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dc.description.abstractPhytoplasmas are wall-less bacterial plant pathogens. They are restricted to the cytoplasm of phloem sieve cells, and can not be cultured in artificial culture medium. Phytoplasmas are spread by phloem-feeding insects including leafhoppers, planthoppers and psyllids. The disease symptoms caused by phytoplasmas including yellowing leaf, greening flowers, proliferation of stems and phyllody. Those morphological changes are mainly caused by the secreted effectors produced by phytoplasmas. The whole genome of Aster Yellows phytoplasma strain witches’ broom (AY-WB) has been completely sequenced, and 56 candidate effectors has been identified and named secreted AY-WB proteins (SAPs). Among them, SAP11 contains a nuclear localization signal (NLS) for nucleus targeting. SAP11 has been shown to destabilize plant TCP transcription factor, in order to repress the expression of LIPOXYGENASE2, an essential enzyme for jasmonate (JA) biosynthesis. Although SAP11 is a small protein with a molecular weight of approximately 14 kDa, the molecular mechanism in disease developing is still not clear. Here, we found that the purified recombinant SAP11 protein was unstable and could not be crystallized. However, after gel-filtration chromatography analysis, we showed that SAP11 formed a multimeric protein. With RNA-seq analysis, we showed that the expression of Pi deficiency-induced genes such as IPS1, PS2, PHT1;4, IPS2... were triggered by SAP11. Because Pi is an essential element for intracellular pathogens, the regulation of Pi homeostasis may contribute to the growth of phytoplasmas in host cells. In addition, we found that the expression of defense response genes such as PR1, WIN3, PAD4… were also suppressed by SAP11. This data suggests that phytoplasmas repress plant immune responses via secreted SAP11.en_US
dc.description.abstract植物菌質體(phytoplasma)為一種無細胞壁植物病原菌,寄生在宿主篩管細胞內,且至今尚無法以人工方式培養。菌質體主要藉由葉蟬、飛蝨等媒介昆蟲吸食植物韌皮部組織汁液而傳播。菌質體感染的植株具有發育遲緩、植株矮化、節間縮短、莖葉叢生、葉片捲曲、黃化、花器綠化或葉化等外表型態的改變。目前已知這些病徵的表現和菌質體分泌的作用因子(effector)有關。本實驗研究的材料為翠菊黃萎病(aster yellows)中發現的植物菌質體(Aster Yellows phytoplasma strain Withches’ Broom,AY-WB)。此菌質體的序列已被解序,而其基因組可轉譯出56個分泌性的AY-WB蛋白,簡稱SAPs。SAP11為其中之一,帶有細胞核定位信號(nuclear localization signals,NLSs)且可進入宿主細胞的細胞核,會促進TCP轉錄因子(TCP transcription factors)的降解,進而抑制LOX2的表現,使茉莉酸(jasmonate,JA)生合成量下降。SAP11大小約14 kDa,致病機制尚未清楚。因此本實驗期望能藉不同的實驗方法以更進一步了解SAP11的功能。首先在大腸桿菌中表現重組SAP11蛋白並純化後,藉由分子篩選層析管柱分析可知SAP11會形成聚分子。但因蛋白分子結構不穩定,不易形成結晶。進一步利用次世代定序方式進行SAP11轉殖株與野生種擬南芥的轉錄體分析,發現SAP11的表現會誘發植物產生缺磷反應,其中IPS1、PS2、PHT1;4、IPS2、…等缺磷誘導的基因皆大量表現。由於磷是許多病原菌的必需生存元素,因此調控磷的吸收可能有助於菌質體的繁殖。另一方面,發現表達SAP11會抑制植物產生免疫反應,PR1、WIN3、PAD4、…等水楊酸相關基因有表現量降低或表現時間延後的現象。而這與先前研究指出表現SAP11可提高菌質體昆蟲宿主繁殖後代量之結果相符。因此推測菌質體可能透過分泌SAP11到植物體內降低免疫反應,以提高自身以及傳播之昆蟲宿主繁殖與存活率。zh_TW
dc.description.tableofcontents謝誌 中文摘要 i Abstract ii 目次 iii 圖表目次 v 附錄目次 vi 第一章 緒論 1 一、前言 1 二、文獻回顧 2 植物菌質體(phytoplasma)簡介 2 翠菊黃萎病菌質體 4 翠菊黃萎病菌質體之分泌系統與作用因子 4 其他植物菌質體與所分泌之作用因子 7 磷營養源對植物與病原菌的影響 9 植物auxin生長素生合成與防禦機制之關聯性 13 植物誘導水楊酸之防禦機制 14 三、研究目的與策略 17 第二章、材料與方法 18 ※材料 18 一、研究中所使用之菌株、質體、抗生素、增幅SAP11AYWB的核苷酸引子、各式培養基與溶液 18 二、研究中所使用之植物 (詳見附錄九) 18 ※方法 18 一、載體之構築 18 二、擬南芥轉殖株之建立 20 三、菸草的螢光蛋白短暫表現系統之農桿菌注射法(Agroinfiltration) 22 四、分析擬南芥轉殖株之RNA表現量 22 五、重組蛋白質之表現 24 六、蛋白質結晶 27 七、抗體製備與純化 27 八、擬南芥之水耕系統與缺磷反應 28 第三章 結果 29 一、表現重組SAP11蛋白並純化及anti-SAP11多株抗體製備與測試 29 二、利用次世代定序進行SAP11轉殖株與野生種擬南芥之轉錄體分析 30 三、利用qRT-PCR驗證次世代定序結果 31 四、SAP11誘發植物產生缺磷反應 31 五、miR319a並非缺磷訊號傳遞鍊的標靶 32 六、SAP11表現會使PHO2基因表現被抑制 32 七、SAP11抑制植物產生免疫反應 33 八、SAP11與其他菌質體之作用因子 34 九、SAP11之 C端保留性序列缺失不影響其誘導植物產生缺磷反應及抑制植物防禦的能力 34 第四章 討論 36 一、SAP11蛋白是否具有proteasome的活性 36 二、SAP11誘導植物產生缺磷反應 36 三、SAP11抑制植物的免疫反應 36 四、SAP11之C端保留性序列不影響外表型與缺磷反應 37 五、SAP11PnWB是否也會誘導植物產生缺磷反應 37 六、Auxin對於SAP11影響缺磷反應以及免疫反應所扮演的角色 38 第五章 參考文獻 39 第六章 圖、表 48 第七章 附錄 58zh_TW
dc.subjectPi deficiencyen_US
dc.subjectdefense responseen_US
dc.titleThe Phytoplasma Effector Modulates Phosphate Homeostasis and Defense Response in Arabidopsisen_US
dc.typeThesis and Dissertationzh_TW
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