草鱼与赤眼鳟杂交F_1遗传特征及对草鱼呼肠孤病毒抗性的研究
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摘要
草鱼,中国四大家鱼之一,在我国大宗淡水鱼类养殖中占据举足轻重的地位。目前,草鱼种质资源衰退较为严重,疾病频发。草鱼出血病是草鱼养殖中至今没有得到有效控制的难题,制约着我国水产养殖业的健康可持续发展。随着生物技术发展,着眼草鱼种质资源改良,提高草鱼抗病力可望成为从根本上解决草鱼出血病这一难题的治本途径。远缘杂交是鱼类育种的重要方法之一,并且取得了较好的成果。赤眼鳟,与草鱼同属雅罗鱼亚科,不仅外形上与草鱼相似,还具有抗病力强等优点,是理想杂交对象。为此,本课题研究团队从2010年起开展了草鱼与赤眼鳟的杂交试验,获得了草鱼(♀)×赤眼鳟(♂)正交Fl和赤眼鳟(♀)×草鱼(♂)反交F12个杂交子代群体。为了探索杂交的可行性和杂交子代的杂种优势,本研究对草鱼与赤眼鳟正、反交F1及其父母本子代的外形与红细胞特征、一年龄内的生长、基因组DNA和线粒体基因组DNA多态性及抗草鱼呼肠孤病毒的特性进行比较分析。主要研究结果如下：
1.草鱼与赤眼鳟正、反交F1形态、生长与细胞遗传学比较
5月龄正、反交F1体型偏向于赤眼鳟,而二年龄正、反杂交F1体型、体色和鳞被与草鱼更相似。8个可量可数性状的杂种指数显示正交F1有4个性状偏向草鱼,有4个性状偏向赤眼鳟,而反交F1有5个性状偏向赤眼鳟,3个性状偏向草鱼。
红细胞长径与短径比值以反交F1最小,其余3种鱼无显著差异,说明反交F1红细胞最圆。
池塘养殖条件下,一年龄正、反交F1体重和体长的绝对增长优于父母本子代；正、反交F1增重率介于父母本子代之间,比赤眼鳟小,但优于草鱼。
2.草鱼与赤眼鳟正、反交F1分子遗传学比较
RAPD分析基因组DNA多态性表明赤眼鳟的基因多样性系数和Shannon信息指数最大,正交F1和草鱼次之,反交Fl的最小。遗传同源性和UPGMA聚类分析表明正、反交F1遗传偏向赤眼鳟。
线粒体基因组结构与多态性分析表明,正、反交F1及赤眼鳟线粒体基因组均包括2个rRNA基因,13个蛋白编码基因,22个tRNA基因和2个非编码区；同源性和聚类分析表明正、反交F1与母本的同源性更高。湘江赤眼鳟与长江下游地区(采样点为江苏省)赤眼鳟线粒体基因组序列长度相同,同源性为99%。
3.草鱼与赤眼鳟正、反交F1对草鱼呼肠孤病毒抗性比较
结合RT-PCR和改良RACE技术克隆了正、反交F1Mx基因,cDNA全长分别为2306和2330bp；二者开放阅读框和氨基酸序列长度一致,分别为1884bp和627aa。二者序列结构具有发动蛋白家族典型结构域：N-端发动蛋白GTP酶结构域(DYNc)和C-端发动蛋白GTP酶效应结构域(GED)。多序列比对发现正、反交F1的Mx基因氨基酸序列只有3个差异位点,与草鱼Mx2的同源性高于赤眼鳟Mx。
感染草鱼呼肠孤病毒后,正、反交F1成活率显著高于草鱼,也高于赤眼鳟,赤眼鳟的成活率又高于草鱼。各个血液生理生化指标在4种鱼中的变化趋势基本一致,正、反交F1的T-SOD和IL-1β浓度介于父母本子代之间,C3和IFN-α活力偏母性。感染后96h内,T-SOD整体呈下调趋势,4种鱼对超氧化物的歧化作用降低；至感染后96h,正交F1C3以及正、反交F1IL-1β含量显著提高；4种鱼IFN-α含量在感染后36h内基本处于下调状态,之后上调至正常水平或高于正常水平。
Mx基因和IFN基因在健康草鱼、赤眼鳟及其正、反交F1肝脏、脾脏、肠、鳃、头肾和肌肉中都有表达，但表达量不同；感染草鱼呼肠孤病毒后96h内,Mx基因和1FN基因在4种鱼6种组织中的表达呈波动变化。整体上赤眼鳟和反交F1Mx和IFN基因的表达量高于草鱼和正交F1；杂交子代IFN基因的表达模式与母本同源子代相似,而至感染96h，正交F1IFN基因在各组织中的表达均高于草鱼。Mx基因能被草鱼呼肠孤病毒诱导上调表达,Mx基因上调表达后对IFN基因的表达有一定的抑制作用。
综合以上养殖成活率、血液生理生化指标和抗病毒相关基因的克隆与表达特征的比较结果,赤眼鳟的抗草鱼呼肠孤病毒的能力优于草鱼,正、反交F1抗草鱼呼肠孤病毒的能力也优于草鱼。
Grass carp (Ctenophaiyngodon idellus), one of four chinese carps, occupies a pivotal position in conventional freshwater fish industries of China. However, to date, the germplasm resource of grass carp has been declining, along with a lot of diseases. Grass carp hemorrhagic disease, an epidemic disease caused by grass carp reovirus, is still no effective methods to control and seriously restricts the develpoment of aquaclture in China. With the development of biological techonology, enhancing the disease resistance of grass carp by inproving germplasm resource must be a useful approach to slove the tough problem. Distant hybridization, an important way for fish breeding, has been used in many cases and has good effect.
Barbel chub (Squaliobarbus curriculus), belonging to Leuciscinae as grass carp, is the ideal hybrid object for grass carp. Not only is barbel chub similar to grass carp in appearance but also the barbel chub possesses outstanding capability of disease resistance. Therefore, we carry out the expriment of distant hybridization between grass carp and barbel chub from2010, and we obtain two populations of hybrids:the F1hybrid of grass carp (♀) x barbel chub (♂)(direct cross F1)and the F1hybrid of barbel chub (♀)×grass carp (♂)(reverse cross F1). To explore the feasibility of hybridization and the heterosis of hybrids, the characteristics of appearance, erythrocyte, growth, genome DNA, mitochondrial DNA and the Grass Carp Reovirus (GCRV) resistance of reciprocal cross F1between grass carp and barbel chub were studied. The major results are as follows:
1. Comparative analysis of morphology, growth and cytogenetics
The somatotype of5months old direct cross F1and reverse cross F1were more similar to barbel chub by visual inspection. With age, the characteristics of somatotype, body color, squamation of two years old reciprocal cross F1were more similar to grass carp. Hybrid indexes indicated that four of the eight measurable and numerable characters of direct cross F1were biased toward grass carp, the others were biased toward barble chub; while five of eight of reverse cross F1were biased toward barble chub, three were biased toward grass carp. The size of erythrocyte was no significant difference among grass carp, barbel chub and their direct cross F1, but the ratio erythrocyte long diameter/short diameter of reverse cross F1was minimum.which indicated the erythrocyte of reverse cross F1was most round.
By pond culture, the absolute mean body weight and mean body length of reciprocal cross F1were superior to the F1of grass carp and the F1of barbel chub. The hybrid indexes of weight gain rate of reciprocal cross F1,which were superior to the F1of grass carp but worse than the F1of barbel chub, were intermediate type between two parents
2. Comparative analysis of molecular genetics
Genetic diversity of grass carp, barbel chub and their reciprocal cross F1was analyzed by Random Amplified Polymorphic DNA (RAPD). These results illustrated the Nei's gene diversity and Shannon's Information index of barble chub were highest, then direct cross F and grass carp, while the Nei's gene diversity and Shannon's Information index of reverse cross F1were lowest. The genetic identity and UPGMA analysis indicated the reciprocal cross F1were more similar to barbel chub.
Genetic structure and genetic diversity analysis of mitogenomes showed the composition and organization of mitogenomes of the reciprocal cross F1and barbel chub were the same as that of other cyprinid fish, including two ribosomal RNA genes,13protein-coding genes,22transfer RNA genes, and two non-coding control region. The results indicated genetic structure of fish mitogenome was highly conserved sequence.Genetic identity and cluster analysis results showed the reciprocal cross F1were more similar to female parent, although there were some mutations which were mainly base substitution. The genetic identity between Xiangjiang River barbel chub and the lower reaches of Yangtze River (Jiangsu province) barbel chub was99%
3. Comparative analysis of the characteristic of GCRV resistance
The Mx gene cDNA sequences of the direct cross F1and reverse cross F1,2306bp and2330bp in length respectively, were cloned by RT-PCR and improved RACE methods. The length of their open reading frames and amino acid sequences were both1884bp and627aa. respectively. The two Mx genes were found containing DYNc domain in the N-terminal and GED domain in the C-terminal. There are only three variation sites in amino acid sequences between direct cross F1and reverse cross F1. Multiple sequence alignment showed the genetic identity of Mx between hybrids F1and grass carp was higher than that between hybrids F1and barbel chub.
After challenge with GCRV, the survival rates of reciprocal cross F1were higher than that of barbel chub and significant higher than that of grass carp, while the survival rate of barbel chub was higher than that of grass carp.each blood physiological and biochemical index showed the similar variation trends in the four kinds of fish within96h, respectively. The concentrations of T-SOD and IL-1β were intermediate type between two parents, while the enzyme activities of C3and IFN-a were more similar to female parent. The results indicated the concentrations of T-SOD of grass carp, barbel chub and their reciprocal cross F1were down-regulated within96h after infection by GCRV, which implied the dismutation of superoxide of the four kinds of fish were reduced. At the time point of96h, the enzyme activities of C3of direct cross F1and the concentrations of IL-1β of direct cross F1and reverse cross F1were significantly increased.The enzyme activities of IFN-a of the four kinds of fish were down-regulated in the first36h after infection, then up-regulated to normal level or higher than normal.
qPCR assays showed expression of Mx and IFN transcripts was observed in liver, spleen, intestine, gill, head kidney and muscle of grass carp, barbel chub and their reciprocal cross F1although the levels were different. Within96h after challenge with GCRV, overall pattern of expression of Mx and IFN transcripts was fluctuated in all the tested tissues of four kinds of fish, and the expression of Mx and IFN gene transcripts of reverse cross F1and barble chub were higher than that of direct cross F1and grass carp.The expression pattern of IFN gene of reciprocal cross F1were similar with that of the maternal homologous offspring, but the expression of IFN gene transcripts of direct cross F1was higher than of grass carp in all detected tissues at the time point of96h. It was concluded that Mx gene was inducibly up-regulated by GCRV, but the up-regulated Mx could inhibit the expression of IFN to a certain extent.
Comprehensive analysis of the results of the survival rate, blood physiological and biochemical indexes, cloning and expression of anti-virus genes, showed the GCRV resistance of barbel chub was better than that of grass carp, so were the reciprocal cross F1.
1.草鱼与赤眼鳟正、反交F1形态、生长与细胞遗传学比较
5月龄正、反交F1体型偏向于赤眼鳟,而二年龄正、反杂交F1体型、体色和鳞被与草鱼更相似。8个可量可数性状的杂种指数显示正交F1有4个性状偏向草鱼,有4个性状偏向赤眼鳟,而反交F1有5个性状偏向赤眼鳟,3个性状偏向草鱼。
红细胞长径与短径比值以反交F1最小,其余3种鱼无显著差异,说明反交F1红细胞最圆。
池塘养殖条件下,一年龄正、反交F1体重和体长的绝对增长优于父母本子代；正、反交F1增重率介于父母本子代之间,比赤眼鳟小,但优于草鱼。
2.草鱼与赤眼鳟正、反交F1分子遗传学比较
RAPD分析基因组DNA多态性表明赤眼鳟的基因多样性系数和Shannon信息指数最大,正交F1和草鱼次之,反交Fl的最小。遗传同源性和UPGMA聚类分析表明正、反交F1遗传偏向赤眼鳟。
线粒体基因组结构与多态性分析表明,正、反交F1及赤眼鳟线粒体基因组均包括2个rRNA基因,13个蛋白编码基因,22个tRNA基因和2个非编码区；同源性和聚类分析表明正、反交F1与母本的同源性更高。湘江赤眼鳟与长江下游地区(采样点为江苏省)赤眼鳟线粒体基因组序列长度相同,同源性为99%。
3.草鱼与赤眼鳟正、反交F1对草鱼呼肠孤病毒抗性比较
结合RT-PCR和改良RACE技术克隆了正、反交F1Mx基因,cDNA全长分别为2306和2330bp；二者开放阅读框和氨基酸序列长度一致,分别为1884bp和627aa。二者序列结构具有发动蛋白家族典型结构域：N-端发动蛋白GTP酶结构域(DYNc)和C-端发动蛋白GTP酶效应结构域(GED)。多序列比对发现正、反交F1的Mx基因氨基酸序列只有3个差异位点,与草鱼Mx2的同源性高于赤眼鳟Mx。
感染草鱼呼肠孤病毒后,正、反交F1成活率显著高于草鱼,也高于赤眼鳟,赤眼鳟的成活率又高于草鱼。各个血液生理生化指标在4种鱼中的变化趋势基本一致,正、反交F1的T-SOD和IL-1β浓度介于父母本子代之间,C3和IFN-α活力偏母性。感染后96h内,T-SOD整体呈下调趋势,4种鱼对超氧化物的歧化作用降低；至感染后96h,正交F1C3以及正、反交F1IL-1β含量显著提高；4种鱼IFN-α含量在感染后36h内基本处于下调状态,之后上调至正常水平或高于正常水平。
Mx基因和IFN基因在健康草鱼、赤眼鳟及其正、反交F1肝脏、脾脏、肠、鳃、头肾和肌肉中都有表达，但表达量不同；感染草鱼呼肠孤病毒后96h内,Mx基因和1FN基因在4种鱼6种组织中的表达呈波动变化。整体上赤眼鳟和反交F1Mx和IFN基因的表达量高于草鱼和正交F1；杂交子代IFN基因的表达模式与母本同源子代相似,而至感染96h，正交F1IFN基因在各组织中的表达均高于草鱼。Mx基因能被草鱼呼肠孤病毒诱导上调表达,Mx基因上调表达后对IFN基因的表达有一定的抑制作用。
综合以上养殖成活率、血液生理生化指标和抗病毒相关基因的克隆与表达特征的比较结果,赤眼鳟的抗草鱼呼肠孤病毒的能力优于草鱼,正、反交F1抗草鱼呼肠孤病毒的能力也优于草鱼。
Grass carp (Ctenophaiyngodon idellus), one of four chinese carps, occupies a pivotal position in conventional freshwater fish industries of China. However, to date, the germplasm resource of grass carp has been declining, along with a lot of diseases. Grass carp hemorrhagic disease, an epidemic disease caused by grass carp reovirus, is still no effective methods to control and seriously restricts the develpoment of aquaclture in China. With the development of biological techonology, enhancing the disease resistance of grass carp by inproving germplasm resource must be a useful approach to slove the tough problem. Distant hybridization, an important way for fish breeding, has been used in many cases and has good effect.
Barbel chub (Squaliobarbus curriculus), belonging to Leuciscinae as grass carp, is the ideal hybrid object for grass carp. Not only is barbel chub similar to grass carp in appearance but also the barbel chub possesses outstanding capability of disease resistance. Therefore, we carry out the expriment of distant hybridization between grass carp and barbel chub from2010, and we obtain two populations of hybrids:the F1hybrid of grass carp (♀) x barbel chub (♂)(direct cross F1)and the F1hybrid of barbel chub (♀)×grass carp (♂)(reverse cross F1). To explore the feasibility of hybridization and the heterosis of hybrids, the characteristics of appearance, erythrocyte, growth, genome DNA, mitochondrial DNA and the Grass Carp Reovirus (GCRV) resistance of reciprocal cross F1between grass carp and barbel chub were studied. The major results are as follows:
1. Comparative analysis of morphology, growth and cytogenetics
The somatotype of5months old direct cross F1and reverse cross F1were more similar to barbel chub by visual inspection. With age, the characteristics of somatotype, body color, squamation of two years old reciprocal cross F1were more similar to grass carp. Hybrid indexes indicated that four of the eight measurable and numerable characters of direct cross F1were biased toward grass carp, the others were biased toward barble chub; while five of eight of reverse cross F1were biased toward barble chub, three were biased toward grass carp. The size of erythrocyte was no significant difference among grass carp, barbel chub and their direct cross F1, but the ratio erythrocyte long diameter/short diameter of reverse cross F1was minimum.which indicated the erythrocyte of reverse cross F1was most round.
By pond culture, the absolute mean body weight and mean body length of reciprocal cross F1were superior to the F1of grass carp and the F1of barbel chub. The hybrid indexes of weight gain rate of reciprocal cross F1,which were superior to the F1of grass carp but worse than the F1of barbel chub, were intermediate type between two parents
2. Comparative analysis of molecular genetics
Genetic diversity of grass carp, barbel chub and their reciprocal cross F1was analyzed by Random Amplified Polymorphic DNA (RAPD). These results illustrated the Nei's gene diversity and Shannon's Information index of barble chub were highest, then direct cross F and grass carp, while the Nei's gene diversity and Shannon's Information index of reverse cross F1were lowest. The genetic identity and UPGMA analysis indicated the reciprocal cross F1were more similar to barbel chub.
Genetic structure and genetic diversity analysis of mitogenomes showed the composition and organization of mitogenomes of the reciprocal cross F1and barbel chub were the same as that of other cyprinid fish, including two ribosomal RNA genes,13protein-coding genes,22transfer RNA genes, and two non-coding control region. The results indicated genetic structure of fish mitogenome was highly conserved sequence.Genetic identity and cluster analysis results showed the reciprocal cross F1were more similar to female parent, although there were some mutations which were mainly base substitution. The genetic identity between Xiangjiang River barbel chub and the lower reaches of Yangtze River (Jiangsu province) barbel chub was99%
3. Comparative analysis of the characteristic of GCRV resistance
The Mx gene cDNA sequences of the direct cross F1and reverse cross F1,2306bp and2330bp in length respectively, were cloned by RT-PCR and improved RACE methods. The length of their open reading frames and amino acid sequences were both1884bp and627aa. respectively. The two Mx genes were found containing DYNc domain in the N-terminal and GED domain in the C-terminal. There are only three variation sites in amino acid sequences between direct cross F1and reverse cross F1. Multiple sequence alignment showed the genetic identity of Mx between hybrids F1and grass carp was higher than that between hybrids F1and barbel chub.
After challenge with GCRV, the survival rates of reciprocal cross F1were higher than that of barbel chub and significant higher than that of grass carp, while the survival rate of barbel chub was higher than that of grass carp.each blood physiological and biochemical index showed the similar variation trends in the four kinds of fish within96h, respectively. The concentrations of T-SOD and IL-1β were intermediate type between two parents, while the enzyme activities of C3and IFN-a were more similar to female parent. The results indicated the concentrations of T-SOD of grass carp, barbel chub and their reciprocal cross F1were down-regulated within96h after infection by GCRV, which implied the dismutation of superoxide of the four kinds of fish were reduced. At the time point of96h, the enzyme activities of C3of direct cross F1and the concentrations of IL-1β of direct cross F1and reverse cross F1were significantly increased.The enzyme activities of IFN-a of the four kinds of fish were down-regulated in the first36h after infection, then up-regulated to normal level or higher than normal.
qPCR assays showed expression of Mx and IFN transcripts was observed in liver, spleen, intestine, gill, head kidney and muscle of grass carp, barbel chub and their reciprocal cross F1although the levels were different. Within96h after challenge with GCRV, overall pattern of expression of Mx and IFN transcripts was fluctuated in all the tested tissues of four kinds of fish, and the expression of Mx and IFN gene transcripts of reverse cross F1and barble chub were higher than that of direct cross F1and grass carp.The expression pattern of IFN gene of reciprocal cross F1were similar with that of the maternal homologous offspring, but the expression of IFN gene transcripts of direct cross F1was higher than of grass carp in all detected tissues at the time point of96h. It was concluded that Mx gene was inducibly up-regulated by GCRV, but the up-regulated Mx could inhibit the expression of IFN to a certain extent.
Comprehensive analysis of the results of the survival rate, blood physiological and biochemical indexes, cloning and expression of anti-virus genes, showed the GCRV resistance of barbel chub was better than that of grass carp, so were the reciprocal cross F1.
引文
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