气孔运动过程中保卫细胞液泡的动态及其与微丝骨架的关系
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摘要
气孔运动调节植物与环境之间气体和水分的交换。有研究表明,气孔运动过程中保卫细胞的液泡大小和数目随气孔开度的变化而改变,微丝骨架在气孔运动过程中参与多种信号途径。但是还不清楚液泡和微丝的动态变化细节,以及微丝骨架与液泡动态之间的关系。本论文主要利用共聚焦显微镜技术,结合荧光标记、绿色荧光蛋白等技术研究了保卫细胞液泡的动态变化及其在气孔运动中的作用,以及微丝骨架动态变化及其与液泡动态变化之间的关系。
研究结果表明:蚕豆气孔关闭时,保卫细胞内存在许多球形的和管状的小液泡。在气孔开放过程中,小液泡相互融合形成体积较大的液泡。当气孔关闭时,大液泡又重新变成许多小液泡。液泡融合的抑制剂E-64d抑制蚕豆气孔保卫细胞液泡的融合,同时也抑制气孔的开放速度。拟南芥AtVAM3(SGR3)基因编码蛋白SYP22定位在液泡膜上,参与小液泡的融合。利用拟南芥突变体sgr3-1的研究结果表明,由于SGR3基因的突变,使保卫细胞液泡的融合受到抑制,气孔的开放速度也明显比野生型的低。以上结果说明,小液泡相互融合有利于气孔的开放。
研究发现蚕豆关闭气孔的保卫细胞中包含复杂的液泡膜结构。光切片的3D重组以及光漂白实验发现蚕豆保卫细胞的液泡可能是相互连通的。推测关闭气孔保卫细胞中包含的复杂的液泡膜结构可能作为液泡膜的储存形式,液泡相互连通的特点可能有利于水分和溶质在液泡间的流动,以适应气孔快速开闭运动中液泡体积快速变化的需要。
利用转基因技术,研究了Gfp-mTalin标记的烟草气孔保卫细胞的微丝骨架。发现无论在关闭还是开放气孔的保卫细胞中,既存在网状的微丝骨架系统,也存在放射状排列的微丝骨架系统,但关闭与开放气孔保卫细胞中微丝排列与分布也有明显差异。保卫细胞微丝排列与分布存在多样性的特点可能与其功能有关。
利用液泡和微丝的双标记技术和药理学方法研究了微丝骨架动态与液泡动态变化之间的关系,发现CB处理能使微丝骨架解聚,也明显加快液泡数目和体积的变化过程,同时加快了气孔的开放速度。而Phalloidin处理则抑制微丝骨架解聚,减缓液泡体积和数量的变化过程和降低了气孔的开放速度。进一步的研究发现烟草气孔保卫细胞中微丝骨架和液泡可能是共定位的。因此微丝的解聚可能有利于液泡的融合,从而促进气孔的开放;而抑制微丝的解聚可能抑制液泡的融合,从而抑制气孔的开放速度。
本研究较详细地研究了保卫细胞液泡的动态变化及其在气孔运动过程中的作用,以及微丝骨架的动态变化与液泡动态变化间的关系,为保卫细胞液泡的动态变化和微丝骨架参与气孔运动的研究提供了新的证据和新的研究思路。
Stomatal movement is important for plants to exchange gas with environment. The process contributes to the optimization of the photosynthesis and transpiration. Studies have showed that the change of vacuolar volume in guard cells is involved in the stomatal movement. However, little is known about the regulatory mechanism for the rapid changes of the vacuolar volume in guard cells during stomatal movement. Actin cytoskeleton of guard cells also is dynamics and involved in various signal pathway in regulating stomatal movement. However, the relationship between the dynamic of vacuoles and that of actin cytoskeleton, and their interacting mechanism during stomatal movement are unclear. Using CLSM and GFP-labeling technology, we studied the dynamics and function of vacuole and the relationship between the dynamic of vacuoles and that of actin cytoskeleton in guard cell during stomatal movement.There are many small spherical vacuoles and tubular vacuoles in guard cells of closed Vicia faba stomata, and the tubular vacuoles link the spherical vacuoles. The small vacuoles are irregular and harbor many complex membrane structures in lumen of vacuoles, such as ingrowths of tonoplast and vesicles. The small vacuoles and complex membrane systems are fused with each other or with the bigger vacuoles to generate the large vacuoles during stomatal opening. Reversely, the large vacuoles in the guard cells split into smaller vacuoles and generate many complex membrane structures in the closing and closed stomata. Vacuole fusion inhibitor, E-64d, can significantly inhibit the stomatal opening of Viciafaba. Furthermore, mutation of the Arabidopsis thaliana SGR3 also leads to retardation of vacuolar fusion and stomatal opening. These evidences indicated that vacuolar fusion is required for stomatal movement. From the disappearance of the complex membranes in vacuolar lumen in the opened stomata, we also can infer that these membranes serve as a reservoir of tonoplast for the changes of vacuolar volume during stomatal movement. From the 3D projection of many optical sections and photobleaching by CLSM, we found that the lumens of different vacuoles in a guard cell of Viciafaba are interconnected and their saps can flow freely. Vacuolar continuum in guard cells might function in stomatal movement via facilitating the diffusion of solutes and water among the different parts of guard cells to maintain the balance of osmotic potential and turgor pressure in the whole guard cell.Using transgenic tobacco (Nicotiana tobacuum) plant overexpressing gfp-mTalin driven by 35s, we observed the F-actin arrangement and distribution in guard cells of stable and moving stomata. The web and radial pattern of the F-actin arrangement in closed and opened stomata were observed. The difference of distribution of actin of guard cells in the closed stomata and the opened was also found. The actin was found in the cortical plasma and cytoplasm in closed stomata, whereas it was mainly located in the cortical plasma in opened ones. We also found the endoplasmic F-actin increased and cortical F-actin decreased accordingly during stomatal movement induced stomatal closing by ABA and dark.Pharmacological experiments showed that cytochalasin B, which inhibit the polymerization of F-actin in guard cells, facilitates not only light and FC-induced stomatal opening, but also the changes of vacuolar number and volume during stomatal opening. In contrast, phalloidin, which polymerize the
F-actin, inhibits stomatal opening and vacuolar changes induced light and FC. By dobule labeling method, we found that the vacuoles and F-actin in guard cells of tobacco stomata are colocalization probably. Combining with the pharmacological results, we infered that F-actin depolymerization in guard cells facilitates the vacuolar fusion, thus enhance stomatal opening, and the retardation the vacuolar fusion by inhibiting F-actin depolymerization leads to downregulated stomatal opening.The dynamics and function of vacuoles of guard cells during stomatal movement, and the dynamics relationshi
研究结果表明:蚕豆气孔关闭时,保卫细胞内存在许多球形的和管状的小液泡。在气孔开放过程中,小液泡相互融合形成体积较大的液泡。当气孔关闭时,大液泡又重新变成许多小液泡。液泡融合的抑制剂E-64d抑制蚕豆气孔保卫细胞液泡的融合,同时也抑制气孔的开放速度。拟南芥AtVAM3(SGR3)基因编码蛋白SYP22定位在液泡膜上,参与小液泡的融合。利用拟南芥突变体sgr3-1的研究结果表明,由于SGR3基因的突变,使保卫细胞液泡的融合受到抑制,气孔的开放速度也明显比野生型的低。以上结果说明,小液泡相互融合有利于气孔的开放。
研究发现蚕豆关闭气孔的保卫细胞中包含复杂的液泡膜结构。光切片的3D重组以及光漂白实验发现蚕豆保卫细胞的液泡可能是相互连通的。推测关闭气孔保卫细胞中包含的复杂的液泡膜结构可能作为液泡膜的储存形式,液泡相互连通的特点可能有利于水分和溶质在液泡间的流动,以适应气孔快速开闭运动中液泡体积快速变化的需要。
利用转基因技术,研究了Gfp-mTalin标记的烟草气孔保卫细胞的微丝骨架。发现无论在关闭还是开放气孔的保卫细胞中,既存在网状的微丝骨架系统,也存在放射状排列的微丝骨架系统,但关闭与开放气孔保卫细胞中微丝排列与分布也有明显差异。保卫细胞微丝排列与分布存在多样性的特点可能与其功能有关。
利用液泡和微丝的双标记技术和药理学方法研究了微丝骨架动态与液泡动态变化之间的关系,发现CB处理能使微丝骨架解聚,也明显加快液泡数目和体积的变化过程,同时加快了气孔的开放速度。而Phalloidin处理则抑制微丝骨架解聚,减缓液泡体积和数量的变化过程和降低了气孔的开放速度。进一步的研究发现烟草气孔保卫细胞中微丝骨架和液泡可能是共定位的。因此微丝的解聚可能有利于液泡的融合,从而促进气孔的开放;而抑制微丝的解聚可能抑制液泡的融合,从而抑制气孔的开放速度。
本研究较详细地研究了保卫细胞液泡的动态变化及其在气孔运动过程中的作用,以及微丝骨架的动态变化与液泡动态变化间的关系,为保卫细胞液泡的动态变化和微丝骨架参与气孔运动的研究提供了新的证据和新的研究思路。
Stomatal movement is important for plants to exchange gas with environment. The process contributes to the optimization of the photosynthesis and transpiration. Studies have showed that the change of vacuolar volume in guard cells is involved in the stomatal movement. However, little is known about the regulatory mechanism for the rapid changes of the vacuolar volume in guard cells during stomatal movement. Actin cytoskeleton of guard cells also is dynamics and involved in various signal pathway in regulating stomatal movement. However, the relationship between the dynamic of vacuoles and that of actin cytoskeleton, and their interacting mechanism during stomatal movement are unclear. Using CLSM and GFP-labeling technology, we studied the dynamics and function of vacuole and the relationship between the dynamic of vacuoles and that of actin cytoskeleton in guard cell during stomatal movement.There are many small spherical vacuoles and tubular vacuoles in guard cells of closed Vicia faba stomata, and the tubular vacuoles link the spherical vacuoles. The small vacuoles are irregular and harbor many complex membrane structures in lumen of vacuoles, such as ingrowths of tonoplast and vesicles. The small vacuoles and complex membrane systems are fused with each other or with the bigger vacuoles to generate the large vacuoles during stomatal opening. Reversely, the large vacuoles in the guard cells split into smaller vacuoles and generate many complex membrane structures in the closing and closed stomata. Vacuole fusion inhibitor, E-64d, can significantly inhibit the stomatal opening of Viciafaba. Furthermore, mutation of the Arabidopsis thaliana SGR3 also leads to retardation of vacuolar fusion and stomatal opening. These evidences indicated that vacuolar fusion is required for stomatal movement. From the disappearance of the complex membranes in vacuolar lumen in the opened stomata, we also can infer that these membranes serve as a reservoir of tonoplast for the changes of vacuolar volume during stomatal movement. From the 3D projection of many optical sections and photobleaching by CLSM, we found that the lumens of different vacuoles in a guard cell of Viciafaba are interconnected and their saps can flow freely. Vacuolar continuum in guard cells might function in stomatal movement via facilitating the diffusion of solutes and water among the different parts of guard cells to maintain the balance of osmotic potential and turgor pressure in the whole guard cell.Using transgenic tobacco (Nicotiana tobacuum) plant overexpressing gfp-mTalin driven by 35s, we observed the F-actin arrangement and distribution in guard cells of stable and moving stomata. The web and radial pattern of the F-actin arrangement in closed and opened stomata were observed. The difference of distribution of actin of guard cells in the closed stomata and the opened was also found. The actin was found in the cortical plasma and cytoplasm in closed stomata, whereas it was mainly located in the cortical plasma in opened ones. We also found the endoplasmic F-actin increased and cortical F-actin decreased accordingly during stomatal movement induced stomatal closing by ABA and dark.Pharmacological experiments showed that cytochalasin B, which inhibit the polymerization of F-actin in guard cells, facilitates not only light and FC-induced stomatal opening, but also the changes of vacuolar number and volume during stomatal opening. In contrast, phalloidin, which polymerize the
F-actin, inhibits stomatal opening and vacuolar changes induced light and FC. By dobule labeling method, we found that the vacuoles and F-actin in guard cells of tobacco stomata are colocalization probably. Combining with the pharmacological results, we infered that F-actin depolymerization in guard cells facilitates the vacuolar fusion, thus enhance stomatal opening, and the retardation the vacuolar fusion by inhibiting F-actin depolymerization leads to downregulated stomatal opening.The dynamics and function of vacuoles of guard cells during stomatal movement, and the dynamics relationshi
引文
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