白三叶对干旱胁迫的适应性研究
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摘要
当前,环境恶化严重威胁人类的生存与发展,干旱是世界危害最为严重的灾害之一。近几年来由于园林绿化事业被日益重视,白三叶在园林绿化和水土保持方面起到越来越大的作用。白三叶根系不发达,对干旱胁迫很敏感。本项目以不同品种白三叶为供试植物,采用植物生态、生理及生物化学方法,研究白三叶对干旱胁迫的反应及反应的品种间差异,研究成果可为我国干旱半干旱地区人工草场建造、城市园林绿化以及恢复与重建退化生态系统提供理论依据和科学指导,也可为将来抗旱植物的培育与筛选提供理论参考。主要研究结果如下:
1品比试验
收集原产地不同的白三叶12个品种进行盆栽对比试验,观察其生长状况和耐旱情况。结果表明,不同品种白三叶的农艺性状有较大差异,其耐旱能力也各不相同,其中,‘碧盛’、‘草地胡以阿’、‘纳努克’、‘考拉’、‘海发’、‘拉丁诺’、‘百霸’的耐旱能力较强,‘缪克罗文’、‘盖达尔夫’、‘萨克雷门特’、‘泰坦’、‘瑞文德’的耐旱能力较差。从农艺性状方面来看,‘海发’、‘百霸’、‘考拉’、‘拉丁诺’、‘纳努克’、‘瑞文德’较适合于园林栽培。
2不同强度干旱胁迫对白三叶叶片光合作用和叶绿素荧光参数日变化的影响
以品种‘海发’为试材,研究了不同强度干旱胁迫对白三叶叶片光合作用和叶绿素荧光参数日变化的影响。结果表明,土壤相对含水量为75%和50%时,白三叶的光合参数日变化呈双峰趋势,叶绿素荧光参数受到的影响不大,说明白三叶在这两种水分条件下能够正常生长;土壤相对含水量为25%时,PSⅡ反应中心遭到破坏,原初光能转化效率(Fv/Fm)、PSⅡ电子传递量子产量(φPSⅡ)和光化学猝灭系数降低;净光合速率、蒸腾速率下降明显,日变化曲线表现为单峰特征;气孔导度显著降低,而胞间CO2浓度升高,非气孔限制成为Pn下降的主要原因;说明白三叶在重度干旱(HD)条件下,生理机能遭到破坏,不能正常生长。
3白三叶光合特性和根系空间分布对土壤水分梯度的动态响应
通过对两个品种白三叶‘海发’和‘瑞文德’盆栽试验,模拟3种不同的土壤水分状况(无水分胁迫:保持植株良好的水分供应;轻度胁迫:表层0cm～20cm土壤处于干旱状态;重度胁迫:表层0cm～20cm土壤处于极干旱状态,20cm～40cm土壤处于干旱状态)对白三叶光合作用和根系生长的影响。结果表明,当植株未遭受水分胁迫时,两个品种白三叶的光合作用和根系生长状况没有明显差异;当表层0cm～20cm处于干旱状态时,‘海发’在处理后期的净光合速率和水分利用效率高于对照,根系生长量增大,表现出适宜性调节作用,‘瑞文德’受到的影响不显著;当表层0cm～20cm处于极干旱、20cm～40cm处于干旱状态时,‘海发’在处理前期受到轻微影响,随后恢复正常状态,‘瑞文德’则受到较严重的影响。随着干旱程度的加深和时间的延长,白三叶的根冠比逐渐增大。与‘瑞文德’相比,在相同时期相同胁迫程度下,‘海发’的根冠比没有显著差异,但深根数量大大超过‘瑞文德’,因而,‘海发’的耐旱能力强于‘瑞文德’。
4持续干旱对白三叶渗调物质积累和水分利用的影响
土壤水分变化首先影响到生物量在植物不同器官的分配,即累积性的CO2净同化产物的分配,白三叶能够通过提高匍匐茎、根系的干物质分配比例来规避干旱。同时,在干旱胁迫发生时,植株内可溶性糖含量、脯氨酸含量都大幅度增加。抗旱品种通过增加可溶性糖、脯氨酸含量提高渗透势的能力显著大于不抗旱品种。在水分胁迫发生时,白三叶叶片相对含水量不断下降,甚至干枯或死亡;土壤水分变化还影响到白三叶根系在土壤中的生长状况,使不同层次土壤中的根量和根活力发生改变,从而改变了白三叶的吸水状况,抗旱性较强的品种通过增加深根数量提高对深层土壤水分的吸收能力显著大于抗旱性较差的品种。
5白三叶抗氧化清除系统对干旱胁迫的响应
在轻度水分胁迫下,白三叶幼苗可以通过提高酶活性来清除因胁迫产生的过量活性氧,减少氧化胁迫伤害,使MDA含量、膜透性降低。耐旱性不同的品种间差异不显著;在重度胁迫下,抗旱性差的品种酶活性在胁迫初期有一定程度的提高,但提高的幅度不足以维持活性氧产生与清除之间的平衡,使植株产生氧化伤害,最终导致酶活性下降。而抗旱性强的品种酶活性提高的幅度能够维持这种平衡,使植株在一定程度上避免了伤害。在干旱胁迫下,抗旱品种利用ASC、MDHAR、ASPX、SOD等的调节能力较不抗旱品种强,这表明在不同干旱时期白三叶体内起主要作用的抗氧化物质不同。
6干旱胁迫下白三叶的激素调节
在轻度干旱胁迫下,白三叶叶片、根系中的ABA含量都显著升高。在重度胁迫下,‘海发’和‘瑞文德’根系中的ABA含量都迅速升高,而在叶片中,‘瑞文德’的ABA含量升高后有迅速下降。干旱条件下,两种白三叶叶片内的CTK含量均下降,根系中CTK的表现则相反。轻度干旱胁迫条件下,‘海发’叶片和根系中的IAA含量均呈上升的趋势,而根系上升的幅度更大。‘瑞文德’根系中的IAA含量也上升,但上升幅度较小。在重度胁迫下,‘海发’叶片中的IAA含量前期上升,随后开始下降,而根系中的IAA含量始终保持上升趋势。‘瑞文德’叶片中IAA含量迅速上升,但很快即开始下降,根系中的IAA含量始终保持上升趋势。在轻度胁迫下,白三叶‘海发’叶片内的GA含量在胁迫期间变化不明显;在重度胁迫下,仅在胁迫后期有小幅升高。‘瑞文德’叶片中GA含量在轻度胁迫时缓慢升高,在重度胁迫时小幅升高后又表现为缓慢下降的趋势。在根系中,‘海发’GA含量随着胁迫时间的延长小幅上升,而‘瑞文德’在轻度胁迫时变化不明显,在重度胁迫时表现为下降的特征。干旱胁迫下四种激素之间协调的总趋势是向着气孔关闭,促进根系发展的方向进行,但不同抗旱强度的白三叶其内源激素的调节机制有区别。
7渗透胁迫对白三叶幼苗根系离子分泌和质膜ATP酶活性的影响
用PEG-6000渗透胁迫处理,白三叶根系质膜透性升高,抗旱品种‘海发’增幅小于不抗旱品种‘瑞文德’。根系分泌物中K+、Ca2+、H+的含量变化与根系质膜透性的变化呈显著正相关。水分胁迫早期,白三叶根系PM H+-ATPase活性升高,胁迫时间延长,不抗旱品种‘瑞文德’的PM H+-ATPase酶活性下降,但抗旱品种‘海发’的酶活性一直较高。PM Ca2+-ATPase酶的活性与PM H+-ATPase酶表现一致。说明PM Ca2+-ATPase酶和PM H+-ATPase酶活性与白三叶的抗旱性有关。
Currently, drought is one of the most serious environmental problems. And in arid andsemi-arid regions, drought is a major constraint imposed on plant survival and growth. In China, White clover (Trifolium repens L) has been planting as the pasture all along and plays more and more important role in city landscape and soil and water conservation in recently years. The shallow root system of white clover is sensitive to drought. Under dry condition,The root system of white clover is small, it is sensitive to aridity. Under the circumstance of aridity, the changes of growth and physiology and biochemistry character were more evident. The different varieties of White clover were used as the mode plants in the item, to study the reaction to the aridity of white clover and the discrepancy among different varieties by the methods of ecology, physiology and biochemistry. To provide the theory basis and scientific guidance for building the artificial pasture in the arid and semiarid zone, garden virescence in cities, and resume and reconstruction the degenerating ecology system. To provide the theory references for culturing and selecting the anti-aridity plants.
1. Comparing among different varieties of white clover
12 varieties of white clover were planted in the pots. To observe the growth status and drought endurance. It showed that the agricultural characters played clearly discrepancy among different varieties of white clover. Their capacities of drought endurance were also difference. Among which,‘Zapican’,‘Grasslands Huia’,‘Nanouk’,‘Korla’,‘Haifa’,‘Ladino’,‘Beber’showed stronger capacity of drought endurance than‘Milkanova’,‘Gandalf’,‘Sacremento’,‘Titan’,‘Rivendel’played a poor capacity of drought endurance. From the agricultural characters,‘Haifa’,‘Beber’,‘Korla’,‘Ladino’,‘Nanouk’,‘Rivendel’were capable of garden planting.
2. Effect of daily dynamics of photosynthesis and chlorophyll fluorescence parameters
in Trifolium repens leaves on the different drought stresses Daily dynamics of photosynthesis and chlorophyll fluorescence parameters in Trifolium repens cv. Haifa leaves under drought stress were studied. The results showed that under stress for 7 days, the curves of diurnal variation in Pn, Tr and Gs of Trifolium repens leaves were observed, and the chlorophyll fluorescence parameters such as conversion efficiency of primary light energy (Fv/Fm) of PSⅡ, the photochemical quenching coefficient (qP) and the quantum yield of PSⅡelectron transport (φPSⅡ) were decreased respectively when the relative soil moisture content was 25%. Nonstomatal limitation was the main reason responsible for Pn declined.
3. Effects of soil water gradient on photosynthesis and root spatial distribution in two Trifolium repens cultivars
The plant drought tolerance is one of the important parameters for selecting sustainable landscape plants. Trifolium repens Cultivars,‘Haifa’and‘Rivendel’, were studied for their photosynthesis and root spatial distribution under different drought stresses. The results showed that there were no differences in plant photosynthesis or root growth between the two cultivars under adequate water supply. Net photosynthetic rates and water use efficiency increased, and the root growth was enhanced for‘Haifa’with moderate drought stress in the 0 cm– 20 cm soil layer, but there was no significant change for‘Rivendel’. Under extreme drought condition in 0 cm– 20 cm soil, and drought condition in 20 cm– 40 cm soil, the cultivar‘Haifa’was affected only slightly at the first stage of the experiment and later recovered to a normal state, while‘Rivendel’was much more adversely effected. Degree and time of the moisture stress increased the root/shoot ratios gradually for plants of‘Rivendel’. Compared to‘Rivendel’, the root/shoot ratio for‘Haifa’did not increase significantly given the same experiment conditions as‘Haifa’, but there were more deeper roots for‘Haifa’than for‘Rivendel’. Therefore, it is believed that‘Haifa’was more drought resistant than‘Rivendel’.
4. Effects of continuous drought stress on osmotic adjustment substance and water metabolism of white clover
The effects of continuous drought stress on osmotic adjustment substance and water metabolism of white clover (Trifolium repens‘Haifa’) were studied. The result proved, leaf dry matter was lower in water - stressed plants than in irrigated plants, but stolon and root dry matter were higher. During drought time, the water - stress plants had higher soluble sugar and proline content than irrigated plants. Relative water content in leaves of white clovers declined gradually during drought stress compared with controls(with water supply during treatment), the leaves wilted and died at last. Root distribution changed with soil water decreased, but the response varied with different varieties, the droght resistant variety had higher root lenghth densities in deeper soil, so it had higher ability to absorb soil water.
5.Effects of drought stress on lipid peroxidation and clearing mechanism in white clovers leaves
The effects of drought stress on RPMP, MDA contents and the activities of MDHAR, DHAR, ASPX, GST, SOD, POD in white clovers leaves were studied. After submerging white clovers roots with PEG-6000 solution, drought stress increased MDA content and permeability of membrane in white clovers leaves. In slight drought stress, the white clovers could keep the balance of production and scavenging of active oxygen by heightening activities of the enzymes, and reduced the oxidic stress resulting from drought stress, At the same time reduced the content of MDA and permeability of cell membrane. In severe stress, activities of the enzymes in variety which was not resistant to drought increased at first, but they could not keep the balance of production and scavenging of active oxygen, white clovers were damaged. but the variety which was resistant to drought could keep the balance by increasing activities of the enzymes, then white clovers were not damaged.
6. The change endogenesis hormones in white clover under drought stress
The content of ABA and IAA in the leaves of two white clover increased at different extent and the content CTK and GA were all down trend under drought strss. The general trend of coordinatiion among four endogenesis hormones was closeing the stoma and promoting the growth of root. Whereas the coordinatin mechanism of different white clover germplasm resources was different. Under serious drought strss, drought-resistance white clover had higher coordination ability in reducing grow rate to ensure surviving to avoid the damage of lack of water effectively.
7. Effects of PEG stress on nutrient contents in root exudates and plasma membrane ATPase activity of white clovers
The plasma membrane (PM) permeability increased in white clover roots under PEG-6000 osmotic stress, and the amplitude of changes in drought-resistant cultivar was more sensitive than that in the drought-sensetive one. the contents of K+, Ca2+ and H+ in root exudates changed with the plasma membrane permeability. PM H+-ATPase activity increased at early stage under stress, but dropped at late stage under serious stress in drought-resistant cultivar. PM Ca2+-ATPase activity changed like PM H+-ATPase activity. These results show considerable correlation between the activities of PM H+-ATPase or PM Ca2+-ATPase and the drought tolerance of white clover.
1品比试验
收集原产地不同的白三叶12个品种进行盆栽对比试验,观察其生长状况和耐旱情况。结果表明,不同品种白三叶的农艺性状有较大差异,其耐旱能力也各不相同,其中,‘碧盛’、‘草地胡以阿’、‘纳努克’、‘考拉’、‘海发’、‘拉丁诺’、‘百霸’的耐旱能力较强,‘缪克罗文’、‘盖达尔夫’、‘萨克雷门特’、‘泰坦’、‘瑞文德’的耐旱能力较差。从农艺性状方面来看,‘海发’、‘百霸’、‘考拉’、‘拉丁诺’、‘纳努克’、‘瑞文德’较适合于园林栽培。
2不同强度干旱胁迫对白三叶叶片光合作用和叶绿素荧光参数日变化的影响
以品种‘海发’为试材,研究了不同强度干旱胁迫对白三叶叶片光合作用和叶绿素荧光参数日变化的影响。结果表明,土壤相对含水量为75%和50%时,白三叶的光合参数日变化呈双峰趋势,叶绿素荧光参数受到的影响不大,说明白三叶在这两种水分条件下能够正常生长;土壤相对含水量为25%时,PSⅡ反应中心遭到破坏,原初光能转化效率(Fv/Fm)、PSⅡ电子传递量子产量(φPSⅡ)和光化学猝灭系数降低;净光合速率、蒸腾速率下降明显,日变化曲线表现为单峰特征;气孔导度显著降低,而胞间CO2浓度升高,非气孔限制成为Pn下降的主要原因;说明白三叶在重度干旱(HD)条件下,生理机能遭到破坏,不能正常生长。
3白三叶光合特性和根系空间分布对土壤水分梯度的动态响应
通过对两个品种白三叶‘海发’和‘瑞文德’盆栽试验,模拟3种不同的土壤水分状况(无水分胁迫:保持植株良好的水分供应;轻度胁迫:表层0cm～20cm土壤处于干旱状态;重度胁迫:表层0cm～20cm土壤处于极干旱状态,20cm～40cm土壤处于干旱状态)对白三叶光合作用和根系生长的影响。结果表明,当植株未遭受水分胁迫时,两个品种白三叶的光合作用和根系生长状况没有明显差异;当表层0cm～20cm处于干旱状态时,‘海发’在处理后期的净光合速率和水分利用效率高于对照,根系生长量增大,表现出适宜性调节作用,‘瑞文德’受到的影响不显著;当表层0cm～20cm处于极干旱、20cm～40cm处于干旱状态时,‘海发’在处理前期受到轻微影响,随后恢复正常状态,‘瑞文德’则受到较严重的影响。随着干旱程度的加深和时间的延长,白三叶的根冠比逐渐增大。与‘瑞文德’相比,在相同时期相同胁迫程度下,‘海发’的根冠比没有显著差异,但深根数量大大超过‘瑞文德’,因而,‘海发’的耐旱能力强于‘瑞文德’。
4持续干旱对白三叶渗调物质积累和水分利用的影响
土壤水分变化首先影响到生物量在植物不同器官的分配,即累积性的CO2净同化产物的分配,白三叶能够通过提高匍匐茎、根系的干物质分配比例来规避干旱。同时,在干旱胁迫发生时,植株内可溶性糖含量、脯氨酸含量都大幅度增加。抗旱品种通过增加可溶性糖、脯氨酸含量提高渗透势的能力显著大于不抗旱品种。在水分胁迫发生时,白三叶叶片相对含水量不断下降,甚至干枯或死亡;土壤水分变化还影响到白三叶根系在土壤中的生长状况,使不同层次土壤中的根量和根活力发生改变,从而改变了白三叶的吸水状况,抗旱性较强的品种通过增加深根数量提高对深层土壤水分的吸收能力显著大于抗旱性较差的品种。
5白三叶抗氧化清除系统对干旱胁迫的响应
在轻度水分胁迫下,白三叶幼苗可以通过提高酶活性来清除因胁迫产生的过量活性氧,减少氧化胁迫伤害,使MDA含量、膜透性降低。耐旱性不同的品种间差异不显著;在重度胁迫下,抗旱性差的品种酶活性在胁迫初期有一定程度的提高,但提高的幅度不足以维持活性氧产生与清除之间的平衡,使植株产生氧化伤害,最终导致酶活性下降。而抗旱性强的品种酶活性提高的幅度能够维持这种平衡,使植株在一定程度上避免了伤害。在干旱胁迫下,抗旱品种利用ASC、MDHAR、ASPX、SOD等的调节能力较不抗旱品种强,这表明在不同干旱时期白三叶体内起主要作用的抗氧化物质不同。
6干旱胁迫下白三叶的激素调节
在轻度干旱胁迫下,白三叶叶片、根系中的ABA含量都显著升高。在重度胁迫下,‘海发’和‘瑞文德’根系中的ABA含量都迅速升高,而在叶片中,‘瑞文德’的ABA含量升高后有迅速下降。干旱条件下,两种白三叶叶片内的CTK含量均下降,根系中CTK的表现则相反。轻度干旱胁迫条件下,‘海发’叶片和根系中的IAA含量均呈上升的趋势,而根系上升的幅度更大。‘瑞文德’根系中的IAA含量也上升,但上升幅度较小。在重度胁迫下,‘海发’叶片中的IAA含量前期上升,随后开始下降,而根系中的IAA含量始终保持上升趋势。‘瑞文德’叶片中IAA含量迅速上升,但很快即开始下降,根系中的IAA含量始终保持上升趋势。在轻度胁迫下,白三叶‘海发’叶片内的GA含量在胁迫期间变化不明显;在重度胁迫下,仅在胁迫后期有小幅升高。‘瑞文德’叶片中GA含量在轻度胁迫时缓慢升高,在重度胁迫时小幅升高后又表现为缓慢下降的趋势。在根系中,‘海发’GA含量随着胁迫时间的延长小幅上升,而‘瑞文德’在轻度胁迫时变化不明显,在重度胁迫时表现为下降的特征。干旱胁迫下四种激素之间协调的总趋势是向着气孔关闭,促进根系发展的方向进行,但不同抗旱强度的白三叶其内源激素的调节机制有区别。
7渗透胁迫对白三叶幼苗根系离子分泌和质膜ATP酶活性的影响
用PEG-6000渗透胁迫处理,白三叶根系质膜透性升高,抗旱品种‘海发’增幅小于不抗旱品种‘瑞文德’。根系分泌物中K+、Ca2+、H+的含量变化与根系质膜透性的变化呈显著正相关。水分胁迫早期,白三叶根系PM H+-ATPase活性升高,胁迫时间延长,不抗旱品种‘瑞文德’的PM H+-ATPase酶活性下降,但抗旱品种‘海发’的酶活性一直较高。PM Ca2+-ATPase酶的活性与PM H+-ATPase酶表现一致。说明PM Ca2+-ATPase酶和PM H+-ATPase酶活性与白三叶的抗旱性有关。
Currently, drought is one of the most serious environmental problems. And in arid andsemi-arid regions, drought is a major constraint imposed on plant survival and growth. In China, White clover (Trifolium repens L) has been planting as the pasture all along and plays more and more important role in city landscape and soil and water conservation in recently years. The shallow root system of white clover is sensitive to drought. Under dry condition,The root system of white clover is small, it is sensitive to aridity. Under the circumstance of aridity, the changes of growth and physiology and biochemistry character were more evident. The different varieties of White clover were used as the mode plants in the item, to study the reaction to the aridity of white clover and the discrepancy among different varieties by the methods of ecology, physiology and biochemistry. To provide the theory basis and scientific guidance for building the artificial pasture in the arid and semiarid zone, garden virescence in cities, and resume and reconstruction the degenerating ecology system. To provide the theory references for culturing and selecting the anti-aridity plants.
1. Comparing among different varieties of white clover
12 varieties of white clover were planted in the pots. To observe the growth status and drought endurance. It showed that the agricultural characters played clearly discrepancy among different varieties of white clover. Their capacities of drought endurance were also difference. Among which,‘Zapican’,‘Grasslands Huia’,‘Nanouk’,‘Korla’,‘Haifa’,‘Ladino’,‘Beber’showed stronger capacity of drought endurance than‘Milkanova’,‘Gandalf’,‘Sacremento’,‘Titan’,‘Rivendel’played a poor capacity of drought endurance. From the agricultural characters,‘Haifa’,‘Beber’,‘Korla’,‘Ladino’,‘Nanouk’,‘Rivendel’were capable of garden planting.
2. Effect of daily dynamics of photosynthesis and chlorophyll fluorescence parameters
in Trifolium repens leaves on the different drought stresses Daily dynamics of photosynthesis and chlorophyll fluorescence parameters in Trifolium repens cv. Haifa leaves under drought stress were studied. The results showed that under stress for 7 days, the curves of diurnal variation in Pn, Tr and Gs of Trifolium repens leaves were observed, and the chlorophyll fluorescence parameters such as conversion efficiency of primary light energy (Fv/Fm) of PSⅡ, the photochemical quenching coefficient (qP) and the quantum yield of PSⅡelectron transport (φPSⅡ) were decreased respectively when the relative soil moisture content was 25%. Nonstomatal limitation was the main reason responsible for Pn declined.
3. Effects of soil water gradient on photosynthesis and root spatial distribution in two Trifolium repens cultivars
The plant drought tolerance is one of the important parameters for selecting sustainable landscape plants. Trifolium repens Cultivars,‘Haifa’and‘Rivendel’, were studied for their photosynthesis and root spatial distribution under different drought stresses. The results showed that there were no differences in plant photosynthesis or root growth between the two cultivars under adequate water supply. Net photosynthetic rates and water use efficiency increased, and the root growth was enhanced for‘Haifa’with moderate drought stress in the 0 cm– 20 cm soil layer, but there was no significant change for‘Rivendel’. Under extreme drought condition in 0 cm– 20 cm soil, and drought condition in 20 cm– 40 cm soil, the cultivar‘Haifa’was affected only slightly at the first stage of the experiment and later recovered to a normal state, while‘Rivendel’was much more adversely effected. Degree and time of the moisture stress increased the root/shoot ratios gradually for plants of‘Rivendel’. Compared to‘Rivendel’, the root/shoot ratio for‘Haifa’did not increase significantly given the same experiment conditions as‘Haifa’, but there were more deeper roots for‘Haifa’than for‘Rivendel’. Therefore, it is believed that‘Haifa’was more drought resistant than‘Rivendel’.
4. Effects of continuous drought stress on osmotic adjustment substance and water metabolism of white clover
The effects of continuous drought stress on osmotic adjustment substance and water metabolism of white clover (Trifolium repens‘Haifa’) were studied. The result proved, leaf dry matter was lower in water - stressed plants than in irrigated plants, but stolon and root dry matter were higher. During drought time, the water - stress plants had higher soluble sugar and proline content than irrigated plants. Relative water content in leaves of white clovers declined gradually during drought stress compared with controls(with water supply during treatment), the leaves wilted and died at last. Root distribution changed with soil water decreased, but the response varied with different varieties, the droght resistant variety had higher root lenghth densities in deeper soil, so it had higher ability to absorb soil water.
5.Effects of drought stress on lipid peroxidation and clearing mechanism in white clovers leaves
The effects of drought stress on RPMP, MDA contents and the activities of MDHAR, DHAR, ASPX, GST, SOD, POD in white clovers leaves were studied. After submerging white clovers roots with PEG-6000 solution, drought stress increased MDA content and permeability of membrane in white clovers leaves. In slight drought stress, the white clovers could keep the balance of production and scavenging of active oxygen by heightening activities of the enzymes, and reduced the oxidic stress resulting from drought stress, At the same time reduced the content of MDA and permeability of cell membrane. In severe stress, activities of the enzymes in variety which was not resistant to drought increased at first, but they could not keep the balance of production and scavenging of active oxygen, white clovers were damaged. but the variety which was resistant to drought could keep the balance by increasing activities of the enzymes, then white clovers were not damaged.
6. The change endogenesis hormones in white clover under drought stress
The content of ABA and IAA in the leaves of two white clover increased at different extent and the content CTK and GA were all down trend under drought strss. The general trend of coordinatiion among four endogenesis hormones was closeing the stoma and promoting the growth of root. Whereas the coordinatin mechanism of different white clover germplasm resources was different. Under serious drought strss, drought-resistance white clover had higher coordination ability in reducing grow rate to ensure surviving to avoid the damage of lack of water effectively.
7. Effects of PEG stress on nutrient contents in root exudates and plasma membrane ATPase activity of white clovers
The plasma membrane (PM) permeability increased in white clover roots under PEG-6000 osmotic stress, and the amplitude of changes in drought-resistant cultivar was more sensitive than that in the drought-sensetive one. the contents of K+, Ca2+ and H+ in root exudates changed with the plasma membrane permeability. PM H+-ATPase activity increased at early stage under stress, but dropped at late stage under serious stress in drought-resistant cultivar. PM Ca2+-ATPase activity changed like PM H+-ATPase activity. These results show considerable correlation between the activities of PM H+-ATPase or PM Ca2+-ATPase and the drought tolerance of white clover.
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