臺灣博碩士論文加值系統

自行政院衛生署預防醫學研究所,蒐集1991至1994年鼠傷寒沙門氏菌(Salmonella enterica subsp. enterica servar Typhimurium)人體分離株共45株，主要分離自臨床下痢者與食物中毒案例患者之血液與糞便檢體，小鼠動物毒性結果可將菌株毒性分成兩群，即高毒性群(high virulence)與低毒性群(low virulence)。隨機挑選五株脈衝式電場膠體電泳法(pulsed-field gel electrophoresis, PFGE)分型之主要次分型(subtype)菌株(X5A4S4)，分析其動物毒性，五株主要次分型之菌株皆為高毒性菌株。並非所有臨床S. Typhimurium菌株皆具有毒性質體，PFGE主要次分型菌株13株 (28.9 %)帶有毒性質體，臨床來源菌株有毒性質體共有30株(66.7 %)，無毒性質體者為33.3 % (15/45)。45株S. Typhimurium均具有inv與stn基因，且stn基因之限制酶片段多型性(restriction fragment length polymorphism, RFLP)於菌株間並無不同，S. Typhimurium菌株之腸毒素力價(titer of enterotoxin)介於4至32之間，三株PFGE主要次分型菌株力價最高。S. Typhimurium於小鼠來源Raw 264.7巨噬細胞內中侵入、抗吞噬能力與菌株毒性並不相關，因此，S. Typhimurium於Raw 264.7巨噬細胞侵入與抗吞噬能力能力，不適合用以鑑別菌株之毒性。S. Typhimurium菌株對巨噬細胞Raw264.7之毒殺能力與毒性無關，屬於低毒性次分型X6A7S6與X17之菌株對H2O2敏感性較高。S. Typhimurium高毒性菌株之耐酸性 (acid tolerance response, ATR) 能力較強，以SDS-聚丙烯醯胺膠體電泳 (polyacrylamide gel electrophoresis, PAGE) 分析S. Typhimurium胞外分泌蛋白質之類型，結果發現：無法找出胞外分泌蛋白質類型與菌株毒性之相關性，PFGE主要次分型菌株於耐酸性、抗H2O2、腸毒素力價等毒性因子之整體表現較其他菌株毒性高，可能為此次分型菌株成為高毒性與主要致病流行菌株之主要原因。S. Typhimurium之毒性與刺激巨噬細胞Raw264.7產生腫瘤壞死因子-a(tumor necrosis facrtor alpha, TNF-a) 與生物膜(biofilm)產生能力無關，無法取代動物試驗。
近年來多重抗藥性(multidrug resistant) S. Typhimurium菌株逐年增加，本研究收集並分析比較台灣地區動物與人體分離之S. Typhimurium抗微生物藥劑圖譜(antibiogram)，透過這些抗藥基因之研究以了解抗藥基因之起源，結果顯示本地區分離之人體與家畜等動物來源S. Typhimurium抗微生物藥劑圖譜相近。台灣地區人體與動物分離菌株對第一線抗微生物藥劑如tetracycline 、sulfisoxazole 、ampicillin、streptomycin、chloramphenicol具高抗藥性，對gentamicin、fluoroquinolone類抗微生物藥劑如norfloxacin與第三代頭芽胞菌素cefoperazone則為敏感性，93~100 %之本地區菌株受其抑制。台灣S. Typhimurium菌株多重抗藥性比例高，動物來源佔58.6 %，人體來源菌株亦有68.9 %為多重抗藥性，且其PFGE主要次分型菌株為對ampicillin、chloramphenicol、streptomycin、sulfonamide、tetracycline (ACSSuT)之多重抗藥性菌株。以5’CS/3’CS引子組進行聚合酶鏈鎖反應 (polymerase chain reaction, PCR)分析integron，PFGE次分型與抗微生物藥劑圖譜均相同之兩來源菌株，二者之抗藥基因integron大小相同，台灣人體來源分離之多重抗藥性S. Typhimurium菌株，可能是由感染之動物經食物鏈媒介傳播。
多重抗藥性之S. Typhimurium菌株，逐年增加成為世界各地沙門氏菌病之元兇，成為 Salmonella 感染時，臨床治療上的一大問題，因此，醫生與科學家將治療病原菌感染由抗微生物藥劑之施用，轉為使用益生菌強化腸黏膜防禦機制，保護腸道免於病原菌之侵襲。乳酸菌為人體腸道之益生菌，作為益生菌之基本條件，包括具有耐酸性、耐膽鹽性、抑制病原菌的生長，且能附著於宿主的腸道上皮細胞。本研究先以磷酸鹽緩衝系統 (pH 2) 以及牛膽汁 (0.3 ﹪oxgall)，篩選出動物與植物來源之耐酸、耐膽鹽之乳酸菌菌株LA5、LF33、MP012與CCRC 10069，再以人體腸道細胞株Int 407及Caco-2來進行體外附著試驗，菌株LA5、LF33具有與腸上皮細胞附著能力。受試乳酸菌株經連續口服7天後，四株乳酸菌對S. Typhimurium於小鼠體內的感染提供了部分保護作用，以LA5效果最佳，CCRC 10069與LF33再次之，MP012再次之，由於CCRC 10069在人體腸道細胞株Int 407及Caco-2不具附著性，而於小鼠活體具有保護 Salmonella 侵入之功效。因此，進一步探討 CCRC 10069於活體具保護效果之原因。在小鼠腸道吸附性試驗中，CCRC 10069具有吸附性，推測CCRC 10069於小鼠感染S. Typhimurium時可提供保護作用與其吸附能力有關。動物性來源之乳酸菌LA5、LF33於小鼠腸道中具高吸附現象，與其對小鼠有腸道保護作用有關。
由於 Salmonella spp.與Listeria monocytogenes為引起人類食品中毒重要之食品病原菌，本研究發展同時檢測Salmonella spp.與Listeria monocytogenes之方法，此方法為免疫磁珠分離法(immunomagnetic separation method, IMS)合併聚合酶連鎖反應之方法(polymerase chain reaction, PCR)，並將此方法應用於食品中Salmonella與Listeria monocytogenes之同時檢測。若只使用多套式PCR，雖然兩目標細菌菌數都在檢測靈敏度之上，若菌數差異在兩個對數值以上，例如n x 107 與 n x 104 或是n x 106 與 n x 103，則無法檢測出菌數較低之目標菌；若在PCR反應前，使用等量anti-Salmonella 與anti-Listeria免疫磁珠吸附目標細菌菌體，則可以同時檢測出兩種目標細菌。經過增菌步驟，IMS-mPCR可成功應用於牛奶、乳製品、肉類等食品樣品中Salmonella 與Listeria monocytogenes之檢測。

45 Salmonella Typhimurium strains isolated between 1991-1994 were collected from the National Institute of Preventive Medicine, Department of Health, Executive Yuan. These S. Typhimurium strains were divided into two groups，ie, high and low virulence groups, according to the virulence study to the mice. Five strains randomly selected from the major PFGE subtypes were subjected to the virulence study to mice. It was found that they were all high virulent strains. Virulence plasmids could not be isolated from all of these 45 strains. However, 13 S. Typhimurium strains (28.9 %) of the major PFGE subtype carry the virulence plasmid and 30 strains (66.7 %) of the clinical isolates carry the virulence plasmid. Virulence plasmid were not isolated from 33.3 % (15/45) of S. Typhimurium strains studied. On the other hand, the inv and stn genes were found in all of the 45 strains. The restriction fragment length polymorphism (RFLP) patterns of stn gene show no difference for these 45 Salmonella Typhimurium strains. The relationships between invasive ability to human and the enterotoxin titers of S. Typhimurium strains were not found. Also, the relationship between invasion ability and replication capability to the virulence of S. Typhimurium cells in Raw 264.7 macrophage cell line was not found. Thus, the abilities of invasion and replication of S. Typhimurium to Raw 264.7 were not useful for the analysis of Salmonella virulence The relationship between cytotoxic to macrophage and virulence was not found. Strains in PFGE subtypes X6A7S6 and X17 were belonging to low virulence strains and were more susceptible to H2O2. Strains of high virulence subtypes were more acid tolerant. SDS-polyacrylamide gel electrophoresis (SDS-PAGE) patterns of the secretion proteins for those Salmonella Typhimurium strains were not relative to their virulence. Strains from PFGE major subtype were more acid tolerant, more resistant to H2O2, and possess higher enterotoxin titer, which might be the reasons of being virulent and epidemic. Tumor necrosis factor alpha (TNF-a) formations in macrophage and biofilm formation ability were also not related to the virulence of these S. Typhimurium strains. The method mentioned above could not replace the method of animal models for the study of the virulence of S. Typhimurium.
In recently years, the number of multidrug resistant Salmonella Typhimurium strains was increasing. In this study, the antibiograms of 45 human isolates and 87 animal isolates of Salmonella Typhimurium in Taiwan was investigated. Analysis of the antibiotic resistant genes in molecular level may help us to realize the origins of antibiotic resistance strains. It was found that the antibiograms for human and animal isolates are quite similar. Both the human and animal isolates are highly resistant to first-line antibiotics, such as tetracycline, sulfisoxazole, ampicillin, streptomycin and chloramphenicol; but are sensitive to gentamicin, fluoroquinolone antibiotics, such as norfloxacin and the third generation antibiotic of cephalosporin, such as cefoperazone. Between 93 % and 100 % of the local strains is inhibited by these antibiotics. Also, a significant fraction of these S. Typhimurium isolates are multidrug resistant strains. For example, 58.6 % of the animal isolates and 68.9 % of the human isolates are multidrug-resistant. Strains from PFGE major subtype were resistant to ampicillin, chloramphenicol, streptomycin, sulfonamide, tetracycline (ACSSuT). When sizes of the integrons were analyzed using PCR primers 5’CS/3’CS, the integron sizes were the same for human and animal isolates, which show identical PFGE and antibiogram patterns. Thus, antibiotic resistant human isolates may be circulated through foods from the infectious animals.
The increasing of multidrug resistant Salmonella Typhimurium strains in recent years and the predominance of these multidrug resistant strains as major infectious strains of Salmonellosis all over the world become a serious problem. Such fact has induced the interest of probiotic use. Probiotic are capable of protecting the gastrointestinal tract from invasion by pathogens. The effectiveness of probiotic could be assessed by their prevention of pathological conditions and by their form as biotherapeutic agents. Due to the probiotic properties of lactic acid bacteria (LAB), we have used some LAB strains for the study of their prevention of Salmonella invasion. The basic probiotic properties evaluated for LAB strains are their capability to tolerate to gastric acid and bile salts, the ability to inhibit pathogenic bacteria and the adherence capability to human intestinal epithelium cells. In this study, four LAB strains LA5, LF33, MP012 and CCRC 10069 have been shown to be tolerant to pH 2.0 buffer, and resistant to 0.3% bile salts. However, for the adhesive ability study, only LA5 and LF33 strains were found adhesive to the human Int 407 or Caco-2 cell lines. The protective effectiveness for the orally administered LAB strains to the invasion of mice spleen and liver organs by S. Typhimurium was investigated. Results showed that the cell numbers of S. Typhimurium in liver and spleen organs of mice measured at the 6th day after Salmonella infection was lower for LAB administered mice as compared to the control mice. Of them, strain LA5 strain was most effective; followed by strains CCRC 10069 and LF33. CCRC 10069 was not able to adhere to Int 407 or Caco-2 cell lines but it provides the protective effect too. These four LAB strains were also studied for their adhesion ability to the mice intestinal cells. CCRC 10069 show the adherence to the mice intestinal cells, and this may be the reason why CCRC 10069 provides the protective effect for S. Typhimurium infection too.
Owing to Salmonella and Listeria monocytogenes to human health, in this study, we also tried to develop a method for the simultaneous and rapid detection of these two kinds of organisms. The method we developed was the immunomagnetic separation followed by PCR method.
A method that combined the immunomagnetic separation (IMS) technique and the multiplex polymerase chain reaction (PCR) method (i. e., the IMS-mPCR method) were developed for simultaneous detection of Listeria monocytogenes and Salmonella spp. in food samples. When only the multiplex PCR method was used, it was found that if cell numbers of each of the two target organisms (L. monocytogenes and Salmonella spp.) were above the detection limit, but differed by more than 2 logs-e. g., n x 107 to n x 104 or n x 106 to n x 103 — the organism presenting the lower numbers might go undetected. Listeria and anti-Salmonella immunomagnetic beads was performed prior to PCR, both pathogens could be detected unambiguously. Such results could be obtained for target organisms in food samples, such as milk, dairy, and meat products, if similar enrichment and IMS steps were performed prior to PCR.

目次
頁次
中文摘要………………………………………………………………………...1
Abstract……………………………………………………………………4
第一章、研究背景與目的…………………………………………………8
第二章、文獻整…………………………………………………………….12
第一部分、Salmonella……………………………………………………12
一、 Salmonella 之型態及生理特性……………………………………12
二、Salmonella與食品中毒之關係……………………………………….13
三、Salmonella 感染之症狀………………………………………………13
四、Salmonella之傳統檢測………………………………………………14
五、S. Typhimurium之致病性……………………………………………14
(一)病原菌、病原性、毒性、毒性因子之定義………………………14
(二)Salmonella 成為病原菌之相關基因及機制……………………16
(三)毒性基因之調控……………………………………………………20
六、活體內測定Salmonella毒性方法……………………………………20
七、S. Typhimurium之抗藥性研究………………………………………21
(一)多重抗藥性菌株之增加與傳播…………………………………….21
(二)integron與抗藥基因……………………………………………...…21
(三)多重抗藥性S. Typhimurium之研究……………………………….22
八、乳酸菌對Salmonella之抑制………………………………………….23
(一)乳酸菌作為益生菌之特性……………………………………….…23
(二)乳酸菌對Salmonella之抑制…………………………………….…24
(三)、乳酸菌抑制Salmonella侵入或感染之相關研究…………………25
第二部分Listeria monocytogenes之簡介及其與Salmonella之快速檢驗法…………………………………………………25
一、Listeria monocytogenes簡介…………………………………………25
(一)、Listeria monocytogenes之一般特性………………………………25
二、Listeria monocytogenes之傳統檢驗法………………………………28
三、Salmonella、Listeria monocytogenes之快速檢驗法……………28
(一)DNA探針分析法…………………………………………………...28
(二)免疫分析……………………………………………………………29
(三)免疫磁珠分離法……………………………………………………30
(四)聚合酶鏈鎖反應及其應用…………………………………………31
表 2-1~表2-5…………………………………………………………33
圖2-1~圖.2-4………………………………………………………….41
第三章、人體來源S. Typhimurium毒性質體、致病基因、表型與動物毒性之關係………………………………………………….…………………45
壹、前言………………………………………………………… …………47
貳、材料與方法……………………………………………………………47
一、 實驗材料………………………………………………………………47
二、 實驗方法………………………………………………………………55
(一)小鼠模式毒性試驗…………………………………………………...55
(二)以聚合酶鏈鎖反應確認spv gene之存在……………………57
(三)以DNA雜合法確認spv位於質體上………………………………..58
(四)90 kb毒性質體之確認………………………………………………..60
(五)以聚合酶鏈鎖反應確認inv基因………………………………60
(六)S. Typhimurium對巨噬細胞株Raw 264.7侵入能力之測定………..60
(七)S. Typhimurium於巨噬細胞株Raw 264.7抗吞噬能力之測定……..61
(八)S. Typhimurium對巨噬細胞株Raw 264.7之細胞毒性測定………..61
(九)S. Typhimurium對H2O2之敏感性測定……………………………...62
(十)以聚合酶鏈鎖反應確認stn基因與PCR產物之限制酶片段多型性62
(十一) Salmonella腸毒素對CHO細胞毒性之分析…………………….63
(十二)S. Typhimurium之耐酸反應(Acid tolerance response; ATR)試驗.64
(十三)以SDS-聚丙烯醯胺膠體電泳分析胞外分泌蛋白質類型………..64
(十四)S. Typhimurium對巨噬細胞株Raw 264.7 產生TNF-a 能力之測定…………………………………………………………………………..…65
(十五)於adherence test medium(ATM)培養，生物膜(biofilm)產生能力之測定……………………………………………………………………65
(十六) 統計分析與繪圖………………………………………………….65
參、結果與討論……………………………………………………………66
(一)小鼠模式中S. Typhimurium之毒性…………………………………66
(二)S. Typhimurium90 kb毒性質體存在之確認…………………………67
(三)90 kb毒性質體與動物毒性、人類之致病性之關係…………………68
(四)以聚合酶鏈鎖反應確認inv基因………………………….…69
(五)S. Typhimurium對巨噬細胞株Raw 264.7………………………….69
(六)S. Typhimurium對H2O2敏感性測定………………………………..71
(七)S. Typhimurium菌株之stn基因檢測及其RFLP……………………72
(八)S. Typhimurium腸毒素對CHO細胞毒性分析………………………72
(九)耐酸反應(acid tolerance response; ATR)………………………73
(十)S. Typhimurium胞外分泌蛋白質類型………………………………74
(十一)S. Typhimurium之基因與毒性相關表型之綜合討論……………74
(十二)S. Typhimurium刺激巨噬細胞株Raw 264.7 產生TNF-a 能力之測定……………………………………………………………………………..75
(十三)於ATM培養S. Typhimurium生物膜產生能力之測定………….75
肆、結論…………………………………………………………………………76
表3-1~表3-10…………………………………………………………...78
圖3-1~圖3-13…………………………………………………………...98.
第四章、人體與家畜等動物來源S. Typhimurium抗微生物藥劑圖譜與抗藥性來源之分析………………………………………………………………106
壹、前言…………………………………………………………………….106
貳、材料與方法…………………………………………………………….108
一、實驗材料……………………………………………………………..108
二、方法…………………………………………………………………..109
(一)抗微生物藥劑感受性試驗…………………………………………..109
(二)以聚合鏈鎖反應檢測integron……………………………………110
(三)由洋菜瓊脂膠體上回收PCR產物…………………………………..110
(四)PCR產物定序……………………………………………………….111
(五) PCR產物核苷酸序列比對………………………………….111
參、結果與討論………………………………………………………….111
肆、結論………………………………………………………………….117
表4-1 ~表4-8 ………………………………………………………….118
圖4-1 ~圖4-3 ………………………………………………………….130
第五章乳酸菌抑制Salmonella之感染……………………………………133
壹、前言…………………………………………………………………133
貳、材料與方法…………………………………………………………….134
一、實驗材料……………………………………………………………..134
二、實驗方法……………………………………………………………..134
(一)耐酸性試驗…………..……………………………………………...136
(二)耐膽鹽性試驗…………………..…………………………………...136
(三) 腸道細胞株附著性 (adhesion)試驗…………………………….136
(四)乳酸菌抑制S. Typhimurium侵入腸道細胞株Int 407之試驗……137
(五) 乳酸菌於BALB/c小鼠模式中抑制S. Typhimurium之感染試驗..139
(六)乳酸菌在小鼠腸道上皮細胞之吸附……………………………….140
(七)統計分析……………………….……………………...140
參、結果與討論…………………………………………………………….140
一、耐酸性之評估………………………………………………………..140
二、耐膽鹽特性之評估…………………………………………………..141
三、乳酸菌於Int407與Caco-2腸上皮細胞株之吸附…………………….142
四、乳酸菌抑制S. Typhimurium侵入腸道細胞株Int 407之試驗………143
五、乳酸菌於小鼠活體對Salmonella之抑制作用……………………..145
六、乳酸菌吸附於小鼠腸道上皮細胞之試驗…………………………..147
肆、結論…………………………………………………………………….148
表5-1 ~表5-4…………………………………………………………..150
圖5-1 ~圖5-9………………………………………………………….154
第六章、結合免疫磁珠分離技術與多套式PCR同時檢測食品中Salmonella及Listeria monocytogenes…………………………………154
壹、前言…………………………………………………………...163
貳、材料與方法…………………………………………………………...165
一、實驗材料……………………………………………………………..165
二、實驗方法……………………………………………………………...167
(一)菌株之培養方法……………………………………………………….167
(二)多套式聚合酶鏈鎖反應……………………………………………….167
(三)免疫磁珠分離法之應用……………………………………………….168
(四)UP broth增殖培養後，以多套式PCR檢測Listeria monocytogenes及 Salmonella……………………………………………………………...169
(五)多套式PCR法應用於食品樣品中Listeria monocytogenes及 Salmonella之檢測………………………………………………………...169
參、結果與討論……………………………………………………………169
一、多套式PCR特異性之探討…………………….170
二、多套式PCR引子組之檢測靈敏度…………………………………….170
三、以UP 增殖培養步驟提高多套式PCR之檢測靈敏度……………….171
四、UP broth增殖培養後，以IMS-mPCR檢測Listeria monocytogenes及Salmonella……………………………………………………………….172
肆、結論………………………………………………………………...174
表 6-1~表6-4…………………………………………………………..175
圖6-1 圖6-4……………………………………………………………179
參考文獻……………………………………………………………………183

台灣省政府農林廳編印，民國八十七年，飼料管理手冊：飼料添加物詳細品目，p.300-301.
中國國家標準：食品微生物檢驗法-食品中 Listeria monocytogenes之檢驗標準CNS12542.
行政院衛生署乳品類衛生標準，88衛署食字第88017907號
邱政洵。民國86年。北台灣地區沙門氏菌感染之現況並以分子生物學與生物學方法探討沙門氏菌之毒性質體與其致病性和宿主特異性之關係。私立長庚大學臨床醫學研究所博士論文。
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