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研究生:馮一郎
研究生(外文):Yi-Lang Feng
論文名稱:Pseudomonas vesicularis MA103 carIII 基因轉殖至 Lactococcus lactis NZ3900 中表現及其所產 k-紅藻膠酶水解 k-紅藻膠所得寡醣之生理活性探討
論文名稱(外文):Studies on Cloning and Expression of carIII Gene from Pseudomonas vesicularis MA103 to Lactococcus lactis NZ3900 and Bioactivities of k-Carrageenan Hydrolyzed by Cloned k-Carrageenase
指導教授:潘崇良
指導教授(外文):Chorng-Liang Pan
學位類別:碩士
校院名稱:國立臺灣海洋大學
系所名稱:食品科學系
學門:農業科學學門
學類:食品科學類
論文種類:學術論文
論文出版年:2012
畢業學年度:100
語文別:中文
論文頁數:172
中文關鍵詞:轉殖Pseudomonas vesicularis MA103Lactococcus lactis NZ3900乳酸鏈球菌素調控表現系統k-紅藻膠k-紅藻膠酶
外文關鍵詞:TransformationPseudomonas vesicularis MA103Lactococcus lactis NZ3900Nisin Controlled Expression systemk-Carrageenank-Carrageenase
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本研究之目的為將 Pseudomonas vesicularis MA103 所產紅藻膠酶 carIII 基因以食品級載體 pNZ8149 轉形至 Lactococcus lactis NZ3900 中表現,並探討轉殖株所產 k-Carrageenase (Pv-Car-C) 水解紅藻膠所得寡醣產物 (Car-F5) 之組成成分以及抗氧化與抗凝血生理活性。首先於引子對兩端分別加入 NcoI 與 PstI 限制酶切位,再以菌株 MA103 染色體 DNA 為模板進行聚合酶反應 (Polymerase chain reaction, PCR) 擴增 carIII 基因,將所得 carIII 基因以 TA-cloning 方式與 pGEM-TTM 載體製備成重組載體 I 並轉形至菌株 DH5a。再以三種限制酶將重組載體 I 進行剪切,再與 pNZ8149 載體進行連結反應 (Ligation) 形成重組載體 II 並轉形至菌株 NZ3900 中,篩選出 4 株 lac+ 轉殖株,以核苷酸序列定序經比對確認後,將轉殖株命名為 Lc. lactis NZ3900-rcarIII。將轉殖株 NZ3900-rcarIII 以 10 ng/mL Nisin 誘導 6 hr 後,而胞內 Pv-Car-C 可測得比活性 0.042 U/mg,且經 SDS-PAGE 試驗結果顯示,此 Pv-Car-C 之分子量約為 106 kDa,與自 carIII 基因序列所推估之分子質量 106 kDa 相近。胞外 Pv-Car-C 則以分子篩大小 10 kDa、30 kDa 與 100 kDa 進行區分並收集大於 100 kDa 之酵素液,可測得比活性 0.034 U/mg。在基質特異性測試中 Pv-Car-C 對於 k-Carrageenan 較具有專一性。而將 Pv-Car-C 水解 0.15% k-Carrageenan 96 hr 後之寡醣水解液進行 TLC 分析,可推測主要水解寡醣產物為聚合度 (Degree of polymerization, DP) DP5 之寡醣。而後將寡醣水解液利用超過濾系統區分出小於 3 kDa 分子量之寡醣,再經 Sephadex G-10 膠體過濾層析區分,並經 HPLC 分析比對標準品後,推測為聚合度 DP5 之寡醣。FT-IR 分析下可以發現於 1,066 cm-1 有明顯之吸收波峰,推測為 SO42- 基團,硫酸基含量為 17.70%,經計算後寡醣總分子量為 890 Da,將此寡醣命名為 Car-F5。測試 Car-F5 之抗氧化活性變化,發現因不具有可溶性多酚類含量,故對於 DPPH 自由基清除能力及螯合亞鐵離子能力無明顯活性,但在還原力方面則可測得 13.95 TE g/mL。在兔血漿抗凝血之凝血酶原時間 (Prothrombin time, PT) 實驗中,20 mg/mL Car-F5 可明顯增加兔血漿 PT 之延遲時間 132 sec,且延遲時間較低劑量 10 mg/mL 之 Heparin 長。而活化部分凝血活酶時間 (Activated partial thromboplastin time, APTT) 實驗中,只需以 5 mg/mL Car-F5 就可明顯增加兔血漿 APTT 之延遲時間 319 sec,且其凝集程度效果也較低劑量 10 mg/mL 之 Heparin 佳。而本研究主要建立食品級轉殖株 NZ3900-rcarIII 所產 Pv-Car-C 並用於製備 Car-F5 之生產技術,且在抗凝血實驗中顯示較佳的活性,未來可能有做為預防血栓性心血管疾病保健功能食品之發展潛力。
The purpose of this thesis is using carrageenase carIII gene from Pseudomonas vesicularis MA103 as a reporter gene combine with plasmid pNZ8149 from nisin-controlled expression system and transgenic to Lactococcus lactis NZ3900 host cells, then evaluate the characterization of nisin-induced k-carrageenase crude enzyme (PV-Car-C), the composition of oligosaccharide (Car-F5) from Pv-Car-C hydrolyzed k-carrageenan of the antioxidant bioactivity and anticoagnlant bioactivity. The carIII forward primer encode NcoI restriction site and the reverse primer encode PstI restriction site. Then using the gene of carrageenase carIII as a template for PCR reaction, and this PCR amplified product as ligated into pGEM-TTM vector, and then transformed into the competent cells of E. coli DH5α. Then pGEM-TTM vector, which carrying the cloned carIII gene, was digested with NcoI and PstI, and ligated into the LAB vector pNZ8149. The recombined vector II (pNZ8149-palIII) was electroporated into the Lc. lactis NZ3900. And the lac+ yellow colony was picked and the punified and examined it’s plasmid profile 4 successful transformed carIII cloned were identified by designated. It was proved that the sequence of the cloned DNA fragments was completely identical to the DNA sequence of the carIII gene, and the succeed clone was designated as cloned strain NZ3900-rcarIII. The clones induced by 10 ng/mL nisin for 6 hr could not obtain extracellular carrageenase activity. The intracellular crude enzyme solution could obtain higher carrageenase activity (0.042 U/mg), which revealed a 106 kDa protein band by SDS-PAGE. The extracellular crude enzyme solution with molecular mass > 10 kDa, > 30 kDa, and > 100 kDa were collected by UF system. It could obtain higher carrageenase activity (0.034 U/mg). On substrate specificity, Pv-Car-C can hydrolyze k-Carrageenan. Analyzing the hydrolysate of Pv-Car-C by TLC, compared to the standards, discovered that the main hydrolysate were assumed to be DP5 oligosaccharide. Oligosaccharides hydrolysates with molecular mass < 3 kDa were collected by a UF system, then though GPC and HPLC to obtain DP5 oligosaccharide (Car-F5). FTIR chromatograms showed that was obvious signal enhancement in wave numbers 1066 cm-1 resulted from sulfate contents, and the sulfate contents of Car-F5 was 17.70%. In the evaluation of the antioxidation effect of Car-F5, soluble total phenol, DPPH free radical scavenging, Fe2+ chelating effect and of Car-F5 did not increase, but the reducing power of Car-F5 had significant increase (13.95 TE mg/mL). In the experiment of anticoagulant activities of rabbit plasma, prothrombin time analysis of Car-F5 (20 mg/mL) was delayed to 132 sec, and activated partial thromboplastin time analysis of Car-F5 (5 mg/mL) was delayed to 319 sec, and it was tendency for reduction on absorbance. This study provides optimal parameters for the NICE system in cloned strain NZ3900-rcarIII as well as a means to produce functional Car-F5, and serving as the basis for development of health food for preventing the high risk of thrombotic cardiovascular disease in the future.
目錄 i
表目錄 vii
圖目錄 viii
附錄目錄 x
中文摘要 xi
英文摘要 (Abstract) xiii
壹、前言 1
貳、文獻整理 3
一、紅藻膠 (Carrageenan) 3
1-1. 來源 3
1-2. 成分及化學結構 3
1-3. 黏度特性及安定性 4
1-4. 成膠機制 4
1-5. 與蛋白質的交互作用 5
1-6. 紅藻膠之應用 6
二、紅藻膠酶 (Carrageenase) 6
三、紅藻膠酶之生化特性與純化 7
3-1. 溫度與 pH 值對 Carrageenase 之影響 7
3-2. 金屬離子對 Carrageenase 之影響 8
3-3. Carrageenase 之分子量變化 9
3-4. 紅藻膠酶之純化 10
四、乳酸菌 (Lactic acid bacteria) 10
4-1. 乳酸菌演化的起源 10
4-2. 乳酸菌之定義 11
4-3. 乳酸菌之特性與對碳水化合物之代謝途徑 12
4-4. 乳酸菌做為益生菌 (Probiotic) 之應用 12
4-5. 乳酸菌於遺傳工程上之應用價值 16
4-6. 乳酸菌基因轉殖之相關方法 17
4-7. 乳酸鏈球菌素調控表現系統 20
4-8. 乳酸菌宿主細胞、乳酸菌質體、及乳酸鏈球菌素 22
4-9. 乳酸菌轉殖株胞內外蛋白質之表現與胞內物萃取方法 23
五、紅藻膠酶之基因選殖 26
5-1. 紅藻膠酶基因以 E. coli 做為宿主之轉殖系統 26
5-2. 紅藻膠酶基因以 B. subtilis 做為宿主之轉殖系統 26
六、紅藻膠寡醣之製備生產 26
七、紅藻寡醣組成成分之分析與鑑定 27
7-1. 薄層層析法 (Thin-layer chromatography, TLC) 27
7-2. 高效能液相層析法 (High performance liquid
chromatography, HPLC) 28
7-3. 傅立葉轉換紅外線光譜法 [Fourier-transform infrared
(FTIR) spectroscopy] 29
7-4. 核磁共振光譜法 (Neuclear magnetic resonance, NMR) 30
八、紅藻膠酶水解產物之生理活性 31
8-1. 抗氧化 31
8-2. 抗凝血 32
8-3. 抗病毒 32
參、實驗設計 34
肆、實驗材料與方法 35
一、實驗材料 35
1-1. 實驗菌株 35
1-2. 試驗藥品 35
1-2-A. 藥品 35
1-2-B. 載體 38
1-2-C. 引子 38
1-2-D. 聚合酶鏈鎖反應試劑 38
1-2-E. 限制酶及蛋白酶抑制劑 39
1-2-F. 蛋白質及 DNA 萃取純化套裝試劑 39
1-2-G. 電泳標準品 39
1-2-H. 培養基組成 40
1-2-I. 紅藻膠酶反應基質 42
1-2-J. DNS 溶液 42
1-2-K. 電泳膠片配製 42
1-2-L. 電泳溶液 43
1-2-M. 除菌過濾 44
1-2-N. 薄層層析矽膠片 (TLC silica gel) 44
1-2-O. 離心式超過濾濾膜 44
1-3. 儀器設備 44
二、實驗方法 46
2-1. 菌種保存與活化 46
2-1-A. 菌株保存 46
2-1-B. 菌株活化 47
2-2. P. vesicularis MA103 之紅藻膠酶基因選殖至 Lc. lactis NZ3900 47
2-2-A. 菌株 MA103 染色體 DNA 之萃取純化 47
2-2-B. 引子對之設計 48
2-2-C. 聚合酶鏈鎖反應擴增 carIII 基因 48
2-2-D. 重組載體 I (pGEM-TTM vector-carIII) 之製備 49
2-2-E. E. coli DH5a 勝任細胞之製備 50
2-2-F. E. coli DH5a 之電穿孔轉形 50
2-2-G. 貝爾塔尼 (Luria-Bertani, LB-ampicillin) 鑑別培養基初次確認轉殖株 DH5a-rcarIII 51
2-2-H. PCR 擴增反應二度確認轉殖株 DH5a-rcarIII 51
2-2-I. 限制酶剪切三度確認轉殖株 DH5a-rcarIII 51
2-2-J. 轉殖株 DH5-rcarIII 核苷酸序列定序比對 52
2-2-K. 重組載體 II 之製備 (pNZ8149 vector-rcarIII) 52
2-2-L. Lc. lactis NZ3900 接受細胞之製備 53
2-2-M. Lc. lactis NZ3900 之電穿孔轉形 54
2-2-N. 溴甲酚紫 (Brom cresol purple-lactose, BCP-L)
鑑定培養基初次確認轉殖株 NZ3900-rcarIII 54
2-2-O. PCR 擴增反應二度確認轉殖株 NZ3900-rcarIII 54
2-2-P. 限制酶剪切三度確認轉殖株 NZ3900-rcarIII 55
2-2-Q. 轉殖株 NZ3900-rcarIII 核苷酸序列定序比對 55
2-3. 轉殖株 NZ3900-rcarIII 之特性分析 56
2-3-A. 革蘭氏染色法檢測 56
2-3-B. 石蕊牛乳檢測 57
2-3-C. 生長曲線 57
2-3-D. 質體穩定性測試 58
2-3-E. 不同濃度 Nisin 對轉殖株 NZ3900-rcarIII 生長之影響 58
2-3-F. 培養液 pH 值之調控 58
2-4. 轉殖株 NZ3900-rcarIII 所產 Pv-Car-C 之表現 59
2-4-A. 超音波細胞破碎法萃取轉殖胞內 Pv-Car-C 59
2-4-B. 液態氮細胞破碎法萃取轉殖胞內 Pv-Car-C 59
2-4-C. 轉殖 Pv-Car-C 之活性測定 60
2-4-D. 轉殖 Pv-Car-C 之蛋白質定量 61
2-4-E. 轉殖胞內外 Pv-Car-C 之區分純化 61
2-4-F. 轉殖 Pv-Car-C 之分子量確認 62
2-4-G. 基質特異性測試 63
2-5. 轉殖 Pv-Car-C水解紅藻膠所得寡醣產物之組成分析 63
2-5-A. 全醣量與還原醣量分析 63
2-5-B. 紅藻寡醣之分離製備 64
2-5-C. 薄層色層分析 (TLC) 64
2-5-D. 高效能液相層析分析 (HPLC) 65
2-5-E. 傅立葉轉換紅外線光譜儀分析 (FTIR) 65
2-5-F. 硫酸酯含量分析 66
2-6-G. 可溶性多酚類含量分析 66
2-6. 紅藻五糖 Car-F5 之抗氧化生理活性測定 66
2-6-A. 清除 DPPH 自由基能力測定 67
2-6-B. 亞鐵離子螯合能力測定 68
2-6-C. 還原力之測定 68
2-7. 紅藻五糖 Car-F5 之抗凝血生理活性測定 69
2-6-A. 凝血酶原時間 69
2-6-B. 活化部份凝血活酶時間 69
三、統計分析 69
伍、結果與討論 71
一、紅藻膠酶 carIII 基因轉殖至菌株 NZ3900 中 71
1-1. 菌株 MA103 紅藻酶基因篩選 71
1-2. carIII 基因以菌株 DH5 做為宿主之轉殖系統 72
1-3. carIII 基因以菌株 NZ3900 做為宿主之轉殖系統 73
二、轉殖株 NZ3900-rcarIII 74
2-1. 轉殖株 NZ3900-rcarIII 之特性分析 74
2-2. 轉殖 Pv-Car-C 之表現 78
2-3. 轉殖 Pv-Car-C 胞內外粗酵素液之區分純化 79
三、轉殖 Pv-Car-C 水解紅藻膠所得寡醣之組成分析 80
3-1. 基質特異性測試 80
3-2. 全醣量與還原醣量分析 81
3-3. 薄層色層分析 (TLC) 81
3-4. 高效能液相層析分析 (HPLC) 82
3-5. 膠體過濾層析分析 (GPC) 83
3-6. 傅立葉轉換紅外線光譜分析 (FTIR) 84
3-7. 硫酸酯含量分析 84
3-8. 可溶性多酚類含量分析 85
四、紅藻五糖 Car-F5 之生理活性分析 86
4-1. 清除 DPPH 自由基能力之分析 86
4-2. 螯合亞鐵離子能力之分析 86
4-3. 還原力之分析 87
4-4. 凝血酶原時間之分析 87
4-5. 活化部份凝血活酶時間之分析 88
陸、結論 89
柒、參考文獻 91
捌、附錄 158
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