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標題: 阿拉伯芥中NAC-like基因與ERD基因之功能性分析
Characterization and Functional Analysis of NAC-like and ERD Genes in Arabidopsis thaliana
作者: 吳秋嫺
Ng, Chiu-Sien
關鍵字: NAC;NAC;gene redundant;Chimeric repressor silence technology;功能重複;CRES-T
出版社: 生物科技學研究所
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NAC (NAM, ATAF1, 2, CUC2)為一群植物特有的轉錄因子,被發現參與多種植物發育過程之調控。本研究從阿拉伯芥中選殖兩個NAC-like基因,分別為AtNAC3次群中的AtNACL2及NAC2次群中的AtNACL10。根據基因表現分析, AtNACL2和AtNACL10在葉片和花苞都有較高的表現量,而AtNACL10在根部及花序亦有較高的表現量。在性狀分析方面,AtNACL2和AtNACL10各別的T-DNA插入突變株並沒有明顯性狀,推測此二基因有功能重複(gene redundant)的現象,所以本實驗利用Chimeric repressor silence technology (CRES-T)技術來解決基因功能重疊的問題。本研究選殖上述兩個基因之序列,並構築至帶有抑制子(repressor) SRDX序列之載體上,進一步轉殖至阿拉伯芥中,藉由其表現之性狀來分析這些AtNACL基因之功能。結果顯示,在35S::AtNACL2-SRDX轉殖株中出現鋸齒葉性狀;而35S::AtNACL10-SRDX轉殖株則出現葉片上捲及叢生花序等性狀。在後期發育上,35S::AtNACL2-SRDX和35S::AtNACL10-SRDX的轉殖株也出現了花絲較短及花藥不開裂的現象。進一步利用real-time PCR分析顯示,與頂芽分生組織相關基因FAS (FASCIATA), WUS (WUSCHEL)及葉片發育相關基因KNAT1(Knotted-like from Arabidopsis thaliana1), IAMT1(IAA carboxyl methyltransferase1),在突變株中的表現量皆有受到影響。綜合上述分析結果,認為AtNACL2和AtNACL10透過不同路經,達到共同參與調控葉片及花器的發育。阿拉伯芥中的AtEPF1為Cys2/His2型鋅指基因,屬於具完整保守序列CX2CX3FX5LX2HX3H的EPF基因家族,而將之異位表現會造成植株極度矮小及近似乾燥的性狀。為了深入研究受到AtEPF1調控的基因,藉由生物晶片(microarray)分析,得到許多受AtEPF1調控的基因,本研究分別挑選了ERD4 (EARLY-RESPONSIVE TO DEHYDRATION 4)及ERD7 (EARLY-RESPONSIVE TO DEHYDRATION 7)進行深入研究。ERD4和ERD7基因分別屬於不同基因家族,但皆是一群在乾旱逆境早期被迅速誘導表現的基因。ERD4和ERD7在阿拉伯芥各個生長時期及組織器官都有表現。經過斷水逆境處理得知此二基因的表現量確實會受到外界環境水分缺乏的影響。在啟動子分析方面,在ERD4::GUS植株,子葉、根部、葉緣排水孔及成熟花花萼、花柱、花粉皆偵測到GUS活性;而在ERD7::GUS植株,子葉、根部、花柱及花粉都具GUS活性。另外當ERD4::GUS植株處於斷水逆境,葉緣排水孔的GUS活性會延伸至葉脈維管束。在35S::ERD7轉殖株中,亦觀察到植株矮小及葉片黃化皺縮的性狀。若進一步分析,發現35S::ERD7轉殖株葉綠素含量降低,丙二醛含量上升,且受乙烯誘導表現的ERF1 (ETHYLENE-RESPONSE-FACTOR 1)基因表現上升。據文獻指出,當植物處於淹水逆境,葉綠素會因崩解而降低,且植物體內乙烯含量會上升。基於上述,認為受AtEPF1負調控的ERD4及ERD7在阿拉伯芥中可能參與水分的調控,以達到保護植物的目的。

NAC (NAM, ATAF1,2 and CUC2 ) proteins have been considered as one of the plant-specific transcription factors that are involved in the regulation of diverse plant developmental processes. Two Arabidopsis NAC-like genes AtNACL2 and AtNACL10 were isolated and analyzed in this study. AtNACL2 and AtNACL10 were highly expressed in the leaves and flowers. Interestingly, AtNACL10 is also highly detected in root and inflorescence in Arabidopsis thaliana. T-DNA mutants of AtNACL2 and AtNACL10 were phenotypically indistinguishable from wild-type. To solve the problem caused by functional redundancy, the chimeric repressor silencing technology (CRES-T) was used. In 35S::AtNACL2-SRDX plants, the abnormal phenotype with serrated leaves was observed. 35S::AtNACL10-SRDX plants exhibited dramatic leaf curvature and multiple branches with secondary inflorescence or flowers at the same position of an internode. In addition, 35S::AtNACL2-SRDX and 35S::AtNACL10-SRDX showed serious sterility due to the production of short stamen and indehiscence of the anther. Further analysis indicated that these mutant phenotypes were correlated with the alteration of the expression for genes involved in meristematic development such as FAS (FASCIATA), WUS (WUSCHEL) and leaf development such as KNAT1(Knotted-like from Arabidopsis thaliana1), IAMT1(IAA carboxyl methyltransferase1). The results suggest that AtNACL2 and AtNACL10 are key regulators in regulating leaf morphology and floral development in Arabidopsis thaliana. A conserved sequence of CX2CX3FX5LX2HX3H identified in the Cys2/His2 zinc finger region of the AtEPF1 was present in EPF family genes. Ectopic expression of AtEPF1 in transgenic Arabidopsis plants showed phenotypes by significantly reducing the plant size and producing curled and dehydrated leaves. To investigate the target genes regulated by AtEPF1, microarray analysis was performed. Two of AtEPF1 regulated genes, ERD4 (EARLY-RESPONSIVE TO DEHYDRATION 4) and ERD7, were further analyzed. ERD4 and ERD7 were originally described as a rapidly drought-responsive gene in Arabidopsis. When exposing plants to progressive drought stress, we obtained elevated levels of the ERD4 and ERD7. In ERD4::GUS plants, GUS activity was detected in cotyledons, roots, hydathodes, sepal, style and pollens. In ERD7::GUS plants, the GUS activity was detected in cotyledons, roots, style and pollens. Interestingly, GUS activity in ERD4::GUS plants was detected in leaf veins by drought treatment. Ectopic expression of ERD7 showed novel phenotypes by reducing the plant size and producing yellow wrinkled leaf. Further analysis indicated that the chlorophyll relative contents also decreased, lipid peroxidation tended to increase and ERF1 (ETHYLENE-RESPONSE-FACTOR 1) expression up-regulated. The present evidence indicated that ERD4 and ERD7 may play roles in protection of the plants with water retention. These results support our speculation for the ERD4 and ERD7 function based on its regulation by AtEPF1.
其他識別: U0005-2607201120090700
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