臺灣博碩士論文加值系統

本論文以台農57號甘藷為實驗材料，建構甘藷受傷葉片之cDNA基因庫，以傷害誘導之DNA片段為探針，選殖其cDNA全長，並經基因序列之分析而得知此傷害誘導DNA為Ipomoelin (IPO) 基因，並且其氨基酸序列與凝集素(lectin)具有高度相似性。IPO基因全長為720個鹼基對，可轉譯成含154個胺基酸之蛋白質，利用南方氏墨點法可得知台農57號甘藷中具有兩個IPO基因。然而IPO基因在植物中並非普遍存在的，阿拉伯芥、菸草、蕃茄之受傷葉片中均無法測得IPO基因之表現，即使在甘藷中不同的品系其IPO基因表現量亦不相同。本論文中更進一步製備IPO蛋白質之抗體，以西方墨點法得知IPO蛋白質在甘藷葉片受傷後一天即會開始累積，並在七天後累積量大增。
在IPO蛋白質的功能分析實驗中，利用大腸桿菌大量表現IPO蛋白質，將此經純化之蛋白質以不同稀釋濃度塗抹於桑葉表面，餵食予二齡幼蠶，經過觀察發現幼蠶食用塗抹IPO蛋白質的桑葉會有體重減輕及存活率降低的現象，並且此現象與IPO蛋白質濃度相關，此結果證明IPO蛋白質為防禦相關蛋白質之一。在血液凝集試驗中發現IPO蛋白質可將人類紅血球凝集，並且此凝集作用可被某些醣類抑制，此結果更進一步證實IPO為醣類結合蛋白質之一。
IPO基因在甘藷中的訊息傳導途徑亦為本論文之重要課題， IPO基因在傷害之本片葉及系統葉片中均會被誘導表現，此外，茉莉酸甲酯 (MeJA)、多聚半乳糖醛酸 (PGA)、及脫乙醯殼多糖 (CHI) 等物質亦會誘發IPO基因之表現。甘藷受物理性傷害而誘導表現IPO基因經北方墨點法及共軛焦顯微鏡對鈣離子的觀察得知，其訊息傳遞途徑依序為鈣離子進入細胞、胞器內鈣離子釋放、甲基化茉莉酸生成、去磷酸化蛋白質亦參與其中之調控。然而在IPO基因受PGA誘導的訊息傳導途徑中，則發現蛋白質激脢也參與誘導IPO基因的表現。乙烯為植物荷爾蒙之一，並可誘發甘藷葉片IPO基因的大量表現，其訊息傳導途徑為乙烯促使鈣離子經由細胞膜上對鈣離子高親和性孔徑蛋白質進入細胞，引發細胞內胞器大量釋放鈣離子至細胞質，而後經由去磷酸化蛋白質、過氧化氫、蛋白質激脢MAPK等傳導信息，進而啟動IPO基因之表現。

In order to investigate the functions and applications of wound inducible genes in the plant defense response, sweet potato (Ipomoea batatas cv. Tainung 57) was used as the experimental materials in this study. In the previous study from our laboratory indicated that ipomoelin (IPO) gene from sweet potato was wound- and methyl jasmonate (MeJA)-inducible, but the function and signal transduction of IPO protein was not clear. To understand the basic traits of IPO gene, the full-length cDNA of IPO gene was obtained by screening the cDNA library of the wounded sweet potato leaves. The full-length cDNA of IPO gene contains 720 base pairs, and encodes 154 amino acids. Also, the amino acid sequence of IPO protein showed high homology with those of lectin proteins. Southern blotting assay demonstrated that IPO gene existed two copies within sweet potato. However, IPO gene induced by wounding was only present in sweet potato of Tainung 57 and Tainung 25, but not in Taoyuan 1 or Taoyuan 2. Moreover, IPO gene was not expressed in the wounded leaves of Arabidopsis, tobacco, and tomato. The result of Western blotting assay indicated that the IPO protein was accumulated in the day one after the treatment of wounding, and the maximum accumulation was in the day seven after wounding.
To study the function of IPO protein, His-IPO fusion protein was expressed in E.coli BL21(DE3), and two-week-old silkworms were fed the mulberry leaves smeared with His-IPO protein. The feeding experiment showed that His-IPO fusion protein reduces silkworm growth and survival rates. Additionally, the reduction of silkworm growth and survival rates was dosage-dependent. This result demonstrated that IPO protein was one of the defense-related proteins. Furthermore, His-IPO fusion protein agglutinates human blood and this agglutination was inhibited by some carbohydrates. This result further demonstrated that IPO protein was a carbohydrate-binding protein.
The regulation of IPO gene in sweet potato was also an important project in this study. IPO gene was induced in both local and systemic leaves of sweet potato after wounding. In addition, IPO gene was induced by MeJA, polygalacturonic acid (PGA), and chitosan (CHI) in sweet potato. Moreover, several factors, including calcium ion influx, calcium release from organelles, biosynthesis of MeJA, and protein dephosphorylation, were involved in the expression of wound-inducible IPO gene, and analyzed by Northern blotting assay and confocal scanning microscope. However, when IPO gene was induced by PGA, protein kinase was involved in the signal transduction pathway. On the other hand, ethylene, a plant hormone, induced large expression of the IPO gene in sweet potato leaves. The signals involved in the ethylene-induced IPO gene expression were via calcium ions influx through high affinity calcium channels into cytoplasma, calcium ions released from organelles, and the activation of protein phosphatase, hydrogen peroxide, and MAPK to promote the expression of IPO gene.
In this study, the DNA of IPO gene was cloned and the function of IPO protein was demonstrated to be involved in the plant defense response. Moreover, the regulation of IPO gene was also studied. These data help us understand how the stress-related IPO gene regulated by environmental factors, and further elucidate the complex regulation of gene expression with plants.

Abbreviations.................................................1
Chapter one:
Literature Review.............................................3
Abstract......................................................4
References...................................................11
Figures......................................................17
Chapter two:
Cloning and characterization of the wound-inducible and carbohydrate-binding protein ipomoelin from sweet potato.....18
Abstract.....................................................19
Introduction.................................................20
Materials and methods........................................22
Plant materials and assay conditions.........................22
mRNA isolation and cDNA library construction.................22
cDNA library screening.......................................23
DNA isolation and analysis...................................23
RNA isolation and analysis...................................24
Protein isolation and analysis...............................25
Anti-IPO serum preparation...................................25
Protein expression and purification..........................26
Hemagglutination test........................................26
Insect feeding experiment....................................27
Results......................................................28
IPO gene was cloned by cDNA library screening................28
Sweet potato contains two copies of the IPO gene.............28
IPO gene expression is restricted to certain varieties of sweet potato.......................................................29
Expression of IPO protein following wounding.................30
His-IPO fusion protein agglutinates human blood..............30
His-IPO fusion protein binds carbohydrates...................31
His-IPO fusion protein reduces silkworm growth and survival rates........................................................32
Discussion...................................................34
References...................................................38
Figures......................................................42
Chapter three:
Expression of the ipomoelin gene from sweet potato is regulated by dephosphorylated proteins, calcium ion, and ethylene......52
Abstract.....................................................53
Introduction.................................................54
Materials and methods........................................57
Plant materials and assay conditions.........................57
RNA isolation and analysis...................................57
Ca2+ detection by confocal scanning microscope...............58
Results......................................................60
Mechanical damage induces IPO gene expression................60
MeJA induces IPO gene expression via dephosphorylated proteins.....................................................60
Effects of ABA and SA on IPO gene expression.................61
Ca2+ is involved in the activation of IPO gene...............62
MeJA does not induce the accumulation of Ca2+................62
Ethylene is involved in the induction pathway of IPO gene....63
Discussion...................................................65
References...................................................70
Figures......................................................77
Chapter four:
Involvement of polygalacturonic acid and chitosan in the induction of a Wound-Inducible Gene, IPO, from Sweet Potato..86
Abstract.....................................................87
Introduction.................................................88
Materials and methods........................................91
Plant materials and assay conditions.........................91
RNA isolation and analysis...................................91
Results......................................................93
PGA and CHI stimulate the expressionof the IPO gene..........93
Protein phosphorylation and dephosphoryltion are involved in the expression of the IPO gene induced by PGA and CHI........93
Effects of calcium ion on the expression of the IPO gene induced by PGA and CHI.......................................94
MeJA is not a signal transducer for the induction of the IPO gene via PGA or CHI..........................................95
Discussion...................................................96
References...................................................98
Figures.....................................................102
Chapter five:
Ethylene induces the expression of the IPO gene via calcium ion, dephosphorylated protein, hydrogen peroxide, and MAPK..106
Abstract....................................................107
Introduction................................................108
Materials and methods.......................................111
Plant materials and assay conditions........................111
RNA isolation and analysis..................................111
Ca2+ Detection by confocal scanning microscope..............112
Results.....................................................114
Ethylene induces the expression of the IPO gene.............114
A23187 and ethylene induce the expression of Ca2+...........114
The accumulation of Ca2+induced by ethylene was inhibited by Neo, LaCl3 DILT, VERA, RR, but not EGTA.....................115
The accumulation of Ca2+ was not inhibited by PD98059.......115
Ethylene induces IPO gene expression via calcium ion releasing from organelle..............................................116
Effects of PD98059 on IPO gene expression...................116
A23187 does not induce the accumulation of IPO gene.........117
Hydrogen peroxide and MAPK are involved in the signal pathway of IPO gene expression......................................117
Discussion..................................................119
References..................................................125
Figures.....................................................130

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