic acid increased with increasing salinity and then decreased when the salinity increased to greater than 1.2%.
引文
[1]夏江宝,赵西梅,刘俊华,赵自国,刘庆,陈印平.黄河三角洲莱州湾湿地柽柳种群分布特征及其影响因素.生态学报,2016,36(15):4801-4808.
[2]Cao D,Shi F C,Ruan W B,Lu Z H,Chai M.Seasonal changes in and relationship between soil microbial and microfaunal communities in a Tamarix chinensis community in the Yellow River Delta.African Journal of Biotechnology,2011,10(80):18425-18432.
[3]Xia J B,Zhang S Y,Zhang G C,Xu J W,Liu J T,Li C R,Shao H B.Growth dynamics and soil water ecological characteristics of Tamarix Chinensis Lour.forests with two site types in coastal wetland of Bohai golf.Journal of Food Agriculture and Environment,2013,11(2):1492-1498.
[4]Sun L K,Liu W Q,Liu G X,Chen T,Zhang W,Wu X K,Zhang G S,Zhang Y H,Li L,Zhang B G,Zhang B L,Wang B,Yang R Q.Temporal and spatial variations in the stable carbon isotope composition and carbon and nitrogen contents in current-season twigs of Tamarix chinensis Lour.and their relationships to environmental factors in the Lanzhou Bay wetland in China.Ecological Engineering,2016,90:417-426.
[5]何秀平,王保栋,谢琳萍.柽柳对盐碱地生态环境的影响.海洋科学,2014,38(1):96-101.
[6]马玉蕾,王德,刘俊民,温小虎,高猛,邵宏波.黄河三角洲典型植被与地下水埋深和土壤盐分的关系.应用生态学报,2013,24(9):2423-2430.
[7]李峰,谢永宏,覃盈盈.盐胁迫条件下湿地植物的适应策略.生态学杂志,2009,28(2):314-321.
[8]Rich S M,Ludwig M,Pedersen O,Colmer T D.Aquatic adventitious roots of the wetland plant Meionectes brownii can photosynthesize:implications for root function during flooding.New Phytologist,2011,190(2):311-319.
[9]Pan Y,Xie Y H,Deng Z M,Tang Y,Pan D D.High water level impedes the adaptation of Polygonum hydropiper to deep burial:Responses of biomass allocation and root morphology.Scientific Reports,2014,4:5612.
[10]Jiang Z M,Chen Y X,Bao Y.Population genetic structure of Tamarix chinensis,in the Yellow River Delta,China.Plant Systematics and Evolution,2012,298(1):147-153.
[11]Zhu Z,Zhang L Y,Gao L X,Tang A Q,Zhao Y,Yang J.Local habitat condition rather than geographic distance determines the genetic structure of Tamarix chinensis populations in Yellow River Delta,China.Tree Genetics and Genomes,2016,12(1):1-9.
[12]薛苹苹,曹春辉,何兴东,张宁,邬畏.盐碱地绒毛白蜡与柽柳展叶期生理生化特性比较.南开大学学报:自然科学版,2009(4):18-23.
[13]朱金方,陆兆华,夏江宝,陈曦,张萌,刘京涛.盐旱交叉胁迫对柽柳幼苗渗透调节物质含量的影响.西北植物学报,2013,33(2):0357-0363.
[14]Liu J H,Xia J B,Fang Y M,Li T,Liu J T.Effects of salt-drought stress on growth and physiobiochemical characteristics of Tamarix chinensis seedlings.The Scientific World Journal,2014,2014(6):765840.
[15]Rong Q Q,Liu J T,Cai Y P,Lu Z H,Zhao Z Z,Yue W C,Xia J B.“Fertile island”effects of Tamarix chinensis,Lour.on soil N and P stoichiometry in the coastal wetland of Laizhou Bay,China.Journal of Soils and Sediments,2016,16(3):864-877.
[16]贺强,崔保山,赵欣胜,付华龄.水、盐梯度下黄河三角洲湿地植物种的生态位.应用生态学报,2008,19(5):969-975.
[17]凌敏,刘汝海,王艳,罗先香,周凤琴.黄河三角洲柽柳林场湿地土壤养分的空间异质性及其与植物群落分布的耦合关系.湿地科学,2010,8(1):92-97.
[18]朱金方,刘京涛,陆兆华,夏江宝,柳海宁,金悦.盐胁迫对中国柽柳幼苗生理特性的影响.生态学报,2015,35(15):5140-5146.
[19]刘克东,郑彩霞,郝建卿.甘蒙柽柳对Na Cl胁迫的生理响应.广东农业科学,2012,39(10):38-42.
[20]徐先英,张孝仁.柽柳耐盐性水培试验研究.甘肃林业科技,1993(4):1-3.
[21]Wilson J B.A review of evidence on the control of shoot:root ratio,in relation to models.Annals of Botany,1988,61(4):433-449.
[22]Lloret F,Casanovas C,Pe1uelas J.Seedling survival of Mediterranean shrubland species in relation to root:shoot ratio,seed size and water and nitrogen use.Functional Ecology,1999,13(2):210-216.
[23]Passioura J B.Root signals control leaf expansion in wheat seedlings growing in drying soil.Australian Journal of Plant Biology,1988,15(5):687-693.
[24]Gedroc J J,Mc Connaughay K D M,Coleman J S.Plasticity in root/shoot partitioning:optimal,ontogenetic,or both?Functional Ecology,1996,10(1):44-50.
[25]刘莹,盖钧镒,吕彗能.作物根系形态与非生物胁迫耐性关系的研究进展.植物遗传资源学报,2003,4(3):265-269.
[26]Mou P,Jones R H,Tan Z Q,Bao Z,Chen H M.Morphological and physiological plasticity of plant roots when nutrients are both spatially and temporally heterogeneous.Plant and Soil,2013,364(1/2):373-384.
[27]弋良朋,王祖伟.盐胁迫下3种滨海盐生植物的根系生长和分布.生态学报,2011,31(5):1195-1202.
[28]梁建生,庞佳音,陈云.渗透胁迫诱导的植物细胞中脱落酸的合成及其调控机制.植物生理学报,2001,37(5):447-451.
[29]Fujita Y,Fujita M,Satoh R,Maruyama K,Parvez M,Seki M,Hiratsu K,Ohme-Takagi M,Shinozaki K,Yamaguchi-Shinozaki K.AREB1 is a transcription activator of novel ABRE-dependent ABA signaling that enhances drought stress tolerance in Arabidopsis.The Plant Cell,2005,17(12):3470-3488.
[30]郝格格,孙忠富,张录强,杜克明.脱落酸在植物逆境胁迫研究中的进展.中国农学通报,2009,25(18):212-215.
[31]陈娟,潘开文,辜彬.逆境胁迫下植物体内脱落酸的生理功能和作用机制.植物生理学报,2006,42(6):1176-1182.
[32]闫艳华,姜国斌,侯和胜,金华,马金龙,王颖.杨树内源激素对Na Cl胁迫的响应.西北农业学报,2011,20(9):160-164.
[33]孙洪刚,陈益泰.沿海防护林四个树种根系分布对盐胁迫的响应.生态学杂志,2010,29(12):2365-2372.
[34]梁艳荣,胡晓红,张颍力,刘湘萍.植物过氧化物酶生理功能研究进展.内蒙古农业大学学报:自然科学版,2003,24(2):110-113.
[35]陈金峰,王宫南,程素满.过氧化氢酶在植物胁迫响应中的功能研究进展.西北植物学报,2008,28(1):188-193.
[36]李汝佳,李雪梅.水杨酸、脱落酸和过氧化氢对镉胁迫小麦幼苗光合及抗氧化酶活性的影响.生态学杂志,2007,26(12):2096-2099.
[37]刘小金,徐大平,杨曾奖,张宁南.脱落酸对檀香幼苗生长、光合及叶片抗氧化酶活性的影响.南京林业大学学报:自然科学版,2016,40(3):57-62.
[2]Cao D,Shi F C,Ruan W B,Lu Z H,Chai M.Seasonal changes in and relationship between soil microbial and microfaunal communities in a Tamarix chinensis community in the Yellow River Delta.African Journal of Biotechnology,2011,10(80):18425-18432.
[3]Xia J B,Zhang S Y,Zhang G C,Xu J W,Liu J T,Li C R,Shao H B.Growth dynamics and soil water ecological characteristics of Tamarix Chinensis Lour.forests with two site types in coastal wetland of Bohai golf.Journal of Food Agriculture and Environment,2013,11(2):1492-1498.
[4]Sun L K,Liu W Q,Liu G X,Chen T,Zhang W,Wu X K,Zhang G S,Zhang Y H,Li L,Zhang B G,Zhang B L,Wang B,Yang R Q.Temporal and spatial variations in the stable carbon isotope composition and carbon and nitrogen contents in current-season twigs of Tamarix chinensis Lour.and their relationships to environmental factors in the Lanzhou Bay wetland in China.Ecological Engineering,2016,90:417-426.
[5]何秀平,王保栋,谢琳萍.柽柳对盐碱地生态环境的影响.海洋科学,2014,38(1):96-101.
[6]马玉蕾,王德,刘俊民,温小虎,高猛,邵宏波.黄河三角洲典型植被与地下水埋深和土壤盐分的关系.应用生态学报,2013,24(9):2423-2430.
[7]李峰,谢永宏,覃盈盈.盐胁迫条件下湿地植物的适应策略.生态学杂志,2009,28(2):314-321.
[8]Rich S M,Ludwig M,Pedersen O,Colmer T D.Aquatic adventitious roots of the wetland plant Meionectes brownii can photosynthesize:implications for root function during flooding.New Phytologist,2011,190(2):311-319.
[9]Pan Y,Xie Y H,Deng Z M,Tang Y,Pan D D.High water level impedes the adaptation of Polygonum hydropiper to deep burial:Responses of biomass allocation and root morphology.Scientific Reports,2014,4:5612.
[10]Jiang Z M,Chen Y X,Bao Y.Population genetic structure of Tamarix chinensis,in the Yellow River Delta,China.Plant Systematics and Evolution,2012,298(1):147-153.
[11]Zhu Z,Zhang L Y,Gao L X,Tang A Q,Zhao Y,Yang J.Local habitat condition rather than geographic distance determines the genetic structure of Tamarix chinensis populations in Yellow River Delta,China.Tree Genetics and Genomes,2016,12(1):1-9.
[12]薛苹苹,曹春辉,何兴东,张宁,邬畏.盐碱地绒毛白蜡与柽柳展叶期生理生化特性比较.南开大学学报:自然科学版,2009(4):18-23.
[13]朱金方,陆兆华,夏江宝,陈曦,张萌,刘京涛.盐旱交叉胁迫对柽柳幼苗渗透调节物质含量的影响.西北植物学报,2013,33(2):0357-0363.
[14]Liu J H,Xia J B,Fang Y M,Li T,Liu J T.Effects of salt-drought stress on growth and physiobiochemical characteristics of Tamarix chinensis seedlings.The Scientific World Journal,2014,2014(6):765840.
[15]Rong Q Q,Liu J T,Cai Y P,Lu Z H,Zhao Z Z,Yue W C,Xia J B.“Fertile island”effects of Tamarix chinensis,Lour.on soil N and P stoichiometry in the coastal wetland of Laizhou Bay,China.Journal of Soils and Sediments,2016,16(3):864-877.
[16]贺强,崔保山,赵欣胜,付华龄.水、盐梯度下黄河三角洲湿地植物种的生态位.应用生态学报,2008,19(5):969-975.
[17]凌敏,刘汝海,王艳,罗先香,周凤琴.黄河三角洲柽柳林场湿地土壤养分的空间异质性及其与植物群落分布的耦合关系.湿地科学,2010,8(1):92-97.
[18]朱金方,刘京涛,陆兆华,夏江宝,柳海宁,金悦.盐胁迫对中国柽柳幼苗生理特性的影响.生态学报,2015,35(15):5140-5146.
[19]刘克东,郑彩霞,郝建卿.甘蒙柽柳对Na Cl胁迫的生理响应.广东农业科学,2012,39(10):38-42.
[20]徐先英,张孝仁.柽柳耐盐性水培试验研究.甘肃林业科技,1993(4):1-3.
[21]Wilson J B.A review of evidence on the control of shoot:root ratio,in relation to models.Annals of Botany,1988,61(4):433-449.
[22]Lloret F,Casanovas C,Pe1uelas J.Seedling survival of Mediterranean shrubland species in relation to root:shoot ratio,seed size and water and nitrogen use.Functional Ecology,1999,13(2):210-216.
[23]Passioura J B.Root signals control leaf expansion in wheat seedlings growing in drying soil.Australian Journal of Plant Biology,1988,15(5):687-693.
[24]Gedroc J J,Mc Connaughay K D M,Coleman J S.Plasticity in root/shoot partitioning:optimal,ontogenetic,or both?Functional Ecology,1996,10(1):44-50.
[25]刘莹,盖钧镒,吕彗能.作物根系形态与非生物胁迫耐性关系的研究进展.植物遗传资源学报,2003,4(3):265-269.
[26]Mou P,Jones R H,Tan Z Q,Bao Z,Chen H M.Morphological and physiological plasticity of plant roots when nutrients are both spatially and temporally heterogeneous.Plant and Soil,2013,364(1/2):373-384.
[27]弋良朋,王祖伟.盐胁迫下3种滨海盐生植物的根系生长和分布.生态学报,2011,31(5):1195-1202.
[28]梁建生,庞佳音,陈云.渗透胁迫诱导的植物细胞中脱落酸的合成及其调控机制.植物生理学报,2001,37(5):447-451.
[29]Fujita Y,Fujita M,Satoh R,Maruyama K,Parvez M,Seki M,Hiratsu K,Ohme-Takagi M,Shinozaki K,Yamaguchi-Shinozaki K.AREB1 is a transcription activator of novel ABRE-dependent ABA signaling that enhances drought stress tolerance in Arabidopsis.The Plant Cell,2005,17(12):3470-3488.
[30]郝格格,孙忠富,张录强,杜克明.脱落酸在植物逆境胁迫研究中的进展.中国农学通报,2009,25(18):212-215.
[31]陈娟,潘开文,辜彬.逆境胁迫下植物体内脱落酸的生理功能和作用机制.植物生理学报,2006,42(6):1176-1182.
[32]闫艳华,姜国斌,侯和胜,金华,马金龙,王颖.杨树内源激素对Na Cl胁迫的响应.西北农业学报,2011,20(9):160-164.
[33]孙洪刚,陈益泰.沿海防护林四个树种根系分布对盐胁迫的响应.生态学杂志,2010,29(12):2365-2372.
[34]梁艳荣,胡晓红,张颍力,刘湘萍.植物过氧化物酶生理功能研究进展.内蒙古农业大学学报:自然科学版,2003,24(2):110-113.
[35]陈金峰,王宫南,程素满.过氧化氢酶在植物胁迫响应中的功能研究进展.西北植物学报,2008,28(1):188-193.
[36]李汝佳,李雪梅.水杨酸、脱落酸和过氧化氢对镉胁迫小麦幼苗光合及抗氧化酶活性的影响.生态学杂志,2007,26(12):2096-2099.
[37]刘小金,徐大平,杨曾奖,张宁南.脱落酸对檀香幼苗生长、光合及叶片抗氧化酶活性的影响.南京林业大学学报:自然科学版,2016,40(3):57-62.