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研究生:楊忠達
研究生(外文):Chung-Dar Yang
論文名稱:弓蟲SAG1及SAG2次單位疫苗對BALB/c小鼠保護率之分析
論文名稱(外文):Analysis of the protective capacity of SAG1 and SAG2 subunit vaccines in BALB/c mice
指導教授:趙大衛趙大衛引用關係張甘楠
指導教授(外文):David ChaoGan-Nan Chang
學位類別:博士
校院名稱:國立中山大學
系所名稱:生物科學系研究所
學門:生命科學學門
學類:生物學類
論文種類:學術論文
論文出版年:2004
畢業學年度:92
語文別:中文
論文頁數:256
中文關鍵詞:緩殖子弓蟲速殖子
外文關鍵詞:SAG2Toxoplasma gondiiSAG1
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II
中文摘要
弓蟲表面膜蛋白分子SAG1 與SAG2 對蟲體入侵宿主細胞非常
重要,本研究以這兩個蛋白質分子為主角,來發展不同組合的次單位
疫苗,並以BALB/c 小鼠為實驗動物,來評估所發展的各種次單位疫
苗的保護效果。本研究首先根據免疫網路學說製備具有SAG2 內像
(SAG2 internal image)的anti-idiotype IgG(aId-IgG)來作為疫苗,
然後將2 µg、4 µg、8 µg 的aId-IgG 以皮下免疫三組小鼠後,分析小
鼠的免疫反應,發現小鼠淋巴球以弓蟲抗原刺激後可造成明顯的增殖
反應,並分泌大量的IFN-γ,上述的免疫小鼠以1×103 速殖子
( tachyzoites)作皮下感染攻擊,結果小鼠存活率為75%~88%。另
將弓蟲SAG1 基因與SAG2 基因分別選殖於載體pGEX-6P-1,並以大
腸桿菌BL21 star(DE-3)來表現重組蛋白rSAG1 與rSAG2,然後
將rSAG1 與rSAG2 分別以不同劑量及免疫方式進行動物實驗,發現
小鼠以腹腔免疫重組蛋白10 µg rSAG1 或10 µg rSAG2 所引起的淋巴
球增殖反應較明顯,且可使免疫後的小鼠淋巴球分泌大量IFN-γ及少
量的IL-4,因此反應是偏向Th1 相關的反應;反之以其他不同劑量
的抗原經皮下免疫小鼠後,卻導致IFN-γ分泌量的降低,經免疫反應
分析後,每隻免疫後的小鼠以1×103 速殖子作皮下攻擊感染,結果發
現小鼠腹腔免疫10 µg rSAG1 或10 µg rSAG2 均有64%小鼠可存活。
更進一步將10 µg rSAG1 與10 µg rSAG2 混合後腹腔免疫小鼠,再以
1×103 速殖子進行皮下攻擊感染,小鼠的存活率可達71%,說明混合
重組蛋白rSAG1 與rSAG2 的保護效果比單僅免疫rSAG1 或rSAG2
的結果更好,這結果鼓勵我們製備嵌合蛋白(chimeric protein),即將
SAG1 基因與SAG2 基因先後選殖在同一載體上,使兩段基因接合在
一起,因此所表現的重組蛋白rSAG1/2 將同時具有SAG1 與SAG2
蛋白質的抗原性,以不同劑量的rSAG1/2 進行腹腔免疫小鼠,分析
這些小鼠的免疫反應後,發現以腹腔免疫10 µg rSAG1/2 的小鼠淋巴
III
球在弓蟲抗原TsoAg 的刺激下可造成明顯的增殖反應,而淋巴球也
可以分泌大量的IFN-γ,更驚訝的是在進行皮下感染攻擊之後,竟可
造成80%小鼠的存活,因此rSAG1/2 的保護效果會稍高於混合10 µg
rSAG1 與10 µg rSAG2 的效果。另外,本研究也探討以忌熱性腸毒素
(heat-labile enterotoxin, LT)LTA2B 或LTB 作為腸道黏膜系統的刺激
物來提高黏膜系統抵抗自然感染弓蟲的能力,即將LTA2B 基因與LTB
基因分別選殖後,獲得重組蛋白rLTA2B 與rLTB,然後將rLTA2B 或
rLTB 混合rSAG1/2 進行動物實驗並分析免疫反應,除混合型的抗原
組合外,也討論將LTA2B 基因或LTB 基因和SAG1 基因與SAG2 基因
接合在一起,形成重組蛋白rLTA2B-SAG2/1 或rLTB-SAG1/2 的融合
型抗原,以鼻腔內免疫方式將上述混合型或融合型抗原免疫動物,結
果發現在所有鼻腔內免疫的條件中以4 µg rLTB 混合10 µg rSAG1/2
免疫組以及10 µg rLTB-SAG1/2免疫組的小鼠所展現的淋巴球增殖反
應較好,而這兩組小鼠的腸道IgA 抗體的分泌量也較高,雖然這兩組
小鼠的淋巴球只分泌大量的IL-4,但在以1×103 速殖子皮下感染攻擊
後,在這兩組展現的保護效果(67%)明顯高於其他免疫組,而其餘
的抗原組合方式所表現出來的保護效果均在27%以下,所以利用
rLTB 作為黏膜刺激物的效果優於rLTA2B,且rLTB 不論混合rSAG1/2
或融合rSAG1/2 均可促進黏膜對弓蟲抗原的反應。以不同方式製備
的抗原可因處理方式、抗原組合方式及劑量不同而造成不同的結果,
其中以aId-IgG 的保護效果最好,其次是rSAG1/2,再其次是rSAG1
混合rSAG2,雖然rLTB 混合rSAG1/2 以及rLTB-SAG1/2 所展現的保
護效果不比上述三者高,但卻顯示重大的意義,即自然感染弓蟲的途
徑仍可利用黏膜刺激物來提高抵抗力,所以這些抗原將來均可成為疫
苗發展時的優良候選抗原。
IV
英文摘要
SAG1 and SAG2 are important surface molecules of T. gondii for the
invasion of tachyzoites into host-cells. Previous studies have been
demonstrated they are good candidates for development of vaccines
against toxoplasmosis. Therefore, we used SAG1 and SAG2 to generate
subunit vaccines and evaluated the protective capacity in BALB/c mice.
Anti-idiotype IgG (aId-IgG) with the SAG2 internal image was prepared
from anti-SAG2 monoclonal antibody in accordance with the network
theory. Lymphocytes from mice immunized subcutaneously twice with 2,
4 or 8 µg aId-IgG showed great proliferations and secreted high level of
IFN-γ, which is an important cytokines secreted by Th1 cells. After
challenged subcutaneously with 1×103 tachyzoites, the highest survival
rate reached 88%. Further, SAG1 and SAG2 genes were respectively
cloned and recombinant proteins rSAG1 and rSAG2 were prepared.
Lymphocytes from mice immunized intraperitoneally twice with rSAG1
or rSAG2 displayed apparently Th1-associated responses, while
lymphocytes from mice immunized subcutaneously twice with rSAG1 or
rSAG2 did not. Mice immunized intraperitoneally twice with rSAG1 or
rSAG2 had a survival rate of 64% which was higher than those mice
immunized subcutaneously with rSAG1 or rSAG2. When mice
immunized intraperitoneally twice with rSAG1 mixed with rSAG2,
survival rate was even higher (71%). Therefore, mixed antigens induced
higher protection. Subsequently, SAG1 gene was fused with SAG2 gene
and a recombinant protein rSAG1/2 was expressed from the fused gene.
Th1-associated responses were detected from lymphocytes of mice
immunized intraperitoneally twice with 10 µg rSAG1/2. Interestingly,
80% rSAG1/2-immunized mice survived and it was extremely higher
V
than rSAG1- or rSAG2-immunized mice (64%). In an attempt to
stimulate immune responses against T. gondii infections in the mucosal
system, we chose heat-labile enterotoxin (LT) secreted from toxigenic E.
coli as the stimulator. LTA2B and LTB genes were respectively cloned and
then rLTA2B and rLTB were obtained. Moreover, LTA2B gene or LTB
gene fused with SAG1 and SAG2 genes was performed and then
recombinant proteins rLTA2B-SAG2/1 and rLTB-SAG1/2 were prepared.
Subsequently, mice were immunized intranasally twice with
rLTA2B-SAG2/1, rLTB-SAG1/2, rLTA2B mixed with rSAG1/2, or rLTB
mixed with rSAG1/2. A strong protection (67%) was shown by the group
of mice immunized intranasally with 10 µg rLTB-SAG1/2 or 4 µg rLTB
mixed with 10 µg rSAG1/2. rLTB, rather than rLTA2B, will be a better
candidate for stimulation the mucosal system. In summary, different
survival rates caused by antigens prepared in the study may be attributed
to many factors such as the treatment, the preparation and the dose of
antigen. The highest survival rate is caused by aId-IgG (88%); the second
is shown by rSAG1/2 (80%); the third is resulted from rSAG1mixed
rSAG2 (71%). Although the survival rate raised by rLTB-SAG1/2 or
rLTB mixed rSAG1/2 is slightly low (67%), these data demonstrate
stimulators such as LT could induce anti-Toxoplasma immune response
of the mucosal system. Antigens capable of inducing higher survival rates
in mice should be worthy of further investigation for searching better
vaccine candidates.
總目錄
中文摘要………………………………………………………I
英文摘要…………………………………………………….. III
總目錄………...……………………………………………….V
表目錄………………………………………………………...XI
圖目錄………………………………………………………..XII

第一章 總論…...……………………………………………....1
第1節 弓蟲的發現………………………………………………1
第2節 弓蟲之形態及生活史……………………………………2
第3節 弓蟲之傳播與流行病學…………………………………5
第4節 臨床症狀與病變…………………………………………7
第5節 弓蟲症之治療…………………………………………..10
第6節 弓蟲症之診斷方式……………………………………..12
第7節 弓蟲症之疫學研究……………………………………..16

第二章 研究動機與目的…………………………………..25

第三章 研究內容…………………………….....................26
第1節 Anti-idiotype antibody對小鼠之保護率分析..............27
前言……………….……………………………………………27
材料與方法…………………………………………………….30
1. 單株抗體之製備…………………………………..........30
2. Idiotypic vaccine(anti-idiotype IgG)之製備………...35
3. aId-IgG免疫計劃…………………………………….....36
4. 淋巴球增殖試驗………………………………………..36
5. IFN-�蚺坐尷R…………………………………………….38
6. 活蟲攻擊及保護率之評估……………………………..39
結果…………………………………………………………….40
1. 單株抗體之Fab(或Id)……………………………..40
2. aId-IgG之抗原性……………………………………....40
3. 免疫血清之專一性……………………………………..41
4. 明顯之淋巴球增殖反應………………………………..41
5. 提高IFN-�蚺尷c…………………………………………42
6. 評估存活率……………………………………………..42
討論…………………………………………………………….43
附表…………………………………………………………….49
附圖…………………………………………………………….50

第2節 重組蛋白rSAG2對小鼠之保護率分析…………..…..56
前言…………………………………………………………….56
材料與方法…………………………………………………….59
1. SAG2基因第一階段選殖…………………....................59
2. SAG2基因第二階段選殖………………………………63
3. 重組蛋白rSAG2之製備……………………………….65
4. 免疫小鼠rSAG2…………………………………..........67
5. 分析接種rSAG2之免疫免應………………………….67
結果……………………………………………………………..71
1. 重組蛋白rSAG2之抗原性分析……………………….71
2. 小鼠免疫rSAG2之血清分析………….………..……..72
3. 小鼠免疫rSAG2之淋巴球增殖反應...………………..72
4. 小鼠免疫rSAG2之細胞素分析...……………………..73
5. 小鼠免疫rSAG2之存活率...…………………………..73
討論…………………………………………………………......75
附表……………………………………………………………..82
附圖……………………………………………………………..84

第3節 重組蛋白rSAG1對小鼠之保護率分析………………..97
前言……………………………………………………………..97
材料與方法……………………………………………………101
1. SAG1基因第一階段選殖……………………………...101
2. SAG1基因第二階段選殖……………………………...102
3. 重組蛋白rSAG1之製備………………………………102
4. 免疫小鼠rSAG1……………………………………….103
5. 分析接種rSAG1之免疫反應…………………………103
結果……………………………………………………………104
1. 重組蛋白rSAG1之抗原性分析………………………104
2. 小鼠免疫rSAG1之血清分析…………………………105
3. 小鼠免疫rSAG1之淋巴球增殖反應……….………...105
4. 小鼠免疫rSAG1之細胞素分析.………...……………106
5. 小鼠免疫rSAG1之存活率.......……………………….106
討論…………………………………………………………...108
附表…………………………………………………………...113
附圖…………………………………………………………...115

第4節 rSAG1混合rSAG2對小鼠之保護率分析……………128
前言……………………………………………………………128
材料與方法……………………………………………………130
1. 抗原來源……………………………………………….130
2. 免疫小鼠及活蟲攻擊………………………………….130
3. 分析接種rSAG1混合rSAG2之免疫反應…………..130
結果……………………………………………………………131
1. 小鼠免疫rSAG1混合rSAG2之血清分析……………131
2. 小鼠免疫rSAG1混合rSAG2之淋巴球增殖試驗.….131
3. 小鼠免疫rSAG1混合rSAG2之細胞素分析…….….132
4. 小鼠免疫rSAG1混合rSAG2之存活率......................132
討論……………………………………………………………133
附表……………………………………………………………135
附圖……………………………………………………………137

第5節 SAG1與SAG2之嵌合蛋白對小鼠之保護率分析…..140
前言……………………………………………………………140
材料與方法……………………………………………………142
1. 以pGexSAG1選殖SAG2(HindIII / HindIII)基因片段.………………………………………………………142
2. 純化rSAG1/2蛋白…………………………………….143
3. 免疫小鼠及活蟲攻擊………………………………….143
4. 分析接種rSAG1/2之免疫反應……………………….143
結果……………………………………………………………144
1. 表現融合基因SAG1/2…………………………………144
2. 小鼠免疫rSAG1/2之血清分析……………………….144
3. 小鼠免疫rSAG1/2之淋巴球增殖試驗.………………145
4. 小鼠免疫rSAG1/2之細胞素分析…………………….146
5. 小鼠免疫rSAG1/2之存活率分析…………………….146
討論……………………………………………………………147
附表……………………………………………………………154
附圖……………………………………………………………156

第6節 忌熱性腸毒素提高小鼠黏膜系統抵抗弓蟲感染………………………………………………………...167
前言……………………………………………………………167
材料與方法……………………………………………………173
1. 分子選殖LTB基因……………………………………173
2. 分子選殖LTB-SAG1/2基因…………………………...174
3. 分子選殖LTA2B-SAG2/1基因…………………………174
4. 純化重組蛋白rLTB-SAG1/2…………….……………176
5. 純化重組蛋白rLTA2B-SAG2/1…………………….…176
6. 免疫小鼠……………………………………………….177
7. 免疫反應分析………………………………………….177
結果……………………………………………………………179
1. 重組蛋白rLTB與rLTB-SAG1/2………………………179
2. 重組蛋白rLTA2B與rLTA2B-SAG2/1…………………179
3. 以腸毒素為免疫加強劑之血清分析………………….181
4. 以腸毒素為免疫加強劑之腸道IgA分析…………….182
5. 以腸毒素為免疫加強劑之淋巴球增殖試驗………….183
6. 以腸毒素為免疫加強劑之細胞素分析……………….183
7. 以腸毒素為免疫加強劑之存活率…………………….184
討論……………………………………………………………186
附表……………………………………………………………191
附圖……………………………………………………………194

第七節 結論……………………………………………………..223
附表……………………………………………………………225
附錄…………………………………………………………..226
參考文獻……………………………………………………..236



表目錄

表1-1:分析小鼠免疫aId-IgG之淋巴球增殖反應及IFN-�蚺尷c……..49
表2-1:分析小鼠免疫rSAG2之血清IgG2a / IgG1……….…………..82
表2-2:小鼠免疫rSAG2之淋巴球增殖反應及細胞素分泌..................83
表3-1:分析小鼠免疫rSAG1之血清IgG2a / IgG1………………….113
表3-2:小鼠免疫rSAG1之淋巴球增殖反應及細胞素分泌…………114
表4-1:分析小鼠免疫rSAG1混合rSAG2之血清IgG2a / IgG1……135
表4-2:小鼠免疫rSAG1混合rSAG2之淋巴球增殖反應及細胞素分泌………………………………………………….………...136
表5-1:分析小鼠免疫rSAG1/2之血清IgG2a / IgG1………………..154
表5-2:小鼠免疫rSAG1/2之淋巴球增殖反應及細胞素分泌……….155
表6-1:以腸毒素為免疫加強劑之血清IgG2a / IgG1分析………….191
表6-2:以腸毒素為免疫加強劑之淋巴球增殖反應與細胞素分泌....192
表7-1:融合蛋白對BALB/c小鼠之預備試驗……………………….225


圖目錄
圖一:弓蟲之生活史……..………………………………………......…23
圖二:速殖子的外形……………………………………………….......24
圖1-1:單株抗體TGCM#5以12% SDS-PAGE分析………............…50
圖1-2:單株抗體TGCM#5之亞型鑑定…………………….…………51
圖1-3:分析單株抗體TGCM#5之抗原專一性…………..………….52
圖1-4:分析anti-idiotype IgG(aId-IgG)之抗原性……….………..53
圖1-5:分析免疫aId-IgG之血清專一性………………..……………54
圖1-6:小鼠免疫aId-IgG之存活率………………………..………….55
圖2-1:兩階段選殖SAG2基因之流程圖……………………..………84
圖2-2:弓蟲表面抗原SAG2基因之PCR增幅……..……………….85
圖2-3:以載體pET32a選殖SAG2基因………………………………86
圖2-4:製備SAG2(BamHI / NotI)(I)..……………………………87
圖2-5:製備SAG2(BamHI / NotI)(II)..…………………….…….88
圖2-6:以載體pGEX-6P-1來選殖SAG2(BamHI / NotI)…………89
圖2-7:SAG2基因之核酸定序結果…………………………………..90
圖2-8:純化重組蛋白rSAG2之流程…………………………………91
圖2-9:以12% SDS-PAGE分析rSAG2……………………………..92
圖2-10:分析rSAG2之抗原性………………………………………93
圖2-11:分析小鼠免疫rSAG2之血清專一性……………….……...94
圖2-12:分析各組小鼠免疫rSAG2之血清…………………………95
圖2-13:小鼠免疫rSAG2之存活率………………………….……...96
圖3-1:兩階段選殖SAG1基因的流程圖.…………………………….115
圖3-2:弓蟲表面抗原SAG1基因之PCR增幅.………………….….116
圖3-3:以載體pET32a選殖SAG1基因.…………………….………117
圖3-4:製備SAG1(EcoRI / XhoI)(I).……………………………118
圖3-5:製備SAG1(EcoRI / XhoI)(II).………………………….119
圖3-6:以載體pGEX-6P-1來選殖SAG1(EcoRI / XhoI).……….120
圖3-7:SAG1基因的核酸定序結果…………………………………122
圖3-8:以12% SDS-PAGE分析rSAG1……………………………..123
圖3-9:分析rSAG1之抗原性…..………………………………….. 124
圖3-10:分析小鼠免疫rSAG1之血清專一性………………………..125
圖3-11:分析各組小鼠免疫rSAG1之血清..……………………….126
圖3-12:小鼠免疫rSAG1之存活率…..……………………..………127
圖4-1:小鼠免疫rSAG1混合rSAG2之血清專一性………………137
圖4-2:分析小鼠免疫rSAG1混合rSAG2之血清………………….138
圖4-3:小鼠免疫rSAG1混合rSAG2之存活率…………………….139
圖5-1:以pGexSAG1選殖SAG2基因(HindIII / HindIII)………156
圖5-2:SAG1基因接合SAG2基因之菌落篩選…………………….157
圖5-3:SAG1/2基因之核酸定序結果…………………….…………158
圖5-4:重組蛋白rSAG1/2之分子量分析…………………………..161
圖5-5:小鼠感染弓蟲血清分析rSAG1/2之抗原性………………..162
圖5-6:以單株抗體TGCM#4與TGCM#5分析rSAG1/2之抗原性...163
圖5-7:小鼠免疫rSAG1/2之血清專一性...…………………………164
圖5-8:分析小鼠免疫rSAG1/2之血清……………………………..165
圖5-9:小鼠免疫rSAG1/2之存活率………………………………..166
圖6-1:LT毒素……………………………………………………….194
圖6-2:LTB基因之PCR增幅……………………………………….195
圖6-3:以pGEX-6P-1載體選殖LTB基因(BamHI / BamHI)……….196
圖6-4:LTB基因之核酸定序結果…………………………………..197
圖6-5:rLTB之抗原性分析………………………………………….198
圖6-6:以pGexLTB作為載體來選殖SAG1/2基因(EcoRI / NotI)…199
圖6-7:LTB-SAG1/2基因的定序結果……………………………….202
圖6-8:rLTB-SAG1/2之抗原性分析………………………………..203
圖6-9:LTA2B基因之PCR增幅……………………………………..204
圖6-10:以pGEX-6P-1作為載體來選殖LTA2B基因(BamHI / BamHI)…………………………………………………..205
圖6-11:LTA2B基因定序的結果…………………………………….206
圖6-12:rLTA2B之抗原性分析……………………………………..207
圖6-13:LTA2B、SAG1與SAG2三段基因的接合的順序…………208
圖6-14:以pGexSAG2作為載體來選殖LTA2B基因(BamHI / BamHI)…………………………………………………….209
圖6-15:SAG1基因(HindIII / HindIII)之PCR增幅……………….210
圖6-16:以pGexLTA2B-SAG2來選殖SAG1基因(HindIII / HindIII)……………………………………………………..211
圖6-17:LTA2B-SAG2/1基因之核酸定序結果………………………214
圖6-18:rLTA2B-SAG2/1的抗原性分析……………………………..215
圖6-19:小鼠鼻腔內免疫腸毒素及重組抗原的血清專一性分析…216
圖6-20:以rLTA2B為免疫加強劑之血清分析……………………..217
圖6-21:以rLTB為免疫加強劑之血清分析………………………..218
圖6-22:小鼠腸道IgA分析…………………………………………219
圖6-23:小鼠鼻腔內免疫rLTA2B-SAG2/1或rLTA2B混合rSAG1/2之存活率……………………………………………………220
圖6-24:分析小鼠鼻腔內免疫rLTB-SAG1/2或rLTB混合rSAG1/2之存活率..…………………………………….……………221
圖6-25:分析小鼠鼻腔內免疫rSAG1/2之存活率………………..222
236
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