臺灣博碩士論文加值系統

E3 RING finger的基因在ubiquitin蛋白質降解過程以及控制植物生長發育和分化扮演重要的角色。APC8是E3蛋白複合體中的其中一個成員，控制有絲分裂的進行以及完成。APC8基因擁有TPR-repeat-containing domain，它的功能是蛋白質與蛋白質之間的交互作用。雖然APC/C在真菌和動物中廣泛被研究，但是在植物中的功能仍未知。實驗發現一個阿拉伯芥T-DNA突變株，其外表型呈現不孕和不正常的葉片。透過inverse PCR (IPCR)的檢查發現T-DNA插在APC8基因的5’端調控區，距離起始密碼有199 bp。有趣的是所有的apc8突變植株皆以heterozygotes型存在，但其APC8 mRNA和蛋白質表現量高於正常野生阿拉伯芥。這樣的結果顯示apc8突變外表型有可能是因為T-DNA插入造成APC8基因被活化所致，為了證明此可能性，我們利用35S promoter表現APC8 cDNA sense和antisense並分析外表型。結果發現antisense的植物外表型與野生型並無明顯差異，而sense的植物外表型則相似於apc8突變的植株。近一步研究發現35S::APC8 antisense可以降低apc8突變植株中APC8的表現量，並減緩其不孕的現象。接著promoter::GUS轉基因植物分析試驗顯示，290 bp長度的APC8 promoter 片段所驅動GUS基因的表現量高於410 bp 或 1500 bp長度片段所驅動。此結果說明在原始APC8 promoter於起始密碼前199 到 410 bp間可能含一負調控區，會抑制APC8之表現，在apc8突變植株中這個區域被T-DNA插入，所以阻止了負調控子之結合在正常的區域，造成APC8表現提高而形成突變。除了APC8 這個基因外，本研究包括另外兩個E3 RING finger 基因 AtRING3 和 AtRING4的特性分析，其中AtRING3 和 AtRING4 mRNA在成株的葉和花中表現量較高。為了研究AtRING3 和 AtRING4的功能，我們利用yeast two-hybrid assay，釣取到三個與AtRING4有結合影響的蛋白，分別是COR47, At2g17190 與 CSN5B，其中CSN5B亦與AtRING3結合。

E3 RING finger genes played key roles in the ubiquitin-mediated proteolysis and controlled many processes of plant development and differentiation. APC8 is one of the multiple-subunit of E3 protein that controls important transitions during mitotic progression and exit. A TPR-repeat-containing domain that controlled the protein-protein interaction was identified in APC8 protein. Although APC/C has been extensively studied in both fungal and animal, the structure and function of plant APC/C’s remained unexplored. An Arabidopsis T-DNA insertional mutant caused germ abortion and abnormal leaf formation was characterized in this research. Through inverse PCR (IPCR), it has been found that the T-DNA was inserted in the 199 bp of 5’ regulation regions from start codon of APC8 gene. Interestingly, all the apc8 mutants are heterozygotes for T-DNA insertion and the APC8 mRNA and protein were up-regulated in the apc8 mutants . This result revealed that the apc8 mutant phenotype may be caused by the activation of APC8 gene after T-DNA insertion. To explore this possibility, sense and antisense of APC8 cDNA driven by 35S promoter were transformed into Arabidopsis and phenotypic analyzed. The result indicated that antisense plants were phenotypically indistinguishable from wild-type plants whereas sense plants exhibited similar phenotype resembled T-DNA insertional apc8 mutants. Furthermore, 35S::APC8 antisense down-regulated the APC8 expression and rescued the germ abortion in apc8 mutants. Further promoter analysis showed that transgenic plants contained 290 bp length of APC8 promoter fusing with GUS exhibited much higher GUS expression and GUS activity than the transgenic plants contained 410 bp and 1500 bp length of APC8 promoter fusing with GUS. Our results revealed that a negative regulator may bind to the region between 199 to 410 bp of APC8 promoter and suppressed the APC8 expression in wild-type plant. This region was interrupted by the T-DNA insertion in apc8 mutants and prohibited the binding of negative regulator resulting in the increase of APC8 expression in the mutants. In addition to APC8 gene, two E3 RING finger genes, AtRING3 and AtRING4 were also cloned and characterized. AtRING3 and AtRING4 mRNA were expressed in leaf and flowers. To investigate the function of AtRING3 and AtRING4 action, yeast two-hybrid assays was performed. Three proteins (COR47, At2g17190 and CSN5B) that interacted with AtRING4 and one protein (CSN5B) that interacted with RING3 were identified.

目次

總摘要(中文)
總摘要(英文)

第一章阿拉伯芥中apc8 突變株之定性分析

摘要 ---------------------------------------------------------------------------------------------2
壹、前言-----------------------------------------------------------------------------------------3
貳、材料與方法--------------------------------------------------------------------------------9
参、結果----------------------------------------------------------------------------------------18
一、apc8 突變株性狀分析---------------------------------------------------------------18
二、IPCR結果分析-----------------------------------------------------------------------18
三、apc8 突變株基因型的分析---------------------------------------------------------18
四、apc8 突變株與野生阿拉伯芥中，APC8 基因表現量的比較---------------19
五、apc8 突變株與野生阿拉伯芥中，APC8 蛋白表現量的比較---------------19
六、APC8 的轉基因植物功能性分析------------------------------------------------- 20
(1) 35S:: APC8與35S:: APC8 antisense構築體之構築--------------------------20
(2)35S:: APC8 轉入阿拉伯芥野生型性狀分析-----------------------------------21
(3) 35S:: APC8轉基因植物的基因表現分析----------------------------------------21
(4)野生阿拉伯芥、apc8 突變株及.35S:: APC8轉殖株中APC8的表現量----
分析-----------------------------------------------------------------------------------------22
(5) 35S:: APC8 antisense轉入apc8 突變株型性狀分析與表現量分析--------22
七、APC8 5端的調控區的分析--------------------------------------------------------------23
(1)選殖三段不同長度APC8基因的啟動子---------------------------------------- 23
(2) APC8 promoter 290bp::GUS、APC8 promoter 410bp::GUS與APC8 promoter 1500bp::GUS的轉殖株，GUS染色結果與GUS基因表現量分析----------24
八、apc8 突變株、野生阿拉伯芥中及35S:: APC8轉殖株中，APC2基因和 APC11基因表現量的比較-----------------------------------------------------------------------25
肆、討論-----------------------------------------------------------------------------------------27
伍、參考文獻-----------------------------------------------------------------------------------31
陸、圖表---------------------------------------------------------------------------------------35






第二章 阿拉伯芥RING finger基因(AtRING3及AtRING4)之特性與功能性分析

摘要 ----------------------------------------------------------------------------------------------62
壹、前言-----------------------------------------------------------------------------------------63
貳、材料與方法--------------------------------------------------------------------------------69
参、結果-----------------------------------------------------------------------------------------73
一、AtRING3、AtRING4與AtRING1基因資訊的比較----------------------------73
二、AtRING3、AtRING4於阿拉伯芥野生型的表現情形--------------------------73
三、AtRING3、AtRING4之酵母雙雜合( yeast two-hybrid )分析-----------------74
四、AtRING3與AtRING4轉基因植物的性狀分析----------------------------------75
五、AtRING3與AtRING4 啟動子(promoter)的選殖---------------------------------76
肆、討論------------------------------------------------------------------------------------------78
伍、參考文獻------------------------------------------------------------------------------------82
陸、圖表------------------------------------------------------------------------------------------87

圖表目次

--表1-1 第一章PCR反應所使用引子之序列-----------------------------------------35
--圖1-1 野生型阿拉伯芥與apc8突變株之性狀分析--------------------------------37
--圖1-2 野生型阿拉伯芥與apc8突變株之性狀分析--------------------------------38
--圖1-3 IPCR鑑定結果與apc8突變株基因型分析----------------------------------39
--圖1-4 推測apc8突變株同型接合子(homozygote)會致死-------------------------40
--圖1-5 野生型阿拉伯芥與apc8突變株，APC8 基因表現量的比較------------41
--圖1-6 野生型阿拉伯芥與apc8突變株，APC8 蛋白表現量的比較 -----------42
--圖1-7 以PCR方法選殖APC8 (At3g48150)cDNA----------------------------------43
--圖1-8 35S:: APC8 sense 與 antisense clone之確認---------------------------------44
--圖1-9 35S::APC8轉殖阿拉伯芥野生型之性狀分析--------------------------------45
--圖1-10 35S:: sense APC8轉基因植物的基因表現情形------------------------------46
--圖1-11 野生型阿拉伯芥、 apc8突變株與35S:: sense APC8轉基因植物的APC8表現情形---------------------------------------------------------------------------47
--圖1-12 偵測35S::antiAPC8/apc8轉殖株APC8表現量與性狀分析--------------48
--圖1-13 選殖三段不同長度APC8基因的啟動子(promoter)之確認----------------49
--圖1-14 三段不同長度APC8基因的啟動子融合報導基因分析--------------------50
--圖1-15 APC2和APC11在野生阿拉伯芥、 apc8 突變株中及 35S:: APC8轉殖株中的表現量分析----------------------------------------------------------------51
--圖1-16 阿拉伯芥T-DNA insertion mutant(apc8 mutant)的模式圖------------------52
--附表1-1 分裂的酵母菌、脊椎動物以及阿拉伯芥APC/C的組成-------------------53
--附圖1-1 ubiquitin (Ub)/26S proteasome pathway示意圖-----------------------------54
--附圖1-2 apc2 insertion mutant 的性狀與T-DNA插入的位子----------------------55
--附圖1-3 APC/C在cell cycle的功能-----------------------------------------------------56
--附圖1-4 pGEM®-T Easy vector之載體圖譜--------------------------------------------57
--附圖1-5 pBI121之載體圖譜--------------------------------------------------------------58
--附圖1-6 pBI101 和pEpyon-01K之載體圖譜------------------------------------------59
--附圖1-7 pEpyon-12H之載體圖譜--------------------------------------------------------60
--表2-1 第二章PCR反應所使用引子之序列-------------------------------------------87
--表2-2 yeast two-hybrid實驗篩選出與AtRING1、AtRING3和AtRING4有交互作用的蛋白表-------------------------------------------------------------------89
--圖2-1 阿拉伯芥AtRING1(At3g10910 )mRNA序列及相對應氨基酸------------90
--圖2-2 阿拉伯芥AtRING3(At5g05280)mRNA序列及相對應氨基酸-------------91
--圖2-3 阿拉伯芥AtRING4(At5g01880)mRNA序列及相對應氨基酸-------------92
--圖2-4 AtRING1、AtRING3與AtRING4氨基酸序列比對--------------------------93
--圖2-5 AtRING基因之演化樹圖譜------------------------------------------------------94
--圖2-6 利用RT-PCR偵測AtRING3和AtRING4基因於野生型阿拉伯芥中的表現情況-------------------------------------------------------------------------------95
--圖2-7 用於yeast two-hybrid實驗之AtRING3和AtRING4基因的ORF片段之分子選殖-------------------------------------------------------------------------------96
--圖2-8 酵母菌中pGBKT7 構築體鑑定及高營養篩選篩選出與AtRING3、AtRING4交互作用的蛋白------------------------------------------------------97
--圖2-9 35::AtRING3 和35::AtRING4突變株之性狀分析---------------------------98
--圖2-10 選殖AtRING3和AtRING4基因的啟動子(promoter)之確認--------------99
--圖2-11 與AtRING1、AtRING3與AtRING4有相互作用的蛋白模式圖--------100
--附圖2-1 (a)HECT、(b)RING/U-box、(c)SCF及(d)anaphase-promoting complex(APC)E3蛋白接合酵素的構造圖-----------------------------------101
--附圖2-2 35S::AtRING1 antisense植株之異常性狀-----------------------------------102
--附圖2-3 35S::Cor47-histag sense植株之異常性狀-----------------------------------103
--附圖2-4 35S::STZ sense 植株之異常性狀---------------------------------------------104
--附圖2-5 SCF complex受CSN及CAND1調控的示意圖--------------------------105
--附圖2-6 cullins 的neddylation與deneddylation示意圖----------------------------106
--附圖2-7 阿拉伯芥CSN5A和CSN5B的點突變對於cullin neddylation 和植物生長外表型的影響-------------------------------------------------------------107
--附圖2-8 pGBKT7之載體圖譜-----------------------------------------------------------108
--附圖2-9 pGAD10之載體圖譜------------------------------------------------------------109
--附圖2-10 （A）Yeast two-hybrid系統原理示意圖。（B）AH109及Y187兩種Yeast strain所使用的報導基因-----------------------------------------------------110

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