臺灣博碩士論文加值系統

土壤環境中鹽類含量過高會導致植物處於鹽逆境與滲透逆境，甚至進一步產生氧化逆境。肌醇(inositol)為一相容質，於水份含量相關之鹽、乾旱及低溫逆境下會大量累積於細胞質以協助植物適應逆境，此外肌醇衍生物也參與植物適應逆境的過程，例如pinitol、galactinol、raffinose與ascorbate等，除了具有相容質之功能外，也具有清除ROS (reactive oxygen species)之能力。文獻中指出，耐鹽模式植物冰花(Mesembryanthemum crystallinum)於鹽逆境下可藉由肌醇運輸蛋白(inositol transporter; INT) McINT4.1/MITR1與McINT4.2/MITR2使鈉離子於植物體內重新分佈，以降低根部鈉離子累積，為冰花適應鹽逆境的重要機制之一。經由冰花轉錄體搜尋出7個INT基因，以親緣關係和序列相似性依照阿拉伯芥INT家族重新命名為：McINT1.1、McINT1.2、McINT2、McINT3、McINT4.1、McINT4.2與McINT4.3。本論文主要探討外源性肌醇對鹽逆境下冰花小苗生長、鈉鉀離子含量及肌醇運輸機制以及McINT家族表現量差異。依據本實驗室先前培養無菌冰花小苗的條件，冰花小苗容易有玻璃質化的現象，離開無菌環境後容易脫水造成實驗誤差，故本論文測試了培養基的組成與培養條件，發現藉由通氣可降低小苗玻璃質化的現象，成功獲得健康且均質化的小苗進行後續實驗。此外，本實驗室先前處理冰花小苗的方式有小苗處理不完全與使小苗處於淹水逆境之疑慮，而本論文也改以直立浸泡的方式解決以上問題，並且可以使同一培養皿內之冰花小苗同時進行多種處理，以降低可能因培養於不同培養皿所造成的誤差。本論文觀察到鹽逆境下添加外源性肌醇可以減緩冰花小苗脫水的現象，而檢測McINT表現量，發現於不同溶液處理下，地上部與地下部McINT有不同的表現趨勢。地上部所有McINT皆受到鹽逆境而表現量上升，其中地上部McINT3、McINT4.2與McINT4.3受到肌醇負調控有表現量下降的趨勢，而於地下部，鹽逆境下無論有無外源性肌醇，McINT皆有表現量上升的趨勢，且沒有任何McINT受到肌醇負調控，顯示不同群的McINT有特定的功能與調控機制。於追蹤外源性肌醇與分析鈉離子累積之實驗中發現，冰花小苗具有吸收環境中微量肌醇的能力，且鈉離子可以促進外源性肌醇之吸收，於長時間鹽逆境下，添加外源性肌醇可以降低根部鈉鉀比值。植物於鹽逆境下會導致氧化逆境的發生，而本論文發現冰花小苗於各處理下其ROS的累積程度並沒有明顯的差異，此結果可能因背景值過高而需要進一步進行定量分析。綜合上述，本論文初步鑑定了冰花小苗McINTs之功能及參與調控機制，冰花小苗可藉由增加基因表現或蛋白活性促進鈉離子累積於地上部並降低地下部的鈉鉀比值，以增加小苗對鹽逆境的耐受性。

Plants develop many mechanisms to adapt to the change of environment, but if the environment becomes more extreme, plants fail to respond and result in the stresses. Stresses can be distinguished into two classes, one is the biotic stress that is involved in interaction with other organisms, and the other is the abiotic stress caused by the change of the environmental factors. Excessive salt content in the soil environment will cause plants encountering salt, osmotic and even oxidative stress. Inositol is a compatible solute that accumulates in the cytosol and organelles to facilitate plant to adapt to water-deficit related stresses. Inositol derivatives such as pinitol, galactinol, raffinose and ascorbate are also involved in the processes of stress adaptation. These derivatives are also compatible solutes and have the ability to scavenge the ROS (reactive oxygen species). Halophyte ice plant (Mesembryanthemum crystallinum) is a model organism to study plant salt tolerance. Ice plant can redistribute the sodium ions in the plant under the salt stress via inositol transporter (INT) McINT4.1/MITR1 and McINT4.2/MITR2. It is one of the vital mechanisms for ice plant to adapt to salt stress. We identified seven INT genes form ice plant transcriptome and, according to the classification of Arabidopsis thaliana AtINT family, renamed as McINT1.1, McINT1.2, McINT2, McINT3, McINT4.1, McINT4.2 and McINT4.3. According to the previous culture condition of the ice plant seedlings, the seedlings tended to develop vitrification in the sealed condition. To solve the problem, we modified the compositions of the culture medium and culture conditions and successfully obtained healthy and uniform seedlings. Furthermore, the previous method of seedling treatment has the doubts about the incomplete treatment of roots and seedlings were under flooding stress. Therefore, a vertical immersion of seedlings was used to avoid the problem described. I found that the supply of exogenous myo-inositol can reduce the dehydration effects of salt-stressed seedlings. Differential expression analysis of McINTs showed that all McINTs were induced by salt stress at the shoot, and the expressions of McINT3, McINT4.2 and McINT4.3 were down-regulated by inositol. In root, the expression of all McINTs also induced under salt stress with or without exogenous inositol supply, and none of McINTs was down-regulated by inositol. This result indicated that different McINTs might have different functions and regulation in ice plant seedlings. Analyses of the uptake of exogenous inositol and accumulation of sodium ions revealed that ice plant seedlings had intrinsic ability to take up a trace amount of external inositol and the presence of sodium could facilitate inositol uptake. Under prolong salt stress, an exogenous supply of inositol decreased the Na/K ratio in roots. Oxidative stress is usually induced when plant suffer in salt stress. Ice plant seedlings have no significant differences in ROS accumulation by salt treatments. Quantification of ROS is needed for further confirmation. In conclusion, I identified the possible functions and regulatory mechanisms of ice plant McINT. Ice plant seedlings can accumulate more sodium in shoot and decrease the Na/K ratio in the root via induced specific McINT gene expression or activated transport activity of specific member of McINT to enhance the tolerance of salt stress.

摘要 i
Abstract ii
目次 iv
表目次 vi
圖目次 vii
壹、前言 1
一、鹽逆境對植物生長之影響 1
二、植物適應鹽逆境之機制 2
(一) 離子運輸與儲存 2
(二) 小分子相容質之累積 3
(三) 清除活性氧之途徑 3
三、肌醇在植物對抗逆境之角色位置 4
(一) 肌醇生合成與甲基化的肌醇 5
(二) 活性氧清除分子 6
(三) 肌醇運輸蛋白 7
四、模式生物冰花 9
五、研究目的 10
貳、材料與方法 11
一、實驗材料 11
(一) 冰花(Mesembryanthemum crystallinun)小苗無菌栽培 11
(二) 培養基與處理溶液配方 11
(三) 冰花小苗測試處理 11
二、實驗方法 12
(一) McINT之基因表現量分析 12
1. 冰花total RNA萃取 12
2. 反轉錄聚合酵素連鎖反應(Reverse transcription- polymerase chain reaction, RT-PCR) 13
3. 即時定量反轉錄聚合酵素連鎖反應(Quantitative reverse transcription polymerase chain reaction, qRT-PCR) 13
4. 洋菜膠體電泳分析 (DNA agarose electrophoresis) 14
(二) 檢測放射性肌醇[3H]myo inositol吸收與運輸 14
(三) 檢測冰花小苗植體內Na及K含量 15
1. 坩鍋前置處理 15
2. 樣品高溫灰化及強酸萃取 15
(四) 活性氧(reaction oxygen species, ROS)檢測 15
1. Nitroblue tetrazolium (NBT)染色分析 15
2. 3,3’-diaminobenzidine (DAB)染色分析 15
參、結果 17
一、冰花McINT序列比對與命名 17
二、無菌栽培冰花小苗最適生長條件 18
三、醣類分子影響冰花小苗McINT表現量 19
四、冰花小苗經鹽和myo-inositol處理後之外表型變化 19
五、冰花小苗鹽逆境下添加外源性肌醇對McINTs表現分析(RT-PCR) 20
六、冰花小苗鹽逆境下添加外源性肌醇對McINTs表現定量分析(qRT-PCR) 20
七、鈉離子運輸累積位置分析 21
八、外源性肌醇運輸累積位置分析 21
九、過氧化氫與超氧陰離子累積分佈位置 22
肆、討論 23
一、McINTs功能與調控機制之探討 23
二、鹽逆境下肌醇與Na分佈 24
三、外源性肌醇與ROS清除之探討 25
四、總結與未來展望 26
伍、參考文獻 27
陸、附錄 61

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