三叶草对干旱胁迫的反应及适应性研究
摘要
三叶草属植物根系发达,茎叶茂密、花色鲜艳,适应性较强,是半干旱地区重要的草坪和牧草,被世界各地广泛应用。在绿化美化城市,道路护坡、保护江河湖泊的堤坝方面显示出独有的作用。三叶草应用最多的是白三叶(Trifolium repens L.)、红三叶(Trifolium pratenseL.)、杂三叶(Trifolium hybridum L.)等几个种。本试验选用海法(Haifa)、瑞文德(Rivendel)、普通红三叶(Common red clover)和杂三叶(Alsike clover)为研究对象,通过盆栽控水,形成5个干旱梯度,在此干旱梯度的基础上,测定几种三叶草的形态指标、生理指标的动态变化规律,用PEG6000模拟干旱处理植株,测定RT2抗旱基因的表达差异,并比较干旱胁迫下4个草种抗旱性的强弱,旨在为选育抗旱性强的三叶草,在北方的大面积种植和干旱半干旱地区节水型草坪品种的选育提供重要的理论依据。测定指标包括：土壤相对含水量、叶片相对含水量、水势、离体叶片失水速度、游离脯氨酸含量、可溶性蛋白质含量、可溶性糖含量、过氧化物酶活性、叶绿素含量和相对电导率；气孔面积、气孔密度、气孔开关比和气孔导度；RT2基因的克隆和表达。研究结果及创新点如下：
1水分状况方面
随着土壤干旱时间的延长叶片相对含水量、叶水势和离体叶片失水速率都呈下降的趋势。并且在相同的水分梯度下,草种间的差异性不同。普红(Cr)的抗旱能力最强,海法(Hf)最弱,其它2个草种介于之间。4个草种的叶片相对含水量、叶水势,离体叶片失水速率与十壤相对含水量都呈极显著的正相关关系,草种不同相关性表现不同。说明土壤水分是引起三叶草植株水分变化的主要环境因子之一,当土壤水分降低时,植株中的水分会受到直接的影响。抗旱性强的草种水分变化小,保水能力强,弱的的草种则相反。
2渗透物质调节方面
随着土壤干旱时间的延长,三叶草叶片游离脯氨酸含量、可溶性蛋白质含量和可溶性糖含量都呈现出明显增加的趋势,在增加的过程中因草种不同和水分胁迫程度的不同而有所波动,草种间表现差异不同。但在处理期间三种物质总体的变化规律均是普红(Cr)和瑞文德(Rd)优于杂三叶(Ac)和海法(Hf)。试验结果说明：当土壤含水量降低时三种可溶性物质的含量都会增加。增加幅度越大的品种抗旱能力越强。
三种渗透调节物质与叶片相对含水量、叶水势呈负相关关系,各渗透调节物质之间呈正相关关系。说明：4种三叶草在水分胁迫下,均会调节增加体内的三种渗透调节物质的含量,以维持细胞内低的基础渗透式,来保证细胞吸水,进行正常的生命活动。调节能力的大小能在一定程度上表示植物抗旱能力的大小。渗透调节物质之间的相关性也说明三种物质在水分胁迫时的的渗透调节作用关系比较复杂,应该是协同的、综合的,各物质之间的相关性需做更深入的探讨。
3叶绿素、过氧化物酶活性和细胞质膜透性变化方面
随着干旱程度的加重叶片叶绿素的含量变化整体表现出慢-快-慢增加,最后减小的趋势。叶绿素含量与叶片含水量0-8d呈负相关关系,第8-16d表现正相关关系,各草种表现相关显著和极显著。试验结果显示：植物在水分胁迫时,叶绿素含量增加,轻度胁迫变化较快,中度胁迫变化较慢,而重度胁迫下,叶绿体遭到了破坏,叶绿素增加的速度明显减慢或低于开始处理水平。说明干旱前期叶片含水量相对较高,叶绿素仍处于合成状态,随着叶片含水量的减少,则呈现出了合成受阻而使合成的量减少。
4个草种POD活性整体呈现出先增加后减少的变化趋势。由此表明：4种三叶草在水分胁迫下,POD表现了很高的活性,轻度胁迫表现明显,中重度胁迫活性减小。叶片POD活性与叶片相对含水量、叶片水势0-8d呈负相关关系,处理8-16d呈正相关关系。
随着干旱程度的加重,4个草种的电导率都在不同程度上显著增加,前期较慢,后期较快。叶片电导率与叶片相对含水量、叶片水势呈负相关关系,与过氧化物酶活性呈止相关关系,并且均表现相关显著和极显著。从这一指标比较草种的抗旱顺序：普红(Cr)>瑞文德(Rd)>杂三叶(Ac)>海法(Hf)。
4气孔因素方面
干旱胁迫使三叶草气孔面积、开关比和导度减小,气孔密度增大。气孔面积轻度和中度干旱胁迫下降缓慢,重度胁迫下降较快,总体变化平缓；气孔密度增大速度平缓。开关比干旱前后期下降较快,中期缓慢；气孔导度干旱前期下降缓慢,后期迅速。气孔密度与叶片相对含水量和叶水势呈负相关；气孔开关比、气孔面积与叶水势、含水量呈正相关关系。气孔导度与叶片水势和土壤水分含量呈正相关关系。
5抗旱基因RT2的表达
干旱胁迫下普红(Cr)RT2基因的时间表达结果显示干旱胁迫可诱导三叶草RT2基因的表达,暗示三叶草的抗旱性可能和活性氧清除系统的相关基因的诱导表达存在密切关系。
在干旱处理24h时,不同草种间三叶草RT2基因的表达差异显著,在4个草种中,普红(Cr)和瑞文德(Rd)RT2基因的表达量显著高于海法(Hf)和杂三叶(Ac)。其中,普红(Cr)的RT2基因表达量分别是海法(Hf)和杂三叶(Ac)的2.5倍和2.8倍。结果表明,RT2基因在不同草种间的基因表达存在明显差异,耐旱性强的草种的RT2基因的表达量相对较高。显示了普红(Cr)和瑞文德(Rd)较海法(Hf)和杂三叶(Ac)具有较强的抗旱能力。
6品种间抗旱性比较
4种三叶草的抗旱顺序是普通红三叶(Cr)>瑞文德(Rd)>杂三叶(Ac)>海法(Hf)
7研究的创新点
1)对气孔动态变化与叶片水分状况在干旱胁迫下的相互关系进行综合研究,在三叶草上尚属首次。
2)对叶片水分状况、土壤水分状况变化与细胞渗透调节物质、细胞膜透性,保护酶活性,及其之间关系进行综合分析,在三叶草抗旱研究上非常全面。
3)首次在干旱条件下克隆了三叶草抗旱基因RT2,并比较其在草种间的表达情况,揭示了三叶草的抗旱性与细胞活性氧清除系统相关基因表达的密切关系。
Clovers with developed roots, dense stems and leaves, bright-coloured flowers, are the most important turf and forage grass species of the temperate regions, used around the world widely. Which show unique role in beautifying cities, roads, mixly protecting dams of rivers and lakes, ect. Especially for the cultivars of Trifolium repens L. Trifolium pratense L. Trifolium hybridum L. In this study, Haifa, Rivendel, Common red clover, Alsike clover as materials to be researched under drought conditions. Potted, with5water gradients, on the basis of drought gradient, measured the dynamic changes of morphological and physiological indexes of the clover. PEG6000treated plants, determined the expression difference of RT2drought-resistant, compared drought-resistant ability of different cultivars under drought stress. The purpose of this study is to provide important theoretical basis on selecting drought tolerance cultivars and cultivating clovers in large areas and in the arid and semi-arid areas. Measurements include: relative water content of soil, leaf relative water content, leaf water potential, leaf free proline content, leaf leaf soluble protein content, leaf soluble sugar content, leaf peroxidase activities, leaf chlorophyll content and leaf relative conductivity rate, stomatal areas、stomatal densities、stomatal opening and closing ratio、stomatal conductance, the cloning and expression of RT2. The results and the innovation were as follows:
1Moisture aspects:With the lengthening of stress durations, the trends of the RWC, water potential of leaves, lost water rate of cutting leaves decreased, and appeared differences among clovers in the same moisture gradient. Common red clover was strongest and Haifa was weakest in drought resistance, the others were between them. The correlation was very significant positive between3moisture indexes and soil relative water content of4clovers. The correlation was different among clovers. Showed:soil moisture was a main environmental factor, which caused water changes of clovers. When the soil moisture decreased, plant moisture would be directly affected. The clovers of drought-resistance changed alittle in moisture.
2Osmoregulation aspects:with the lengthening of stress duration, free proline contents, soluble protein contents and soluble sugar contents of leaves were presenting obvious trend of increase. They were of different fluctuation due to the different clovers and the degree of water stress during the process. Differences were significant among clovers. But The whole changes rules of three substances were Common red clover> Rivendel> Alsike clover> Haifa during the process. The test results showed:when the soil moisture reducing, the contents of three kinds of soluble substances increased, the more increased, more stronger in drought-resistance.
From correlationship between osmoregulation substances and RWC, water potential analysis, three kinds of substances were negatively correlated between RWC and water potential, positive correlativity among three substances in the whole process. T he results showed:4kinds of clovers would increased three osmoregulation substances contents under water stress to ensure to maintain lowly basic penetration of cells, water absorption, and activities of cells, that related to the cultivars drought-resistant ability of a plant. The correlationship amang osmoregulation substances in different cultivars showed the relationship was coordinated and integrated.
3The contents of chlorophyll, POD and cytoplasmic membrane permeability: With the increasing of drought stress, chlorophyll contents of leaves increased by slow-fast-slow tendency first, decreased at last. The chlorophyll contents and the relative water contents of leaves negatively correlated in the0d-8d, positive correlation in the8d-16d.
The activities of POD in4clovers increased first and then decreaced at last during the treatment process. POD showed high activity in4clovers under water stress, the performance was obvious under mild stress, but the activities decreaced from moderate to severe stress. Activities of PODwere negatively correlated between RWC and water potential in the0d-8d, positive correlation in the8d-16d. The activities of POD in clovers were regulated by the changes of leaf water contents greatly, increased in the early stage of drought, with the increased stress, the activities decreased. The protecting cell membrane abilities of POD were limited, when clovers were under drought stress.
With the increasing degree of drought stress, the conductivity of4clovers increased significantly, increased slowly in the mild stress, and fast at the late stage, and showed negative correlation between the RWC, water potential of leaves, positive correlated between the activities of POD, and related significant and a very significant. The drought resistant orders:Common red clover> Rivendel> Alsike clover> Haifa.
4Stomatal factors:Stomatal area, opening and closing ratio and stomatal conductance decreased under drought stress, but stotnata density increased. Stomatal area declined slowly in the mild and moderate drought stress, declined fast in the severe stress. Stomatal density increased gently. Stomatal opening and closing ratio declined faster in the first and last stage s, slowly in the interim; Stomatal conductance declined slowly in the first stage, but rapidly in the later stage. Stomata density showed negative correlation between the RWC, water potential of leaves. Stomatal areas, opening and closing ratio correlated positively between the RWC, water potential of leaves. Stomatal conductance correlated positively between water potential of leavesN Relative water contents of soil.
5Drought-resistant gene RT2expression:analyzed the changes of RT2expression in Common red clover with the stressed point. The results showed that drought stress could induce the gene RT2expression of Common red clover, that suggested:the drought resistance of clovers might associate with the gene induction and expression of scavenging system of active oxygen.
With24h of drought stress, the RT2expression was significant difference among4clovers. RT2expression of Common red clover and Rivendel were significantly higher than those of Alsike clover and Haifa. The results showed that the RT2expression was different among different clovers, the aboundances of RT2expression were relatively higher in stronger drought-resistant clovers. Common red clover and Rivendel displayed stronger tolerance to drought than Alsike clover and Haifa.
6Drought resistance comparison in different cultivars:comparatively analyzed the changes of drought-resistant indexes in clovers under drought stress, the drought-resistance orders:Common red clover> Rivendel> Alsike clover> Haifa.
7Innovation points of research:
1) It was the first that researched comprehensivly stomatal dynamic changes and leaf water status under drought stress in the relationship in the clovers
2) It was very comprehensive research on leaf water status, soil moisture status change, cell osmotic regulation substances, cell membrane permeability, protective enzyme activity, and the relationship among them analysised comprehensivly in clover drought resistance
3) For the first time, cloned clover RT2under drought condition and compared their expression, revealsed:the close relationship between the drought resistance in clovers and the expression of active oxygen removal system geneof cells
1水分状况方面
随着土壤干旱时间的延长叶片相对含水量、叶水势和离体叶片失水速率都呈下降的趋势。并且在相同的水分梯度下,草种间的差异性不同。普红(Cr)的抗旱能力最强,海法(Hf)最弱,其它2个草种介于之间。4个草种的叶片相对含水量、叶水势,离体叶片失水速率与十壤相对含水量都呈极显著的正相关关系,草种不同相关性表现不同。说明土壤水分是引起三叶草植株水分变化的主要环境因子之一,当土壤水分降低时,植株中的水分会受到直接的影响。抗旱性强的草种水分变化小,保水能力强,弱的的草种则相反。
2渗透物质调节方面
随着土壤干旱时间的延长,三叶草叶片游离脯氨酸含量、可溶性蛋白质含量和可溶性糖含量都呈现出明显增加的趋势,在增加的过程中因草种不同和水分胁迫程度的不同而有所波动,草种间表现差异不同。但在处理期间三种物质总体的变化规律均是普红(Cr)和瑞文德(Rd)优于杂三叶(Ac)和海法(Hf)。试验结果说明：当土壤含水量降低时三种可溶性物质的含量都会增加。增加幅度越大的品种抗旱能力越强。
三种渗透调节物质与叶片相对含水量、叶水势呈负相关关系,各渗透调节物质之间呈正相关关系。说明：4种三叶草在水分胁迫下,均会调节增加体内的三种渗透调节物质的含量,以维持细胞内低的基础渗透式,来保证细胞吸水,进行正常的生命活动。调节能力的大小能在一定程度上表示植物抗旱能力的大小。渗透调节物质之间的相关性也说明三种物质在水分胁迫时的的渗透调节作用关系比较复杂,应该是协同的、综合的,各物质之间的相关性需做更深入的探讨。
3叶绿素、过氧化物酶活性和细胞质膜透性变化方面
随着干旱程度的加重叶片叶绿素的含量变化整体表现出慢-快-慢增加,最后减小的趋势。叶绿素含量与叶片含水量0-8d呈负相关关系,第8-16d表现正相关关系,各草种表现相关显著和极显著。试验结果显示：植物在水分胁迫时,叶绿素含量增加,轻度胁迫变化较快,中度胁迫变化较慢,而重度胁迫下,叶绿体遭到了破坏,叶绿素增加的速度明显减慢或低于开始处理水平。说明干旱前期叶片含水量相对较高,叶绿素仍处于合成状态,随着叶片含水量的减少,则呈现出了合成受阻而使合成的量减少。
4个草种POD活性整体呈现出先增加后减少的变化趋势。由此表明：4种三叶草在水分胁迫下,POD表现了很高的活性,轻度胁迫表现明显,中重度胁迫活性减小。叶片POD活性与叶片相对含水量、叶片水势0-8d呈负相关关系,处理8-16d呈正相关关系。
随着干旱程度的加重,4个草种的电导率都在不同程度上显著增加,前期较慢,后期较快。叶片电导率与叶片相对含水量、叶片水势呈负相关关系,与过氧化物酶活性呈止相关关系,并且均表现相关显著和极显著。从这一指标比较草种的抗旱顺序：普红(Cr)>瑞文德(Rd)>杂三叶(Ac)>海法(Hf)。
4气孔因素方面
干旱胁迫使三叶草气孔面积、开关比和导度减小,气孔密度增大。气孔面积轻度和中度干旱胁迫下降缓慢,重度胁迫下降较快,总体变化平缓；气孔密度增大速度平缓。开关比干旱前后期下降较快,中期缓慢；气孔导度干旱前期下降缓慢,后期迅速。气孔密度与叶片相对含水量和叶水势呈负相关；气孔开关比、气孔面积与叶水势、含水量呈正相关关系。气孔导度与叶片水势和土壤水分含量呈正相关关系。
5抗旱基因RT2的表达
干旱胁迫下普红(Cr)RT2基因的时间表达结果显示干旱胁迫可诱导三叶草RT2基因的表达,暗示三叶草的抗旱性可能和活性氧清除系统的相关基因的诱导表达存在密切关系。
在干旱处理24h时,不同草种间三叶草RT2基因的表达差异显著,在4个草种中,普红(Cr)和瑞文德(Rd)RT2基因的表达量显著高于海法(Hf)和杂三叶(Ac)。其中,普红(Cr)的RT2基因表达量分别是海法(Hf)和杂三叶(Ac)的2.5倍和2.8倍。结果表明,RT2基因在不同草种间的基因表达存在明显差异,耐旱性强的草种的RT2基因的表达量相对较高。显示了普红(Cr)和瑞文德(Rd)较海法(Hf)和杂三叶(Ac)具有较强的抗旱能力。
6品种间抗旱性比较
4种三叶草的抗旱顺序是普通红三叶(Cr)>瑞文德(Rd)>杂三叶(Ac)>海法(Hf)
7研究的创新点
1)对气孔动态变化与叶片水分状况在干旱胁迫下的相互关系进行综合研究,在三叶草上尚属首次。
2)对叶片水分状况、土壤水分状况变化与细胞渗透调节物质、细胞膜透性,保护酶活性,及其之间关系进行综合分析,在三叶草抗旱研究上非常全面。
3)首次在干旱条件下克隆了三叶草抗旱基因RT2,并比较其在草种间的表达情况,揭示了三叶草的抗旱性与细胞活性氧清除系统相关基因表达的密切关系。
Clovers with developed roots, dense stems and leaves, bright-coloured flowers, are the most important turf and forage grass species of the temperate regions, used around the world widely. Which show unique role in beautifying cities, roads, mixly protecting dams of rivers and lakes, ect. Especially for the cultivars of Trifolium repens L. Trifolium pratense L. Trifolium hybridum L. In this study, Haifa, Rivendel, Common red clover, Alsike clover as materials to be researched under drought conditions. Potted, with5water gradients, on the basis of drought gradient, measured the dynamic changes of morphological and physiological indexes of the clover. PEG6000treated plants, determined the expression difference of RT2drought-resistant, compared drought-resistant ability of different cultivars under drought stress. The purpose of this study is to provide important theoretical basis on selecting drought tolerance cultivars and cultivating clovers in large areas and in the arid and semi-arid areas. Measurements include: relative water content of soil, leaf relative water content, leaf water potential, leaf free proline content, leaf leaf soluble protein content, leaf soluble sugar content, leaf peroxidase activities, leaf chlorophyll content and leaf relative conductivity rate, stomatal areas、stomatal densities、stomatal opening and closing ratio、stomatal conductance, the cloning and expression of RT2. The results and the innovation were as follows:
1Moisture aspects:With the lengthening of stress durations, the trends of the RWC, water potential of leaves, lost water rate of cutting leaves decreased, and appeared differences among clovers in the same moisture gradient. Common red clover was strongest and Haifa was weakest in drought resistance, the others were between them. The correlation was very significant positive between3moisture indexes and soil relative water content of4clovers. The correlation was different among clovers. Showed:soil moisture was a main environmental factor, which caused water changes of clovers. When the soil moisture decreased, plant moisture would be directly affected. The clovers of drought-resistance changed alittle in moisture.
2Osmoregulation aspects:with the lengthening of stress duration, free proline contents, soluble protein contents and soluble sugar contents of leaves were presenting obvious trend of increase. They were of different fluctuation due to the different clovers and the degree of water stress during the process. Differences were significant among clovers. But The whole changes rules of three substances were Common red clover> Rivendel> Alsike clover> Haifa during the process. The test results showed:when the soil moisture reducing, the contents of three kinds of soluble substances increased, the more increased, more stronger in drought-resistance.
From correlationship between osmoregulation substances and RWC, water potential analysis, three kinds of substances were negatively correlated between RWC and water potential, positive correlativity among three substances in the whole process. T he results showed:4kinds of clovers would increased three osmoregulation substances contents under water stress to ensure to maintain lowly basic penetration of cells, water absorption, and activities of cells, that related to the cultivars drought-resistant ability of a plant. The correlationship amang osmoregulation substances in different cultivars showed the relationship was coordinated and integrated.
3The contents of chlorophyll, POD and cytoplasmic membrane permeability: With the increasing of drought stress, chlorophyll contents of leaves increased by slow-fast-slow tendency first, decreased at last. The chlorophyll contents and the relative water contents of leaves negatively correlated in the0d-8d, positive correlation in the8d-16d.
The activities of POD in4clovers increased first and then decreaced at last during the treatment process. POD showed high activity in4clovers under water stress, the performance was obvious under mild stress, but the activities decreaced from moderate to severe stress. Activities of PODwere negatively correlated between RWC and water potential in the0d-8d, positive correlation in the8d-16d. The activities of POD in clovers were regulated by the changes of leaf water contents greatly, increased in the early stage of drought, with the increased stress, the activities decreased. The protecting cell membrane abilities of POD were limited, when clovers were under drought stress.
With the increasing degree of drought stress, the conductivity of4clovers increased significantly, increased slowly in the mild stress, and fast at the late stage, and showed negative correlation between the RWC, water potential of leaves, positive correlated between the activities of POD, and related significant and a very significant. The drought resistant orders:Common red clover> Rivendel> Alsike clover> Haifa.
4Stomatal factors:Stomatal area, opening and closing ratio and stomatal conductance decreased under drought stress, but stotnata density increased. Stomatal area declined slowly in the mild and moderate drought stress, declined fast in the severe stress. Stomatal density increased gently. Stomatal opening and closing ratio declined faster in the first and last stage s, slowly in the interim; Stomatal conductance declined slowly in the first stage, but rapidly in the later stage. Stomata density showed negative correlation between the RWC, water potential of leaves. Stomatal areas, opening and closing ratio correlated positively between the RWC, water potential of leaves. Stomatal conductance correlated positively between water potential of leavesN Relative water contents of soil.
5Drought-resistant gene RT2expression:analyzed the changes of RT2expression in Common red clover with the stressed point. The results showed that drought stress could induce the gene RT2expression of Common red clover, that suggested:the drought resistance of clovers might associate with the gene induction and expression of scavenging system of active oxygen.
With24h of drought stress, the RT2expression was significant difference among4clovers. RT2expression of Common red clover and Rivendel were significantly higher than those of Alsike clover and Haifa. The results showed that the RT2expression was different among different clovers, the aboundances of RT2expression were relatively higher in stronger drought-resistant clovers. Common red clover and Rivendel displayed stronger tolerance to drought than Alsike clover and Haifa.
6Drought resistance comparison in different cultivars:comparatively analyzed the changes of drought-resistant indexes in clovers under drought stress, the drought-resistance orders:Common red clover> Rivendel> Alsike clover> Haifa.
7Innovation points of research:
1) It was the first that researched comprehensivly stomatal dynamic changes and leaf water status under drought stress in the relationship in the clovers
2) It was very comprehensive research on leaf water status, soil moisture status change, cell osmotic regulation substances, cell membrane permeability, protective enzyme activity, and the relationship among them analysised comprehensivly in clover drought resistance
3) For the first time, cloned clover RT2under drought condition and compared their expression, revealsed:the close relationship between the drought resistance in clovers and the expression of active oxygen removal system geneof cells
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