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研究生： 鄭明龍
Cheng, Ming-Lung
論文名稱： 由T-DNA插入突變種源中發掘調控水稻發育之新穎基因
Mining for novel genes regulating rice development from the T-DNA insertion mutant population
指導教授： 余淑美
Yu, Su-May
賀端華
Ho, Tuan-Hua
黃浩仁
Huang, Hao-Jen
學位類別： 博士
Doctor
系所名稱： 生物科學與科技學院 - 生命科學系
Department of Life Sciences
論文出版年： 2017
畢業學年度： 105
語文別： 英文
論文頁數： 54
中文關鍵詞： TRIM資料庫 、肌動蛋白絲 、生長激素運輸 、胞間連絲 、水稻 、捲葉 、WIN1基因
外文關鍵詞： TRIM database, Actin filament, Auxin transport, Plasmodesmata, Rice, Spiral phenotype, WIN1
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    • 水稻為世界上主要的糧食作物之一，超過50% 的人口以水稻為主食。水稻已於2005年完成全基因體定序，並預測至少有三萬六千個基因存在水稻的基因體中。利用這些已定序的資料及配合農桿菌轉殖技術，Taiwan Rice Insertional Mutants (TRIM) database已建立水稻基因剔除或活化的T-DNA插入突變種原庫。我們利用此種原庫篩選出一個造成水稻鞘葉呈螺旋狀生長的基因（WINDING 1, WIN1）。此基因在突變株中會被T-DNA活化。WIN1 為具有BTB及NPH3 domains及一個coiled coil motif的蛋白質，其與阿拉伯芥的NPH3蛋白質家族具有同源性。WIN1的表現主要在根部、根與鞘葉間的組織，但當WIN1於鞘葉過量表現時，即會造成兩種不同方向的螺旋狀捲葉。在根部的eGFP-WIN1會送到細胞膜上下位置，並與原生質絲共同形成點狀的分佈；但在葉部則分佈在細胞膜上下左右的位置，且會導致生長激素集中分佈在螺旋捲葉外曲的位置。此外，在螺旋捲葉外曲的細胞其絲狀肌動蛋白（F-actin）的量也較螺旋捲葉內曲的細胞多，顯示WIN1會與肌動蛋白作用，造成生長激素不對稱分佈進而造成捲葉的生長。

    Rice is the most important staple food in the world and supports over 50% of the human population. Sequencing of the whole genome of rice was finished in 2005, which predicted at least 30,000 genes. The sequence data provide good bioinformatics information for rice gene functional studies. The Taiwan Rice Insertional Mutants (TRIM) population contains at least 100,000 T-DNA insertion mutant lines, making this a valuable resource and efficient tool for rice gene discoveries. By adopting a forward genetics approach, we screened the TRIM population and found an interesting T-DNA insertion mutant, WINDING 1 (WIN1). The WINDING 1 (WIN1) gene encodes a protein containing BTB and NPH3 domains and a coiled-coil motif, which is homologous to the Arabidopsis NPH3 protein. The rice homolog of NPH3, CPT1, has been shown to be important for coleoptile phototropism and regulation of lateral auxin transport. WIN1 is mainly expressed in roots and leaf/root junctions. Subcellular localization studies show that WIN1 is primarily associated with plasmodesmata along the horizontal edges of plasma membranes. However, upon ectopic expression, WIN1 is distributed on all edges of leaf blade and sheath cells. The excurvature side of the spiral leaf sheath in WIN1Act and WIN1-Ox plants contains more auxin and F-actin than the incurvature side, indicating that WIN1 regulates F-actin organization and causes asymmetric auxin distribution. This dissertation demonstrates the utility of using the genetic resources in the TRIM population for gene discovery and functional studies in rice.

    Chapter 1 1 Introduction to rice functional genomics research using TRIM 1 References 5 Chapter 2 7 Ectopic expression of WINDING 1 leads to asymmetrical distribution of auxin and a spiral phenotype in rice 7 Introduction 7 Materials and Methods 10 Plants materials 10 Primers 10 Plasmid construction 10 RT-PCR analyses 11 Phylogenetic analysis of WIN1 homologous proteins 12 GUS staining 12 Subcellular localization studies 12 Microscopic examination of actin filaments 13 Protein extraction and immunoblotting analyses 13 GUS activity assays 14 Measurement of epidermal cell size and guard cell size 14 Statistical analysis 14 Accession number of WIN1 15 Result 16 Ectopic expression of WIN1 leads to a spiral phenotype in rice shoots 16 WIN1 is a member of the BTBN protein family with novel functions 17 WIN1 is specifically expressed in roots and shoot/root junctions 18 WIN1 primarily localizes on plasma membrane and partially co-localizes with plasmodesmata 18 The spiral phenotype in WIN1Act and WIN1-Ox plants is not altered by growth under constant darkness or treatment with exogenous auxins or auxin transporter inhibitors 20 Ectopic expression of WIN1 leads to asymmetric distribution of auxin in leaf sheaths 20 Ectopic expression of WIN1 leads to differential cell expansion and a reduction of F-actin in leaf sheaths 21 Discussion 23 Ectopic expression of WIN1, which encodes a unique BTBN protein, is responsible for the shoot spiral phenotype 23 WIN1 is localized to the plasma membrane in a punctate pattern and also partially co-localized to plasmodesmata 24 Ectopic expression of WIN1 leads to an unequal horizontal distribution of auxin in shoots 25 Ectopic expression of WIN1 causes changes in the levels of F-actin and differential cell enlargement between the excurvature and incurvature sides of the shoot spiral structure 27 Reference 47 Chapter 3 52 Perspectives: Future research for crop improvement with WIN1 52 Perspectives for WIN1 52 References 54 Table 1. Primers list for WIN1 study 30 Table 2. Accession number of WIN1 homologous proteins used for phylogenetic analysis 31 Figure 1. Overexpression of WIN1 confers the spiral shoot phenotype 33 Figure 2. Ectopic expression of WIN1 in aboveground tissues confers twisted abnormal development 34 Figure 3. Multiple alignment of WIN1, AtNPH3 and CPT1 showing the low identity of amino acid sequences 35 Figure 4. WIN1 overexpression does not alter the phototropism of coleoptiles 36 Figure 5. Phylogenetic tree of WIN1 homologous proteins in plants 37 Figure 6. Overexpression of WIN1 in Arabidopsis does not cause a spiral shoot or twisted leaf phenotype 38 Figure 7. WIN1 is highly expressed in roots and the shoot/root junction region 39 Figure 8. WIN1 is localized on the plasma membrane and partially co-localizes with a plasmodesmata marker protein 40 Figure 9. Treatment with auxins, auxin efflux transporter inhibitor or growth under constant darkness do not alter the spiral phenotype of WIN1-overexpressing plants 41 Figure 10. Abscisic acid (ABA), brassinolide and gibberellin could not recover the spiral phenotype in rice ectopically-expressing WIN1 42 Figure 11. Uneven auxin distribution correlates with the winding phenotype in leaf sheaths of the WIN1Act line 43 Figure 12. Ectopic WIN1 expression causes differential cell expansion in leaf sheaths 44 Figure 13. F-actin networks are affected by ectopic expression of WIN1 45 Figure 14. Expression of two other genes causing rice spiral phenotype is not affected by ectopic expression of WIN1 46

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