水孔蛋白与植物对干旱和低温胁迫响应关系的研究
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摘要
水孔蛋白(Aquaporin)属于跨膜通道MIP(Membrane Intrinsic Protein)大家族,是一种能介导水分跨膜运输的功能性的膜通道蛋白,它广泛地存在于从微生物到高等动物和植物的各种生物体中。水孔蛋白在转录、翻译、翻译后修饰等各个水平上受到调控。水孔蛋白的发现对植物生理学是一个概念性的发展。
在本研究中,我们就稻(Oryza sativa)水孔蛋白对干旱和低温的响应关系展开探讨,实验主要取得如下结果:
(1)通过对一个水稻品种(秀水63)和一个陆稻品种(中旱3号)在12%PEG(聚乙二醇) 2000(?0.45 MPa)渗透胁迫处理7?8天后的水分生理比较,来研究稻的抗旱生理机制。渗透胁迫处理后,两个品种均表现生长缓慢,植株矮小,新叶叶尖卷曲甚至干枯,但中旱3号要比秀水63表现得更为严重。相对秀水63,相同强度的渗透胁迫处理对中旱3号根系和地上部的生长均有较强的抑制作用。在渗透胁迫处理后,两个品种的叶子的相对含水量和水势都有所下降,但下降程度中旱3号强于秀水63。渗透胁迫处理使秀水63叶的渗透势显著下降,但中旱3号没有明显变化,同时秀水63的叶膨压高于中旱3号,说明秀水63可能具有更强的渗透调节能力。中旱3号在渗透胁迫7?8天后的气孔导度和蒸腾速率下降显著,但秀水63却变化不大。中旱3号和秀水63在渗透胁迫后的根水导度都显著下降。通过对不定根的徒手切片的荧光显微镜检测,两个品种的不定根均未表现出在渗透胁迫后内皮层和外皮层的木栓化增加。胁迫处理能使中旱3号和秀水63的根水导降显著下降。汞离子是许多水孔蛋白的抑制剂,通过汞离子抑制实验可以初步研究水孔蛋白的功能。在未受胁迫时,汞处理能使中旱3号和秀水63的根水导度显著下降;在胁迫处理后,中旱3号根水导度对汞离子的敏感性更高,而秀水63的根水导度对汞离子的敏感性却几乎消失。综合以上结果,水稻和陆稻可能采取不同的机制抵御干旱,水稻以耐旱为主(如渗透调节),陆稻以避旱为主(如增加根系吸水能力、减少水分散失),水孔蛋白则可能参与了陆稻的避旱过程。
(2)为了进一步确证水孔蛋白在植物避旱中的作用,我们将稻水孔蛋白RWC3基因遗传转化一个我们认为避旱机制较弱的水稻品种(中花11号)中,然后观测转基因植株在PEG诱导的干旱条件下的生理表型。在本研究中,首先构建SWPA2::RWC3结构,将其构入双元载体pCAMBIA 1301中,SWPA2是来自于红薯(Ipomoea batatas)的逆境
Aquaporins, which show the ability to facilitate passive exchange of water acrossmembranes, belong to a high conserved membrane protein family MIP (Major IntrinsicProtein). In plants, the activities of aquaporins can be controlled at transcription, translationand post-translation modification levels. The discovery of aquaporins in plants has resulted ina paradigm shift in plant physiology.
In this study, we focused on the role of rice aquaporins when respond to drought andchilling stress. The main results are as follows:
(1) The mechanism of drought tolerance was studied by comparing water relationsbetween an upland rice cultivar (Zhonghan 3) and a lowland rice cultivar (Xiushui 63) duringthe water deficit mediated by PEG. After 7?8 days with osmotic stress treatment, bothcultivars showed retardation of plant growth, dwarf plants and young leaf rolling or evendrying. These symptoms were more serious in Zhonghan 3 than Xiushui 63. The osmoticstress greatly retarded the growth of roots and aerial parts in Zhonghan 3 than Xiushui 63.After initiation of the osmotic stress, relative water content (RWC) and water potential ofleaves decreased significantly in both cultivars, and Xiushiu 63 had higher leaf RWC andwater potential than Zhonghan 3. Xiushui 63 probably showed higher osmotic adjustmentbecause Xiushui 63 had greater drop of leaf osmotic potential and higher turgor compared toZhonghan 3 when subjected to the osmotic stress. After initiation of the osmotic stress,Zhonghan 3 showed decreased transpiration rate and stomata conductivity, while they wereunchanged in Xiushiu 63. Root hydraulic conductivity (Lp) decreased significantly in bothcultivars after initiation of the osmotic stress. Moreover, by view of the freehandcross-sections of adventitious roots under a fluorescent microscope, no any differentlignifications at exodermis or endodermis in the roots of both cultivars was observed.Mercurial compounds are widely used to study the function of aquaporins because they are
inhibitors for aquaporins. Mercury treatment greatly decreased root hydraulic conductivity ofnon-stressed plants in both cultivars. However, during the osmotic stress, mercury showedmore serious inhibition effects in Zhonghan 3, while showed alleviating inhibition effects inXiushiu 63. To summarize, all these results suggested that upland rice and lowland mobilizeddistinct mechanism to resist drought, i.e. upland rice was drought avoidance (e.g. increasingroot water uptake and decreasing water evaporation) while lowland rice was drought tolerance(e.g. osmotic adjustment). Aquaporins might play role in drought avoidance of upland rice.(2) To demonstrate the role of aquporins in plant drought avoidance in fine detail,aquaporin RWC3 from rice (Oryza sativa L.) was transferred into a lowland rice cultivars(Zhonghua 11), which was thought to be weak in drought avoidance, and consequently somephysiological parameters were measured during the water deficit mediated by PEG.SWPA2::RWC3 was constructed into a binary plasmid pCAMBIA 1301, and was transferred tolowland rice mediated by Agrobacterium tumefaciens. SWPA2 promoter, which was isolatedin sweetpotato, was a stress inducible promoter. By the assays of GUS histochemical staining,PCR and PCR-Dig-ELISA, the presence of SWPA2::RWC3 was confirmed in the lowland rice.By RT-PCR and Western blot analysis, it was found that SWPA2 promoter-RWC3 transgenicplant and WT (wild type) had similar RWC3 expression levels under normal condition,nevertheless, under water deficit mediated by 20% PEG, the transgenic plant had higherRWC3 expression levels than WT, suggesting that RWC3 expression in transgenic plants couldbe induced by drought. However, no visible phenotypic difference was found between thetransgenic plants and the WT plants, either under normal condition or during water deficit.After the water deficit, SWPA2 promoter-RWC3 transgenic plant showed higher root hydraulicconductivity and leaf water potential than WT, and the decrease of relative accumulativetranspiration rate during the water deficit was alleviated in the transgenic plant. These resultswere summarized that compared to WT, SWPA2 promoter-RWC3 transgenic plant wasendowed with higher water transport of roots and improved water status under water deficit,suggesting aquaporin RWC3 probably play a role during drought avoidance of rice(3) Plant chilling injury was partially caused by water imbalance between water uptakein root and transpiration in leaf. In this study, two rice cultivars (Somewake and Wasetoitsue)were exposed to 7 ± 1°C for up to 24 h and following temperature recovery (28 ± 1°C) for 3
h. Throughout the chilling stress, Wasetoitsu showed more serious injury symptoms thanSomewake. By measuring the relative electrical conductivity of rice leaves, Wasetoitsu hadhigher membrane electrolyte leakage than Somewake at the low temperature. These resultsindicated that Wasetoitsue was a less chilling tolerant cultivar compared to Somewake. Thetwo cultivars were in different water status, i.e. Wasetoitsu had lower leaf RWC thanSomewake in the duration of chilling, although both cultivars had decreasing RWC at thesame time RWC. In both cultivars, accumulative transpiration rate and root Lp decreasedobviously during the chilling treatment, but the decrease of Lp was more rapid and acute thanthat of accumulative transpiration rate, suggesting that the water imbalance occurred betweenthe water uptake of root and the water evaporation of aerial parts. These physiologicalobservations prompted us to investigate the expression of water channel proteins. At mRNAlevel, three rice aquaporin genes, RWC1、OsPIP2a and RWC3 showed the similar response tothe chilling treatment in both cultivars, i.e. remained approximate constant within the 9 h withchilling treatment while were decreased after 24 h with chilling treatment. The analysis ofWestern blot with antibody against PIP1 was consistent with the mRNA expression. In bothcultivars, PIP1 protein decreased when the plants were subjected to the chilling treatment for24 h. As we know, this is the first report on aquaporin responses to chilling stress in plantroots. Surprisingly, the temperature recovery for 3 h following chilling treatment resulted incontinued decreases of root PIPs mRNA expression. Aquaporins might be related to ricechilling adaptation but not a key factor of chilling resistance at the early stage of the chillingstress.
在本研究中,我们就稻(Oryza sativa)水孔蛋白对干旱和低温的响应关系展开探讨,实验主要取得如下结果:
(1)通过对一个水稻品种(秀水63)和一个陆稻品种(中旱3号)在12%PEG(聚乙二醇) 2000(?0.45 MPa)渗透胁迫处理7?8天后的水分生理比较,来研究稻的抗旱生理机制。渗透胁迫处理后,两个品种均表现生长缓慢,植株矮小,新叶叶尖卷曲甚至干枯,但中旱3号要比秀水63表现得更为严重。相对秀水63,相同强度的渗透胁迫处理对中旱3号根系和地上部的生长均有较强的抑制作用。在渗透胁迫处理后,两个品种的叶子的相对含水量和水势都有所下降,但下降程度中旱3号强于秀水63。渗透胁迫处理使秀水63叶的渗透势显著下降,但中旱3号没有明显变化,同时秀水63的叶膨压高于中旱3号,说明秀水63可能具有更强的渗透调节能力。中旱3号在渗透胁迫7?8天后的气孔导度和蒸腾速率下降显著,但秀水63却变化不大。中旱3号和秀水63在渗透胁迫后的根水导度都显著下降。通过对不定根的徒手切片的荧光显微镜检测,两个品种的不定根均未表现出在渗透胁迫后内皮层和外皮层的木栓化增加。胁迫处理能使中旱3号和秀水63的根水导降显著下降。汞离子是许多水孔蛋白的抑制剂,通过汞离子抑制实验可以初步研究水孔蛋白的功能。在未受胁迫时,汞处理能使中旱3号和秀水63的根水导度显著下降;在胁迫处理后,中旱3号根水导度对汞离子的敏感性更高,而秀水63的根水导度对汞离子的敏感性却几乎消失。综合以上结果,水稻和陆稻可能采取不同的机制抵御干旱,水稻以耐旱为主(如渗透调节),陆稻以避旱为主(如增加根系吸水能力、减少水分散失),水孔蛋白则可能参与了陆稻的避旱过程。
(2)为了进一步确证水孔蛋白在植物避旱中的作用,我们将稻水孔蛋白RWC3基因遗传转化一个我们认为避旱机制较弱的水稻品种(中花11号)中,然后观测转基因植株在PEG诱导的干旱条件下的生理表型。在本研究中,首先构建SWPA2::RWC3结构,将其构入双元载体pCAMBIA 1301中,SWPA2是来自于红薯(Ipomoea batatas)的逆境
Aquaporins, which show the ability to facilitate passive exchange of water acrossmembranes, belong to a high conserved membrane protein family MIP (Major IntrinsicProtein). In plants, the activities of aquaporins can be controlled at transcription, translationand post-translation modification levels. The discovery of aquaporins in plants has resulted ina paradigm shift in plant physiology.
In this study, we focused on the role of rice aquaporins when respond to drought andchilling stress. The main results are as follows:
(1) The mechanism of drought tolerance was studied by comparing water relationsbetween an upland rice cultivar (Zhonghan 3) and a lowland rice cultivar (Xiushui 63) duringthe water deficit mediated by PEG. After 7?8 days with osmotic stress treatment, bothcultivars showed retardation of plant growth, dwarf plants and young leaf rolling or evendrying. These symptoms were more serious in Zhonghan 3 than Xiushui 63. The osmoticstress greatly retarded the growth of roots and aerial parts in Zhonghan 3 than Xiushui 63.After initiation of the osmotic stress, relative water content (RWC) and water potential ofleaves decreased significantly in both cultivars, and Xiushiu 63 had higher leaf RWC andwater potential than Zhonghan 3. Xiushui 63 probably showed higher osmotic adjustmentbecause Xiushui 63 had greater drop of leaf osmotic potential and higher turgor compared toZhonghan 3 when subjected to the osmotic stress. After initiation of the osmotic stress,Zhonghan 3 showed decreased transpiration rate and stomata conductivity, while they wereunchanged in Xiushiu 63. Root hydraulic conductivity (Lp) decreased significantly in bothcultivars after initiation of the osmotic stress. Moreover, by view of the freehandcross-sections of adventitious roots under a fluorescent microscope, no any differentlignifications at exodermis or endodermis in the roots of both cultivars was observed.Mercurial compounds are widely used to study the function of aquaporins because they are
inhibitors for aquaporins. Mercury treatment greatly decreased root hydraulic conductivity ofnon-stressed plants in both cultivars. However, during the osmotic stress, mercury showedmore serious inhibition effects in Zhonghan 3, while showed alleviating inhibition effects inXiushiu 63. To summarize, all these results suggested that upland rice and lowland mobilizeddistinct mechanism to resist drought, i.e. upland rice was drought avoidance (e.g. increasingroot water uptake and decreasing water evaporation) while lowland rice was drought tolerance(e.g. osmotic adjustment). Aquaporins might play role in drought avoidance of upland rice.(2) To demonstrate the role of aquporins in plant drought avoidance in fine detail,aquaporin RWC3 from rice (Oryza sativa L.) was transferred into a lowland rice cultivars(Zhonghua 11), which was thought to be weak in drought avoidance, and consequently somephysiological parameters were measured during the water deficit mediated by PEG.SWPA2::RWC3 was constructed into a binary plasmid pCAMBIA 1301, and was transferred tolowland rice mediated by Agrobacterium tumefaciens. SWPA2 promoter, which was isolatedin sweetpotato, was a stress inducible promoter. By the assays of GUS histochemical staining,PCR and PCR-Dig-ELISA, the presence of SWPA2::RWC3 was confirmed in the lowland rice.By RT-PCR and Western blot analysis, it was found that SWPA2 promoter-RWC3 transgenicplant and WT (wild type) had similar RWC3 expression levels under normal condition,nevertheless, under water deficit mediated by 20% PEG, the transgenic plant had higherRWC3 expression levels than WT, suggesting that RWC3 expression in transgenic plants couldbe induced by drought. However, no visible phenotypic difference was found between thetransgenic plants and the WT plants, either under normal condition or during water deficit.After the water deficit, SWPA2 promoter-RWC3 transgenic plant showed higher root hydraulicconductivity and leaf water potential than WT, and the decrease of relative accumulativetranspiration rate during the water deficit was alleviated in the transgenic plant. These resultswere summarized that compared to WT, SWPA2 promoter-RWC3 transgenic plant wasendowed with higher water transport of roots and improved water status under water deficit,suggesting aquaporin RWC3 probably play a role during drought avoidance of rice(3) Plant chilling injury was partially caused by water imbalance between water uptakein root and transpiration in leaf. In this study, two rice cultivars (Somewake and Wasetoitsue)were exposed to 7 ± 1°C for up to 24 h and following temperature recovery (28 ± 1°C) for 3
h. Throughout the chilling stress, Wasetoitsu showed more serious injury symptoms thanSomewake. By measuring the relative electrical conductivity of rice leaves, Wasetoitsu hadhigher membrane electrolyte leakage than Somewake at the low temperature. These resultsindicated that Wasetoitsue was a less chilling tolerant cultivar compared to Somewake. Thetwo cultivars were in different water status, i.e. Wasetoitsu had lower leaf RWC thanSomewake in the duration of chilling, although both cultivars had decreasing RWC at thesame time RWC. In both cultivars, accumulative transpiration rate and root Lp decreasedobviously during the chilling treatment, but the decrease of Lp was more rapid and acute thanthat of accumulative transpiration rate, suggesting that the water imbalance occurred betweenthe water uptake of root and the water evaporation of aerial parts. These physiologicalobservations prompted us to investigate the expression of water channel proteins. At mRNAlevel, three rice aquaporin genes, RWC1、OsPIP2a and RWC3 showed the similar response tothe chilling treatment in both cultivars, i.e. remained approximate constant within the 9 h withchilling treatment while were decreased after 24 h with chilling treatment. The analysis ofWestern blot with antibody against PIP1 was consistent with the mRNA expression. In bothcultivars, PIP1 protein decreased when the plants were subjected to the chilling treatment for24 h. As we know, this is the first report on aquaporin responses to chilling stress in plantroots. Surprisingly, the temperature recovery for 3 h following chilling treatment resulted incontinued decreases of root PIPs mRNA expression. Aquaporins might be related to ricechilling adaptation but not a key factor of chilling resistance at the early stage of the chillingstress.
引文
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