Please use this identifier to cite or link to this item: http://hdl.handle.net/11455/36265
標題: 光訊息傳遞因子LZF1/BBX22參與阿拉伯芥幼苗發育之分子研究
LZF1/BBX22 conveys light signals for optimal seedling development in Arabidopsis
作者: 張瓊穗
Chang, Chiung-Swey Joanne
關鍵字: light
暗型態發育
HY5
LZF1
COP1
photomorphogenesis
skotomorphogenesis
26S proteasome
Arabidopsis
光訊息傳遞因子
轉錄調控因子
蛋白酶體
後轉譯作用
生長優勢
出版社: 生物科技學研究所
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摘要: “光”調控許多植物生長與發育的過程。不論是面對微小或是劇烈的“光”環境變化,植物都可以作出適當偵測與回應,讓植物可以在多變化的“光”環境中獲得最佳生長優勢。在自然環境中,在土壤中發芽的阿拉伯芥幼苗進行暗型態發育,藉由持續延長的下胚軸來突破土表。在適當的光照環境中,植物則會進行光型態發育並開始自營生活。植物藉由光受體分子來接受“光”訊息,並經由訊息傳導和轉錄體的調整來因應“光”環境的改變。目前針對這些“光”訊息傳遞因子的了解,大多來自突變株的篩選,而且幾乎所有的“光”訊息傳遞因子均作用於HY5—一個很重要的光訊息傳遞轉錄因子—的上游。是否有其它的光訊息傳遞轉錄因子,能承接從HY5來的光訊號,進而調控下游的光型態生長,仍屬未知;相關研究將可以進一步開拓我們對於“光”如何有效改變植物轉錄體的知識。 我們藉由比對野生型和hy5突變株的差異性基因表現,發現了一個尚未被研究過的C2C2-CO B-box轉錄調控因子—LZF1/BBX22。在光照環境下,HY5會直接結合至LZF1的啟動子來活化LZF1基因表現。針對LZF1的研究顯示:LZF1是一個“光”訊息的正向調控因子,透過活化下游的基因表現—例如MYB75及多個葉綠體蛋白質相關基因,來抑制下胚軸的延長,促進花青素的生合成及葉綠體的發育等幼苗發育歷程。在短日照環境中,LZF1也會抑制阿拉伯芥下胚軸在黑暗中的延長。 我們更進一步探討LZF1蛋白質在黑暗及特定的光照環境中受到差異性調控的特性。在暗型態及光型態發育中,轉錄及後轉譯作用同時精確的控制LZF1的轉錄子及蛋白質的量。不論在黑暗或光照下,LZF1蛋白質都會經由26S蛋白酶體調控而快速降解,LZF1蛋白質的半衰期介於二十與六十分鐘之間。在黑暗中,LZF1蛋白質的降解需要COP1但不需要HY5的參與。在光照下,HY5則會參與部分LZF1蛋白質的降解作用。當生長在黑暗或短日照下,過量的LZF1蛋白質的產生會造成cop1突變株過度進行光型態發育,不利幼苗的生長。過量的LZF1蛋白質也會影響光訊息及賀爾蒙訊息因子的基因表現。所以在黑暗及短日照下,正確調控LZF1蛋白質適時的降解與適當的累積對植物維持最佳的生長優勢是非常重要的。 本研究發現一個新的光訊息傳遞因子—LZF1,藉由承接HY5所帶來的光訊息,進一步改變轉錄體來調控下游的光型態生長。LZF1基因的轉錄子與蛋白質的總量均受到嚴格的控管,使得阿拉伯芥幼苗中的光訊息傳導得以順利執行,以達到最佳的幼苗生長狀態。
Light regulates multiple aspects of plant growth and development in plants. To achieve an optimal growth control, plants sophistically adjust their developmental programs in response to both the tiny changes of environmental light compositions and the rapid fluctuation of light intensities. In nature, plants proceed with skotomorphogenesis before emerging from the soil. When exposed to the light, plants start to use photomorphogenesis to establish the architecture in support of their autotrophic lives. Proper regulation of these processes is important for plants to achieve their optimal growth. It is known that transcriptomic changes govern the expression of signaling molecules upon the perception of light. However, the identifications of most light signaling molecules were from forward genetic studies and most of these molecules function upstream of HY5, an influential light signaling transcriptional regulator. The discovery of new component(s) conveying light signals from HY5 to photomorphogenic growth will no doubt broaden our knowledge of transcriptional cascades in light signaling pathways. I used global transcriptome comparisons to survey genes differentially expressed during early photomorphogenesis in wild-type but not in hy5 mutant. These gene expression data revealed LZF1 (light-regulated zinc finger protein 1)/BBX22, a gene encoding a previously uncharacterized C2C2-CO B-box transcriptional regulator, whose induction is both light-regulated and HY5-dependent. HY5 activates LZF1 via direct binding to LZF1 promoter. HY5 is necessary but not sufficient for the induction of LZF1. Functional studies indicate that LZF1 is a positive regulator for diverse light-mediated seedling growth, including the inhibition of hypocotyl elongation, anthocyanin biogenesis and chloroplast development. LZF1 achieves these functions via activating the expression of MYB75 and the genes encoding chloroplast proteins. In the absence of HY5, mutation of LZF1 leads to further reduced light sensitivity, supporting a possible synergistic impact of HY5 and LZF1 in Arabidopsis. We also confirmed that LZF1 enhances the inhibition of hypocotyl growth under short-day (SD) via attenuating the hypocotyl elongation during the dark-grown period. I also characterized the tight post-translational regulation of LZF1 in response to light/dark environment. We found the expression of LZF1 is tightly regulated at both the transcriptional and post-translational levels in skotomorphogenesis and during the process of de-etiolation. LZF1 is a short-lived protein degraded by 26S proteasome under both dark and light conditions. The degradation of LZF1 in the dark depends on the presence of COP1 but not HY5. The degradation of LZF1 in the light partially depends on HY5. To assess whether the rigid LZF1 degradation control is required for optimizing plant growth, we analyzed the seedling growth while LZF1 is over-produced in the cop1 mutant. The over-produced LZF1 negatively affects the Arabidopsis seedling development in the dark and short-day environments. Over-produced LZF1 exaggerates light-mediated seedling growth via regulating the expression of light and hormone-responsive genes. Therefore, the proper accumulation of LZF1 is crucial for plants to maintain optimal growth fitness when growing in the dark as well as under short-day. This research demonstrates that, by combining the rapid transcriptional activation and rigid surveillance of protein abundance via post-translational degradation, LZF1 functions to convey light signal from HY5 for an optimal seedling development in Arabidopsis.
URI: http://hdl.handle.net/11455/36265
其他識別: U0005-2910201001495900
文章連結: http://www.airitilibrary.com/Publication/alDetailedMesh1?DocID=U0005-2910201001495900
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