铝胁迫对西藏野生垂穗披碱草种子萌发及幼苗生长的影响
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摘要
采用水培的方法,以采自西藏高寒区域不同地区的野生垂穗披碱草种子为材料,初步探讨了酸性环境下(pH4.5)不同浓度的Al 3+(0、0.5、1.0和1.5mmol·L-1)胁迫对垂穗披碱草种子萌发及幼苗生长的影响,以期为酸铝土壤的改良提供优良的种质资源。研究结果显示,低浓度Al 3+(0.5、1.0mmol·L-1)对3种垂穗披碱草种子的发芽率无显著影响(P>0.05),但Al 3+浓度为1.5mmol·L-1时,3种垂穗披碱草种子的发芽率均被显著抑制(P<0.05)。同时,3种野生牧草的根长、株高、根重及茎叶重在Al 3+胁迫下受到了不同程度的抑制,且根系被抑制的程度大于茎叶。此外,植物幼苗中丙二醛和可溶性糖作为受胁迫程度的重要指标,其含量在3种垂穗披碱草幼苗中均随Al 3+浓度的升高呈现出上升的趋势。结果还表明,1.5mmol·L-1 Al 3+处理时,那曲、申扎、浪卡子垂穗披碱草根系中铝含量分别是地上部的12.2、15.3、17.5倍。铝毒和磷缺失往往是酸性土壤中同时存在的问题,这严重影响了植物生长,研究表明Al 3+抑制了垂穗披碱草对磷的吸收及转运。通过隶属函数法对3种垂穗披碱草的抗铝特性评估结果显示,采自申扎县的垂穗披碱草对Al 3+的耐性较高,可驯化育种,为酸性土壤的改良提供优质的种质资源。
In general,Elymus nutans grows well in hostile environments at high altitudes.Aluminum(Al)stress is one of the serious factors limiting plant productivity in acidic soils.However,studies of the influence of aluminum on this plant are still lacking.The seeds of wild E.nutans from three different areas in Tibet(Naqu,Shenza and Langkazi Counties)were used as materials in this study.The major experimental method involved hydroponic culture with four concentrations of Al 3+(0,0.5,1.0 and 1.5 mmol·L-1)under acidic conditions(pH 4.5).In order to provide robust germplasm resources for the improvement of acid-aluminum soil,the effects of Al 3+stress on the seed germination and seedling growth were initially explored.Our investigations indicated that there were no significant differences(P>0.05)in the germination rate of the three samples of wild E.nutans seeds at lower concentrations of Al 3+stress(0.5 and 1.0 mmol·L-1)compared to the control,but that results were adverse at higher concentrations(1.5 mmol·L-1).The results showed that root length,plant height,root weight and shoots' weight in the three samples decreased with increasing Al 3+concentration.In addition,the MDA and soluble sugar contents of seedlings substantially increased with Al 3+application.Data showed that Al was poorly translocated from the roots to aerial parts of the three seedling samples.In the treatment groups,the Al content of roots increased 12.2,15.3 and 17.5 times more than that of the aerial parts.Al toxicity and phosphorus(P)deficiency often co-exist in acidic soils and limit plant growth and crop production.In this study,we observed that Al stress induced a reduction in P accumulation in the plants.Using eight indicators for seedlings under Al stress,we performed a comprehensive evaluation of Al tolerance of the three E.nutans samples and showed that the Shenza County sample was highly tolerant to aluminum and so could be used for domestication and breeding to provide good germplasm resources for the improvement of acid-aluminum soils.
In general,Elymus nutans grows well in hostile environments at high altitudes.Aluminum(Al)stress is one of the serious factors limiting plant productivity in acidic soils.However,studies of the influence of aluminum on this plant are still lacking.The seeds of wild E.nutans from three different areas in Tibet(Naqu,Shenza and Langkazi Counties)were used as materials in this study.The major experimental method involved hydroponic culture with four concentrations of Al 3+(0,0.5,1.0 and 1.5 mmol·L-1)under acidic conditions(pH 4.5).In order to provide robust germplasm resources for the improvement of acid-aluminum soil,the effects of Al 3+stress on the seed germination and seedling growth were initially explored.Our investigations indicated that there were no significant differences(P>0.05)in the germination rate of the three samples of wild E.nutans seeds at lower concentrations of Al 3+stress(0.5 and 1.0 mmol·L-1)compared to the control,but that results were adverse at higher concentrations(1.5 mmol·L-1).The results showed that root length,plant height,root weight and shoots' weight in the three samples decreased with increasing Al 3+concentration.In addition,the MDA and soluble sugar contents of seedlings substantially increased with Al 3+application.Data showed that Al was poorly translocated from the roots to aerial parts of the three seedling samples.In the treatment groups,the Al content of roots increased 12.2,15.3 and 17.5 times more than that of the aerial parts.Al toxicity and phosphorus(P)deficiency often co-exist in acidic soils and limit plant growth and crop production.In this study,we observed that Al stress induced a reduction in P accumulation in the plants.Using eight indicators for seedlings under Al stress,we performed a comprehensive evaluation of Al tolerance of the three E.nutans samples and showed that the Shenza County sample was highly tolerant to aluminum and so could be used for domestication and breeding to provide good germplasm resources for the improvement of acid-aluminum soils.
引文
[1]Kochian L V,Pi1eros M A,Hoekenga O A.The physiology,genetics and molecular biology of plant Al resistance and toxicity.Plant and Soil,2005,274:175-195.
[2]Sade H,Meriga B,Surapu V,et al.Toxicity and tolerance of aluminum in plants:tailoring plants to suit to acid soils.Biometals,2016,29:187-210.
[3]Matsumoto H,Motoda H.Aluminum toxicity recovery processes in root apices;possible association with oxidative stress.Plant Science,2012,186:1-8.
[4]Tahara K,Yamanoshita T,Norisada M,et al.Aluminum distribution and reactive oxygen species accumulation in root tips of two Melaleucatrees differing in aluminum resistance.Plant and Soil,2008,307:167-178.
[5]Poschenrieder C,Gunse B,Corrales I,et al.A glance into aluminum toxicity and resistance in plants.Science of the Total Environment,2008,400(1/3):356-368.
[6]Samac D A,Tesfaye M.Plant improvement for tolerance to aluminum in acid soils-a review.Plant Cell Tissue and Organ Culture,2003,75(3):189-207.
[7]Takeda K,Ma J F.An aluminum-activated citrate transporter in barley.Plant and Cell Physiology,2007,48:1081-1091.
[8]Lu B R.A study on systematic relationships between Elymus and Hordeum.Journal of Systematics and Evolution,1997,35(3):193-207.卢宝荣.披碱草属与大麦属系统关系的研究.植物分类学报,1997,35(3):193-207.
[9]Dong Y C,Liu X.Chinese crops and their wild relatives.Beijing:China Agricultural Press,2007:142-144.董玉琛,刘旭.中国作物及其野生近缘植物.北京:中国农业出版社,2007:142-144.
[10]Miao Y J.Study and utilization on high-quality wild grass germplasm resources in Tibet.Yangling:Northwest Agriculture and Forestry University,2007.苗彦军.西藏几种野生优质牧草种质资源研究与利用.杨凌:西北农林科技大学,2007.
[11]Zhang S X,Ni M P C,Xu Y M,et al.Physiological responses to low temperature stress and cold tolerance evaluation in three Elymus species.Pratacultural Science,2016,33(6):1154-1163.张尚雄,尼玛平措,徐雅梅,等.3个披碱草属牧草对低温胁迫的生理响应及苗期抗寒性评价.草业科学,2016,33(6):1154-1163.
[12]Fu J J,Liu J,Sun Y F,et al.Effects of cold stress on the growths and physiological characteristics of two Elymus nutans varieties.Acta Agrestia Sinica,2014,22(4):789-795.付娟娟,刘建,孙永芳,等.冷胁迫对2种垂穗披碱草生长和生理特性的影响.草地学报,2014,22(4):789-795.
[13]Zhang C N.Study on drought resistance of twenty Elymus L.germplasm resources.Xining:Qinghai University,2010.张晨妮.披碱草属(Elymus L.)20种牧草种质资源的抗旱性研究与评价.西宁:青海大学,2010.
[14]Jia Y X,Li X L,Yuan Q H,et al.Evaluation of salt-tolerance and the related physiological characteristics of wild Elymus spp.Scientia Agricultura Sinica,2008,41(10):2999-3007.贾亚雄,李向林,袁庆华,等.披碱草属野生种质资源苗期耐盐性评价及相关生理机制研究.中国农业科学,2008,41(10):2999-3007.
[15]Ma J F,Ryan P R,Delhaize E.Aluminium tolerance in plants and the complexing role of organic acids.Trends in Plant Science,2001,6(6):273-278.
[16]Silva S.Aluminium toxicity targets in plants.Journal of Botany,2012,(8):219-262.
[17]Ahn S J,Rgngel Z,Matsumoto H.Aluminum-induced plasma membrane surface potential and H+-ATPase activity in nearisogenic wheat lines differing in tolerance to aluminum.New Phytologist,2004,162:71-79.
[18]Guo T,Zhang G,Zhou M,et al.Effects of aluminum and cadmium toxicity on growth and antioxidant enzyme activities of two barley genotypes with different Al resistance.Plant and Soil,2004,258(1):241-248.
[19]Chen J X,Wang X F.Plant physiology experiment guidance.Guangzhou:South China University of Technology Press,2006:74.陈建勋,王晓峰.植物生理学实验指导.广州:华南理工大学出版社,2006:74.
[20]Chen C,Song Y H,Gao X Q.Dry and wet digestion determination of aluminum content in laver.Journal of Food Safety and Quality,2010,27(3):118-123.陈昌,宋颖华,高晓强.干法消解与湿法消解测定紫菜中铝含量.食品安全质量检测学报,2010,27(3):118-123.
[21]Guo Y P,Mi F G,Run L J,et al.Physiological response to drought stresses and drought resistances evaluation of different Kentucky bluegrass varieties.Acta Prataculturae Sinica,2014,23(4):220-228.郭郁频,米福贵,闰利军,等.不同早熟禾品种对干旱胁迫的生理响应及抗旱性评价.草业学报,2014,23(4):220-228.
[22]Alvim M N,Ramos F T,Fran9a M G C.Seed storage period reduces aluminum tolerance in rice(Oryza sativa).Seed Science and Technology,2007,35:688-697.
[23]Alamgir A N M,Akhter S.Effects of aluminium(Al 3+)on seed germination and seedling growth of wheat(Triticum aestivum L.).Bangladesh Journal of Botany,2009,38:1-6.
[24]Horst W J,Wang Y,Eticha D.The role of the root apoplast in aluminium-induced inhibition of root elongation and in aluminium resistance of plants:a review.Annals of Botany,2010,106:185-197.
[25]Motoda H,Kano Y,Hiragami F,et al.Morphological changes in the apex of pea roots during and after recovery from aluminum treatment.Plant and Soil,2010,333:49-58.
[26]Zhang H H,Jiang Z,Qin R,et al.Accumulation and cellular toxicity of aluminum in seedling of Pinus massoniana.BMC Plant Biology,2014,14:2-16.
[27]Kochian L V,Hoekenga O A,Pi1eros M A.How do crop plants tolerate acid soils?Mechanisms of aluminum tolerance and phosphorous efficiency.Annual Review of Plant Biology,2004,55:459-493.
[28]Xu F J,Li G,Jin C W,et al.Aluminum-induced changes in reactive oxygen species accumulation,lipid peroxidation and antioxidant capacity in wheat root tips.Biologia Plantarum,2012,56:89-96.
[29]Ou Z L,Cao F L,Zheng J.The changes of physio-biochemistry and morphological characteristics of Ginkgo biloba L.subjected to high temperature stress.Journal of Nanjing Forestry University(Natural Sciences Edition),2008,32(3):31-34.欧祖兰,曹福亮,郑军.高温胁迫下银杏形态及生理生化指标的变化.南京林业大学学报(自然科学版),2008,32(3):31-34.
[30]Su H M,He B H,Cai X H,et al.Effects of water stress on the morphological and physiological characteristics of Scaevola aemulacutting seedling.Chinese Journal of Ecology,2011,30(10):2185-2190.苏慧敏,何丙辉,蔡兴华,等.水分胁迫对太阳扇扦插苗形态和生理特征的影响.生态学杂志,2011,30(10):2185-2190.
[31]Tabaldi L A,Cargnelutti D,Goncalves J F,et al.Oxidative stress is an early symptom triggered by aluminum in Al-sensitive potato plantlets.Chemosphere,2009,76:1402-1409.
[32]AL-Qahtani K M.Assessment of heavy metals accumulation in native plant species from soils contaminated in Riyadh City,Saudi Arabia.Life Science Journal,2012,9:384-392.
[33]Ma J F,Hiradate S,Nomoto K,et al.Internal detoxification mechanism of Al in hydrangea:identification of Al form in the leaves.Plant Physiology,1997,113:1033-1039.
[34]Matsumoto H,Hirasawa E,Morimura S,et al.Localization of aluminum in tea leaves.Plant and Cell Physiology,1976,17:627-631.
[35]Liao H,Wan H,Shaff J,et al.Phosphorus and aluminum interactions in soybean in relation to aluminum tolerance:Exudation of specific organic acids from different regions of the intact root system.Plant Physiology,2006,141(2):674-684.
[36]Chen W R,Liu P,Huang C B,et al.Effects of aluminum on influx and translocation of aluminum and other essential elements in Buck wheat(Fagopyrum esculentum Monench).Journal of Soil and Water Conservation,2006,20(3):173-186.陈文荣,刘鹏,黄朝表,等.铝对荞麦铝和其它营养元素运输的影响.水土保持学报,2006,20(3):173-186.
[37]Mugwira L M,Patel S U,Fleming A L.Aluminium effects on growth and Al,Ca,Mg,K and P levels in triticale,wheat and rye.Plant and Soil,1980,57(2/3):467-470.
[38]Sun Q B,Dong X Y,Shen R F.Effects of phosphorus or lime applications on growth and mineral compositions of two Lespedezaspecies.Soils,2009,41(2):206-211.孙清斌,董晓英,沈仁芳.施用磷、钙对红壤上胡枝子生长和矿质元素含量的影响.土壤,2009,41(2):206-211.
[2]Sade H,Meriga B,Surapu V,et al.Toxicity and tolerance of aluminum in plants:tailoring plants to suit to acid soils.Biometals,2016,29:187-210.
[3]Matsumoto H,Motoda H.Aluminum toxicity recovery processes in root apices;possible association with oxidative stress.Plant Science,2012,186:1-8.
[4]Tahara K,Yamanoshita T,Norisada M,et al.Aluminum distribution and reactive oxygen species accumulation in root tips of two Melaleucatrees differing in aluminum resistance.Plant and Soil,2008,307:167-178.
[5]Poschenrieder C,Gunse B,Corrales I,et al.A glance into aluminum toxicity and resistance in plants.Science of the Total Environment,2008,400(1/3):356-368.
[6]Samac D A,Tesfaye M.Plant improvement for tolerance to aluminum in acid soils-a review.Plant Cell Tissue and Organ Culture,2003,75(3):189-207.
[7]Takeda K,Ma J F.An aluminum-activated citrate transporter in barley.Plant and Cell Physiology,2007,48:1081-1091.
[8]Lu B R.A study on systematic relationships between Elymus and Hordeum.Journal of Systematics and Evolution,1997,35(3):193-207.卢宝荣.披碱草属与大麦属系统关系的研究.植物分类学报,1997,35(3):193-207.
[9]Dong Y C,Liu X.Chinese crops and their wild relatives.Beijing:China Agricultural Press,2007:142-144.董玉琛,刘旭.中国作物及其野生近缘植物.北京:中国农业出版社,2007:142-144.
[10]Miao Y J.Study and utilization on high-quality wild grass germplasm resources in Tibet.Yangling:Northwest Agriculture and Forestry University,2007.苗彦军.西藏几种野生优质牧草种质资源研究与利用.杨凌:西北农林科技大学,2007.
[11]Zhang S X,Ni M P C,Xu Y M,et al.Physiological responses to low temperature stress and cold tolerance evaluation in three Elymus species.Pratacultural Science,2016,33(6):1154-1163.张尚雄,尼玛平措,徐雅梅,等.3个披碱草属牧草对低温胁迫的生理响应及苗期抗寒性评价.草业科学,2016,33(6):1154-1163.
[12]Fu J J,Liu J,Sun Y F,et al.Effects of cold stress on the growths and physiological characteristics of two Elymus nutans varieties.Acta Agrestia Sinica,2014,22(4):789-795.付娟娟,刘建,孙永芳,等.冷胁迫对2种垂穗披碱草生长和生理特性的影响.草地学报,2014,22(4):789-795.
[13]Zhang C N.Study on drought resistance of twenty Elymus L.germplasm resources.Xining:Qinghai University,2010.张晨妮.披碱草属(Elymus L.)20种牧草种质资源的抗旱性研究与评价.西宁:青海大学,2010.
[14]Jia Y X,Li X L,Yuan Q H,et al.Evaluation of salt-tolerance and the related physiological characteristics of wild Elymus spp.Scientia Agricultura Sinica,2008,41(10):2999-3007.贾亚雄,李向林,袁庆华,等.披碱草属野生种质资源苗期耐盐性评价及相关生理机制研究.中国农业科学,2008,41(10):2999-3007.
[15]Ma J F,Ryan P R,Delhaize E.Aluminium tolerance in plants and the complexing role of organic acids.Trends in Plant Science,2001,6(6):273-278.
[16]Silva S.Aluminium toxicity targets in plants.Journal of Botany,2012,(8):219-262.
[17]Ahn S J,Rgngel Z,Matsumoto H.Aluminum-induced plasma membrane surface potential and H+-ATPase activity in nearisogenic wheat lines differing in tolerance to aluminum.New Phytologist,2004,162:71-79.
[18]Guo T,Zhang G,Zhou M,et al.Effects of aluminum and cadmium toxicity on growth and antioxidant enzyme activities of two barley genotypes with different Al resistance.Plant and Soil,2004,258(1):241-248.
[19]Chen J X,Wang X F.Plant physiology experiment guidance.Guangzhou:South China University of Technology Press,2006:74.陈建勋,王晓峰.植物生理学实验指导.广州:华南理工大学出版社,2006:74.
[20]Chen C,Song Y H,Gao X Q.Dry and wet digestion determination of aluminum content in laver.Journal of Food Safety and Quality,2010,27(3):118-123.陈昌,宋颖华,高晓强.干法消解与湿法消解测定紫菜中铝含量.食品安全质量检测学报,2010,27(3):118-123.
[21]Guo Y P,Mi F G,Run L J,et al.Physiological response to drought stresses and drought resistances evaluation of different Kentucky bluegrass varieties.Acta Prataculturae Sinica,2014,23(4):220-228.郭郁频,米福贵,闰利军,等.不同早熟禾品种对干旱胁迫的生理响应及抗旱性评价.草业学报,2014,23(4):220-228.
[22]Alvim M N,Ramos F T,Fran9a M G C.Seed storage period reduces aluminum tolerance in rice(Oryza sativa).Seed Science and Technology,2007,35:688-697.
[23]Alamgir A N M,Akhter S.Effects of aluminium(Al 3+)on seed germination and seedling growth of wheat(Triticum aestivum L.).Bangladesh Journal of Botany,2009,38:1-6.
[24]Horst W J,Wang Y,Eticha D.The role of the root apoplast in aluminium-induced inhibition of root elongation and in aluminium resistance of plants:a review.Annals of Botany,2010,106:185-197.
[25]Motoda H,Kano Y,Hiragami F,et al.Morphological changes in the apex of pea roots during and after recovery from aluminum treatment.Plant and Soil,2010,333:49-58.
[26]Zhang H H,Jiang Z,Qin R,et al.Accumulation and cellular toxicity of aluminum in seedling of Pinus massoniana.BMC Plant Biology,2014,14:2-16.
[27]Kochian L V,Hoekenga O A,Pi1eros M A.How do crop plants tolerate acid soils?Mechanisms of aluminum tolerance and phosphorous efficiency.Annual Review of Plant Biology,2004,55:459-493.
[28]Xu F J,Li G,Jin C W,et al.Aluminum-induced changes in reactive oxygen species accumulation,lipid peroxidation and antioxidant capacity in wheat root tips.Biologia Plantarum,2012,56:89-96.
[29]Ou Z L,Cao F L,Zheng J.The changes of physio-biochemistry and morphological characteristics of Ginkgo biloba L.subjected to high temperature stress.Journal of Nanjing Forestry University(Natural Sciences Edition),2008,32(3):31-34.欧祖兰,曹福亮,郑军.高温胁迫下银杏形态及生理生化指标的变化.南京林业大学学报(自然科学版),2008,32(3):31-34.
[30]Su H M,He B H,Cai X H,et al.Effects of water stress on the morphological and physiological characteristics of Scaevola aemulacutting seedling.Chinese Journal of Ecology,2011,30(10):2185-2190.苏慧敏,何丙辉,蔡兴华,等.水分胁迫对太阳扇扦插苗形态和生理特征的影响.生态学杂志,2011,30(10):2185-2190.
[31]Tabaldi L A,Cargnelutti D,Goncalves J F,et al.Oxidative stress is an early symptom triggered by aluminum in Al-sensitive potato plantlets.Chemosphere,2009,76:1402-1409.
[32]AL-Qahtani K M.Assessment of heavy metals accumulation in native plant species from soils contaminated in Riyadh City,Saudi Arabia.Life Science Journal,2012,9:384-392.
[33]Ma J F,Hiradate S,Nomoto K,et al.Internal detoxification mechanism of Al in hydrangea:identification of Al form in the leaves.Plant Physiology,1997,113:1033-1039.
[34]Matsumoto H,Hirasawa E,Morimura S,et al.Localization of aluminum in tea leaves.Plant and Cell Physiology,1976,17:627-631.
[35]Liao H,Wan H,Shaff J,et al.Phosphorus and aluminum interactions in soybean in relation to aluminum tolerance:Exudation of specific organic acids from different regions of the intact root system.Plant Physiology,2006,141(2):674-684.
[36]Chen W R,Liu P,Huang C B,et al.Effects of aluminum on influx and translocation of aluminum and other essential elements in Buck wheat(Fagopyrum esculentum Monench).Journal of Soil and Water Conservation,2006,20(3):173-186.陈文荣,刘鹏,黄朝表,等.铝对荞麦铝和其它营养元素运输的影响.水土保持学报,2006,20(3):173-186.
[37]Mugwira L M,Patel S U,Fleming A L.Aluminium effects on growth and Al,Ca,Mg,K and P levels in triticale,wheat and rye.Plant and Soil,1980,57(2/3):467-470.
[38]Sun Q B,Dong X Y,Shen R F.Effects of phosphorus or lime applications on growth and mineral compositions of two Lespedezaspecies.Soils,2009,41(2):206-211.孙清斌,董晓英,沈仁芳.施用磷、钙对红壤上胡枝子生长和矿质元素含量的影响.土壤,2009,41(2):206-211.