抗WSSV口服VP28亚单位疫苗研究及促克氏螯虾生长口服DNA药物研究
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摘要
白斑综合症病毒（white spot syndrome virus，WSSV）是危害水产养殖业的高致病病原体。1992年白斑综合症病毒首次在中国台湾爆发，而后分别在日本、东南亚、北美等国家和地区爆发，造成虾大面积死亡，对水产养殖业造成了巨大的经济损失。各国研究学者不断探索防治WSSV感染的方法和措施，但是到目前为止还没有实际可行的防治方法问世。
Tat蛋白是HIV-1的转录激活因子，研究发现该蛋白结构域中包含一个11个氨基酸序列的短肽，该短肽能够介导外源物质进入多种细胞或组织中，将其称为蛋白转导域TAT。蛋白转导域TAT作为转运载体已在哺乳动物细胞、昆虫细胞、小鼠中进行了广泛研究，但目前还未见其用于甲壳类动物——克氏螯虾。
在本研究中构建重组质粒pET-28a-VP28; pET-28a-TAT-VP28;pET-28a-VP28-EGFP； pET-28a-TAT-VP28-EGFP，将上述质粒转化到基因工程表达菌株E.coli BL21(DE3)中，诱导表达目的蛋白。将纯化后的VP28-EGFP与TAT-VP28-EGFP融合蛋白喂食克氏螯虾，采用流式细胞术的方法证实，蛋白转导域TAT能够携带与其融合的外源蛋白穿过克氏鳌虾肠壁组织到达血淋巴中；通过荧光免疫组化方法证实TAT-VP28-EGFP融合蛋白存在于克氏螯虾中肠，肌肉和心脏等组织中。
克氏螯虾分别免疫VP28，TAT-VP28蛋白质亚单位疫苗后，测定了血淋巴中酚氧化酶（PO）和超氧化物歧化酶（SOD）酶活力变化。实验结果表明克氏螯虾免疫TAT-VP28蛋白2小时后，血淋巴中的PO及SOD酶酶活均有显著提高；而克氏螯虾免疫VP28蛋白后的血淋巴中PO酶酶活没有变化，24小时后SOD酶酶活有一定提高。这一结果再次证实了TAT-VP28进入克氏螯虾血淋巴后，可以引发克氏螯虾体内的免疫反应。
为了研究蛋白质亚单位疫苗的免疫效果，将VP28，TAT-VP28蛋白包被饲料后，以口服的方式、100μg/只的日剂量免疫克氏螯虾。分别连续免疫7天和14天后，进行病毒攻击实验。实验结果表明与阳性对照组在10天内达到90%以上的死亡率相比，TAT-VP28，VP28组其死亡率分别为43.3%、36.7%；60%、50.0%。为了研究蛋白质亚单位疫苗的免疫时效性。克氏螯虾连续免疫VP28，TAT-VP28蛋白14天，在免疫后的第3和7天分别进行病毒攻击进行病毒攻击实验。实验结果表明与阳性对照组在10天内达到90%以上的死亡率相比，TAT-VP28，VP28组其死亡率分别为43.3%、53.3%%；60%、66.7%。
本研究首次报道和证实以克氏螯虾作为动物模型，VP28蛋白在蛋白转导域TAT介导下能够穿过虾肠壁组织到达血淋巴中。使亚单位疫苗能不通过肌肉注射的方式直接进入血淋巴中，通过血淋巴在虾体内的循环，使其到达到不同的组织中，同时也增强了虾血淋巴中酚氧化酶和超氧化物歧化酶酶活，从而增强克氏螯虾的免疫防御体系。该研究为预防和控制白斑综合症病毒提供了一种实际可行的新思路。
鱼类生长激素(fish growth hormone, fGH)是由鱼类脑垂体合成并分泌的一种蛋白多肽。鱼类生长激素参与蛋白质合成和脂类代谢等生理功能，能够提高鱼类的生长速率，在水产养殖业中具有着重要的研究意义和应用价值。2008年研究学者将罗非鱼生长激素基因构建在真核表达载体中，通过胚胎电穿孔方法，将上述质粒转入到南方滨对虾胚胎受精卵中，与对照组比较幼虾的平均体重有32%的增加。
减毒的鼠伤寒沙门氏菌作为DNA疫苗转运载体目前已经在鼠、雏鸡、鱼和克氏螯虾等体内得到广泛研究和应用，本研究首次将对虾抗菌肽Penaeidin-2信号肽与鲤鱼生长激素基因融合，将其构建到真核表达载体pcDNA3.1-中。用脂质体法将该质粒转染COS-7细胞，用RT-PCR和Western blot的方法证实SGH在人巨细胞病毒(CMV)启动子的控制下在COS-7细胞中得到了高效表达。克氏螯虾幼虾分别注射50μg pcDNA3.1-和pcDNA3.1-SGH后的第3，7，14和21天，实时荧光定量PCR方法检测克氏螯虾血淋巴中Astacidin; Chymotryspin和Trypsin三种酶mRNA表达水平的变化。与对照组pcDNA3.1-比较，pcDNA3.1-SGH组血淋巴中Astacidin; Chymotryspin和Trypsin三种酶的转录水平显著上升。分别将50μg pcDNA3.1-, pcDNA3.1-SGH肌肉注射免疫克氏螯虾幼虾后，与对照组比较，在25天内克氏螯虾幼虾平均体重增加了22.2%以上。
将重组质粒pcDNA3.1-SGH和质粒pcDNA3.1-通过电转的方法转化到减毒鼠伤寒沙门氏菌SV4089中，重组菌分别命名为SV/pcDNA3.1-SGH和SV/pcDNA3.1-。将重组菌SV/pcDNA3.1-SGH和重组菌SV/pcDNA3.1-包被饲料后，以109CFU/只日剂量连续喂食克氏螯虾幼虾十天。为了研究减毒鼠伤寒沙门氏菌SV4089能否有效携带DNA质粒到克氏螯虾幼虾体内，分别在免疫后的第1、3、7、10和14天，分别从克氏螯虾幼虾体内收集血淋巴、游泳足、腮、心脏、肝胰腺和尾肌等不同的组织，提取总RNA，结果在以上组织中检测到了gh的转录。将重组菌SV/pcDNA3.1-SGH和重组菌SV/pcDNA3.1-包被饲料后，以109CFU/只日剂量连续喂食克氏螯虾幼虾十天后，在免疫后的一个月内克氏螯虾幼虾体重增加了21%以上。
本研究首次报道了生长激素口服DNA疫苗促进克氏螯虾幼虾生长，证实了减毒鼠伤寒沙门氏菌SV4089能够将质粒DNA运送到克氏螯虾幼虾体内。这项研究为促进甲壳类动物——克氏螯虾生长提供了一种新思路。
White spot syndrome virus（WSSV） is a highly pathogenic virus that infectsshrimp and other crustaceans. Since the first report of WSSV in Taiwan in1993, it hasrapidly spread to all the major shrimp farming areas in Japan, Southeast Asia, NorthAmerica and other countries. It resulted in a great loss annually. Researchers try tofind effective strategies to prevent and control WSSV. So far there's been no effectiveproduct or practical measure to deal with WSSV.HIV-1Tat protein is a transcriptional activator. The study found that the proteindomain structure, including11amino acid peptide sequence, the short peptides coulddeliver heterologous proteins across most biomembranes without losing bioactivity.TAT protein transduction domain as a delivery vector in mice, rabbits, mammaliancells, insect cells were studied extensively, but has not been reflected for thecrustaceans-Crayfish.
In this study we constructed recombinant plasmids and transformed them intoE.coli strain BL21(DE3) to express TAT-VP28, VP28, TAT-VP28-EGFP andVP28-EGFP fusion proteins. Crayfish in experimental groups were individually fedthe same amount of purified proteins (TAT-VP28-EGFP, VP28-EGFP) via a feedingtube. Immunohistochemistry and flow cytometry methods confirmed that TAT-fusionproteins are able to transduce across shrimp the intestinal wall tissue and go intohemolymph. Strong green-fluorescent signals of EGFP were detected in the inner layer of the mid gut, muscle and heart sections from crayfish fed withrTAT-VP28-EGFP.The purified proteins VP28and TAT-VP28were used to orally immunize C.clarkii.The supernatants of their hemolymph were collected and processed individually toassay the activities of phenoloxidase (PO) and superoxide dismutase (SOD). It wasfound that the rTAT-VP28resulted in the most pronounced increase of PO and SODactivity after2hours immunization. After immunization rVP28, the PO enzymeactivity did not change, SOD activity increase after24hours. This result once againconfirms that TAT-fusion protein are able to transduce across shrimp the intestinalwall tissue and go into hemolymph, the shrimp can cause the immune response.
To evaluate the potential of the subunit vaccine, vaccination experiment wasconducted after shrimps were fed for7and14days respectively. The pelletscontained a daily dose of100μg rVP28and100μg rTAT-VP28per shrimp. Thechallenge test was conducted with an injection of100μl WSSV at7and14dayspost-last feeding. The results showed that with the positive control group at10daysmore than90%mortality.TAT-VP28, VP28group the mortality was43.3%,36.7%;60%,50.0%. In the vaccination experiments, TAT-VP28, VP28and control groupsafter the14days of vaccination, which were subsequently challenged at3and7dayspost vaccination. The results showed that with the positive control group at10daysmore than90%mortality.TAT-VP28, VP28group the mortality was43.3%,53.3%;60%,66.7%.
Our study here demonstrates, for the first time, the C. clarkia is as an animal model,we attempted to utilize the Tat-derived peptide as an oral delivery method for subunitvaccine against WSSV in C. clarkia. TAT-fusion proteins are able to transduce acrossshrimp gut and go into hemolymph and other tissues. After immunization, it wasfound that the rTAT-VP28resulted in the most pronounced increase of PO and SODactivity. So it enhanced the immune defense system of shrimp. The study provides anew way to design more practical strategies for prevent and control WSSV.
Fish growth hormone(fGH)is a polypeptide produced by the pituitary cells ofteleosts, which involved in protein synthesis and lipid metabolism and otherphysiological functions in fish. fGH has been demonstrated to promote growth in fish,it has high economic value in aquaculture areas. Electroporation of Litopenaeusschmitti embryos was used to transfer the plasmid that contains the tilapia growthhormone gene(tiGH), the plasmid was introduced shrimp embryos fertilized eggs, thefollow-up study confirmed that compared with the control group, juvenile shrimpweight increase of32%.
Live attenuated attenuated Salmonella typhimurium have been used as a deliverysystem for DNA vaccines in vertebrate and invertebrate such as mice, poultry, fishand shrimp. The signal peptide of Penaeidin-2and common carp growth hormonegene fusion construct into the pcDNA3.1-. The plasmid was transferred into COS-7,using RT-PCR and Western blot methods confirmed SGH was highly expression.Crayfish were injected intramuscularly with pcDNA3.1-, pcDNA3.1-SGH, usingreal-time fluorescence quantitative method to detect the levels of transcription ofAstacidin; Chymotryspin and Trypsin in hemolymph. The levels of transcription ofAstacidin; Chymotryspin and Trypsin in hemolymph are increased in pcDNA3.1-SGH group comparing to pcDNA3.1-group. So juvenile crayfish were increased more than22.2%body weigh in25days.
The plasmid pcDNA3.1-and pcDNA3.1-SGH were transformed to SV4089byelectroporation and the produced recombinant bacteria were named SV/pcDNA3.1-and SV/pcDNA3.1-SGH, respectively. Recombinant bacteria and recombinantSV/pcDNA3.1-SV/pcDNA3.1-SGH coated feed, to109CFU/daily dose ofcontinuous feeding crayfish juveniles ten days. To investigate whether S.Typhimurium strain SV4089could orally deliver the plasmid pcDNA3.1-SGH intocrayfish and how long the recombinant bacteria could exist in the crayfish,hepatopancreas, swimming foot, gills, heart, hepatopancreas and tail muscle werecollected for isolation of the recombinant bacteria SV/pcDNA3.1-SGH at1,3,7,10and14days post oral administration. For RNA extraction, results in the above tissuesdetected gh transcription. Juvenile crayfish were increased more than21%bodyweigh in one month.
This is the first report of the oral growth hormone oral DNA vaccine, confirming theattenuated Salmonella typhimurium DNA vaccine plasmid to transfer to the crayfishbody. It provides a new way to design more practical strategies for the growth ofjuvenile crayfish.
Tat蛋白是HIV-1的转录激活因子，研究发现该蛋白结构域中包含一个11个氨基酸序列的短肽，该短肽能够介导外源物质进入多种细胞或组织中，将其称为蛋白转导域TAT。蛋白转导域TAT作为转运载体已在哺乳动物细胞、昆虫细胞、小鼠中进行了广泛研究，但目前还未见其用于甲壳类动物——克氏螯虾。
在本研究中构建重组质粒pET-28a-VP28; pET-28a-TAT-VP28;pET-28a-VP28-EGFP； pET-28a-TAT-VP28-EGFP，将上述质粒转化到基因工程表达菌株E.coli BL21(DE3)中，诱导表达目的蛋白。将纯化后的VP28-EGFP与TAT-VP28-EGFP融合蛋白喂食克氏螯虾，采用流式细胞术的方法证实，蛋白转导域TAT能够携带与其融合的外源蛋白穿过克氏鳌虾肠壁组织到达血淋巴中；通过荧光免疫组化方法证实TAT-VP28-EGFP融合蛋白存在于克氏螯虾中肠，肌肉和心脏等组织中。
克氏螯虾分别免疫VP28，TAT-VP28蛋白质亚单位疫苗后，测定了血淋巴中酚氧化酶（PO）和超氧化物歧化酶（SOD）酶活力变化。实验结果表明克氏螯虾免疫TAT-VP28蛋白2小时后，血淋巴中的PO及SOD酶酶活均有显著提高；而克氏螯虾免疫VP28蛋白后的血淋巴中PO酶酶活没有变化，24小时后SOD酶酶活有一定提高。这一结果再次证实了TAT-VP28进入克氏螯虾血淋巴后，可以引发克氏螯虾体内的免疫反应。
为了研究蛋白质亚单位疫苗的免疫效果，将VP28，TAT-VP28蛋白包被饲料后，以口服的方式、100μg/只的日剂量免疫克氏螯虾。分别连续免疫7天和14天后，进行病毒攻击实验。实验结果表明与阳性对照组在10天内达到90%以上的死亡率相比，TAT-VP28，VP28组其死亡率分别为43.3%、36.7%；60%、50.0%。为了研究蛋白质亚单位疫苗的免疫时效性。克氏螯虾连续免疫VP28，TAT-VP28蛋白14天，在免疫后的第3和7天分别进行病毒攻击进行病毒攻击实验。实验结果表明与阳性对照组在10天内达到90%以上的死亡率相比，TAT-VP28，VP28组其死亡率分别为43.3%、53.3%%；60%、66.7%。
本研究首次报道和证实以克氏螯虾作为动物模型，VP28蛋白在蛋白转导域TAT介导下能够穿过虾肠壁组织到达血淋巴中。使亚单位疫苗能不通过肌肉注射的方式直接进入血淋巴中，通过血淋巴在虾体内的循环，使其到达到不同的组织中，同时也增强了虾血淋巴中酚氧化酶和超氧化物歧化酶酶活，从而增强克氏螯虾的免疫防御体系。该研究为预防和控制白斑综合症病毒提供了一种实际可行的新思路。
鱼类生长激素(fish growth hormone, fGH)是由鱼类脑垂体合成并分泌的一种蛋白多肽。鱼类生长激素参与蛋白质合成和脂类代谢等生理功能，能够提高鱼类的生长速率，在水产养殖业中具有着重要的研究意义和应用价值。2008年研究学者将罗非鱼生长激素基因构建在真核表达载体中，通过胚胎电穿孔方法，将上述质粒转入到南方滨对虾胚胎受精卵中，与对照组比较幼虾的平均体重有32%的增加。
减毒的鼠伤寒沙门氏菌作为DNA疫苗转运载体目前已经在鼠、雏鸡、鱼和克氏螯虾等体内得到广泛研究和应用，本研究首次将对虾抗菌肽Penaeidin-2信号肽与鲤鱼生长激素基因融合，将其构建到真核表达载体pcDNA3.1-中。用脂质体法将该质粒转染COS-7细胞，用RT-PCR和Western blot的方法证实SGH在人巨细胞病毒(CMV)启动子的控制下在COS-7细胞中得到了高效表达。克氏螯虾幼虾分别注射50μg pcDNA3.1-和pcDNA3.1-SGH后的第3，7，14和21天，实时荧光定量PCR方法检测克氏螯虾血淋巴中Astacidin; Chymotryspin和Trypsin三种酶mRNA表达水平的变化。与对照组pcDNA3.1-比较，pcDNA3.1-SGH组血淋巴中Astacidin; Chymotryspin和Trypsin三种酶的转录水平显著上升。分别将50μg pcDNA3.1-, pcDNA3.1-SGH肌肉注射免疫克氏螯虾幼虾后，与对照组比较，在25天内克氏螯虾幼虾平均体重增加了22.2%以上。
将重组质粒pcDNA3.1-SGH和质粒pcDNA3.1-通过电转的方法转化到减毒鼠伤寒沙门氏菌SV4089中，重组菌分别命名为SV/pcDNA3.1-SGH和SV/pcDNA3.1-。将重组菌SV/pcDNA3.1-SGH和重组菌SV/pcDNA3.1-包被饲料后，以109CFU/只日剂量连续喂食克氏螯虾幼虾十天。为了研究减毒鼠伤寒沙门氏菌SV4089能否有效携带DNA质粒到克氏螯虾幼虾体内，分别在免疫后的第1、3、7、10和14天，分别从克氏螯虾幼虾体内收集血淋巴、游泳足、腮、心脏、肝胰腺和尾肌等不同的组织，提取总RNA，结果在以上组织中检测到了gh的转录。将重组菌SV/pcDNA3.1-SGH和重组菌SV/pcDNA3.1-包被饲料后，以109CFU/只日剂量连续喂食克氏螯虾幼虾十天后，在免疫后的一个月内克氏螯虾幼虾体重增加了21%以上。
本研究首次报道了生长激素口服DNA疫苗促进克氏螯虾幼虾生长，证实了减毒鼠伤寒沙门氏菌SV4089能够将质粒DNA运送到克氏螯虾幼虾体内。这项研究为促进甲壳类动物——克氏螯虾生长提供了一种新思路。
White spot syndrome virus（WSSV） is a highly pathogenic virus that infectsshrimp and other crustaceans. Since the first report of WSSV in Taiwan in1993, it hasrapidly spread to all the major shrimp farming areas in Japan, Southeast Asia, NorthAmerica and other countries. It resulted in a great loss annually. Researchers try tofind effective strategies to prevent and control WSSV. So far there's been no effectiveproduct or practical measure to deal with WSSV.HIV-1Tat protein is a transcriptional activator. The study found that the proteindomain structure, including11amino acid peptide sequence, the short peptides coulddeliver heterologous proteins across most biomembranes without losing bioactivity.TAT protein transduction domain as a delivery vector in mice, rabbits, mammaliancells, insect cells were studied extensively, but has not been reflected for thecrustaceans-Crayfish.
In this study we constructed recombinant plasmids and transformed them intoE.coli strain BL21(DE3) to express TAT-VP28, VP28, TAT-VP28-EGFP andVP28-EGFP fusion proteins. Crayfish in experimental groups were individually fedthe same amount of purified proteins (TAT-VP28-EGFP, VP28-EGFP) via a feedingtube. Immunohistochemistry and flow cytometry methods confirmed that TAT-fusionproteins are able to transduce across shrimp the intestinal wall tissue and go intohemolymph. Strong green-fluorescent signals of EGFP were detected in the inner layer of the mid gut, muscle and heart sections from crayfish fed withrTAT-VP28-EGFP.The purified proteins VP28and TAT-VP28were used to orally immunize C.clarkii.The supernatants of their hemolymph were collected and processed individually toassay the activities of phenoloxidase (PO) and superoxide dismutase (SOD). It wasfound that the rTAT-VP28resulted in the most pronounced increase of PO and SODactivity after2hours immunization. After immunization rVP28, the PO enzymeactivity did not change, SOD activity increase after24hours. This result once againconfirms that TAT-fusion protein are able to transduce across shrimp the intestinalwall tissue and go into hemolymph, the shrimp can cause the immune response.
To evaluate the potential of the subunit vaccine, vaccination experiment wasconducted after shrimps were fed for7and14days respectively. The pelletscontained a daily dose of100μg rVP28and100μg rTAT-VP28per shrimp. Thechallenge test was conducted with an injection of100μl WSSV at7and14dayspost-last feeding. The results showed that with the positive control group at10daysmore than90%mortality.TAT-VP28, VP28group the mortality was43.3%,36.7%;60%,50.0%. In the vaccination experiments, TAT-VP28, VP28and control groupsafter the14days of vaccination, which were subsequently challenged at3and7dayspost vaccination. The results showed that with the positive control group at10daysmore than90%mortality.TAT-VP28, VP28group the mortality was43.3%,53.3%;60%,66.7%.
Our study here demonstrates, for the first time, the C. clarkia is as an animal model,we attempted to utilize the Tat-derived peptide as an oral delivery method for subunitvaccine against WSSV in C. clarkia. TAT-fusion proteins are able to transduce acrossshrimp gut and go into hemolymph and other tissues. After immunization, it wasfound that the rTAT-VP28resulted in the most pronounced increase of PO and SODactivity. So it enhanced the immune defense system of shrimp. The study provides anew way to design more practical strategies for prevent and control WSSV.
Fish growth hormone(fGH)is a polypeptide produced by the pituitary cells ofteleosts, which involved in protein synthesis and lipid metabolism and otherphysiological functions in fish. fGH has been demonstrated to promote growth in fish,it has high economic value in aquaculture areas. Electroporation of Litopenaeusschmitti embryos was used to transfer the plasmid that contains the tilapia growthhormone gene(tiGH), the plasmid was introduced shrimp embryos fertilized eggs, thefollow-up study confirmed that compared with the control group, juvenile shrimpweight increase of32%.
Live attenuated attenuated Salmonella typhimurium have been used as a deliverysystem for DNA vaccines in vertebrate and invertebrate such as mice, poultry, fishand shrimp. The signal peptide of Penaeidin-2and common carp growth hormonegene fusion construct into the pcDNA3.1-. The plasmid was transferred into COS-7,using RT-PCR and Western blot methods confirmed SGH was highly expression.Crayfish were injected intramuscularly with pcDNA3.1-, pcDNA3.1-SGH, usingreal-time fluorescence quantitative method to detect the levels of transcription ofAstacidin; Chymotryspin and Trypsin in hemolymph. The levels of transcription ofAstacidin; Chymotryspin and Trypsin in hemolymph are increased in pcDNA3.1-SGH group comparing to pcDNA3.1-group. So juvenile crayfish were increased more than22.2%body weigh in25days.
The plasmid pcDNA3.1-and pcDNA3.1-SGH were transformed to SV4089byelectroporation and the produced recombinant bacteria were named SV/pcDNA3.1-and SV/pcDNA3.1-SGH, respectively. Recombinant bacteria and recombinantSV/pcDNA3.1-SV/pcDNA3.1-SGH coated feed, to109CFU/daily dose ofcontinuous feeding crayfish juveniles ten days. To investigate whether S.Typhimurium strain SV4089could orally deliver the plasmid pcDNA3.1-SGH intocrayfish and how long the recombinant bacteria could exist in the crayfish,hepatopancreas, swimming foot, gills, heart, hepatopancreas and tail muscle werecollected for isolation of the recombinant bacteria SV/pcDNA3.1-SGH at1,3,7,10and14days post oral administration. For RNA extraction, results in the above tissuesdetected gh transcription. Juvenile crayfish were increased more than21%bodyweigh in one month.
This is the first report of the oral growth hormone oral DNA vaccine, confirming theattenuated Salmonella typhimurium DNA vaccine plasmid to transfer to the crayfishbody. It provides a new way to design more practical strategies for the growth ofjuvenile crayfish.
引文
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