臺灣博碩士論文加值系統

巴東體屬細菌 (Bartonella spp.)為培養與生長條件均具挑剔性的革蘭氏陰性桿菌，其自然宿主以哺乳類動物為主且涵蓋種類廣泛，此細菌屬中已有許多細菌種為人畜共通病原，瞭解地區性不同的感染動物宿主流行病學狀況，對防治此類病原對人類之威脅具有重要性。本研究於2014年12月至2017年9月期間，在台灣8個城市的16個地點採集了4個不同目別動物，總計184個血液檢體樣本，進行細菌培養及分子檢測。所有培養及分子檢測陽性之檢體依巴東體屬菌特定之gltA基因進行PCR。結果顯示，巴東體屬細菌在7種動物中檢測到，包括30.2％ (19/63) 的鬼鼠(Bandicota indica)，100％ (1/1) 的田鼷鼠(Mus caroli)，35.7％ (5/14) 的小黃腹鼠（Rattus losea），50％ (1/2) 的家鼠(Rattus rattus)，50％（1/2）的錢鼠（Suncus murinus），5.3％（3/57）的食蟹獴(Herpestes urva)和18.2％（2/11）的台灣獼猴(Macaca cyclopis)。PCR陽性樣品進加一步使用gltA基因片段定序之部份序列結果以親緣樹進行分析，共鑑定出八種巴東體屬菌種，其中包括七種過去已知的巴東體菌種(B. coopersplainsensis, B. elizabethae, B. henselae, B. queenslandensis, B. quintana, B. rattimassiliensis and B. tribocourm)和一種潛在的新型巴東體菌種(B. phoceensis-like)，此結果顯示了巴東體屬細菌在台灣具有多樣性。依據血液培養分離陽性檢體進一步於培養基上隨機挑選菌落進行分析之結果，發現3.4％ (3/86) 的小型哺乳動物具有不同巴東體屬細菌之混合感染。本研究也進行兩屬新興病媒傳播病原，即無形體屬菌(Anaplasma spp.)和伯氏疏螺旋體(Borrelia spp.)的調查分析，方法上為以常規或巢式PCR檢測與序列分析。結果顯示40.7％（35/86）的小型哺乳動物感染了A. bovis，18.4％ (16/87）的肉食性動物也有無形體屬菌感染（當中包括十五個A. bovis和一個A. platys）。然而，PCR結果顯示，本研究中並未發現任何檢測動物感染伯氏疏螺旋體。此外，進一步交叉分析同一動物個體中感染不同病原之結果，發現在所有的動物樣本中，8.7％共感染巴東體屬菌及無形體屬菌。
本研究的調查有助於瞭解台灣野生哺乳動物感染三個新興人畜共通病原細菌屬之節肢動物病媒傳染病流行病學情況，研究結果可為未來預防和控制這些傳染病之重要資訊。本研究為國際首次由食蟹獴(Herpestes urva)和台灣獼猴(Macaca cyclopis)中分離到人畜共通巴東體屬細菌，為重要研究成果。

Bartonella spp. are fastidious, gram-negative, rod-shaped bacteria that are highly adapted to a wide range of mammalian hosts. In this study, a total of 184 blood samples from 4 animal orders were collected at 16 locations among eight cities in Taiwan during December 2014 to September 2017. Based on the PCR analysis of the Bartonella-specific gltA gene using the cultured isolates and PCR-positive samples, Bartonellae were detected in seven animal species, including 30.2% (19/63) of Bandicota indica, 100% (1/1) of Mus caroli, 35.7% (5/14) of Rattus losea, 50% (1/2) of Rattus rattus, 50% (1/2) of Suncus murinus, 5.3% (3/57) of Herpestes urva and 18.2% (2/11) of Macaca cyclopis. Through phylogenetic analysis of Bartonella PCR-positive samples using the partial sequence of the gltA gene, eight Bartonella spp. were identified, including seven known Bartonella species (B. coopersplainsensis, B. elizabethae, B. henselae, B. queenslandensis, B. quintana, B. rattimassiliensis and B. tribocourm) and one potentially novel Bartonella species (B. phoceensis-like); it showed the large diversity of Bartonella spp. in Taiwan. By direct blood culture results, 3.4% (3/86) of the small mammals were co-infected with different Bartonella spp. To study other emerging vector-borne agents, such as Anaplasmataceae and Borrelia burgdorferi, conventional or nested polymerase chain reaction assays and sequence analysis were further conducted. The results show that 40.7% (35/86) of the small mammals were infected with A. bovis, and 18.4% (16/87) of the carnivores were with Anaplasma spp. (including 15 A. bovis and one A. platys). Furthermore, 8.7% of the animals positive for Anaplasma were co-infected with Bartonella. However, none of the animals showed PCR positivity for B. burgdorferi. The investigation in this study could help to understanding the epidemiology of these infections in wild mammals in Taiwan and provide a groundwork to prevent and control these vector-borne diseases. To the best of our knowledge, this is the first report on isolation of Bartonella from Herpestes urva and Macaca cyclopis worldwide.

摘要 i
Abstract iii
Contents v
List of Tables viii
List of Figures ix
Chapter 1 Literature review 1
1. 1. Characteristics of Bartonella spp. and main reservoir hosts 1
1.1.1 Characteristic and classification 1
1.1.2 Bartonella in small mammals 2
1.1.3 Bartonella in carnivores 4
1.1.4 Bartonella in non-human Primates 5
1.2. Characteristics of Anaplasma spp. and main reservoir hosts 6
1.2.1 Characteristic and classification 6
1.2.2 Main animal reservoirs 7
1.3. Characteristics of Borrelia burgdorferi and main reservoir hosts 8
1.3.1 Classification 8
1.3.2 Main animal reservoirs 9
1.4. Objectives of this study 11
Chapter 2 Materials and Methods 12
2.1. Animal collection and capture sites 12
2.2. Blood culture 13
2.3. DNA extraction 13
2.3.1 From bacterial colonies 13
2.3.2 From whole blood 14
2.4. Identification of coinfection of Bartonella spp. in the animals 14
2.5. Polymerase chain reaction 15
2.5.1 Detection of Bartonella spp. by the citrate synthase (gltA) gene 15
2.5.2 Detection of Borrelia spp. by the 5S-23S interspacer region 16
2.5.3 Detection of Ehrlichia spp. by the 16S rRNA gene 17
2.5.4 Detection of Anaplasmataceae by the 16S rRNA gene 18
2.6. Sequencing and phylogenetic analysis 19
2.7. Statistical analysis 20
Chapter 3 Results 21
3.1. Animal samples 21
3.2. Bartonella infection 22
3.2.1 Isolation of Bartonella using whole blood culture 22
3.2.2 Molecular detection of Bartonella using PCR assay 23
3.2.3 Co-infection of Bartonella spp.. 24
3.2.4 Sequence comparison and phylogenetic analyses 24
3.2.5 Understanding the host range of Bartonella spp. using the sequences identified in this study to compare the information collected all over the world 26
3.3. Anaplasma infection 28
3.3.1 PCR detection of Anaplasmataceae bacteria in animals 28
3.3.2 Sequence comparison and phylogenetic analyses 28
3.3.3 Understanding the host range of Anaplasma spp. using the sequences identified in this study to compare the information collected all over the world 29
3.4 Borrelia burgdorferi 30
3.4.1 Prevalence of Borrelia burgdorferi using PCR detection in wild animals 30
3.5. Co-infection of multiple vector-borne pathogens 30
3.6. Epidemiological analysis for factors associated with Bartonella, Anaplasma and Borrelia infections 31
Chapter 4 Discussions 32
4.1 Bartonella infection 32
4.1.1 Small mammals 32
4.1.2 Carnivore animals 34
4.1.3 Non-human primates 37
4.2 Anaplasma infection 39
4.3 Borrelia burgdorferi infection 41
4.4 Co-infection 42
4.4.1 Co-infection of different Bartonella spp. 42
4.4.2. Co-infection of multiple vector-borne pathogens 43
4.5 Statistical analysis 44
References 72

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