高羊茅不同基因型抗旱性评价及其生理调节机理研究
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摘要
干旱胁迫是一种限制植物生长的环境因素。研究植物抗旱机理与植物适应干旱胁迫的调节机制，对选育抗旱新品种和缓解水资源短缺具有重大意义。本文以44份来自非洲、欧洲、美洲和澳洲的不同基因型高羊茅(Festucaarundinacea)为材料，首先测定干旱胁迫下的生理差异，结合分子标记结果研究高羊茅抗旱性聚类；其次，以抗旱差异较大的两个高羊茅基因型，分析高羊茅抗旱性能与整株生理、生化、形态特征之间的关系。论文还以美国7个田间抗旱差异较大的高羊茅基因型为材料，研究不同基因型地上部水分利用、根系生长、根系活力的差异，以及干旱响应蛋白质表达的差异。主要研究内容及结果如下：
1.不同高羊茅基因型抗旱性聚类分析
44份高羊茅基因型在温室干旱处理了13d。根据两次草坪质量（Turfquality, TQ）、叶片相对含水量（Relative water content, RWC）、电解液渗透率（Electric leakage）、叶绿素含量（chlorophyll content, Chl）、以及蒸散率（Evapotranspiration rate，ET）的试验结果，确定44份材料的抗旱性差异。聚类结果发现12份材料抗旱能力强，15份基因型对干旱敏感。在44份材料中，来自美洲的‘SH-3-AM-US’和欧洲的‘PI598949-EU-IT’抗旱能力均较强，但两者间的遗传差异较大。选择抗旱且有较大遗传差异的基因资源作为育种计划的候选亲本材料，将有助于培育抗旱性能强的品种。
2.根系抗氧化酶活性与高羊茅抗旱性关系的研究
抗旱基因型‘Focus’和旱敏感基因型‘PI150156’在温室干旱处理50d。测定TQ、RWC、EL、根系活力(Root Viability, RV)和根系抗氧化酶活性，判断根系抗氧化酶活性与高羊茅抗旱性的关系。TQ、RWC与RV、根系超氧化物歧化酶（Superoxide Dismutase, SOD）和过氧化氢酶（Catlase, CAT）酶活性变化成正相关关系；EL与RV、根系SOD和CAT酶活性成负相关关系，EL与以及根系谷胱甘肽还原酶（Glutathione Reductase, GR）和抗坏血酸还原酶（Ascorbate Peroxidase, APX）酶活性呈负相关性关系。结果说明维持根系底部抗氧化酶(SOD、CAT、GR和APX)活性能促进高羊茅在干旱胁迫下保持较高的草坪质量(用TQ衡量)、冠层水状态(用RWC衡量)、膜质稳定性(用EL衡量)和根系活力(RV)，提高高羊茅的抗旱性。
3.高羊茅水分代谢、光合特征及根系生长与高羊茅抗旱性关系的研究
试验以‘Focus’和‘PI150156’，7个高羊茅基因型(‘RU5’、‘RU7’、‘RU9’、‘RU10’、‘RU18’、‘RU19’和‘RU21’)为材料，分2个试验分别研究了干旱胁迫下冠层水分代谢指标、光合特征、以及根系形态和根系活力的变化。干旱后期，抗旱基因型的水分利用率（water use effciency, WUE）、蒸散率、以及冠层与空气温度差（canopy-air temperature, CTD）显著高于干旱敏感型；光合作用受到抑制，但抗旱基因型的净光合速率（net photosyntheticrate, Pn）、蒸腾速率（Transpiration rate, Tr）、气孔导度（stomatal conductance,gs）和气孔开度(stomatal aperture, SA)都显著高于干旱敏感基因型；抗旱基因型的深层根系活力（Root viability, RV）、根系干重(Total root weight)与长度(Total root length)，以及相对根茎（Specific root length, SRL）均显著高于干旱敏感基因型。结果发现优异的草坪质量、较高的叶片含水量与气孔开放程度（高气孔导度）、更有效的水分利用（高蒸散量和蒸腾速率有效降低了草坪冠层温度，高水分利用率提高了植物水分利用效率）、高光合效率、以及根系活力与根系吸水能力有关。另外，相对根茎值大的根系也是植物应对干旱的策略。
4.干旱胁迫对高羊茅干旱响应蛋白的影响
抗旱高羊茅‘RU9’和干旱敏感型‘RU18’干旱处理12d，研究了TQ、RWC、EL、Pn和光化学效率(photosynthesis efficiency, Fv/Fm)，以及蛋白表达谱的变化。试验选取干旱第12d与浇水对照的叶片，进行了全蛋白的提取和双向电泳分离。凝胶图像分析发现处理间、材料间共有31个差异蛋白点，质谱结果显示其中的22个蛋白点的主要功能涉及植物代谢、能量合成与分解、蛋白合成和胁迫防御。结合生理指标差异，发现与能量代谢相关的蛋白表达量差异确保‘RU9’在蛋白降解情况下，依然保持较高的光合反应、碳固定途径等重要生理活动。
Drought stress is a major factor limiting turfgrass growth. Understandingmechanisms for drought resistance in turfgrass species is important for selecting andbreeding drought-tolerant cultivars and for water conservation in turfgrass management.The objectives of the dissertation research were1) to determine genetic variations inmajor physiological traits associated with drought tolerance in a collection of tall fescue(Festuca arundinacea) germplasms collected from different geographic regionsworldwide;2) to examine genetic diversity and relationship among germplasm varying indrought tolerance via molecular marker classification and physiological/biochemical traitanalysis; and3) to determine metabolic processes controlled by specific proteinsimparting superior drought tolerance in tall fescue using germplams contrasting indrought tolerance with high genetic diversity and through two-dimensionalelelctropheresis and mass spectrometry. These objectives were addressed by conductingfour independent experiments. The main results of different experiments are summarizedas the following:
1Clustering analysis of different Tall Fescue genotypes for droughtresistance
Plants of44accessions of tall fescue were exposed to well-watered conditions ordrought stress for13day. Visual evaluation of turf quality, leaf relative water content,electrolyte leakage, chlorophyll content, and evapotranspiration rate demonstrated widephenotypic variation contributing to drought tolerance in this collection of tall fescuegermplasm. A hierarchical cluster analysis was conducted based on physiological traitsand among44tall fescue accessions. Among the44accessions,12accessions wereclassified as most tolerant,17as moderately tolerant, and15as susceptible. Among the44accessions,‘SH-3-AM-US’from America and‘PI598949-EU-IT’from Europe were genetically distinct but both were classified as drought-tolerant germplasm based onphysiological data. These two accessions could be used as potential parents in breedingdrought-tolerant tall fescue genotypes.
2Root antioxidant enzyme activities associated with drought tolerancein tall fescue
Two tall fescue genotypes ‘Focus’ and ‘PI150156’ contrasting in droughttolerance were examined in the greenhouse without irrigation for50days. Evaluation ofwhole-plant drought tolerance in different tall fescue genotypes was based on thevariations in turf quality (TQ) and leaf relative water content (RWC). Root physiologicaltraits, cell membrane stability (expressed as electrolyte leakage, EL), root viability (RV)and the activities of antioxidant enzymes, were determined to evaluate root activitiesassociated with drought tolerance. The results demonstrated that drought-tolerant traits ofleaves (TQ, RWC) were positively associated with root viability and superoxidedismutase (SOD) and catalase (CAT) activities while root membrane stability waspositively associated with root viability and activities of all four antioxidant enzymes(SOD, CAT, glutathione reductase and ascorbate peroxidase). This study suggested thatmaintaining high antioxidant enzyme activities of roots may help to maintain root cellmembrane integrity and root viability, and thus ultimately contribute to better droughttolerance of the whole plant.
3Water use, photosynthetic traits and root development associatedwith drought resistance in tall fescue genotypes
The characteristics of water use, photosynthesis, and root morphological features androot viability were examined and compared among different genotypes of tall fescuediffering in drought tolerance. Plants of ‘Focus’, PI150156’,‘RU5’,‘RU7’,‘RU9’,‘RU10’,‘RU18’,‘RU19’, and ‘RU21’ were exposed to drought stress in greenhouse andcontrolled-environment growth chambers. The results demonstrated that the superior turfperformance (higher TQ) and leaf hydration status (higher RWC) under drought stresswere associated with sustained stomatal opening (greater stomatal aperature and stomatal conductance) and water use rate (higher transpiration rate, evapotranspiration rate andwater use efficiency) and maintaining higher photosynthetic rate (Pn), as well as thedevelopment of root system with higher viability (lower root EL) and higher ability ofextraction water. In addition, maintaining photosynthesis under long-term drought stressdepended on larger stomatal aperture, but not stomatal density. Higher specific rootlength or finer roots was positively related to the maintenance of higher TQ and RWC indrought-tolerant genotypes. This study suggested that the combined traits of developingfine (small diameter) and viable roots (lower EL) and leaves with higher stomatalopening are critical for controlling photosynthesis and water use under drought stressconditions.
4. Identification of proteins associated with drought tolerance in tallfescue
Drought tolerant‘RU9’and drought sensitive‘RU18’were exposed to droughtstress without irrigation for12days. The TQ, RWC, Pn, photochemical efficiency (Fv/Fm)of two tall feacue genotypes decreased, and EL increased under drought stress. Two-dimensional electrophoresis analysis identified31protein spots differentially expressedin the two genotypes under drought stress compared to their respective well-wateredcontrol plants. Mass spectrophotometry analysis revealed that differential expression ofproteins with functions controlling carbon fixation and glycolytic energy metabolismcould at least partially account for genotypic variations and contributes to superiordrought tolerance in tall fescue.
1.不同高羊茅基因型抗旱性聚类分析
44份高羊茅基因型在温室干旱处理了13d。根据两次草坪质量（Turfquality, TQ）、叶片相对含水量（Relative water content, RWC）、电解液渗透率（Electric leakage）、叶绿素含量（chlorophyll content, Chl）、以及蒸散率（Evapotranspiration rate，ET）的试验结果，确定44份材料的抗旱性差异。聚类结果发现12份材料抗旱能力强，15份基因型对干旱敏感。在44份材料中，来自美洲的‘SH-3-AM-US’和欧洲的‘PI598949-EU-IT’抗旱能力均较强，但两者间的遗传差异较大。选择抗旱且有较大遗传差异的基因资源作为育种计划的候选亲本材料，将有助于培育抗旱性能强的品种。
2.根系抗氧化酶活性与高羊茅抗旱性关系的研究
抗旱基因型‘Focus’和旱敏感基因型‘PI150156’在温室干旱处理50d。测定TQ、RWC、EL、根系活力(Root Viability, RV)和根系抗氧化酶活性，判断根系抗氧化酶活性与高羊茅抗旱性的关系。TQ、RWC与RV、根系超氧化物歧化酶（Superoxide Dismutase, SOD）和过氧化氢酶（Catlase, CAT）酶活性变化成正相关关系；EL与RV、根系SOD和CAT酶活性成负相关关系，EL与以及根系谷胱甘肽还原酶（Glutathione Reductase, GR）和抗坏血酸还原酶（Ascorbate Peroxidase, APX）酶活性呈负相关性关系。结果说明维持根系底部抗氧化酶(SOD、CAT、GR和APX)活性能促进高羊茅在干旱胁迫下保持较高的草坪质量(用TQ衡量)、冠层水状态(用RWC衡量)、膜质稳定性(用EL衡量)和根系活力(RV)，提高高羊茅的抗旱性。
3.高羊茅水分代谢、光合特征及根系生长与高羊茅抗旱性关系的研究
试验以‘Focus’和‘PI150156’，7个高羊茅基因型(‘RU5’、‘RU7’、‘RU9’、‘RU10’、‘RU18’、‘RU19’和‘RU21’)为材料，分2个试验分别研究了干旱胁迫下冠层水分代谢指标、光合特征、以及根系形态和根系活力的变化。干旱后期，抗旱基因型的水分利用率（water use effciency, WUE）、蒸散率、以及冠层与空气温度差（canopy-air temperature, CTD）显著高于干旱敏感型；光合作用受到抑制，但抗旱基因型的净光合速率（net photosyntheticrate, Pn）、蒸腾速率（Transpiration rate, Tr）、气孔导度（stomatal conductance,gs）和气孔开度(stomatal aperture, SA)都显著高于干旱敏感基因型；抗旱基因型的深层根系活力（Root viability, RV）、根系干重(Total root weight)与长度(Total root length)，以及相对根茎（Specific root length, SRL）均显著高于干旱敏感基因型。结果发现优异的草坪质量、较高的叶片含水量与气孔开放程度（高气孔导度）、更有效的水分利用（高蒸散量和蒸腾速率有效降低了草坪冠层温度，高水分利用率提高了植物水分利用效率）、高光合效率、以及根系活力与根系吸水能力有关。另外，相对根茎值大的根系也是植物应对干旱的策略。
4.干旱胁迫对高羊茅干旱响应蛋白的影响
抗旱高羊茅‘RU9’和干旱敏感型‘RU18’干旱处理12d，研究了TQ、RWC、EL、Pn和光化学效率(photosynthesis efficiency, Fv/Fm)，以及蛋白表达谱的变化。试验选取干旱第12d与浇水对照的叶片，进行了全蛋白的提取和双向电泳分离。凝胶图像分析发现处理间、材料间共有31个差异蛋白点，质谱结果显示其中的22个蛋白点的主要功能涉及植物代谢、能量合成与分解、蛋白合成和胁迫防御。结合生理指标差异，发现与能量代谢相关的蛋白表达量差异确保‘RU9’在蛋白降解情况下，依然保持较高的光合反应、碳固定途径等重要生理活动。
Drought stress is a major factor limiting turfgrass growth. Understandingmechanisms for drought resistance in turfgrass species is important for selecting andbreeding drought-tolerant cultivars and for water conservation in turfgrass management.The objectives of the dissertation research were1) to determine genetic variations inmajor physiological traits associated with drought tolerance in a collection of tall fescue(Festuca arundinacea) germplasms collected from different geographic regionsworldwide;2) to examine genetic diversity and relationship among germplasm varying indrought tolerance via molecular marker classification and physiological/biochemical traitanalysis; and3) to determine metabolic processes controlled by specific proteinsimparting superior drought tolerance in tall fescue using germplams contrasting indrought tolerance with high genetic diversity and through two-dimensionalelelctropheresis and mass spectrometry. These objectives were addressed by conductingfour independent experiments. The main results of different experiments are summarizedas the following:
1Clustering analysis of different Tall Fescue genotypes for droughtresistance
Plants of44accessions of tall fescue were exposed to well-watered conditions ordrought stress for13day. Visual evaluation of turf quality, leaf relative water content,electrolyte leakage, chlorophyll content, and evapotranspiration rate demonstrated widephenotypic variation contributing to drought tolerance in this collection of tall fescuegermplasm. A hierarchical cluster analysis was conducted based on physiological traitsand among44tall fescue accessions. Among the44accessions,12accessions wereclassified as most tolerant,17as moderately tolerant, and15as susceptible. Among the44accessions,‘SH-3-AM-US’from America and‘PI598949-EU-IT’from Europe were genetically distinct but both were classified as drought-tolerant germplasm based onphysiological data. These two accessions could be used as potential parents in breedingdrought-tolerant tall fescue genotypes.
2Root antioxidant enzyme activities associated with drought tolerancein tall fescue
Two tall fescue genotypes ‘Focus’ and ‘PI150156’ contrasting in droughttolerance were examined in the greenhouse without irrigation for50days. Evaluation ofwhole-plant drought tolerance in different tall fescue genotypes was based on thevariations in turf quality (TQ) and leaf relative water content (RWC). Root physiologicaltraits, cell membrane stability (expressed as electrolyte leakage, EL), root viability (RV)and the activities of antioxidant enzymes, were determined to evaluate root activitiesassociated with drought tolerance. The results demonstrated that drought-tolerant traits ofleaves (TQ, RWC) were positively associated with root viability and superoxidedismutase (SOD) and catalase (CAT) activities while root membrane stability waspositively associated with root viability and activities of all four antioxidant enzymes(SOD, CAT, glutathione reductase and ascorbate peroxidase). This study suggested thatmaintaining high antioxidant enzyme activities of roots may help to maintain root cellmembrane integrity and root viability, and thus ultimately contribute to better droughttolerance of the whole plant.
3Water use, photosynthetic traits and root development associatedwith drought resistance in tall fescue genotypes
The characteristics of water use, photosynthesis, and root morphological features androot viability were examined and compared among different genotypes of tall fescuediffering in drought tolerance. Plants of ‘Focus’, PI150156’,‘RU5’,‘RU7’,‘RU9’,‘RU10’,‘RU18’,‘RU19’, and ‘RU21’ were exposed to drought stress in greenhouse andcontrolled-environment growth chambers. The results demonstrated that the superior turfperformance (higher TQ) and leaf hydration status (higher RWC) under drought stresswere associated with sustained stomatal opening (greater stomatal aperature and stomatal conductance) and water use rate (higher transpiration rate, evapotranspiration rate andwater use efficiency) and maintaining higher photosynthetic rate (Pn), as well as thedevelopment of root system with higher viability (lower root EL) and higher ability ofextraction water. In addition, maintaining photosynthesis under long-term drought stressdepended on larger stomatal aperture, but not stomatal density. Higher specific rootlength or finer roots was positively related to the maintenance of higher TQ and RWC indrought-tolerant genotypes. This study suggested that the combined traits of developingfine (small diameter) and viable roots (lower EL) and leaves with higher stomatalopening are critical for controlling photosynthesis and water use under drought stressconditions.
4. Identification of proteins associated with drought tolerance in tallfescue
Drought tolerant‘RU9’and drought sensitive‘RU18’were exposed to droughtstress without irrigation for12days. The TQ, RWC, Pn, photochemical efficiency (Fv/Fm)of two tall feacue genotypes decreased, and EL increased under drought stress. Two-dimensional electrophoresis analysis identified31protein spots differentially expressedin the two genotypes under drought stress compared to their respective well-wateredcontrol plants. Mass spectrophotometry analysis revealed that differential expression ofproteins with functions controlling carbon fixation and glycolytic energy metabolismcould at least partially account for genotypic variations and contributes to superiordrought tolerance in tall fescue.
引文
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