Ramarao Nalini, Nielsen-Leroux Christina, Lereclus Didier
INRA, Micalis UMR1319, France.
J Vis Exp. 2012 Dec 11(70):e4392. doi: 10.3791/4392.
The study of bacterial virulence often requires a suitable animal model. Mammalian models of infection are costly and may raise ethical issues. The use of insects as infection models provides a valuable alternative. Compared to other non-vertebrate model hosts such as nematodes, insects have a relatively advanced system of antimicrobial defenses and are thus more likely to produce information relevant to the mammalian infection process. Like mammals, insects possess a complex innate immune system(1). Cells in the hemolymph are capable of phagocytosing or encapsulating microbial invaders, and humoral responses include the inducible production of lysozyme and small antibacterial peptides(2,3). In addition, analogies are found between the epithelial cells of insect larval midguts and intestinal cells of mammalian digestive systems. Finally, several basic components essential for the bacterial infection process such as cell adhesion, resistance to antimicrobial peptides, tissue degradation and adaptation to oxidative stress are likely to be important in both insects and mammals(1). Thus, insects are polyvalent tools for the identification and characterization of microbial virulence factors involved in mammalian infections. Larvae of the greater wax moth Galleria mellonella have been shown to provide a useful insight into the pathogenesis of a wide range of microbial infections including mammalian fungal (Fusarium oxysporum, Aspergillus fumigatus, Candida albicans) and bacterial pathogens, such as Staphylococcus aureus, Proteus vulgaris, Serratia marcescens Pseudomonas aeruginosa, Listeria monocytogenes or Enterococcus faecalis(4-7). Regardless of the bacterial species, results obtained with Galleria larvae infected by direct injection through the cuticle consistently correlate with those of similar mammalian studies: bacterial strains that are attenuated in mammalian models demonstrate lower virulence in Galleria, and strains causing severe human infections are also highly virulent in the Galleria model(8-11). Oral infection of Galleria is much less used and additional compounds, like specific toxins, are needed to reach mortality. G. mellonella larvae present several technical advantages: they are relatively large (last instar larvae before pupation are about 2 cm long and weight 250 mg), thus enabling the injection of defined doses of bacteria; they can be reared at various temperatures (20 °C to 30 °C) and infection studies can be conducted between 15 °C to above 37 °C(12,13), allowing experiments that mimic a mammalian environment. In addition, insect rearing is easy and relatively cheap. Infection of the larvae allows monitoring bacterial virulence by several means, including calculation of LD50(14), measurement of bacterial survival(15,16) and examination of the infection process(17). Here, we describe the rearing of the insects, covering all life stages of G. mellonella. We provide a detailed protocol of infection by two routes of inoculation: oral and intra haemocoelic. The bacterial model used in this protocol is Bacillus cereus, a Gram positive pathogen implicated in gastrointestinal as well as in other severe local or systemic opportunistic infections(18,19).
对细菌毒力的研究通常需要合适的动物模型。感染的哺乳动物模型成本高昂,且可能引发伦理问题。使用昆虫作为感染模型提供了一种有价值的替代方法。与线虫等其他非脊椎动物模型宿主相比,昆虫具有相对先进的抗菌防御系统,因此更有可能产生与哺乳动物感染过程相关的信息。与哺乳动物一样,昆虫拥有复杂的先天免疫系统(1)。血淋巴中的细胞能够吞噬或包裹微生物入侵者,体液反应包括诱导产生溶菌酶和小抗菌肽(2,3)。此外,在昆虫幼虫中肠上皮细胞与哺乳动物消化系统的肠道细胞之间存在相似之处。最后,细菌感染过程中几个基本组成部分,如细胞黏附、对抗菌肽的抗性、组织降解和对氧化应激的适应,在昆虫和哺乳动物中可能都很重要(1)。因此,昆虫是用于鉴定和表征参与哺乳动物感染的微生物毒力因子的多用途工具。大蜡螟幼虫已被证明能为包括哺乳动物真菌(尖孢镰刀菌、烟曲霉、白色念珠菌)和细菌病原体,如金黄色葡萄球菌、普通变形杆菌、粘质沙雷氏菌、铜绿假单胞菌、单核细胞增生李斯特菌或粪肠球菌等多种微生物感染的发病机制提供有用的见解(4 - 7)。无论细菌种类如何,通过角质层直接注射感染大蜡螟幼虫所获得的结果始终与类似的哺乳动物研究结果相关:在哺乳动物模型中减毒的细菌菌株在大蜡螟中显示出较低的毒力,而导致严重人类感染的菌株在大蜡螟模型中也具有高毒力(8 - 11)。大蜡螟的口服感染较少使用,且需要额外的化合物,如特定毒素,才能导致死亡。大蜡螟幼虫具有几个技术优势:它们相对较大(化蛹前的末龄幼虫约2厘米长,重250毫克),因此能够注射确定剂量的细菌;它们可以在不同温度(20°C至30°C)下饲养,感染研究可以在15°C至37°C以上进行(12,13),从而允许进行模拟哺乳动物环境的实验。此外,昆虫饲养容易且成本相对较低。幼虫感染允许通过多种方式监测细菌毒力,包括计算半数致死剂量(14)、测量细菌存活率(15,16)以及检查感染过程(17)。在此,我们描述昆虫的饲养,涵盖大蜡螟的所有生命阶段。我们提供了两种接种途径感染的详细方案:口服和血腔注射。本方案中使用的细菌模型是蜡样芽孢杆菌,一种革兰氏阳性病原体,与胃肠道以及其他严重的局部或全身机会性感染有关(18,19)。