嗜水气单胞菌非质粒依赖性菌蜕疫苗的研制及评价
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摘要
嗜水气单胞菌在自然界中分布广泛,由致病性嗜水气单胞菌引起的疾病在世界海、淡水养殖中也普遍存在,给养殖生产带来了巨大损失,如何对其所致的各类疾病进行有效控制一直为研究者所重视。
菌蜕是革兰氏阴性菌被噬菌体phiX174的裂解基因E裂解后所形成的不含胞质内含物的完整细菌空壳。由于保留了细菌原有的表面结构和特性,因此是一种比常规疫苗更理想的新型疫苗体系。当前菌蜕疫苗的制备均是通过转化携带有裂解基因E及其控制单元的质粒并诱导其表达而完成,存在潜在抗性基因扩散及由于残存活菌所致的安全问题。
为解决以上问题,本研究通过将裂解基因E及其控制单元整合进嗜水气单胞菌染色体的aroA基因内,构建出兼具减毒特性的菌蜕疫苗制备候选突变株,具有更高的安全性。
1.嗜水气单胞菌疫苗候选菌株的验证
通过形态学观察、选择性培养基筛选、生理生化分析及16SrDNA和gyrB序列测定等方法对本室保存的一株分离自患出血性败血症鲫鱼的细菌J-1进行验证,证明该细菌为嗜水气单胞菌。人工感染实验显示该细菌具有致病性,对异育银鲫的半数致死量为1.2×106CFU。
2.基于pR/pL分别启动裂解基因E和核酸酶A基因共表达载体的构建菌蜕是一种新的疫苗平台,它的安全高效生产主要依赖于合适的功能载体。本研究中,我们构建了一系列含phiX裂解基因E和/或核酸酶A的温控质粒并在大肠杆菌中进行了评价。结果表明,核酸酶A的直接产物(pBV220-SNA可以降解质粒及大肠杆菌的基因组,而基因E和部分Cro基因的融合产物(pKF396M-2)则不能裂解宿主菌。通过在两者间插入增强子元件T7g10和SD盒(pKF396M-3)可以恢复基因E的功能。利用基于pR/pL分别启动裂解基因E和核酸酶A基因的共表达载体pKF396M-4,残留质粒及大肠杆菌基因组被清除,灭活效率较单纯使用E介导的裂解质粒要高2个数量级。将基因E和SNA用设计合成的多克隆位点替换即构建成具有两个多克隆位点的双表达载体pKF396M-5。
3.嗜水气单胞菌aroA无标记缺失突变株的构建及其特性
克隆嗜水气单胞菌J-1株aroA基因及其上下游序列,最终测序拼接出长度为3704bp的核苷酸序列,该序列与嗜水气单胞菌ATCC7966同源性相似度最高,达97%。在此基础上重新设计合成两对引物分别扩增出aroA基因上下游1200bp左右的序列,酶切后连入经多克隆位点置换改造的自杀质粒pDS132,构建出含缺失1103bp aroA基因的重组自杀质粒,将其接合转导至嗜水气单胞菌J-1,通过2次同源重组及PCR鉴定筛选出aroA无抗性标记缺失株,该嗜水气单胞菌aroA缺失株能在LB固体培养基、选择性培养基Rimler-Shoots琼脂上和鉴别性培养基AHM琼脂平板上正常生长,呈现的形态特征也与J-1株类似,但在LB液体培养基中的生长速度则较J-1为慢。在所测试的15项生理生化反应指标中,仅VP反应指标一项与与J-1株不同,为阴性,胞外蛋白酶活性检测呈阳性,对异育银鲫的LD50为2.8×108CFU。
4.基于染色体裂解基因E介导的嗜水气单胞菌菌蜕疫苗的制备
利用同源重组技术将裂解基因E及其控制单元整合进嗜水气单胞菌染色体的aroA基因内,并通过升温诱导裂解基因E表达成功制备出非质粒依赖性菌蜕疫苗。溶菌动力学试验显示,在诱导后的1h内,其OD600快速上升,但2h后OD600开始持续下降,6h后开始趋于平稳,此时培养物活菌数从诱导前的6.4×107CFU/mL下降至1.7×102CFU/mL,下降了105倍。扫描电镜观察可见绝大部分菌体经诱导后形成菌蜕,跨膜通道位于细胞中央或两端。
5.嗜水气单胞菌菌蜕疫苗对异育银鲫的免疫保护
于27.2℃～29.8℃水温下,分别用菌蜕疫苗(AHG)、福尔马林全菌灭活疫苗(FKC)和PBS通过腹腔注射免疫异育银鲫,比较其免疫效果。结果显示,AHG免疫组血清凝集指数(1：277.3)高于FKC免疫组(1：170.7)和PBS免疫组(1：9.6),对同源菌也显示出了更高的杀菌活性,但溶菌酶活力与FKC免疫组相当,均为0.089,显著高于空白组的0.032。同源菌攻击试验显示,AHG、FKC和PBS组的存活率分别为90%,65%和15%。这一切表明,AHG疫苗能诱导更为强烈的免疫反应,提高免疫保护能力。
Aeromonas hydrophila is widely distributed in the world.The diseases caused by the pathogenic strains of this organism have also prevailed in seawater or freshwater culture and brought about a great loss to aquaculture production, which arose great concern about how to control these diseases effectively among scientific researchers.
Bacterial ghosts are empty and intact bacterial envelopes of Gram-negative bacteria,which are produced by controlled expression of phage phiX174lysis gene E and devoid of cytoplasmic content. As the surface structure and properties of their original state remained, it is a more ideal vaccine system than conventional vaccines. Current bacterial ghosts production depends on the transformation and induction of plasmids with lysis gene E and its regulatory elements, which brings in the potential risk of antibiotic resistance gene spread and safety problem from the remanent inactivated cells during the ghost preparation.
To solve this problem, the lysis gene E and its regulatory elements (cI857/pR/E/rrnb T1T2) were integrated into the aroA gene of A.hydrophila in the present study and the constructed mutants could be used as a vaccine candidate for bacterial ghost production with less virulence and higher safety.
1. Identification of Aeromonas hydrophila Strain as Vaccine Candidate
A bacterial strain J-1, which was isolated from the moribund crucian carp with the symptom of hemorrhagic septicemia, was identified as aeromonas hydrophila by morpholgy, selective medium,biochemical tests, sequence comparison of16S rDNA and gyrB. Artificial infection experiment showed that the isolate was virulent to allogynogenetic sliver crucian carp with an LD50of1.2X106CFU.
2. A Novel Dual Vector Coexpressing PhiX174Lysis E Gene and Staphylococcal Nuclease A Gene on the basis of Lambda Promoter pR and pL, Respectively
Bacterial Ghost is a novel vaccine platform and its safe and efficient production depends largely upon a suitable and functional vector. In this study, a series of temperature-inducible plasmids, carrying phix174lysis gene E and/or staphylococcal nuclease A (SNA) gene, were constructed and evaluated in Escherichia coli. The results showed that the direct product of SNA (pBV220-SNA) could degrade the plasmid and genomic DNA of E.coli while the fusion product of gene E and partial Cro gene (pKF396M-2) lost the ability to lyse the host strain. The insertion of enhancer T7g10elements and Shine-Dalgarno box (ESD) between them (pKF396M-3) could resume the function of gene E. Using plasmid pKF396M-4with gene E and SNA respectively under the immediate control of promoter pR and pL, the remnant plasmids and genomic DNA of E.coli were eliminated and the rates of inactivation increased by2orders of magnitude over that obtained with the exclusive use of E-mediated lysis plasmid. By substituting these two genes with customized multiple cloning sites (MCS) sequences, the plasmid could be modified to a dual expression vector (pKF396M-5).
3. Construction and Characterization of Aeromonas hydrophila aroA-Non-marked Deletion Mutant
The aroA gene and its flanked region of aeromonas hydrophila J-l was cloned, and the sequences was determined. The nucleotide sequence of3704bp showed a high degree of identity of97%with the aeromonas hydrophila ATCC7966. On the basis of this the upstream and downstream region of aroA was amplified respectively with redesigned and synthesized primers, and the PCR products were cut with restriction endonuclease and ligated to the corresponding sites of suicide vector pDS132with the MCS modification, therefore constructing a recombinant suicide vector with the1103bp deletion aroA gene. This suicide vector was transformed to aeromonas hydrophila J-l by conjunction and the non-marked aroA deletion mutant which has experienced two homologous recombination was selected by PCR identification.The aroA deletion mutant could normally grow on LB, Rimler-Shoots selective medium and differential medium AHM, but grew slowly in the LB broth than J-l. Among the15kinds of biochemical and physiological tests, only the result of the item VP test was different with J-l (negative). The ECP activity test was positive and the LD50to allogynogenetic sliver crucian carp was2.8×108CFU.
4. Aeromonas hydrophila Ghosts Production by Expression of Chromosomally-Based Lysis Gene E
Using recombinant DNA technology, the lysis gene E and its regulatory elements (cI857-pRM/pR/-E-rrnbT1T2) were intergrated into the aroA gene of A.hydrophila J-l and the plasmid-independent bacterial ghosts were produced sussessfully by induction of temperature upshift from28℃to42℃. The lysis kinetics showed that the OD600value of culture media increased rapidly one hour after induction and then began to decrease gradually two hours later and finally came to a steady state at the time of six hour when the number of the viable cells declined by5orders of magnitude, from6.4X107CFU/ml to1.7×102CFU/ml. Scanning electron microscopy observation proved that most lysed bacteria shrinked and the transmembrane tunnel located either in the middle of the cells or at the polar sites.
5. Evaluaion of Induced Immune by Aeromonas hydrophila Ghosts in silver carps, Carassius auratus gibelio
At temperature of27.2℃～29.8℃in water, the immunisity induced by aeromonas hydrophila ghosts (AHG), formalin-killed A.hydrophila (FKC) and PBS were compared in silver carps (Carassius auratus gibelio) by intraperitoneal immunization. The results showed that the serum agglutination titers induced by AHG group (1:277.3) was highest compared to FKC (1:170.7) and PBS control (1:9.6). AHG also induced higher bactericidal activity than either FKC or control, while in lysozyme activity, AHG had similar effect to FKC (0.089),only showing significantly higher level than control (0.032). In challenge test, the AHG-immunized fishes had best protection against A.hydrophila (90%) compared to65%and15%survival in FKC and PBS control, respectively.These findings indicated that AHG could induce stronger immune response and could confer enhanced protection against infection.
菌蜕是革兰氏阴性菌被噬菌体phiX174的裂解基因E裂解后所形成的不含胞质内含物的完整细菌空壳。由于保留了细菌原有的表面结构和特性,因此是一种比常规疫苗更理想的新型疫苗体系。当前菌蜕疫苗的制备均是通过转化携带有裂解基因E及其控制单元的质粒并诱导其表达而完成,存在潜在抗性基因扩散及由于残存活菌所致的安全问题。
为解决以上问题,本研究通过将裂解基因E及其控制单元整合进嗜水气单胞菌染色体的aroA基因内,构建出兼具减毒特性的菌蜕疫苗制备候选突变株,具有更高的安全性。
1.嗜水气单胞菌疫苗候选菌株的验证
通过形态学观察、选择性培养基筛选、生理生化分析及16SrDNA和gyrB序列测定等方法对本室保存的一株分离自患出血性败血症鲫鱼的细菌J-1进行验证,证明该细菌为嗜水气单胞菌。人工感染实验显示该细菌具有致病性,对异育银鲫的半数致死量为1.2×106CFU。
2.基于pR/pL分别启动裂解基因E和核酸酶A基因共表达载体的构建菌蜕是一种新的疫苗平台,它的安全高效生产主要依赖于合适的功能载体。本研究中,我们构建了一系列含phiX裂解基因E和/或核酸酶A的温控质粒并在大肠杆菌中进行了评价。结果表明,核酸酶A的直接产物(pBV220-SNA可以降解质粒及大肠杆菌的基因组,而基因E和部分Cro基因的融合产物(pKF396M-2)则不能裂解宿主菌。通过在两者间插入增强子元件T7g10和SD盒(pKF396M-3)可以恢复基因E的功能。利用基于pR/pL分别启动裂解基因E和核酸酶A基因的共表达载体pKF396M-4,残留质粒及大肠杆菌基因组被清除,灭活效率较单纯使用E介导的裂解质粒要高2个数量级。将基因E和SNA用设计合成的多克隆位点替换即构建成具有两个多克隆位点的双表达载体pKF396M-5。
3.嗜水气单胞菌aroA无标记缺失突变株的构建及其特性
克隆嗜水气单胞菌J-1株aroA基因及其上下游序列,最终测序拼接出长度为3704bp的核苷酸序列,该序列与嗜水气单胞菌ATCC7966同源性相似度最高,达97%。在此基础上重新设计合成两对引物分别扩增出aroA基因上下游1200bp左右的序列,酶切后连入经多克隆位点置换改造的自杀质粒pDS132,构建出含缺失1103bp aroA基因的重组自杀质粒,将其接合转导至嗜水气单胞菌J-1,通过2次同源重组及PCR鉴定筛选出aroA无抗性标记缺失株,该嗜水气单胞菌aroA缺失株能在LB固体培养基、选择性培养基Rimler-Shoots琼脂上和鉴别性培养基AHM琼脂平板上正常生长,呈现的形态特征也与J-1株类似,但在LB液体培养基中的生长速度则较J-1为慢。在所测试的15项生理生化反应指标中,仅VP反应指标一项与与J-1株不同,为阴性,胞外蛋白酶活性检测呈阳性,对异育银鲫的LD50为2.8×108CFU。
4.基于染色体裂解基因E介导的嗜水气单胞菌菌蜕疫苗的制备
利用同源重组技术将裂解基因E及其控制单元整合进嗜水气单胞菌染色体的aroA基因内,并通过升温诱导裂解基因E表达成功制备出非质粒依赖性菌蜕疫苗。溶菌动力学试验显示,在诱导后的1h内,其OD600快速上升,但2h后OD600开始持续下降,6h后开始趋于平稳,此时培养物活菌数从诱导前的6.4×107CFU/mL下降至1.7×102CFU/mL,下降了105倍。扫描电镜观察可见绝大部分菌体经诱导后形成菌蜕,跨膜通道位于细胞中央或两端。
5.嗜水气单胞菌菌蜕疫苗对异育银鲫的免疫保护
于27.2℃～29.8℃水温下,分别用菌蜕疫苗(AHG)、福尔马林全菌灭活疫苗(FKC)和PBS通过腹腔注射免疫异育银鲫,比较其免疫效果。结果显示,AHG免疫组血清凝集指数(1：277.3)高于FKC免疫组(1：170.7)和PBS免疫组(1：9.6),对同源菌也显示出了更高的杀菌活性,但溶菌酶活力与FKC免疫组相当,均为0.089,显著高于空白组的0.032。同源菌攻击试验显示,AHG、FKC和PBS组的存活率分别为90%,65%和15%。这一切表明,AHG疫苗能诱导更为强烈的免疫反应,提高免疫保护能力。
Aeromonas hydrophila is widely distributed in the world.The diseases caused by the pathogenic strains of this organism have also prevailed in seawater or freshwater culture and brought about a great loss to aquaculture production, which arose great concern about how to control these diseases effectively among scientific researchers.
Bacterial ghosts are empty and intact bacterial envelopes of Gram-negative bacteria,which are produced by controlled expression of phage phiX174lysis gene E and devoid of cytoplasmic content. As the surface structure and properties of their original state remained, it is a more ideal vaccine system than conventional vaccines. Current bacterial ghosts production depends on the transformation and induction of plasmids with lysis gene E and its regulatory elements, which brings in the potential risk of antibiotic resistance gene spread and safety problem from the remanent inactivated cells during the ghost preparation.
To solve this problem, the lysis gene E and its regulatory elements (cI857/pR/E/rrnb T1T2) were integrated into the aroA gene of A.hydrophila in the present study and the constructed mutants could be used as a vaccine candidate for bacterial ghost production with less virulence and higher safety.
1. Identification of Aeromonas hydrophila Strain as Vaccine Candidate
A bacterial strain J-1, which was isolated from the moribund crucian carp with the symptom of hemorrhagic septicemia, was identified as aeromonas hydrophila by morpholgy, selective medium,biochemical tests, sequence comparison of16S rDNA and gyrB. Artificial infection experiment showed that the isolate was virulent to allogynogenetic sliver crucian carp with an LD50of1.2X106CFU.
2. A Novel Dual Vector Coexpressing PhiX174Lysis E Gene and Staphylococcal Nuclease A Gene on the basis of Lambda Promoter pR and pL, Respectively
Bacterial Ghost is a novel vaccine platform and its safe and efficient production depends largely upon a suitable and functional vector. In this study, a series of temperature-inducible plasmids, carrying phix174lysis gene E and/or staphylococcal nuclease A (SNA) gene, were constructed and evaluated in Escherichia coli. The results showed that the direct product of SNA (pBV220-SNA) could degrade the plasmid and genomic DNA of E.coli while the fusion product of gene E and partial Cro gene (pKF396M-2) lost the ability to lyse the host strain. The insertion of enhancer T7g10elements and Shine-Dalgarno box (ESD) between them (pKF396M-3) could resume the function of gene E. Using plasmid pKF396M-4with gene E and SNA respectively under the immediate control of promoter pR and pL, the remnant plasmids and genomic DNA of E.coli were eliminated and the rates of inactivation increased by2orders of magnitude over that obtained with the exclusive use of E-mediated lysis plasmid. By substituting these two genes with customized multiple cloning sites (MCS) sequences, the plasmid could be modified to a dual expression vector (pKF396M-5).
3. Construction and Characterization of Aeromonas hydrophila aroA-Non-marked Deletion Mutant
The aroA gene and its flanked region of aeromonas hydrophila J-l was cloned, and the sequences was determined. The nucleotide sequence of3704bp showed a high degree of identity of97%with the aeromonas hydrophila ATCC7966. On the basis of this the upstream and downstream region of aroA was amplified respectively with redesigned and synthesized primers, and the PCR products were cut with restriction endonuclease and ligated to the corresponding sites of suicide vector pDS132with the MCS modification, therefore constructing a recombinant suicide vector with the1103bp deletion aroA gene. This suicide vector was transformed to aeromonas hydrophila J-l by conjunction and the non-marked aroA deletion mutant which has experienced two homologous recombination was selected by PCR identification.The aroA deletion mutant could normally grow on LB, Rimler-Shoots selective medium and differential medium AHM, but grew slowly in the LB broth than J-l. Among the15kinds of biochemical and physiological tests, only the result of the item VP test was different with J-l (negative). The ECP activity test was positive and the LD50to allogynogenetic sliver crucian carp was2.8×108CFU.
4. Aeromonas hydrophila Ghosts Production by Expression of Chromosomally-Based Lysis Gene E
Using recombinant DNA technology, the lysis gene E and its regulatory elements (cI857-pRM/pR/-E-rrnbT1T2) were intergrated into the aroA gene of A.hydrophila J-l and the plasmid-independent bacterial ghosts were produced sussessfully by induction of temperature upshift from28℃to42℃. The lysis kinetics showed that the OD600value of culture media increased rapidly one hour after induction and then began to decrease gradually two hours later and finally came to a steady state at the time of six hour when the number of the viable cells declined by5orders of magnitude, from6.4X107CFU/ml to1.7×102CFU/ml. Scanning electron microscopy observation proved that most lysed bacteria shrinked and the transmembrane tunnel located either in the middle of the cells or at the polar sites.
5. Evaluaion of Induced Immune by Aeromonas hydrophila Ghosts in silver carps, Carassius auratus gibelio
At temperature of27.2℃～29.8℃in water, the immunisity induced by aeromonas hydrophila ghosts (AHG), formalin-killed A.hydrophila (FKC) and PBS were compared in silver carps (Carassius auratus gibelio) by intraperitoneal immunization. The results showed that the serum agglutination titers induced by AHG group (1:277.3) was highest compared to FKC (1:170.7) and PBS control (1:9.6). AHG also induced higher bactericidal activity than either FKC or control, while in lysozyme activity, AHG had similar effect to FKC (0.089),only showing significantly higher level than control (0.032). In challenge test, the AHG-immunized fishes had best protection against A.hydrophila (90%) compared to65%and15%survival in FKC and PBS control, respectively.These findings indicated that AHG could induce stronger immune response and could confer enhanced protection against infection.
引文
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