臺灣博碩士論文加值系統

工業革命後，重金屬被大量利用及排放，加劇了存在已久的重金屬污染 問題。在科學家持續的研究下，對於重金屬對人類及動植物的危害，甚至物 種多樣性的改變，均有較透徹的了解;除此之外，包括生物復育等環境整治 策略也應運而生。然而，隨著光電與半導體產業等科技興起，使用量大增的 技術關鍵元素 (Technology-critical elements, TCE) 因生產過程或衍生的廢 棄物可能造成的污染疑慮，也開始受到人們的注意。其中，半導體產業的重 要原料，如鎵 (Gallium, Ga) 及銦 (Indium, In) 等，對人體及動植物皆具毒 性，但對環境中大量存在的微生物影響為何所知有限。之前的研究透過將平 鎮土系 (Pinchen series; Pc series) 之土樣以 100 mg/L 的銦進行增殖培養後， 分離出一細菌菌株 PcI1001，且發現 PcI1001 同時具有抗鎵、銦以及同屬於 3A 族金屬元素鉈 (Thallium, Tl) 的能力。以 16S rRNA 基因序列初步進行 物種鑑定及經由 PacBio SMRT 技術進行全基因體定序分析，確認其屬於 Luteibacter spp.，基因組大小為 4,550,000 bp，且與 L. rhizovicinus LJ96 的 相似度達 94%。分析添加不同濃度銦土樣中的微生物相發現，PcI1001 所屬 的 Xanthomonadales 目及 Rhodanobacteraceae 科之數量均會隨著添加銦濃 度增加而增加。PcI1001 對鎵、銦及鉈的最小抑制濃度 (Minimal inhibition concentration,MIC) 分別約為 150mg/L、25mg/L 和 1200mg/L。透過穿透 式電子顯微鏡觀察，發現 PcI1001 在鎵、銦和鉈的處理下，其細胞膜變得不 穩定，菌體在分裂時期的狀態也與未處理組不同。利用 ICP-MS 檢測發現， 以 200 mg/L 鎵和銦處理後，PcI1001 的菌體能吸附兩者的量分別為 17.3±1.8mg/kg 和 36.3±10.8mg/kg。接著，以鎵、銦和鉈處理 PcI1001 跳 躍子插入突變庫 (transposon-insertion mutant library)，篩選出 6 個對這些 TCE 敏感度改變的突變株，經過定序後發現跳躍子插入的位置分別是在 Spermine/spermidine acetyltransferase (bltD)、DUF4398-domain containing protein、Polyhydroxyalkanoate depolymerase、Phosphoglycerol transferase (MdoB)、Cocaine esterase (cocE_4) 等基因中及 copper resistance/multidrug resistance 基因組的啟動子序列上。透過基因刪除突變株的建構，觀察到 Spermine/spermidine acetyltransferase (bltD) 及 DUF4398-domain containing protein 的突變株對鎵更具耐受性，而 Phosphoglycerol transferase (MdoB) 突 變株則對鉈耐受性有降低的趨勢。基因表現上， bltD 在鎵、銦及鉈的處理 下皆有較高表現量，DUF4398-domain containing protein 和 polyhydroxy- alkanoate depolymerase 的基因在鉈處理下表現量較高，而 mdoB 在銦處理 下有較高的表現量。然而，以上這些基因缺失後的突變株對鎵、銦和鉈的耐 受性改變皆與野生菌株無統計上差異。雖然突變株篩選及分析結果並未得到 PcI1001 耐受鎵、銦或鉈的可能機制，然而本研究所觀察到 PcI1001 受鎵、 銦和鉈處理後的型態變化以及所建立的實驗方法皆有助於日後研究 PcI1001 對鎵、銦或鉈耐受性反應之相關基因上。

Since the first industrial revolution, utilization and discharge of heavy metals have increased dramatically, exacerbating the long-standing problem of heavy metal pollution. Continuous works by the scientists expand our knowledge more profoundly on the detrimental effects of heavy metals to humans, animals and plants, and even alterations of biodivisity. In addition, environmental remediation strategies, including bioremediation, have also been developed. However, with the development of optoelectronic and semiconductor industries, people start to concern about that the technology-critical elements (TCE), which have increased their utilization lately, may cause pollution due to the manufacturing processes or waste derived. Among those TCE, major elements for the semiconductor industry, such as gallium (Ga) and indium (In), are toxic to humans, animals and plants, but there is limited knowledge about their effects on microbiome in the environment. In previous studies, a bacterial strain PcI1001 was isolated from enrichment culture using soil samples of the Pinchen (Pc) series supplemented with 100 mg/L indium. PcI1001 is tolerant to gallium and indium as well as thallium, an element also belonging to the Group 3A in the periodic table. Both 16S rRNA gene sequencing and whole genome sequencing by PacBio SMRT technology indicate PcI1001 belongs to genus Luteibacter. The genome size of PcI1001 is 4,550,000 bp, which shares 94% identity to that of L. rhizovicinus LJ96. Metagenomic analysis using the data obtained from soil samples amended with different concentrations of indium revealed that abundance of Order Xanthomonadales and Family Rhodanobacteracea, which PcI1000 belongs to, increased along with the increased concentrations of indium. The minimum inhibition concentration (MIC) of PcI1001 for Ga, In, and Tl is about 150 mg/L, 25 mg/L, and 1200 mg/L, respectively. Transmission electron microscope observation revealed that cell membrane became unstable and fragile after treatment of Ga, In, and Tl, and the morphology of cells under division was also different from that of the untreated group. After detection by inductively coupled plasma mass spectrometry (ICP- MS), it was found that the adsorption capacity of PcI1001 on Ga and In was 17.3±1.8 mg/kg and 36.3±10.8 mg/kg, respectively, after incubation in 200 mg/L gallium and indium. Using PcI1001 mutant library generated by EZ-Tn5-based transposon insertion, six mutants whose susceptibility to Ga, In or Tl was altered were identified. In these six mutants, EZ-Tn5 inserts at coding sequences for spermine/spermidine acetyltransferase (BltD), DUF4398-domain containing protein, polyhydroxyalkanoate depolymerase, phosphoglycerol transferase (MdoB), Cocaine esterase (CocE_4) and the promoter region of copper resistance/multidrug resistance operon, respectively. Mutants with in-frame deletion of genes coding for BltD and DUF4398-domain containing protein are seemingly more tolerant to gallium. A mutant deleted of mdoB is slightly more sensitive to thallium. Transcriptional analysis showed that the expression of bltD was higher after treatment of gallium, indium and thallium. The expressions of the genes coding for DUF4398-domain containing protein and polyhydroxyalkanoate depolymerase were higher in response to thallium treatment whereas that of mdoB was higher after indium treatment. However, compared to wild type, no statistical difference on tolerance to gallium, indium and thallium can be observed in these in-frame deletion mutants. Nevertheless, the morphological changes observed on gallium, indium or thallium-treated cells and the experimental procedures developed in this study can help us better understand the currently unknown areas on genes involved in bacterial tolerance to gallium, indium, and thallium.

致謝 i
中文摘要 iii
Abstract v
目錄 viii
表目錄 xii
圖目錄 xiii
附表目錄 xv
附圖目錄 xvi
壹、文獻回顧 1
一、重金屬污染研究之重要性 1
1. 重金屬與科技關鍵元素 1
2. 重金屬之污染之問題 2
二、鎵、銦和鉈的簡介及應用 6
1. 鎵的基本性質介紹與應用 6
2. 銦的基本性質介紹與應用 7
3. 鉈的基本性質介紹與應用 8
三、鎵、銦及鉈對生物的毒害情形 9
1. 鎵對動物之影響： 10
2. 鎵對植物之影響： 11
3. 鎵對微生物之影響： 12
4. 銦對動物之影響 14
5. 銦對植物之影響： 15
6. 銦對微生物之影響： 16
7. 鉈對動物之影響： 16
8. 鉈對植物之影響： 17
9. 鉈對微生物之影響： 17
五、微生物之重金屬抗性的相關機制與應用 19
1. 細菌細胞壁的屏障 20
2. 對金屬離子的主動運輸 22
3. 細胞外的金屬螯合 25
4. 細胞內的金屬螯合 26
5. 金屬離子的還原 27
貳、研究動機與目的 28
參、材料及方法 30
一、細菌菌株與培養條件 30
二、細菌菌株鑑定和全基因體序列分析 30
三、平鎮土系土壤樣品總體基因體數據再分析 31
四、檢測 PcI1001 對重金屬的抗性 31
五、穿透式電子顯微鏡 (Transmission electron microscopy, TEM) 樣品之製備32
六、掃描式電子顯微鏡 (Scanning electron microscopy, SEM) 樣品之製備 33
七、PcI1001 重金屬吸附量分析 34
八、以 PcI1001 突變體庫篩選對鎵、銦和鉈抗性改變之菌株 34
九、鑑定候選突變菌株基因序列中跳躍子插入之位置 35
十、抽取 PcI1001 全基因體 DNA 36
十一、電穿孔法勝任細胞的製備 37
十二、電穿孔轉型 37
十三、小量質體 DNA之萃取 38
十四、膠體內DNA純化 (gel elution) 38
十五、以 GENEzol 抽取細菌之 RNA 39
十六、重金屬壓力下基因表現量測定 40
十七、半定量反轉錄 PCR 40
十八、PcI1001 目標基因剔除突變株之建構 41
十九、目標基因回補株之製備 42
肆、結果 43
一、PcI1001 物種鑑定及全基因體序列分析 43
二、土壤環境微生物總體基因之分析 44
三、測定 PcI1001 對鎵、銦和鉈的最小抑制濃度 (minimum inhibitory concentration, MIC) 46
四、以穿透視顯微鏡觀察 PcI1001 型態及鎵、銦和鉈對細胞之影響 47
五、PcI1001 吸附鎵與銦之能力 48
六、突變株的篩選以及跳躍子插入序列之鑑定 48
七、將目標基因剔除後及將突變菌株基因回補後之抗性狀態 50
八、檢測在重金屬壓力下篩選之目標基因的表現量 50
伍、討論 53
陸、參考文獻 69
柒、圖表 87
捌、附圖表 116

法務部，光電材料及元件製造業放流水標準，民國 101 年 10 月 12 日
法務部，土壤及地下水污染整治法，民國 89 年 2 月 2 日
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