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標題: 榖類種子糊粉層細胞離層酸信息傳導之新解析
New insights into ABA signaling pathways in cereal aleurone cells
作者: 黃冠穎
Huang, Kuan-Ying
關鍵字: 榖類
出版社: 生物科技學研究所
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摘要: The phytohormone, abscisic acid (ABA) regulates many aspects in plant development including seed formation, dormancy and germination. Several putative ABA receptors have been reported in recent years; however the validity of some of these reports has been seriously questioned. We have investigated the role of these proteins in cereal aleurone tissue where well-defined ABA responses can be analyzed with precision. These putative ABA receptors or regulatory proteins include FCA (flowering time control protein a), CHLH (H subunit of chloroplast Mg2+-chelatase), GTG (GPCR-Type G protein), and RPK (receptor-like protein kinase). All these genes were overexpressed or suppressed by RNAi in cereal aleurone cells to investigate their effects on hormone signaling in aleurone tissue consisting of a single cell type responsive to gibberellin (GA) and ABA. GFP fusion proteins of these regulators were also expressed to study their cellular localization. ABA induction of LEA (late embryogenesis abundant) protein genes was enhanced by over-expression of FCA, but significantly suppressed by FCA RNAi or CHLH RNAi. Simultaneous repression of the expression of all four regulatory proteins further blocked the ABA induction of LEA genes. However, the other two signaling pathways, i.e. GA induction and ABA suppression of α-amylase, were not affected by overexpression or repression of FCA, RPK, GTG or CHLH genes. Furthermore, the suppressive effect of GTG RNAi was partially overcome by overexpression of FCA, suggesting that this nucleus-localized regulator may work downstream from GTG. This independent pathway could be blocked by the mutant form of protein phosphatase 2C, ABI1, suggesting interplay between the PYR/PYL/RCAR receptor system and these regulators. The role of FCA has been more extensively investigated in the cereal aleurone cells because it was first isolated from this tissue. Following a multidisciplinary approach combining molecular genetics, biochemistry, cell biology, and physiology, we have obtained strong evidence to suggesting that FCA is an important enhancer in ABA signaling in cereal grain germination. Epistasis studies indicated that FCA works upstream from VP1/ABI5,which are important signaling molecules/transcription factors involved in ABA up-regulation of LEA protein expression. The FCA-GFP fusion protein is initially localized in the cytoplasm with a punctate pattern but then gradually translocated into the nucleus. This cytosol-to-nucleus translocation of FCA was further enhanced by ABA treatment. Both yeast and in planta two-hybrid studies revealed that FCA interacts with VP1. In vitro pull-down assay also confirmed that VP1 and FCA physically interact with each other. Mutation of the highly conserved WW domain in FCA suppressed nuclear translocation, disrupted FCA-VP1 interaction, and also suppressed ABA signaling. Our results indicate that FCA plays a pivotal role in ABA signaling by transducing the signaling from cytosol to the nucleus where it then interacts with the transcription factor complex of VP1/ABI5 that are required for ABA up-regulation of gene expression. Furthermore, our observations suggest although FCA regulates flowering in dicotyledonous plants, its role in cereals is centered on the regulation of seed germination, an intruging divergence in function for this regulatory protein. The role of SnRK2 protein kinase family in mediating the ABA suppression of gene expression has also been explored in this project. Among the ten family members, SAPK1 and SAPK2 were sufficient in mimicking ABA in suppressing the GA-induced α-amylase expression. Neither of these SnRK2 kinases appears to have significant inhibition on ABA induction. None of the other SnRK2 genes exert the same effect. However, knocking down of the expression of either SAPK1 or SAPK2 gene, or both of them, had no significant effect on the action of ABA. An additional, yet unidentified, regulator is likely to be involved in the ABA suppression of gene expression. In summary, the results obtained in this thesis project suggest the existence of multiple ABA signaling pathways leading to the ABA regulation of gene expression. The role of one of signaling molecules, FCA, have been investigated.
近年來許多植物可能的ABA (abscisic acid, 離層酸) 受器 (receptor) 陸續被發表, 然而某些成員正確性被嚴重質疑, 本實驗主要目的在探討幾個可能的ABA調控因子在穀類糊粉層細胞中離酸信息傳導所扮演的角色, 這些基因包括: FCA (Flower time Control protein A), CHLH (H subunit of Chloroplast Mg2+-chelatase), GTG (GPCR-type G protein), 以及RPK (receptor-like protein kinase). 藉由在糊粉層中持續性表現或是RNA干擾(RNAi)這些基因, 我們嘗試找出它們對ABA及GA(Gibberline, 吉貝素)信息傳導的影響. 我們也將這些調控蛋白融合綠色螢光蛋白(GFP)來觀察牠們在細胞中分佈的情況. 結果顯示FCA的持續性表現會增益30% ABA 所誘發的LEA(Late Embrygenesis Abundant protein)基因表現, 而FCA 及CHLH的RNAi 會顯著的抑制此反應. 如同時將此四個調控因子以RNAi處理抑制它們的表現, 那麼ABA 所誘發的LEA表現將會更進一步的被抑制. 然而, 其他的一些賀爾蒙信息傳導作用, 像是GA 所誘發且會被ABA所抑制的α澱粉酶(α-amylase)則不受此四個調控蛋白所影響. 我們發現RPK蛋白分佈於細胞膜ㄧ帶, CHLH分布在質體(plastid)中, GTG分布於液泡狀物體的表面, 顯示這些調控蛋白可能擁有不同功能. 而這作用會被ABI1基因所抑制, 顯示PYR/PYL/RCAR系統亦與這些調控因子有關聯. 藉由各種不同的研究策略, 我們發現FCA為一個重要的ABA信息強化因子(enhancer). 如前面所述, FCA其作用對象僅針對ABA信息系統, 持續性表現會增益超過30% ABA 所誘發的LEA基因表現. 在ABA信息系統中, FCA作用於ABI5/VP1, 這兩個對ABA 誘發LEA表現至關重要的轉錄因子的上游. FCA-GFP蛋白一開始以小點狀分佈於細胞質中, 隨著時間增加, FCA逐漸移往細胞核中. 這種運動可以被ABA處理所加速. 且為ABI1所抑制. 不管在以酵母菌或是植物細胞為背景的雙雜合(two-hybrid)系統中, FCA都會與VP1這個轉錄因子發生交互反應(Interaction). 在活體外進行的pull-down assay也呈現相同結果. 我們更發現當雙色胺酸功能區(WW domain)發生變異時, FCA向細胞核進行的運動, 與VP1的交互反應以及對ABA信息的增益能力都會被抑制. 結合以上結果, 我們證明FCA 蛋白具有將ABA信息自細胞質傳導到細胞核內, 並與VP1/ABI5發生交互反應, 從而促進ABA誘導基因之表現. 為了解ABA與GA 之間的拮抗關係, 我們針對水稻SAPK(屬於SnRK2群)的基因群進行研究, 發現其中SAPK1與SAPK2對GA誘發澱粉酶的表現具有某種程度的抑制效果. 但即使將此兩基因基因同時以RNAi技術進行干擾, ABA依舊抑制澱粉酶的表現. 總體而言, 本實驗所獲得的結果證實有多條ABA信息傳遞的途徑存在於穀類糊粉層細胞之中, 其中FCA所扮演的角色在本研究中已獲得闡明.
其他識別: U0005-2208201214043600
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