Ryan E T, Crean T I, John M, Butterton J R, Clements J D, Calderwood S B
Division of Infectious Diseases, Massachusetts General Hospital, Boston, Massachusetts 02114, USA.
Infect Immun. 1999 Apr;67(4):1694-701. doi: 10.1128/IAI.67.4.1694-1701.1999.
Vibrio cholerae secretes cholera toxin (CT) and the closely related heat-labile enterotoxin (LT) of Escherichia coli, the latter when expressed in V. cholerae. Both toxins are also potent immunoadjuvants. Mutant LT molecules that retain immunoadjuvant properties while possessing markedly diminished enterotoxic activities when expressed by E. coli have been developed. One such mutant LT molecule has the substitution of a glycine residue for arginine-192 [LT(R192G)]. Live attenuated strains of V. cholerae that have been used both as V. cholerae vaccines and as vectors for inducing mucosal and systemic immune responses directed against expressed heterologous antigens have been developed. In order to ascertain whether LT(R192G) can act as an immunoadjuvant when expressed in vivo by V. cholerae, we introduced a plasmid (pCS95) expressing this molecule into three vaccine strains of V. cholerae, Peru2, ETR3, and JRB14; the latter two strains contain genes encoding different heterologous antigens in the chromosome of the vaccine vectors. We found that LT(R192G) was expressed from pCS95 in vitro by both E. coli and V. cholerae strains but that LT(R192G) was detectable in the supernatant fraction of V. cholerae cultures only. In order to assess potential immunoadjuvanticity, groups of germfree mice were inoculated with the three V. cholerae vaccine strains alone and compared to groups inoculated with the V. cholerae vaccine strains supplemented with purified CT as an oral immunoadjuvant or V. cholerae vaccine strains expressing LT(R192G) from pCS95. We found that mice continued to pass stool containing V. cholerae strains with pCS95 for at least 4 days after oral inoculation, the last day evaluated. We found that inoculation with V. cholerae vaccine strains containing pCS95 resulted in anti-LT(R192G) immune responses, confirming in vivo expression. We were unable to detect immune responses directed against the heterologous antigens expressed at low levels in any group of animals, including animals that received purified CT as an immunoadjuvant. We were, however, able to measure increased vibriocidal immune responses against vaccine strains in animals that received V. cholerae vaccine strains expressing LT(R192G) from pCS95 compared to the responses in animals that received V. cholerae vaccine strains alone. These results demonstrate that mutant LT molecules can be expressed in vivo by attenuated vaccine strains of V. cholerae and that such expression can result in an immunoadjuvant effect.
霍乱弧菌分泌霍乱毒素(CT)以及大肠杆菌的密切相关的不耐热肠毒素(LT),后者在霍乱弧菌中表达时也会分泌。这两种毒素也是有效的免疫佐剂。已经开发出了突变型LT分子,当在大肠杆菌中表达时,它们保留免疫佐剂特性,同时具有明显降低的肠毒性活性。一种这样的突变型LT分子是用甘氨酸残基取代精氨酸-192 [LT(R192G)]。已经开发出了霍乱弧菌减毒活菌株,它们既用作霍乱弧菌疫苗,也用作诱导针对表达的异源抗原的黏膜和全身免疫反应的载体。为了确定LT(R192G)在霍乱弧菌体内表达时是否能作为免疫佐剂,我们将表达该分子的质粒(pCS95)导入三种霍乱弧菌疫苗菌株,即秘鲁2株、ETR3株和JRB14株;后两种菌株在疫苗载体的染色体中含有编码不同异源抗原的基因。我们发现,LT(R192G)在体外可由大肠杆菌和霍乱弧菌菌株从pCS95中表达,但仅在霍乱弧菌培养物的上清液部分中可检测到LT(R192G)。为了评估潜在的免疫佐剂活性,将无菌小鼠组分别接种三种霍乱弧菌疫苗菌株,并与接种补充有纯化CT作为口服免疫佐剂的霍乱弧菌疫苗菌株或从pCS95中表达LT(R192G)的霍乱弧菌疫苗菌株的组进行比较。我们发现,口服接种后,小鼠至少在评估的最后一天持续排出含有携带pCS95的霍乱弧菌菌株的粪便达4天。我们发现,接种含有pCS95的霍乱弧菌疫苗菌株会引发抗LT(R192G)免疫反应,证实了其在体内的表达。我们无法在任何一组动物中检测到针对低水平表达的异源抗原的免疫反应,包括接受纯化CT作为免疫佐剂的动物。然而,与仅接受霍乱弧菌疫苗菌株的动物相比,我们能够测量到接受从pCS95中表达LT(R192G)的霍乱弧菌疫苗菌株的动物对疫苗菌株的杀弧菌免疫反应增强。这些结果表明,突变型LT分子可由霍乱弧菌减毒疫苗菌株在体内表达,且这种表达可产生免疫佐剂效应。