臺灣博碩士論文加值系統

臺灣能源短缺，常需依靠國外進口，而且四面環海，所能使用的陸地資源有限，若能充分利用海洋資源，如海洋生物、礦產等，可舒緩自然資源的匱乏。近年來發現新型天然資源甲烷(天然氣)水合物，廣泛地分布於極區永凍土層、海底底泥中及陸緣海域等處。目前全球甲烷水合物的蘊藏量估計為所有傳統已知化石燃料等能源之有機碳儲量之兩倍。經濟部中央地質調查所與臺灣地質物理與化學家自93年開始以多頻道反射震測區域性調查台灣西南海域，結果顯示有50 %以上之震測線有明顯之海底仿擬反射(BSR)信號，顯示此區可能有天然氣水合物賦存。台大地質系楊燦堯教授實驗室以碳同位素研究顯示這些甲烷大多是生物性的，由絕對厭氧的甲烷太古生物產生與國際類似研究結果相似。賴美津教授實驗室自2008年起參與經濟部中央地質調查所與國科會國家能源主軸計畫，探討台灣西南海域甲烷水合物潛力區微生物相。本研究分析於2009年及2010年以海洋研究船(Ocean Reasearch I and III, ORI and ORIII)於KP-9鑽井區96泥火山群區及枋寮峽谷的岩心底泥樣品。經過增殖、序列稀釋及滾管(Roll-tube)方式培養後，部份純化與Methanofollis、Methanosarcina、Methanococcoides及Methanogenium等屬親緣型相關甲烷太古生物。其中一株純化自96泥火山群底泥另一個已純化的Methanoculleus sp. S3Fa菌株，經由親緣關係比對，發現與純化自黑海表層底泥的Methanoculleus marisnigri及自日本南開海槽 (Nankai Trough)甲烷水合物賦存區底泥的Methanoculleus submarinus皆有99 %相似度。將純化及部份純化菌株與親緣關係相近的可培育及未培育的環境轉殖株的16S rRNA基因序列分析，由演化樹顯示與海底底泥、石油蘊藏區、甲烷水合物賦存區及海底泥火山等棲地的甲烷太古生物有關，因此，菌株S3Fa可能是海底泥火山甲烷水合物區域的重要甲烷太古生物。

To solve the energy shortage problem, the alternate energy methane (gas) hydrates which are estimated at about twice the amount of all other known fossil fuel reserves, are distributed along coastal margins, like SW offshore of Taiwan, as well as in terrestrial regions and marine sediments. With 74% of methane on earth are biogenic origin, this represents a significant source of natural gas from biological methanogenesis. Isotopic data from professor Yang Tsan-Yao at National Taiwan University suggested that most of the methane produced in Taiwan methane hydrate potential area is biogenic origin. To explore the methanogen diversity at methane seep or hydrate habitats, sediment samples obtained from piston and gravity cores at 96 Mud Volcano groups, Fangliao Canyon and KP-9 region by ORI and ORIII cruises were anaerobic enriched for methanogens. After serial sub-transfer and anaerobic roll-tube incubation, genus as Methanoculleus, Methanofollis, Methanosarcina, Methanococcoides and Methanogenium were partially purified. Among them, the irregular coccoid cells of Methanoculleus sp. S3Fa from the core sediment of 96 Mud Volcano Groups were purified. The 16S rDNA sequence of the strain S3Fa showed 99 % similarity with Methanoculleus marisnigri and M. submarinus (isolated from methane hydrate at Nankai Trough, Japan). Comparison of the 16S rRNA sequences of sediment samples with related uncultured environment samples showed that methanogenic phylotypes are close related with clones from marine sediment of submarine mud volcanos at East Mediterranean Sea, methane hydrate-bearing sites and oil reservoir sites. The results suggests that those phylotype of methanogens maybe important in methane hydrate formation.

中文摘要i
英文摘要ii
目次iii
表目次 ix
圖目次 x
壹、前言 1
貳、前人研究 2
一、甲烷太古生物 2
二、甲烷太古生物分類 2
(一) Order Methanobacteriales 3
1. Family Methanobacteriaceae 3
(1) Genus Methanobacterium 3
(2) Genus Methanobrevibacter 4
(3) Genus Methanosphaera 4
(4) Genus Methanothermobacter 5
2. Family Methanothermaceae 5
(二) Order Methanococcales 6
1. Family Methanococcaceae 6
(1) Genus Methanococcus 6
(2) Genus Methanothermococcus 6
2. Family Methanocaldococcaceae 7
(1) Genus Methanocaldococcus 7
(2) Genus Methanotorris 7
(三) Order Methanocellales 8
(四) Order Methanomicrobiales 8
1. Family Methanocorpusculaceae 8
2. Family Methanomicrobiaceae 9
(1) Genus Methanoculleus 9
(2) Genus Methanofollis 11
(3) Genus Methanogenium 11
(4) Genus Methanolacinia 12
(5) Genus Methanomicrobium 12
(6) Genus Methanoplanus 12
3. Family Methanoregulaceae 13
(1) Genus Methanolinea 13
(2) Genus Methanoregula 13
(3) Genus Methanosphaerula 14
4. Family Methanospirillaceae 14
5. Genera incertae sedis 14
(1) Genus Methanocalculus 14
(五) Order Methanosarcinales 15
1. Family Methanosarcinaceae 15
(1) Genus Methanimicrococcus 15
(2) Genus Methanococcoides 16
(3) Genus Methanohalobium 16
(4) Genus Methanohalophilus 17
(5) Genus Methanolobus 17
(6) Genus Methanomethylovorans 18
(7) Genus Methanosalsum 18
(8) Genus Methanosarcina 19
2. Family Methanosaetaceae 19
3. Family Methermicoccaceae 20
(1) Genus Methermicoccus 20
(六) Order Methanopyrales 21
1. Family Methanopyraceae 21
(七) Order“Methanoplasmatales” 22
三、深海底泥、泥火山、冷泉與甲烷水合物賦區的甲烷生成與氧化太古生物 22
(一) 深海底泥環境及甲烷生成與氧化太古生物 22
(二) 海底泥火山環境及甲烷生成與氧化太古生物 23
(三) 深海冷泉環境及甲烷生成與氧化太古生物 25
(四) 甲烷水合物賦區及甲烷生成與氧化太古生物 26
四、台灣西南海域甲烷水合物賦存區海底底泥微生物相關研究 28
五、研究目的策略與重要結果 29
叁、材料與方法 31
一、採樣時間、地點與樣品處理 31
二、藥劑與培養基 31
(一) 除氧操作系統 (Hungate station) 31
(二) 厭氧培養液及固態培養基的配製：MB/W 及MM/W培養基 32
(三) 還原劑與碳源及抗生素溶液之製備 33
(四) TGC (thioglycollate) 培養基 34
三、厭氧轉殖接種與甲烷菌增殖(enrichment)培養 34
(一) 厭氧轉殖接種 34
(二) 以氣相層析儀偵測樣本的甲烷氣體 35
(三) Roll tube方法 35
(四) 序列稀釋方法 36
(五) 抗生素添加 36
四、Roll tube 菌株篩選 36
(一) 厭氧操作箱(Coy chamber) 36
(二) 挑取roll-tube內的菌落 37
五、微生物形態觀察 37
(一) 以位相差顯微鏡觀察細胞形態及記錄 37
(二) 以穿透式電子顯微鏡負染方式觀察細胞形態及記錄 38
六、生長與生理生化特性分析 38
(一) 甲烷化基質的配製 38
(二) 含不同NaCl濃度厭氧培養液MB/W的配製 39
(三) 不同酸鹼值的厭氧培養液MB/W的配製 39
(四) 最適生長溫度範圍測試 39
(五) 抗生素抗性測試 39
(六) SDS 感受性測試 40
(七) 生長曲線的測定 40
七、甲烷太古生物染色體DNA萃取 41
八、核酸純度與定量分析 41
九、核酸膠體電泳分析 42
十、聚合酶連鎖反應 (Polymerase chain reaction, PCR)增幅16S rRNA基因 42
十一、PCR增幅產品回收純化 43
十二、DNA 黏合反應 44
十三、勝任細胞的製備與質體轉形作用 44
(一) 勝任細胞的製備 44
(二) 質體轉形作用 44
十四、質體的抽取與純化 45
十五、DNA定序樣品之製備 46
十六、核酸定序與序列分析47
十七、表面蛋白(Surface layer)之製備 48
十八、全細胞蛋白質圖譜之製備 48
十九、蛋白質電泳分析 (SDS- PAGE) 49
肆、結果與討論 51
一、深海底泥甲烷菌之增殖培養與純化 51
二、自海洋96泥火山群純化的甲烷太古生物Methanoculleus sp. S3Fa 形態生理生化特性分析與與系統演化分類 52
1. 細胞形態 52
2. 16S rRNA基因序列分析與系統演化分析分析 53
3. 生理生化與生長特性分析 54
4. 抗生素抗性 55
5. 表面蛋白(Surface layer)與全細胞蛋白質分析 56
6. 全細胞蛋白質分析 56
三、部分純化的甲烷菌增殖樣品 56
1. ORI-934 樣本 56
(1) 96 泥火山群 (96 Mud volcano groups ) 56
(2) 枋寮峽谷 (Fangliao canyon) 57
2. ORIII-1368 樣本 (KP-9甲烷水合物賦存區) 58
3.台灣西南外海甲烷水合物賦存區甲烷菌的多樣性 59
伍、結論與展望62
陸、表與圖 64
柒、參考文獻 79

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