NaCl胁迫对牧草种子萌发与幼苗生理生化的影响及耐盐性评价
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摘要
牧草种植是改良盐碱地的有效途径,而耐盐性的评价则是筛选、推广利用优质牧草种质的重要基础,对于改良与利用滨海盐渍化土壤具有重要意义。本研究以草木樨、苜蓿王、紫云英、沙打旺和小冠花5种豆科牧草为试验材料,采用0.0%(CK)、0.2%、0.4%、0.6%、0.8%、1.0%、1.2%、1.4%和1.6%九个NaCl盐浓度梯度,对5种牧草种子萌发和幼苗生长两个阶段进行盐胁迫处理。通过对牧草种子的相对发芽率、相对发芽势、耐盐指数、幼苗生长势及相对含水量、细胞膜透性、叶绿素含量、脯氨酸含量、丙二醛和可溶性糖含量以及保护酶活性等生理生化指标的测定,观测其在盐胁迫下的一系列生理生化反应,并进行综合评价得出其耐盐性强弱。结果表明:
(1)供试的5种牧草种子萌发对NaCl盐胁迫的敏感浓度不一致。低盐浓度(0.2%,0.4%和0.6%)对牧草种子萌发具有促进作用,草木樨、苜蓿王和沙打旺的相对发芽率高于对照;随着盐浓度的升高,供试牧草的相对发芽率和相对发芽势呈明显的下降趋势。盐浓度>0.8%时相对发芽势下降幅度增大,盐浓度为1.6%时,相对发芽率和相对发芽势分别下降到72%和70%以下。二者与盐浓度之间均呈显著至极显著相关关系。
(2)低浓度盐胁迫对幼苗生长具有一定的促进作用,在盐浓度为0.2%～0.8%之间,5种牧草的幼苗生长势均高于对照。幼苗生长势与耐盐指数均随盐浓度增大呈现先增加后降低的趋势。
(3)随着盐浓度的增加,5种牧草叶片相对含水量下降,细胞膜透性增加;盐胁迫下,其叶绿素总量、叶绿素a、叶绿素b含量变化一致,随着盐浓度梯度的升高表现为先增大后降低的趋势,其开始降低的NaCl浓度>0.4%,叶绿素含量的变化主要归因于叶绿素a含量的变化;供试牧草叶绿素a含量与盐浓度之间呈显著至极显著相关关系,叶绿素b含量与盐浓度之间的相关关系依品种不同而有差别。
(4)5种牧草幼苗MDA含量和可溶性糖含量依品种及盐浓度的不同而不同,总体上呈现出低浓度(<0.4%)时升高,随着盐浓度增加而显著降低。在低盐浓度(0.2%和0.4%)时,草木樨、沙打旺、苜蓿王和紫云英的可溶性糖含量高于对照,小冠花变化不明显;不同牧草可溶性糖含量差异明显,MDA含量和可溶性糖含量与盐浓度间均呈极显著负相关关系。
(5)5种牧草体内游离脯氨酸含量和三种酶活性的变化随着盐处理浓度的增大呈现出先增大后降低的变化趋势。不同牧草体内脯氨酸的变化趋势各异,草木樨和沙打旺在不同浓度下游离脯氨酸含量均高于对照。NaCl浓度>1.0%时,苜蓿王、小冠花和紫云英的SOD活性明显下降,草木樨和沙打旺的SOD活性随着盐浓度的增大仅有小幅度的减小。供试牧草的POD和CAT活性在盐浓度为0.4%左右时出现峰值,随着盐浓度的增大而明显下降。
(6)应用主成分分析法对耐盐性评价指标的筛选表明,在牧草种子萌发阶段,相对发芽率、相对发芽势、耐盐指数及幼苗生长势可作为牧草种子耐盐性的评价指标;在幼苗生长阶段,游离脯氨酸、叶绿素含量、SOD活性和可溶性糖含量可作为幼苗生长期耐盐性评价的首选指标。
(7)应用隶属函数法对5种牧草进行综合评价,种子萌发阶段耐盐性大小顺序为苜蓿王>沙打旺>草木樨>小冠花>紫云英;牧草幼苗生长阶段耐盐性大小为草木樨>沙打旺>小冠花>苜蓿王>紫云英;对两个生育阶段耐盐性的综合评价为草木樨>沙打旺>苜蓿王>小冠花>紫云英。草木樨、沙打旺可在轻度、中度盐化土壤上种植,苜蓿王、小冠花可在轻度及潜在盐化土壤上种植,而紫云英不耐盐碱,仅可在潜在盐化土壤上种植。
Herbages growing is an effective way to modify the saline land,and assessment of salt tolerance is the important foundation of screening、utilizing and popularizing germplasms of high quality herbages. It plays an important role in modifying and utilizing coastal salinization soil. In this study, 5 leguminous herbages such as Melilotus albus L.、Medicago stiva L.、Astragalus sinicus L.、Astragalus adsurgens Pall. and Coronilla variable L. were employed to analyze as test materials. Using 9 salt concentration gradients that was 0.0%(CK)、0.2%、0.4%、0.6%、0.8%、1.0%、1.2%、1.4% and 1.6%,we made salt stress treatment to 5 herbages seeds in two stages that was seed germination and seedling growth. With determining some physiological and biochemical indexes,including relative germination rate、relative germination potential、salt tolerance index、seedling growth potential、relative water content、cell membrane permeability、Chlorophyll content、free proline content、MDA content、soluble sugar content and protective enzyme activity and so on, observing a series of physiological and biochemical response on salt stress treatment,and evaluating comprehensively,the order of salt tolerance could be obtained.The results showed that:
(1) The sensitive concentration of of NaCl stress for 5 tested herbages were different.Low salt concentration (0.2%,0.4% and 0.6%) had promoting effect on seed germination of herbages.The relative germination rate of Melilotus albus、Medicago stiva and Astragalus adsurgens were higher than that in the control group.With the increase of salt concentration,relative germination rate and relative germination potential showed apparent downtrend.When the salt concentration was higher than 0.8%,relative germination rate and relative germination potential of tested herbages decreased a lot. When the salt concentration was 1.6%,relative germination rate and relative germination potential of tested herbages decreased to less than 72% and 70% respectively.Both showed significantly and extremely significant correlation with salt concentrations.
(2) Low salt stress concentration had promotion effect on seedling growth of herbages.Between 0.2% and 0.8% salt concentrations,the seedling growth potential of 5 herbages were all higher than those in the control group.With the increase of salt concentration,seedling growth potential and salt tolerance index had the same change tendency as first increasing and then decreasing.
(3) With the increase of salt concentration,the leaf relative water content of 5 herbages decreased,cell membrane permeability increased;Chlorophyll content、Chlorophyll a content and Chlorophyll b content showed similar changes on salt stress,the trend was first increasing and then decreasing.When the salt concentration was higher than 0.4%,the change of Chlorophyll content mainly attributed to the change of Chlorophyll a content. Chlorophyll a content showed significantly and extremely significant correlation with salt concentrations.The correlation between Chlorophyll b content and salt concentration were different relaying on the varieties of herbages.
(4) The MDA content and soluble sugar content of 5 herbages varied according to varieties of herbages and salt concentration,both showed increasing at low salt concentration(less than 0.4%),and then decreasing significantly with the increasing of salt concentration.At the low salt concentration(0.2% and 0.4%),soluble sugar content of Melilotus albus、Astragalus adsurgens、Medicago stiva and Astragalus sinicus were higher than that in the control group,but Coronilla variable did not change significantly;different herbages had obviously different soluble sugar content, MDA content and soluble sugar content all showed extremely negatively significant correlation with salt concentrations.
(5) The change of free proline content and three kinds of protective enzyme activity in 5 herbages showed first increasing and then decreasing trend with the increase of salt concentration.The change of free proline content showed different trends in different herbages,free proline content in Melilotus albus and Astragalus adsurgens were higher than that in the control group. When the salt concentration was higher than 1.0%, SOD activity of Medicago stiva、Coronilla variable and Astragalus sinicus decreased obviously,and SOD activity of Melilotus albus and Astragalus adsurgens had only small range of decreasing with the increase of salt concentration.When the salt concentration was 0.4%, POD activity and CAT activity appeared peak value,and with the increase of salt concentration,they decreased obviously.
(6) Using principal component analysis to assess the index of salt tolerance,it showed that on the germination stage of herbages,the assessment index of salt tolerance of herbages seeds were relative germination rate、relative germination potential、salt tolerance indexand seedling growth potential;on the seedling growth stage of herbages,the first choice to assess salt tolerance were free proline content、Chlorophyll content、SOD activity and soluble sugar content.
(7) Using membership function to assess comprehensively 5 herbages,it showed that the sequence of salt tolerance on the germination stage of herbages was Medicago stiva>Astragalus adsurgens > Melilotus albus > Coronilla variable > Astragalus sinicus;on the seedling growth stage of herbages,it was Melilotus albus>Astragalus adsurgens>Coronilla variable>Medicago stiva>Astragalus sinicus;on the two stages of growing, it was Melilotus albus>Astragalus adsurgens> Medicago stiva>Coronilla variable>Astragalus sinicus. Melilotus albus and Astragalus adsurgens could be planted on mild and moderate salinization land, Medicago stiva and Coronilla variable could be planted on mild or potential salinization land, Astragalus sinicus could not be tolerant of saline and alkaline,it could only be planted on potential salinization land.
(1)供试的5种牧草种子萌发对NaCl盐胁迫的敏感浓度不一致。低盐浓度(0.2%,0.4%和0.6%)对牧草种子萌发具有促进作用,草木樨、苜蓿王和沙打旺的相对发芽率高于对照;随着盐浓度的升高,供试牧草的相对发芽率和相对发芽势呈明显的下降趋势。盐浓度>0.8%时相对发芽势下降幅度增大,盐浓度为1.6%时,相对发芽率和相对发芽势分别下降到72%和70%以下。二者与盐浓度之间均呈显著至极显著相关关系。
(2)低浓度盐胁迫对幼苗生长具有一定的促进作用,在盐浓度为0.2%～0.8%之间,5种牧草的幼苗生长势均高于对照。幼苗生长势与耐盐指数均随盐浓度增大呈现先增加后降低的趋势。
(3)随着盐浓度的增加,5种牧草叶片相对含水量下降,细胞膜透性增加;盐胁迫下,其叶绿素总量、叶绿素a、叶绿素b含量变化一致,随着盐浓度梯度的升高表现为先增大后降低的趋势,其开始降低的NaCl浓度>0.4%,叶绿素含量的变化主要归因于叶绿素a含量的变化;供试牧草叶绿素a含量与盐浓度之间呈显著至极显著相关关系,叶绿素b含量与盐浓度之间的相关关系依品种不同而有差别。
(4)5种牧草幼苗MDA含量和可溶性糖含量依品种及盐浓度的不同而不同,总体上呈现出低浓度(<0.4%)时升高,随着盐浓度增加而显著降低。在低盐浓度(0.2%和0.4%)时,草木樨、沙打旺、苜蓿王和紫云英的可溶性糖含量高于对照,小冠花变化不明显;不同牧草可溶性糖含量差异明显,MDA含量和可溶性糖含量与盐浓度间均呈极显著负相关关系。
(5)5种牧草体内游离脯氨酸含量和三种酶活性的变化随着盐处理浓度的增大呈现出先增大后降低的变化趋势。不同牧草体内脯氨酸的变化趋势各异,草木樨和沙打旺在不同浓度下游离脯氨酸含量均高于对照。NaCl浓度>1.0%时,苜蓿王、小冠花和紫云英的SOD活性明显下降,草木樨和沙打旺的SOD活性随着盐浓度的增大仅有小幅度的减小。供试牧草的POD和CAT活性在盐浓度为0.4%左右时出现峰值,随着盐浓度的增大而明显下降。
(6)应用主成分分析法对耐盐性评价指标的筛选表明,在牧草种子萌发阶段,相对发芽率、相对发芽势、耐盐指数及幼苗生长势可作为牧草种子耐盐性的评价指标;在幼苗生长阶段,游离脯氨酸、叶绿素含量、SOD活性和可溶性糖含量可作为幼苗生长期耐盐性评价的首选指标。
(7)应用隶属函数法对5种牧草进行综合评价,种子萌发阶段耐盐性大小顺序为苜蓿王>沙打旺>草木樨>小冠花>紫云英;牧草幼苗生长阶段耐盐性大小为草木樨>沙打旺>小冠花>苜蓿王>紫云英;对两个生育阶段耐盐性的综合评价为草木樨>沙打旺>苜蓿王>小冠花>紫云英。草木樨、沙打旺可在轻度、中度盐化土壤上种植,苜蓿王、小冠花可在轻度及潜在盐化土壤上种植,而紫云英不耐盐碱,仅可在潜在盐化土壤上种植。
Herbages growing is an effective way to modify the saline land,and assessment of salt tolerance is the important foundation of screening、utilizing and popularizing germplasms of high quality herbages. It plays an important role in modifying and utilizing coastal salinization soil. In this study, 5 leguminous herbages such as Melilotus albus L.、Medicago stiva L.、Astragalus sinicus L.、Astragalus adsurgens Pall. and Coronilla variable L. were employed to analyze as test materials. Using 9 salt concentration gradients that was 0.0%(CK)、0.2%、0.4%、0.6%、0.8%、1.0%、1.2%、1.4% and 1.6%,we made salt stress treatment to 5 herbages seeds in two stages that was seed germination and seedling growth. With determining some physiological and biochemical indexes,including relative germination rate、relative germination potential、salt tolerance index、seedling growth potential、relative water content、cell membrane permeability、Chlorophyll content、free proline content、MDA content、soluble sugar content and protective enzyme activity and so on, observing a series of physiological and biochemical response on salt stress treatment,and evaluating comprehensively,the order of salt tolerance could be obtained.The results showed that:
(1) The sensitive concentration of of NaCl stress for 5 tested herbages were different.Low salt concentration (0.2%,0.4% and 0.6%) had promoting effect on seed germination of herbages.The relative germination rate of Melilotus albus、Medicago stiva and Astragalus adsurgens were higher than that in the control group.With the increase of salt concentration,relative germination rate and relative germination potential showed apparent downtrend.When the salt concentration was higher than 0.8%,relative germination rate and relative germination potential of tested herbages decreased a lot. When the salt concentration was 1.6%,relative germination rate and relative germination potential of tested herbages decreased to less than 72% and 70% respectively.Both showed significantly and extremely significant correlation with salt concentrations.
(2) Low salt stress concentration had promotion effect on seedling growth of herbages.Between 0.2% and 0.8% salt concentrations,the seedling growth potential of 5 herbages were all higher than those in the control group.With the increase of salt concentration,seedling growth potential and salt tolerance index had the same change tendency as first increasing and then decreasing.
(3) With the increase of salt concentration,the leaf relative water content of 5 herbages decreased,cell membrane permeability increased;Chlorophyll content、Chlorophyll a content and Chlorophyll b content showed similar changes on salt stress,the trend was first increasing and then decreasing.When the salt concentration was higher than 0.4%,the change of Chlorophyll content mainly attributed to the change of Chlorophyll a content. Chlorophyll a content showed significantly and extremely significant correlation with salt concentrations.The correlation between Chlorophyll b content and salt concentration were different relaying on the varieties of herbages.
(4) The MDA content and soluble sugar content of 5 herbages varied according to varieties of herbages and salt concentration,both showed increasing at low salt concentration(less than 0.4%),and then decreasing significantly with the increasing of salt concentration.At the low salt concentration(0.2% and 0.4%),soluble sugar content of Melilotus albus、Astragalus adsurgens、Medicago stiva and Astragalus sinicus were higher than that in the control group,but Coronilla variable did not change significantly;different herbages had obviously different soluble sugar content, MDA content and soluble sugar content all showed extremely negatively significant correlation with salt concentrations.
(5) The change of free proline content and three kinds of protective enzyme activity in 5 herbages showed first increasing and then decreasing trend with the increase of salt concentration.The change of free proline content showed different trends in different herbages,free proline content in Melilotus albus and Astragalus adsurgens were higher than that in the control group. When the salt concentration was higher than 1.0%, SOD activity of Medicago stiva、Coronilla variable and Astragalus sinicus decreased obviously,and SOD activity of Melilotus albus and Astragalus adsurgens had only small range of decreasing with the increase of salt concentration.When the salt concentration was 0.4%, POD activity and CAT activity appeared peak value,and with the increase of salt concentration,they decreased obviously.
(6) Using principal component analysis to assess the index of salt tolerance,it showed that on the germination stage of herbages,the assessment index of salt tolerance of herbages seeds were relative germination rate、relative germination potential、salt tolerance indexand seedling growth potential;on the seedling growth stage of herbages,the first choice to assess salt tolerance were free proline content、Chlorophyll content、SOD activity and soluble sugar content.
(7) Using membership function to assess comprehensively 5 herbages,it showed that the sequence of salt tolerance on the germination stage of herbages was Medicago stiva>Astragalus adsurgens > Melilotus albus > Coronilla variable > Astragalus sinicus;on the seedling growth stage of herbages,it was Melilotus albus>Astragalus adsurgens>Coronilla variable>Medicago stiva>Astragalus sinicus;on the two stages of growing, it was Melilotus albus>Astragalus adsurgens> Medicago stiva>Coronilla variable>Astragalus sinicus. Melilotus albus and Astragalus adsurgens could be planted on mild and moderate salinization land, Medicago stiva and Coronilla variable could be planted on mild or potential salinization land, Astragalus sinicus could not be tolerant of saline and alkaline,it could only be planted on potential salinization land.
引文
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