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dc.description.abstractRice (Oryza sative L.), a graminaceous plants, uses the Strategy II mechanism to absorb Fe from soil, but previous studies also demonstrated that rice can take up Fe (II) directly through OsIRT1 and OsIRT2. During rice growing season, paddy soil is submerged. The submerging conditions result in the temperal and spatial variations in the concentrations of Fe (II) and Fe (III) in the soils. Therefore, we hypothesized that rice might be able to regulate the uptake of Fe from paddy soil. Accordingly, Fe uptakes of rice seedlings in different Fe (II)/Fe (III) conditions were investigated. Rice plants were grown in hydroponic solutions with different concentrations of Fe (II)/ Fe (III), ranging from deficient conditions to toxic conditions. FeSO4:NH2OH=1:1and Fe (III)-EDTA was used as the sources of Fe (II) and Fe (III), respectively. In the treatments of lower Fe concentrations (0.17~1.7 mg Fe L-1), the result showed that rice can take up Fe more efficiency in Fe (II) system than the Fe (III) counterparts. Therefore the OsIRO2 gene expression in the Fe (II) system is lower than that in the Fe (III) system. In the treatments of higher Fe concentrations (17~85 mg Fe L-1), Fe concentrations in rice plants increased with increasing Fe concentration in hydroponic solution, especially in the Fe (II) system. Moreover, the Fe concentrations of plants in the Fe (II) system are much higher than their Fe (III) counterparts. The Fe concentrations in iron plaque and in shoot are highly correlated and the formation of iron plaque changed with increasing Fe concentration in the solution. The content of ferrihydrite in the iron plaque is also increased. These results indicated that the concentration of Fe in iron plaque and the formation of iron plaque determine the iron uptake of rice. Nonetherless, the Fe concentration in iron plaque was not correlated to root oxidative capacity/ hydrogen peroxide content. The results of stable Fe isotope fractionation showed that Fe (II) undergoes an oxidative precipitation mechanism on rice root surface but can also be taken up directly by rice root. Thus, Fe (II) is oxidized to Fe (III), which is then bound to phytosiderophores and the resultant Fe (III)-PS complexes were taken by rice root. In the Fe (II) system, rice can use more than one uptake strategies and therefore exhibits a higher Fe uptake efficiency.en_US
dc.description.abstract水稻 (Oryza sative L.)屬於禾本科植物,主要是利用策略II吸收Fe (III),但水稻也被證實能透過OsIRT1和OsIRT2的載鐵蛋白,直接的吸收Fe (II)。由於水田土壤中Fe (II)/ Fe (III)的濃度為一動態變化,水稻為適應此土壤環境,應有一套調控鐵吸收的機制,因此本研究為探討水稻幼苗在不同Fe (II)/ Fe (III)濃度 (包含鐵缺乏到鐵毒害) 的水耕條件下對鐵的吸收機制,所使用的水稻品種為台梗九號 (Tai- keng 9),並分別使用FeSO4:NH2OH=1:1和Fe (III)-EDTA做為水耕液中的Fe(II)和Fe(III)來源。其結果顯示在低濃度 (0.17~1.7 mg Fe L-1)處理下,水稻在Fe (II)系統中有較好的吸收效率,OsIRO2的基因表現量較Fe (III)系統低;在高濃度 (17~85 mg Fe L-1)處理下,Fe (II)/ Fe (III)系統水稻植體中的鐵濃度皆隨水耕液鐵濃度的增加而增加,尤以Fe (II)系統中植體莖部的鐵濃度為最。然而,在各濃度處理下,Fe (II)系統植體中的鐵濃度卻大於Fe (III)系統。從水稻根表鐵膜的生成與水耕液的鐵濃度有顯著的正相關、鐵膜的鐵濃度和莖部鐵濃度顯著正相關,以及Fe (II)系統中鐵膜的水鐵礦組成會隨鐵濃度的增加而增加的結果,此皆顯示鐵膜的鐵濃度和礦物組成會影響水稻對鐵的吸收。不過,鐵膜的生成量與根部的總氧化力和H2O2的含量卻無顯著的相關性。以鐵的穩定同位素來探討水稻對鐵的吸收機制上發現,在根部除了有鐵的氧化沉澱外,在Fe (II)系統中也發現有Fe (II)直接吸收的現象。在莖部則顯示鐵膜中的鐵會與載鐵物質 (phytosiderophores)鉗合後,形成Fe (III)-PS的鉗合物再吸收,以及鐵膜還原溶解後再吸收的機制,但隨水耕液中的鐵濃度增加,則顯示鐵會直接進入水稻植體中。相較於Fe(III)系統,生長於Fe (II)系統的水稻具有較多的鐵吸收策略,因此在低鐵濃度的情況下鐵的吸收效率較高;但隨鐵濃度的增加,無論是在Fe (II)/ Fe (III)系統,皆有大量的鐵被吸收進入水稻植體當中。zh_TW
dc.description.tableofcontents摘要 I Abstract III 目錄 V 圖次 IX 表次 XII 第一章 前言 1 1.1 研究緣起 1 1.2 研究目的 3 第二章 文獻回顧 4 2.1鐵 4 2.1.1 土壤中的鐵 4 2.1.2 植物中的鐵 5 2.2 水田土壤鐵化學 6 2.2.1 浸水土壤氧化還原 6 2.3植物鐵缺乏和鐵毒害的相關機制 9 2.3.1 鐵缺乏 9 2.3.2 水稻鐵吸收機制 9 2.3.3水稻缺鐵診斷-缺鐵基因 (OsIRO2)分析 12 2.3.4 鐵毒害 14 2.3.5活性氧化物質(Reactive Oxygen Stress, ROS) 15 2.4 水稻根部的氧化能力 18 2.5 水稻根表鐵膜的生成 19 2.6 鐵同位素組成和分化作用 20 第三章 材料方法 23 3.1 水稻幼苗生長試驗 23 3.1.1育苗方法 23 3.2 試驗方法 26 3.2.1 前置實驗 26 試驗一:不同還原劑 (FeSO4:還原劑=1:90)添加抑制水耕液Fe (II)氧化 26 試驗二: 還原劑 (NH2OH)的添加對水稻生理的影響 27 試驗三: 不同濃度Fe (II)水耕液配置 28 試驗四: 缺鐵處理時間和缺鐵基因 (OsIRO2)表現時間 29 試驗五: 不同Fe (III)濃度下,誘導水稻缺鐵基因表現時間和濃度 30 3.3水稻在不同Fe (II)和Fe (III)濃度下,對鐵的吸收和運輸機制 31 3.4實驗分析方法 32 3.4.1 水耕液收集和Fe (II)比色法 32 3.4.2 植體採收和量測 32根表鐵膜鐵含量測定:DCB萃取法 33 植體鐵含量分析 33 3.4.3統計分析 34 3.4.4 鐵穩定同位素分析 34 水稻樣品前處理 34 鐵穩定同位素純化分析 35 3.4.5 根部總氧化力分析 35 3.4.6 H2O2定量分析 37 3.4.7 基因分析 37 RNA萃取 37 RNA定量 38 DNAase處理 38 合成cDNA 39 Real Time PCR 39 3.4.8水稻切片製成、鐵膜分佈分析和穿透式X光顯微鏡分析 40 鐵膜礦物組成分析 40 3.4.9轉移因子 (Translocation Factor, TF)以及生物濃度係數 (Bioconcentration Factor, BCF) 41 第四章 實驗結果 42 4.1前置實驗 42 試驗一: 不同還原劑(FeSO4:還原劑=1:90)添加抑制水耕液Fe (II)氧化 42 試驗二: 還原劑 (NH2OH)的添加對水稻生理的影響 46 試驗三: 不同濃度Fe (II)水耕液配置 55 試驗四: 缺鐵處理時間和缺鐵基因 (OsIRO2)表現時間 57 試驗五: 不同鐵濃度處理下誘導水稻缺鐵基因 (OsIRO2)表現時間 59 4.2 水稻在不同Fe (II)和Fe (III)濃度下,對鐵的吸收和運輸機制 61 4.2.1 Fe (II)/ Fe (III)水耕液殘留和移除之總鐵濃度 61 4.2.2 以不同濃度Fe (II)/ Fe (III)水耕液處理之水稻生長情形 64 4.2.3 水稻植體對鐵的累積與轉移 71 4.2.4水稻植體內的鐵含量及其分佈百分比 80 4.2.5 水稻缺鐵基因 (OsIRO2)表現 85 4.2.6 水稻幼苗根部IRT1、YSL15和TOM1的基因表現 87 4.2.7 水稻鐵膜分布和鐵膜的礦物組成 90 4.2.8 鐵膜生成與地上部鐵濃度累積之關係 97 4.2.9 水稻根部氧化力和H2O2與鐵膜生成之影響 101 4.2.10 水稻根部和莖部H2O2生成 104 4.2.11 水稻同位素變化 106 第五章 討論 110 第六章 結論 126 第七章 參考文獻 128zh_TW
dc.subjectRice (Oryza sative L.)en_US
dc.subjectIron uptakeen_US
dc.subjectIron redoxen_US
dc.subjectRoot oxidative activityen_US
dc.subjectFe stable isotopeen_US
dc.subject水稻 (Oryza sative L.)zh_TW
dc.titleIron Uptake Mechanisms in Rice Seedlingsen_US
dc.typeThesis and Dissertationen_US
item.openairetypeThesis and Dissertation-
item.fulltextwith fulltext-
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