電解水 (Electrolyzed water) 作為良好的抑菌劑且廣泛應用於食品工業上。不足的殺菌條件下可能會導致菌體進入存活但不可培養狀態 (Viable but non-culturable, VBNC state) ，該狀態下的菌體已失去在平板上長出菌落的能力，但仍表現出可被檢測的代謝活性。李斯特菌 (Listeria monocytogenes) 作為輕度加工水產品及未經充分加熱的即食性食品中常見的污染菌，雖不是前五名的食品中毒病原菌，但卻能引起近 20% 的致死率。因該菌體對於乾燥、高溫、酸性環境、鹽等具有耐受性，且在該環境壓力下有使菌株進入 VBNC 狀態的可能性。因為食品業廣用標準平板計數方法無法偵測到 VBNC 菌體，這將導致嚴重的食安風險。因此，本研究將著重於 VBNC 菌體的生成機制探討，以避免食品病原菌進入 VBNC 狀態造成的食品中毒事件發生。加上，近年已有學者對革蘭氏陰性菌進入 VBNC 狀態做蛋白質體學的探究，但革蘭氏陽性菌的研究較為少數。本研究主要是使用微酸性電解水誘導 L. monocytogenes 進入 VBNC 狀態，再利用平板計數法結合流式細胞儀確認菌體的狀態，及使用掃描式電子顯微鏡去觀察形態變化，後續以 TMT 定量蛋白質體學分析 VBNC 狀態下菌體蛋白質的變化。研究結果表示使用有效氯為 8-10 mg/L 的微酸性電解水可以誘導 L. monocytogenes 進入 VBNC 狀態，且在電子顯微鏡的觀察下發現細胞膜有皺褶的現象。透過TMT標記定量串聯式質譜分析得到微酸性電解水誘導 VBNC 菌體的蛋白質體學資訊，總共鑑定到 203 個差異性表現蛋白 (Differential expressed proteins) ，包括 78 上調的蛋白及 125 下調蛋白。經過 GO 功能性註解及 KEGG 代謝路徑分析，差異性表現蛋白顯著與核醣體、次級代謝物的合成及胺醯 tRNA 合成酶有關。此外，在本研究中還鑑定了 31 段氯化胜肽 (22 個氯化蛋白質)，驗證在微酸性電解水的處理下會使蛋白質被氯化的現象。該氯化蛋白質主要為延長因子 Tu及伴護蛋白。從氯化胜肽的結構模體上 (sequence motif)，可以觀察到 6 段氯化胜肽 (pcrB in Y112, groEL in Y201, rpsS in Y61, 和 tuf in Y45, Y130, Y161) 在其酪胺酸上游 6 個胺基酸位點若為精胺酸 (RxxxxxcY motif)，則有利於被氯化的趨勢。因此，L. monocytogenes 在有效氯為 8-10 mg/L 的微酸性電解水處理下得以進入 VBNC 狀態，且該狀態下的蛋白質主要影響轉譯作用，其中延長因子 Tu 及伴護蛋白為主要被氯化的蛋白質。

Electrolyzed water (EW) is a good bactericidal agent widely used in food industry for sanitization. Insufficiency sanitization could induce foodborne pathogens to enter viable but non-culturable (VBNC) state, i.e. the cells that loss the ability to form colonies during traditional culturing but exhibit detectable metabolic activity. Listeria monocytogenes as a fatal foodborne pathogen generally found in minimal processing seafood and ready-to-eat (RTE) food. Although L. monocytogenes is not the top five germs that cause foodborne illness, listeriosis even causes nearly 20% fatality. Moreover, L. monocytogenes could endure various stress such as drought, extreme temperature, acid, and salinity etc. Thus, the stress may induce them to tentatively enter VBNC state and possibly lead to foodborne outbreaks due to their invisibility under conventional plate counting technique. Therefore, we have to concentrate on the formation mechanism of VBNC bacteria in order to prevent the bacteria enter VBNC state during the food processing. Several studies conducted the proteomic analysis on the VBNC state of Gram negative bacteria recently, but rarely for Gram positive bacteria. For this study, we use the slightly acidic electrolyzed water (SAEW) to trigger the VBNC state of L. monocytogenes, then plating method combing with flow cytometry is used to detect the VBNC bacteria and observe the morphology with scanning electron microscopy (SEM). Additionally, we explore the protein profiles of the VBNC bacteria using tandem mass tag (TMT) labelled-LC-MS/MS technique. The chlorine concentration of SAEW to develop VBNC state was 8-10 mg/L and bacteria showed the shrinkage of the cell membrane under SEM observation. Besides, the proteomic results indicated 203 differential expressed proteins (DEPs), including 78 up-regulated and 125 down-regulated DEPs. After gene ontology (GO) functional annotation and Kyoto encyclopedia of genes and genomes (KEGG) metabolic pathway analysis, the significant DEPs related to ribosome, biosynthesis of secondary metabolites and aminoacyl-tRNA biosynthesis. Additionally, we presently have identified 31 chlorinated peptides in the 22 chlorinated proteins to validate that SAEW induced the protein chlorination during the treatment. The most prominent chlorinated proteins were identified as the elongation factor Tu and chaperone proteins. The 6 peptides, namely pcrB in Y112, groEL in Y201, rpsS in Y61, and tuf in Y45, Y130, Y161, recognized as the RxxxxxcY motif. In conclusion, L. monocytogenes could enter VBNC state by 8-10 mg/L of ACC in SAEW. The critical protein of VBNC bacteria primarily involved in translation process, and furthermore the elongation factor Tu and chaperon were mainly chlorinated.

摘要 I
Abstract II
List of Figures V
List of Tables VI
List of Supplementary Figures and Tables VII
Abbreviations IX
Chapter 1 Introduction 1
Chapter 2 Literature Review 3
2.1 Foodborne pathogens 3
2.1.1 Listeria monocytogenes 3
2.1.2 Viable but non-culturable (VBNC) bacteria 4
2.2 Slightly acidic electrolyzed water (SAEW) 6
2.3 Proteomic analysis 8
2.3.1 Quantitative proteomic analysis 9
Chapter 3 Experimental Design 10
3.1 The formation of viable but non-culturable (VBNC) bacteria 10
3.2 The mechanism of formation VBNC bacteria 11
Chapter 4 Materials and Methods 12
4.1 Materials 12
4.1.1 Bacteria strain 12
4.1.2 Chemical reagents 12
4.2 Experimental produces 12
4.2.1 Bacterial cultures and preparations 12
4.2.2 Preparation of slightly acidic electrolyzed water (SAEW) 12
4.2.3 Pathogens treatment 13
4.2.4 Flow cytometry analysis of treated samples 13
4.2.5 Scanning electron microscopy (SEM) analysis 13
4.2.6 Total protein extraction of VBNC cells 14
4.2.7 Sodium dodecyl sulfate polyacrylamide gel electrophoresis (SDS-PAGE) 14
4.2.8 Whole-protein extraction for proteomic analysis 14
4.2.9 Mass spectrometric analysis 15
4.2.9.1 In solution digestion 15
4.2.9.2 Tandem Mass Tags (TMT) labelling 15
4.2.9.3 SDB fractionation and desalted 15
4.2.9.4 LC-MS/MS 16
4.2.9.5 Data analysis and protein network construction 16
4.2.10 Statistical analysis 16
Chapter 5 Experimental Results 17
5.1 The culturability of L. monocytogenes after treatment with slightly acidic electrolyzed water (SAEW) 17
5.2 Determination of cell viability of L. monocytogenes by flow cytometry 17
5.3 Morphological changes of VBNC L. monocytogenes by scanning electron microscopy 18
5.4 SDS-PAGE analysis 18
5.5 Differential expressed proteins (DEPs) of L. monocytogenes after treating with SAEW 18
5.6 GO and UniProtKB Keywords analysis of up/down regulated proteins 19
5.7 KEGG pathway analysis of L. monocytogenes DEPs exposed to SAEW 19
5.8 The chlorination protein of SAEW induced VBNC L. monocytogenes 20
Chapter 6 Discussion 22
Chapter 7 Conclusion 28
Chapter 8 References 29
Figures and Tables 38
Appendix 72

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