石斑魚為我國高經濟價值魚種，魚苗產值高達三億五千萬元；然而，神經壞死病毒感染卻造成其仔稚魚死亡率最高達100％，對養殖漁業造成嚴重的經濟損失。疫苗為病毒性疾病最佳防治方式，利用本研究室發表之龍膽石斑神經壞死病毒似病毒顆粒，施打龍膽石斑、瑪拉巴石斑與棕點石斑，並以八種不同免疫學原理方法分析石斑魚抗血清，其中抗原捕捉式酵素連結免疫吸附法(Antigen-capture ELISA)不僅可定性分析，更可量化抗體力價。當免疫一週後，三種魚均可偵測血清特異性抗體存在，且抗體力價隨時間增加而提高。龍膽石斑施打10 μg後，一個月抗體力價可達0.95，予追加注射一次可提升力價35.7％。瑪拉巴石斑施打50 μg兩劑六週後，力價強度較單次施打者，可提升33.3％。而棕點石斑施打450 μg一劑者，體內血清抗體強度在五週後可高達1.57。魚體之血清抗體力價可維持平穩達四個月，於七個月後魚體內仍可偵測抗體存在。當病毒力價為103.5 TCID50/mL時，in vitro中和實驗結果顯示，可保護SSN-1細胞不發生細胞病變，三魚種免疫前之抗體中和力價均小於1：50，而龍膽石斑免疫四週後的抗體中和力價可由免疫一週時的1：200，提升至1：1600，而十一週仍可高達1：6400；瑪拉巴石斑免疫後六週力價可提升至1：3200；而棕點石斑力價則可提升至1：400。佛氏完全佐劑的添加對抗體強度並無提升效果。綜合以上結果顯示，龍膽石斑神經壞死病毒之似病毒顆粒，可誘發魚體產生高力價之特異性抗體，並具良好的保護效力。

The groupers are vital fish in Taiwan, the market of grouper fry over 300 million dollars. While grouper nervous necrosis virus (NNV) has caused mass mortality, especially 100% in larvae and juveniles, which economically impacts on culture of marine fish. The vaccination is one of the best methods to against viral diseases. The dragon grouper (Epinephelus lanceolatus), malabar grouper (E. malabaricus) and brown-marbled grouper (E. fuscoguttatus) were injected with different dosages and injection frequencies of virus-like particles (VLPs) of DGNNV, which is by the first claimed. The anti-sera of vaccinated fish were analyzed with eight kinds of immunology methods, among which antigen-capture ELISA was the best choice for qualitative and quantitative assays. The signal of antibodies in the vaccinated fish was detected in all groupers in one week after primary immunization, and the antibody titers increased markedly in one month. In dragon grouper, fish was injected with 10 μg of VLPs, the antibody titer reached 1.05. To given booster injection once, antibody titers were raised to 35.7%. In malabar grouper, after injected twice with 50 μg of VLPs, the antibody titer raised 33.3% than inoculation once in six weeks. After brown-marbled grouper was injected with 450 μg of VLPs, the high antibody titer reached to 1.57 at five weeks post primary immunization. Specific antibodies still can be detected after seven months. In the in vitro assay with MGNNV of 103.5 TCID50/mL, neutralizing antibody titer of control fish were all lower than 1:50. The neutralizing antibody titer of anti-serum of dragon grouper was detected at 1:200 at one week, and raised to 1:1600 at four weeks and 1:6400 at eleven weeks after primary vaccination. In malabar grouper and brown-marbled grouper, the neutralizing antibody titers were 1:3200 and 1:400, respectively, in one month. The antibody titer can not increased by Freund’s complete adjuvant. The fish produced high antibody titer and high protection by immunization with VLPs.

致謝 --------------------------------------------------------------------------- Ⅰ
中文摘要 --------------------------------------------------------------------- Ⅱ
Abstract ------------------------------------------------------------------------ Ⅲ
目錄 --------------------------------------------------------------------------- Ⅳ
表目錄 ------------------------------------------------------------------------ Ⅶ
圖目錄 ------------------------------------------------------------------------ Ⅷ
壹、前言
一、台灣石斑養殖現況 ------------------------------------------------------------- 1
二、魚類神經壞死病毒(Piscine nervous necrosis virus) ----------------------- 2
三、病毒性神經壞死症 (viral nervous necrosis)之臨床表徵 ---------------- 5
四、溫度對神經壞死病毒感染能力之影響 ------------------------------------- 5
五、Nodavirus之基因體 ------------------------------------------------------------ 6
六、Betanodavirus之分子生物學 ------------------------------------------------- 7
七、Betanodavirus之檢測法 ------------------------------------------------------- 12
八、增殖Betanodavirus之細胞株 ------------------------------------------------- 15
九、魚類之免疫系統 ---------------------------------------------------------------- 18
(1)非特異性免疫系統 ----------------------------------------------------------- 19
(2)特異性免疫系統 -------------------------------------------------------------- 21
十、魚類疫苗給予途徑與種類----------------------------------------------------- 22
十一、似病毒顆粒(virus-like particles, VLPs)------------------------------------ 29
十二、研究目的 ---------------------------------------------------------------------- 33
貳、材料與方法
一、石斑魚之來源 ------------------------------------------------------------------- 35
二、SSN-1細胞培養 --------------------------------------------------------------- 35
三、似病毒顆粒之製備與純化 --------------------------------------------------- 36
四、SDS-PAGE蛋白質電泳 ------------------------------------------------------- 37
五、蛋白質定量(Lowry method) -------------------------------------------------- 38
六、抗VLP之小鼠血清製備 ------------------------------------------------------ 38
七、石斑魚抗血清力價檢測法之建立 ------------------------------------------ 39
(1)抗原捕捉式點墨法(Antigen-capture dot blotting)偵測抗體 ----------- 39
(2)競爭型西方點墨法(Competitive Western blotting) ----------------------- 41
(3)間接阻斷型西方點墨法(Indirect blocking Western blotting) ----------- 42
(4)競爭型間接螢光抗體染色法(Competitive indirect fluorescent antibody test, Competitive IFAT) --------------------------------------------
43
(5)間接阻斷型間接螢光抗體染色法(Indirect blocking indirect
fluorescent antibody test, Indirect blocking IFAT) ------------------------
45


(6)競爭型酵素連結免疫吸附法(Competitive Eenzyme linked immunosorbent assay, Competitive ELISA) -------------------------------
48
(7)間接阻斷型酵素連結免疫吸附法(Indirect blocking Eenzyme linked immunosorbent assay, Indirect blocking ELISA) -------------------------
49
(8)抗原捕捉式酵素連結免疫吸附法(Antigen–capture enzyme linked immunosorbent assay, antigen-capture ELISA)測定抗體力價 --------
50
八、VLPs誘發石斑魚之免疫效力 ------------------------------------------------ 52
(1)龍膽石斑之免疫 --------------------------------------------------------------- 52
(2)添加佛氏完全佐劑對VLPs誘發龍膽石斑免疫效價之影響 ---------- 53
(3)瑪拉巴石斑之免疫 ------------------------------------------------------------ 53
(4)添加佛氏完全佐劑對VLPs誘發瑪拉巴石斑免疫效價之影響 ------- 54
(5)棕點石斑之免疫 --------------------------------------------------------------- 55
九、TCID50之測定 ------------------------------------------------------------------- 55
十、抗VLP之石斑血清之中和試驗(Neutralization test) ---------------------- 56
十一、中和指數(Neutralization Index) -------------------------------------------- 57
參、結果
一、建立免疫學的方法，檢測魚類產生抗體的程度 ---------------------------- 59
(1) Antigen-capture dot blotting (抗原捕捉式點墨法)偵測抗體生成 ------ 60
(2) Competitive western blotting (競爭型西方點墨法)與Indirect blocking western blotting (間接阻斷型西方點墨法) -------------------------------
60
(3) Competitive IFAT (競爭型螢光抗體染色法)與Indirect blocking IFAT (間接阻斷型螢光抗體染色法) ----------------------------------------------
61
(4) Competitive ELISA (競爭型酵素連結免疫吸附法)與Indirect blocking ELISA (間接阻斷型酵素連結免疫吸附法) --------------------
62
(5) Antigen-capture ELISA (抗原捕捉式酵素連結免疫吸附法) ------------ 62
二、VLPs誘發魚類產生抗體力價之強度及抗體的持續有效性 -------------- 63
(1)以VLPs誘發龍膽石斑產生的免疫效力 ---------------------------------- 63
(2)佛氏完全佐劑(FCA)對VLPs誘發龍膽石斑免疫效價之影響 -------- 65
(3)龍膽血清抗體之中和病毒能力 --------------------------------------------- 66
(4)以VLPs誘發瑪拉巴石斑產生的免疫效力 -------------------------------- 67
(5)佛氏完全佐劑(FCA)對VLPs誘發瑪拉巴石斑免疫效價之影響 ------ 67
(6)瑪拉巴石斑之抗體中和病毒能力 ------------------------------------------- 68
(7)以VLPs誘發棕點石斑產生的免疫效力 ----------------------------------- 68
(8)棕點石斑之抗體中和病毒能力 ---------------------------------------------- 69
(9)以VLPs誘發石斑魚產生抗體之持久性 ----------------------------------- 70
肆、討論 ----------------------------------------------------------------------- 71
一、VLPs誘發魚體特異性抗體生成時間與持久性 --------------------------- 71
二、免疫劑量、劑數影響石斑魚之免疫力價 ------------------------------------- 73
三、佛氏完全佐劑(FCA)對VLPs誘發石斑魚免疫效價之影響 ----------- 76
四、VLPs誘發石斑魚抗血清之中和病毒能力 --------------------------------- 78
五、VLPs對魚體、魚種間之免疫差異性 ---------------------------------------- 80
六、石斑魚抗血清檢測法之優缺 ------------------------------------------------- 83
伍、參考文獻 ---------------------------------------------------------------- 89
陸、圖表 ---------------------------------------------------------------------- 115
附錄A ------------------------------------------------------------------------- 145
附錄B ------------------------------------------------------------------------- 146
附錄C ------------------------------------------------------------------------- 147

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