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研究生：	南宮乖 Nainggolan, Paul Fernando Haposan
論文名稱：	bHLH2 轉錄因子對水稻花粉發育之探討 Role of bHLH2 in Rice Pollen Development
指導教授：	郭瑋君 Guo, Woei-Jiun
共同指導教授:	辜瑞雪 Ko, Swee-Suak
學位類別：	碩士 Master
系所名稱：	生物科學與科技學院 - 熱帶植物科學研究所 Institute of Tropical Plant Sciences
論文出版年：	2017
畢業學年度：	105
語文別：	英文
論文頁數：	72
中文關鍵詞：	水稻、轉錄因子bHLH 、雄性不孕、花粉發育、絨氈層細胞程序化凋亡
外文關鍵詞：	Oryza sativa, bHLH, male sterility, pollen development, tapetal PCD
相關次數：	點閱：201 下載：15
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摘要

無論是在植物或動物的染色體中，bHLH轉錄因子都是很大的基因家族。bHLHs轉錄因子的功能通常是參與調控代謝作用以及組織的發育。像是UDT1 (bHLH164)、TDR (bHLH5) 、EAT1 (bHLH141) 、bHLH142 (TIP2) 這些轉錄因子，在花粉發育過程中扮演極重要的角色。轉錄因子 bHLH2的基因註解為「與ICE1 相近的轉錄因子」。ICE1轉錄因子已被證實具有抗逆境的功能，但目前並無參花粉發育之報導。故本研究就bHLH2 可能參與花粉發育做進一步探討。
qRT-PCR的結果指出bHLH2基因表達在葉鞘、幼穂、花葯、子房及種子中。進一步利用原位雜交技術發現 bHLH2 基因表達在花葯發育早期之花粉壁和減數分裂細胞中；且該基因持續表達在子房發育過程中的心皮及胚珠。從基因表達模式指出bHLH2可能涉及水稻雄性和雌配子體的發育。藉由韓國T-DNA突變系 (bhlh2)進行詳細的研究。剔除了bHLH2 的突變系的花粉為部份雄不稔且結實不良。基因分型證實T-DNA插入在bHLH2 基因的第3個外顯子。Southern blot數據顯示，該突變系分離成具有一個或三個T-DNA插入的個體，惟插入一個T-DNA的純合子用於進一步的分生研究。TUNEL測定發現bhlh2突變體有延長絨氈層細胞程序化凋亡(PCD)的現象。一些RNAi-bHLH2轉基因水稻明顯降低了bHLH2 基因全長的表達量，且造成不結實的情形很嚴重。在小孢子期，控制PCD標記基因（AP37、AP25 和CP1）和花粉脂質生物合成標記基因（MS2、CYP704B2 和C6）在RNAi-bHLH2轉殖株明顯減少基因的表達量。我們推測T-DNA突變系插在bHLH2下游，具有截短的bHLH2蛋白（保留完整的bHLH結構域），相較於RNAi-bHLH2轉殖株有更好的稔實度，意味著bHLH domain之重要。TRIM　pTag8激活之突變系，持續增加bHLH2 基因的表達量，但其花粉活力佳。總而言之，本研究發現bHLH2有參與水稻的花粉發育。

Abstract
The basic helix loop helix (bHLH) transcription factors (TFs) is a large gene family and diverse in animal and plant genomes. Consistent with functions of TFs, bHLHs play many important roles in regulating metabolism and tissues development. bHLH TFs such as UDT1 (bHLH164), TDR (bHLH5), EAT1 (bHLH141), bHLH142 (TIP2) had been reported play an essential role in pollen development. bHLH2 is annotated as “Similar to Transcription factor ICE1”. ICE1 had been reported related to stresses tolerance but so far not been reported an association with pollen development. This study aims to characterize the role of bHLH2 in rice pollen development.
The bHLH2 is broadly expressed in leaf sheath, young panicles, anther, ovary, and seed. In situ hybridization data indicated that bHLH2 expressed highly in the anther walls, meiocyte at early stage of anther development. Moreover, bHLH2 is highly expressed in carpel and ovule during ovary development. The expression pattern suggested that bHLH2 might involve in reproductive organs development. A Korea Postech T-DNA mutant (bhlh2) was carried out detail study. The bhlh2 mutant showed partial pollen viability and grain fertility. Genotyping confirmed T-DNA was inserted in the 3rd exron of bHLH2. Southern blot data indicated that this mutant has three T-DNA insertions but single insertion homozygote was used for the further molecular study. TUNEL assay indicated that bhlh2 mutant has prolonged tapetal PCD. RNAi-bHLH2 transgenic lines indicated that some RNAi lines expressed a low level of FL cds bHLH2 and have more severe in grain infertility. PCD markers (AP37, AP25, and CP1) and lipid biosynthesis markers (MS2, CYP704B2, and C6) were down-regulated in RNAi lines at YM. We hypothesize that T-DNA homozygote line has truncated form of bHLH2 (retains conserve bHLH domain) has less sterility problem than RNAi transgenic line. That implies the importance of bHLH domain for normal biological function. TRIM activation homozygous line showed constitutively upregulated of bHLH2 and it maintains good pollen fertility. Overall, this study indicated that bHLH2 contributes to pollen development in rice.

Chinese abstract	1
Abstract	2
Acknowledgement...	3
Index...	4
I.Introduction	11
1. Reproductive organs development	11
2. Basic helix-loop-helix transcription factor	12
3. Tapetal programmed cell death (PCD)	13
4. Sporopollenin biosynthesis	13
5. Biological function of  ICE1	13
6. Preliminary results of bhlh2	14
II. Material and Methods	15
1. Phylogenetic analysis	15
2. Information of Vector of bhlh2 mutants	15
3. Sample collection	15
4. Total RNA isolation method	16
5. RNA In situ Hybridization	16
6. Genotyping to confirm T-DNA insertion site	17
7. Southern blot analysis	17
8. Agronomy traits evaluation	18
9. qRT-PCR analysis	18
10. TUNEL (TdT-mediated dUTP Nick-End Labeling) assay	18
11. Pollen viability test	19
12. RNAi-mediated gene silencing of bHLH2	19
13. bHLH2 activation tagging TRIM	20
III. Results	21
1. Phylogenetic analysis of bHLH2	21
2. Gene expression patterns of bHLH2 in various tissues	21
3. bHLH2 expression in the anther and ovary analyzed by In situ hybridization (ISH)	22
4. Genotyping confirmed T-DNA inserted in bHLH2	22
5. Southern blot analysis T-DNA copy number in mutant	23
6. bhlh2 mutant defective in seed development	23
7. Agronomy traits and grain yield loss in bhlh2 mutant	23
8. Defective of anther development in bhlh2 mutant	24
9. Prolonged PCD in bhlh2 anther	24
10. bHLH2 reduced PCD in ovary	25
11. bHLH2 RNAi transgenic line defected in pollen development	25
12. Agronomic traits and grain yield loss in RNAi-bHLH2	26
13. Alteration in the expression of key genes associated with PCD  
      and lipid biosynthesis	26
14. Gain of function using activation-bHLH2	27
15. bHLH2 might regulates PSS1 expression 	28
IV. Discussion	29
1. bHLH2 involves in rice pollen development	29
2. Prolong PCD in bhlh2 mutant cause male sterility	30
3. Future prospective	30
4. Summary of this study	31
5.Table	32
6. Figure	37
7. Supplementary data	61
References	63
Appendix	68



Table content 
Table. 1 Primer List of this study	32
Table. 2 Agronomy traits and grain yield loss in bhlh2 mutant	34
Table. 3 Agronomy traits and grain yield in RNAi-bHLH2  
               transgenic rice.	35
Table. 4 Agronomy traits and grain yield loss activation   
               tagged mutant line	36

Fig.1   Scheme of bHLH2 gene and vector of pGA2715 and pGA2707	37
Fig. 2  Multiple alignments of bHLH domain	38
Fig. 3  Phylogenetic tree of bHLH2, its homologues and pollen   
           marker  gene	39
Fig. 4  Gene expression pattern of bHLH2 in various tissues    
            of Hwa Yang.	40
Fig.5   In situ hybridization of bHLH2 expression pattern in the
            anther  and ovary of Hwa Yang during spikelet    
             development.	41
Fig.6  Genotyping of bhlh2 mutant	42
Fig.7  Southern blot analysis T-DNA copy number in bhlh2   
           mutant.	43
Fig.8  Phenotype of bhlh2 mutant	44
Fig.9  bhlh2 mutant reduced grain fertility and slightly reduced seeds size.	45
Fig.10  bhlh2 mutant reduced in pollen viability	46
Fig.11 Loss function of bHLH2 defects in anther wall and   
            microspore development	47
Fig. 12 Prolonged tapetal programmed cell death in bhlh2   
             anther by TUNEL assay.	48
Fig 13 Comparison of PCD in ovary between the wild-type and   
             bhlh2 mutant.	49
Fig. 14 Alter PCD markers genes in bhlh2 mutant.	50
Fig. 15 Alter lipid biosynthesis markers genes in bhlh2 mutant.	51
Fig. 16 Knockdown bHLH2 using RNAi transgenic approach.	52
Fig. 17 Phenotype of RNAi-bHLH2.	53
Fig. 18 RNAi-bHLH2 transgenic plants reduced pollen fertility	54
Fig. 19 Altered PCD marker gene expression in the anther of RNAi-  
              bHLH2 transgenic lines.	55
Fig. 20 Altered lipid biosynthesis marker gene expression in the anther   
            of RNAi-bHLH2 transgenic lines.	56
Fig. 21 TRIM mutant activated bHLH2	57
Fig. 22 Phenotype of activation bHLH2	58
Fig. 23 Binding site prediction of bHLH2 on PSS1 promoter region and altered expression of PSS1 in knock out mutant and RNAi silencing of bHLH2.	59
Fig. 24 Proposed model of bHLH2 in this study.	60 
Supplementary Data
Supplementary Fig. 1  Phenotype of bhlh2 mutant.	61
Supplementary Fig. 2  Southern blot indicated bhlh2 mutant has   
                                two copies of T-DNA insertion.	62 
Appendix  

Appendix. 1  Rice flower anatomy .	68
Appendix. 2  Rice anther  morphology ……………………….……..........69 
Appendix. 3  Known genes involved in pollen development pathway.	70
Appendix. 4  Rice expression Profile by RiceXpro database .	71
Appendix. 5  OE bHLH2 causing lethal transgenic plants.	72

                                    

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