探究徒長病對水稻苗期轉錄體之影響

鄭安伯

doi:10.6342/NTU201800550

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探究徒长病对水稻苗期转录体之影响

Exploring the Transcriptome of Rice Seedlings Affected by Bakanae Disease

郑安伯 , 硕士　　导师：锺嘉绫　　

英文 DOI： 10.6342/NTU201800550

水稻徒长病；转录体分析；防御机制； CRISPR/Cas9 ； bakanae disease ； transcriptome analysis ； defense-related mechanisms ； CRISPR/Cas9

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摘要 │ 参考文献 (352) │ 国际计量

摘要〈TOP〉

水稻徒长病 (bakanae disease) 为Fusarium fujikuroi所引起的种子及土壤传播性真菌病害，其侵染水稻後会造成植株徒长及/或叶片夹角加大等不正常生理现象，严重时可导致植株死亡或谷粒不充实。据报导，水稻徒长病曾在许多大规模水稻种植国家如日本、印度、泰国等地造成严重损失；台湾近年则於东部有较严重发病之情形。由於水稻抵御徒长病菌的相关机制目前研究甚少，本研究利用Illumina次世代定序技术，分析水稻抗病品系Tainung 67 (TNG67)及感病品系Zerawchanica karatals (ZK) 之幼苗基部受徒长病菌感染7天後之转录体，透过比对水稻与阿拉伯芥资料库进行基因功能预测，探讨与贺尔蒙及防御相关之基因表现变化。结果显示，F. fujikuroi接种後有显着表现差异之基因 (differential expression genes, DEGs)，在抗病品系TNG67有118个 [包括WRKY、plant pattern recognition receptor (PRR)及peroxidase]，在感病品系ZK有169个 (包括cytochrome P450、WRKY、PRR、glycoside hydrolase、thaumatin-like protein及peroxidase)。检视贺尔蒙相关之基因网路，发现吉贝素生合成、茉莉酸生合成、茉莉酸讯号传递路径於接种前後并无整体上升或下降趋势；而在抗性品系TNG67中，参与吉贝素讯号传递之OsGID1 (吉贝素受体；促进DELLA蛋白之降解) 表现量下降、OsSLR1 (DELLA蛋白；吉贝素讯息传递抑制因子) 则显着上升，显示抗病品种对吉贝素有负调控的现象。针对16个被注解为抗性相关之DEGs，利用real-time qRT-PCR测定额外三次独立实验中，TNG67与ZK两品种接种7、14、21天後之基因表现，其中八个基因在接种後7天与转录体分析结果具有一致性，且於不同次独立实验之趋势一致。本研究选择以CRISPR/Cas9技术进行基因功能丧失验证，首先以实验室前人研究所得之三个抗稻热病候选基因为对象，建立水稻CRISPR/Cas9系统，再应用於四个抗徒长病候选基因之突变。上述候选基因之single guide RNA (sgRNA) 质体，经T7核酸内切酶I进行水稻原生质体短暂表现之突变体检测後，筛选出sgRNA突变率超过10%者，继续进行水稻转殖株之建立。目前已获得其中一个基因之五个T0转殖品系，经基因型测定後，确认有四个品系於目标序列被成功突变。期望透过本研究了解水稻对徒长病之抗性网路及有潜力的抗性基因，为水稻育种及病害防治找到新的发展方向。

并列摘要〈TOP〉

Bakanae disease is a seedborne and soilborne fungal disease caused by Fusarium fujikuroi. The plants infected with the pathogen show symptoms such as abnormal stem elongation and/or wider angle of the leaf with the stem. Severe infection can cause seedling blight and sterile grains. In rice-growing countries such as Japan, India, and Thailand, there are reports on bakanae disease causing yield losses and great impact on rice production. In recent years, outbreaks of bakanae disease were observed in Taiwan, particularly in the eastern areas. Since little is known about the mechanisms involved in rice against F. fujikuroi, the research focuses on applying Illumina next-generation sequencing technology to analyze the transcriptome profiles of a resistant cultivar Tainung 67 (TNG67) and a susceptible cultivar Zerawchanica karatals (ZK), using the basal stems of rice seedlings collected after 7 days post inoculation (dpi) of F. fujikuroi. Genes were annotated according to the reference databases of Oryzae sativa (MSU7) and Arabidopsis thaliana (TAIR10). Expression profiles of genes associated with phytohormones and defense responses were examined. There were 118 differential expression genes (DEGs) in the resistant TNG67 [including WRKYs, plant pattern recognition receptor (PRR), and peroxidases] and 169 DEGs in the susceptible ZK (including cytochrome P450, WRKY, PRR, glycoside hydrolase, thaumatin-like protein, and peroxidase). For phytohormone biosynthesis and signaling pathways, overall up- or down-regulations were not observed in gibberellin (GA) biosynthesis, jasmonic acid (JA) biosynthesis, and JA signaling pathways. In TNG67, the GA signaling pathway seemed to be suppressed according to the down-regulation of OsGID1 (gibberellin receptor; induced degradation of DELLA proteins) and up-regulation of OsSLR1 (DELLA protein; GA-insensitive gene homolog) after F. fujikuroi inoculation. Real-time quantitative RT-PCR was conducted to evaluate the expression levels of 16 defense-related DEGs at 7, 14, and 21 dpi in additional three independent inoculation trials. A set of 8 candidate genes, with 7-dpi expression patterns similar to the transcriptome data and consistency among different replicated trials, were selected for loss-of-function verification using CRISPR/Cas9 system. Three blast-resistance candidate genes identified in our another study were used as targets to set up CRISPR/Cas9 system, which was subsequently applied to four candidate genes associated with bakanae resistance. Mutation efficiencies of single guide RNA (sgRNA) vectors were tested in rice protoplasts using T7 endonuclease I (T7E1) assay, and the constructs showed mutation efficiencies of >10% were sent for generating transgenic rice plants. Currently five T0 mutant lines for one candidate gene were obtained. These lines have been genotyped and four lines were successfully mutated at the target site. Uncovering potential resistance-related genes and mechanisms will provide new insights into resistance breeding and disease control of bakanae disease.

参考文献 ( 352 ) 〈TOP〉

Amatulli, M., Spadaro, D., Gullino, M., and Garibaldi, A. 2010. Molecular identification of Fusarium spp. associated with bakanae disease of rice in Italy and assessment of their pathogenicity. Plant Pathology 59:839-844.
连结：
Anders, S., and Huber, W. 2010. Differential expression analysis for sequence count data. Genome Biology 11:R106.
连结：
Athwal, D. 1971. Semidwarf rice and wheat in global food needs. The Quarterly Review of Biology 46:1-34.
连结：
Belhaj, K., Chaparro-Garcia, A., Kamoun, S., and Nekrasov, V. 2013. Plant genome editing made easy: targeted mutagenesis in model and crop plants using the CRISPR/Cas system. Plant Methods 9:39.
连结：
Boatwright, J. L., and Pajerowska‐Mukhtar, K. 2013. Salicylic acid: an old hormone up to new tricks. Molecular Plant Pathology 14:623-634.
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使用DOI作为永久连结

引用含有DOI的文献

DOI可强化引用精确性、增强学术圈连结，并给予使用者跨平台的良好使用经验，目前在全世界已有超过五千万个对象申请DOI。 如想对DOI的使用与概念有进一步了解，请参考 （ ） 。

DOI可强化引用精确性、增强学术圈连结，并给予使用者跨平台的良好使用经验，目前在全世界已有超过五千万个对象申请DOI。如想对DOI的使用与概念有进一步了解，请参考（）。