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標題: The Different Response to Oxidative Stress for GIGANTEA(GI) Orthologues from Arabidopsis and Ferns
作者: 黃品瑄
Pin-Syuan Huang
關鍵字: 蕨類;阿拉伯芥;基因;GI
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GIGANTEA(GI)基因在阿拉伯芥中是受光週期(circadian clock)調控並促進開花的重要基因,此外也被研究出與調控老化相關。在阿拉伯芥中,GI基因突變會使植株延遲開花並提高植株對氧化壓力的抗性。而研究顯示在開花途徑中GI蛋白藉由和FLAVIN-BINDING, KELCH REPEAT, F-BOX 1(FKF1)蛋白進行交互作用,進而調控下游的開花基因,如CONSTANS(CO)等基因。實驗室從屬於非開花植物的銀脈鳳尾蕨選殖到GI的同源基因,並命名為PcGI。PcGI的基因表現和GI同樣會受光週期調控。進一步研究發現,將阿拉伯芥GI基因(AtGI)、鳳尾蕨GI基因(PcGI)、含有阿拉伯芥GI基因的C端和鳳尾蕨GI的N端互相置換的重組基因(PcAt-GI)以及含阿拉伯芥GI基因的N端和鳳尾蕨GI的C端互相置換的重組基因(AtPc-GI)大量暫時性表現於菸草表皮細胞,發現AtGI蛋白只位於細胞核中,並呈現斑點狀分布。PcGI則會出現在細胞核和過氧化小體(peroxisome)中。PcAt-GI會位於在核及過氧化小體,且情況比AtPc-GI好。顯示AtGI的C端相較於N端在入細胞核及開花調控的功能上扮演較重要角色。PcGI的N端則會使PcGI進入過氧化小體,並參與氧化逆境之調控。進一步發現,於紅光以及微光處理下PcGI進入過氧化小體的情形比藍光處理的組別明顯,PcGI在藍光較少的環境促使PcGI進入過氧化小體。而藍光下PcGI與AtGI均會受到FKF1影響而造成核中的圖形分佈從斑點狀變成均勻分布,而在紅光下則影響較小,顯示AtGI與PcGI均會受藍光誘導與FKF1交互作用。而PcGI與catalase3(CAT3)互為競爭關係,其使CAT3無法進入過氧化小體分解過氧化氫而造成過氧化氫累積。綜合上述結果顯示,開花植物AtGI受藍光誘導並藉由AtGI之C端使AtGI進入核中與FKF1交互作用並調控開花。而鳳尾蕨類PcGI可受光照影響,判斷是否藉由PcGI之N端進入過氧化小體與CAT3競爭並造成植株過氧化氫濃度上升進而調控氧化逆境。因而推測生活於較陰暗潮濕之蕨類植物,其GI功能為調節氧化逆境已達到抗菌抗病的功能,隨著演化為生活於光亮乾燥的開花植物,GI之功能轉變為以調控開花為主。

In Arabidopsis, GIGANTE (GI) is known as an important flowering time regulatory gene, which is regulated by circadian clock. Mutation in GI caused the delay of the flowering time and the increase of the tolerance to oxidative stress in Arabidopsis. GI has been reported to interact with FLAVIN-BINDING, KELCH REPEAT, F-BOX 1(FKF1) to regulate downstream genes such as CONSTANS (CO) in flowering pathway. We have cloned GI orthologue PcGI from non-flowering ferns Cretan Brake. The expression of PcGI also showed a circadian rhythm. Furthermore, transient expression of AtGI, PcGI, PcAt-GI which contained the C-terminus of AtGI and N-terminus of PcGI, and AtPc-GI which contained the C-terminus of PcGI and N-terminus of AtGI were performed in tobacco cells. The results indicated that the localization of AtGI was detected only in nucleus, which had the speckled nucleus. The PcGI was detected in nucleus as well as in peroxisome. The localization of PcAt-GI in peroxisome and nucleus was stronger than that for AtPc-GI. These results revealed that the C-terminus of the AtGI is functionally more important than N-terminus in nucleus localization and flowering. The N-terminal portion of PcGI is the major motif for PcGI to localize in peroxisome and is associated with senescence and oxidative stress. PcGI was detected in peroxisome in red and weak light, but not in blue light. The AtGI and PcGI were changing from speckled to disperse in nucleus by interacting with FKF1 in blue light but with less effect in red and weak light. PcGI competed and prevented the entering of catalase 3 (CAT3) into peroxisome and resulted in the hydrogen peroxide accumulation. Our result implied that PcGI played a major role in regulating oxidative stress and antibiotic tolerance in ferns whereas GI orthologues become more important in regulating flowering time in flowering plants during evolution.
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