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研究生：	陳沭勝 Chen, Shu-Sheng
論文名稱：	臺灣侵襲性綠膿桿菌對碳氫黴烯類抗藥性之特性分析 Characteristics of carbapenem-resistance among invasive Pseudomonas aeruginosa in Taiwan
指導教授：	吳俊忠 Wu, Jiunn-Jong
學位類別：	碩士 Master
系所名稱：	醫學院 - 微生物及免疫學研究所 Department of Microbiology & Immunology
論文出版年：	2011
畢業學年度：	100
語文別：	中文
論文頁數：	116
中文關鍵詞：	綠膿桿菌、碳氫黴烯抗生素、OprD 、AmpC 、碳氫黴烯酶
外文關鍵詞：	Pseudomonas aeruginosa, carbapenem, OprD, AmpC, carbapenemase
相關次數：	點閱：57 下載：13
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綠膿桿菌（Pseudomonas aeruginosa）不只造成嚴重的公共衛生問題，其對於革蘭氏陰性菌感染最後一線抗生素的碳氫黴烯抗生素（carbapenems）感受性亦在減少。我們先前的研究發現VIM-2 變異型的VIM-3 是臺灣最常見的carbapenemase。本研究中，我們收集2000 年元月至2011 年2 月間，臺灣兩間醫學中心全數82 株自血流感染所分離出抗carbapenem 綠膿桿菌，以調查臺灣carbapenem 抗藥性綠膿桿菌的盛行率與特徵。以瓊脂稀釋法得知菌株對不同抗生素之感受性，帶有carbapenemase 的菌株以表現型檢驗與PCR基因篩選測定，CCCP 用於輸出幫浦抑制試驗，AmpC 大量表現與廣效性AmpC 活性的菌株以cloxacillin 抑制試驗調查，及imipenem 通道蛋白oprD 基因的定序與重組分析。Aztreonam、cefepime、ceftazidime、ciprofloxacin、gentamicin 及levofloxacin 抗藥性比率，各為53.7、61.0、57.3、47.6、42.7 及50.0%。共有3 株VIM-2、3 株VIM-3、2 株OXA-10 及2 株OXA-17 菌株；AmpC 大量表現與ESAC（extended-spectrum AmpC）菌株各有31.7%及11.0%；僅1 株為CCCP 輸出幫浦抑制試驗陽性；OprD 胺基酸多樣性、蛋白質截斷、蛋白質序列延長及插入序列，各為34.1%、32.9%、12.2%及6.1%。後天獲得的碳氫黴烯水解酶（11.0%），並非本研究主要抗藥機轉，OprD 功能性缺失可能為主要的抗藥機轉（81.7%），但ESAC 活性、AmpC 大量表現及CCCP 敏感的輸出幫浦亦參與臺灣綠膿桿菌對carbapenem 抗藥機轉。

Pseudomonas aeruginosa not only causes serious public health problems but also becomes less susceptibility to carbapenems, the last line of antimicrobial agents against multi-drug resistant Gram-negative bacterial infection. Our previous study has found that the acquired VIM-3 carbapenemase, a variant of VIM-2, was the most prevalent carbapenemase in P. aeruginosa in Taiwan. In this study, we investigated the prevalence and characteristics of carbapenem-resistant P. aeruginosa isolates in Taiwan. A total of 82 carbapenem-resistant P. aeruginosa isolates obtained from bloodstream infection were collected from two medical centers in Taiwan between 2000 January and 2011 February. The susceptibility of various antimicrobial agents was determined by agar dilution method. The phenotypic tests and genotypic PCR screening detected carbapenemase producers. Carbonyl cyanide m-chlorophenylhydrazone was used for the efflux pump inhibition test. AmpC overproducer and extended-spectrum AmpC activity were investigated by the cloxacillin inhibition assay. Gene sequencing and open reading frame alignment of imipenem channel porin gene, oprD, was analyzed. The resistance rate to aztreonam, cefepime, ceftazidime, ciprofloxacin, gentamicin, and levofloxacin were 53.7, 61.0, 57.3, 47.6, 42.7, and 50.0%, respectively. Overall, 2 VIM-2, 3 VIM-3, 2 OXA-10, and 2 OXA-17 isolates were determined, respectively. AmpC overproducer and ESAC (extended-spectrum AmpC) strain were 31.7% and 11.0%, respectively. Only one efflux pump inhibition assay positive strain was identified. Amino acid polymorphisms, truncation, elongation and insertion sequences in OprD were 34.1%, 32.9%, 12.2%, and 6.1%, respectively. Acquired carbapenem hydrolyzing enzymes (11.0%) are not the dominant carbapenem resistant mechanism in our strains. The functional loss of OprD is possible the major resistant mechanism (81.7%), but ESAC activity, AmpC overproduction and CCCP-sensitive efflux pumps also contribute to carbapenem resistance in P. aeruginosa in Taiwan.

中文摘要 ..................... i
英文摘要 .................... ii
誌謝 .................. iii
目錄 ................... iv
表目錄 ............ viii
圖目錄 ............... ix
符號與縮寫 .............. x
第一章 緒論 ............. 1
1-1 綠膿桿菌 ..................................... 1
1-2 抗生素 ............................................ 2
1-2-1 乙內醯胺類（β-lactams） ................ 2
1-2-2 氟喹諾酮類（Fluoroquinolones） ...................... 5
1-2-3 胺基醣苷類（Aminoglycosides） ............... 5
1-2-4 四環黴素類（Tetracyclines） ................ 6
1-2-5 福利黴素類（Rifamycins） .................. 6
1-2-6 巨內酯環類（Macrolides） ................... 6
1-2-7 脂肽類（Lipopeptides） .................... 7
1-3 抗藥基因 ............................................. 7
1-3-1 Class A β-lactamases ........................... 8
1-3-2 Class B β-lactamases ........................ 11
1-3-3 Class D β-lactamases .......................... 14
1-4 綠膿桿菌對carbapenem 的內生性抗藥機轉 ................ 17
1-4-1 Efflux pumps ...................................... 17
1-4-2 AmpC overproduction ........................ 19
1-4-3 Porin loss ............................ 19
1-4-4 Penicillin binding proteins（PBPs） .............. 20
1-5 研究目的 ............................... 21
第二章 材料與方法 .................................... 22
2-1 菌株來源、培養與保存 ....................... 22
2-2 抗生素感受性測試 ................................. 22
2-2-1 最小抑制濃度（minimum inhibitory concentration, MIC）測定 ................ 22
2-2-2 Modified Hodge test (MHT) ............... 23
2-2-3 2-MPA 雙紙錠協同試驗（2-mercaptopropionic acid double-disk synergy test） ....... 24
2-2-4 DPA 結合紙錠試驗（dipicolinic acid combined disk test, DPA-CDT） ..... 24
2-2-5 Efflux pump inhibition (EPI)assay ................. 25
2-2-6 Cloxacillin inhibitory assay ................. 25
2-3 DNA 實驗操作 ................................. 25
2-3-1 DNA 粗萃取 .......................... 25
2-3-2 聚合酶連鎖反應（Polymerase chain reaction, PCR)......25
2-3-3 DNA 定序（DNA sequencing） .............. 26
2-3-4 序列重組與分析（Sequence alignment and analysis） ............ 26
2-4 蛋白質實驗操作 .................................. 27
2-4-1 綠膿桿菌外膜蛋白質萃取（Extraction of P. aeruginosa outer membrane proteins） ................ 27
2-4-2 蛋白質濃度測定 .............................. 28
2-4-3 SDS-PAGE (sodium dodecyl sulfate polyacrylamide gel electrophoresis) .... 28
第三章 結果 .................... 29
3-1 綠膿桿菌之抗生素感受性測試 .................. 29
3-1-1 Antimicrobial agent susceptibilities ............. 29
3-1-2 Prevalence of carbapenem non-susceptible P. aeruginosa in NCKUH .......... 29
3-2 綠膿桿菌抗藥基因 ..................................... 30
3-2-1 Class A β-lactamases .................... 30
3-2-2 Class B β-lactamases ..................... 30
3-2-3 Class D β-lactamases ..................... 31
3-3 Efflux pump inhibition assay .................... 32
3-4 AmpC 大量表現與ESAC .................... 32
3-4-1 AmpC overproduction ....................... 32
3-4-2 ESAC characteristics ........................ 33
3-5 Porin profile 與OprD 蛋白質序列分析 .................. 33
3-5-1 oprD 基因的PCR 增幅 ...................... 33
3-5-2 OprD 蛋白的序列分析 ..................... 34
3-5-3 綠膿桿菌的outer membrane profile .......... 35
第四章 討論 ....................................... 36
4-1 臺灣與全球carbapenem 抗藥性綠膿桿菌之比較與感染控制 .......... 36
4-2 侵襲性carbapenem 抗藥性綠膿桿菌在臨床的威脅 ............................................. 37
4-3 帶有carbapenemase 的綠膿桿菌表現型檢定 ................................... 38
4-4 Efflux pump inhibition assay 藥物選擇與其他可能的臨床用途 ..... 40
4-5 AmpC 與carbapenem 抗藥性綠膿桿菌 .......... 41
4-6 OprD 與carbapenem 抗藥性綠膿桿菌 ...................... 42
4-7 回顧與展望 ........................................ 44
4-7-1 綠膿桿菌已知的carbapenem 抗藥機轉 ............... 44
4-7-2 綠膿桿菌其他可能的carbapenem 抗藥機轉 ................ 45
4-7-3 Insertion sequences 所造成的抗藥性問題 ............. 46
第五章 總結 ............................................. 48
參考文獻 .......... 49
表 ............. 83
圖 .................. 93
Appendix ........... 100
自述 ................................................ 116

表目錄
Table 1 . The primers used in this study ................ 83
Table 2 . Antimicrobial agent susceptibilities of 82 carbapenem non-susceptible P . aeruginosa strains from NCKUH and NTUH ...................................... 86
Table 3 . Carbapenem-resistant mechanisms and MICs of 82 carbapenem-non-susceptible P . aeruginosa clinical isolates in Taiwan .......................................... 87
Table 4 . Phenotypic screening of class A and B carbapenemases of carbapenem-nonsusceptible P . aeruginosa clinical isolates ................................ 89
Table 5 . Cloxacillin inhibition assay for AmpC overproduction and extended-spectrum AmpC β - lactamase activity .................... 90
Table 6. oprD inactive mutation in P. aeruginosa in Taiwan ............................................ 91
Table 7. Correlation of MIC and carbapenem-resistant mechanisms in P. aeruginosa ..... 92

圖目錄
Figure 1 . Prevalence of carbapenem non-susceptible Pseudomonas aeruginosa in National
Cheng Kung University Hospital from 2000 to February , 2011 ...................... 93
Figure 2 . The size of β - lactamase PCR products .......... 94
Figure 3. Modified Hodge test for class A carbapenemase detection .............................. 95
Figure 4 . Detection of metallo - β - lactamase-producer by 2-MPA synergy disk test and DPA-combined disk test ...... 96
Figure 5 . Genetic characterization of acquired β - lactamases in carbapenem non-susceptible P. aeruginosa clinical isolates in Taiwan ............... 97
Figure 6 . Outer membrane protein profile of 82 carbapenem-non-susceptible P . aeruginosa clinical isolates ..... 98
Figure 7 . Summary of carbapenem-resistant mechanisms in P. aeruginosa in this study 99
                                    

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