浒苔硫酸多糖对NaCl胁迫下植物的影响
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摘要
通过外源物质增强植物的抗盐能力，对于合理开发和利用盐碱地，增加粮食产量，具有重要意义。许多研究表明，浒苔硫酸多糖是一种具有多种生物活性的物质，但是对于其在农业方面的研究和应用还未见报道。本文使用微波辅助降解的方法制备了不同分子量的浒苔硫酸多糖，并对其提高植物的抗盐性方面进行了研究，通过各种生理生化指标及分子生物学方法对其增强植物抗盐性的效果进行了比较，初步揭示了部分浒苔硫酸多糖提高植物抗盐作用机理。本研究对浒苔硫酸多糖增强植物抗盐性进行了有益的探索同时也为“绿潮”藻的高值利用提供了新思路。获得主要结论如下：
1．化学组成分析显示浒苔硫酸多糖(EP)为高分子量杂多糖，总糖含量为52.83%，硫酸根含量和糖醛酸含量分别达到21.98%和26.46%，。鼠李糖和葡萄糖是主要的单糖组分（含量为1:1），同时还含有木糖，少量的甘露糖和半乳糖以及痕量的岩藻糖。红外光谱显示，在842和1248以及1642cm-1有典型的C-O-S键以及S=O和C=O键吸收峰。以EP为原料，通过微波辅助等技术制备了不同分子量的浒苔硫酸多糖DEP-1、DEP-2和DEP-3，分子量分别为234KDa、19.1KDa和5.1KDa，理化分析显示，DEP-1、DEP-2和DEP-3与EP的硫酸根含量和糖醛酸含量基本相同，单糖组成基本相似，暗示降解过程只是破坏了糖苷键，没有对多糖的重要化学基团产生影响。
2．研究了EP对NaCl胁迫下玉米和小麦的影响，结果表明：1mg/mL EP可以提高100mmol/L NaCl胁迫条件下玉米、小麦幼苗地上部分和根系生长，减少MDA含量，降低细胞膜透性。活性氧清除酶系SOD、CAT、POD和APX的活性都有不同程度增加，增强了活性氧清除能力，避免氧化胁迫对玉米和小麦幼苗的损伤。同时，还降低了叶绿素的分解，能够保护叶绿体，增强盐胁迫下的光合作用。
3．不同分子量浒苔硫酸多糖对于NaCl胁迫下玉米、小麦的影响表明：在盐胁迫条件下，与EP相比，1mg/L的DEP-1、DEP-2和DEP-3具有更好的促进玉米、小麦幼苗生及根生长，提高生物量积累的作用。通过对快速叶绿素荧光动力学曲线及K点相对可变荧光、PSⅡ最大光化学效率以及光合性能指数PI等测定，证明浒苔硫酸多糖能够减少盐胁迫时PSⅡ的损伤，保持光合机构正常状态。推测浒苔硫酸多糖缓解NaCl的胁迫机理可能是减轻了光抑制作用。
4．不同分子量浒苔硫酸多糖在盐胁迫下对拟南芥的作用表明：0.2mg/mL,1mg/mL以及5mg/mL EP、DEP-1、DEP-2和DEP-3对于诱导拟南芥抗盐胁迫均具有一定作用，但是1mg/mL的效果较明显。浓度相同时，低分子量浒苔硫酸多糖DEP-1与DEP-3对于提高拟南芥抗盐性效果较好。1mg/mL EP、DEP-1、DEP-2、DEP-3处理拟南芥后，拟南芥中抗盐胁迫相关基因SOS1与rd29A的表达均有不同程度上调，其中DEP-1与DEP-3处理后，SOS1基因表达分别是对照的1.23倍和1.22倍，rd29A基因表达分别是对照的5.6倍和2.5倍，说明DEP-1与DEP-3可增强拟南芥抗盐能力，推测浒苔硫酸多糖可以通过增强Na+区隔化以及渗透调节能力来增强植物抗盐性。
Efficient foreign substances could improve salt tolerance in plants. That was very meaningfulfor increasing grain yields and utilizing of saline-alkali land efficiently. Many study resultsshowed that sulfated polysaccharides from Enteromorpha prolifera was a kind of bioactivesubstances, however there were little work in the literature is related to the agricultural uses ofthem. We obtained sulfated polysaccharides from E. prolifera with different molecular weights bymicrowave-assistance. Using some physiological and biochemical methods, we estimated theefforts of sulfated polysaccharides from E. prolifera on salt tolerance of plants and wanted toinvestigate the related mechanism. Our studies not only had significant meaning to improve thesalt tolerance in plants with foreign substances but also gave a way for potential use of ‘green tideseaweed’. The conclusions are as below:
1. The chemical compositions showed EP was a kind of heteropolysaccharide with huge Mwand the content of total sugar was52.83%. The contents of sulfate and uronic acid in sulfatedpolysaccharide from E. prolifera (EP) were respectively21.98%and26.46%. The majorproportions of monosaccharide were rhamnose and glucose (almost1:1) and there were also smallamounts of xylose, mannose and galactose and trace amounts of fucose. The characteristicabsorption of EP in the FT-IR suggested that typical absorptions at842,1248and1642cm-1werecontributed to the stretching vibration of C-O-S, S=O and C=O respectively. A method wasestablished to degrade EP with microwave-assistance acid hydrolysis and prepared EPs withdifferent Mws. DEP-1、DEP-2and DEP-3were prepared and their Mws were234KDa,19.1KDaand5.1KDa respectively. Analysis of chemical compositions and FT-IR spectra indicated that thecontents of sulfate and uronic acid in DEP-1、DEP-2and DEP-3were similar to EP. The resultindicated that only glycosidic linkages were cleft psecially and the significant structural units inEP were not broken.
2. We investigated the influences of EP on maize and wheat under slat stress and the resultshowed:1mg/mL EP could increase plant and rootstock height both in maize and wheat under100mmol/L NaCl stress. The contents of MDA and the membrane pemeability decreased.Compared to NaCl stress, the SOD, CAT, POD and APX activity were stimulated to higher levelsunder both EP and NaCl treatment. Those free radical scavenging enzymes could avoid thedamage by accumulation of reactive oxygen species in maize and wheat. Also the degradation ofchlorophyll was declined which implied the using of EP could protect chloroplast functions andsustain photosynthesis under salt stress.
3. We also studied the changes of salt tolerance in maize and wheat after being treated withdifferent Mws samples:1mg/L DEP-1, DEP-2and DEP-3could increase more plant and rootstockheight than EP both in maize and wheat under100mmol/L NaCl stress. We got the same result onbiomass accumulation. We investigated chlorophyll a fluorescence transient and some importantparameters such as Wk%, φPoand PI. Results showed that EP, DEP-1, DEP-2and DEP-3couldprotect PSⅡ from severely damage and keep photosynthetic apparatus maintain normal functionunder slat stress. It also implied that the possible mechanism may relate to alleviating effect onphotoinhibition.
4. EP, DEP-1, DEP-2and DEP-3with concentrations of0.2mg/mL,1mg/mL or5mg/mLcould improve salt tolerance in arabidopsis thaliana while the samples of1mg/mL had moresignificant effort. Compared to EP, DEP-1and DEP-3had better efforts under the sameconcentration. The plants showed upregulation of SOS1and rd29A after treated with1mg/mL EP,DEP-1, DEP-2, and DEP-3by RT-PCR. Compared to CK, SOS1upregulated to1.23and1.22times after treated with DEP-1and DEP-3, while rd29A upregulated to5.6and2.5timesrespectively. Results showed that DEP-1and DEP-3increased salt tolerance well in Arabidopsisthaliana. We concluded that the EPs could increase intracellular compartmentation of Na+andosmotic adjustment to improve slat tolerance in arabidopsis thaliana.
1．化学组成分析显示浒苔硫酸多糖(EP)为高分子量杂多糖，总糖含量为52.83%，硫酸根含量和糖醛酸含量分别达到21.98%和26.46%，。鼠李糖和葡萄糖是主要的单糖组分（含量为1:1），同时还含有木糖，少量的甘露糖和半乳糖以及痕量的岩藻糖。红外光谱显示，在842和1248以及1642cm-1有典型的C-O-S键以及S=O和C=O键吸收峰。以EP为原料，通过微波辅助等技术制备了不同分子量的浒苔硫酸多糖DEP-1、DEP-2和DEP-3，分子量分别为234KDa、19.1KDa和5.1KDa，理化分析显示，DEP-1、DEP-2和DEP-3与EP的硫酸根含量和糖醛酸含量基本相同，单糖组成基本相似，暗示降解过程只是破坏了糖苷键，没有对多糖的重要化学基团产生影响。
2．研究了EP对NaCl胁迫下玉米和小麦的影响，结果表明：1mg/mL EP可以提高100mmol/L NaCl胁迫条件下玉米、小麦幼苗地上部分和根系生长，减少MDA含量，降低细胞膜透性。活性氧清除酶系SOD、CAT、POD和APX的活性都有不同程度增加，增强了活性氧清除能力，避免氧化胁迫对玉米和小麦幼苗的损伤。同时，还降低了叶绿素的分解，能够保护叶绿体，增强盐胁迫下的光合作用。
3．不同分子量浒苔硫酸多糖对于NaCl胁迫下玉米、小麦的影响表明：在盐胁迫条件下，与EP相比，1mg/L的DEP-1、DEP-2和DEP-3具有更好的促进玉米、小麦幼苗生及根生长，提高生物量积累的作用。通过对快速叶绿素荧光动力学曲线及K点相对可变荧光、PSⅡ最大光化学效率以及光合性能指数PI等测定，证明浒苔硫酸多糖能够减少盐胁迫时PSⅡ的损伤，保持光合机构正常状态。推测浒苔硫酸多糖缓解NaCl的胁迫机理可能是减轻了光抑制作用。
4．不同分子量浒苔硫酸多糖在盐胁迫下对拟南芥的作用表明：0.2mg/mL,1mg/mL以及5mg/mL EP、DEP-1、DEP-2和DEP-3对于诱导拟南芥抗盐胁迫均具有一定作用，但是1mg/mL的效果较明显。浓度相同时，低分子量浒苔硫酸多糖DEP-1与DEP-3对于提高拟南芥抗盐性效果较好。1mg/mL EP、DEP-1、DEP-2、DEP-3处理拟南芥后，拟南芥中抗盐胁迫相关基因SOS1与rd29A的表达均有不同程度上调，其中DEP-1与DEP-3处理后，SOS1基因表达分别是对照的1.23倍和1.22倍，rd29A基因表达分别是对照的5.6倍和2.5倍，说明DEP-1与DEP-3可增强拟南芥抗盐能力，推测浒苔硫酸多糖可以通过增强Na+区隔化以及渗透调节能力来增强植物抗盐性。
Efficient foreign substances could improve salt tolerance in plants. That was very meaningfulfor increasing grain yields and utilizing of saline-alkali land efficiently. Many study resultsshowed that sulfated polysaccharides from Enteromorpha prolifera was a kind of bioactivesubstances, however there were little work in the literature is related to the agricultural uses ofthem. We obtained sulfated polysaccharides from E. prolifera with different molecular weights bymicrowave-assistance. Using some physiological and biochemical methods, we estimated theefforts of sulfated polysaccharides from E. prolifera on salt tolerance of plants and wanted toinvestigate the related mechanism. Our studies not only had significant meaning to improve thesalt tolerance in plants with foreign substances but also gave a way for potential use of ‘green tideseaweed’. The conclusions are as below:
1. The chemical compositions showed EP was a kind of heteropolysaccharide with huge Mwand the content of total sugar was52.83%. The contents of sulfate and uronic acid in sulfatedpolysaccharide from E. prolifera (EP) were respectively21.98%and26.46%. The majorproportions of monosaccharide were rhamnose and glucose (almost1:1) and there were also smallamounts of xylose, mannose and galactose and trace amounts of fucose. The characteristicabsorption of EP in the FT-IR suggested that typical absorptions at842,1248and1642cm-1werecontributed to the stretching vibration of C-O-S, S=O and C=O respectively. A method wasestablished to degrade EP with microwave-assistance acid hydrolysis and prepared EPs withdifferent Mws. DEP-1、DEP-2and DEP-3were prepared and their Mws were234KDa,19.1KDaand5.1KDa respectively. Analysis of chemical compositions and FT-IR spectra indicated that thecontents of sulfate and uronic acid in DEP-1、DEP-2and DEP-3were similar to EP. The resultindicated that only glycosidic linkages were cleft psecially and the significant structural units inEP were not broken.
2. We investigated the influences of EP on maize and wheat under slat stress and the resultshowed:1mg/mL EP could increase plant and rootstock height both in maize and wheat under100mmol/L NaCl stress. The contents of MDA and the membrane pemeability decreased.Compared to NaCl stress, the SOD, CAT, POD and APX activity were stimulated to higher levelsunder both EP and NaCl treatment. Those free radical scavenging enzymes could avoid thedamage by accumulation of reactive oxygen species in maize and wheat. Also the degradation ofchlorophyll was declined which implied the using of EP could protect chloroplast functions andsustain photosynthesis under salt stress.
3. We also studied the changes of salt tolerance in maize and wheat after being treated withdifferent Mws samples:1mg/L DEP-1, DEP-2and DEP-3could increase more plant and rootstockheight than EP both in maize and wheat under100mmol/L NaCl stress. We got the same result onbiomass accumulation. We investigated chlorophyll a fluorescence transient and some importantparameters such as Wk%, φPoand PI. Results showed that EP, DEP-1, DEP-2and DEP-3couldprotect PSⅡ from severely damage and keep photosynthetic apparatus maintain normal functionunder slat stress. It also implied that the possible mechanism may relate to alleviating effect onphotoinhibition.
4. EP, DEP-1, DEP-2and DEP-3with concentrations of0.2mg/mL,1mg/mL or5mg/mLcould improve salt tolerance in arabidopsis thaliana while the samples of1mg/mL had moresignificant effort. Compared to EP, DEP-1and DEP-3had better efforts under the sameconcentration. The plants showed upregulation of SOS1and rd29A after treated with1mg/mL EP,DEP-1, DEP-2, and DEP-3by RT-PCR. Compared to CK, SOS1upregulated to1.23and1.22times after treated with DEP-1and DEP-3, while rd29A upregulated to5.6and2.5timesrespectively. Results showed that DEP-1and DEP-3increased salt tolerance well in Arabidopsisthaliana. We concluded that the EPs could increase intracellular compartmentation of Na+andosmotic adjustment to improve slat tolerance in arabidopsis thaliana.
引文
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